Palm Tree Susceptibility to Hemi-Epiphytic Parasitism by Ficus

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Palm Tree Susceptibility to Hemi-Epiphytic Parasitism by Ficus
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english
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Kramer,Gregory T
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University of Florida
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Master's ( M.S.)
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University of Florida
Degree Disciplines:
Horticultural Science, Environmental Horticulture
Committee Chair:
Moore, Kimberly A
Committee Members:
Fitzpatrick, George E
Vendrame, Wagner A

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Environmental Horticulture -- Dissertations, Academic -- UF
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Horticultural Science thesis, M.S.
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Abstract:
A survey of palm species located at the Montgomery Botanical Gardens in Miami FL was conducted to investigate if palms with fronds that wither but remained attached to the stem were more susceptible to have Ficus species in the crown and on the plant. The retention of the leaf base allowed for the accumulation of soil and debris which appeared to create a suitable habitat for Ficus seeds to germinate. However, palms with prominent crownshafts (a smooth elongated leaf base) and leaf bases that cleanly fall to the ground were less susceptible to Ficus infestations. The most susceptible palms observed in this study were in the genera Sabal and Phoenix. Palms in these two genera have no crownshaft (no elongated smooth leaf base) and have fronds that remain attached to the stems providing a habitat for Ficus seed germination.
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In the series University of Florida Digital Collections.
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by Gregory T Kramer.
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Thesis (M.S.)--University of Florida, 2011.
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Adviser: Moore, Kimberly A.
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RESTRICTED TO UF STUDENTS, STAFF, FACULTY, AND ON-CAMPUS USE UNTIL 2013-08-31

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1 PALM TREE SUSCEPTIBILITY TO HEM I EPIPHYTIC PARASITISM BY FICUS BY GREGORY KRAMER A THESIS PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PART IAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE UNIVERSITY OF FLORIDA 2011

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2 2011 Gregory Kramer

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3 To my parents for always supporting my curiosity for the sciences and allowing me to follow that curiosity through education

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4 ACKNOWLEDGMENTS I would like to sincerely thank my entire supervisory committee, Dr. Kimberly Moore, Dr George E. Fitz patrick and Dr. Wagner Vendrame for making my learning experience at UF an exceptional one. A special acknowledgment to Dr. Moore, for encouraging me to pursue my degree, and, and for being a constant source of guidance throughout my studies. I would al so like to thank the staff of Mont gomery Botanical Center for allowing me to use the facility to conduct my research, in particular Dr. Patrick Griffith, Executive D irector ; Arantza A. Strader, Database S upervisor ; and Vickie Murphy, Nursery Curator A nd finally t o my entire family who have supported my curiosity for the sciences from a young age, in particular Dave Nancy and Emil

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5 TABLE OF CONTENTS page ACKNOWLEDGMENTS .................................................................................................. 4 LIST OF TABLES ............................................................................................................ 6 LIST OF FIGURES .......................................................................................................... 7 ABSTRACT ..................................................................................................................... 8 CHAPTER 1 INTRODUCTION .................................................................................................... 10 2 LITERATURE REVIEW .......................................................................................... 13 Epiphytes ................................................................................................................ 14 Ficus Hemi epiphytes ............................................................................................. 17 3 MATERIALS AND METHODS ................................................................................ 19 4 RESULTS AND DISCUSSION ............................................................................... 25 5 SUMMARY AND CONCLUSION ............................................................................ 44 LIST OF REFERENCES ............................................................................................... 45 BIOGRAPHICAL SKETCH ............................................................................................ 48

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6 LIST OF TABLES Table page 3 1 List of palm species examined at Montgomery Botanical Center in spring 2011. .................................................................................................................... 21 3 2 F icus species planted at the Montgomery Botanical Center. ............................... 24 4 1 Incidence of Ficus spp. growing as a hemi epiphytic parasite on palms growing at the Montgomery Botanical Center, as influenced by the palm species gross structural characteristics. .............................................................. 29 4 2 Incidence of Ficus spp. growing as a hemi epiphytic parasite on palms growing at Montgomery Botanical Center, as influenced by the palm species gross structural characteristics. .............................................................. 31 4 3 Incidence of Ficus spp. growing as a hemi epiphytic parasite on palms growing at the Montgomery Botanical Center, as influenced by the palm species taxonomi c assignment. ........................................................................... 31 4 4 Incidence of Ficus spp. growing as hemi epiphytic parasite on palms growing at Montgomery Botanical Center, as influenced by the taxonomic tribe of palm species. ........................................................................................... 31 4 5 Incidence of Ficus spp. growing as hemi epiphytic parasites on palms growing at Montgomery Botanical Center, as influenced by taxonomic tribe and age range of palm species. ........................................................................... 32 4 6 Incidence percentage of Ficus spp. growing as a hemi epiphytic parasite on palms growing at Montgomery Botanical Center, as influenced by the palm species, genus, tribe and subfamilies gross structural characteristics. ....... 33 4 7 Percentage of noncrownshaft palms with marcescent leaf bases and hemi epiphytes and the total percentage of palms within all three palm groups sampled with hemi epiphytes ............................................................................... 33 4 8 Average age of non crownshaft, marcescent leaf based palms at Montgomer y Botanical Center which harbor Ficus hemi epiphytes. The ages of non crownshaft marcescent leaf based palms ranged from 52 years old to 10 years old when using plant accession numbers as a gau ge of palm age. ......................................................................................................... 34

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7 LIST OF FIGURES Figure page 3 1 Montgomery Botanical Center, 11901 Old Cutler Road, Coral Gables FL 331564242. ......................................................................................................... 23 4 1 Self sheathing crownshaft palm. Royal palm ( Roystonea regia). ........................ 35 4 2 Self sheathing non crownshaft palm. Triangle palm ( Dypsis decaryi ). ................ 36 4 3 Non self sheathing, non crownshaft palm with marcescent leaf bases Sabal palm ( Sabal palmetto), with Ficus ....................................................................... 37 4 4 Early Ficus hemi epi phytic recruitment in Sabal palmetto leaf base. ................... 38 4 5 Advanced Ficus development upon Sabal palmetto. ........................................... 39 4 6 Ariel root initiation of Ficus growing on Sabal palmetto. ....................................... 40 4 7 Advanced hemi epiphytic parasitism of Ficus tree on Sabal palmetto. ................ 41 4 8 Free standing Ficus tree engulfing Sabal palmetto. ............................................. 42 4 9 Phoenix canariensis with two different Ficus spp. hemi epiphytic parasites. ....... 43

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8 Abstrac t of Thesis Presented to the Graduate School o f the University of Florida in Partial Fulfillment of the Requirements for the Degree Master of Science PALM TREE SUSCEPTIBILI TY TO HEMI EPIPHYTIC PARASITISM BY FICUS By Gregory Kramer August 2011 Chair: Kimberly Moore Major : Environmental Horticulture Palm tree susceptibility to hem i epiphytic parasitism can be attributed to marces cent leaf bases that are typical of certain palm tree species. Palms with marce scent leaf bases seem more susceptible to infestation by the hemi epiphytic parasi tes in the genus Ficus My hypothesis was that palms with cleanly abscissioning leaf bases were e xpressing a phenological adaptation to preclude hemi epiphytic Ficus parasitism. Leaf base retention in palm trees affects establishment of hemi epiphytic parasites by allowi ng the accumulation of detritus, which creates a substrate for Ficus seeds to gem inate within The results from this work showed that palm trees in the group possessing no crownshafts and with marcescent leaf bases were the most susceptible to Ficus hemi epiphytes. The phenological trait of leaf base retention allows for the accumulation of detritus and the subsequent nursery needed for Ficus seed retention and germination. Palms with prominent crownshafts and cleanly abscising leaf bases were observed to be less susceptible, as did the group of non crownshaft palms with cleanly abscising leaf bases. Palm tree susceptibility to Ficus hemi epiphytic parasitism showed varying degrees of parasitism even within a susceptible group such

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9 as non crownshaft palms with marcescent leaf bas es. The two palm genera that seemed the most susceptible to hemi epiphytes were Sabal and Phoenix

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10 CHAPTER 1 INTRODUCTION In the plant kingdom epiphytes are known as pl ants that grow on other plants T he phorophytes ( the support trees) provide the support The epiphytes may or may not obtain their nutrients or water from the plant on which they are growing, as there are many different types of epiphytes found in nature. There are over 28,000 species of epiphytes known in 84 different families (Wilson, 1999). Some of the plant families include cacti, arums, ferns and the most numerous being the orchids. The humid tropical regions of the earth, house the greatest diversity of epiphytes t han anywhere else (Ricklefs, 2007) Some plants seem to be more s usceptible to epiphytes than others. Plant morphology seems to play a big role in epiphyte and hemi epi phyte recruitment and establishment ( Roberts and Male, 2005). One feature that could facilitate epiphyte recruitment could be epidermal texture and roughness. For example, the smooth, hard outer surface of Delonix regina (Boj. Ex Hook.) Raf (a dicotyledonous plant) and Roystonea regia (Kunth.) O.F. Cook (a monocotyledonous plant) seem s to discourage the accumulation of detritus. It is the detritus m aterial which is needed for epiphyte seeds to take hold and germinate. Within the monocotyledonous plants, particularly in the family Arecaceae, an important feature for epiphytic recruitment might be whether leaf bases are marcescent or if leaf bases are senescent with the fronds. Recruitment of hemi epiphytic parasites in palm trees seems to be related to leaf base retention, but not always. In the study conducted by J.R. Mc Pherson (1999) tw o groups of palms were examined: self cleaning palms and palm s with marcescent leaf bases. The number of individual self cleaning palms was 95 and the number of palms

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11 with marcescent leaf bases w as 780. Of the self cleaning palms, 2 were Ficus phorophytes while 90 of the palms with marcescent leaf bases were Ficus phorophytes. O f the studied population of self cleaning palms 2.11% were Ficus phorophytes ; and, o f palms with marcescent leaf bases 11.54% were Ficus phorophytes ( Mc Pherson 1999) In a similar study with tree ferns leaf and leaf base retention was shown to preclude epiphytes and climbing vines in their native habitat of New Zealand (Brownsey and Page, 1986) The retained dead leaves or skirt protect the crown area and subsequent growth bud. Much like the ferns, palms also have one area at t he top of their stems which contains the growth bud and any subsequent damage can kill the plant. U nlike ferns which tend to produce leaves synchronously or in growth spurts palms continually produce leaves during the growing season. Two fern species mentioned, that cleanly sheath their leaves and leaf bases are Cyathea alata (E Fourn.) Copel. of New Caledonia and Cyathea cooperi (W.J. Hook ex F. von Mueller) Domin. of easter n Australia. After the leaves are s hed the naked trunk is smooth ( Brownsey and Page, 1986) Although palms are in the phylum Angiospermophyta and ferns are in the phylum Pteridophyta, there could be a convergent evolution benefit for both plants in diff erent phyla, to cleanly sheath old leaf bases to preclude hemi epiphytic parasites. South Florida is home to many ornamental palm species (Stevenson, 1996) and houses a huge palm nursery industry. In one nursery publication, there are over 46 nurseries l isting for sale over 147 different species of palms in South Florida (Betrocks, 2011) Understanding palm morphology and how it could relate to palm susceptibility concerning hemi epiphytic parasite recruitment could save municipalities and nursery own er s money in their continuing need for hemi epiphytic Ficus removal Little research

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12 has been conducted on the number of hemi epiphytic parasites found growing on palms in South Florida. Palm tree morphology may either induce recruitment or preclude recruitment of epiphytes including hemi epiphyt ic parasites. Ultimately, the survival of the palm could be influenced by those criteria. P alms exhibiting prominent crownshafts and palms without crownshafts yet wit h cleanly abscising leaf bases may have a phenological adaptation to preclude hemi epiphytic parasite recruitment. If this were true, h emi epiphytic recruitment would most like ly be found on palm trees with marcescent leaf bases. The objective of this research was to examine the different phenological structures in palm leaf abscission or retention to answer some of the questions as to why certain palm species seem more susceptible to hemi e piphytic parasites than other palm species.

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13 CHAPTER 2 LITERATURE REVIEW The class of Monocotyledons in the order Arecales conta ins the palm family. I n Arecaceae, there are 250 genera and wi thin those genera there are 2520 species (Dransfeld, et al ., 2008) Palm species can be found on every continent other than Antarctica. Palms trees can also be found growing in a variety of habitats from desert, cool temperate, deciduous subtropical and wet tropical lowlands. The highest number of palm species occurs in wet tropical lowlands ( Craft and Riffle, 2003) The phenological structures within the Arecaceae family vary in many ways The size of the palm, leaf shape, and the presence or absence of a crownshaft are some of the variations. One of t he most important distinguishing features of palm trees relative to hemi epiphytic parasitism is the presence of marcescent leaf bases An example of a palm species commonly found in South Florida with marcescent leaf bases is the cabbage palm, Sabal palmetto (Walt.) Lodd ex Schultes In the absence of leaf bases hemi epiphytic parasites recruitment seems unlikely on palms. The presence or absence of a crownshaft is another important dis tinguishing feature that is used in determining the susceptibility of palms to hemi epiphytic parasites Cleanly leaf sheathing palm tree s with crownshafts are not subjected to hemi epiphytic parasiti sm because they lack leaf bases. An example of a palm species commonly found in South Florida with a crownshaft would be the royal palm, Roystonea regia. Both old leaf bases and the attached leaf frond senescent as the palm continues new leaf production. However some palms do not have crownshafts and do not have marcescent leaf bases. T hese palms may be unsusceptible to hemi epiphytic parasite recruitment. This could be attributed to t he cleanly sheathing leaf bases, leaving no

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14 significant for epiphyte attachment. A commonly found species in South Florida with out a crownshaft and sheathing leaf bases in Bismarckia nobilis Hildebrandt and H. Wendl. commonly called the bismark palm Epiphytes T here are many different types of epiphytic plants that have different types of relationships to the plant on which t hey are growing. For example, some epiphytes have a commensalistic relationship with their host This means that one plant will benefit from the host plant but the host plant is neither affected positively or negatively by the epiphytes presence ( Wilson 1999) A good example of this type of commensalistic relationship would be Ti llandsia usneoides (L.) L., S p anishmoss growing on Quer cus virginiana L., live o ak trees Taxodium distichum (L.) A. Rich, baldcypress trees and other tree species. S panishmoss having no functional root structures, grows on the braches of the trees to gain access to sunlight It obtains nutrients from the air and rain leachate, which is absorbed thr oughout the leaf scales (Garth, 1964) T here are some epiphytes that have a parasitic relationship with their host plant. These epiphytes not only live on their host tree for support but derive inorganic minerals and water from their host plant and cannot live without them ( Poindexter and Thompson, 2005) A good example of a parasitic epiphyte is Phoradendron leucarpum (Raf.) Reveal and M.C. Johnst. A merican mistletoe. American mistletoe is an evergreen hemi parasite that can grow to three feet tall and three feet round (A lden et al ., Nelson, 1999) American mistletoe can be found parasitizing an array of deciduous trees in its native range of eastern North Americas temperate c limate (USDA, 2011) In one study conducted in the state of Kentucky these s pecies were found to be most susceptible in descending order : Juglans nigra L. black walnut ; Prunus serotina Ehrh.

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15 black cherry ; Ulmus americana L. A merican elm ; Robinia pseudoacacia L. black locust ; and, Fraxinus a mericana L. white ash (Poindexter and Thompson, 2005 ) Mistletoe is usually found on the stems and branches of these trees being an epiphyte and having no true roots Phoradendron l eucarpum (Raf .) Reveal and M.C. Johnst. has specialized root like structures called haustori (Hemmerly et al 1987) that penetrate the host cambium to derive the needed minerals and water for plant metabolism as well as stabilizing the plant in the tree canopy. E p iphytes that never grow in a soil medium and complete their life cycle having never grown in a soil substrate are called holoepiphytes. All of a holoep ip hyte s nutritional needs are sources obtained either through water or leachate from the host plant A good example of this type of plant is Tillandsia usneoides ( Garth 1964). Primary hemi epiphytes begin life as an epi phyte on another plant and their roots grow subsequently dow nw ard toward the soil They began their lif e having germinated and started their growth in a small pocket of humus that had accumulated somewhere on the host plant. The location of humus accumulation on the host plant could have been a branch crotch, branch or trunk wound, or in the case of palm trees leaf bases retained on the trunk. Some examples of plant genera that are considered primary hemi epiphytes would be Ficus Clusia and Shefflera. Secondary hemi epiphytes initially began life rooted in the soil and secondarily became h emi epiphytes as they matured disconnecting their root system from the terr estrial soil substrate (Kato and Tsutsumi, 2006). Some examples of genera which are considered secondary hemi epiphytes are Philodendron in the Araceae family (Cro at, 1997) and Nephrolepis in the Lomariopsidaceae family.

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16 It is the primary hemi epiphytes predominately in the genera Ficus that are of concern to palm growers Parasitic hemi epiphytes like Ficus begin their life as an epiphyte grow ing on another plant and then produce roots that continually grow downward toward the soil as the plant matures Once the roots touch the ground the plant is no longer an epiphyte. This relationship usually does not quickly kill the host plant ; but as the hemi epiphyte continues growing, added weight can cause limb breakage, and subsequent shading of the host plant leaves can decrease photosynthesis. A hemi epiphytic parasite begins its life cycle as an epiphyte and continues to grow and produce aeria l roots toward the ground. Yet unlike all of the previous groups of epiphytes thi s group parasitizes and eventually kills its host tree. This method of parasi tization is different from the A merican mistletoe, in that there is not direct parasitization of the host trees vascular system. The hemi epiphyte uses the host plant (phorophyte) for support and access to sunlight. Through competition for sunlight, nutrients and vascular constriction of t he phloem and xylem tissues (Argo, 1 964) the host plant is overwhelmed and perishes (Todzia, 1986) There are some ways in which being an epiphyte could be advantageous. Growing in the tree canopy allows t he epiphytes access to sunlight that might be reduced when growing on the ground surrounded by large trees. Growing high in the tree canopy may also protect epiphytes from fire that could occur when growing on the ground ( Putz and Susilo 1994) When a study in the Venezuelan llanos was conducted, along with other fire maintained plant communities, the ground fires did not reach into the palm canopies and kill the hemi epiphytes. Any aerial root damage resulting from the fire was replaced by new downward root production.

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17 There are also many challenges to growing as a hemi epiphyte. Hemiepiphytes are subjected to extremes of moisture availability and temperature. One particular type of hem i epiphytic parasite, Clusia rosea Jacq. has been shown to employ c rassulacean acid metabolism (CAM). The uptake of CO occurs when plant stomata are open at night. This adaptation makes the plant less susceptible to desiccation by having stomata closure during the day and t he CAM method of carbon sequestion is more commonly found in arid adapted plants like cacti Yet hemi epiphytic plants are not in contact with the soil medi um and are subjected to drought like conditions even when growing in a wet environment (DeNiro et al ., 1985) Ficus H emi epiphytes The genus Ficus in the family Moraceae has approximately 750 species worldwide and is mostly distributed in the tropics and subtropics (Frank et al 1992). In Florida, there are two native species restricted to th e southern portion of the state. The species native to Florida are Ficus citrifolia Mill. the shortleaf fig (Frank et al ., 1992) and, Ficus aurea Nutt the strangler fig ( Alden et al 1998). There are numerous nonnative species of Ficus that have been planted as ornamental plants and are naturalizing around the southern portion of the state of Florida. In one study over 60 nonnative Ficus species were documented to have naturalized, including, Ficus altissima Blume, Ficus benghalensis L. and Ficus microcarpa L. T here is the potential for two other Ficus species to natur alize Ficus benjamina L. and Ficus religiosa L. (Frank et al ., 1992) There are several methods for hemi epiphytes dispersal agents and subsequent recruitment into susceptible palm host species. Ficus fruit is mammalian and avian

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18 dispersed. Many animals that are frugivors after feasting on Ficus fruits will seek shelter for roosting in a thick canopy of leaves suc h as a palm crown (Todzia, 1986) When the animals defecate, if the palm has marcescent leaf bases that have accumulated detritus the Ficus seeds will have a substrate in which to germinate. Also, being in a palm canopy allows for the high light requirements that some species of Ficus need in order to germinate, thus possibly giving them a competitive advantage over seeds that fall to the ground. Not all Ficus fruits survive pass ing through the digestive tracks of birds in fact some species of birds can be considered seed predators. Columba leucocephala L. the whitecrowned pigeon can digest both the fruit and many of the seeds ingested to produce pigeons milk that is fed to the ir young birds. In a study conducted on crop content of whitecrowned pigeons, it was found that early in the season, Ficus spp. dominated the contents (both Ficus aurea Nutt. and Ficus citrifolia Mill. ) of the birds crop, by as much as 60% As the season progressed, the content of Ficus seed s dropped to 2% (Bancroft and Bowman, 1994) Another agent of seed dispersal of hemi epiphytic parasites in palm crowns are seed harvesting ants. Ants in the genus Pheidole were found to be Ficus seed predators ; but in the process of carrying seeds back to the nest the ants may drop or discard seeds. If by chance a Ficus seed was to land in a favorable area like a palm crown with marcescent leaf bases and an accumulation of humus hemi epiphytic establishment could ensue. When Pheidole spp. n ests in tree canopies were examined, seed r eserves were f ound to be uneaten (Laman, 1996. ) This could possibl y increase Ficus hemi epiphytic establishment in those trees with uneaten seeds

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19 CHAPTE R 3 MATERIALS AND METHOD S The research site was t he Montgomery Botanical Center, in the c ity of Cora l Gables F lorida (11901 Old Cutler Road, Coral Gables, FL 33156) Latitude: 25 39' 46.9" N Longitude: 80 16' 51.1" W This botanical garden is 120 acres in area and is noted for its collection palms and cycads (Fig 3 1) The garden was established in 1932 as the private residence of Col onel and Mrs. Robert H. Montgomery and it is noted for its programs in plant conservation, advancing science and education of tropical plants. The Montgomery Botanical Center was established as a not for profit organization in 1959, after Col. Montgomerys death. The southeastern and northeastern portions of t he property are know n as the lowland s Parts of the lowlands had been land fil led in the 1930s There were reported to have been over 300 different varieties and species of palms planted on the property during the 1930s (Montgomery 1935). Currently, existing portions in the lowland region not land filled are mangrove estuaries co nsisting of Rhizophora mangle L. red mangrove; Laguncularia racemosa (L.) Gaertn. f. white mangrove; and Schinus terebinthifolius Raddi Brazilian pepper M ost of the land area of the garden is located on the Miami Ridge and is around 21 feet above sea level. The other surrounding portions of the property outside the garden are highly suburbanized and planted with an array of both native and exotic plants. Each plant located within Montgomery Botanical Ce nter has an accession number and can be found using a formatted grid system A survey was conducted by walking the grounds of the garden and observing palm trees with hemi epiphytic Ficus recruitment (Table 3.1) W hen a species of palm was found to have a hemi-epiphytic Ficus growing on it it was recorded and added to the survey, which included the plant accession

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20 number A data print out of all palm species being studied was provided by the garden administration. Palms examined included: Prominent c rownshaft, non crownshaft abscis sioning leaf bases and non crownshaft marcescent leaf bases (Table 31) A comparison was conducted and individual palm species found to have hemi epiphytic Ficus recruitment were noted and compared to the total number of palms species without hemi epiphytes found on the property. In addition, if one individual palm was found to have hemi epiphytic Ficus recruitment it was added to the survey list and compared to other palms within the same species found on the property Differe nces among groups were detected by using chi square analysis. T he survey only noted the presence or absence of Ficus hemi epiphytes and not the species of Ficus Some palms had well established large Ficus hemi epiphytes and other palms had smal l plants beginning initial recruitment. A list of Ficus plants accessioned was provided by Montgomery Botanical Center (Table 32). There are 14 species of Ficus planted on the Montgomery Botanical Centers property. Of those 14 species, 2 are native to the state of Florida: Ficus aurea L. and Ficus citrifolia L. Palm species that where short enough to observe directly for hemi epiphytic Ficus species were done so without the aid of binoculars. For t rees that wer e tall er the aid of binoculars was needed to look for hemi epiphytes. The binoculars that were used were the brand Konus Vue with a field r esolution of 7x50. This resolution was sufficient to clearly look into the tallest of the gardens palm crowns.

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21 Table 3 1 L ist of palm species examined at Montgomery Botanical Center in spring 2011. Palm species are listed based on difference in specific phenological characteristics. Number of individuals observed Prominent crownshaft 1) Archontophoenix alexandrae 6 2) Archontophoenix cunninghamia 2 3) Archontophoenix myolensis 18 4) Archontophoenix purpurea 1 5) Archontophoenix tuckeri 2 6) Carpentaria acuminata 19 7) Chambeyronia macrocarpa 16 8) Dictyosperma album 21 9) Dypsis lutescens 7 10) Hyophorbe indica 10 11) Hyophorbe lagenicaulis 22 12) Hyophorbe verschaffeltii 16 13) Normanbya normanbyi 8 14) Roystonea altissima 14 15) Roystonea borinquena 32 16) Roystonea lenis 4 17) Roystonea oleracea 58 18) Roystonea princeps 1 19) Roystonea regia 152 20) Pseudophoenix ekmanii 1 21) Pseudophoenix sargentii ssp. s argentii 2 22) Pseudophoenix sargentii ssp. var navassana 4 23) Pseudophoenix vinifera 33 24) Veitchia arecina 43 25) Veitchia joannis 1 26) Veitchia spiralis 13 27) Veitchia winn 10 No prominent crownshaft; abscis sion ing leaf bases 1) Acoelorraphe wrightii 46 2) Bismarckia noblis 47 3) Coccothrinax argentea 12 4) Coco nucifera 14 5) Dypsis decaryi 25 6) Heterospathe elata 25 7) Washingtonia robusta 12 No prominent crownshaft; marcescent leaf bases 1) Attalae butyracea 40 2) Attalea cohune 62 3) Attalea phalerata 31 4) Butia capitates var odorata 14 5) Butia eriospatha 10 6) Caryota rumphiana 23 7) Copernicia baileyana 14 8) Elaeis guineensis 31 9) Hyphaene coriacea 48 10) Hyphaene petersiana 7

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Table 31. Continued. 22 Number of individuals observed 11) Hyphaene thebaica 16 12) Livistona chinensis 59 13) Livistona decora 98 14) Livistona saribus 50 15) Phoenix canariensis 46 16) Phoenix reclinata 14 17) Sabal domingensis 67 18) Sabal maritima 29 19) Sabal mexicana 3 20) Sabal palmetto 357 21) Sabal uresana 1 22) Syagrus coronata 30 23) Thrinax radiata 156

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23 Figure 31. Montgomery Botanical Center, 11901 O ld Cutler Road, Coral Gables, Florida. Latitude: 25 40' N Longitude: 80 17' W Photo courtesy of Montgomery Botanical Center.

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24 Table 32 Ficus species planted at the Montgomery Botanical Center Name Number Observed Ficus afghanistanica 1 Ficus altissima 2 Ficus aurea 5 Ficus benjamina 2 Ficus citrifolia 1 Ficus dammaropsis 2 Ficus fraseri 1 Ficus hirta 1 Ficus lyrata 3 Ficus microcarpa var nitida 1 Ficus mysorensis 1 Ficus racemosa 2 Ficus subcordata 1 Ficus trigonata 1

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25 CHAPTER 4 RESULTS AND DISCUSSION Palms with prominent crownshafts and cleanly senescing leaves were not observed to host any Ficus hemi epiphytic parasites (Table 41; Fig. 4 1). Palm trees without crownshafts and senescing leaves also were not observed to host Ficus hemi epiphytic parasites (Table 41; Fig. 4 2). The palms that hosted the most Ficus hem i epiphytic parasites were palms with marcescent leaves. For example, 38 out of the 357 Sabal palmettos at Montgomery Botanical Center had Ficus hemi epiphytes (Fig. 4 3) Even among palms with marcescent lea f bases, some palms seemed to favor hem i epiphytic parasitism (Table 4 2). The old world genera of palms Hyphaene and Phoenix appear to frequently host hemi epiphytes Of the new world palm genera, Sabal appears to be the most susceptible. Of the sampled population of non crownshaft palms with marcescent leaf bases, 10% were found to host hemi epiphytes (Table 4 7 ). When compared to the total sampled palm population, including prominent crownshaft palms, non c rownshaft senescing leaf palms and non crowns haft marcescent leaf palms 6.5% of the palm population hosted hemi epiphytic parasites (Table 4 7 ). When all three palm categories were examined, there was no evidence of hemi epiphytic Ficus establishment occurring either on the palm trunks or within the palm crowns of the 516 crownshaft palms surveyed. In addition to the crownshaft palms surveyed, the 181 non crownshaft palms with absci ssion ing leaf bases did not show Ficus establishment (Table 4 2). Wi thin the non crownshaft palms with marcescent leaf bases there was varying degrees of establishment (Table 4 1). Within the non crownshaft, marcescent leaf based palms; it was quite surprising to find two species of palms that harbored hemi epiphytic par asit es The two species

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26 included the native Thrinax radiata Lodd ex Schultes and Shultes f. and the introduced Caryota rumphiana Mart. Through research and eye witness accounts these two palm species had not appeared to be susceptible to hemi epiphytic parasitism before this study. Although the morphological requirements of leaf base retention were present, hemi epiphytes were rarely seen. In the palm genera Sabal and Phoenix hemi epiphytic recruitment was not surprising as in the palm genera Thrinax and Caryota. Both Sabal (Fig. 45) and Phoenix are two palm genera that are commonly found functioning as phorophytes for Ficus hemi epiphytes in South Florida. In the palm population at Montgomery Botanical Center 10.6% of the population of Sabal palmettos had Ficus hemi epiphyte present while Phoenix canariensis had 43% Ficus present (Table 46 ). For Sabal palmetto, 1 out of every 9 trees had Ficus recruitment either occurring within the palm crown or along the palm trunk. For Phoenix canariensis it was almost 1 out of every 2 trees had Ficus recruitm ent either in the pal m crown or along the palm trunk. Both of thes e palm species are commonly planted around the State of Florida including South Florida where hemi epiphytes are very common. Given such a high incidence of hemi epiphytic parasitism occurring on Sabal palmetto and Phoenix canariensis maintenance requirements will be required to remove Ficus hemi epiphytes regularly to insure palm aesthetics in la ndscapes ( Fig 4 7) and palm survival (Fig 4 8). The elimination of Ficus hemi epiphytes in South Florida would be almost impossible given the nature of Ficus seed dissemination, which is mainly by birds and ants. Birds continually use palm crowns for roosting and often defecate seeds from a previously e aten Ficus meal T his behavior may insure a continual influx of seeds upon a palm susceptible to hemi epiphytes (Argo 1964). The following birds were observed

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27 eating the fruit of Ficus trees in the study plot of Montgomery Botanical Center one native species and two introduced species. The native species was Mimus polyglottos L., northern mockingbird and the two introduced species were Sturnus vulgaris L. ,starling and Gracula religiosa L. hill mynah. One species of parrot Bro togeris vericol u rus Muller., canary winged parakeet was also noted in the Ficus tree; but, it could not be determined if the parrots were indeed eating the fruit. Some birds may spread Ficus seeds by a behav ior know n as scatter hoarding (Grubb et al 1997). This method of hiding or hoarding food could play an important dispersal agent for Ficus seeds into tree crotches and palm tree cr owns. Yet some birds are seed predators and would be unlikely candidates for Ficus seed dispersal. Columba leucocephala L. white crowned pigeon, is a Ficus seed predator. This could include the white crown pigeons more common cousin Zenaida macroura L mourning dove (Hayslette and Pol ing 2006) and the introduced Streptopelia decaocto Frivaldszky, E urasian collared dove. Thus, by the nature of being seed predators these three bird species would be unlikely candidates for spreading Ficus seeds into palm crowns. With the many choices of palm species that can be planted in South Florida, palms with cleanly abscising leaf bases m a y be a better option to plant in the landscape than palms with marcescent leaf bases. When crownshaft palms and non crownshaft, leaf based abscis sioning palms are compared to non crownshaft leaf base marcescent palms with Ficus hemi epiphytes, there is a statistically significant difference between Ficus hemi epiphytic s usceptibility in taxomomic tribes (Table 44) Additionally, there was a 95% chance that these groups were statistically different based on species (Table 4 2 ) This statist ical outcome seems to parallel the initial survey data that showed no Ficus hemi epiphytic recruitment

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28 occurring on either crownshaft palms or non crownshaft pal ms with abscis sioning leaf bases. The phenological adaptation of abscising leaf bases does not allow either detritus to collect under leaf bases or Ficus seed recruitment. The question of whether cleanly sheathing leaf bases are a direct phenological adaptation to preclude hemi epiphytic Ficus recruitment would require more research. However, it a ppear s that palms that lose their leaf bases have an ecological advantage over palms that retain their leaf bases in precluding hemi epiphytic parasiti sm in environments where Ficus species are present Whereas non crownshaft palms with marcescent leaf bases may act as nurseries (Dirzo and Lopez, 2007) (Fig 4 4) for Ficus hemi epiphyti c parasitism putting them at a disadvantage. In one study conducted in Mexico on Sabal mexicana, the epiphytic soi l was found to contain 40times the carbon, 14times the nitrogen and 12times the phosphorus than the soil in which the palm was growing (Aguirre et al ., 2010). It is not unusual to find several different species of Ficus occurring on the same palm when looking at non crownshaft ma r ces cent leaf based palms around South Florida (Fig 4 9). O lder taller palms had the largest and most established Ficus hemi epiphyt ic r ecruitment, some already in the process of producing aerial roots ( Table 4 8; Table 45; Fig. 4 6). It is probable that Ficus hemi epiphytes are not removed as often from taller palms due, to the difficulty in reaching great er heights as those hemi epiphytes that can be reached from the ground or with the aid of a ladder. In some of the palms surveyed, easily a ccessible p alms had Ficus that was flushing out new growth after previously having been cut if thick established roots were retained on the palm.

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29 Table 41 Incidence of Ficus spp. growing as a hemi epiphytic parasite on palms growing at the Montgomery Botanical Center, as influenced by the palm species gross structural characteristics. Number of individuals observed with no hemi epiphytes Number of individuals observed with hemi epiphytes Prominent crownshaft; abscis sion ing leaf bases 1) Archo ntophoenix alexandrae 6 0 2) Archontophoenix cunninghamiana 2 0 3) Archontophoenix myolensis 18 0 4) Archontophoenix purpurea 1 0 5) Archontophoenix tuckeri 2 0 6) Carpentaria acuminata 19 0 7) Chambeyronia macrocarpa 16 0 8) Dictyosperma album 21 0 9) Dypsis lutescens 7 0 10) Hyophorbe indica 10 0 11) Hyophorbe lagenicaulis 22 0 12) Hyophorbe verschaffeltii 16 0 13) Normanbya normanbyi 8 0 14) Roystonea altissima 14 0 15) Roystonea borinquena 32 0 16) Roystonea lenis 4 0 17) Roystonea oleracea 58 0 18) Roystonea princeps 1 0 19) Roystonea regia 152 0 20) Pseudophoenix ekmanii 1 0 21) Pseudophoenix sargentii ssp s argentii 2 0 22) Pseudophoenix sargentii ssp var navassana 4 0 23) Pseudophoenix vinifera 33 0 24) Veitchia arecina 43 0 25) Veitchia joannis 1 0 26) Veitchia spiralis 13 0 27) Veitchia winn 10 0 No prominent crownshaft; absci s s ion ing leaf bases 1) Acoelorraphe wrightii 46 0 2) Bismarckia noblis 47 0 3) Coccothrinax argentea 12 0 4) Coco nucifera 14 0 5) Dypsis decaryi 25 0 6) Heterospathe elata 25 0 7) Washingtonia robusta 12 0 No prominent crownshaft; marcescent leaf bases 1) Attalea butyracea 40 4 2) Attalea cohune 62 1 3) Attalea phalerata 31 7 4) Butia capitata var. odorata 14 3 5) Butia eriospatha 10 1 6) Caryota rumphiana 23 1 7) Copernicia baileyana 14 1

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Table 4.1. Continued. 30 Number of individuals observed with no hemi epiphytes Number of individuals observed with hemi epiphytes 8) Elaeis guineensis 31 3 9) Hyphaene coriacea 48 9 10) Hyphaene petersiana 7 6 11) Hyphaene thebaica 16 1 12) L ivistona chinensis 59 4 13) Livistona decora 98 1 14) Livistona saribus 50 5 15) Phoenix canariensis 46 20 16) Phoenix reclinata 14 1 17) Sabal domingensis 67 4 18) Sabal maritima 29 4 19) Sabal mexicana 3 3 20) Sabal palmetto 357 38 21) Sabal uresana 1 1 22) Syagrus coronata 30 5 23) Thrinax radiata 156 1

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31 Table 42 Incidence of Ficus spp. growing as a hemi epiphytic parasite on palms growing at Montgomery Botanical Center, as influenced by the palm species gross structural characteristics. The distribution of palm species with Ficus spp. i nfestation is significantly different between groups as determined by the test with 4 degrees of freedom ( =76.7; P<0.05). Structural characteristics Number of individuals observed with no epiphytes Number of individuals observed with epiphytes Total number of individuals observed Prominent crownshaft 516 0 516 No prominent crownshaft; abscis sion ing leaf bases 181 0 181 No prominent crownshaft; m arcescent leaf bases 1,082 124 1,206 Table 43 Incidence of Ficus spp. g rowing as a hemi epiphytic parasite on palms growing at the Montgomery Botanical Center, as influenced by the palm species taxonomic assignment The distribution of palm species with Ficus spp. i nfestation is not significantly different between these subfamilies as determined by the test with 2 degrees of freedom ( =0.153; P>0.05). Subfamily within the family Arecaceae Numbers of individuals observed with no epi phytes Number of individuals o bserved with epiphytes Total number of individuals observed Arecoideae 194 24 218 Coryphoideae 888 100 988 Table 44 Incidence of Ficus spp. g rowing as hemi epiphytic parasite on palms growing at Montgomery Botanical Center, as influenced by the taxonomic tribe of palm species. The distribution of palm tribes with Ficus spp. i nfestation is significantly different between palm tribes as determined by the test with 12 degrees of freedom ( =74.9; P<0.05). Tribes Number of individuals observed with no epiphytes Number of individuals observed with epiphytes Total number of individuals observed Broasseae 55 16 71 Cocesea 194 24 218 Corypheae 22 1 23 Cysophileae 155 1 156 Phoeniceae 39 21 60 Sabaleae 407 50 457 Trachycarpeae 210 11 221

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32 Table 45. Incidence of Ficus spp. growing as hemi epiphytic parasites on palms growing at Montgomery Botanical Center, as influenced by taxonomic tribe and age range of palm species. The distribution of palm tribes with Ficus spp. infestation is not significantly different between ages as determined by the test with 24 degrees of freedom ( =35.8; P<0.05) Tribes Number of Individuals observed with epiphytes from 0 10 years of age Number of individuals observed with epiphytes from 10 20 years of age Number of individuals observed with epiphytes from 20 30 years of age Number of individuals observed with epiphytes from 30 years of age and greater Total numbers observed Broasseae 0 12 1 3 16 Cocesea 0 12 2 10 24 Corypheae 0 1 0 0 1 Cysophileae 0 1 0 0 1 Phoeniceae 4 7 4 6 21 Sabaleae 0 40 2 8 50 Trachycarpeae 0 7 1 3 11

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33 Table 4 6 Incidence p ercentage of Ficus spp. growing as a hemi epip hytic parasite on palms growing at Montgomery Botanical Center, as influenced by the palm species genus, tribe and subfamilies gross structural characteristics No prominent crownshaft; marcescent leaf bases Number of individuals observed with hemi epiphyt ic parasites Percent incidence of palms within a species with marcescent lea f bases and hemi epiphytic p arasites Percent incidence of palms with hemi epiphytic parasites within a genus Percent incidence of palms with hemi epiphytic parasites within a tribe Percent incidence of palms with hemi epiphytic parasites within a subfamily 1) Thrinax radiata 1 0.64% Livistona 4.8% Cysophileae 0.64% C oryphoideae 10.1% 2) Livistona decora 1 1.0 % Attalea 9.0% Corypheae 4.3 % A recoideae 11.0% 3) Attalea cohune 1 1.6 % Sabal 10.9% Trachycarpeae 4.9 % 4) Caryota rumphiana 1 4.3 % Butia 16.6% Cocoseae 11.0 % 5) Sabal domingensis 4 5.9 % Hyphaene 22.5% Sabaleae 10.9 % 6) Hyphaene thebaica 1 6.2 % Phoenix 35 % Borasseae 22.5 % 7) Livistona chinensis 4 6.7 % Phoeniceae 35 % 8) Copernicia baileyana 1 7.1 % 9) Phoenix reclinata 1 7.1 % 10) Elaeis guineensis 3 9.6 % 11) Attalea butyracea 4 10 % 12) Butia eriospatha 1 10 % 13) Livistona saribus 5 10 % 14) Sabal palmetto 38 10.6 % 15) Sabal maritima 4 13.7 % 16) Syagrus coronata 5 16.6 % 17) Hyphaene coriacea 9 18.7 % 18) Butia capitata var odorata 3 21.4 % 19) Attalea phalerata 7 22.5 % 20) Phoenix canariensis 7 43 % 21) Hyphaene petersiana 20 85.7 % 22) Sabal mexicana 6 100 % 23) Sabal uresana 3 100 % 0 Table 47 Percentage of noncrownshaft palms with marcescent leaf bases and hemi epiphytes and the total percentage of palms within all three palm groups sampled with hemi e piphytes Percentage of non crownshaft palms with marcescent leaf bases and hemi epiphytes 10 % Total percentage of palms sampled with hemi epiphytes within all three palm groups 6.5%

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34 Table 4 8 Average age of non crownshaft, marcescent leaf based palm s at Montgomery Botanical Center which harbors Ficus hemi epiphytes The ages of non crownshaft marcescent leaf based palms ranged from 52 years old to 10 years old when using plant accession numbers as a gage of palm age. The actual accession numbers may n ot accurately depict the age of the palms, only when they were recorded as being planted on the garden property. A 00 depicts that the palms were planted when Montgomery Botanical Center was Colonel Montgomerys private residence, and are part of the original plant collection and the accession date of the planting is between19321959. Non crownshaft, marcescent leaf based palms Average age of palms (years) 1) Attalea butyracea 13 2) Attalea cohune 20 3) Attalea phalerata 28 4) Butia capitates var odorata 00 5) Butia eriospatha 19 6) Caryota rumphiana 15 7) Copernicia baileyana 18 8) E laeis guineenis 15 9) Hyphaene coriacea 20 10) Hyphaene petersiana 16 11) Hypaene thebaica 46 12) Livistona chinensis 22 13) Livistona decora 00 14) Livistona saribus 16 15) Phoenix canariensis 20 16) Phoenix reclinata 32 17) Sabal domingensis 15 18) Sabal maritima 18 19) Sabal mexicana 26 20) Sabal palmetto 23 21) Sabal uresana 38 22) Syagrus coronat a 13 23) Thrinax radiat a 20

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35 Figure 4 1 Self sheathing crownshaft palm, Royal palm ( Roystonea regia) Photo by Greg ory Kramer Photo taken at Montgomery Botanical Center.

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36 Figure 4 2 Self sheathing non crownshaft palm Triangle palm ( Dypsis decaryi ) Photo by Greg ory Kramer Photo taken at Montgomery Botanical Center.

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37 Figure 43. Non self sheathing, non crownshaft palm with marcescent leaf bases Sabal palm ( Sabal palmetto) with Ficus Photo by Greg ory Kramer Photo taken at Dade County P ark.

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38 Figure 44 Early stage infestation of Ficus hem i epiphytic on Sabal palmetto le af base. Photo by Greg ory Kramer Photo taken at Montgomery Botanical Center

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39 Figure 45 Advanced Ficus development on Sabal palmetto. Photo by Greg ory Kramer Photo taken at Montgomery Botanical Center.

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40 Figure 46 Ariel root initiation of Ficus growing on Sabal palmetto. Photo by Greg ory Kramer Photo taken at Dade County Park.

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41 Figure 47 Advanced hemi epiphytic parasitism of Ficus tree on Sabal p almetto. Photo by Greg ory Kram er Photo taken a Dade County Park.

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42 Figure 4 8 Free standing Ficus tree engulfing S abal palmetto. Photo by Greg ory Kramer Photo taken at Pinecrest Gardens.

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43 Figure 49 Phoenix canariensis with two different Ficus spp. hemi epiphytic parasites. Photo by Greg ory Kramer Photo taken at Dade County Park.

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44 CHAPTER 5 SUMMARY AND CONCLUSI ON P alm trees in the group with no crownshaft s and with marcescent leaf bases had the highest level of Ficus hemi epiphytes The phenological trai t of leaf base retention allows for the accumulation of detritus and the subsequent physical support needed for Ficus seed retention and germination. Palms with prominent crownshafts and cleanly abscis sioning leaf bases and the group of non crownshaft palms with cleanly abscis sioning leaf bases were not observed to have Ficus hemi epiphytes. Palm tree susceptibility to Ficus hemi epiphytic parasitism showed varying degrees of parasitism even with in the susceptible group. The two genera t hat seemed the most susceptible to hemi epiphytes were Sabal and Phoenix Both of these pal ms are commonly planted throughout the state of Florida but only support Ficus hemi epiphytes in the southern portion of the state where Ficus grow.

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45 LIST OF REFERENCES Addo, M.G.,Anning, A.K., Osei, F.O. and Patrick A. F. 2009. Composition and distribution of vascular epiphytes in a tropical semideciduous f orest, Ghana. The Authors J Comp. 47: 767773. Aguirre, A., Guevara, R., Garcia, M. and Lopez J.C 2010. Fate of epiphytes on phorophytes with different architectural characteristics along the perturbation gradient of Sabal mexicana forests in Veracruz, Mexico. J. Veg. Sci. 21: 6 15. Alden, P., Cech R. and Nelson, G. 1999. National Audubon Society Fiel d G uide to the Southeastern States, Alabama, Arkansas, Georgia, Kentucky, Louisiana, Mississippi, North Carolina, South Carolina, Tennessee Chanticleer Press, Inc., New York Alden, P., Cech, R., and Nelson, G. 1998. National Audubon Society : Field Gui de to Florida. ( Eighth edition) Chaticleer Press, Inc., New York Argo, V. N. 1 964. St r angler f ig ( Ficus aurea). Nat Hist 73(9) : 2629. Bancroft, G T and Bowman, R. 1994. Temporal p atterns in d iet of n esting w hite crowned p igeons: Implications for conservation of Frugivorous columbids The Auk 111(4) : 844852 Betrock's Plant Finder 2011. Betrock Information Systems, Inc. Hollywood, Florida. January 15, 2011 Brownsey, J P and Page, C.N 1986. Treef ern skirts: A defense against climbers and l ar ge e piphytes. J. Eco. 74: 787796. Craft, P. and Riffle, L. R. 2003. An Encyclopedia of Cultivated P alms Timber Press, Inc. Portland, Oregon. Croat, T. B. 1997. A revision of Philodendron subgenus Philodendron (Araceae) for Mexico and Central America. Ann Missouri Bot Garden 84: 311704. DeNiro, M.J Lord, E M ., Sternberg, L, S. and Ting, I. P. 1985. Crassulacean Acid Metabolsim in the Strangler Clusia rosea Jacq. Science 229( 4717 ):969971 Dirzo, R. and Lopez, J C 2007. Floristic diversity of S abal palmetto w oodland: A n endemic and endangered vegetation type from Mexico. Biodiversity and Conservation 16: 807825. Dransfield, J., Uhl, N.W., Asmussen, C.B., Baker, W.J., Harley M.M. and Lewis, C E. 2008. Genera Palmarum: The Evolution and Classification of Palms. Royal Botanic Gardens, Kew Richmond, Surrey, UK

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46 Eric N. and Swagel, A. V. 1997. Sub strate w ater p otential constraints o n g ermination o f the s trangler f ig (Moraceae). Amer J. Bot 84(5) : 716722. Frank, R.J., Knight R.J. and Nadel, H. 1992. Escapees and Accomplices: The Naturalization of Exotic Ficus and Their Associated Faunas in Florida. Florida Entom 75 (1) : 2938 Garth, R E 1964. The e cology of Spanis h Moss ( Tillandsia u sneoides ): Its g rowth and d istribution Ecology 45( 3 ) : 470481. Gerhard, V. B. 2003. The epiphyte vegetation of the palm Socratea exorrhizaC orrelations with tree size, tree age and bryophyte cover. J Trop Ecol. 19: 8190. Grubb, T. C Jr., Schuett, K.C. and Volman, S F 1997. Relative h ippocampal volume in r elation to f oodstoring b ehavior in four species of w oodpeckers. Brain, Behavior Evol. 49(2) : 110120. Harrison, R.D. 2006. Mortality and rec r uitment of hemi epi phytic figs in the canopy of a Borean rain forest. J Trop. Ecol. 20: 477480. Hayslette, S E 2006. Seedsize selection in m ourning d oves and e urasian c ollaredd oves. Wilson J Ornithology 118(1) : 6469 Hayslette, S.E. and Poling, T D 2006. Dietary o verlap and f oraging competition between m ourning d oves and e urasian collaredd oves. J. Wildlife Man 70(4) : 9981004. Hemmerly, M.N ., Gasperson, D. G., and McAlister A. A. 1987. Parasitic flowering plants. P roceeding of the 4th International Symposium on Parasitic Flowering Plants. Middle Tennessee St ate Universit y, Murfreesboro, TN Herre, E. A., Sandra P, and Tyree, M. T. 1995. Comparison of h ydraulic a rchitecture of w oody p lants of d iffering p hylogeny and g rowth f orm with special r eference to f reestanding a nd h emi e piphytic Ficus species from Panama. Blackwe ll Pub 129(1):125134. Holbrook, F. E. and Putz F.E. 1989. St r angler f ig r ooting h abits and n utrient r elations in the Llanos of Venezuela. Amer J Bot 84: 781788. Kato, M. and Tsutsumi C. 2006. Evolution of epiphytes in Davalliaceae and related ferns Bot J. Linnean Soc 151: 495510 Laman, T.G. 1996. The i mpact of seed harvesting a nts ( Pheidole sp. nov.) on Ficus e stablishment in the c anopy. Biotropica 28( 4 ):777781. Laman, T.G 1995. Ficus stupenda g ermination and seedling e stablishment in a Borean r ain f orest canopy Ecology 76(8) : 26172626.

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47 Male, T.D. and Roberts, G. E 2005. Host as sociations of the strangler fig, Ficus watkinsiana, in a subtropical Queensland rain forest. Aust Ecol. 30 : 229236. McPherson, J.R 1999. Studies in u rban e cology: Strangler f igs in u rban p arklands of Brisane, Q ueensland, Australia. Inst Aust Geographers 37(3) : 214 229. Montgomery, R. 1935. A Palmetum in South Florida. Proc. Fla State Hort. Soc 48: 172173. Mucunguzi, P. 2007. Diversity and distribution of hemi epiphytes and facultative herbaceous epiphytes in Kibale National Park, Uganda. African J Ecol. 45: 5764. Poindexter, D B and Thompson, R.L 2005. Host specifi city of A merican m istletoe ( Phoradendron leucrpum Viscaceae) in Garrard County, Kentucky. J Kentucky Acad Sci. 66( 1) : 4043. Putz, F.E and Susilo, A. 1994. Figs and f ire. Biotropica 26( 4 ):468469. Ricklefs, R.E. 2007. The E conomy of Nature ,Data Analysis Update. Tropical rain forest biome. W.H. Freem an and Company, New York, N.Y. Stevenson, G.B 1996) Palms of South Florida. University Press of Florida, Gainesville, F lorida Timothy, L. G. 1996. The i mpact of seed h arvesting a nts ( Pheidole sp. nov.) on Ficus e stablishment in the c anopy Biotropica 28(4b):777781. Todzia, C. 1986. Growth h abits, h ost t ree s pecies, and d ensity of h emiepiphytes on Barro Colorado Island, Panama. 18(1): Biotropica 18: 22 27. Tomlinson, P B 2001. Ficus aurea, Ficus citrifolia Laguncularia racemosa, Rhizophora mangle and Schinus terebinthifolius (2nd edition ) The Biology of Trees Native to Tropical Florida. Harvard Printing and Publication Services Allston, Massachusetts United States Department of Agriculture (USDA). 2011. Plant Profiles < http://plants.usda.gov/java/profile?symbol=PHLE14> 18 Mar. 2011 Wilson, E. O 1999. The Diversity of Life. W. W. Norton and Company, Inc. New York, N.Y.

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48 BIOGRAPHICAL SKETCH Gregory T. Kramer was born in New York City in 1968. He g raduated from the University of Miami with a Bachelor of Arts in b iology. He has 20 years of working experience in the horticultural profession, having worked in both the private and public sector. Early in his ca reer he held a horticulturist position with both the Wildlife Conservation Society and the City of New York Parks Department Before returning to school and Florida, he was the Director of Horticulture for the New York Restoration Project. He successful ly completed his M aster of S cience in environmental horticulture at the U niversity of F lorida (UF), in the summer of 2011, and is pursuing a d octorate degree in the Plant Medicine Program at UF in the fall of 2011.