ECOLOGY OF NATIVE OIL-PRODUCING PALMS AND THEIR POTENTIAL FOR

BIOFUEL PRODUCTION IN SOUTHWESTERN AMAZONIA

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By
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JOANNA MARIE TUCKER LIMA
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A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL

OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT
OF THE REQUIREMENTS FOR THE DEGREE OF
DOCTOR OF PHILOSOPHY

UNIVERSITY OF FLORIDA

2010

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© 2010 Joanna Marie Tucker Lima
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To my parents who taught me
to appreciate and marvel at the natural world.
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 ACKNOWLEDGMENTS

This work is the culmination of a long journey and the encouragement and support

of many people throughout my academic career. As this chapter of my life comes to a

close, I wish to extend special thanks to my PhD advisor, Karen Kainer, who was a

continuous and consistent inspiration and source of encouragement to me. Her

expertise and insight enriched my research from beginning to end. I also thank Evandro

Ferreira for his contagious love for palms and mix of practical and scientific advice. His

students, Janice and Anelena, as well as the research team at the Parque Zoobotanico

at UFAC, especially Plinio, Lira, and Edir, were always willing to help with field work and

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logistics, for which I am very grateful. Anelise Regiane and her chemistry students at
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UFAC (Thayna, Nubia and Marcia) gave unselfishly of their time to help me process

palm fruits and run chemical analysis that I could never have done alone. I am indebted
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to Francis “Jack” Putz for sharing his curiosity for the natural world and his persistent

search for answers to both basic and complex ecological issues that affect our daily

lives. His prodding stretched my ideas and encouraged me to look beyond the easy
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answers. I wish to thank Emilio Bruna and Jane Southworth for their insightful feedback

as I developed my dissertation, and for sharing their deep understanding in their fields

of expertise. I am also grateful to Meghan Brennan and Christina Staudhammer for

their brilliant help with statistical analyses.

I am truly grateful for all the financial support I received during my degree from

various sources—NSF Integrative Graduate Education Research and Training Program,

Tropical Conservation and Development Program (UF), International Palm Society, and

the Environmental Protection Agency. I cannot forget to mention the debt I owe to the

principle figures during my academic formation, including Emilio Moran, Eduardo

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Brondizio, Andrew Henderson, and Daniel Zarin. These men inspired me, awakened

my zeal for the Amazon, Brazil, and palm trees, and contributed the foundation, both

academic and personal, that brought me to this point. Friends and family have given

me unconditional support throughout this journey, and I thank them from the bottom of

my heart for their patience, wisdom, listening ears, and encouragement. I especially

thank my parents who unwaveringly stood by me in the good and the difficult times.

Last but not least, I thank my husband, Evandro, without whom this work would have

been impossible. I thank him for his reliability and wisdom as he helped with my

fieldwork, and for sharing his ideas and intimate understanding of Amazonian forests. I

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thank him for his patience and willingness to stick it out during the “last year” of
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finishing-up, and for supporting me unselfishly during that time. Finally, I thank God, for

bringing all these wonderful people into my life, for never letting me go, and for
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providing me the strength, hope, perseverance, intelligence, and faith to complete my

dissertation.
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 TABLE OF CONTENTS

page

ACKNOWLEDGMENTS .................................................................................................. 4 

ABSTRACT ................................................................................................................... 12 

CHAPTER

1 INTRODUCTION .................................................................................................... 15 

2 FLOWER COLOR VARIATION IN ATTALEA PHALERATA (ARECACEAE) ......... 19 

Field Observations .................................................................................................. 21 

Phenology ........................................................................................................ 21 

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Flower Color Variation ...................................................................................... 22 
Color Polymorphism in Palms ................................................................................. 24 
Possible Explanations for Color Polymorphism in Attalea phalerata Flowers ......... 27 
Final Considerations ............................................................................................... 28 

3 DOES LANDSCAPE CHANGE ALTER REPRODUCTIVE PHENOLOGY, SEX

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EXPRESSION AND PRODUCTIVITY OF ATTALEA PALMS IN
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SOUTHWESTERN AMAZONIA? ............................................................................ 30 

Introduction ............................................................................................................. 30 

Materials and Methods............................................................................................ 34 
Study Area........................................................................................................ 34 
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Study Species .................................................................................................. 35 

Site Selection ................................................................................................... 37 
Data Collection ................................................................................................. 37 
Reproductive phenology ............................................................................ 37 
Size and illumination measurements ......................................................... 38 
Palm dissections ........................................................................................ 38 
Statistical Analyses .......................................................................................... 39 
Results .................................................................................................................... 40 
Comparison of Flower and Fruit Production in Forest and Pasture .................. 40 
Flowering and fruiting phenology ............................................................... 40 
Flowering and fruiting frequency ................................................................ 43 
Associations between Reproduction, Light Availability, and Tree Size ............. 47 
Palm Dissections .............................................................................................. 48 
Discussion .............................................................................................................. 49 
Pasture Palms Out-Produced Forest Palms ..................................................... 49 
Phenology ........................................................................................................ 49 
Hidden Mechanisms Control Sex Expression................................................... 50 
Sunlight and Plant Size Linked to Sex Expression and Productivity................. 52 
Sex Expression and Male Dominance .............................................................. 53 

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 Pollen Availability may Alter Operational Sex Ratios ........................................ 55 
Conclusions ............................................................................................................ 56 

4 RECOVERY OF ATTALEA PHALERATA MART. EX SPRENG. PALM

POPULATIONS AFTER SLASH-AND-BURN AGRICULTURE IN
SOUTHWESTERN AMAZONIA .............................................................................. 58 

Introduction ............................................................................................................. 58 

Study Area .............................................................................................................. 61 
Study species ......................................................................................................... 62 
Materials and Methods............................................................................................ 63 
Research Design .............................................................................................. 63 
Data Collection ................................................................................................. 64 
Statistical Analysis ............................................................................................ 65 
Results .................................................................................................................... 66 
Pre-Burn Palm Populations .............................................................................. 66 
Immediate Effects of Slash-and-Burn ............................................................... 68 

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Recovery .......................................................................................................... 69 
Recruitment and survival ........................................................................... 69 
Effect of slash-and-burn on A. phalerata populations ................................ 71 
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Palm demography ...................................................................................... 73 
Leaf production and growth........................................................................ 75 
Discussion .............................................................................................................. 77 
Palm Colonization or Persistence? ................................................................... 77 
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Impediments to seedling recruitment ......................................................... 78 
Mechanisms of persistence........................................................................ 79 
Continued Post-Fire Recovery ......................................................................... 81 
Palm growth ............................................................................................... 82 
Plant demography ...................................................................................... 84 
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Disturbance promotes palm dominance ..................................................... 86 

Ecological Pressures Build Resistance ............................................................ 87 

5 CAN EXISTING POPULATIONS OF NATIVE OLEAGINOUS PLANTS

CONTRIBUTE TO BIOFUEL PRODUCTION IN AMAZONIA? FRUITING
PHENOLOGY AND OIL YIELDS FROM TWO NATIVE ATTALEA PALMS IN
SOUTHWESTERN AMAZONIA .............................................................................. 89 

Introduction ............................................................................................................. 89 

Methods .................................................................................................................. 92 
Site Description ................................................................................................ 92 
Study Species .................................................................................................. 92 
Site Selection ................................................................................................... 94 
Data Collection ................................................................................................. 95 
Reproductive phenology ............................................................................ 95 
Fruit bunch collection and processing ........................................................ 95 
Chemical Analysis ............................................................................................ 96 
Data Analysis ................................................................................................... 97 

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 Results .................................................................................................................... 97 
Comparison of Fruit Production in Forest and Pasture ..................................... 97 
Inflorescence production ............................................................................ 97 
Infructescence production .......................................................................... 98 
Fruiting phenology ..................................................................................... 99 
Productivity and Oil Estimates ........................................................................ 101 
Fruit morphology and yields ..................................................................... 101 
Oil production estimates .......................................................................... 102 
Discussion ............................................................................................................ 104 
Pasture Palms Out-Produced Forest Palms ................................................... 104 
Attalea Species Comparison .................................................................... 107 
Variation, Predictability and Uncertainty .................................................. 107 
What can Attalea Contribute to Energy Portfolios in Rural Amazonia? .......... 109 
Harvest Considerations: Advantages and Disadvantages ...................... 111 
Local-based consumption versus market sale ......................................... 113 
Conclusion ............................................................................................................ 114 

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6 CONCLUSION ...................................................................................................... 115 

LIST OF REFERENCES ............................................................................................. 118 

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BIOGRAPHICAL SKETCH .......................................................................................... 140 
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 LIST OF TABLES
Table page

2-1 Number of yellow and purple-shaded staminate inflorescences of Attalea

phalerata observed in anthesis in pasture and old-growth forest during
monthly visits. ..................................................................................................... 25 

3-1 Comparison of Attalea phalerata and A. speciosa mean (±1 SE) leaf,
inflorescence and infructescence production in old-growth forest and pasture
in Acre and Rondônia, Brazil.. ............................................................................ 46 

3-2 Number of inflorescences (staminate and pistillate) followed through time

until fruit maturity, and percentage of pistillate inflorescence of total
inflorescence for Attalea phalerata and A. speciosa ........................................... 47 

4-1 Attalea phalerata plot densities and mean treatment densities (± 1 SD)
measured in six 50 x 20m control (CTL) and treatment (SLB) plots. .................. 67 

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4-2 Densities per hectare (ha) and percent (%) of original A. phalerata palms that
died within five weeks of forest clearing and burning (T1) in each control plot
(CTL-1, CTL-2, CTL-3) and treatment plot (SLB-4, SLB-5, SLB-6).. .................. 69 

4-3
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Number of newly germinated A. phalerata seedlings ha-1 in each control
(n=3) and treatment plot (n=3) at 2 (T1), 5 (T2), 9 (T3), and 14 months (T4)
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after forest clearing and burning.. ....................................................................... 70 

4-4 Proportion of original A. phalerata individuals surviving per study plot 14

months after slash-and-burn (T5), separated by size class. ............................... 71 
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4-5 Repeated-measures analysis of variance with fixed effects for density by time
and size class using PROC MIXED procedure in SAS ....................................... 72 

4-6 Repeated-measures analysis of variance with fixed effects for relative

abundance by time and size class using PROC MIXED procedure in SAS. ....... 72 

5-1 Mean (± SE) infructescence and fruit characteristics based on seven Attalea
phalerata and ten Attalea speciosa fruit bunches and fruit sub-samples. ......... 102 

5-2 Estimated Attalea phalerata and Attalea speciosa oil production per
infructescence and annually per palm in old-growth forests and pastures in
Southwestern Amazonia. .................................................................................. 103 

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 LIST OF FIGURES
Figure page

2-1 Color variation from yellow to dark purple in Attalea phalerata staminate
inflorescences observed in eastern Acre, Brazil. ................................................ 19 

2-2 Proportion of Attalea phalerata palms with staminate inflorescence and

monthly rainfall (http://www.acrebioclima.pro.br/) from January 2006 until
December 2007. ................................................................................................. 22 

2-3 Purple coloration of petal tips on flowers of an Attalea phalerata pistillate

inflorescence in eastern Acre, Brazil. ................................................................. 23 

3-1 A) Attalea phalerata and B) Attalea speciosa at pasture study sites in Acre
and Rondônia, Brazil, respectively. .................................................................... 36 

3-2 Monthly staminate (male) and pistillate (female) flowering intensities of

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Attalea phalerata in A) old-growth forests and B) pastures with monthly
rainfall (mm) from January 2006 to December 2007 (INMET 2008). .................. 41 

3-3 Monthly staminate (male) and pistillate (female) flowering intensities of

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Attalea speciosa in A) old-growth forests and B) pastures with monthly
rainfall (mm) from January 2006 to June 2007 (original data). ........................... 42 
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4-1 Size class distributions before slash-and-burn (T0) shown as mean
proportion (±1 SD) for control (CTL) (n=3) and treatment (SLB) plots (n=3). ... 68 

4-2 Mean relative abundance of Attalea phalerata size classes in 50 x 20 m plots

at each census before (T0) and 2 (T1), 5 (T2), 9 (T3), and 14 months (T4)
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after forest clearing and burning. ........................................................................ 74 

4-3 Size class distributions on November 1, 2007 (T4), one year after slash-and-
burn disturbance shown as mean proportion (±1 SD) for control (n=3) and
slash-and-burn plots (n=3).................................................................................. 75 

4-4 Proportion of non-reproductive Attalea phalerata individuals that grew at least

one new leaf per census interval in slash-and-burn (n=3) and control plots
(n=3). .................................................................................................................. 76 

5-1 A) A. phalerata and B) A. speciosa infructescences from study sites in Acre

and Rondonia, Brazil, respectively. .................................................................... 94 

5-2 Comparison of mean female inflorescence and mature infructescence

production per palm ±1 SE in old-growth forest and pasture. ............................. 99 

5-3 Monthly fruiting intensity of Attalea phalerata (with mature fruit) and monthly
rainfall from January 2006 to December 2007 (INMET 2008) .......................... 100 

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5-4 Monthly fruiting intensity of Attalea speciosa (with mature fruit) and monthly
rainfall from January 2006 to June 2007 .......................................................... 100 

5-5 Estimated 2006 oil yields (L) per 12 palms from A) A. phalerata endosperm
(kernel) and mesocarp (pulp) and B) A. speciosa endosperm at each study
site in Acre and Rondônia, Brazil, respectively. ................................................ 104 

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 Abstract of Dissertation Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Doctor of Philosophy

ECOLOGY OF NATIVE OIL-PRODUCING PALMS AND THEIR POTENTIAL FOR

BIOFUEL PRODUCTION IN SOUTHWESTERN AMAZONIA

By

Joanna Marie Tucker Lima

May 2010

Chair: Karen A. Kainer

Major: Interdisciplinary Ecology

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In the midst of a worldwide explosion of interest in biofuels, concern has arisen

over displaced food crops, greenhouse gas emissions, forest conversion and
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biodiversity loss. These issues notwithstanding, biofuels are promising as a local fuel

source in rural areas of Amazonia, where communities depend on fossil fuels for
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transportation and electricity generation. This study evaluates the potential of two

native Attalea palm species in southwestern Amazonia as a source of oil for biofuel, and
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generates ecological knowledge to facilitate sustainable management of these species

for oil. Attalea palms are common to forests as well as pastures in Amazonia, and their

fruits contain copious amounts of oil. I compared flowering and fruiting phenologies of

Attalea phalerata Mart. ex Spreng. and Attalea speciosa Mart. ex Spreng. (Arecaceae)

in actively grazed pastures and old-growth forests of Acre and Rondônia, Brazil, over 24

months, and combined these observations with data on palm densities, fruit

characteristics and oil contents to assess biofuel potential. To better understand the

persistence of A. phalerata palms in disturbed landscapes, we also studied the recovery

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of A. phalerata palms after slash-and-burn agriculture using a before-after-control-

impact (BACI) deforestation experiment.

We found greater inflorescence and infructescence production in pastures than in

forests, and generally male-biased inflorescence sex ratios among individuals of both

Attalea species. At the population level, pasture palms demonstrated elevated

reproductive activity, bearing flowers and mature fruits year-round, while reproductive

phenophases of forest palms were more cyclic, with isolated flowering peaks and

consecutive months of inactivity. Light availability was positively correlated with

inflorescence production, and specifically with pistillate inflorescence, although less so

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in A. phalerata than in A. speciosa. The size of the leaf crown (i.e., number of leaves)
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was positively correlated with reproductive activity for both species in pastures, but not

in forests. Total palm height had no apparent effect on inflorescence production.

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Results suggested that female function is more sensitive than male function to

environmental stress and resource limitations. Increased access to light and larger
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crown size shifted Attalea sex expression towards increased female function and

greater productivity primarily during the early stages of bud initiation and development

through the regulation of sex determination and bud abortion.

Just 14 months after deforestation and burning, our analyses showed no overall

effect of slash-and-burn on A. phalerata palm population densities or size class relative

abundances. Pre-adult palms persisted in the slash-and-burn area through resprouting

of new leaves, but there was relatively little germination. Protection of the apical

meristem below-ground and the capacity to continue leaf production following damage

characterizes resprouting in palms were critical to Attalea survival. Within slash-and-

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burn plots, we detected accelerated growth rates in younger palms. Leaf production by

surviving A. phalerata seedlings, post-seedlings, and pre-juveniles was so successful in

slash-and-burn plots that 73% returned to or surpassed their original number of leaves

14 months after disturbance. Our results confirm that A. phalerata populations quickly

recover after slash-and-burn agriculture, and will likely constitute a principal component

of the future regenerating forests in the Southwestern Amazon.

In isolated rural Amazonian communities, naturally occurring populations of

oleaginous plants represent a largely underexploited energy resource and can serve as

an important and immediate source of fuel for electrification and transportation (e.g.,

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river boats, small trucks, or even small airplanes). In a single year, an average A.
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phalerata palm produced more than three times as many infructescences in pasture

than in forest, and A. speciosa produced more than twice as many. A. phalerata palms
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in old-growth forest produced an average 1.17 L palm-1 yr-1 of oil, and in pasture, 4.55 L

palm-1 yr-1, while A. speciosa growing in forest produced 0.93 L palm-1 yr-1 and in

pasture, 2.59 L palm-1 yr-1. Energy production from locally available plant resources
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promises greater independence and improved livelihoods for remote communities, while

assigning greater value to standing forests and diversifying pasture systems already

under use.

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 CHAPTER 1
INTRODUCTION

Over the past decade, interest in alternative energies has soared and biofuels

have quickly expanded into a global commodity (Koh and Ghazoul 2008; FAO 2008;

Tilman et al. 2009). National agendas for energy independence has pushed biofuels to

the forefront of political policies, and proponents have lauded biofuels as a clean

renewable energy source to substitute waning supplies of petroleum-based diesel and

gasoline. At the same time, critics have called for a more careful examination of the

total carbon energy balance of biofuel production, arguing that biofuels can actually

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increase emissions of global warming gases (Fargione et al. 2008; Searchinger et al.

2008), accelerate tropical rain forest destruction and biodiversity loss (Koh and Wilcove
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2007; Groom et al. 2008), and inflate world food prices (Escobar et al. 2009; Rathmann

et al. 2010).
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Against the backdrop of a biofuel boom around the world, this work explores the

ecology of two native Amazonian palm species (Attalea phalerata and A. speciosa) that
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are common across both forested and cleared landscapes in southwestern Brazilian

Amazon, and assesses their potential as sources of locally-derived biofuel. Brazil has

risen to the forefront of liquid biofuel production, use, and technology—particularly

ethanol and biodiesel (da Costa 2004; Brandon 2005; Morgan 2005; Pousa et al. 2007;

Garcez and Vianna 2009), and provides a useful case for exploring the potential of local

and regional biofuel production. In addition to research investments to improve liquid

biofuel crops like castor bean, jatropha, soybean, and African oil palm, the Brazilian

government has funded basic research to identify native oil-producing plants in the

Amazon with potential for biodiesel production.

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 Attalea phalerata and A. speciosa occur abundantly in various habitats of

southwestern Amazonia and contain copious amounts of oil in their fruits. To evaluate

the potential of these species as sources of biofuel, understanding of key ecological

aspects is fundamental. Such knowledge can also offer insights into management

potential, and help understand the impacts of different extraction practices and

intensities on oil yields. These data are largely unavailable for native oleaginous plants

in tropical forest regions (but see Feil 1996; Moraes et al. 1996; Lima et al. 2007),

where energy access looms as an unfulfilled dream for many rural settlements and

isolated communities.

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The potential of native Amazonian oleaginous plants, and in particular palms, has
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been recognized for many years, and has a long history of research and investigation

(Lee 1930; Hodge 1975; Balick 1979; Pesce 1985). Nonetheless, this alternative fuel
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source is largely underutilized and overlooked by local populations as well as

governments. In Amazonia, not only can oils from native plants provide a local and
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reliable source of fuel, they can improve the livelihoods of those who can ill afford

conventional fuels. In the rural areas of Amazonia electricity is rare, although some

homes use diesel generators to provide intermittent electricity. Poor road access

plagues many areas, and rural towns are often isolated from cultural and commercial

centers. Biofuel production from native palm populations could circumvent high fuel

transportation costs and supply locally available fuel for riverboats, trucks, and even

small airplanes frequently used in very remote Amazonian regions. Biofuels extracted

from native plants could bring energy self-sufficiency to remote areas and rural

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settlements, as well as offer the benefits of a clean and sustainable energy source at a

time when climate change and global warming threatens our planet.

African oil palm plantations are frequently cited as a strategy for bringing energy to

remote areas and rural communities in tropical countries. Plantation-based production

of biodiesel in remote rural areas faces various obstacles not only due to isolation, but

also due to scarce economic resources for necessary chemical inputs, technology and

infrastructure, and unpredictable agriculture extension assistance. Plantations also

experience a lag time before fruit production begins, whereas native populations are

already producing. Forest extraction and silvo-pastoral systems that draw upon existing

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populations of native palm species can provide additional income to local producers
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with low additional investment and promote the valuation of standing forests in the

Amazon. Political rhetoric in favor of forest conservation and forest-based development,

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particularly strong in Acre, Brazil, offers hope that the government will support the

extraction of vegetable oil from native palm populations for biodiesel production.
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This research is part of a collaborative effort with an on-going research program at

the Federal University of Acre, Brazil (UFAC), where researchers are investigating the

potential of various native plant populations for the production of biodiesel. The overall

objective of my research was to increase understanding of the ecology of native oil-

producing palms with potential for biofuel production as a local energy source. To

assess biofuel potential, I studied the intricacies of flowering and fruiting phenology of

Attalea phalerata and A. speciosa over two years in pasture and forest—two habitats

where the palms often occur abundantly—to evaluate productivity within each

environment and better understand the factors affecting reproductive biology and

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phenology of these palms (Chapters 2 and 3). I also examined the persistence of A.

phalerata through anthropogenic disturbances, and more specifically, slash-and-burn

agriculture, to understand the regenerative capacity of this species and its

predominance in disturbed landscapes (Chapter 4). Finally, I integrated data on fruiting

phenology, fruit biometry, and oil quantification to estimate oil production potential at the

local scale (Chapter 5). Chapters 2-5 were written as independent, stand-alone

manuscripts for publication, and together they contribute to the overall aim of this work.

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 CHAPTER 2
FLOWER COLOR VARIATION IN ATTALEA PHALERATA (ARECACEAE)

Over the past few years, I have been conducting a phenology study of the

arborescent palm Attalea phalerata Mart. ex Spreng. in southwestern Amazonia.

Although botanical records have consistently reported yellow flowers on A. phalerata

inflorescences, I observed multiple cases of non-yellow staminate flowers. Flower color

varied from dark purple to violet, or a mixture of yellow-orange to magenta flowers

within the same male inflorescence (Figure 2-1; hereafter I refer to non-yellow flowers

as purple). This article reports on field observations of flower color polymorphism in A.

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phalerata and discusses possible explanations for this anomaly.

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Figure 2-1. Color variation from yellow to dark purple in Attalea phalerata staminate
inflorescences observed in eastern Acre, Brazil.

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Figure 2-1. Color from yellow to dark purple in Attalea phalerata, continued.

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 Field Observations

Phenology

I monitored flowering phenology of Attalea phalerata Mart. ex Spreng. in Acre,

Brazil, between January 2006 and December 2007. Using binoculars, I observed

flowering from the ground at monthly intervals at six study sites (three actively grazed

pastures and three areas of old-growth tropical moist forest). At each site I observed 12

reproductive palms. Between July and December 2007, observations were reduced to

two sites per habitat. For each individual I recorded sex and reproductive phase of all

inflorescences – closed inflorescence buds (bracts), inflorescences in anthesis (open

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flowers) and dried post-anthesis inflorescence structures. I also categorized crown
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illumination on a scale of zero to five by counting the number of sides of the palm crown

directly exposed to sunlight (four lateral sides plus top) (Bechtold 2003). Within old-
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growth forest, A. phalerata is mainly a lower canopy palm.

Attalea species alternate between pistillate, staminate, and sometimes

hermaphroditic inflorescences on the same plant. During 24 months of observations of

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72 A. phalerata palms, I registered only four instances of hermaphroditic inflorescences.

The remaining inflorescences were either exclusively pistillate or exclusively staminate.

The majority of A. phalerata staminate inflorescences initiated flowering at the beginning

of the dry season between May and June, peaked in September at the end of the dry

season, and dwindled during the wet season (Figure 2-2). Palms growing in old-growth

forest were more likely to suspend inflorescence production for a short period each year

between February and April, whereas pasture palms produced inflorescences

continuously year-round. Still, pasture palms mimicked the overall seasonal patterns of

flowering peaks and lulls in the forest.

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 2006 2007

Figure 2-2. Proportion of Attalea phalerata palms with staminate inflorescence and

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monthly rainfall (http://www.acrebioclima.pro.br/) from January 2006 until
December 2007 in pastures and old-growth forests in Acre, Brazil.

Flower Color Variation

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To my surprise, of 55 male inflorescences observed in anthesis, the majority
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(55%) produced purple flowers rather than the familiar yellow flowers. During two years

of monthly phenological observations, I recorded 19 purple staminate inflorescences in

anthesis in pastures (on 15 different palms) and 11 in forests (on nine different palms).
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Over the same two-year period, I observed 14 yellow staminate inflorescences in

pastures (on 12 different palms) and 11 in forest (on ten different palms). Some A.

phalerata individuals alternated between purple and yellow flowers, while a few palms

(n=5) repeatedly produced purple flowers. Of the 22 A. phalerata individuals with purple

flowers, more than half (n=12) also produced the better known yellow flowers either

before or after a purple flowering event, indicating phenotypic plasticity within individual

palms. In one case, a single palm exhibited one yellow and one purple inflorescence

simultaneously. Both yellow and purple inflorescences were observed in anthesis, and

the colors remained constant as the flowers developed. I also observed purple

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coloration at the tips of creamy yellow petals of A. phalerata pistillate flowers (Fig. 2-3),

but only three pistillate inflorescences were observed in anthesis during the entire study

period.

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Figure 2-3. Purple coloration of petal tips on flowers of an Attalea phalerata pistillate
inflorescence in eastern Acre, Brazil.

Staminate flower color variation occurred not only within and among individual

palm trees, but also in both the wet (November to April) and dry seasons (May to

October), across different habitats (pasture and forest), and on a regional scale

dispersed over 100 km2. Purple flowers appeared at various times throughout the year,

although mostly during the dry season, which corresponds to A. phalerata’s peak

flowering season (Figure 2-2, Table 2-1). Most flowering anthesis events occurred

between observation visits, and for these I was unable to determine flower color.

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