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Who Dung It? The Impact of Seed Dispersal by Mammals on the fate of MILIUSA LINEATA (Annonaceae) Seeds in a Dry Evergree...

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Title: Who Dung It? The Impact of Seed Dispersal by Mammals on the fate of MILIUSA LINEATA (Annonaceae) Seeds in a Dry Evergreen Forest, Huai Kha Khaeng Wildlife Sanctuary, Thailand
Physical Description: Book
Language: English
Creator: Wheeler, Jessica H.
Publisher: New College of Florida
Place of Publication: Sarasota, Fla.
Creation Date: 2009
Publication Date: 2009

Subjects

Subjects / Keywords: Seed Dispersal
Southeast Asia
Germination
Genre: bibliography   ( marcgt )
theses   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
born-digital   ( sobekcm )
Electronic Thesis or Dissertation

Notes

Abstract: This thesis assesses the impact of seed dispersal by mammals on the fate of seeds from a common Annonaceae species in Huai Kha Khaeng Wildlife Sanctuary, Thailand. I experimentally evaluated three hypotheses: 1) that seeds experimentally placed within mammal dung have higher survivorship than control (bare) seeds; 2) that seeds in higher density clumps (25 as opposed to 5) suffer a higher mortality rate; and 3) that mortality rates decrease with distance from nearest adult conspecific. Seeds dispersed in high density piles (25 seeds) actually had a higher survival rate than seeds dispersed in low density (5 seed) piles. Additionally, seeds embedded in dung had the highest survival rate and distance from nearest adult conspecific had little effect on the survival of seeds through germination. Scolytinae beetles caused the most seed mortality, which differs from many other studies where rodents were the main seed predators. Understanding natural forest regeneration patterns is crucial for implementing successful forest restoration projects. More research over larger spatial and temporal scales is needed to elucidate the complicated processes underlying the fate of seeds in tropical forests.
Statement of Responsibility: by Jessica H. Wheeler
Thesis: Thesis (B.A.) -- New College of Florida, 2009
Electronic Access: RESTRICTED TO NCF STUDENTS, STAFF, FACULTY, AND ON-CAMPUS USE
Bibliography: Includes bibliographical references.
Source of Description: This bibliographic record is available under the Creative Commons CC0 public domain dedication. The New College of Florida, as creator of this bibliographic record, has waived all rights to it worldwide under copyright law, including all related and neighboring rights, to the extent allowed by law.
Local: Faculty Sponsor: Gilchrist, Sandra; Lowman, Margaret

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Source Institution: New College of Florida
Holding Location: New College of Florida
Rights Management: Applicable rights reserved.
Classification: local - S.T. 2009 W56
System ID: NCFE004198:00001

Permanent Link: http://ncf.sobek.ufl.edu/NCFE004198/00001

Material Information

Title: Who Dung It? The Impact of Seed Dispersal by Mammals on the fate of MILIUSA LINEATA (Annonaceae) Seeds in a Dry Evergreen Forest, Huai Kha Khaeng Wildlife Sanctuary, Thailand
Physical Description: Book
Language: English
Creator: Wheeler, Jessica H.
Publisher: New College of Florida
Place of Publication: Sarasota, Fla.
Creation Date: 2009
Publication Date: 2009

Subjects

Subjects / Keywords: Seed Dispersal
Southeast Asia
Germination
Genre: bibliography   ( marcgt )
theses   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
born-digital   ( sobekcm )
Electronic Thesis or Dissertation

Notes

Abstract: This thesis assesses the impact of seed dispersal by mammals on the fate of seeds from a common Annonaceae species in Huai Kha Khaeng Wildlife Sanctuary, Thailand. I experimentally evaluated three hypotheses: 1) that seeds experimentally placed within mammal dung have higher survivorship than control (bare) seeds; 2) that seeds in higher density clumps (25 as opposed to 5) suffer a higher mortality rate; and 3) that mortality rates decrease with distance from nearest adult conspecific. Seeds dispersed in high density piles (25 seeds) actually had a higher survival rate than seeds dispersed in low density (5 seed) piles. Additionally, seeds embedded in dung had the highest survival rate and distance from nearest adult conspecific had little effect on the survival of seeds through germination. Scolytinae beetles caused the most seed mortality, which differs from many other studies where rodents were the main seed predators. Understanding natural forest regeneration patterns is crucial for implementing successful forest restoration projects. More research over larger spatial and temporal scales is needed to elucidate the complicated processes underlying the fate of seeds in tropical forests.
Statement of Responsibility: by Jessica H. Wheeler
Thesis: Thesis (B.A.) -- New College of Florida, 2009
Electronic Access: RESTRICTED TO NCF STUDENTS, STAFF, FACULTY, AND ON-CAMPUS USE
Bibliography: Includes bibliographical references.
Source of Description: This bibliographic record is available under the Creative Commons CC0 public domain dedication. The New College of Florida, as creator of this bibliographic record, has waived all rights to it worldwide under copyright law, including all related and neighboring rights, to the extent allowed by law.
Local: Faculty Sponsor: Gilchrist, Sandra; Lowman, Margaret

Record Information

Source Institution: New College of Florida
Holding Location: New College of Florida
Rights Management: Applicable rights reserved.
Classification: local - S.T. 2009 W56
System ID: NCFE004198:00001


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Who Dung It? The impact of seed dispersal by mammals on the fate of Miliusa lineata (Annonaceae) seeds in a dry evergreen forest, Huai Kha Khaeng Wildlife Sanctuary, Thailand By Jessica H. Wheeler In collaboration with Trevor Caughlin and Thawut Wisate A Thesis Submitted to the Division of Natural Sciences New College of Florida in partial fulfillment of the requirements for the degree Bachelor of Arts Under the sponsorship of Dr. Sandra Gilchrist and Dr. Margaret Lowman Sarasota, Florida May, 2009

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iiAcknowledgements Thanks to my thesis committee for their advice and support. I especially want to thank my advisor, Meg, who has been incred ibly helpful and provided me with countless opportunities while Ive been at New College. Dr. Gilchrist, my cosponsor, was able to set me on track and helped me keep things in perspective and I thank her for that. I also want to express my admiration and respect for Dr. Clore, who has been an important role model for me. Jono Miller, Julie Morris a nd Susan Cerulean kept me feeling both imaginative and grounded during my time at New College and their support has been invaluable. Thanks to Dr. Cooper for enthus iastically helping me with organizing and analyzing my data. I also want to th ank my peer committee member and fellow ecology major, Chris Wilson, who kindly read my thesis in its draftiest forms and lent an ear to my rambling thoughts. Dr. Ellen Andresen deserves special thanks for first sparking my interest in the influence of dung on seed fate and later assisti ng me in developing my thesis project. P Thawut Wisate, my collaborator, made this thesis possible with his diligence and intelligence. Thanks for being a wonderful fr iend. Sheila Hershey, P Noi and P Wit also deserve gratitude for making me feel more at home in Thailand and lending a hand when I needed it most. I also wish to express my gratitude to those students and professors at King Mongkuts University of Technology, Thonburi who gave me support and assistance. ( Kop kun ma kaa, Thank you very much ). I want to thank my parents for enc ouraging my love of nature and being supportive of the many paths I have considered. Whether I become a scientist or just a crazy woods-woman, I know I can count on you to be supportive (and maybe even join me!) I especially want to thank my sister Emma, whose creative and imaginative nature has always been an in spiration to me. Trevor Caughlin has been a supportive friend, innovative collaborator and energetic dance partner. Thanks for believing in me and for knowing just when a spontaneous wilderness excursion was in orde r. Danielle Smith, Carli Cooper, Lauren Caprio, Rachel Renne, David Anderson and Je nna Ervin have been the best surrogate family anyone could ask for. Everythi ng about you is good and nothing is bad. I especially want to thank Bethany Highsmith, my best friend of eighteen years I cant imagine life without our collective memory. Thanks to ev eryone else who made the world a more beautiful place these last four years in particular Ashley Johnson, Sarah McAwesome and Amy Ortiz. Thanks to the New College Alumnae A ssociation, Natural Science Department and the Council on Academic Affairs for their financial support.

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iii Table of Contents I: Introduction .....................................................................................................................1 II: Literature Review ...........................................................................................................4 2.1 Closing the seed dispersal loop..........................................................................4 2.2 Does the presence of dung impact the post dispersal fate of seeds? ...............10 2.2.1 Seed predation ...............................................................................10 2.2.2 Secondary seed dispersal................................................................11 2.2.3 Germination ...................................................................................12 III: Methods ......................................................................................................................14 3.1 Study Site .....................................................................................................14 3.2 Focal Species ...............................................................................................16 3.3 Procedure .....................................................................................................17 IV: Results ................................................................................................................... .....22 4.1 Hypothesis I .................................................................................................23 4.2 Hypothesis II ...............................................................................................23 4.3 Hypothesis III................................................................................................23 V: Discussion ...................................................................................................................28 5.1 Seed removal and predation .........................................................................28 5.1.1 Scolytinae infestation .....................................................................28 5.1.2 Rodent removal .............................................................................29 5.1.3 Land crab predation .......................................................................31 5.1.4 Secondary seed dispersal by dung beetles.....................................32 5.2 Discussion of Hypotheses.............................................................................33 5.2.1 Hypothesis I....................................................................................33 5.2.2 Hypothesis II...................................................................................34 5.2.3 Hypothesis III.................................................................................35 5.3 Primary Seed Dispersal Agents ....................................................................35 5.5.1 Sun bear..........................................................................................36 5.5.2 Gibbons..........................................................................................38 5.5.3 Civets .............................................................................................38 5.5.4 Hogs...............................................................................................39 5.4 Forest Restoration .......................................................................................42 5.5 Conclusions..................................................................................................43 VI: Bibliography ...............................................................................................................45 Appendix I: Raw Data ......................................................................................................49

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iv List of Figures and Tables Figure 2.1 Seed dispersal loop..........................................................................................4 Figure 2.2 Germinated seeds in latrine at base of tree......................................................6 Table 2.1 Progress in seed fate ecology............................................................................7 Figure 2.3 Flow chart of potential seed fates ...................................................................9 Figure 3.1 Map of Thailand Western Forest Complex...................................................15 Figure 3.2 Map of Map of Khlong Pluu Research Station.............................................15 Figure 3.3 View of dry evergreen forest.........................................................................16 Figure 3.4 Miliusa lineata seeds.....................................................................................17 Figure 3.5 Depulping a nd marking seeds........................................................................18 Figure 3.6 Germination tests betw een control and marked seeds...................................18 Figure 3.7 Author setting transect and close up of prepared seeds and dung..................19 Figure 3.8 Examples of seeds at se cond check, beginning to germinate .......................20 Figure 3.9 Miliusa lineata seedlings at final check .......................................................21 Figure 4.1 Histogram of the percent of seeds surviving.................................................22 Figure 4.2 Percent of seeds germin ated and surviving at each check.............................25 Figure 4.3 Box plot of germinated and surviving seeds by final check..........................26 Figure 4.4 Box plot of seeds attacked by scolytinae beetles at first and last check and distance from adult conspecific....................................................27 Figure 5.1 Seed pile infested by sc olytinae and close up of endosperm of infected seed..............................................................................................29 Figure 5.2 Maxomys surifer visiting Miliusa lineata seeds.............................................30 Figure 5.3 Crab hole with germinated seed inside and land crab with eggs found near site.......................................................................................31 Figure 5.4 Roller dung beetles strugg ling with dung ball caught on seed marked with string.................................................................................33 Figure 5.5 Recruitment of seeds in sun bear scat............................................................37 Figure 5.6 Gibbon dung with Miliusa lineata and Mitrephora sp. seeds........................38 Figure 5.7 Civet defecating in latrine..............................................................................39 Figure 5.8Hog dung with Miliusa lineata and Mitrephora sp. seeds..............................40 Table 5.1 Composition and location of dung..................................................................41 Figure 5.9 Example of dung analysis seeds separated from dung and counted...........42

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v Who Dung It? The impact of seed dispersal by mammals on the fate of Miliusa lineata (Annonaceae) seeds in a dry evergreen forest, Thailand Jessica H. Wheeler In collaboration with Trevor Caughlin and Thawut Wisate New College of Florida, 2009 ABSTRACT This thesis assesses the impact of seed disp ersal by mammals on the fate of seeds from a common Annonaceae species in Huai Kha Khaeng Wildlife Sanctuary, Thailand. I experimentally evaluated three hypotheses: 1) that seeds experimentally placed within mammal dung have higher survivorship than cont rol (bare) seeds; 2) that seeds in higher density clumps (25 as opposed to 5) suffer a higher mortality rate; and 3) that mortality rates decrease with distance from nearest ad ult conspecific. Seeds dispersed in high density piles (25 seeds) actually had a higher survival rate than seeds dispersed in low density (5 seed) piles. Additionally, seed s embedded in dung had the highest survival rate and distance from nearest adult conspecific had little effect on the survival of seeds through germination. Scolytinae beetles cause d the most seed mortality, which differs from many other studies wher e rodents were the main seed predators. Understanding natural forest regeneration patterns is crucial for implementing successful forest restoration projects. More research over larger spatial and temporal scales is needed to elucidate the complicated processes underlying the fate of seeds in tropical forests. Dr. Sandra Gilchrist and Dr. Margaret Lowman Division of Natural Sciences _______________________________ ______________________________ Dr. Sandra Gilchrist Dr. Margaret Lowman

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1Though I do not believe that a plant will spring up where no seed has been, I have great faith in a seed. Convince me that you have a seed th ere, and I am prepared to expect wonders. Henry David Thoreau I: Introduction The structure of a tropical forest is the culmination of myriad ecological processes. One crucial process in structuring forest communities is seed dispersal. Seeds, the mature ovules of angiospe rms and the carriers of their genetic information, represent a plants most mobile phase. The seed contains all of the necessary components to begin a new life far from home an embryo and food reserves contained within a protective seed coat. It disperses through space and across time, and is capable of colonizing new habitat. Compar ed to adults, seeds ar e small, making them easily transportable while at the same time leaving them susceptible to hostile environments. Seeds disperse from place to place in space and time via wind, water, and animals. Seed dispersal is critical in coloni zation, succession, forest regeneration, population dynamics, and even landscape level va riations. Of the many gaps which exist in our knowledge of seed dispersal, the most prominent is linking the initial deposition of seeds with the demography of adult tree s (Wang and Smith, 2002). In fact, recent literature has called into question the idea th at initial seed deposit ion has anything to do with the final distribution of individuals and plant communities (e.g. McConkey, 2005). More information is required to understand th e fate of seeds once they reach the forest floor, i.e. the post-dispersal fate of seeds. The life history of the seed di spersal of every species is distinctly different. Some seeds may be deposited only feet from their pa rent while other seeds, carried far in the

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2 gut of a far ranging animal, may be dispersed miles from their origin, with a chance of colonizing new ecosystems (Nathan et al., 2003) Seed ecologists hope to determine a relatively consistent set of variables that improve a seeds chance of germinating and growing to reproductive maturity. When is a seed most vulnerable? How many seeds survive from stage to stage from an animals gut to the forest floor, from seed to germination, from germination to seedling? As a seedling, how long will it persist before light, or some other limiting factor, facilitates re cruitment? Seed mortality is intensive, as can be seen by the vast quantities of seeds produced by adult trees in comparison to the significantly less abundan t supply of seedlings. Is it true that on ly one seed needs to survive to adulthood? If this is the case, is the dispersal of seeds as important as ecologists suggest? Can the survival of one seed, out of potential millions, actually be traced to a distinct mechanism that determined its fate (Crawley, 2000)? Without understanding the role of seeds in forest regeneration, it is difficult to assess the health and status of mature forests. A forest can look relatively healthy for a time, even if it is not successf ully regenerating, because trees have such long life-spans. If we know about natural forest regeneration, we can then apply this knowledge to forest restoration projects. How can land manage rs maintain important ecological processes involved in regeneration in a naturally-indu ced manner? As Andresen (2002) suggested, it is sometimes necessary to determine whic h seed-dispersing speci es should become the focus of conservation efforts wh en the overall goal is to ma intain the regeneration ability of a forest patch (pg. 262). As degraded la nds begin to flourish, can seeds be protected from predation by common seed predators, such as rodents and beetles? How many seeds need to survive for a forest to regenerate?

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3 In this thesis, I focus on large-seede d, fleshy-fruited, tropica l tree species and the fates of their seeds once they reach the fore st floor after being defecated by a mammal. These seeds must survive passage through the mammals gut and subsequently endure attacks from fungi, insects, rodents, as well as extreme drought or flooding and variation in light. I explore the effect that dispersal in dung (i.e. the impact of being clumped, embedded in feces, and removed from below the canopy of the parent tree) on a seeds survival and germination. While I will only be looking at the survival patterns, this research invites further speculation about s eed ecology Does dung actually remain intact long enough to nurture the s eed with moisture and nutr ients? Does dung attract secondary seed dispersers, such as dung beetles, to secure the seeds for germination? And does dung attract potential pred ators and provide a substrate for fungal infection? I address three spec ific hypotheses: I) Seeds experimentally placed within mamma l dung will have higher survivorship than control (bare) seeds. II) Seeds in higher density clumps will suffer a higher mortality rate. III) Mortality rates will decrease with distance from nearest adult conspecific. The first hypothesis is based on resear ch on mammal dung and seed survival by McConkey (2005) and Andresen (1999, 2000, 2004). The other two hypotheses are based on the Janzen-Connell hypothesis of densit y-dependent mortality in which a seed is predicted to have a greater ch ance of being attacked by pred ators foraging in areas with high concentrations of seeds (Janzen, 1970; Co nnell, 1971). I will present data on the species of mammals dispersing the seeds of focal species and the composition of their dung in my discussion.

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4 II: Literature Review 2.1: Closing the seed dispersal loop Wang and Smith (2002) synthesized curre nt knowledge of seed dispersal and called on ecologists to close the seed dispersal loop (Fig. 2.1) This loop is frequently broken into simpler portions for ease of study. This thesis is limited to the portion of the seed dispersal loop between seed rain a nd seedling distribution in which secondary dispersal, predation, and germination take place. This is frequently referred to as seed fate ecology. Figure 2.1: The seed dispersal loop ( reproduced from Wang and Smith, 2002) A seeds fate begins long before it reach es the forest floor. Seeds are susceptible to predation from the beginning, even while still encased in pulp and hanging from a branch of the parent tree. Some seeds ar e attacked by predatory insects and animals curious to test the fruits ripeness. Other seed s fall to the ground from disturbances to the canopy. Seeds that retain their pulp after they reach the ground rarely germinate

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5 (Caughlin and Wisate, unpublished data). This is due in part to chemicals in some fruit that prevent seeds from germinating on th e parent tree (Kitajim a and Fenner, 2000). These seeds may still germinate if they happen to be secondarily dispersed by insects or animals foraging for fallen fruits (personal observation). As fruits ripen, a wide variety of anim als come to feed on them. In Thai evergreen forests an impressive 89% of trees rely on the abundant frugivorous animals to disperse their seeds (Elliot et al., 2003). Unlike many flow ers, which often have tight mutualisms with their pollinator, fruits and seeds are open to dispersal by different species of animals (Andresen, 2002). Some animals swallow fruit whole while others chew off the pulp, dropping the seeds to the fo rest floor. Birds will frequently regurgitate seeds after removing pulp in their gizzards, and other animals will chew both fruit and seeds. Swallowed seeds will be defecated either under or away from the canopy of the parent tree. The various seeds disperse d throughout the forest in cluding the seeds below the parent tree, seeds collected in animal latrines, seeds scatte red with and without dung are considered to be a trees primary seed sha dow. The effectiveness of dispersal is based on quality and quantity of dispersal serv ices. As defined by Schupp (1993), The quantity of dispersal depends on A) the number of visits made to the plant by a disperser and B) the number of seeds dispersed per visit. The quality of seed dispersal depends on A) the quality of treatment given a seed in the mouth and in the gut and B) the quality of seed deposition as de termined by the probability that a deposited seed will survive and become an adult (pg. 15). Some animals may disperse a large number of seeds (high quantity dispersal) but may disperse them in a relatively unfavorab le location for germination (low quality

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6 dispersal). An example of this from my study site are binturongs which disperse large numbers of seeds but did so in designated latrines at the bases of trees where few, if any, of these seeds might grow to adulthood (see germinated seeds in figure 2.2). The quality of dispersal also takes into account the handli ng behavior of the dispersing animal how its guts treated the seed, am ong other factors (Schupp, 1993). Figure 2.2: Germinated seeds in latrine at ba se of tree (photo courtesy of T.Wisate) Even seeds in latrines have a chance of survival if they are secondarily dispersed by dung beetles or rodents. Some of th ese seeds may be moved to favorable microhabitats away from the base of the tree. Clearly, the pr imary seed shadow is not the whole picture a seeds fate does not end here. During most of the 1960s and 70s, seed ecologists mainly studied frugivore seed handling behavior (with the majority of work done on birds and bats) and the primary dispersal of these seeds (Table 2.1). Janzen (1982) analyzed the removal of seeds by ants from horse dung. The study of post dispersal seed fate was not fully set into motion until

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7 the mid-1980s when ecologists began to look beyond the consumption and deposition of seeds by frugivores to examine the complex we b of interactions occu rring between biotic and abiotic factors on the forest floor. Table 2.1: Review of prominent papers in seed ecology Year Authors Location Result 1971 Dan Janzen Review Seed predation is important and is ideal for evolutionary and ecological studies. Before this review, herbivory and granivory were frequently grouped together for study. 1972 Don Wilson and Dan Janzen Puerto Rico Scolytid beetles can attack up to 100% of palm seeds. 1977 Henry Howe Costa Rica Examined bi rd feeding assemblages and fruit/seed handling behavior. 1979 Henry Howe Costa Rica Examined fruit removal by birds, determined vireo as important disperser based on percent crop removed. 1981 Theodore Fleming Costa Rica Observed fruit production and removal by bats. Distances moved by bats rarely exceeded 300m. 1982 Dan Janzen Costa Rica A small number of seeds in many smaller piles of dung are more likely to survive than many seeds in large piles of dung. 1982 Henry Howe and Judith Smallwood Review What is the ecological advantage of seed dispersal? How do fruit characteristics impact frugivore fruit choice? 1990 Eugene W. Shupp BCI, Panama More seed predation of specific seed when there is less overall fruit production in the forest. Overall high seedling recruitment rate. 1991 Alejandro Estrada and Rosamond Coates-Estrada Los Tuxtlas, MX 90% of seeds dispersed by howler monkeys are destroyed by rodents. By relocating seeds to favorable locations, dung beetles play an important role in forest regeneration. 1993 Pierre-Michel Forget BCI, Panama During periods of high fruit production, rodents cache more seeds. There was a low rate of survival in scatter-hoarded seeds. 1993 Doug Levey and Margaret Byrne Costa Rica Ants harvested seeds from frugivore feces and cache them in nests. The majority of seeds are killed, but some seeds are significantly benefited. 1996 Colin and Lauren Chapman Uganda Examined relationship between seedling recruitment and distance from parent tree. 30% more dispersed seeds survived than seeds under adults. 1998 Virginia Sheperd and Colin Chapman Uganda Through burying seeds, dung beetles significantly protect seeds from predation

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8 Today ecologists are beginning to connect the primary seed shadow with the establishment of seedlings. It appears that the primary seed shadow has little relevance to the final distribution of seeds. This is not to say that the initial seed shadow does not impact the fate of seeds both spatially and te mporally, but rather to say that it my not necessarily reflect the distribution of s eedlings, much less trees (McConkey, 2005). In addition to the distribution of seeds, the state of these seeds has a great impact on their fate whether a seed is defecated, spit, or falls from the canopy with pulp intact, will alter the series of events that will lead, ev entually, to death, germination, or dormancy. Seed dormancy is relatively uncommon in tropi cal tree species, and very little is known about it when it does exist in the tropics (Vasquez-Yanes and Orozco-Segovia, 1996). Therefore, a seed generally has one of two fates, germination or death (Fig. 2.3). The general question is this: what spatial and temporal, biotic and abiotic factors lead to favorable conditions for seed germination? When a seed is moved from its original locat ion, it either surviv es, or does not. In addition, if it is moved after primary dispersal, it will either be moved to a more favorable microhabitat, less favorable microhabitat, or a microhabitat of similar quality. If the seed survives, it is considered secondarily disper sed. Frequently, secondary dispersal can be helpful, such as when seeds are buried by dung beetles (see sect ion on secondary seed dispersal, below). Secondary dispersal can also mean that the seed was hoarded by a rodent, an encounter that it may or may not survive (Vander Wall et al., 2005). Some seeds are even adapted for secondary dispersal, using external lipid-packets to attract ants (Kaufmann et al., 1991).

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9 Figure 2.3: Flow chart of potent ial seed fates (Andresen, 2005) At the other end of the spectrum, seeds can be either attacked or preyed upon after dispersal. A predated seed is dead. An attack ed seed might have a fungal infection that causes the seed leaves to be damaged, but st ill survive. Alternatively, an animal might

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10 remove some of the seed reserves, but not damage it enough to prevent it from germinating (Andresen, 2002). Seed predation is an umbrella term for many fates, as many different organisms can destroy seeds. They can be preyed upon by fungi, boring beetles, rodents and other mammals. Three ecologists have intensively studied the impact of survival in dung and its impact on seed survival: 1) Kim McConkey, wh o works in Borneo, studies the impact of gibbon dung on seed predation; 2) Ellen Andresen who focu ses on secondary dispersal by dung beetles in the Neotropics; 3) and Anna Traveset who studies the impact of dung composition on germination success, mainly in Alaska. 2.2 Does the presence of feces alter the fate of a seed? 2.2.1 Seed Predation McConkey (2005) published two studies on seed removal one short-term and one long-term. The short-term study had two main goals to determine if feces improved the ultimate location of seeds and to identify the animals that interacted with dispersed seeds. She had three treatments seeds encased in dung, bare seeds, and dung without seeds. The number of seeds in each pile mimicked the actual composition of gibbon droppings, and the droppings were less than a day old. To examine the seed predator species she covered the area surrounding each replicate with sand and identified the track s of animals visiting the dung piles. She checked the piles at 1, 2, and 4 hours and then once a day for a week. She found that most predation by rodents occurre d within the first week after dispersal. Tropical seeds

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11 do not typically have a period of dormancy, so seeds that are not preyed upon usually germinate. In her long term study (11 months), she analyzed the influence of the primary seed shadow on the fate of seeds. McC onkey (2005) strongly advocates using natural primary seed shadows instead of experimental placements. One reason for this is that gibbons (and many other mammals) use simila r foraging paths on a regular basis. Because of this, they provide a constant s upply of food for foragi ng predators which can recognize the paths of the gibbons. If experiments haphazardly place dung pats and seeds, there may be very different predation levels. The majority of seed mortality was cau sed by vertebrates (86%), while only 2% were destroyed by insects. Almost all of the piles (98%) were vi sited by rodents, and 75% of this occurred within the first week. Conversely, most insect predation occurred relatively late, one or two months after the seeds were placed. McConkey also noted that when many tree s were fruiting, the predation levels of seeds in dung were significantly lower than when fewer trees were fruiting. In other words, when more appetizing foods were pr esent, those seeds in dung were safer. This has implications for restoration projects, which could use the fruiting of nearby species to offset the predation levels on broadcast seeds used in restoration. 2.2.2 Secondary Seed Dispersal Andresen (1999) st udied the fates of seeds disper sed in howler and spider monkey dung in the Peruvian Amazon. In the course of allocating dung for breeding and feeding, it is not uncommon for dung beetles to relocate and sometimes bury seeds. In her study

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12 site, dung beetles buried an average of 41% of seeds, but rodent removal ranged from 63 to 97%. Rodents were sometimes able to find seeds that were buried by dung beetles, but not if the dung beetles burie d the seeds deeply. Rapid removal of seeds and dung by dung beetles ultimately reduced the level of rode nt attraction. These buried seeds have a per capita increased survival rate of 80% a nd seeds were only buried if they were in dung. If, as this study implies, dispersal in dung leads to much higher recruitment rates, over-hunting of important dispersers could shift tr ee recruitment. Andresen and Levey (2004) examined how the presence and amount of dung affected the long-term fate of seeds a nd seedlings, how the burial depth affected seedlings, and how seed size a ffected the interac tion between seed size, dung beetles, and rodents. In this study, nearly one-half of dung-encased seeds were secondarily dispersed by dung beetles. Andresen (1999) concluded th at deep burial was beneficial, because it protected the seeds from preda tion by rodents. In this la ter study, Andresen and Levey (2004) observed that the deeper a seed was buried, the higher its rate of mortality; presumably the seed was unable to germinate, or successfully push through the soil. The optimal depth for burial was around 3 cm. Though seeds dispersed in mammal dung and seeds deposited without mammal dung underwent very different fates, each treatment ultimately produced the same proportion of seedlings. 2.2.3 Germination Traveset and colleagues (2004) examined the impact of manure composition of brown bear dung on germination and survival. This was one of the first germination studies where dung composition was tested experimentally. One species of Rubus

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13 examined actually had a higher germination rate in bear dung th an in dung composed mainly of fiber, fruit pulp or potting soil. Though they germinated more quickly in the dung composed mainly of animal material, th e final germination success of seeds in the different dung compositions were the same. Other seeds tested from a species of Vaccinium, were not as affected by manure composition but, like the Rubus seeds, the Vaccinum seeds grew more quickly and produced mo re leaves in the animal material dung. An omnivorous animal might have a positive impact on the seeds it disperses. Germination is sometimes viewed as a mode of escape from predation (Andresen, 2002), whereby quickly germinating species are able to outrun foraging pred ators. If this is the case, omnivorous animals, while perhaps pr oviding a lower quantity of dispersal than highly frugivorous birds or primates, may pr ovide a very high quality of dispersal, provided they disperse the seed s away from the parent tree.

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14 III: Methods 3.1 Study Site This study was conducted in the Huai Kha Khaeng Wildlife Sanctuary (HKK) in West-Central Thailand (Fig. 3.1). HKK is part of the Tha iland Western Forest Complex, the largest continuous preserved area in mainland SE Asia, covering 1,870,000 hectares of land along the Thai-Burmese border. Data were collected in th e area surrounding the Khlong Phuu research station (Fig. 3.2). This site was chosen because it has a rich and intact mammal population. In addition, it is a prime example of a naturally functioning dry evergreen forest, an endange red forest type at which Thai restoration efforts are being aimed. The sites altitude ranges from appr oximately 550-650 m and is composed of gently sloping terrain with ephemeral stre ams in the wet season. The surface soil is classified as sandy-loam. While HKK is com posed of many forest types, we limited our study to an area of seasonal dry evergreen forest. The annual rainfall there is 1,476 mm/year and has a distinct dry season from No vember to April. This forest, which has canopy ranging from 40-55 m in height, is dominated by emergent Dipterocarpaceae and various Annonaceae, mesic species and figs (Fig 3.3). The species richness in this area is relatively low, but contains a relatively high number of rare species (Bunyavejchewin et al., 2004). The understory is relatively open, except near stre am beds where it is dominated by Zingiberaceae (personal observation ). The animal community is very rich, with 159 species of mammals, 379 species of bi rds, and 70 species of reptiles including viable populations of elephants, tigers and leopards. Several troops of gibbons range in the area (appr. 5.4 ind/km2; Bunyavejchewin et al., 2004).

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15 Figure 3.1: Thailand Western Forest Complex, circled in red (from UNESCO website, http://whc.unesco.org) Figure 3.2: Map of Khlong Pluu Research St ation in relation to HKK headquarters (Map created using Google Earth)

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16 Figure 3.3: View of typical area of dr y evergreen forest near Khlong Phuu Focal Species Miliusa lineata (Craib) Ast (Annonaceae) is a re latively abundant tree, with approximately 33 trees >1 cm-dbh/ha (Bunyavejc hewin et al., 2004). The fruits vary in size, containing 1-5 seeds (average of 3.8 seed s/fruit). The seeds averaged at 11.2 x 7.4 x 5.8 mm. The fruits were 23.1x19.4x17.0 mm in size on average (Fig. 3.4).

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17 a. b. Figure 3.4: a) Miliusa lineata fruit on tree. b) Variation in M. lineata fruit size and seed content 3.3 Procedure Seeds (n= 1,750) were collected from Miliusa lineata both directly from the tree when possible or fallen below the tree. All fruits were checked for insect entrance holes before being collected. Seeds were manually de pulped and then float-tested for viability. Floating seeds were discarded. Each seed was then dried overnight and marked. Marking consisted of making a red line with a permanent marker on both sides of the seed. Then small strips of blue flagging ta pe were attached to the seed coat of each, carefully avoiding the suture, using Ethyl-2-Cyanoacrylic super glue (Fig. 3.5). Germination success between control seeds and ma rked seeds was tested. Thirty seeds of control and marked seeds were planted approximately two inches deep in small plastic bags filled with sandy soil from the stream bank (Fig. 3.6). No significant difference in germination success was observed (60% of control vs. 71% of marked seeds germinated.)

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18 a. b. Figure 3.5: Collaborators T. Caughlin and T. Wisate (a) depulping and (b) marking seeds a. b. Figure 3.6: Germination tests be tween control (a) and marked (b). Sac stands for Saccopetulum Miliusa lineatas previous scientific name. Dung was collected on a daily basis and the composition of dung varied from transect to transect. All dung had some quan tity of primate dung as well as one or more dung pats from sun bear, hog and/or civet. Th is was a confounding aspect of our research

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19 and ideally only one species dung should be used. Howeve r, it was difficult to find enough dung from any given species to create e nough replicates. We hope that the seeds and predators reacted similarly to the mixe d dung as they would have the single-species dung. Andresen (personal communication, 200 8) said that mixing dung or using dung from human latrines or pig farmers was acceptable, though not ideal. We randomly placed 25, 30 m transects thr oughout the forest. Each transect was composed of four replicates (1) five seeds with no dung, (2) five seeds in dung, (3) 25 seeds with no dung, and (4) 25 seeds in dung (Fig. 3.7). Each replicate was placed in a randomly chosen order along the transect. Site s were ten meters apart, which has been shown to allow for independence (Andres en, personal communication, 2008). While ideally we would have mimicked a natural primary seed shadow (as in McConkey, 2005) we were not able to do so because of time constraints. Gibbons and civets did not appear to defecate along the distinct animal trails in the forest, and therefore their movements and defecations would have to have been mapped over a longer time period. a. b. Figure 3.7: a) Author setting transect and b) close up of prepared seeds and dung

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20 Transects were set in mid-July and every ten days for a total of three checks. During each check, the seeds were examined for rodent, fungus or insect damage, removal, germination, or seedling establishment. Germinated (Fig. 3.8) and established seedlings (Fig. 3.9) were checked for dama ge to the cotyledons, leaves and roots. Presence of ant piles, crab burrows, and scattering of seeds was also noted. a. b. c. d. Figure 3.8: Examples of seeds at second ch eck, beginning to germinate. A) 25 seeds in dung, b) 5 seeds not in dung, c) 5 seeds in dung and d) 25 seeds not in dung.

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21 a. b. Figure 3.9: a) Miliusa lineata seedlings at final check and b) flagged seedlings

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22 IV: Results Because my data were not normally distributed (Fig. 4.1), I chose to use the nonparametric Kruskal-Wallis one-way analysis of variance. Though it would be possible to transform my data into a normal distribution and use a parametric statistical analysis, I believe I would have lost important informa tion by doing so. A large percentage of seed piles did not survive, which skewed my data to have the distribution seen below. The Kruskal-Wallis Test is in many ways simila r to ANOVA, but the dependent variable is ranked (in this case germination success), with the smallest value getting a rank of one, the next highest value getting a rank of two, and so on. Ultimately, it compares the median ranks of two sets of data. For this reason, box plots were used to display data, as they show the median and range. Data we re analyzed using SAS and graphs created using R2.4.1. Figure 4.1: Histogram of the per cent of surviving seedlings.

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23 4.1: Hypothesis I: Experimentally pla ced within mammal dung will have higher survivorship than control (bare) seeds The presence of dung significantly impacted the cumulative survival ra tes (p=0.06), with 41% of seeds surviving in dung piles, and onl y 29% of bare seeds surviving. Much of this difference in survival can be attributed to the success of germinated seeds. Bare seeds germinated at a higher rate than those in dung at first, but late r those seedlings died (Fig. 4.2). In the end, a higher percentage of seeds embedded in dung survived (Fig. 4.3). Bare seeds and seeds embedded in dung had simila r levels of scolytinae beetle infestation (dung seeds = 63%, bare seeds = 73%). 4.2: Hypothesis II: Seeds in higher density clumps (25 as opposed to 5) will suffer a higher mortality rate Seeds dispersed in large piles actually had a significantly higher percentage of seeds survive (47%) as opposed to the small piles, in which only 22% of seeds survived (p=0.0001). Small piles of seeds suffered a higher mortality rate due to scolytinae infestation (85% for small piles oppo sed to 51% in large piles). 4.3: Hypothesis III: Mortality rates decrease with distance from near est adult conspecific The design of this experiment was not created to account for this hypothesis, so seed piles were not placed at regular inte rvals from parent trees. Seed s were, coincidentally, placed from 0-27 m from adult Miliusa lineata trees. There was no consistent increase or decrease in seed survival as the distance from adult conspe cific increased (p=0.53; Fig.

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24 4.4b). However, seeds within 3 m of an adu lt conspecific were at tacked more quickly than distant seeds (Fig. 4.4). Rodents were not a significant contribu tor to overall seed mortality (removing only 2% of seeds), nor did rode nt preferences appear to be correlated to the presence of dung, or the number of seeds per replicate. Over all, the most biologically relevant result was the percentage of seeds germinated and surviving.

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25 Percent of seeds germinated and surviving0% 5% 10% 15% 20% 25% 30% 35% 40% 45% 50% 55% 60% 65% 70% 75% 80% 85% 90% 95% 100% startcheck 1check 2check 3percent germinated 5 seeds in dung 5 seeds, no dung 25 seeds in dung 25 seeds, no dung Figure 4.2: Average percent of seeds germin ated and surviving at each check. Note that the seeds embedded in dung germinated steadily and had a higher survival rate than bare seeds, which germinat ed quickly, but later died.

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26 a. b. Figure 4.3: Box plot of germinated and surv iving seeds by final check. From top to bottom the lines represent the sample maximum, upper quartile, median, lo wer quartile and sample minimum.

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27 a. b. Figure 4.4: Box plot of seeds attacked by scolytinae beetles on first check (a) by day 36 (b) and distance from adult conspecif ic (1 = 0-3m, 2 = 3.1-6m, 3 = 6.1-9m etc)

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28 V: Discussion 5.1: Seed removal and predation Overall, regardless of treatment, the major ity of seedling mortality as attributed to scolytinae infestation. Very few seeds were removed or atta cked by other organisms. Based on 24 hour camera trapping conducted by Caughlin and Wisate (unpublished data) the most likely seed removers were rodent s, which took seeds at night. Based on two, real-time dung observations I c onducted (see section 5.1.4) it is also likely that some seeds were secondarily dispersed by dung beetles though the impact of this was most likely insignificant. 5.1.1: Scolytinae infestation The beetles observed attacking seeds were tentatively identified as Scolytinae, a subfamily of the weevil family, Curculionidae. Commonly called bark beetles, scolytinae beetles are well-known pine and elm pests in America. The beetles likely have low dispersal capability, but thei r populations can expand rapidl y once they find a resource, such as a seed pile (Fig. 5.1). Female scolytinae are haplodiploid and can actually produce young without mating (Jordal et al., 2002). Caughlin and Wisate (unpublished data) observed that they can produce multiple generations within a single seed. Though they can disperse only short distances, it is probably not particularly relevant in HKK with regard to Miliusa lineata seeds. M. lineata is a very common forest tree and, even with randomly arranged transects, seed p iles averaged only 9 m from an adult.

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29 Figure 5.1: Seed pile infested by scolytinae and close up of endosperm of infected seed. 5.1.2: Rodent removal I have yet to find a rodent removal rate as low as the one encountered in this study. As reviewed in Notman and Ville gas (2005), seed predation by mammals is considerably higher than predation by insects in most tropical forests. Assuming this removal was caused, for the most part, by rodents (as implied by Caughin and Wisates unpublished camera trapping data) this result has interesting implications (Fig. 5.2). The most abundant rodent species in the evergreen forests of HKK is Maxomys surifer

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30 (Miller, 1900) which composes 90% of the small mammal community (Walker and Rabinowitz, 1992). M. surifer is not highly granivorous, with a large portion of its diet being composed of insects (Walker and Rabinowitz, 1992). Additionally, dry forests, such as this evergreen forest, have a substa ntially lower biomass of small mammals than aseasonal forests, as the ecosystems carrying capacity is limited during the dry season. Though perhaps not directly comparable, a similar aseasonal Malaysian forest had a small mammal biomass of 1,660 g/ha while the dry evergreen forest of Huai Kha Khaeng had a mere 616 g/ha (Nor, 2008; Walker and Rabinowitz, 1992). HKK is the most undisturbed area in mainland SE Asia, and hosts a near inta ct population of animals. Rodents are known to increase when there are few top predators to control their populations often near disturbe d forest edges or secondary forests. In this intact ecosystem, it is likely that their populations remain low. Figure 5.2: Maxomys surifer visiting M. lineata seeds (photo courtesy of Caughlin and Wisate)

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31 5.1.3: Land Crabs The land crabs in Huai Kha Khaeng have yet to be studied, and I was unable to identify them (Fig. 5.3b), though they may be Isolapotamidae, genus Demanietta. They are known to be an important food source for hornbills. Their impact on seedling recruitment in this fo rest is unknown. Land crabs actually create burrows near seed piles and sequester seeds within them (Fig. 5.3a). In some ecosystems, crabs are known to be important seed and seedling predators. Sherman (2002) experimentally excluded crabs for two years and found that the density of seedlings increased 144%. He also observed that naturally crab free areas have twice as many plant species present as areas with high crab densities. His study suggests that crabs can drastically al ter plant community composition. a. b. Figure 5.3: Crab hole with germinated s eed inside and land crab with eggs found near site (photos courtesy of Thawut Wisate)

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32 5.1.4: Secondary seed dispersal by dung beetles I conducted two dung observations ove r the course of field work. I experimentally placed three human dung pats every ten meters and checked the pats three times over the course of the day once in the early morning, once mid-morning and again in the early afternoon. The roller dung beetles observed were relatively large (about 2 cm in length) and carved distinct balls out of the edge of th e dung pats (Fig. 5.4). The hole where they buried their allocated dung was about 3.5 cm away from the original pat. When dung pats were lifted, about 10 tunnels made by small (<0.5 cm) dung beetles were observed. There were also many dwellers that made tunnels directly into the top of the dung pats. Occasionally, dung pats were made into ne sts by ants. These pats were always avoided by other insects and remained intact. This allowed ants to utilize the dung, and likely protected the seeds from predation. On ce I observed a small beetle approaching an ant infested dung pile, the ants attacked it, and it fl ed. Many flies were observed landing on the dung pats. Spiders would sometimes take cover on the pats and attempt to attack visiting flies and other soft-bodied insects. Once, a frog was seen attacking one of these spiders. I marked the seeds placed in the dung pa ts with small pieces of neon thread. Several times, dung beetles attempted to move these seeds, but the thread became tangled (Fig. 5.4). However, this observation leads me to believe that at least some seeds are being secondarily dispersed by dung beetles at my study site. Some dung piles disappeared entirely perhaps being eaten whole by feral hogs.

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33 Figure 5.4: Roller dung beetles struggling wi th dung ball caught on seed marked with string 5.2: Discussion of Hypotheses 5.2.1: Hypothesis I: Experime ntally placed within ma mmal dung will have higher survivorship than control (bare) seeds The overall number of seeds to germinate in piles with and without dung did not differ. However, bare seeds germinated quickly and then died, so that the final number of surviving seeds was relatively low. Seeds in dung germinated later, but almost all of the seeds that germinated survived. Scolytinae be etle predation was similar for both bare and dung-encased seeds. Traveset (1998) f ound that seeds embedded in dung composed mainly of fruit pulp had a slower germina tion rate, possibly because of germination inhibitors present in the pulp. I speculate that this similarly slowed the germination rate

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34 in my study but, once germinated, seedlings had sufficient nutrients to continue growing successfully. 5.2.2: Hypothesis II: Seeds in higher density clumps (25 as opposed to 5) will suffer a higher mortality rate Seeds actually had a significantly reduced survival rate when dispersed in small piles. In approximately half of the of the small seed piles (n=24), seed mortality reached 100%. Conversely, there were few large seed piles (n=3) in which no seeds survived. The gap in survival rates may decrease with time as seedlings compete for resources and suffer from density dependent mortality. Howeve r, it appears more probable that at least one seed from a large pile will survive than one seed from a small seed pile. This is distinctly different from Pizo and Samaos (2001) research in which they found that animals that deposit seeds in faecal clumps ar e less efficient seed dispersers than those that regurgitate seeds individually (pg. 229) Additionally, Russo and Auspurger (2004) concluded that seeds dispersed in piles of five had a 2.8 times worse odd of survival than seeds dispersed singly. Conversely, Andres en (2002) observed that seed predators, specifically rodents, did not re spond to the density of disperse d seeds. In my study, seeds in both treatments suffered high mortality rate s from scolytinae beetles, but it was much higher for seeds in small pile s. These results, combined with the impact of dung on germinating seeds, seems to imply that seed s are more limited by nutri ents (or some other benefit dung provides) than by density-depende nt mortality. As can be seen in my results, piles of five seeds are very likely to be entirely removed or destroyed, whereas

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35 seeds piles of 25 seeds, though survival might be low, are likely to ha ve at least one seed survive per pile. 5.2.3: Hypothesis III: Mortality rates decr ease with distance from nearest adult conspecific It is interesting that no correlation wa s observed between distance from nearest adult Miliusa lineata and seed survival or attack. The only variation I observed was that (during the first check) scolytinae infesta tion was highest near the canopies of adult M. lineata. Previous studies, most recently review ed in Hammond and Brown (1996) found that insect attack varies with distance to parent tree. In Hammonds 1999 paper, however, he found that the total number of seeds attack ed by scolytinae beetles or rodents did not differ with distance to adult conspeci fic. Allmen and colle agues (2004) similarly found that, with palm seeds under very high density, predation is much more closely related to the time a seed is dispersed than where it is dispersed. Notman and Villegas (2005) found that insect predation was not distance-dependent, but that mammal predation was highest under adult trees. 5.3: Primary seed dispersal agents Birds and bats are typically given the most credit for important seed dispersal services. They often disperse seeds singly or in small clumps and these seeds have little dung attached. Seeds are also frequently regu rgitated. While I do think these animals are important for long distance dispersal services data in this study support the notion that animals which disperse large numbers of seeds in a substantial amount if dung may be as important.

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36 When it comes to conservation practices a nd restoration, it can be difficult to incorporate the services of these animals. Organisms such as gibbons and sun bears are not present in disturbed habitats, and therefor e their dispersal servic es cannot be used for restoration. Below, I review th e potential of each of the primar y dispersers at my site. 5.3.1: Sun bear Sun bears ( Helarctos malayanus (Raffles, 1821)) are the smallest living bears, 1.2 m in length and weighing 25-65 kg. Sun bear populations are in decline do to the use of their gall bladders in traditional Chinese medi cine. Their diet consists of honey, termites, small animals and fruits. Because sun be ars are small and do not require the same amount of protein as their larg er relatives, they can most lik ely subsist for longer periods of time on nothing but fruit. Sun bears will create nests in the canopy of a fruiting tree, and remain there for the duration of the trees fruiting period. For this reason, the quality of their dispersal se rvices is disputable (McConkey, 1998). Two sun bears scats were collected at our site, one containing 40 and the other containing 140 seeds. There was a large qu antity of dung surrounding the seeds (Table 5.1).

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37 Figure 5.5: Recruitment of seeds in sun bear scat. Temperate bears have been recognized as important frugivores but the ecology of sun bears is less well understood. Prior to McConkeys (1998) pa per on sun bear ecology the only record of frugivory in sun bears was the consumption of durian fruit. At our study site, we confirmed the role of sun bears as seed dispersers. We found a pile of sun bear scat near a fruiti ng Syzygium; upon returning one month later, we saw that many of the seeds had germinat ed. Paying closer note, we saw the claw marks of sun bears on the Syzygium and realized that this tree was very common in this area, though we had not encountered it el sewhere in the forest (Figure 5.5).

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38 5.5.2: Gibbons Gibbons ( Halobates lar (Linnaeus, 1771)) are considered hi ghly frugivorous. They consume relatively small, unprotected fr uits that do not require much hand/incisor preparation. One attribute that makes gibbons important dispersal agents is that they typically swallow and disperse seeds whole. Gibbon feces usually scatters as it falls through the canopy. While gibbons are important dispersers, they are rare in disturbed habitats (Corlett, 1992). The gibbon dung found at our site was orange and fibrous (Fig. 5.8) likely due to the orange pulp of both Mitrephora sp. and M. lineata fruit. The pats contained from 6-27 seeds apiece (Table 5.1). Figure 5.6: Gibbon dung with Miliusa lineata and Mitrephora sp. seeds 5.5.3 Civets Palm civets (subfamily Paradoxurinae ) and binturongs ( Arctictis binturong (Raffles, 1881)) are the most consistently frugivorous civet species. These species can climb trees and swallow seeds whole, thoug h they are discriminating in their fruit selection. Annonaceae are commonly consumed by civets. One benefit to seed dispersal

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39 by civets is that they trav el long distances. Many spec ies, however, repeatedly use latrines. In out study site, bi nturongs and some civets used la trines (Fig. 5.7). They also frequently defecated on logs, which is likely not highly beneficial for dispersed seeds. Civets are relatively tolerant of human dist urbance (Corlett, 1998). AT our site, civet dung had little fecal matter surrounding each seed. In the dun, we occasionally found bone and insect fragments. The scats consisted of anywhere from 8-171 seeds. Figure 5.7: Civet defecating in latrine (Caughlin and Wisate, unpublished study) 5.5.4: Hogs Eurasian wild pigs, Sus scrofa (Linnaeus, 1758), could be very important, high quality and quantity dispersers. Based on dung content and the germination of removed seeds, it appears that wild pigs provid e good dispersal services. In HKK, each dung pat contained from 41-208 seeds (Table 5.1). S. scrofa can endure a variety of conditions and has a highly adaptable diet though they are known to have some destructive habits.

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40 Matias and colleagues (2008) evaluated th e role of feral hogs and other omnivores as seed dispersers in a Mediterranean climate. They found that, for some plant species, hogs only dispersed viable seeds. Hogs did have higher numbers of cracked seeds than the foxes and martens, but it did not make th em significantly less adept dispersal agents. However, hogs could have negative impacts on seedling recruitment du e to their rooting habits. Little is known about the role of hogs as seed dispersal agen ts. We frequently found hog dung and it always contained some nu mber of viable seeds (Fig. 5.8). The dispersal of seeds by hogs has several implicat ions. First, hogs ar e retrieving their seeds from fallen fruits, which is a form of secondary seed dispersal. In addition, hogs are known to eat the dung of other animals, and the seeds found in their dung could also potentially be coming from that source. As stated in my discussion, hogs could be important as seed dispersal agents in highly disturbed habitats or, perhaps equally likely, they could damage seedlings through their extensive rooting behavior. Figure 5.8: Hog dung with Miliusa lineata and Mitrephora sp. seeds

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41 Table 5.1: Composition and location of dung. M/M stands for Miliusa lineata/Mitrephora sp. which we were unable to identi fy in the dung due to their similar seed shape. Unk. stands for unknown seed species. Gibbon Dung description: orange fibrous fecal material, slightly scattered Seed content Location 17 m/m ground 19 m/m road 6 unk. ground 13 longan trail 14 m/m ground 7 unk. road 27 m/m ground 27 m/m ground 2 m/m road Civet Dung Description: Little fecal matter. Usually in a line. Some bone fragments and insect parts present. Sometimes substantial amounts of pulp left on seeds. Seed content Location 52 m/m in mud 70 m/m log 23 m/m log 42 m/m log 55 m/m log 9 unk. log 8 m/m log 26 m/m log 33 m/m log 68 m/m latrine 171 m/m latrine 29 m/m ditch Sun bear Dung description: Thick, very strong smelling, compact cylinders of dung Seed content Location 40 m/m ground 143 m/m grass near field station Feral Hog Dung description: Dark brown fecal material. Varying seed:dung ratio. Seed content Location 99 m/m; 10 unk. ground 98 m/m ground 40 m/m; 1 unk. ground 113 m/m ground 44 m/m ground 208 m/m ground 68 m/m ground

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42 Figure 5.9: Example of dung analysis seeds separated from dung and counted. 5.4: Forest Restoration Natural patterns of regeneration are influenced by five potentially limiting factors: site resources (i.e. soil and microclimate), competition with weeds, site disturbance, occurrence of established plants and/or propagules, and seed dispersal by animals. When land is deteriorated by poor agricultural practices and subsequently abandoned, it is of little use to local people and of little ecol ogical value. For this reason, Thai government officials recognized defore station as a concern both for conservation and impoverished rural communities. Theref ore, members of the government have promised to double the present forest cover fr om the current 20% forest to a hopeful 40% cover. They also promise to conserve 25 % of that for the pur pose of biodiversity (Hardwick et. al, 1997). The rest will be pl anted with only indigenous tree species, but be available for use by people. The goal is valid and theoretically attain able, given that logging was made illegal in 1989 by the Thai government, and there is land that has potential for reforestation.

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43 However this is not possible with current levels of ecological knowledge. To restore a forest, not just for timber concessions, but for the protection of biodiversity and native ecosystems, a more comprehensive knowledge of original forest regenerative processes is necessary. 5.7: Conclusions As I discussed in my literature review many problems in seed ecology studies stem from issues of accounting for spatial an d temporal scale. Variations in fruiting season, animal feeding patterns, small mammal and insect behavior a ll alter the fate of seeds at any given time. The fate of a seed can also be altered by how it is dispersed in regard to adult conspecifics and animal trails. Through cond ucting this short-term study, I realize why this is such an issue. Each step of the process is difficult from obtaining enough seeds and dung, to designing a study that is both accurate and replicable. As with field studies in any remote, tropical area, last minute changes to methodology are extremely difficult. When I went to Hu ai Kha Khaeng, I planned on studying secondary seed dispersal by dung beetles. When I began to set up my study, I realized it was necessary to start with a less specific topic, i.e. post-dispersal seed fate in general. However, by the time I was in the forest, I had no access to new supplies and had to do what I could with what I had with me. Additionally, my time in the field was very limited. If I continue this study, it will be nece ssary to stay in the forest for at least 6 months, if not more, so that seeds could be tracked through seedling establishment. Also, so little is known about the forest in genera l that many tangential studies would be highly valuable, such as recording gibbon foraging paths, tree phenology, and latrine use.

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44 It is easy enough to exclaim the importance of closing the seed disp ersal loop, but it is much more difficult in practice. Seed ecologists need to collaborate on several larger studies that can actually begin to shed light the process of s eedling recruitment. As it is (and my study is a case in point) each ecologi st has been attempting to re-invent the wheel of seed fate ecology, and many prelim inary studies have b een done. If these scientists were to pu t their ideas together, a more co mplete, longer-term study could be carried out. Additionally, while the conservatio n implications of these studies have been discussed, few of these have made it to the fi eld (but see the work of Thailands Forest Research and Restoration Unit, e.g. Elliot et al., 2003).

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45 VI: Bibliography Andresen, E. 1999. Seed Dispersal by Monkeys and the Fate of Dispersed Seeds in a Peruvian Rain Forest. Biotropica (31)145-158. Andresen, E. 2001. Effects of dung presence, dung amount and secondary dispersal by dung beetles on the fate of Micropholis guyanensis (Sapotaceae) seeds in Central Amazonia. Journal of Tropical Ecology (17)61-78. Andresen, E. 2002. Primary Seed Dispersal by Red Howler Monkeys and the Effect of Defecation Patterns on the Fate of Dispersed Seeds. Biotropica (34)261-272. Andresen, E and Levey, D.J. 2004. Effects of dung and seed size on secondary dispersal, seed predation, and seedling establ ishment of rain forest trees. Oecologia (139)45-54. Bunyavejchewin, S. 2002. Stand structure of a seasonal dry evergreen forest at Huai Kha Khaeng Wildlife Sanctuary, Western Thailand. Silvicultural Research Report http://www.forest.go.th/silvic/WP_Publications/Paper_SVGPub_PDF/ SilvicReport45/t1r45.pdf Byrne, M.M. and Levey, D.J. 1993. Removal of seeds from frugivore defecations by ants in a Costa Rican rain forest. Plant Ecology (107-108)363-374. Chapman, C.A. and Chapman, L.J. 1996. Frugivo ry and the Fate of Dispersed and NonDispersed Seeds of Six African Tree Species. Journal of Tropical Ecology (12)491-504. Connell, J. H. 1971. On the role of natural enemies in preventing competitive exclusion in some marine animals and in rain fo rest trees. In: Den Boer, P. J. and Gradwell, G. editors, Dynamics of populations. PUDOC, pp. 298. Corlett, R.T. 1998. Frugivory and seed dispersal by vertebrates in the Oriental (Indomalayan) Region. Biological Reviews of the Cambridge Philosophical Society (73)413-448. Crawley, M. 2000. Seed predators and plant population dynamics. In: Fenner, Michael, editor. Seeds: the ecology of regeneration in plant communities. 2nd ed. CABI Publishing. p.167. Elliot, S., Navakitbumrung, P., Kuarak, C., Zangkum, S., Anusarnsunthorn, V. and Blakesley, D. 2003. Selecting framework tree species for restoring seasonally dry tropical forests in northern Thai land based on field performance. Forest Ecology and Management (184)177-191. Estrada, A. and Coates-Estra da, R. 1991. Howler Monkeys ( Alouatta palliata ), Dung

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46 Beetles (Scarabaeidae) and Seed Dispersal: Ecological Interacti ons in the Tropical rain Forest of Los Tuxtlas, Mexico. Journal of Tropical Ecology (7)459-474. Fleming, T.H. and Heithaus, E.R. 1981. Frugivorous Bats, Seed Shadows, and the Structure of Tropical Forests. Biotropica (13)45-53. Forget, P. 1993. Post-dispersal pr edation and sca tterhoarding of Dipteryx panamensis (Papilionaceae) seeds by rodents. Oecologia (94)255-261. Hammond, D.S. and Brown, V.K. 1996. Di sturbance, phenology and life-history characteristics: factors influencing dist ance/density-dependent attack on tropical seeds and seedlings. Dynamics of tropical commun ities: the 37th symposium of the British Ecological Society Hammond, D.S., Brown, V.K. and Zagt, R. 1999. Spatial and temporal patterns of seed attack and germination in a large-seeded neotropical tree species. Oecologia (119)208-218. Hardwick, K., Healey, J., Elliot, S., Garwood, N., Anusarnsunthorn, V. 1997. Understanding and assisting natural regene ration processes in degraded seasonal evergreen forests in northern Thailand. Forest Ecology and Management (12)203-214. Howe, H.F. 1977. Bird Activity and Seed Dispersal of a Tropical Wet Forest Tree. Ecology (58) 539-550. Howe, H.F. and Vande Kerckove, G. 1979. Fec undity and seed disp ersal of a tropical tree. Ecology (60)180-189. Howe, H.F. and Smallwood, J. Ec ology of Seed Dispersal. 1982. Annual Review of Ecology and Systematics (13)201-228. Janzen, D.H. 1970. Herbivores and the Number of Tree Species in Tropical Forests. The American Naturalist. (940)501-528. Janzen, D.H. 1971. Seed Predation by Animals. Annual Review of Ecology and Systematics (2)465-492. Janzen, D.H. 1982. Removal of Seeds from Horse Dung by Tropical Rodents: Influence of Habitat and Amount of Dung. Ecology (63)1887-1900. Jordal, B.H., Normark, B.B. and Farrell, B.D. 2000. Evolutionary radiation of an inbreeding haplodiploid b eeltes lineage (Curculionidae, Scolytinae). Biological Journal of the Linnean Society (71)483-499. Kaufman, S., McKey, D.B., Hossaert-McKey, M. and Horvitz, C.C. 1991. Adaptations

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47 for a Two-Phase Seed Dispersal System Involving Vertebrates and Ants in a Hemiepiphytic Fig. American Journal of Botany (7)971-977. Kitajima, K. and Fenner, M. 2000. Ecology of Seedling Regeneration. In: Fenner, Michael, editor. Seeds: the ecology of regeneration in plant communities. 2nd ed. CABI Publishing. Levey, D.J. and Byrne, M.M. 1993. Complex AntPlant Interactions: Ra in-Forest Ants as Secondary Dispersers and Post -Dispersal Seed Predators. Ecology (74)1802-1812. Matias, L., Zamora, R., Mendoza, I., Hodar, J.A. 2008. Seed Dispersal Patterns by Large Frugivorous Mammals in a Degraded Mosaic Landscape. Restoration Ecology. Published online at www3.interscience.wiley.com. McConkey, K.R. and Galetti, M. 1999. Seed dispersal by the sun bear (Helarctos malayanus) in Central Borneo. Journal of Tropical Ecology (15)237-241. McConkey, K.R. 2005. Influence of faeces on seed removal from gibbon droppings in a dipterocarp forest in Central Borneo. Journal of Tropical Ecology (21)117-120. McConkey, K.R. 2005. The influence of gibbon primary seed shadows on post-dispersal seed fate in a lowland dipteroc arp forest in Central Borneo. Journal of Tropical Ecology (21)255-262. Nathan, R., Perry, G., Cronin, J.T., Strand, A.E. and Cain, M.L. 2003. Methods for estimating long-distance dispersal. Oikos (2):261-273. Nor, S.M.D. 2008. Elevational diversity patterns of small mammals on Mount Kinabalu, Sabah, Malaysia. Global Ecology and Biogeography (10)41-62. Notman, E.M. and Villegas, A.C. 2005. Patterns of Seed Predation by Vertebrate versus Invertebrate Seed Predators among Different Plant Species, Seasons and Spatial Distributions. In: Forget, Pierre-Michel, Lambert, Joanna E, Hulme, Philip E. and Vander Wall, Stephen B, editors. Seed fate: predation, dispersal, and seedling establishment Cambridge, MA. CABI Publishing. p. 55-75. Pizo, M.A. and Simao, I. 2001. Seed deposition patterns and the survival of seeds and seedlings of the palm Euterpe edulis. Acta Oecologia (22)229-233. Russo, S. and Augspurger, C.K. 2004. Aggreg ated seed dispersal by spider monkeys limits recruitment to clumped patterns in Virola calophylla. Ecology Letters (7)1058-1067. Schupp, E.W. 1990. Annual Variation in Seedfall, Postdispersal Predation, and Recruitment of a Neotropical Tree. Ecology (71)504-515.

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48 Schupp, E.W. 1993. Quantity, quality and the eff ectiveness of seed dispersal by animals. Plant Ecology (107-108)15-29. Shepherd, V.E. and Chapman, C.A. 1998. Dung beetles as secondary seed dispersers: impact on seed predation and germination. Journal of Tropical Ecology (14)199215. Sherman, P.M. 2002. Effects of land crabs on seedling densities and distributions in a mainland neotropical rain forest. Journal of Tropical Ecology (18)69-89. Traveset, A. 1998. Effect of seed passag e through vertebrate frugivores' guts on germination: a review. Perspectives in Plant Ecology, Evolution and Systematics 1(151-190). Traveset, A., Bermejo, T. and Willson, Ma ry. 2004. Effect of manure composition on seedling emergence and growth of two common shrub species of Southeast Alaska. Plant Ecology (155)29-34. Vander Wall, S.B., Forget, P., Lambert, J.E., and Hulme, P.E. Seed fate pathways: filling the gap between parent and offspring. In: Forget, Pierre-Michel, Lambert, Joanna E, Hulme, Philip E. and Vander Wall, Stephen B, editors. Seed fate: predation, dispersal, and seedling establishment. Cambridge, MA. CABI Publishing. p. 1-8. Vasquez-Yanes, C. and Orozco-Segovia, A. 1996. Physiological Ecology of Seed Dormancy and Longevity. Tropical Forest Plant Ecophysiology. von Allmen, C., Morellato, L.P. and Pizo, M.A. 2004. Seed predation under high seed density condition: the palm Euterpe edulis in the Brazilian Atlantic Forest. Journal of Tropical Ecology (20)471-474. Walker, S. and Rabinowitz, A. 1992. The Sm all-Mammal Community of a Dry-Tropical Forest in Central Thailand. Journal of Tropical Ecology (8)57-71. Wang, B.C. and Smith, T.B. 2002. Cl osing the seed dispersal loop. Trends in Ecology and Evolution (17)379-386. Wilson, D.E. and Janzen, D.H. 1972. Predation on Scheelea palm seeds by bruchid beetles: seed density and di stance from the parent palm. Ecology (53)954-959.

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49 Appendix I: Raw Data LEGEND: ID: Transect number; Seed #: seed number treatment; dung: dung treatment; BEGseed: number of seeds surviving from last check; scolytinae: number of seeds attacked by scolytinae; roden t/crab: number of seeds att acked by rodents or crabs; totalgerm: total number of germinated seeds; intactseed: total number of intact seeds; dead: total number of dead seeds (including scolytinae, crab); survive: total number of surviving seeds (including germ inated and intact seeds). Check 1 ID Seed# dung BEGSeed scolytinae rodent/crab totalgerm intactseed dead survive 26 5 nd 5 0 0 0 3 2 3 26 5 d 5 0 0 0 0 5 0 26 25 d 25 0 0 10 15 0 25 26 25 nd 25 0 0 17 8 0 25 20 5 nd 5 0 0 3 0 2 3 20 25 nd 25 0 0 15 10 0 25 20 5 d 5 0 0 0 5 0 5 20 25 d 25 0 0 10 15 0 25 11 5 d 5 0 0 2 3 0 5 11 5 nd 5 0 0 3 2 0 5 11 25 d 25 0 0 5 20 0 25 11 25 nd 25 0 0 9 16 0 25 18 25 d 25 0 0 12 13 0 25 18 5 d 5 0 0 1 4 0 5 18 5 nd 5 0 0 4 1 0 5 18 25 nd 25 0 0 13 12 0 25 31 5 nd 5 0 0 0 0 5 0 31 25 nd 25 0 0 10 15 0 25 31 5 d 5 0 0 3 2 0 5 31 25 d 25 0 0 13 11 1 24 33 5 d 5 0 0 0 5 0 5 33 25 nd 25 0 0 18 7 0 25 33 25 d 25 0 0 5 20 0 25 33 5 nd 5 5 0 0 0 5 0 27 5 nd 5 0 0 1 4 0 5 27 25 nd 25 0 0 12 13 0 25 27 25 d 25 0 0 11 14 0 25 27 5 d 5 0 0 1 4 0 5 32 25 nd 25 0 0 17 8 0 25 32 5 nd 5 0 0 1 3 1 4 32 5 d 5 0 0 1 4 0 5 32 25 d 25 0 0 10 15 0 25 16 25 d 25 0 0 1 24 0 25 16 25 nd 25 0 0 23 1 1 24 16 5 d 5 0 0 4 0 1 4 16 5 nd 5 0 0 4 0 1 4 15 5 nd 5 0 0 1 4 0 5

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50Check 1 ID Seed# dung BEGSeed scolytinae rodent/crab totalgerm intactseed dead survive 15 25 d 25 0 0 17 8 0 25 15 5 d 5 0 0 4 1 0 5 15 25 nd 25 0 0 1 24 0 25 19 5 nd 5 1 0 1 3 0 4 19 25 nd 25 0 0 10 15 0 25 19 25 d 25 0 0 8 17 0 25 19 5 d 5 0 0 2 3 0 5 23 5 nd 5 0 0 0 5 0 5 23 25 d 25 0 0 2 23 0 25 23 5 d 5 0 0 2 3 0 5 23 25 nd 25 0 0 7 18 0 25 14 25 nd 25 0 0 10 15 0 25 14 5 d 5 0 0 4 1 0 5 14 5 nd 5 0 0 1 1 3 2 14 25 d 25 0 0 1 24 0 25 34 5 d 5 0 0 2 3 0 5 34 25 nd 25 0 0 14 11 0 25 34 25 d 25 0 0 12 13 0 25 34 5 nd 5 0 0 2 3 0 5 37 25 d 25 0 0 0 25 0 25 37 5 nd 5 0 0 0 5 0 5 37 5 d 5 0 0 0 5 0 5 37 25 nd 25 0 1 1 23 1 24 30 5 nd 5 0 0 3 2 0 5 30 25 d 25 0 0 5 20 0 25 30 5 d 5 0 0 0 5 0 5 30 25 nd 25 0 0 11 14 0 25 29 25 nd 25 0 0 17 8 0 25 29 5 d 5 0 0 3 2 0 5 29 5 nd 5 0 0 3 2 0 5 29 25 d 25 0 0 15 10 0 25 13 5 nd 5 0 0 2 3 0 5 13 25 d 25 0 0 10 15 0 25 13 5 d 5 0 0 2 3 0 5 13 25 nd 25 0 0 18 7 0 25 21 5 d 5 0 0 3 2 0 5 21 25 d 25 0 0 0 25 0 25 21 25 nd 25 0 0 15 10 0 25 21 5 nd 5 0 0 1 4 0 5 5 25 nd 25 0 0 20 5 0 25 5 5 d 5 0 0 0 5 0 5 5 5 nd 5 0 0 0 5 0 5 5 25 d 25 0 0 4 21 0 25 36 25 d 25 0 0 12 13 0 25 36 5 d 5 0 0 0 5 0 5 36 25 nd 25 0 0 17 8 0 25

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51Check 1 ID Seed# dung BEGSeed scolytinae rodent/crab totalgerm intactseed dead survive 36 5 nd 5 0 0 1 3 0 4 25 5 d 5 0 0 0 5 0 5 25 5 nd 5 0 0 1 4 0 5 25 25 nd 25 0 0 13 12 0 25 25 25 d 25 0 0 3 22 0 25 1 25 d 25 0 0 3 22 0 25 1 5 d 5 0 0 0 5 0 5 1 25 nd 25 0 0 15 9 1 24 1 5 d 5 0 0 1 4 0 5 Check 2 ID Seed# dung BegSeed scolytinae rodent/crab totalgerm intactseed dead survive 26 5 nd 5 5 0 0 0 5 0 26 5 d 0 5 0 0 0 5 0 26 25 d 25 0 0 10 15 0 25 26 25 nd 25 4 0 18 3 4 21 20 5 nd 3 5 0 0 0 5 0 20 25 nd 25 3 0 19 3 3 22 20 5 d 5 0 0 2 3 0 5 20 25 d 25 0 0 16 9 0 25 11 5 d 5 0 0 2 3 0 5 11 5 nd 5 3 0 2 0 3 2 11 25 d 25 0 0 5 20 0 25 11 25 nd 25 13 0 10 2 13 12 18 25 d 25 1 0 20 0 1 20 18 5 d 5 0 0 3 0 0 3 18 5 nd 5 2 0 3 0 2 3 18 25 nd 25 10 0 12 3 10 15 31 5 nd 0 5 0 0 0 5 0 31 25 nd 25 14 0 7 4 14 11 31 5 d 5 1 0 2 2 1 4 31 25 d 24 1 0 18 6 1 24 33 5 d 5 1 0 1 3 1 4 33 25 nd 25 0 0 19 6 0 25 33 25 d 25 0 0 18 7 0 25 33 5 nd 0 0 0 0 0 0 0 27 5 nd 5 4 0 1 0 4 1 27 25 nd 25 7 0 0 10 7 10 27 25 d 25 0 0 1 24 0 25 27 5 d 5 0 0 3 2 0 5 32 25 nd 25 7 0 18 0 7 18 32 5 nd 5 5 0 0 0 5 0 32 5 d 5 0 0 2 3 0 5 32 25 d 25 0 0 10 15 0 25 16 25 d 25 0 0 0 25 0 25

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52Check 2 ID Seed# dung BegSeed scolytinae rodent/crab totalgerm intactseed dead survive 16 25 nd 24 1 0 22 2 1 24 16 5 d 4 1 0 4 0 1 4 16 5 nd 4 2 0 2 0 2 2 15 5 nd 5 2 0 1 2 2 3 15 25 d 25 0 0 19 6 0 25 15 5 d 5 0 0 5 0 0 5 15 25 nd 25 15 0 4 5 15 9 19 5 nd 4 0 0 1 0 0 1 19 25 nd 25 4 0 10 0 4 10 19 25 d 25 0 0 0 23 0 23 19 5 d 5 0 0 0 3 0 3 23 5 nd 5 5 0 0 0 5 0 23 25 d 25 0 0 1 4 0 5 23 5 d 5 0 0 2 3 0 5 23 25 nd 25 5 0 17 3 5 20 14 25 nd 25 9 0 17 0 9 17 14 5 d 5 0 0 4 1 0 5 14 5 nd 2 3 0 1 1 3 2 14 25 d 25 0 0 1 24 0 25 34 5 d 5 0 0 2 3 0 5 34 25 nd 25 8 1 13 3 9 16 34 25 d 25 0 0 15 10 0 25 34 5 nd 5 3 0 1 1 3 2 37 25 d 25 0 0 1 24 0 25 37 5 nd 5 4 1 0 0 5 0 37 5 d 5 0 0 0 5 0 5 37 25 nd 24 10 0 14 4 10 18 30 5 nd 5 0 0 2 0 0 2 30 25 d 25 0 0 8 17 0 25 30 5 d 5 0 0 2 3 0 5 30 25 nd 25 7 0 16 2 7 18 29 25 nd 25 5 0 20 0 5 20 29 5 d 5 3 0 2 0 3 2 29 5 nd 5 1 0 3 1 1 4 29 25 d 25 0 0 20 5 0 25 13 5 nd 5 4 0 1 0 4 1 13 25 d 25 0 0 12 13 0 25 13 5 d 5 0 0 3 2 0 5 13 25 nd 25 2 0 22 1 2 23 21 5 d 5 0 0 4 1 0 5 21 25 d 25 0 0 15 10 0 25 21 25 nd 25 8 0 15 2 8 17 21 5 nd 5 5 0 0 0 5 0 5 25 nd 25 4 0 17 4 4 21 5 5 d 5 1 0 1 3 1 4 5 5 nd 5 0 0 5 0 0 5 5 25 d 25 0 0 7 18 0 25

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53Check 2 ID Seed# dung BegSeed scolytinae rodent/crab totalgerm intactseed dead survive 36 25 d 25 0 0 14 11 0 25 36 5 d 5 5 0 0 0 5 0 36 25 nd 25 1 0 19 5 1 24 36 5 nd 4 3 0 1 1 3 2 25 5 d 5 5 0 0 0 5 0 25 5 nd 5 5 0 0 0 5 0 25 25 nd 25 13 0 12 0 13 12 25 25 d 25 1 0 4 20 1 24 1 25 d 25 0 0 5 20 0 25 1 5 d 5 5 0 0 0 5 0 1 25 nd 24 10 0 15 0 10 15 1 5 d 5 0 0 1 4 0 5 Check 3 ID Seed# dung BegSeed scolytinae rodent/crab totalgerm intactseed dead survive 26 5 nd 0 0 0 0 0 0 0 26 5 d 0 5 0 0 0 5 0 26 25 d 25 17 0 9 0 17 9 26 25 nd 21 11 0 9 1 11 10 20 5 nd 0 0 0 0 0 0 0 20 25 nd 22 5 0 15 1 5 16 20 5 d 5 3 0 2 0 3 2 20 25 d 25 8 0 16 1 8 17 11 5 d 5 3 0 2 0 3 2 11 5 nd 2 1 0 1 0 1 1 11 25 d 25 9 0 16 0 9 16 11 25 nd 12 2 0 10 0 2 10 18 25 d 20 5 0 20 0 5 20 18 5 d 3 2 0 4 0 1 4 18 5 nd 3 2 0 1 0 2 1 18 25 nd 15 7 0 7 0 7 7 31 5 nd 0 0 0 0 0 0 0 31 25 nd 11 6 0 5 0 6 5 31 5 d 4 4 0 0 0 4 0 31 25 d 24 11 0 13 1 11 14 33 5 d 4 4 0 1 0 4 1 33 25 nd 25 3 0 20 2 3 22 33 25 d 25 0 0 21 4 0 25 33 5 nd 0 0 0 0 0 0 0 27 5 nd 1 1 0 0 0 1 0 27 25 nd 10 5 0 3 2 5 5 27 25 d 25 10 0 15 0 10 15 27 5 d 5 2 0 2 0 2 3 32 25 nd 18 9 0 9 0 9 9 32 5 nd 0 0 0 0 0 0 0 32 5 d 5 3 0 2 0 3 2 32 25 d 25 6 6 8 1 12 9

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54Check 3 ID Seed# dung BegSeed scolytinae rodent/crab totalgerm intactseed dead survive 16 25 d 25 25 0 0 0 25 0 16 25 nd 24 3 0 21 0 3 21 16 5 d 4 3 0 1 0 3 2 16 5 nd 2 1 0 2 0 1 2 15 5 nd 3 4 0 0 0 4 0 15 25 d 25 8 0 17 0 8 17 15 5 d 5 0 0 5 0 0 5 15 25 nd 9 5 0 4 0 5 4 19 5 nd 1 1 0 0 0 1 0 19 25 nd 10 0 0 10 0 0 10 19 25 d 23 23 0 0 0 23 0 19 5 d 3 0 0 3 0 0 3 23 5 nd 0 5 0 0 0 5 0 23 25 d 5 24 0 0 1 24 1 23 5 d 5 5 0 0 0 5 0 23 25 nd 20 3 0 17 0 3 17 14 25 nd 17 11 0 6 0 11 6 14 5 d 5 1 0 4 0 1 4 14 5 nd 2 1 0 1 0 1 1 14 25 d 25 24 0 1 0 24 1 34 5 d 5 5 0 0 0 5 0 34 25 nd 16 11 0 5 0 11 5 34 25 d 25 6 0 17 0 6 19 34 5 nd 2 2 0 0 0 2 0 37 25 d 25 20 0 2 0 20 2 37 5 nd 0 0 0 0 0 0 0 37 5 d 5 5 0 0 0 5 0 37 25 nd 18 4 0 13 1 4 14 30 5 nd 2 4 0 0 1 4 1 30 25 d 25 7 0 17 1 7 18 30 5 d 5 2 0 3 0 2 3 30 25 nd 18 7 0 16 2 7 18 29 25 nd 20 5 0 20 0 5 20 29 5 d 2 3 0 2 0 3 2 29 5 nd 4 3 0 2 0 3 2 29 25 d 25 4 0 20 0 4 20 13 5 nd 1 5 0 0 0 5 0 13 25 d 25 10 0 14 1 10 15 13 5 d 5 1 0 4 0 1 4 13 25 nd 23 2 0 16 5 2 21 21 5 d 5 1 0 4 0 1 4 21 25 d 25 7 0 17 0 7 17 21 25 nd 17 7 0 10 0 7 10 21 5 nd 0 0 0 0 0 0 0 5 25 nd 21 5 0 16 0 5 16 5 5 d 4 3 0 1 1 3 2

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55Check 3 ID Seed# dung BegSeed scolytinae rodent/crab totalgerm intactseed dead survive 5 5 nd 5 5 0 0 0 5 0 5 25 d 25 17 0 7 0 17 7 36 25 d 25 13 0 11 0 13 11 36 5 d 0 5 0 0 0 5 0 36 25 nd 24 2 0 20 0 2 20 36 5 nd 2 1 0 1 0 1 1 25 5 d 0 5 0 0 0 5 0 25 5 nd 0 0 0 0 0 0 0 25 25 nd 12 2 0 10 0 2 10 25 25 d 24 24 0 1 0 24 1 1 25 d 25 24 0 1 0 24 1 1 5 d 0 0 0 0 0 0 0 1 25 nd 15 6 0 9 0 6 9 1 5 d 5 4 0 1 0 4 1 In minimus Natura praestat. Pliny


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