The evolution of plant-insect interactions: Insights from the tertiary fossil record

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THE EVOLUTION OF PLANT-INSECT INTERACTIONS:

INSIGHTS FROM THE TERTIARY FOSSIL RECORD

by Dena Michelle Smith

Copyright © Dena Michelle Smith 2000

A Dissertation Submitted to the Faculty of the

DEPARTMENT OF GEOSCIENCES

In Partial Fulfillment of the Requirements For the Degree of

DOCTOR OF PHILOSOPHY

In the Graduate College

THE UNIVERSITY OF ARIZONA

2000 UMI Number; 9992055

Copyright 2000 by Smith, Dena Michelle

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ACKNOWLEDGMENTS

I received help from several people to complete this dissertation, but I especially want to thank my husband, Cesar Nufio, for his constant encouragement and incredible patience. I am inspired by his intelligence, his great kindness, and his ability to find humor in almost any situation. Without his support I would not have completed this dissertation. I also want to thank my major advisor, Karl Flessa. who has been encouraging and supportive throughout. I especially appreciated his calm demeanor and his ability to diffuse difficult situations with humor and words of wisdom. He created a lab environment that combined good science and lots of fun, a perfect environment for exploring and learning. I want to thank Dan Papaj for inviting me into his lab space in my last year. I would have never gotten a thing done without this wonderful opportunity and I really enjoyed our daily chats. Judy Parrish gave me great advice on a number of occasions and has been a wonderful role model. Maria Teresa Velez helped me to find my home with the community of color at the Univeristy of Arizona and her support made a big difference. I also want to thank the Organization for Tropical Studies and Dr. Deedra Mc Learn for an extremely rewarding experience in the 98-1 tropical biology field course and the opportunity to participate in subsequent field courses. The National Park Service let me use of their collections, facilities and their fossil database. The people working at Florissant Fossil Beds National Monument were great and I especially thank Dr. Herb Meyer for his support, excellent advice, help with fossil plant identification and p>owers of paleo-divination. Special thanks must go to Apama Telang, Simone Alin, Jessa Netting and April Kinchloe. Their friendship has been a source of strength and their intelligence a source of inspiration.

Financial support for this dissertation was provided by a National Science Foundation predoctoral fellowship, the University of Arizona's Research Training Grant for the Analysis of Biological Diversification, the Department of Geological Sciences' Chevron and Exxon summer research grants, the Organization for Tropical Studies. Colorado Natural Areas Program, Sigma Xi, and the Graduate College of the University of Arizona. Kiu-1 Flessa also provided funding, computing equipment, collecting gear, fossil storage space and Paleontologist Barbie. This is dissertation #2 of the Centro de Estudios de Almejas Muertas. 5

DEDICATION

This dissertation is dedicated to my father. Frank A. Smith. 6

TABLE OF CONTENTS

ABSTRACT 8

CHAPTER 1. INTRODUCTION 10

Plant-Insect Interactions; An Overview 10

The Importance of Insect Taphonomy 14

The Importance of Plant Taphonomy 17

Modem Herbivory and Plant Taphonomy 19

Format of the Dissertation 20

CHAPTER 2. PRESENT STUDY 22

Summary of Appendix A: Beetle Taphonomy in a Recent Lake 22

Summary of Appendix B: Herbivory in Modem Tropical Forests 23

Summary of Appendix C: Patterns of Herbivory from the Tertiary 23

CHAPTER 3. CONCLUSIONS AND FUTURE STUDY 25

APPENDIX A: Beetle taphonomy in a recent ephemeral lake.

Southeastern Arizona .35 7

APPENDIX B: Levels of Herbivory in Two Costa Rican

Rainforests: Implications for Studies of Fossil Herbivory 86

APPENDIX C: The Evolution of Plant-Insect Interactions: Insights from the Tertiary

Fossil Record 113

APPENDIX D; SUPPLEMENTARY DATA 167

Locality Information 168

Summary Statistics 168

Herbivory data from Arizona 169

Herbivory data from California 181

Herbivory data from Cerro de la Muerte 187

Herbivory data from Corcovado 198

Herbivory data from La Selva 219

Herbivory data from Palo Verde 250

Herbivory data from Green River 260

Herbivory data from Florissant 280

REFERENCES 304 8

ABSTRACT

Plant-feeding insects are the most species-rich group on the planet today.

Models have been proposed to explain this diversity, but few use the fossil record to evaluate hypotheses. I conduct studies in modem systems to examine (1) taphonomic biases in insect preservation and how this may affect our understanding of insect diversity trends through time and (2) patterns of herbivory in modem ecosystems to improve the comparability of fossil and modem datasets. I then use the Cenozoic fossil record to examine the history of ecological associations between insects and plants and how these interactions respond to environmental change.

I conducted an actualistic study on the preservation of beetles in Willcox Playa. an ephemeral lake in SE Arizona. I compared the insect death assemblage in shoreline sediments to the living beetle assemblage. The sediments captured 56% of the live- collected beetle families, and 28% of the live-collected beetle genera. The relative abundances of living beetles were not reflected in the death assemblage. Beetle dieL feeding habitat, and size influenced the composition of the death assemblage.

Necrophagous, ground-dwelling and smaller beetles were over-represented in the death assemblage. Such biases should be considered in insect paleoecology and in studies of diversity change.

Annual variation in herbivory was compared within and between two lowland neotropical forests Costa Rica. Herbivory did not vary significantly within sites between years, but was significantly different between sites. Modem herbivory data collected with discrete sampling techniques is compartable to herbivory data from fossil 9 forests. Herbivory data from one-time collections of leaf litter are most suitable for comparison with fossil herbivory.

I compared herbivory between two Eocene floras and between the Eocene floras and six modem floras. A decline in levels of herbivory corresponds with a decline in temperature from the middle to the late Eocene. Fossil herbivore damage was significantly lower than modem herbivore damage. This pattern may result from laphonomic bias, environmental differences between the fossil and modem sites or evolutionary change. 10

CHAPTER 1: INTRODUCTION

Are ecological interactions stable through evolutionary/geologic time? I use the

Cenozoic fossil record to examine the history of ecological associations between insects and plants and the evolution of these interactions in response to environmental change.

Due to the paleoecological nature of this work. I found it necessary to explore the taphonomy of insects and plants in lacustrine environments. While the plant taphonomy literature is quite extensive, the insect taphonomy literature is much more limited, and as a result I conducted my own actualistic study of beetle preservation in a playa-lake envirorunent. In addition, I conducted my own study of modem herbivory levels for comparison with the Cenozoic fossil data. Each of these topics and their importance in the dissertation are discussed below.

PLANT-INSECT INTERACTIONS: AN OVERVIEW

Insects constitute the most species-rich group on the planet today. Current estimates of insect species diversity range from 3 to 50 million species (Erwin 1982;

Gaston 1991 and 1992; May 1986). Within the insects, the plant-feeding groups are the most diverse and many workers have suggested that plant feeding has promoted diversification in insects (Farrell and Mitter 1990; Mitter et al. 1988). In addition. 75% of all phytophagous insects specialize on only one or a few related plant species

(Bemays and Chapman 1994). Ehrlich and Raven (1964) first tried to explain the high diversity of plant-feeding insects in their paper on the coevolution of butterflies and their host-plants. They proposed that plants and insects are locked in an arms-race of 11 plant defense and insect counter-defense which is responsible for generating insect and plant diversity. Since that time, many workers have looked for patterns of coevolution in other herbivore-plant associations and have examined patterns of parallel cladogenesis and other important factors in insect-plant interactions, such as the effects of plant secondary compounds (many of which act as toxins and/or feeding deterrents) on insect diet breadth (Berenbaum 1983; Feeny 1975 and 1976; Moran and Hamilton

1980; Rhoades and Gates 1976), plant nutritional quality (Chan et al. 1978; Rausher

1981; Scriber 1977; Scriber and Feeny 1979, Scriber and Slansky 1981), plant and insect population variation in space and time (Feeny 1970; Futuyma and Wasserman

1980; Weis and Berenbaum 1989), and multi-trophic interactions (Flint et al. 1979;

Inouye and Taylor 1979; Whitman 1994 and Wood 1982).

Although there is an extensive body of research on plant-insect interactions, virtually all of these studies have focused on the ecology of associations in modem systems. Only a few studies have sought to understand the evolution of plant and insect interactions at macroevolutionary time-scales. One approach has been to reconstruct the phylogenies of closely associated clades to see if these groups have diversified in parallel (Farrell and Mitter 1990; Funk et al. 1995; Futuyma et al. 1995). Another approach uses sister-group comparisons to see if plant-feeding groups are more diverse than their non-herbivorous sister-clades (Farrell 1998; Farrell et al. 1991; Mitter et al.

1988). These and other analyses have shown evidence for patterns of reciprocal evolution between plants and insects, which would be expected based on the coevolutionary model first proposed by Ehrlich and Raven in 1964. 12

Although the phylogenetic work seems to indicate that many specialized plant and insect interactions have persisted over great evolutionary time-scales, few studies have examined the fossil record for evidence of these associations. The paleontological studies that have been conducted only provide summaries of the history of general patterns of feeding damage (Labandeira and Beall 1990; Scott and Taylor 1983: Smart and Hughes 1972) whereas otliers present isolated occurrences of long-lived, host- specific insect damage (Hickey and Doyle 1977; Hickey and Hodges 1975; Labandeira el al. 1994; Larew 1986; Opler 1973; Upchurch and Dilcher 1990; Wilf et al. 2000).

Only recently have there been any attempts to examine community-wide patterns in herbivory (Ash 1997; Beck and Labandeira 1998. Smith 1998) to document change in herbivory over time (Wilf and Labandeira 1999). This new approach will make it possible to seek out evidence of specific plant-insect interactions and to track changes in plant-herbivore community dynamics through time.

In this dissertation, 1 use a community-based approach to examine the traces of

Insect-feeding damage on fossil plant assemblages from two Tertiary sites, the Green

River Formation (Middle Eocene) and Florissant Fossil Beds (Late Eocene) of

Colorado. It is likely that many modem plant and insect associations had their beginnings during the Tertiary because the first appearances of many modem insect and plant groups occurred at this time (Carpenter 1992; Wing 1987). My work provides documentation of the initiation and early development of modem plant-insect associations. By comparing the damage levels found at these sites to each other and to 13

patterns of modem herbivory, I also attempt to ascertain whether ecological interactions

are stable through evolutionary/geologic time. For example, were plants attacked by

insect herbivores during the past with the same level of intensity as they are today?

Have the associations between particular insects and particular plants changed over

time? Finally, how does climate change affect the associations between plants and

insects? 1 present this work in appendix C, but summarize the results from these studies

here.

In this study, I compared the insect-mediated damage on fossil leaves from the subtropical (46Ma) Green River Formation, fossil leaves from the temperate (35Ma)

Florissant Fossil Beds, and herbivore damage from leaf litter samples collected from six modem forests. The Florissant flora had lower levels of insect-damage than the Green

River flora, with significantly lower amounts of leaf area removed from the assemblage

(1.4% vs. 2.5%) and a lower percentage of leaves with insect damage (23% vs. 34%).

These differences may be the result of taphonomic bias, global climate change, changes in atmospheric CO2 and differences in paleoaltitude between the two sites.

I found that herbivore intensity was much lower in the fossil floras compared to the modem floras. Only 1.4-2.5% of the leaf area was removed by insect herbivores in the fossil assemblage compared to 4.3-10% in the modem floras. Twenty-three to thirty-four percent of the fossil leaves had insect damage compared to the modem samples, which had from 72-90% of the leaves damaged. Damaged leaves in the fossil 14 floras were usually (81-88%) attacked by only one feeding group and leaves in the modem samples were typically (41-67%) attacked by multiple feeding groups.

In addition, there were four examples of a highly specialized damage patterns found in both the Green River and Florissant floras and several examples of damage that appear to be maintained from the Eocene to the Recent. There was also an increase in the diversity of galling damage from the middle to the late Eocene, corresponding with a cooling and drying trend.

THE IMPORTANCE OF INSECT TAPHONOMY

What is known about the accumulation and preservation of insects in the fossil record? Insect fossils are found preserved in a number of environments including lagoonal sediments, amber inclusions, coal swamp concretions and in a variety of forms within lake deposits. Lakes are perhaps the most common environment in which to find insect fossils and the most geologically extensive (Labandeira 1999). However, few studies have addressed the process of insect accumulation and preservation in lakes.

Studies of fossil insect preservation in general have tended to fall into two main categories: (1) studies that examine the preservation of insect cuticle, and (2) those that examine the composition of insects within fossil assemblages. I briefly discuss this work below.

The insect exoskeleton serves as the external structure that protects an insects' internal organs, as a boundary layer that prevents desiccation, and as an attachment site 15

for muscles. The exoskeleton, or cuticle, is composed of both of chitin and protein

fibers that criss-cross and brace each other into a highly strengthened composite

material (Chapman 1982). Although robust and durable, the proteinaceous components

of the insect cuticle often degrade more quickly than the chitinous components. In a

study of Pleistocene insects from California asphalt deposits (Stankiewicz et al. 1997a).

fossil beetles possessed chitin that was almost entirely intact, whereas the protein

components were highly degraded. The oldest documented traces of chitin are found in

an Oligocene (25Ma) deposit in Enspel, Germany (Stankiewicz et al. 1997b). This

lacustrine deposit preserves beetles that still have intact chitin. However, not all of the

insect specimens found at Enspel have traces of chitin and the authors proposed that this

may result from the thicker and more heavily sclerotized exoskeleton of beetles, and

consequently a delayed interval for the complete degradation of chitin. It also has been

suggested that pH may be an important factor in the preservation of chitin and that more

acidic environments may actually enhance chitin degradation (Briggs 1999).

Wilson (1988) utilized the second approach to insect taphonomy when he

examined fossil insect assemblages to determine the preservation of insects in lakes.

Wilson compared the faunal composition and physical conditions of specimens between

near-shore and off-shore environments in several different lake deposits from the

Tertiary of North America. Wilson found that more insects were preserved in near- shore environments compared to off-shore environments and that insects in the near- shore environments were more fragmented. He also found important taxonomic 16 differences between near-shore and ofF-shore deposits, as different insect orders were more abundant in the different settings.

While the chemistry of insect cuticle and the faunal composition of fossil insect assemblages have been examined, no previous studies have explored the role of ecology and morphology in insect preservation. Therefore. I conducted an actualistic study of beetle accumulation and preservation in an ephemeral lake environment. I compared the living beetle assemblage of a playa lake to its accumulating death assemblage by examining the taxonomic composition, relative abundance, diet, feeding habitat and size of the live-collected and dead beetles. Fifty-six percent of the living beetle families and twenty-eight percent of the living beetle genera were found in the sediments. The relative abundances of beetle families in the live assemblage are significantly different from the relative abundances in the death assemblage. Among diet and habitat groups, necrophagous and ground-dwelling beetles are over-represented in the death assemblage, and the sediments contain a greater proportion of smaller, more robust beetles.

Although there have been only a few studies on insect taphonomy, it is clear that there are several taphonomic biases that can influence insect preservation in lacustrine environments. Wilson's (1988) study demonstrated the impwrtance of depositional envirorunents and how they cam affect the taxonomic composition and physical condition of specimens in an insect assemblage, and my work has shown the importance of insect ecology and morphology in the preservation of insects in ephemeral lake environments. 17

THE IMPORTANCE OF PLANT TAPHONOMY

Is the fossil record of plants preserved in lake environments good enough to study ecological interactions between plants and insects? I believe that the answer is yes. Paleobotanists have examined many different preservational environments, different modes of plant preservation (compressions, concretions, permineralized material and other modes) and the preservation potential of different plant parts (such as pollen, spores, seeds, fhiits, wood and leaves). Much of our understanding about the taphonomy of fossil leaves has been acquired through the examination of modem floras and preservational environments, experimental studies that have concentrated on the manipulation of plant material in the lab, and the study of fossil plant assemblages. In the following sections, I provide a brief review of the plant taphonomy literature, focusing on the preservation of leaves in lacustrine environments.

For leaves to become preserved in lake environments, they must first travel from the source plant to the lake. During this stage, leaves can become filtered on the basis of leaf size, the growth form of the source plant and the position of the plant in relation to the lake. Studies of modem leaf accumulations (Bumham et al. 1992, Ferguson

1985. and Greenwood 1992) have shown that leaves do not travel far from their source and in general, leaves will not travel farther than a lateral distance equal to the height of the tree. In addition, fossil leaf assemblages have a tendency to be dominated by smaller "sun" leaves that are derived from the forest canopy (Ferguson 1985;

Greenwood 1992; Spicer 1981). 18

Leaves may also be transported into a lake environment via rivers and streams,

which also affects the potential of leaf preservation. Spicer and Wolfe (1987)

conducted a study of plant preservation in a California lake, whose drainage area

consisted of six separate basins, and they found that the majority of leaves found in the

sediment samples represented species that were abundant along the margins of the lake.

However, there were a few extra-local species that were transported into the lake.

Once a leaf reaches the lake, it must then pass through the water column and

become buried by sediment. Leaf float time, biotic decay and chemical factors also

play a role in the preservation of a leaf. For example, experimental studies (Ferguson

1985; Spicer 1981) that examined leaf floating times in water tanks showed that deciduous leaves float on the water surface for approximately two days, but agitation to

the water surface from wind or rain will decrease a leaTs floating time. Ferguson

(1985) found that herbivore-damaged leaves have shorter floating times as a consequence of the leaves becoming waterlogged more quickly than undamaged leaves.

This suggests that herbivore-damaged leaves may be over-represented in lacustrine assemblages.

The longer time p>eriod that leaves spend in the water column, the more susceptible they are to bacteria, fimgi and other organisms. Biotic degradation can be influenced by the chemical composition of leaves (Spicer, 1977) and factors such as temperature, oxygen and nutrient levels in lakes (Hanlon, 1982). However, there are times when microbial activity can have a positive effect on leaf preservation. Bacteria sometimes precipitate a film that will attach to leaves. This film can protect the leaf 19 from abrasion or attack by other decomposers (Spicer, 1977). In addition, some work suggests that diatom oozes may have played a role in the exceptional preservation of plants and insects at Florissant Fossil Beds (Harding and Chant 2000; O'Brien et al.

1998).

Overall, this is good news for paleoecologists. Lake deposits tend to preserve the leaves of plants from the local environment and the relative abundance of leaves tends to be in proportion to the abundance of trees in the original forest. This means that carefril sampling of a fossil leaf assemblage can provide a good sample of the plants in the original flora.

MODERN HERBIVORY AND PLANT TAPHONOMY

Despite the good news with regard to plant taphonomy, concerns still exist, such as the bias against the preservation of understory leaves and the bias for the preservation of smaller leaves. These biases could be important if there are differences in the levels of insect herbivory between the canopy and the understory and between small and large leaves (a discussion of leaf size and herbivory can be found in the discussion section of appendix C).

The majority of studies on modem insect herbivory have been conducted on understory plants (de la Cruz and Dirzo 1987. Dirzo 1984. Filip et al. 1995, Hendrix and Marquis 1983, Johnstone 1981, Newberry and de Foresta 1985, Odum and Ruiz

1970 and Sterck et al. 1992). One study examined both understory and canopy herbivory and showed that herbivory levels are lower in the canopy than in the 20

understory (Lowman and Heatwole 1992). Because of the potential for this bias. I

collected modem herbivory data from leaf litter samples from a total of six modem

forests, including two tropical lowland forests, one tropical dry forest, one tropical high-

altitude forest and two temperate forests. The raw data collected from these localities

are presented in appendix D and are compared to the fossil data in appendix C.

I examined the levels of herbivory in two Costa Rican rainforests over a two and

three year jjeriod. Comparisons were made between years at each site and between

sites. The mean percent leaf area removed by insect herbivores did not vary

significantly between years, but was different between sites. That there is little

variation in levels of herbivory at ecological time-scales is good news. If herbivory

levels fluctuated greatly within a single forest, then it would be difficult to compare

herbivory levels from fossil floras to modem levels of herbivory. Although additional

studies should be conducted using leaf-litter sampling, current methods of discrete

sampling in the understory may be useful for the comparison of herbivory levels in

modem and fossil assemblages.

FORMAT OF THE DISSERTATION

I present three manuscripts that examine 1) the preservation of insects in lake

environments. 2) herbivory levels and taphonomic processes in modem tropical

ecosystems and 3) long-term change in plant-insect interactions through the comparison of Tertiary floras with modem leaf litter assemblages. All of these manuscripts are

single authored, original works. Appendix A has been published in Palaios and 21 appendix B is in review with Biotropica. Summary statistics and all the herbivory data from the fossil and the modem floras are presented in Appendix D. 22

CHAPTER 2: PRESENT STUDY

The methods, results and conclusions of this work are presented in the papers appended to this thesis. The following is a summary of the most important findings presented in these papers.

Summary of Appendix A: Beetle taphonomy in a recent ephemeral lake

A comparison of the laxonomic composition, relative abundance, diet, feeding habitat, and size of live and dead insects was conducted in Willcox Playa, an ephemeral lake in southeastern Arizona. Death assemblages of beetles, the most abundant group of insects in this study, are preserved in the shallow, subsurface sediments along the shoreline of the lake. Fifty-six percent of the living beetle families and twenty-eight percent of the living beetle genera were found in the sediments; one-hundred percent of the families and ninety-one percent of the genera found dead were also present in the live fauna. Relative abundances of beetle families in the live assemblage are significantly different than relative abundances in the death assemblage. Among diet and habitat groups, necrophagous and ground-dwelling beetles are over-represented in the death assemblage, while wood-inhabitants and aquatics are under-represented. The death assemblage contains a greater proportion of smaller, more robust beetles. Such biases also may occur in fossil assemblages and should be considered in paleoecological and paleoenvironmental reconstructions. 23

Summary of Appendix B: Herbivory in two Costa Rican rainforests

Levels of herbivory in two Costa Rican rainforests were measured over a two and three year period. Comparisons were made between years at each site and between sites. The mean percent leaf area removed by insect herbivores did not vary significantly between years, but was different between sites. Leaf area removed by insect herbivores ranged from 4 to 11%. This is within the range of values typically seen in discrete samples from neotropical rainforests. Patterns in guild structure varied between years and between forests. While the differences between these samples were significant, overall trends remained the same and this may provide useful information when examining long-term change in guild structure. Samples were collected from leaf litter to ensure the comparability of the data in this study to data from studies of herbivory in fossil assemblages. Although additional studies should be conducted using leaf-litter sampling, current methods of discrete sampling in the understory may be useful for the comparison of herbivory levels in modem and fossil assemblages.

Summary of Appendix C: The Evolution of Plant-Insect Interactions.

Has there been dramatic change in plant-insect interactions over geologic time?

1 compared the insect-mediated damage on fossil leaves from the 46 Ma (Middle

Eocene) Green River Formation at Douglas Pass, Colorado to herbivore damage in the

35 Ma (Late Eocene) Florissant Flora of Colorado. Insect damage from these fossil sites was also compared to herbivore damage on leaves that were collected from the 24

leaf-litter of six modem tropical and temperate sites. The intensity of herbivore damage was measured by examining the percentage of leaf area removed by insect herbivores, the number of leaves damaged in the floras, the number of feeding guilds on individual leaves in the floras, and the overall feeding guild structure.

Levels of herbivory declined (from 2.5% to 1.4% leaf area removed and 34% to

23% of leaves damaged) as temperature declined during the Eocene-Oligocene cooling trend. Levels of insect damage in the fossil floras (23-34% of leaves damaged and no more than 3 feeding types on a leaf) were lower than levels found in the modem floras

(72-90% of leaves damaged and greater than 3 types of feeding damage on a leaf). The differences in herbivory between the fossil and the modem floras may due to taphonomic biases, environmental differences between the fossil and modem sites or evolutionary change through time.

Evidence of several long-lived specialized plant-insect associations that were maintained from the middle to the late Eocene and from the late Eocene to the Recent were also observed. Galling diversity increased from the wetter, middle Eocene Green

River flora to the drier, late Eocene Florissant flora. Modem galling insects have greater survivorship and higher diversity in more xeric envirorunents and this may explain the pattern observed in galling diversity during the middle to late Eocene. 25

CHAPTER 3: CONCLUSIONS AND FUTURE STUDY

With this dissertation. I hope to continue the expansion of paleoentomology, a

rather young sub-discipline in paleontology. Through my study on insect taphonomy

(appendix A) I emphasize the need for researchers to be more critical in their examinations of fossil insect assemblages. With the studies that I have conducted on modem (appendix B) and fossil herbivory (appendix C) I have shown that there is the opportunity to test ecological and evolutionary hypotheses with the diverse and abundant fossil record of North America. In this concluding chapter 1 highlight some exciting areas of research that are a natural next step.

INSECT TAPHONOMY

Paleoecological and evolutionary studies that utilize the diversity and abundance of fossil insect taxa require a better understanding of the taphonomic biases that affect the insect fossil record. Why is insect taphonomy so important? Consider the work of

Labandeira and Sepkoski (1993), who constructed an insect diversity' curve that showed the geologic history of insect family-level diversity. While this was an important work in paleoentomology, much of the character of this curve is likely to be defined by sampling and taphonomic biases and not by true diversity. Even the authors mention that much of the variation in their curve may be attributable to "an uneven temporal distribution of rich insect-bearing deposits". Peaks in diversity, like the ones seen in the

Jurassic and in the mid-Tertiary are ascribed, by them, to deposits with exceptional preservation. 26

How could taphonomic biases affect the results of Labandiera and Sepkoski?

Consider amber deposits, which capture very diverse insect assemblages. Although there are biases towards the preservation of smaller insects and insects that are found in the leaf litter and in the bark of trees, amber resins are also known to be an attractant to many insect groups. As a result, a great diversity of insects become trapped in amber

(Henwood 1993a, b and Poinar 1992). How can one then compare the diversity of insects in amber to the diversity of insects in tar — which preserves a greater diversity and abundance of necrophagous insects and aquatic groups, or to lake deposits — which are dominated by small, ground-dwelling herbivores and necrophages and are under- represented in aquatics and wood-inhabiting groups (see appendix A)? Such taphonomically based variation suggests that the insect diversity curve of Labandeira and Sepkoski (1993) may not represent true change in diversity over time.

In this dissertation. 1 examined the effects of taphonomic bias on the accumulation and preservation of insects (appendix A) by comparing the living beetle assemblage of a playa-lake environment to its accumulating death assemblage. I found that only fifty-six percent of the live-collected beetle families and twenty-eight percent of the live-collected beetle genera were preserved in the sediment and the relative abundances of beetle genera in the live and dead assemblages were significantly different. Necrophagous beetles were over-represented in the playa sediments, as were ground-dwelling insects and those groups that were smaller and more robust. In contrast, wood-inhabiting groups and aquatics were under-represented in the death assemblage. 27

This study showed the importance of feeding ecology and morphology on beetle preservation in one ephemeral lake deposit. It is important to expand this type of research to include other lake deposits and other insect taxa. Of greatest priority is the study of taphonomic biases in narrow and deep lake environments - much like the

Florissant and Creede localities of Colorado and the Stewart Valley dep>osits of Nevada.

These deposits are distinctive, as they preserve a variety of rare and delicate species, such as specimens of Lepidoptera (butterflies and moths) and Arachnids (spiders). Not only should actualistic studies be conducted on deep lakes, but a closer examination of fossil lake deposits should be undertaken as well. For example, such studies may examine whether the remains of fossil insects are found primarily in near-shore or offshore environments, the preservationai condition of the insects, and whether certain groups are more abundant than others.

Not only should the preservationai biases that affect insect fossilization is a variety of envirormients be examined, but the biases that affect different groups of insects should also be studied. The Coleoptera, Diptera, Hymenoptera and Orthoptera are also very common in fossil lake deposits and yet very little is known about the factors affect their accumulation and preservation. How does their ecology affect their potential to come in contact with lake environments? Are their softer body-parts more susceptible to biological, chemical and physical degradation? Because many members of these groups are exceptional fliers, are they more likely to be found in offshore environments? 28

Insect taphonomy is poorly understood. A growing number of reseeirchers are

becoming interested in using the insect fossil record to examine diversity trends through

time, to conduct paleoecological studies and to try and interpret changes in climate. If

there are strong biases against the preservation of an insect group, in some depositional

environments, then fossil data cannot be treated at face value.

HERBIVORY IN MODERN FORESTS

In my second paper (appendix B), I studied the patterns and levels of herbivory in two modem tropical forests. Although there are many similar studies on modem herbivory in both temperate and tropical forests, none had been conducted in a manner that allowed for direct comparison with pattems of herbivory preserved in the fossil record. In my study, I collected herbivory data from the leaf litter, which provides a sample dominated by canopy leaves. This is important because canopy leaves also dominate fossil leaf assemblages. I found that the percentage of leaf area removed by insect herbivores did not vary significantly within a two-year period, but did vary significantly between the two forests. Furthermore, the percentage of leaf area removed from the samples (4-11%) was within the range of leaf area reported removed in other studies of neotropical rainforests. These other studies used discrete sampling techniques.

Two areas of study should be pursued from this work. First. I think it is important to examine the pattems of herbivory in other forest types, especially those that may serve as close modem analogs to fossil floras. Data that 1 have collected from 29

both temperate and tropical regions (appendix D) show that there can be a great amount

of variation in the levels and patterns of herbivore damage among different forest types.

Understanding modem variation in herbivory may help to understand the patterns seen

in the fossil record. For example, does a change in feeding guild structure correspond

to a change in climate? Do plant diversity and insect diversity affect overall levels of

herbivory? These questions can be examined through more thorough studies of modem ecosystems.

In addition to conducting studies of modem herbivory that use bulk-sampling

techniques, there is also a need to study modem herbivore damage on specific plant

groups. This would allow for an examination of herbivory within clades and provide a framework for examining changes in damage levels and pattems during the geologic lifetime of a plant host. This approach could shed light on the role that interactions between plants and insects have in affecting the diversification of plants and insects.

For example, if increasing diversity of damage types on a plant host through time corresponds with an increase in the diversity of that plant group, then interactions could be a driving force in plant and insect evolution. In contrast, if plants diversify and insects begin feeding on those groups later in geologic time, the interactions themselves may not be cmcial to the diversification of plants and instead, insects may simply be radiating onto an already diverse food source. PLANT-INSECT INTERACTIONS IN THE FOSSIL RECORD

In my last paper (appendix C) I examined the interactions between insects and

plants during the Eocene - Oligocene climatic cooling and compared the interactions to

patterns of damage seen in modem forests. This allowed me to ask several questions;

First, were plants attacked by insect herbivores with the same level of intensity as they are today? Second, have the associations between particular insects and particular

plants changed over time? Finally, does a dramatic change in climate affect the associations between plants and insects?

I found large differences in the feeding intensity between the fossil and modern

floras. Damage levels were significantly lower (23 - 34% of leaves were damaged and

1.4 - 2.5% of leaf area removed) in the fossil assemblages than in the modem (72 — 90% of leaves were damaged and 4 - 10% of leaf area was removed). I also found evidence of specialized associations that were maintained throughout the Eocene (for example, distinct hole-feeding damage on both Cardiospermum and Syzygiodes) and evidence of fossil associations that are still in existence today (stereotyped aphid galls on several plant genera). In addition. I found that decreasing levels of herbivore damage corresponded with a decrease in temperature during the Eocene-Oligocene cooling trend.

Future studies of plant-insect interactions in the fossil record will be much improved by the pursuit of three main areas of research, these include; (1) taphonomic studies, (2) studies of evolutionary change in plant-insect interactions, (3) further examinations of the affect of climate change on levels of herbivory. 31

First and foremost is the need for a closer look at differential preservation and

differential herbivory. Ferguson's (1985) work on plant taphonomy showed that

damaged leaves sink faster than undamaged leaves, suggesting that insect-damaged

leaves might have a greater p>otential to become preserved than their unblemished

counterparts. However, herbivore damage appears to be universally low in the fossil

record, not just in the Eocene, but also in the Cretaceous (Labandeira et al. 1995 and

Labandeira unpublished data) and the Permian (Beck and Labandeira 1998). This

suggests that other taphonomic factors that may affect the preservation of insect-

damaged leaves. Leaves with specific types of damage (like leaf-mining) may be less

likely to become preserved as fossils. Perhaps damaged leaves do not travel as far from

the source tree and are therefore less likely to come in contact with a lake. Or insect-

damaged leaves may be more susceptible to biotic decay. Actualistic studies that

compare the preservation potential of leaves with each of the different types of insect

damage and leaves without damage will identify the taphonomic filters that may be

contributing to lower herbivory levels in the fossil record.

Differential herbivory is also a concern, and may also contribute to the

universally low levels of herbivory in the fossil record. Fossil leaf assemblages tend to

have an over-representation of leaves that are 63-67% smaller than leaves from the

original source forest (Greenwood 1992, Roth Jind Dilcher 1978). If insects feed less on

smaller leaves than on larger leaves, then herbivory levels will appear lower in fossil

floras than in modem floras. It is important to examine the relationship between leaf size and damage levels. Collecting leaf litter samples that include both small and large 32

leaves would make it possible to compare damage levels on leaves from different size

classes and determine if the size of leaves in an assemblage affects the level of

herbivory.

Although there are taphonomic issues that still need to be dealt with. I think that

one of the most exciting results of this dissertation is that hypotheses about the

evolutionary ecology of plants and insects can be tested through a careful examination

of the fossil record. It is possible to examine the timing of the diversification of plant

groups and the timing of when insects begin feeding on those plants. Does

diversification and the inception of feeding occur at the same time, or is there a time lag

between the two? The outstanding record of Cenozoic plants and insects in North

America provides an oppnirtunity to look at plant-insect interactions on great temporal

and spatial scales.

The second research area that should be pursued is the direct examination of

plant-insect interactions through time. There are two main approaches that I think

should be taken when examining plant-insect interactions through time. First, there is

the community-based approach, which consists of an examination of all the leaves and

their insect damage in a fossil flora. There are two main benefits to using this approach;

(1) the fossil data are comparable to the modem - as the majority of studies on modem

herbivory have also used a community-approach and collected bulk-samples of leaves from modem tropical and temperate forests, and (2) the pattems of diversification of generalist and specialist feeding groups can be examined. Studying the diversification of generalist and specialist feeding groups could be useful in trying to understand the role coevolution may have played in the diversification of plants and insects. If

coevolution is important, then the diversity of specialist feeding types would increase

through time. However, an increase could be explained by other processes as well. For example, insects could have been evolving a range of new specialized feeding strategies

for feeding on an already established assemblage of plants.

In addition to the community-based approach, the taxon-specific approach can

be productive. The benefit of this approach is the ability to look at a specific plant group through time. What was the plant species' distribution and when in geological time do we see an increase in diversity? What is the overall rate of insect damage on these species through time and what are the specific insect attackers? When do the insects begin their associations with the plant? There are several groups that would be ideal for this type of study, for example the Salicaceae (Poplars and Willows) are very well studied in the modem and they have an excellent fossil record in North America.

The Fagaceae (Oaks) and the Ulmaceae (Elms) would also be excellent groups to examine for similar reasons.

The third area of research involves an examination of the role of climate in the evolution of plant-insect interactions. The Tertiary fossil record provides an ideal setting for examining the evolution of plant-insect interactions and for evaluating how climate change affects ecosystems at basic trophic levels. This is a time period with an excellent record of plants and insects, and it provides a window into the early evolution of temp)erate ecosystems. My work and the work of colleagues have focused on earlier portions of the Cenozoic. and we have shown that herbivory appears to be strongly 34 influenced by temperature. When temperature increased in the late Paleocene, so did herbivory levels (Wilf and Labandeira 1999). When temperatures declined in the middle to late Eocene, levels of herbivory also declined (see appendix C). However, this pattern has been demonstrated using only four sites and the need to examine more sites from this time interval is crucial. Are the patterns limited to these particular localities, or are the patterns consistent both spatially and temporally in the Cenozoic of

North America? A large number of Cenozoic lake deposits in North America preserve both plants and insects. It should be possible to track the long-term evolution of specific plant and insect groups and closely examine the effects of climate on these associations.

Studying insects in the fossil record is an exciting new sub-discipline within paleontology. This unlikely, but surprisingly abundant group of fossils can provide information on past ecosystems and insights into modem community interactions. The future study of fossil insects and their ecological interactions with plants will require a multi-disciplinary approach, involving geology, entomology, botany and systematics.

A more rigorous examination of taphonomic bias in both insects and plants, coupled with studies of modem ecology can be integrated with detailed paleoecoiogical analysis.

The result can be a far greater understanding of how life evolves than could be provided by the geological sciences or the biological science alone. APPENDIX A 36

Beetle taphonomy in a recent ephemeral lake, southeastern Arizona.

(Published in PALAIOS, 15(2): 152-160, April 2000)

Dena M. Smith

Department of Geosciences. University of Arizona, Tucson, AZ 85721

ABSTRACT

A comparison of the taxonomic composition, relative abundance, diet, feeding habitat, and size of live and dead insects was conducted in Willcox Playa, an ephemeral lake in southeastem Arizona. Death assemblages of beetles, the most abundant group of insects in this study, are preserved in the shallow, subsurface sediments along the shoreline of the lake. Fifty-six percent of the living beetle families and twenty-eight percent of the living beetle genera were found in the sediments; one-hundred percent of the families and ninety-one percent of the genera found dead were also present in the live fauna. Relative abundances of beetle families in the live assemblage are significantly different than relative abundances in the death assemblage. Among diet and habitat groups, necrophagous and ground-dwelling beetles are over-represented in the death assemblage, while wood-inhabitants and aquatics are under-represented. The death assemblage contains a greater proportion of smaller, more robust beetles. Such biases also may occur in fossil assemblages and should be considered in paleoecological and paleoenvironmental reconstructions. 37

INTRODUCTION

Fossil insects are valuable sources of information on ancient environments

(Coope, 1970), climates (Elias, 1991), evolution (Labandeira and Sepkoski, 1993), and species interactions (Labandeira et al., 1994). But do insect death assemblages accurately reflect the composition and relative abundance of the insect life assemblage?

Despite the diversity and abundance of insects in Tertiary lake deposits (e.g.. Middle

Eocene Green River and Late Eocene Florissant Fossil Beds), little is actually known about insect taphonomy in lacustrine environments.

Understanding the accumulation and preservation of insects is important if we hope to utilize these fossils in paleobiological studies. Although insects often are thought of as being fragile, the chitinous portions of the insect exoskeleton can be quite robust. The oldest evidence of intact chitin was found in beetle fossils from a 25- million-year-old site in Enspel, Germany (Stankiewicz et al.. 1997a). Pleistocene beetles from California tar deposits have extremely high levels of chitin preservation

(Stankiewicz et al., 1997b), with some specimens exhibiting levels close to what is seen in their modem counterparts (Miller et al., 1993). Even fossil insect specimens that have no original chitin remaining can exhibit exceptional preservation of the cuticle morphology and may have features that are preserved in three dimensions (Duncan et al., 1998; McCobb et al., 1998; Park, 1995). Although insect exoskeletons appear to be fairly resistant to post-mortem decay, it is not clear how well fossil insect assemblages reflect the overall species, trophic and life-habit composition of the original living community. 38

Wilson (1988) examined the taphonomy of insects in lacustrine deposits. He

compared nearshore and offshore insect assemblages in several Tertiary lakes in North

America. In general, insects were found in greater numbers in the nearshore deposits.

However, nearshore specimens had tended to be more fragmented than their offshore

counterparts. Wilson also found that Coleoptera and Trichoptera were more common in

the nearshore deposits, whereas the offshore deposits had greater numbers of Diptera,

Hymenoptera and Heteroptera.

In addition to Wilson's work, some studies have examined insect taphonomy in

other preservational environments. For example, Henwood (1993a, b) and Poinar

(1992) noted several taphonomic biases that effect insect preservation in amber. They

showed that there is a bias towards smaller insects (Henwood. 1993a) and those that

frequent leaf litter (Henwood, 1993b) and the bark of trees (Poinar, 1992). Miller and

Peck (1979) examined the preservation of insects in tar deposits. They found a large

number of insects that were attracted to dead organisms already trapped in the tar and

many aquatic insect groups that were likely to have mistaken the oil for p>ools of water.

Elias (1990) examined the preservation of insects in packrat middens. He found that

beetles are the most abundant insect found in the middens and that the majority were

predators or scavengers, many of which were obligate midden inhabitants.

The study addressed herein utilized an actualistic approach to the study of insect taphonomy. Results provide insight into the processes involved in the accumulation and burial of insects with regard to their taxonomic composition, relative abundance, diet, feeding habitat, and size in life and death assemblages. The comparison provides 39

information about the biases likely to occur in fossil beetle assemblages that

accumulated in similar settings.

STUDY AREA

Willcox Playa, an ephemeral lake in southeastern Arizona (Fig. 1), is an ideal

setting for examining the taphonomic processes that effect the preservation of insect

assemblages in the lacustrine fossil record. Willcox Playa is a remnant of Pleistocene

Lake Cochise, which dates from 10,000 years ago (Pine, 1963). The lake originally

covered an area of approximately 130 km^ and may have had water depths of 12-15m

(Pipken, 1964). Today the playa lake is confined to a 300m2 area (Long, 1966).

Modem lake levels fluctuate in response to winter (Dec. - Feb.) and summer (July -

Sept.) rains, with the lake level being the highest (generally less than 2m) following the

summer rains (Pine, 1963).

Willcox Playa is at an elevation of approximately 1260 m and is bounded by

mountain ranges to the east and west. Only two drainages from the surrounding

mountains currently reach the playa and no material greater than sand size is carried

into the lake (Pine, 1963). Lake sediments consist of fine sands, silts, and clays and a

surflcial crust of evaporite minerals (Long, 1966). There are eolian sand dunes on the

northwest - west margins of the playa. These dunes begin approximately 100m from

the shoreline of the lake. The local vegetation is desert grassland and scrub (Martin,

1963). The remains of the diverse insect fauna accumulate along the lake's shoreline, and the environmental setting may be comparable to many ancient lake deposits. 40

MATERIALS AND METHODS

Live Collecting

The summer monsoon season started approximately August 15, 1996 and the living beetle fauna was collected during the summer monsoons, from August 26 through October 13,1996. This time period was chosen because this is when the insect community is most abundant and diverse (University of Arizona Entomology Research

Collections Database) and because this is also the time period in which the greatest amount of sediment is deposited into the playa (some sediment is also deposited during the winter rains). Beetles were collected four times per day; at dawn, mid-day, dusk, and at night. Day collecting was done with the use of sweep nets (for insects found on shrubs, flowers, and grasses), aerial nets (for insects in flight), aquatic nets (for those in the water column and on the bottom of the lake), beat sheets (for those attached to shrubs and sturdy bushes), and ground-scanning methods (to collect insects on the ground, in cracks, or under debris). In addition to these methods, six pitfall traps were set between the lake margin and the dunes. However, all of the traps quickly filled with sediment, making them unusable. Night collecting included the use of a blacklight, sweep nets, and ground-scaiming methods. The blacklight was set up in an open area between the dunes and the lake.

Collections were made from three different microhabitats ~ within the lake, along the lake's shoreline, and in the dunes surrounding the playa.

Beetles were counted and sorted according to genus and stored in vials of isopropanol. Identifications were made using taxonomic keys (Amett et al., 1980; 41

Borror et al., 1989) and the University of Arizona Insect Collection. The beetle feeding-habitat and diet designations were based on the Smithsonian Institution's ETE arthropod paleobiology database (although some of the categories may need to be revised). All beetle specimens were identifiable to family and those that were not easily identifiable to genus were given morphogenus designations. These specimens were determined to morphogenus based on differences in specimen size and color patterns.

The morphotaxon approach is commonly used by paleobiologists and neontologist alike and is a method that is usually considered to be a reasonable estimate of true taxonomic diversity (Oliver and Beattie 1996). However, utilizing this approach increases the risk of over-splitting, as morphological diversity within many insect genera can be great.

It is possible that some of the taxa in this study are over-split, especially the

Carabidae. for which there are seven identified genera and 19 morphogenera designated in the group. If this family is over-split, it will not afifect the comparison of the relative abundance of live and dead beetles, as these data were analyzed at the family level. The analysis of size should also not be affected because the results are based on direct measurements fi-om the specimens. However, over-splitting has the potential to affect the diet and feeding-habitat data, especially if some of the Carabid specimens belong to herbaceous or seed-feeding genera. Fortunately, the Carabid morphogenera in this study comprise only a small proportion of the total number of individuals in the live

(6%) and dead (4%) assemblages and are not likely to alter the overall results of this study. 42

Dead Collecting

Eight 1 -by-3-m quadrats of sediment were collected from the shoreline of the lake. These quadrats were arranged in L-shaped pairs, with one quadrat aligned parallel to the shoreline and the other aligned perpendicular to the shoreline. Each pair of quadrats were spaced 145m apart along the northern (downwind) shoreline of the lake. Only the upper 2-4 cm of surface sediment was collected from the quadrats. This was the sediment that was being deposited into the playa during the summer rainy season. Underlying this muddy active layer was an indurated layer of mud and evaporite minerals. While 2-4 cm may seem like a large amount of sediment to be deposited in a short time period, it is similar to the sedimentation rates observed in other ephemeral saline lakes in the southwest United States (Roberts and Spencer, 1998). The

Willcox Playa sediment samples were collected on October 13. 1996 (same day as the last collection of the living assemblage) at the end of the monsoon season when the lake was beginning to dry. Therefore, the sediment samples only contain insect specimens accumulated during the three-month summer rainy season.

Results from a preliminary study showed that the most abundant insect orders preserved in a 9kg sample of surface sediment were Hymenoptera, the majority of which were ants, and Coleoptera (Table 1, Fig. 2). Soft-bodied insects (those that have cuticles that are not very heavily sclerotized), such as Diptera (flies) and Lepidoptera

(butterflies and moths), are apparently rapidly disarticulated and decomposed, and/or are removed from the shoreline area by ants and necrophagous beetles. Although these 43 soft-bodied groups are a major component of the living assemblage, they are very rare in the death assemblage.

Beetles are the most abundant group at Willcox Playa (University of Arizona

Entomology Research Collections Database) and in the fossil record in general

(Labandeira, 1994); hence they are the focus of this study. Beetles were extracted from the sediment samples by a process outlined by Elias (1994). Each sample consisted of approximately 5.5 kg of sediment that was washed and then wet-sieved to remove clays.

Kerosene was then added and mixed with the remaining sediment and insect parts and allowed to sit for nearly 30 minutes. Then water was stirred into the sediment-kerosene mixture. The kerosene and water layers were allowed to separate and the insect remains were collected off the top layer. This procedure was repeated several times. All complete beetle specimens and/or pairs of elytra (which included unpaired and fragmented elytra that had no matching counterpart) were sorted according to genus and counted. Isolated heads and identifiable thoracic segments were also included, unless there were disartculated elytra from the same individual preserved (this was done to avoid counting disarticulated parts of individuals as multiple individuals). Storage and identification of specimens followed the same protocols used for live-collected specimens.

Statistical Analysis

All statistical analyses were done with the use of JMP IN 3.2.1 (SAS Institute,

1996); data were analyzed using chi-square analysis of contingency tables. In the comparison of dead and live beetle families, those families that had frequencies less

than 5 individuals in the dead and/or live samples were not included in the statistical

analysis. In general, chi-square tests do not work well with expected frequencies that

are less than five and Zar (1996) suggested combining neighboring rows/columns or

removing rows/columns that have low frequencies. Even with the removal of the

families with low frequencies, 84% of the individuals (N=4182) in this study are still

included in the statistical analysis.

RESULTS AND DISCUSSION

Taxonomic Composition

A total of 3943 individuals representing 27 families and 102 genera of

Coleoptera were found in the live assemblage; the sediment samples yielded 239

individuals from 15 families and 32 genera. One-hundred percent of the beetle families

and ninety-one percent, or 30, of the beetle genera found in the dead assemblage also

were found in the live assemblage. Fifty-six percent of the living families and twenty-

eight percent of the living genera were also found in the dead assemblage (Table 2).

The University of Arizona's Entomology Research Collections Database lists 50

families and 395 genera of beetles from the Willcox Playa region. Therefore, only 26%

of the beetle genera known to occur at the playa were captured in the three months of

live collections. It is important to note that the Collections Database is based on collections made during all seasons of the year within the past 50 years. There can be a

large amount of annual variation in insect diversity, as some groups may only emerge once every few years, or only when environmental conditions are right. Live and dead collections made in the fiitvire are likely to yield different insect community compositions. Therefore, the faunal list represents a sample of Willcox Playa insects that has been time- averaged to a far greater degree than the collected live or death assemblage. The two beetle taxa, Canihon (2 specimens) and a Cassidine Leaf Beetle

(1 specimen), in the death assemblage that were not captured in the living are recorded in the Entomology Research Collections Database. It is likely that these species were active during the first week of the monsoons and therefore, were preserved in the sediments but not live collected. This illustrates an important point. Insects can emerge and become active for very short intervals of time. If a short-term emergence occurred between live-sampling periods, then these insects may appear in the sediment sample, but not in the collection of living insects.

There is a common perception that insects are rarely preserved as fossils (e.g..

Carpenter and Bumham. 1985). That 28% of the living genera preserved in the playa sediments is perhaps better than most might expect. In comparison, molluscan death assemblages tend to contain 75-98% of the living species (Kidwell and Bosence. 1991)

- the fidelity is probably even higher at the generic level. Bumham et al. (1992) showed that about 70% of the living plant species (again, the fidelity is probably even higher at the generic level) were found preserved in the forest litter. Although the preservation of living insect species in the death assemblage is higher than most might have expected, it is still considerably lower than values found for other groups. 46

Process of Accumulation

Insects that die on the surface of the lake are rafted to the lake margins by wind.

There they accumulate and become covered with sediment. Intact insect specimens are

found at depths of at least 18 cm in the sediment of Willcox Playa. Ground-dwelling

insects that frequent the shoreline often become trapped in the muddy sediments and are

buried and preserved as well. In addition to insects, other terrestrial arthropods such as

solfligids, vinegaroons (Fig. 3a), and spiders, are also preserved. Branchiopods also are

encountered along the shoreline, and occasionally the mummified remains of small

frogs (Fig. 3b) are found in the sediments as well. Accumulations of dead insects are

greatest along the northern shore of the playa, probably because the monsoon winds

generally blow in a northerly direction and push floating insect remains to this side of

the lake.

Preservation

A preliminary study of insect preservation showed that insects were found in a variety of conditions (Table 3). In that study, 4384 insect parts were found preserved in a 9kg sediment sample. Disarticulated legs were most common (57%), followed by isolated heads, thoracic segments and beetle elytra (Fig. 4).

In the current study, the majority of intact beetle specimens have complete to partially decayed abdominal tissue and disarticulated legs. The head is often disarticulated from the thorax. The most abundant identifiable beetle remains are disarticulated eljira. These are often in excellent condition, with pristine edges, surface 47

architecture, and color patterns (Fig. 5). Such preservation makes it possible to identify

specimens to the genus and often the species level. Brightly colored insects often

undergo some degree of fading. In general, insects that are not covered entirely by

sediment undergo some bleaching of their exposed parts.

Relative Abundance

The proportions of families of live-collected beetles are significantly different

(x~ = 1165.88, P< 0.001) from the proportions of families of dead beetles (Table 2). For

example, while the family Anthicidae comprises the most abundant group (36%) in the

dead sample, it makes up only 6% of the live collected assemblage. The

Chrysomelidae are the most abundant (51%) family in the live-collected assemblage

(Fig. 6). but makes up only 7% of the dead sample.

At the genus level, live-dead proportions also differ. Not only are 70 genera

missing from the shallow subsurface, but their relative abundances are mis-matched as

well. The most abundant genus in the death assemblage is Anthonomus

(Curculionidae), which contributes 29.3% of the individuals in the dead but only 0.2% of the individuals in the living. The genus Diabrotica (Chrysomelidae) is most abundant in the live-collected assemblage, representing 46% of the live individuals, but only 4.2% of individuals in the dead assemblage. Differences in the diversity and abundance of insects in the live and dead collections may be partially attributable to the sampling methods utilized in this study. For example, blacklighting is a technique that attracts night-active insects to a collecting sheet where the collector then removes them. 48

This method is highly effective, but may bias the sample if extra-local taxa are attracted

to the collecting sheet as well. This would add taxa to the live-collected sample that

would not be captured in the sediment sample. Although this may account for some of

the discrepancies between the proportions of dead and live-collected beetle genera at

Willcox Playa, the diet, feeding habitat, and size of the beetles (Appendix) may explain

these differences as well.

The difference in relative abundance between the live-collected and dead

assemblages in this study sharply contrasts with results on the taphonomy of plants in

lacustrine environments. For example, Spicer's (1981) study of Silwood Lake

demonstrated a good correlation between plant species abundance in the lake sediments

and the abundance of plants in the surrounding lakeside plant community. Why such

dramatic differences in plant and insect taphonomy? For one, trees are stationary and

those growing close to a lake margin have a high likelihood of their leaves making it to

the lake environment. Insects on the other hand, are highly mobile and are constantly

moving in and out of the depositional environment. In addition, a single tree can shed

large numbers of leaves, each year for many years. Although all the leaves do not

become fossilized, some of them will make it to the sediment-water interface and may be preserved, recording the presence of the parent tree. Insects have far fewer identifiable parts (as opposed to multiple leaves or pollen from an individual plant) that can become incorporated into the fossil record. 49

Feeding Habitat

Feeding habitat is defined herein as the area where the adult insect feeds and, therefore, spends the majority of its time. Five major feeding habitats are recognized in this study — ground, plants (in general), dung, flowers (specifically), and water (Table

4). Again, the dead and live-collected populations are very different (Fig. 7a, =

75.07. P<0.001). Ground-dwellers were the most abundant (43%) in the dead sample, closely followed by plant dwellers (41%). Plant dwellers are the most abundant (56%) and ground dwellers are the second most abundant group (19%) among the living dataset. Wood-inhabiting beetles also were present in the life assemblage, but in very low numbers (8 individuals); they were absent in the death assemblage. There are very few water inhabitants in either of the two assemblages (1% of the dead and 3% of the live).

The high proportion of plant-dwellers in the death assemblage is expected because this is the most extensive feeding habitat in the Willcox Playa area. It also makes sense that ground-dwellers should be well represented in the sediments. Not only is this the second most extensive feeding habitat in the living assemblage, but the location where entrapment and burial takes place is also on the ground.

The low proportion of aquatic inhabitants in the death assemblage may be due to low proportions of these beetles in the live assemblage. However, the under- representation of aquatic groups is fairly common in fossil lacustrine deposits as well

(Wilson, 1988). Therefore, in addition to overall low numbers in the original living 50

population, there may be another reason why there are low numbers of aquatic insects

in the death assembls^e. Insects that die on the surface of the water often get raited to

the shoreline where they are buried. Insects that die below the water's surface are more

susceptible to biotic decomposition, due to other insects, fungal and bacterial decay, or

branchiopod and other scavengers.

Diet

The proportions of individuals in the living and dead sample were compared

relative to diet (Table 5). Five main diet types occur in the beetle assemblages (Fig. 7b)

~ herbivores, necrophages, predators, dung feeders, and scavengers (non-animal tissue).

In addition, two minor feeding groups also occurred — f\mgivores and wood-feeders.

Overall, proportions in the dead population differ significantly from those in the live population (x~ = 229.514, P<0.001). Herbivorous beetles were the most abundant group in both the living assemblage (60%) and in the dead (42%). However, while necrophagous insects were the second most abundant group in the dead sample (35%), they were only the fourth most abundant group in the live collection (7%). A small number of fimgivores (7 individuals) and wood-feeders (2 individuals) were found in the live-collected assemblage but were not present in the subfossil assemblage.

The large proportion of necrophagous beetles in the death assemblage is explained easily. These beetles feed on decaying organisms that have accumulated along the shoreline; they themselves become trapped. This is analogous to the large numbers of vultures that were trapped in the La Brea tar pits while attempting to feed on 51

the organisms that already were trapjied (Stock and Harris, 1992). The fungivorous

groups foimd in the living assemblage feed predominantly on fiingus found in decaying

wood (Borror et al., 1989). Because fimgivores and wood-feeders spend the majority of

their life within wood, they are less likely to encounter the lake and its shoreline,

explaining the absence of these genera from the subfossil assemblage.

Size

TTie average size of each genus was measured either from the sfjecimens or

collected from the entomology literature. Beetles ranged from 2-40 mm in the live

assemblage and 2-25 mm in the death assemblage (Table 6). The distribution of size

classes (Fig. 7c) in the death assemblage is very different from that found in the live (x^

= 329.28, P<0.001). A large number (62%) of beetles that are 2.5 mm in length occur

in the dead sample. However, this size category accounts for only 7% of the individuals

found in the live assemblage. Beetles that are 5 mm in length are most abundant (49%)

in the living assemblage, whereas only 7% of the dead beetles are this size.

Although all of the size categories of the living assemblage are represented in

the death assemblage, smaller taxa are more abundant in the sediment samples. The

reason for this bias may be one or more of the following: (1) smaller insects are less

able to escape from the mud, (2) small insects are less susceptible to disarticulation, (3)

the live-collected sample may not have captured an accurate representation of all the size classes found at Willcox Playa. 52

Small, ground-dwelling insects that become trapped in the mud may not be able to free themselves from this viscous medium. The same phenomenon has been documented in insects that become trapped in amber. Although many different-sized organisms become preserved in amber, there is an over-representation of smaller groups, which were presumably unable to free themselves from the sticky resin

(Henwood, 1993a). In addition, the small beetles (ground-dwellers and those that were rafted on the lake surface) preserved in Willcox Playa sediments are fairly robust. They are heavily sclerotized relative to their internal soft parts and have a higher surface area

(hard-part) to volume (soft part) ratio, making them less likely to become disarticulated.

It is also likely that smaller beetles will become buried more rapidly, also preventing disarticulation and enhancing their preservation potential.

Finally, the under-representation of smaller sized beetles in the live-collected sample could have been an artifact of collector bias. It is fairly common for collectors to miss the smaller members of an assemblage and some species accumulation curves for beetles demonstrate a lack of insects from smaller size classes (in contrast see Elton

1973; Janzen 1973; Janzen and Schoener 1968; Morse et al. 1988). Although I tried to be cautious and not introduce this type of bias into my sampling, it may have been unavoidable to some degree. If this is the case, then the over-representation of smaller beetles in the dead assemblage may not be as extreme as it first appeared, or may be non-existent. 53

CONCLUSIONS

Live-dead comparisons provide an opportunity to examine the factors that may

effect the burial and preservation of insect remains. This study has shown that there is

low family-level fidelity between the live-collected and dead beetle assemblages of

Willcox Playa. Although finding 28% of the living fauna in the sediment samples was

much higher than expected for insect groups in general, this value is lower than in groups such as molluscs and plants. In addition, there are large differences between the

relative abundance of individual families in the live and dead beetle faunas.

These biases may result firom the diet, feeding habitat, and size of the insects.

Although the most abundant diet (herbivory) in the living assemblage was also the most

abundant among the dead, one should expect an over-representation of necrophagous insects in nearshore fossil insect assemblages. There is also likely to be a bias towards ground-dwelling beetle groups. One can also expect an under-representation of wood- inhabitants and aquatics in lacustrine fossil assemblages, as there was a virtual absence of these groups in both the live and dead communities of Willcox Playa. There is also bias towards beetles that are smaller (2.5 mm) and more robust.

Recognizing these biases in lacustrine fossil deposits may improve interpretations of ancient biotas. For example, much of the Middle Eocene Green River

Formation is thought to have been a playa-like environment with ephemeral shorelines and shallow waters (Eugster and Hardie, 1975; Lundell and Surdam, 1975). Although there are currently no reliable abundance data for insects collected from the Green

River, a few patterns in diversity seem to suggest that these biases may play an important role. For example, 47% of the insects collected from the Green River

Formation were Coleoptera and the majority (58%) of beetle taxa that have been collected are small in size, within the 0-4.5nim size class (Smith, unpublished data).

Collections of fossil insects that include information on relative abundance will allow for a more complete comparison between playa-lake fossil assemblages and the predictions of this actualistic study. In addition to comparative work on fossil biotas, conducting fiirther actualistic studies that examine preservation in other types of lake environments and focus on other insect orders would enhance greatly our understanding of fossil insect preservation.

ACKNOWLEDGMENTS

Financial support for this work was provided by a National Science Foundation

Predoctoral Fellowship, a grant from the University of Arizona's Research Training

Grant for the Analysis of Biological Diversification, and a Chevron Summer Research grant. Cesar Nufio, Jonathan Marcot, and Agata Kowalewska provided valuable assistance with field collecting, and Aspen Garry and Cesar Nufio provided assistance in the lab. Carl Olsen provided assistance with beetle identifications and access to the

University of Arizona's Entomology Research Collections. I greatly appreciate the statistical advice 1 received from Dr. Sean Connolly and Dr. R. J. Frye, and comments from Dr. Peter Allison, Dr. Karl Flessa, Cesar Nufio, and two anonymous reviewers on the early drafts of this manuscript. This is publication #33 of Centro de Estudias de

Almejas Muertas - Division de los Insectos. 55

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TABLE 1. Results from a preliminary study of insect preservation at Willcox Playa.

The number and percent abundance of the insect orders found in the 9kg sediment sample are listed. Although Hymenonptera is the most abundant order, 98% of the indivduals in this group were from one species of ant.

Insect Orders Number Percent

Coleoptera 175 30 Diptera 8 1 Heteroptera 10 2 Homoptera 21 4 Hymenoptera 364 62 Odonata 1 0 Other Arthropods 4 1 TABLE 2. Beetle families from Willcox Playa and the number of live and dead individuals of each family. Families with an asterisk were not included in the statistical analysis. These families had frequencies less than 5 individuals in the dead and/or live samples. Large numbers of such low frequecies are thought to be problematic in chi- square tests. Combining rows or removing rows with low frequencies is a conmion way to deal with this type of problem (Zar, 1996).

Dead Live Family Number Percent Number Percent

Alleculidae* 0 0 2 0 Anthicidae 84 35 232 6 Anthribidae* 0 0 1 0 Bruchidae 13 5 70 2 Cantharidae* 0 0 20 1 Carabidae 9 4 252 6 Cerambycidae* 0 0 1 0 Chrysomelidae 16 7 2010 51 Cicindelidae* 4 2 117 J Cleridae * 2 1 33 1 Coccinelidae 9 4 60 2 Curculionidae 70 29 18 .5 Dytiscidae* 1 0 15 0 Elateridae* 0 0 8 0 Heteroceridae* 0 0 40 I Hydrophilidae* 1 0 56 1 Meloidae* 0 0 14 0 Melyridae * 4 2 43 0 Mordellidae* 0 0 76 2 Mycetophagidae* 0 0 5 0 Nitidulidae* 0 0 1 0 Oedemeridae * 1 0 70 2 Scarabaeidae 19 8 652 17 Scolytidae* 0 0 1 0 Silphidae* 0 0 1 0 Staphylinidae* 2 1 127 3 Tenebrionidae* 4 2 18 .5 TABLE 3. Results from a preliminary study of insect preservation at Willcox Playa.

The number and percent abundance of articulated and disarticulated insect body parts found in the 9kg sediment sample are listed. Seventy-five percent of the disarticulated heads and 79% of the thoracic segments were from ants.

Body Parts Number Percent

3/4 - Entire Body 78 2 Head 638 14 Thoracic Segments 356 8 Abdomen 291 7 Wings-membranous 12 0 Wings-elytra 338 8 Legs 2494 57 Mandibles 132 J Scutella (hemipteroids) 10 0 Genitalia 4 0 Exuvia 28 1 65

TABLE 4. Number and percentage of individuals in the dead and live datasets in the feeding habitats recognized in this study. Feeding habitat is defined as the area where the adult insect feeds and, therefore, spends the majority of its time.

Dead Live Feeding Habitat Number Percent Number Percent

Ground 103 43 786 20 Plant (general) 97 41 2179 55 Dung 19 8 600 15 Flowers 18 8 295 8

Water 2 - 71 2

In Wood 0 - 12 - or Animals 66

TABLE 5. Number and percentage of individuals in the dead and live datasets in the diet type categories recognized in this study. Both major and minor feeding catergories are included in this table.

Dead Live Diet type Number Percent Number Percent

Herbivores 100 42 2338 60

Necrophages 84 35 276 7 Predators 29 12 520 13 Coprophages 19 8 600 15 Scavengers 7 3 200 5

Wood or Fungi 0 - 9 - 67

TABLE 6. Number and percentage of individuals in the dead and live datasets in the different size categories.

Dead Live Size (mm) Number Percent Number Percent

0-4.9 191 80 1012 26 5-9.9 22 9 2106 53 10- 14.9 18 8 661 17 15- 19.9 7 3 145 4

20 and Up 1 - 19 - 68

Figure 1. Location of Willcox Playa and the siirrounding area. 69

MEXICO

o Xfts. Galiu

Tucson Willcox BIVO'X - WILLC Benson PLAYA Drago^^yfts. Chirica^^mMts.

Nobles ARIZONA MEXICO lorso' w 70

Figure 2. Results of a preliminary study on insect preservation at Willcox Playa. This histogram shows the relative abundance of insect orders recovered from a 9kg sample of surface sediment. A total of 583 individuals were recovered from the sample. Abundance(%) K) CJ iw Ol 0> o o o o o Hymenoptera Coleoptera Homoptera Heteroptera Diptera Odonata Other Arthropods Figure 3. Examples of recent preservation. 2A) A vinegaroon (Uropygi) from the shoreline sediments of Willcox Playa. B) A mummified frog, also from the shoreline sediments of Willcox Playa. Scale bar: 5 mm. 73 74

Figure 4. Results of a preliminary study on insect preservation at Willcox Playa. This histogram shows the relative abundance of intact insects and disarticulated insect body parts recovered from a 9kg sample of surface sediment. A total of 4381 body parts were recovered from the sample. Abundance(%) lo <4 ^ (n o> o o o o o —J I I I I 3/4 - Entire Body _ Heads g Thoracic Segements o Abdomens Q Wings - membranous Q. Wings-elytra * Legs ^ Mandibles ^ Scutella Genitalia Exuvia J 76

Figure 5. Elytra from a predaceous diving beetle {Eretes) extracted from one of the sediment samples. Scale bar: 3.25 mm.

78

Figure 6. Histogram showing the relative abundance of families from the dead sample vs. families from the live-collected sample. Pearson's = 1206.86, P< O.OOl. Abundance(%)

Anthicidae

Bruchidae

Carabidae

Chryaomeiidae WWWSWWSSWM Cicindelidae

Cleridae

Coccinelidae

Curculionidae

Dytiacldae

Hydrophillidae

Melyridae [

Oedemeridae

Scarabaeidae s Staphylinidae

Tenebrionidae s S 80

Figure 7. Histograms showing the relative abundances of the (A) feeding habitats of the dead vs. live beetle assemblages, = 75.07, P< 0.001, (B) diet types of the dead vs. live beetle assemblages, x~ = 229.51, P< 0.001, and (C) size categories of the individuals in the dead vs. live beetle assemblages. x~ = 329.288, P< 0.001. DEAD

UVE

Ground Plants Oung Ftowtrs Watar FMding Habitat 60 -1

m UVE

HarbivorM Nacrophagas Pradators Coprophagas Scavangars OiatTypas

OEAO

UVE

0-4.9 S-9.9 10 - 14.9 15-19.9 20 and UP Siza(min) APPENDIX. Record of all live and dead collected beetle taxa from this study. Taxa are identified to genus or tribe. Feeding habitat and diet is determined using the most appropriate taxonomic level (which can vary between groups quite a bit). The size of all sp)ecimens was measured or collected from the entomology literature.

Family Genus/Tribe Feeding Habitat Diet Size (mm)

Alleculidae Hymenorus Flowers Herb 4 Anthicidae Anthicus Ground Necro 2.5 Anthicidae Jsochyropalpus Ground Necro 2.5 Anthicidae Notoxus Ground Necro 2.5 Anthribidae Genus A Wood Fungi 3 Bruchidae Genus A Flowers Herb 4.5 Bruchidae Genus B Flowers Herb 5 Cantharidae Chaul iognathus Flowers Herb 10.5 Carabidae Bembidion Ground Pred 7 Carabidae Brachinus Ground Pred 14 Carabidae Calosoma Ground Pred 25 Carabidae Chlaenius Ground Pred 15 Carabidae Colliurus Ground Pred 7 Carabidae Harpalus Ground Pre 15 Carabidae Pasimachus Ground Pred 21 Czu^bidae Genus A Ground Pred 10 Carabidae Genus B Ground Pred 8 Carabidae Genus C Ground Pred 8 Carabidae Genus D Ground Pred 10 Carabidae Genus E Ground Pred 8 Carabidae Genus F Ground Pred 8 Carabidae Genus G Ground Pred 9 Carabidae Genus H Ground Pred 7 Carabidae Genus I Ground Pred 5 Carabidae Genus J Ground Pred 5 Carabidae Genus K Ground Pred 4.5 Carabidae Genus L Ground Pred 5 Carabidae Genus M Ground Pred 4 83

APPENDIX - Continued

Family Genus/Tribe Feeding Habitat Diet Size (mm)

Carabidae Genus N Ground Pred 4 Carabidae Genus O Ground Pred 4.5 Carabidae Genus P Ground Pred 3.5 Carabidae Genus O Ground Pred 4.5 Carabidae GenusR Ground Pred 2.5 Carabidae Genus S Ground Pred 3 Cerambycidae Megacyllene Wood Wood 18 Chrysomelidae Alticinae (A) Plant Herb 2.5 Chrysomelidae Alticinae (B) Plant Herb 2 Chrysomelidae Cassidinae Plant Herb 7 Chrysomelidae Chlamisinae Plant Herb 3.5 Chrysomelidae Cryptocephalinae Plant Herb 2 Chrysomelidae Diabrotica Plant Herb 5 Chrysomelidae Galerucinae Plant Herb 3.5 Chrysomelidae Genus A Plant Herb 4 Chrysomelidae Genus B Pljint Herb 6 Cicindelidae Cicindela Ground Pred 15 Cleridae Enoclerus Plant Pred 10 Cleridae Phyllobaenus Plant Pred 4 Coccinellidae Hippodamia Plant Pred 6.5 Coccinellidae Hyperaspis Plant Pred 3 Coccinellidae Olla Plant Pred 5 Coccinellidae Scymnus Plant Pred 2 Coccinellidae Genus A Plant Pred 3 Coccinellidae GenusB Plant Pred 3.5 Coccinellidae GenusC Plant Pred 3 Coccinellidae GenusD Plant Pred 2 Curculionidae Anthonomus Plant Herb 3.5 Curculionidae Genus A Plant Herb 6 Curculionidae GenusB Plant Herb 12 Curculionidae Genus C Plant Herb 14 Dytiscidae Eretes Water Pred 17 Dytiscidae Hygroies Water Pred 4 Dytiscidae Genus A Water Pred 3 84

APPENDIX - Continued

Family Genus/Tribe Feeding Habitat Diet Size (mm)

Elateridae Aeolus Plant Herb 10 Elateridae Genus A Plant Herb 10 Elateridae Genus B Plant Herb 4.5 Heteroceridae Heterocerus Ground Necro 5 Hydrophilidae Berosus Water Scav 5 Hydrophilidae Cercyon Water Scav 2 Hydrophilidae Enochrus Water Scav 3.5 Hydrophilidae Hydrophillus Water Fungi 40 Hydrophilidae Genus A Water Scav 2 Meloidae Genus A Flowers Herb 14 Meloidae Genus B Flowers Herb 10 Meloidae Genus C Flowers Herb 6.5 Meloidae Genus D Flowers Herb 10 Melyridae Collops Flowers Pred 5 Melyridae Genus A Flowers Pred 3 Mordellidae Mordella Flowers Herb 4 Mordellidae Mordellistena Flowers Herb 4 Mycetophagidae Typhaea Wood Fungi 3 Nitidulidae Carpophilus Plant Herb 4 Oedemeridae Genus A Flowers Herb 10 Oedemeridae Genus B Flowers Herb 7.5 Scarabaeidae Aphodius Dung Dung 10 Scarabaeidae Canthon Dung Dung 15 Scarabaeidae Cotinis Plant Herb 25 Scarabaeidae Cyclocephala Plant Herb 15 Scarabaeidae Onthophagus Dung Dung 12 Scarabaeidae Oxygrilius Dung Dung 14 Scarabaeidae Phyllophaga Plant Herb 25 Scarabaeidae Trox Animal Necro 15 Scarabaeidae Genus A Plant Herb 7 Scarabaeidae Genus B Plant Herb 7 Scarabaeidae Genus C Dung Dung 4 Scarabaeidae Genus D Dung Dung 9 Scarabaeidae Genus E Plant Herb 3 Scolytidae Scolytus Wood Wood 2 Silphidae Nicrophorus Animal Necro 21 85

APPENDIX - Continued

Family Genus/Tribe Feeding Habitat Diet Size (mm)

Staphyiinidae Genus A Ground Scav 4 Staphylinidae Genus B Ground Scav 3.5 Staphyiinidae Genus C Ground Scav 2.5 Tenebrionidae Eleodes Ground Scav 27 Tenebrionidae Eusattus Ground Scav 13 Tenebrionidae Lobometopon Ground Scav 12 86

APPENDIX B 87

Levels of Herbivory in Two Costa Rican Rainforests: Implications for Studies of Fossil Herbivory. (Submitted to Biotropica on June 22, 2000)

Dena M. Smith

Department of Geosciences, University of Arizona, Tucson. AZ 85721 USA

ABSTRACT

Levels of herbivory in two Costa Rican rainforests were measured over a two and three year period. Comparisons were made between years at each site and between sites.

The mean percent leaf area removed by insect herbivores did not vary significantly between years, but was different between sites. Leaf area removed by insect herbivores ranged from 4 to 11%. This is within the range of values typically seen in discrete samples from neotropical rainforests. Patterns in guild structure varied between years and between forests. While the differences between these samples were significant, overall trends remained the same and this may provide usefiil information when examining long-term change in guild structure. Samples were collected from leaf litter to ensure the comparability of the data in this study to data from studies of herbivory in fossil assemblages. Although additional studies should be conducted using leaf-litter sampling, current methods of discrete sampling in the understory may be usefiil for the comparison of herbivory levels in modem and fossil assemblages. 88

RESUMEN

Los niveles de herbivorismo en dos bosques humedos de Costa Rica flieron midido para dos y tres afios. Las cotnparaciones se hicieron entre aflos en cada sitio y entre sitios. El medio por ciento de ^ea de hoja quitada por insectos herbivorosos no vario signiflcativamente entre aAos, pero era diferente por sitio. Hojee ^ea quitada recorrida de 4 a 11%. Estos valores de herbivorismo son tipicos para estudiosdistinctos en bosques tropicales. Los modeios en la estructura de gremio variaron entre afios y entre bosques. Mientras las diferencias entre muestras fiieran significativas, las tendencias permanecieron simulares y esto puede proporcionar informacion litil cuando examinando el cambio a plazo largo en la estructura de gremio. Muestras de hojarascas fueron colectadon para segurar el comparabilidad de los datos en este estudio a datos de estudios de herbivorismo en colecciones de fosil. Aunque estudios adicionales se deban conducir usando hojarascas de hoja probando, los metodos actuales de muestreo distinto en el sotobosque pueden ser litil para la comparacion de niveles de herbivory en modemo y las colecciones del fosiles.

Keywords: rates of herbivory, fossil herbivory, annual variation, discrete vs. long-term measurement, tropical wet forests.

INTRODUCTION

Modem levels and patterns of herbivore damage have been examined in a variety of forest types including temperate, subtropical and tropical rainforests (excellent 89 summaries can be found in Coley & Barone 1992 and Landsberg & Ohmart 1989).

Although the diversity of these studies has contributed much information about

herbivory levels in modem forests, the different methodologies used make comparisons

between forests very difficult (Lowman 1986). For example, studies have examined

understory and/or canopy foliage (Lowman 1992, Lowman & Heatwole 1992 and

Sterck et al. 1992), the leaves of persisent vs. pioneer plants, and leaves from plants that were at different growth stages (Coley 1983, de la Cruz & Dirzo 1987 and Southwood et al. 1986). Although most studies were conducted in only a single season, a few authors have also done long-term studies to understand both inter- and intra- annual variation in herbivory levels (Coley 1983 and Filip et al. 1995). In addition, researchers have employed a variety of approaches to sampling to quantify levels of herbivore damage: discrete or long-term sampling methods and grid cells, visual estimation techniques, or leaf area meters (Filip et al. 1995 and Lowman 1984, 1985 and 1986).

In addition to the research conducted on herbivory in modem forests, there has been a growing amount of work on herbivory in fossil forests. These studies are conducted with the hope of shedding light on the evolution of plant-insect interactions.

Much of this effort has focused on identifying specialized interactions or tracing the evolution of particular feeding strategies over time. For example, Scott and Taylor

(1983) examined interactions between insects and plants in the Carboniferous, Beck and

Labandeira (1998) studied herbivory on fossilized leaves from the Permian and detailed work on herbivory in Cretaceous floras was conducted by Labandeira et al. (1994 and 1995). Several workers (Lyon et al. 1999, Palmer et al. 1998, Smith 1998. Wilf &

Labandeira, 1999) are now conducting work on the Tertiary of North America.

Tertiary assemblages contain many extant taxa, which allows for the comparison of herbivory in fossil and modem assemblages. Paleobiologists (Beck&

Labandeira 1998, Labandeira et al. 1995 and Palmer et al. 1998) have compared levels and patterns of fossil herbivory with that in modem ecosystems. However, such comparisons are problematic because they do not take into account preservational biases in the plant fossil record. There are two main problems that need to be considered when comparing fossil herbivory to modem herbivory; 1) fossil leaf assemblages are dominated by canopy leaves and 2) leaves that were eaten entirely leave no fossil record.

Fossil leaf assemblages tend to be dominated by canopy leaves (Bumham et al.

1992, Ferguson 1985), but much of the work on modem herbivory has been conducted on understory leaves. Lowman and Heatwole's (1992) work comparing canopy and understory herbivory in Australian Eucalyptus forests suggests that canopy leaves may have lower levels of herbivory than understory leaves. If so, modem studies that examine understory leaves may appear to have greater levels of herbivory than is found in fossil floras, leading to an incorrect assumption that overall herbivory rates were lower in the past.

Several researchers who study modem forests have commented on the benefits of using long-term sampling methods as opposed to discrete sampling methods

(Lowman 1986 and Filip et al. 1995). By marking leaves and tracking herbivory 91 through time, researchers have been able to recognize those leaves that were eaten completely by insects and to correlate seasonal changes in herbivory levels with seasonal changes in leaf chemistry (Coley 1983 and Faeth 1985). The benefit of this type of sampling is that one can gain a better estimate of "true herbivory", whereas discrete sampling (a one time collection of leaves either from the understory or leaf litter) does not allow one to count leaves that were eaten entirely. While long-term sampling is useful for modem ecosystems, these samples are not comparable to those from fossil assemblages. Leaves that were completely eaten by insects are not preserved as fossils. The absence of this part of the record lowers estimates of fossil herbivory and makes fossil assemblages more similar to discrete samples.

Therefore, only herbivory data that are collected from modem leaf-litter samples is truly comparable to fossil leaf data. This is because leaf-litter samples contain a large proportion of canopy leaves (Bumham 1994. 1997), and do not include those leaves that were eaten entirely.

The goals of this paper are as follows: 1) to collect, analyze and present modem herbivory data in a format that is sensitive to plant preservational bias, thereby allowing robust comparisons with herbivory in fossil floras, 2) to examine the temporal variation in herbivore intensity and feeding guild stmcture in modem tropical forests during a 2-3 year period and 3) to see if different lowland tropical wet forests display similar patterns of herbivory. 92

MATERIALS AND METHODS

STUDY SITES. - Leaves were collected from two sites in a tropical lowland rainforest.

The first site was in the La Selva Biological Station in Heredia Province. Costa Rica

(10°26'N, 83 °59'W). This forest has a mean annual temperature (MAT) of 25.9 ^ C

and a mean annual precipitation (MAP) of400cm (McOade & Hartshorn 1994). La

Selva does not have a 'true' dry season, as it never becomes completely dry - mostly

due to the condensation drip that occurs almost nightly (Hartshorn, 1983).

Approximately 1,744 plant species have been described from La Selva thus far

(Hartshorn & Hammel 1994). Leaf litter samples were collected from two sites in the

primary forest. The first site is located perpendicular to the Camino Experimental Sur

(CES), at the 500m marker. The second site is located perpendicular to the Camino

Circular Lejano (CCL), just West of the Quebrada El Salto. One hundred and fifty

leaves were collected from each site for a total of 300 leaves from La Selva in April

1998, April 1999 and February 2000.

Leaves were also collected from the vicinity of the Sirena Field Station, in the

Parque Nacional Corcovado, on the Osa Peninsula, Puntarenas Province, Costa Rica

(8°30'N, 83°35'W). This forest has a mean annual temperatiwe of approximately

25.5®C and a mean annual precipitation of approximately 500 cm (L. Gilbert, pers. comm.). Unlike La Selva, Corcovado does have a distinct dry season that usually lasts for three months (Hartshorn, 1983). The 1998 El Nifio year had an especially long dry season that lasted for five months (L. Gilbert, pers. comm.). Approximately 845 plant species have been identified around the Sirena Field Station, although nearly 2000 93 species have been described from the Osa peninsula overall (L. Gilbert, pers. comm.).

Leaf-litter samples were collected from two sites in the primary forest. The first site is located parallel to the Espervales trail, beginning near tree seven, on tlie self-guided tree path. The second site is located perpendicular to the Ollas trail, seven meters below the top of the ridge. One hundred and fifty leaves were collected from each site for a total of 300 leaves from Corcovado in March 1998 and March 1999.

SAMPLFNG. -Many of the trees found in La Selva and Corcovado shed their leaves just before the onset of the dry season and therefore all the leaf litter samples were collected in the dr>' season. The leaf litter samples did not include leaf fall from previous years, as the decomposition of the leaf litter is fairly rapid (less than one year) in neotropical rainforests (Parker, 1994) and only freshly shed leaves were collected. Samples consisting of 30 dicotyledonous leaves were collected haphazardly from 1 m^ sampling quadrats, at 10m intervals along a 50 m transect, for a total of 5 samples (150 leaves) from each site. Leaf margins and all insect damage on the leaves were traced onto paper.

The number of leaves that had some form of insect damage and the type of damage found on each leaf was documented. Leaves without damage were also counted. Damage was categorized as belonging to one of the following five fimctional feeding groups; I) hole-feeding - an extemal foliage-feeding strategy, in which an insect feeds through the leaf, leaving behind a hole; 2) margin-feeding - an extemal foliage-feeding strategy, in which an insect feeds on the margins of the leaf; 3) 94

skeletonizing - an external foliage-feeding strategy, in which an insect feeds on the soft

tissues of the leaf, but does not feed on the veins; 4) galling - an internal feeding

strategy, in which an insect feeds and lives between the leaf layers and the plant

responds by building-up leaf tissue around the site; and 5) leaf-mining - an internal

feeding strategy, in which an immature insect lives and feeds within the leaf layers,

leaving behind a blotch or serpentine-shaped mine.

All leaf tracings were scanned into a computer using a flatbed scaimer. All

leaves and insect damage were measured using NIH Image software for the Macintosh.

Measurements were made of the leaf area removed by insects, the area of each type of

damage, and the total original leaf area.

STATISTICAL ANALYSIS. - All statistical analyses were performed with JMP IN 3.2.1

(SAS Institute 1996). Comparisons of the mean original leaf area and mean damage

area were conducted with the Tukey-Kramer Honestly Significant Difference test ~ this

allowed for multiple comparisons of the data from the different sites and years utilizing

analysis of variance (Zar 1996). Comparisons of the relative abundance of damaged

leaves, herbivore intensity and the abundance of fimctional feeding groups for all sites and all years were performed using chi-square analysis of contingency tables.

RESULTS

BETWEEN-YEAR VARIATION IN HERBIVORY. - Levels of herbivory for each site and each year are summarized in Table 1. There is no significant difference in mean 95 original leaf area or mean damaged leaf area between the different sample years at La

Selva. The mean percent leaf area removed by insect herbivores from La Selva 1998,

1999 and 2000 was 9%, 11% and 10% respectively. Mean original leaf area is significantly different between years at Corcovado, but the mean leaf area damaged is not significantly different between years. For Corcovado, mean percent leaf area removed was 4% for the 1998 sample and 6% for the 1999 sample.

The percentages of leaves that were damaged at each site in each year are shown in Figure 1. Herbivore intensity is not significantly different between years at La Selva, with 88% - 92% of the leaves in the samples damaged by insect herbivores. Herbivore intensity is significantly different between years at Corcovado, with 83% damaged in the 1998 sample and 62% damaged in the 1999 sample.

The number of damage types per leaf, another measure of feeding intensity, is shown for all sites and years in Figure 2. Although the majority of damaged leaves in the La Selva samples have two damage types per leaf, only the La Selva 1999 and 2000 samples are not significantly different from one another. The pattern of feeding intensity at Corcovado is not significantly different between years.

The relative abundance of the functional feeding groups in each site and year are shown in Figure 3. The La Selva 1998 and 1999 samples have more leaves attacked by hole-feeders than any other functional feeding group and there is no statistical difference in the overall distribution of functional feeding groups between these two samples. However, margin-feeders are the most abundant feeding group in the La Selva

2000 sample. The Corcovado samples are dominated by hole-feeding and margin- 96 feeding damage and there is not a significant difference between the 1998 and 1999 samples.

BETWEEN-SITE VARIATION IN HERBIVORY. - Despite similarities in original mean leaf area, the mean leaf area damaged is significantly different between La Selva and

Corcovado (Table I). Only La Selva 2000 and Corcovado 1998 are not significantly different from each other in the frequency of damaged leaves in the sample. All other comparisons of damage frequency between the sites are statistically different (Figure 1).

Feeding intensities differ between the La Selva and Corcovado sites (Figure 2).

The majority of damaged leaves from La Selva have two damage types per leaf. In contrast, the majority of Corcovado's leaves have only a single damage type. The distribution of functional feeding groups in the 1998 Corcovado sample is indistinguishable from the La Selva 1998 and 1999 samples (Figure 3). The relative abundance of the functional feeding groups in the Corcovado 1999 sample is significantly different from those found in all of the La Selva sites.

DISCUSSION

The lack of variation in percent leaf area removed between years at each of the study sites is impressive in light of the fact that 1998 was an extremely dry year due to the El Nino. This would have affected herbivory levels in the 1999 samples. Leaves that were damaged in 1998 would have been flushed at the beginning of the 1999 dry season, which typically occurs between January and March (Hartshorn 1983). Other 97

authors have also found consistent herbivory levels in their long-term studies of

neotropical forest understories. For example, Filip et al. (1995) found similar between-

year damage levels throughout their three-year study of a tropical dry forest. Their

study also included an extremely dry El Nino year. That there is little variation between

years in herbivory levels in modem forests is good news for paleobiologists. This

means that any differences between analogous modem and fossil sites can be assumed

to be true differences and not due to annual-scale variation in environmental conditions.

In addition to similarity in the overall damaged area, the La Selva samples were all very similar in the percentages of leaves that were damaged. This however, was not

the case at Corcovado. Although, the area removed by insect herbivores was similar

between years, the number of leaves that were damaged was lower. Corcovado experienced a long dry season during the El Niilo year. It is possible that this affected

leaf growth, the chemical content of leaves, and/or the local insect herbivore population,

ail factors that might have affected the nimiber of leaves that were attacked .

Although La Selva and Corcovado are both tropical lowland wet forests, these sites do experience different annual rainfall pattems. Corcovado has a very pronounced dry season and typically exf)eriences weeks to months without rainfall (L. Gilbert, pers. comm.). In contrast, the La Selva dry season is a period of less rain, but not a complete absence of rainfall. Therefore, it is not surprising that there were differences in the levels and pattems of herbivory between these forests. If seasonality has an effect on levels and pattems of herbivory, picking the most appropriate modem forest for comparison with fossil forests may be difficult for paleobiologists. Although the 98 methods used for estimating MAT and MAP of fossil floras (Wiif 1997, 1998) have improved, it is still difficult to interpret the seasonality of a fossil assemblage.

Therefore, it may be necessary to sample several modem forests of the same typ)e to gain an understanding of the modem range of variation and then compare this range to the fossil herbivory data.

The mean area damaged in all of the La Selva samples and the 1999 Corcovado sample are well within the range of herbivore levels found for other studies of neotropical forests that used discrete sampling - 5.5 - 10.5% (de la Cruz & Dirzo 1987,

Dirzo 1984, Filip et al. 1995, Hendrix and Marquis 1983, Johnstone 1981, Newberry & deforesta 1985, Odum & Ruiz 1970 and Sterck et al. 1992). This suggests that the difference in canopy and understory herbivory may not be as great as noted by Lowman and Heatwole (1992). If this is the case, then studies of herbivory in modem forests that utilize discrete sampling methods may be comparable to fossil data.

There is significant variation between years in the number of damage types on leaves and the overall feeding guild structure. These differences may reflect fluctuations in the diversity and abundance of insect herbivores on a year to year basis.

Studies that closely examine the year to year variability in insect populations are likely to shed light on the corresponding variation in patterns of herbivory. Although the annual and site to site differences in patterns of herbivory are significant, there are interesting similarities between the samples. For example, most of the damaged leaves were attacked by insects from two different feeding guilds and most of the damage on the leaves were made by hole-feeders or margin-feeders. By comparing these data to 99 patterns in fossil assemblages we may be able to learn much more about feeding intensity and changes in guild structure over evolutionary time scales.

This study has shown that there is long-term stability in herbivory levels on ecological time-scales. Although there were differences between the samples in guild structure, the overall patterns may be useful in paleobiological studies of herbivory. In addition, studies of modem forests that employ discrete sampling methods may provide useful information for paleobiologists. Conducting additional studies in other modem forests will provide a greater understanding of the type of variation that exists both between and within modem forests. This will allow for the comparison of herbivory in modem and fossil floras and increase our understanding of the evolution of plant and insect interactions.

ACKNOWLEDGMENTS

I thank the Organization for Tropical Studies for post-course funding to begin this study, and for resources and advising throughout their 98-1 tropical biology field course. Dr. Deedra McLeam provided valuable advice and support throughout the project. Larry Gilbert was an invaluable resource at Corcovado. I gr^^atly appreciate

Stacy Forsyth's help with the La Selva 1998 sample and Satya Rhodes-Conways' help with the Corcovado 1998 sample. Many thanks go to Cesar Nuflo, who helped with collecting, tracing leaves and the statistical analyses. I thank K. W. Flessa, C. C.

Labandeira and J. T. Parrish for their comments on the early drafts of this manuscript. 100

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TABLE 1. Mean original leaf area and mean leaf area damaged for all localities.

Samples that have the same letter in the statistical comparison column are not

significantly different from each other (Tukey-Kramer-HSD). For all significant differences P= 0.05.

Original Leaf Area Statistical Leaf Area Damaged Statistical Locality Mean SE Comparison Mean SE % Comparison

La Selva 98 5124.82 217.14 ab 462.70 47.91 9 a La Selva 99 5296.29 217.50 a 559.69 47.99 11 a La Selva 2000 5265.72 217.14 ab 521.63 47.91 10 a

Corcovado 98 4336.44 273.33 b 153.46 29.00 4 b Corcovado 99 5402.76 273.33 a 300J20 29.00 6 b 107

Figure 1. Relative abundance of damaged leaves in the five samples. Samples that share the same letter are not significantly different. For all significant differences P<

0.001. 108

^100

2 60

99 00 98 99 La Selva Corcovado 109

Figure 2. Herbivore Intensity: Relative percentage of leaves with different numbers of

damage types. Samples that share the same letter are not significantly different. For all significant differences P< 0.001. 1001

98 ' 99 La Selva Corcovado

• 1 damage type B 2 damage types Q 3+ damage types ill

Figure 3. Relative abundance of functional feeding guilds in the five samples. Samples that share the same letter are not significantly different. For all significant differences

P< 0.001. 98 99 00 98 ' 99 La Selva Corcovado

• Hole Feeding • Margin Feeding • Skeletonizing B Galling • Leaf Mining 113

APPENDIX C 114

The Evolution of Plant-Insect Interactions; Insights from the Tertiary Fossil Record.

Dena M. Smith

(Formatted for submission of Paleobiology) 115

Abstract. -Has there been dramatic change in plant-insect interactions over geologic

time? I compared the insect-mediated damage on fossil leaves from the 46 Ma (Middle

Eocene) Green River Formation at Douglas Pass, Colorado to herbivore damage in the

35 Ma (Late Eocene) Florissant Flora of Colorado. Insect damage from these fossil

sites was also compared to herbivore damage on leaves that were collected from the

leaf-litter of six modem tropical and temperate sites. The intensity of herbivore

damage was measured by examining the percentage of leaf area removed by insect

herbivores, the number of leaves damaged in the floras, the number of feeding guilds on

individual leaves in the floras, and the overall feeding guild structure.

Levels of herbivory declined (from 2.5% to 1.4% leaf area removed and 34% to

23% of leaves damaged) as temperature declined during the Eocene-Oligocene cooling

trend. Levels of insect damage in the fossil floras (23-34% of leaves damaged and no

more than 3 feeding types per leaf) were lower than levels found in the modem floras

(72-90% of leaves damaged and more than 3 types of feeding damage per leaf). The

differences in herbivory between the fossil and the modem floras may due to

taphonomic biases, environmental differences between the fossil and modem sites or

evolutionary change through time.

Four long-lived specialized plant-insect associations that were maintained from

the middle to the late Eocene and six were maintained from the late Eocene to the

Recent. Galling diversity increased from the wetter, middle Eocene Green River flora

to the drier, late Eocene Florissant flora. Because galling insects have greater survivorship and higher diversity in more xeric environments, the trend may be driven 116 by environmental change. 117

Introduction

Insects constitute approximately 60% of modem species diversity (Stork 1988) and 75% of plant-feeding insects specialize on a single plant species or on a few closely related species (Bemays and Chapman 1994). Ehrlich and Raven (1964) suggested that these specialized interactions are the result of long-lived associations that became established early in the geologic history of these groups. Ehrlich and Raven also proposed that because plants and insects are locked in a coevolutionary arms-race, these highly specialized interactions helped to generate modem levels of insect diversity.

Ehrlich and Raven's model of coevolution has greatly influenced our understanding of plant-insect interactions and has generated a great body of research on the dynamics of host speciali2:ation and how coevolutionary arms races might influence patterns of insect diversity (Farrell 1998; Farrelletal. 1991; Funketal. 1995; Mitteretal. 1988 and 1991).

Fox (1988) modified Ehrlich and Raven's arm-race hypothesis by proposing a model of diffuse coevolution. According to Fox's diffuse coevolution model, plant- insect interactions should be studied not simply as one-to-one interactions between a single plant species and its insect herbivore, but also at the broader community level.

Fox's diffuse coevolution model is thus more inclusive than Ehrlich and Raven's - because it also takes into account the selection pressures imposed on plants by other plant competitors, specialist and non-specialist insect herbivores, and the predators and parasitoids of their insect herbivores. 118

Some studies have examined community level plant-insect interactions in the

geological record (Ash 1997; Beck and Labandeira 1998; Scott and Taylor 1983; Smith

1998; Wilf and Labandeira 1999). Much of the paleontological work on plant-insect

interactions has focused on the the recognition of host-specific insect damage in the

fossil record (Hickey and Doyle 1977; Hickey and Hodges 1975; Labandeira et al.

1994; Larew 1986; Opler 1973; Upchurch and Dilcher 1990; Wilf et al. 2000).

In this study, I use a community level approach to examine changes in insect-

plant interactions on a geological time scale. This approach allows for an examination of the overall impact of insect herbivores on forest communities as a whole. Therefore,

I can examine the long-term trends in overall levels and intensity of herbivory, trends in specialist and generalist feeding patterns and changes in guild structure over evolutionary time. Conducting studies of fossil herbivory at the community-level also allows for comparisons with data already collected from analogous modem forests, because this approach is used by many neontologists to study herbivory in modem

floras (de la Cruz and Dirzo 1987, Dirzo 1984, Filip et al. 1995, Hendrix and Marquis

1983. Johnstone 1981, Newberry & deforesta 1985, Odum and Ruiz 1970 and Sterck et al. 1992).

Here. I examine the traces of insect-mediated damage on fossilized leaves from the Douglas Pass region of the Green River Formation (Middle Eocene) and the

Florissant Fossil Beds (Late Eocene) of Colorado, USA. The comparison of the two fossil floras provides a rare opportunity to examine patterns of change in insect herbivory over an 11 Ma period, to examine the effects of climatic cooling on levels of 119 herbivory and to detect long lived-associations between plants and insects as shown by specialized feeding damage found in both floras, in a study of plant-insect associations during the Paleocene-Eocene warming trend, Wilf and Labandeira (1999) found that herbivory levels increased as mean annual temperatures increased. Here, 1 test if the subsequent decline in temperatures from the Middle to Late Eocene also corresponds with a decline in herbivory levels.

1 also compare the overall levels and patterns of herbivory in the fossil floras to levels and patterns of damage in modem floras. The Eocene is a period of climatic and ecologic change, with remnant Cretaceous and Paleocene plants becoming extinct, and several modem plants making their first appearances (Wing, 1987). Many modem insect groups also make their first appearances in the fossil record during the Tertiary

(Carpenter 1992) and it is thought that many modem plant-insect associations were likely to have become established during this time period (Farrell 1998; Mitter et al

1991). Do these early modem plant and insect communities have herbivory levels that are similar to modem levels of damage? Have modem specialist associations between specific plant and insect groups already become established? Or have there been dramatic changes in insect-damage levels through evolutionary time?

Locations and Environmental Settings

Green River Formation

Fossil plant material was collected from the Douglas Pass region of Garfield

County, west central Colorado (fig. 1). This locality exposes the upper portion of the 46 120

Ma Parachute Creek Member of the Green River Formation (fig. I) in Colorado.

Located on the western boundary of the Piceance Creek Basin, this portion of the lake has been interpreted as a marginal lacustrine environment (Lundell and Surdam 1975;

MacGinite 1969; Picard and High 1972). MacGinitie (1969) interpreted the Green

River flora as a tropical to subtropical environment with a distinct dry season. Studies which have utilized the nearest living relative approach and plant physiognomic characters (MacGinitie 1969; Wolfe 1994) to interpret climate estimate mean annual temperatures (MAT) of 16-18*'C and mean annual precipitation (MAP) of approximately 86 cm for this portion of the Green River Formation. Little work has been done on the paleoelevation of the Green River flora. MacGinitie (1969) estimated that the Green River flora had an elevation of 305m. However, using foliar physiognomic techniques and paleoenthalpy to estimate paleoelevation (Forest et al.

1999; Wolfe et al. 1998), the average estimate of paleoelevation for the middle Eocene

Green River Formation is 2444m (estimates range between 1564 - 2900m).

Collections of fossil plant material were made in the summers of 1997 and 1998. from four sampling areas spaced 35 m apart within the same stratigraphic interval at the

Douglas Pass locality. Data from the four sample areas were combined into one Green

River sample. Leaf fragments smaller than 1 cm^ were not included in the study. All angiosperm specimens were identified to genus using MacGinitie's (1969) classification, with revisions as made by Call and Dilcher (1994), Manchester (1989), and Manchester et al. (1998). Some legume leaflets were not identifiable to the genus level and are listed as "undetermined leaflets" from the family Leguminosae. Other 121 unidentified specimens were not identifiable because they were either poorly preserved or because they were not identified or figured by MacGinitie (1969). All specimens are deposited in the paleontological collections of the University of Colorado Museum of

Natural History, Boulder, Colorado.

Florissant Fossil Beds

Plant macrofossils were collected fi-om the Florissant Fossil Beds National

Monument in Teller County, Colorado (fig. 2). The Florissant lake beds have been dated at 35 Ma by Epis and Chapin (1974) and are within the Florissant Member of the

Florissant Formation (Evanoff and Doi 1992). The Florissant Fossil Beds were deposited in a deep and narrow lake that was formed by the volcaniclastic damming of the drainage basin (McLeroy and Anderson 1966). MacGinitie (1953) interpreted the ancient lake setting as a warm-temperate environment based on the fossil plant assemblage. Subsequent studies have estimated Mean Annual Temperatures (MAT) of

11-14°C (Meyer 1992 and in press; Wolfe 1992 and 1994) based on leaf margin analysis and a Mean Annual Precipitation (MAP) of approximately 63cm based on the composition of the fossil flora (MacGinitie 1953). MacGinitie also utilized floristic methods to estimate the paleoaltitude of the Florissant flora at 300-900m. More recent studies have used foliar physiognomic techniques and either lapse rates or paleoenthalpy to estimate paleoelevations for Florissant (Forest et al. 1995; Gregory

1994; Gregory and Chase 1992; Meyer 1992 and in press; Wolfe 1992 and 1994) and 122 the average estimate of Florissant paleoelevation is 2774m (estimates range from 1900 -

4133m).

The fossil plants were collected from two localities within the boundaries of the national monument. Both lie within the middle shales of the Florissant member (fig. 2) and the data from these localities were combined into one Florissant sample. Because each localit>' only samples a small local area of the source forest, pooling data from the different localities provides a better representation of the source forest overall. These sites are located on the eastern margin of the paleolake and are interpreted here as near- shore environments. National Park Service staff, volunteers and I made the collections in the summers of 1997 and 1998. All angiosperm specimens (non-angiosperm plants were not included in the analysis) were identified to the genus-level when possible.

MacGinitie's (1953) monograph and subsequent taxonomic revisions of the flora

(Manchester 1989; Manchester et al. 1998) were used to identify the plants. Specimens that were not identifiable to the genus level, but were identifiable at the family-level were given a morphotaxon assignment. Unidentified specimens (17% of the specimens) were usually poorly preserved and/or lacked of distinguishing characteristics. All specimens are in the National Peirk Service collections at Florissant Fossil Beds

National Monument, Colorado.

Assessment of Herbivory

Leaves that had some form of insect damage were counted and the type of damage found on each leaf was documented and measured. Only damage that could be 123 attributed to herbivorous insects was included. Damage that may have been due to detritivory was not included. Labandeira (1998) and Beck and Labandeira (1998) outlined four main criteria for distinguishing the damage patterns made by herbivores from damage that was made by detritivores. In this study. I relied on two of the four criteria, including the presence of plant reaction tissue around the damaged area and signs of an insect's presence, such as larval or pupal chambers and the presence of exit holes. This last criterion was also useful for distinguishing damage resulting from insect herbivory as opposed to fiingal damage. Some fungus-induced damage can appear similar to insect damage, especially leaf galls. However, unlike insect produced galls, fungus galls do not have exit holes. The few examples of suspected fimgus damage were not included in this study.

Insect produced damage was categorized into the following five functional feeding groups: 1) hole-feeding - an external foliage feeding strategy, where an insect feeds through the leaf and leaves behind a hole; 2) margin-feeding - an external foliage feeding strategy, where an insect feeds on the margins of the leaf; 3) skeletonizing

(including scraping of leaf tissue)- an external foliage feeding strategy, where an insect feeds on the soft tissues of the leaf, but does not feed on the veins; 4) galling - an internal feeding strategy, where an insect feeds and lives in between the leaf layers and the plant responds by building-up leaf tissue around the site; and 5) leaf-mining - an internal feeding strategy, where an immature insect lives and feeds within the leaf layers and leaves a blotch or serpentine shaped mine. 124

All of the fossil leaves were photographed using a digital camera and the overall

area of each leaf and the area affected by its insect damage were measured using

National Institutes of Health (NIH) Image software for the Macintosh.

Comparison between Fossil Sites

Although the Green River flora is thought to have been deposited in a playa-lake

environment and Florissant in a deep lake environment, the taphonomic signature of the

leaves found in these deposits are similar. Fossilized leaves were collected from near-

shore envirormients from both sites and both assemblages appear to have an over-

representation of small leaves derived primarily from the canopy of the source forest.

MacGinitie (1969) also noted the taphonomic and taxonomic similarity of the two

floras. Sixty-five percent of the families and 43% of the genera in the Florissant flora

are also found in the Green River flora (MacGinitie 1953, 1969) and MacGinitie

suggested that Florissant might contain some of the same species and/or descendants of

species found in the Green River assemblage. Green River and Florissant also have

similarly high altitudes, with mean paleoelevation estimates for the Green River at

2444m and Florissant at 2774m. The range of estimates (Meyer in press) for both sites overlap (between 1900 — 2900m) as well.

The striking similarities between the Green River and Florissant floras make

them ideal sites for conducting paleoecological comparisons. In addition, the deposition of the Parachute Creek member of the Green River Formation during the 125 middle of the Eocene-Oligocene cooling trend and the occurrence of the Florissant flora near the end of this cooling make it possible to examine the effects of cooling on both overall levels of herbivory and on specific damage types in these assemblages.

Comparisons between Fossil and Modem

I compared the levels and patterns of herbivore damage in the Green River and

Florissant floras to those in four modem tropical floras and two temperate floras.

Among those analyzed were La Selva (Heredia Province, Costa Rica) and Corcovado

(Puntarenas Province, Costa Rica), lowland tropical wet forests and Palo Verde

(Guanacaste Province, Costa Rica), a lowland tropical dry forest. These modem tropical floras are good taxonomic analogs for the Green River flora. Sixty-eight percent of the Green River plant families are also found in La Selva, 65% are found in

Palo Verde and 56% are found in Corcovado (compositions of the Costa Rican floras according to Janzen 1983 and McDade et al. 1994). These sites also have fairly equable climates - with little seasonal variation in temperature, a distinct dry season and a lack of frost. These are also characteristics of the climate of the Green River flora

(MacGinitie 1969).

The two temperate sites. Big Basin State Park (Santa Cruz County, California) and Oak Creek Canyon (Coconino County, Arizona), are typical temperate deciduous forests. Quercus and Sequoia — members of plant families that are also prominent in the

Florissant flora, dominate big Basin State Park. Oak Creek Canyon is also dominated 126

by Quercus and has a large proportion of Populus and Acer as well. The third flora.

Cerro de la Muerte (Puntarenas Province, Costa Rica), is a high altitude (~2700m) tropical forest and is another Quercus dominated assemblage. These modem floras

were chosen because they are climatic analogs for the Florissant flora. Florissant has an estimated MAT of 11-14°C (Meyer 1992 and Wolfe 1992) and Big Basin, Oak Creek

Canyon and Cerro de la Muerte (Kappelle et al. 1995) have MATs of approximately

14°C, 15.6°C. and 13.6°C respectively.

Herbivory data from the modem floras were obtained from studies that used leaf litter samples (Smith, in press). Leaf litter samples provide data that is more comparable to fossil leaf samples than data gathered using long-term sampling of understory plants.

Fossil leaf assemblages tend to be dominated by canopy leaves (Bumham et al. 1992;

Ferguson 1985), as are leaf litter samples (Bumham 1994 and 1997; Greenwood 1992).

In addition, leaves that were completely eaten by insects will not be preserved as fossils and they will not be present in modem leaf litter samples. Modem studies that use long-term sampling methods include leaves that were completely consumed in their herbivory assessments. This method results in estimates of herbivory that always appear to be much greater than estimates of leaf litter or fossil herbivory (Smith, in press).

The leaves from the leaf litter samples were assessed for herbivore damage in the same manner as the fossil leaves. The same criteria for distinguishing insect- mediated damage from that produced by detritivores and fungi were employed.

Damage was categorized into one of the five functional feeding groups that was 127

described earlier. Digital images of all the leaves (or their tracings) were taken and then

measured using NIH Image software for Macintosh.

Statistical Analysis

All statistical analyses were performed with JMP IN 3.2.1 (SAS Institute 1996).

The comparisons of fossil and modem leaf area measurements were examined using the

Tukey-Kramer Honestly Significant Difference Test, which allowed for multiple

comparisons utilizing analysis of variance (Zar 1996). Data on plant relative abundance

and damage frequency were analyzed using chi-square analysis of contingency tables.

Plant genera with low sample sizes (less than 25 leaves) may yield inconclusive

statistical and were not included in the analysis. Statistical comparisons between the

fossil sites and between the fossil and modem assemblages in incidence of herbivory,

herbivore intensity and feeding guild stmcture were performed using chi-square analysis of contingency tables.

Results

A total of five-hundred and eighty-four fossil leaves were collected from the

Green River Formation. Five hundred and twenty-two of the specimens were assigned to one of 22 plant genera in 16 families and the remaining 62 specimens remain unidentified (Table 1). Seven genera account for 78% of the leaves in the assemblage and 81 % of the insect damaged leaves are from these genera. Nine plant genera do not 128 have any insect damage. These genera occur at low frequencies ~ only 1-4 specimens

per genus (Table 1). Although the most abundant plants have the majority of insect damage, the distribution of plant abundance is significantly different (x^= 18.89,

P=.0043) from the distribution of the relative abundance of damaged plants.

Approximately 1,200 fossil leaves were collected from Florissant, but only 624 angiosperm leaves were sufficiently well-preserved to be used in this study. Five hundred and fourteen of those leaves were identified to 48 plant genera and morphotaxa within 17 families (Table 2). One hundred and ten specimens were not identifiable.

The four most abundant plant genera (Cedrelospermum, Fagopsis, Staphylea and Rhus) account for 50% of the leaves in the assemblage and 44% of the insect damage in

Florissant is found on these plants. The distributions of overall relative plant abundance and the relative abundance of plants with feeding damage are significantly different

(x^=91.22. P<.0001), indicating that insects were not feeding on plants in proportion to their abundance. Twenty-two percent of the 48 plant genera at Florissant do not show insect damage.

Comparison between fossil sites

Leaf sizes in the Florissant and the Green River floras are not significantly different (f = 1.49, P = .221), but the amount of area damaged by insects is significantly greater (f = 8.87, P = .003) in the Green River flora (2.5% of original leaf area) than in the Florissant flora (1.4% of original leaf area) (Table 3). 129

Only 23% of the leaves were damaged in the Florissant flora, compared to 34%

in the Green River flora (fig. 3a). This difference is statistically significant (x^ = 16.39,

P<.0001). There is no statistical difference (x^ = 3.4, P=. 183) in the number of feeding

types per leaf between the Florissant and Green River floras (fig. 3b) and damaged

leaves usually have only one type of feeding-damage. No leaf from Florissant or the

Green River has more than three types of damage.

Feeding guild structure (fig. 3c) is slightly different (x^=9.97, P=.041) in the two

floras. The main differences between the two assemblages are the greater proportion of

insect galls and the lack of leaf-miners in the Florissant assemblage.

There are only six examples of specialized feeding damage in the Green River

flora. Elongate, elliptical hole-feeding damage is found on the leaves of

Cardiospermum and distinct oval hole-feeding damage is found on Syzygiodes.

Populus and Salix share skeletonizing damage and two distinct hole-feeding patterns.

Rounded hole-feeding damage is also found on the leaves of Cedrelospermum. These

plants are among the most abundant groups in the Green River flora. Legumes are also

abundant, but have little insect damage, the majority of which consists of nondescript

margin-feeding.

In contrast to the Green River flora, thirteen distinct, specialist insect damage

traces occur in the Florissant flora. These specialized damage patterns tend to occur on

the more rare taxa in the assemblage. Fagopsis. the most abundant plant at Florissant,

has no specialized damage and very little damage of any sort. Cedrelospermum and

Staphylea are the only abundant taxa with stereotyped, specialized feeding damage. 130

Staphylea has two types of specialized feeding damage, a large hole-feeding

pattern and gall damage. Distinct galls and rounded hole-feeding patterns are also found on Cedrelospermum leaves and two galls with a very stereotyped arrangement are found on a Cercis leaf. A specimen of Ulmus also has two galls on a single leaf and galls are been found on a specimen of Rhus and a specimen of Robinia. Specialist hole- feeding/window-feeding damage is found on leaves of Trichilia and highly specialized hole-feeding patterns are found on Cardiospermum and Syzygiodes. One Populus specimen has distinct skeletonizing damage and one unidentifiable leaf has very distinct hole-feeding/window-feeding damage.

Four types of specialized insect damage that are found in both the Green River and Florissant floras. One Populus leaf fi-om Florissimt has the traces of skeletonizing damage that is also found on Populus and Salix leaves from the Green River. The rounded hole-feeding patterns that are found on Cedrelospermum leaves in the Green

River flora are also found on Cedrelospermum leaves in Florissant. Finally, distinct, elliptical hole-feeding patterns (fig. 4) are found on the leaves of Cardiospermum and

Syzygiodes in both floras.

Comparison to the Modem

The leaves from both of the fossil sites are significantly smaller (Table 4) than leaves in all of the modem samples (Tukey Kramer Test, P<0.05). The amount of leaf area removed by insect herbivores was significantly lower in the fossil assemblages 131 compared to all of the modem floras, except the Northern California and Cerro de la

Muerte sites (Table 5).

Fewer leaves were damaged in the fossil assemblage (23-34%) than in the modem sites (72-90%) (Table 6) and these differences are significantly different (Table

7). The number of damage types per leaf is also lower in the fossil sites, where the majority (81 -88%) of damaged leaves were attacked by a single feeding group (Table

6). This is in contrast to the modem floras, where the numbers of damage types per leaf are significantly higher (41-67% of damaged leaves attacked by more than one feeding guild).

The overall distribution of functional feeding groups (Table 9) in the fossil floras is statistically different from all of the modem floras with one exception (Table

10). The Green River flora has a feeding guild structure that is similar to the guild stmcture of Corcovado. All of the fossil and modem floras have a high frequency of hole-feeding and margin-feeding damage. In addition, galls and leaf-mines, which tend to be the result of specialized host-plant associations, are uncommon in both the fossil and modem sites.

Several examples of specialist insect damage occur on the leaves of both the fossil and modem plant genera. These are all likely to be examples of long-lived associations between plants and insects. For example, fossil Populus and Salix have skeletonizing damage that is also common on the modem leaves of these genera.

Cedrelospermum is an extinct genus with distinct galls on the leaves. Its closest living relative. Zelkova, has gall damage produced by aphids on its leaves. Fossil Ulmus and 132

Rhus have galls on their leaves and modem leaves from the same genera commonly

have galls produced by aphids as well.

Discussion

Comparison between Fossil Sites.-Levels of herbivory declined from the middle to the late Eocene. Not only did the amount of leaf area removed by insects decrease, but the number of leaves that were attacked also declined. If many of the insect-plant associations were established, or were becoming established, during the middle Eocene, why don't herbivory levels show an increase in the more recent Florissant flora? One explanation may be climate change.

Wilf and Labandeira's (1999) study of herbivory levels and climate change during the Paleocene to Eocene warming trend found a pattern of increasing levels of insect herbivory with incre£ising temperature. They state that this pattern occurs today across latitude: as temperature increases toward the equator, insect and plant diversity increase and herbivory levels increase as well. By their reasoning, one would predict that herbivory levels would decline during periods of time when temperature declined.

Indeed, this is the pattern shown here. Although climate is likely to play a role in plant- insect interactions, the simple pattern of increasing herbivory levels with increasing temperature and decreasing herbivory levels with decreasing temperature is probably not the whole explanation and several factors are likely to confound this relationship.

First, the latitudinal trend in present day insect herbivory has not been clearly demonstrated. Coley and Barone (1996) suggested that there is a latitudinal trend in 1 herbivory levels and found statistically significant differences between levels of herbivory in temperate and tropical forests. However, they did not take into account the different sampling strategies that are generally used in temperate and tropical sites in their statistical analysis. Tropical workers have largely used long-term sampling techniques for assessing herbivory, which are thought to measure 'true" herbivory.

However, most of the work on herbivory in temperate ecosystems has used discrete sampling methods, which may underestimate levels of "true" herbivory. My own data on modem herbivory levels from different temperate and tropical forests (Smith in press; unpublished data), was collected using consistent sampling methods, and I have not found a clear latitudinal trend in levels of insect herbivory. Thus, the correlation between latitude and levels of herbivory is not well established.

Changes in atmospheric COi could also affect levels of insect herbivory. An increase in CO2 may cause decreased levels of herbivory in some taxa (Agrell et al.

2000; Hattenschwiler and Schfellar 1999; Stiling et al. 1999) and increased levels in other taxa (Goverde et al. 1999). In addition, recent analyses have shown a positive correlation between atmospheric CO2 and temperature in the geologic record (Barron

1997; Sloan and Rea 1996; Sloan et al. 1999). Changes in levels of insect herbivory from the Paleocene through the late Eocene could be due to changes in atmospheric

CO2 and not to changes in temperature.

Finally, the differences in levels of herbivory between the Green River and

Florissant floras may be due to differences in paleoelevation between the two sites.

Florissant may have been the same elevation, or as much as 1266m higher than the 134

Green River flora. Although not tested directly, modem herbivory levels may decline with higher elevation, as a wet tropical site at a high elevation had lower levels of herbivory (4.3% of leaf area removed) than two wet tropical sites at lower elevations (5-

10% leaf area removed) (Smith in review and unpublished data). An examination of the relationship between elevation and herbviory levels in modem forests would help to resolve this issue - although plant and insect diversity and temperature are often correlated with elevation change as well. This may confound understanding the factors that most strongly influence herbivory.

Comparison between Fossil and Modern Sites. -Overall, there are dramatic differences between the fossil and modem sites in the percentage of leaf area damaged, the percentage of leaves that were damaged, the number of damage types per leaf and in feeding guild structure. Many factors could explain the lower levels of herbivory observed in the fossil floras compared to the modem floras. Explanations include taphonomic bias, environmental differences and evolutionary change through time.

If damaged leaves are less likely to be preserved, then the low level of fossil herbivory could be a taxonomic artifact. However, Ferguson's (1985) work on plant taphonomy showed that damaged leaves sink faster than undamaged leaves, suggesting that insect-damaged leaves might have a greater potential to become preserved than their unblemished counterparts. However, Ferguson performed his analyses on artificially damaged leaves and not on actual insect-damaged leaves. Other taphonomic factors might affect the preservation of insect-damaged leaves and make them less 135 likely to be preserved as fossils. Insect-damaged leaves could be more susceptible to biotic decay. Leaves with different types of damage may have different preservation potentials. For example, leaves with internal feeding damage have less exposed area than leaves with external feeding damage and therefore they may sink more slowly than the leaves with external feeding. Actualistic studies that compare the preservation potential of leaves with each of the different types of insect damage and leaves without damage will make it possible to identify other taphonomic filters that may be contributing to apparently lower herbivory levels in the fossil record.

There is also a significant difference in the mean original leaf area between the fossil and the modem sites — the mean size of the fossil leaves is only one-tenth that of modem leaves. The small, coriaceous nature of the Green River and Florissant leaves was also noted by MacGinitie (1953 and 1969), and the small size of leaves in fossil deposits has usually been attributed to a taphonomic bias that favors the preservation of the small, thick sun-leaves of forest canopies (Ferguson 1985; Greenwood 1992, Spicer

1981). Actualistic studies have shown that the leaf litter fi*om the bottom of lakes and streams tends to be approximately 63-67% smaller than the leaves in the source canopy

(Greenwood 1992; Roth and Dilcher 1978). Roth and Dilcher (1978) also found that in lacustrine systems, smaller leaves will travel farther from the paleo-shoreline than larger leaves and that an inverse relationship between distance from shore and leaf size may exist. In addition to these taphonomic factors. Tertiary floras that have smaller leaves may have been deposited during drier climates and/or when there may have been lower mean annual ranges of temperature (H. Meyer, per comm.). 136

Smaller mean leaf sizes in the fossil record are problematic only if insects feed

to a different degree on leaves of different sizes. There is some evidence in the modem

herbivory literature to suggest that this is the case (Basset 1991; Moles and Westoby

2000). If insects feed less on smaller leaves than larger leaves, then the low levels of

herbivory in the fossil sample may be an artifact of taphonomic bias or environmental difference. The modem data used in this study do not have a large enough sample of the smaller leaf size classes to perform such a comparison. Detailed studies that examine herbivory levels on leaves from different size classes are needed to resolve this issue.

Another reason why herbivory may be lower in the Eocene compared to the modem floras may be that the modem sites are not the most appropriate analogs for the fossil sites. The modem tropical sites that I examined had much higher mean annual temperatures and dramatically different herbivory levels than the fossil sites. Two of the modem sites that had mean annual temperatures similar to the fossil sites were not significantly different in the mean leaf area damaged by insects, but were significantly different in the other factors that were examined. To avoid this problem, it might be necessary to examine herbivory in modem forests such as the cloud forests of the Sierra

Madre of northeastem Mexico. These cloud forests have similar mean annual temperatures and altitudes to the Green River and Florissant floras (B. Boyle, per. comm.), and there is substantial taxonomic overlap (all but 2 families in the Green

River and all but 4 families in Florissant) between the fossil and Mexican cloud forests

(B. Boyle, per. comm. and Rzedowski and Palacios-Chavez 1977). A taxon-speciflc 137 approach to comparing fossil and modem herbivory would eilso avoid the problems associated with comparing community-level herbivory in non-analogous fossil and modem forests.

Finally, the lower levels of insect herbivory seen in the fossil floras compared to the modem forests could be due to evolutionary change. If there has been an increase in insect diversity from the Eocene to the Recent, then herbivory may also have increased.

Insect diversity may be an important factor that affects levels of herbivory. For example, the greater herbivore damage observed in Australian forests compared to

Neotropical forests may be attributable to the greater species richness of Australia's phytophagous insects, especially beetles (Morrow 1977; Lowman 1982). There is also some evidence to suggest that the diversity of insect herbivores has increased from the

Tertiary to the Recent (Farrell 1998; Labandeira and Sepkoski 1993). However, this trend may be an artifact of taphonomic bias in the insect fossil record and not a true indication of change in insect diversity over time.

If modem plant and insect associations were just becoming established in the

Eocene, as has been suggested by Farrell (1998) and Mitter et al. (1991), then perhaps the lower levels of herbivore damage in the fossil record are due to an evolutionary time lag between the initial introduction of host plants into an area and their eventual colonization by insect herbivores. Time lags are seen on ecological time scales, ranging from 100 — 10.000 years (Strong et al. 1984). Understanding the early colonization of modem insects onto their host plants will be made possible through the examination of 138 other Cenozic fossil floras and will clarify whether ecological scale patterns can be applied to evolutionary time scales.

Specialist interactions. -Four specialized plant-insect associations were maintained fi-om the middle to the late Eocene and there are six long-lived specialist associations that lasted from the Eocene to the Recent. However, the fossil values should be perceived as underestimates. One reason why specialist associations may be underestimated in the fossil record is that it is nearly impossible to identify specialist feeders that make non-stereotyped damage. Only the damage made by insects that feed in a distinct and consistent manner, on the same plant genus, will be recognized as specialist damage. Many external feeding insects, especially margin feeders, specialize on only one plant species, but do not make a distinct and recognizable damage pattern.

For this reason, the number of specialists found in the fossil record is likely to be severely undercounted. Therefore, the record of specialist damage types in the fossil record should be considered an underestimate of the true numbers of specialists and only highly stereotyped damage patterns, like those made by gallers £md leaf-miners should be compared.

Changes in specialist feeding damage on specific plant taxa might also be underestimated due to differences in abundance of plant genera at different fossil sites.

This could be especially problematic for researchers that use a community-based approach to sampling herbviory in the fossil record. Collecting bulk-samples of fossil floras makes it difficult to collect a representative sample of each plant genus, and 139

therefore the full spectrum of insect damage, at each fossil site. For example, the leaves

of Populus and Salix are abundant (a combined total of 64 specimens) in the Green

River flora and they have evidence of damage made by three different specialist insects.

In contrast, Populus and Salix are much less common (a combined total of 13

specimens) in the Florissant flora and include only one example of specialized feeding

damage. A taxon-based approach, which focuses on patterns and levels of herbivory on

specific plant taxa, would make differential sampling less of a problem. In a taxon-

based approach a researcher can focus on collecting large sample sizes of a speciflc

plant genus to attain a sample that represents the full spectrum of insect damage on that

genus.

There is an increase in the diversity of galling damage in the Florissant flora

compared to the Green River flora. Today, there is a pattern of higher species diversity

of galling insects in drier habitats that have nutrient poor soils (Femandes and Price

1991). Galling insects in wetter environments have lower survivorship due to increased

fungal and pathogenic attack. The increase in the diversity of insect galls in the

Florissant flora may be attributable to the more xeric nature of the Florissant

sitecompared to the wetter, middle Eocene Green River flora. If this were the case, 1

would predict that galling insects were even less diverse in the early Eocene compared

to the middle and late Eocene, due to the comparatively warmer and wetter nature of this time period. 140

Summary

A community-based approach to examining insect-plant interactions

demonstrates patterns of change in herbivory from the middle to the late Eocene and

from the Eocene to the Recent. Levels of herbivory declined as temperature declined during the middle to late Eocene cooling trend. Although this trend correlates with

climate change, factors such as change in atmospheric CO2 and paleoelevation differences between the sites may also contribute. Levels of insect damage in the fossil

floras were generally much lower than levels found in the modem floras. The differences in herbivory between the fossil and the modem floras may due to taphonomic biases, differences in leaf size, environmental differences between the fossil and modem sites or evolutionary change through time.

Evidence of several long-lived specialized plant-insect associations that were maintained from the middle to the late Eocene and from the late Eocene to the modem were also observed. Greater numbers of sp>ecialized insects may have been present in the fossil assemblages, but may not be apparent because of a lack of stereotypy or differential sampling of plant taxa at the fossil sites. To avoid this problem in future studies, I recommend that only highly stereotyped damage patterns be examined or that a taxon-specific approach be employed to examine changes in the diversity of specialized damage through time.

Finally, an observed increase in the diversity of insect galls from the middle to the late Eocene may result from the increasingly xeric nature of paleoenvironments 141 during this time period. I predict that galling diversity will be lower in the early

Eocene due to the wetter climates that occurred during that time interval.

Acknowledgments

I thank the National Park Service for permission to collect and for the use of their collections, facilities and the FLFO database. Many thanks to Or. Herb Meyer for his support, advice and help with fossil plant identifications. April IGnchloe also helped with many phases of the Florissant project. I thank Cesar Nufio, Frank Smith,

Mandi Lyon and Rhiannon Grain for assistance with field collecting and the curation of the Green River fossils. Karl Flessa, Judy Parrish and Cesar Nufio provided valuable suggestions on the early drafts of this manuscript.

Financial support for this project was provided by the American Museum of

Natural History, the Colorado Natural Areas Program, the Department of Geosciences and the Graduate College of the University of Arizona, the Geological Society of

America, a National Science Foundation Predoctoral Fellowship, the Research Training

Grant for the Analysis of Biological Diversification-University of Arizona and Sigma

Xi. This is publication #XX of Centro de Estudias de Almejas Muertas - Division de los Insectos. 142

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Table I. Plants found in the Green River sample. Abundance of leaves and damage levels are reported. Parviiegum. = Parvileguminophyllum. Genera with an asterisk are also found in the Florissant sample.

Famiiy/Genus Number Number Total Original Total Damage of Leaves Damaged Leaves Leaf Area (mm^) Area (mm^)

Anacardiaceae Rhus* 45 8 16.173.00 155.52 Araliaceae Araliophyllum 4 0 446.58 0 Burse raceae Bursera* 3 0 1.108.51 0 Euphorbiaceae Aleurites 8 2 8.072.65 217.52 Fagaceae Quercus* 1 0 290.16 0 Leguminosae Leguminosites 9 1 1.459.48 15.01 Parviiegum. 46 8 8.569.80 161.85 Undet. Leaflets 21 4 3.027.20 127.40 Myrtaceae Syzygioides* 64 38 50.874.26 1.862.46 Oieaceae Osmanthus* I 0 176.97 0 Platanaceae iVfacginiiia 12 4 24,539.36 160.79 Rosaceae Rosa* 3 0 427.43 0 Salicaceae Populus* 36 19 29,584.92 1,011.13 Salix* 28 15 13.191.09 408.74 Sapindaceae Allophylus 10 4 16.713.51 291.51 A thy ana* 3 0 283.21 0 Cardiospermum* 107 39 26.975.08 676.30 Sterculiaceae Sterculia 1 0 302.03 0 Styracaceae Styrax 1 0 748.78 0 Symplocaceae Symplocos 1 0 219.84 0 Ulmaceae Cedrelospermum* 115 31 29.304.48 738.65 Celt is* 3 1 692.94 33.56 Unidentified 62 22 20,703.55 535.18

TOTAL 584 196 253,884.87 6495.62 154

Table 2. Plant taxa found in the Florissant Fossil Beds sample. Abundance of leaves and damage levels are reported. Genera with an asterisk are also found in the Green River sample.

Family/Genus Number Number Total Original Total Damage of Leaves Damaged Leaves Leaf Area (mm^) Area (mm^)

Anacardiaceae Cotinus J 0 2.07739 0 Rhus* 29 4 6.144.14 8.64 Schmaltzia 1 1 489.58 24.70 Burseraceae Bursera* 5 0 K946.48 0 Celastraceae Celastrus 2 0 2,818.63 0 Fagaceae Castanea 1 1 532.23 2.55 Fagopsis 123 5 52,636.50 64.60 Quercus* 24 J'S 6.045.05 61.40 Taxon I 1 1 733.87 26.20 Insertae Sedis Populites I I 847.01 17.24 Juglandaceae Carya 5 0 2.39620 0 Leguminosae Caesalpinites 2 0 491.41 0 Cercis 1 1 842.62 3.09 Conzattia 1 0 Ml .03 0 Prosopis 2 0 876.89 0 Robinia 9 3 U28.12 22.81 Taxon / 7 2 1.624.16 3.57 Taxon 2 5 I 247.72 1.08 Meliaceae Trichilia 4 4 3,612.04 363.62 Myrtaceae Sy::ygiodes* 4 3 3.860.14 80.45 Oleaceae Osmanthtts* 1 0 692.08 0 Proteaceae Lomatia 6 0 1.541.55 0 Rhamnaceae Rhamnites 10 0 595.38 0 Ztiphus 1 0 115.57 0 Taxon / 1 0 1.271.36 0 Rosaceae Amelanchier 1 I 128.78 .46 Cercocarpus 20 2 4,26127 16.28 Crataegus 4 0 2246.72 0 Mai us 1 0 740.81 0 Rosa* 2 0 38.98 0 Vauquelinia 8 I 983.00 3.51 Taxon I 3 I 21725 4.96 Taxon 2 1 0 68.35 0 155

Table 2 continued.

Family/Genus Number Number Total Original Total Damage of Leaves Damaged Leaves Leaf Area (mm^) Area (mm )

Rosaceae (continued) Taxon 5 1 0 296.53 0 Taxon 3 1 1 395.86 1.94 Taxon 4 1 0 229.06 0 Salicaceae Populus* 10 1 16.702.80 45-21 Salix* 3 0 2.750.90 0 Sapindaceae A thyana* 14 0 2.701.11 0 Cardiospermum* 3 J 1.512.69 21.53 Koelreuteria 5 2 1.445.37 6.2 Sapindus 19 1 9,048.45 3.87 Staphylaceae Staphylea 31 21 34.170.70 1.352.90 Taxaceae Torreya 3 0 155.23 0 Ulmaceae Cedrelospermum* 127 34 35.116.80 374.11 Celt is* 1 0 410.50 0 Ulmus 5 2 2-094.98 63.89 Taxon I 1 1 803.28 4.27 Unidentified 110 43 39.265.29 937.78

Total 624 144 249^1.86 3^16.86 156

Table 3. Leaf area measurements from the fossil and the modern sites. SE = Standard Error

Site (N) Mean Leaf SE Mean Leaf Area SE Area (mm^) Damaged (mm^) Damaged

Green River (584) 434.73 23.12 10.95 1.49 2.5 Florissant (624) 399.96 16.97 5.64 1.02 1.4

La Selva (899) 5228.84 125.32 514.62 27.68 10 Corcovado (600) 4869.60 194.34 225.94 20.70 5 Palo Verde (300) 3976.19 162.95 316.24 36.02 8 Cerro (300) 2742.12 208.57 118.31 31.59 43 Arizona (300) 5935J8 208.57 435.87 31.55 7.3 California (150) 2968.45 294.97 133.19 44.62 4.5

Table 4. The results of the comparison of mean leaf area between all fossil and modem sites using the Tukey Kramer-HSD Test. a=0.05, ns = not significantly different *• = significantly different. GRN = Green River, FLOR = Florissant, LASE = La Selva, CORC = Corcovado, CERR= Cerro de la Muerte, ARIZ = Arizona, CALI = California.

GRN FLOR LASE CORC PALO CERR ARIZ CALI *« *« ** *« •* *• GRN - ns FLOR - *• •* *« LASE - ns ns •* •• «« *• CORC ' PALO ** ns *» CERR - ns «« ARIZ -

CALI -

Table 5. The results of the comparison of mean leaf area damaged between all fossil and modem sites using the Tukey Kramer-HSD Test. a=0.05, ns = not significantly different ** = significantly different GRN = Green River, FLOR = Florissant LASE = La Selva, CORC = Corcovado Muerte, ARIZ = Arizona, CALI = California.

GRN FLOR LASE CORC PALO CERR ARIZ CALI •« ** GRN - ns •• ns ns FLOR ns •• ns ** «• LASE - ns *« CORC - ns ns ns •« PALO - ns *• CERR - ns ARIZ -

CALI - 157

Table 6. Percentage of leaves with some form of insect damage and the percentage of leaves with the different damage intensities (number of damage types per leaf) in each of the fossil and modem assemblages. GRN = Green River, FLOR = Florissant, LASE = La Selva, CORC = Corcovado. CERR= Cerro de la Muerte, ARIZ = Arizona, CALI = California.

Low damage intensity — High damage intensity

Percent Leaves Percent Leaves Percent Leaves Percent w/ 3 or SITES Damaged w/ 1 damage type w/ 2 damage types more damage types

GRN 34 81 18 1 FLOR 23 88 12 0

LASE 90 33 57 10 CORC 73 51 42 7 PALO 72 59 35 6 CERR 80 59 37 4 ARIZ 78 46 48 6 CALI 79 56 35 9

Table 7. Comparison of the number of leaves with damage between all fossil and modem sites using a chi-square anaylsis for contingency tables, ns = not significantly different, • = significant at P<0.05. *• = significantly at P<0.001. GRN = Green River. FLOR = Florissant LASE = La Selva, CORC = Corcovado. CERR= Cerro de la Muerte. ARIZ = Arizona, CALI = Califomia.

GRN FLOR LASE CORC PALO CERR ARIZ CALI mm mm mm mm mm GRN - mm «» mm mm mm mm mm FLOR - LASE mm mm mm mm mm m CORC - ns ns ns m PALO - ns ns CERR - ns ns ARIZ - ns

CALI - 158

Table 8. Comparison of damage intensity between all fossil and modem sites using a chi-square anaylsis for contingency tables, ns = not significantly different, • = significant at P<0.05, •* = significantly at P<0.001. GRN = Green River, FLOR = Florissant, LASE = La Selva, CORC = Corcovado, CERR= Cerro de la Muerte. ARIZ = Arizona, CALI = California.

GRN FLOR LASE CORC PALO CERR ARIZ CALI GRN ns •* *« «• •* •* «* •« •• «* «« «« •* FLOR - •* «* *• «« « LASE - CORC - ns ns ns ns « PALO - ns ns « CERR - ns

ARIZ - ns

Table 9. Percentage of leaves with one of the five functional feeding groups found in each of the fossil and modem assemblages. GRN = Green River. FLOR = Florissant. LASE = La Selva, CORC = Corcovado. CERR= Cerro de la Muerte, ARIZ = Arizona. CALI = California.

Percentage Percentage Percentage Percentage Percentage SITES Hole-Feeders Margin-Feeders Skeletonizers Callers Leaf-Miners

GRN 44 43 8 4 1 FLOR 53 34 4 9 0

LASE 48 46 3 1 2 CORC 44 44 5 J 3 PALO 32 53 7 3 5 CERR 35 56 6 1 2 ARJZ 41 45 8 1 5 CALI 23 28 25 13 II

Table 10. Comparison of feeding guild structure between all fossil and modem sites using a chi-square anaylsis for contingency tables, ns = not significantly different, • = significant at P<0.05. •* = significantly at P<0.001. GRN = Green River, FLOR = Florissant, LASE = La Selva. CORC = Corcovado, CERR= Cerro de la Muerte, ARIZ = Arizona, CALI = California.

GRN FLOR LASE CORC PALO CERR ARIZ CALI 159

Figure 1. Location of the Douglas Pass locality and a generalized stratigraphy of the

Parachute Creek Member of the Green River Formation, (stratigraphy formatted after

Grande 1984; Young 1995; Coleetal. 1995). •SOI (ZD • i s? Green River Formation i. ^ s i I 2 y 1 St# " S S Garden Parachute Gulch Creek Member Member

091 161

Figure 2. Location of the Florissant Fossil Beds and generalized stratigraphy of the

Florissant Formation (stratigraphy formatted after Evanoff and Doi 1992). OnTTTJ Age (m.y.)

Florissant 34.9 Formation

Study . Telier to Interval Florissant kCoiorado Springs

Wall Mountain Tuff 36.6

PiKes Peak Granite I-2S

Fluvial Conglomerates Lacustrine Units Insect Fossils Volcanigenic Flow Deposit 0 Plant Macrofosslls Welded Tuff Petrified Stumps Granite 163

Figure 3. Florissant flora compared to the Green River floras. (A) Incidence of herbivory: percentage of leaves damaged in each sample. Florissant is significantly different than the Green River (x^=16.389, P<.0001). (B) Herbivore intensity: frequency of one. two or three damage types per leaf in each sample. Florissant is not significantly different that the Green River (x^=3.40, P = .183). (C) Guild Structure: relative abundance of the different functional feeding groups at each site.. Florissant is significantly different than the Green River (x~=9.97, P = .041). 164

A. x2= 16.398 P<.0001

Leaves 30- Damaged (%)

Florissant Green River

B. • 1 damage type • 2 damage types • 3-»> damage types x2=3.40 Frequency \ P = .183 (%) 40H

Florissant Graen River

C. 60n • Hole Feeding • IMargin Feeding • Skeletonizing B Galling • LeaflMining Frequency (%) x2=9.97 P = .041

Florissant Grsen River 165

Figure 4. Specialized hole-feeding damage on Cardiospermum leaves from (A)

Florissant and from (B-D) the Green River flora. Stereotyped hole-feeding damage on

Syzygiodes leaves from (E) Florissant and from (F, G) the Green River Formatioh. 166 167

APPENDIX D Locality informalion.

Localit}' Location - (Latitude & Longitude) Forest Type MAT PPT

N. Arizona Oak Creek Canyon. AZ. USA - (34"54'N, 111"43'W) Temperate Deciduous 15.6''C 44cm N. California Big Basin Slate Park, CA, USA - (37'M I'N, 122"I3'W) Temperate [)eciduous 14"C 127cm Cerro de la Muertc Puntarenas Province, Costa Rica - (0"40'N, 83"50'W) Tropical Wet Montane I3.6''C 250cm Corcovado Puntarenas Province, Costa Rica - (8"3fl'N, I38''30'W) Lowland Wet Tropical 25.5"C 500cm La Sclva Heredia Province, Costa Rica - (I0"26'N, 83"59'W) Lowland Wet Tropical 25.9"C 400cm Palo Verde Province, Costa Rica - (I0"21'N, 85"2rW) Tropical Dry 24"C 125cm

Green River Gartleld County, CO, USA - (39"35'N, I08"48'W) Subtropical-Temperate 16-19''C 83cm Florissanl Teller County, CO, USA - (38"55'N, I05"I6'W) Warm Temperate I4"C 63cm

Summary Statistics.

Mean Leaf Mean Damage Locality N Area (mm^) SE A Area (mm^) SE A

N. Arizona 300 5935.38 450.57 a 435.87 47.94 7 a N. California 150 2968.45 146.63 b 133.19 20.14 4.5 b Cerro dc la Mucrtc 300 2742.12 120.82 b 118.13 12.60 4 b Corcovado 600 4869.60 194.34 c 225.94 20.70 5 be La Sclva 899 5228.84 125.32 ac 514.62 27.68 10 a Palo Verde 300 3976.19 162.95 d 316.24 36.02 8 c Green River 584 434.73 23.12 e 10.95 1.49 2.5 e Florissanl 624 399.96 16.97 e 5.64 1.02 1.4 e

00 Herbivory data from Arizona.

Leaf Original Hole Feed Margin Feed Skeletonizing Galling Leaf Mining Total Number Leaf Area Area Area Area Area Area Damage Area 1 II81.4 0 0 0 0 0 0 2 218.39 0 0 0 0 0 0 3 440.91 0 0 0 0 0 0 4 337.19 0 36.14 0 0 0 36.14 5 529.75 0 0 0 0 0 0 6 681.61 0 0 0 0 0 0 7 1198.55 0 0 0 0 0 0 8 11527.08 0 0 0 0 0 0 9 5049.31 30.56 0 0 0 75 105.56 10 4474.79 27.52 0 0 0 0 27.52 11 3659.21 0 0 0 0 0 0 12 5921.23 621.46 922.05 0 0 0 1543.51 13 20627.6 0 0 0 0 0 0 14 6812.69 126.36 844.2 0 0 0 970.56 15 12686.93 0 0 0 0 0 0 16 1558.37 209.22 0 0 0 0 209.22 17 13686.87 447.7 69.23 0 0 0 516.93 18 13020.83 0 145.14 0 0 0 145.14 19 7890.28 27.08 0 0 0 0 27.08 20 1252.78 0 0 0 0 0 0 21 4521.53 99.31 135.42 0 0 0 234.73 22 4625.6 148.02 110.13 0 0 0 258.15 23 1621.17 8.31 9.35 0 0 0 17.66 24 4335.23 0 156.88 28.05 0 0 184.93 25 1732.58 0 31.74 0 0 0 31.74 26 1802.61 4.67 21.16 0 0 0 25.83 27 2339.78 15.56 14.94 0 0 0 30.5 28 1525.31 10.89 23.65 0 0 0 34.54 29 12385.15 277.28 134.74 0 0 0 412.02 30 4325.05 64.52 136.15 0 0 0 200.67 31 17782.32 365.53 3408.62 0 0 0 3774.15 32 4069.84 60.77 0 0 0 0 60.77 33 1029.48 0 0 0 0 0 0 34 752.58 95.58 0 0 0 44.13 139.71 35 18137.11 1166.05 445.34 0 0 0 1611.39 36 2991.53 0 392.23 0 0 0 392.23 37 11691.95 617.61 0 0 0 55.78 673.39 38 4449.19 255.72 1121.65 0 0 0 1377.37 39 3437.05 57.81 85.71 76.19 0 0 219.71 40 17777.53 264.28 1016.67 0 0 0 1280.95 41 3382.14 26.33 207.59 0 0 0 233.92 42 4241.86 0 0 0 0 0 0 43 12692.59 283.94 259.26 0 0 0 543.2 44 10140.74 38.27 502.47 0 0 0 540.74 45 1359.26 0 0 0 0 0 0 46 16528.77 443.17 286.68 0 0 0 729.85 47 7025.63 157.74 133.96 0 0 0 291.7 48 3803.26 26.29 396.86 0 0 0 423.15 49 5278.47 0 0 0 0 0 0 50 4616.67 38.89 164.59 0 0 0 203.48 51 1182.64 0 0 0 0 0 0 52 919.44 0 0 0 0 0 0 53 12522.04 0 366.57 0 0 0 366.57 54 5153.21 20.25 76.96 0 0 0 97.21 55 2489.02 71.9 906.29 0 0 0 978.19 56 11495 156 0 0 0 0 156 57 9959 287.13 132 0 0 0 419.13 58 16236.32 1077.4 1413.62 0 0 0 2491.02 59 1470.32 0 0 0 0 0 0 60 2777.61 20.25 85.06 0 0 0 105.31 61 3076.33 103.3 0 0 0 0 103.3 62 8077.66 46.59 0 0 0 0 46.59 63 2656.1 210.62 90.12 0 0 0 300.74 64 8614.61 979.19 1856.13 0 0 0 2835.32 65 19524.55 339.27 0 0 0 0 339.27 66 4499.31 0 592.15 92.64 0 0 684.79 67 2268.43 0 0 0 0 0 0 68 10242.2 265.49 162.35 0 0 0 427.84 69 2031.87 19.45 0 0 0 0 19.45 70 4434.09 0 536.93 46.51 0 0 583.44 71 5494.41 0 206.32 0 0 0 206.32 72 10990.19 679.47 362.46 0 0 0 1041.93 73 13126.41 591.57 1536.73 0 0 0 2128.3 74 10272 0 48.61 0 0 0 48.61 75 3868.2 0 60.76 86.07 0 0 146.83 76 2079.92 0 0 0 0 0 0 77 2171.05 25.32 0 0 0 0 25.32 78 1590.82 0 178.22 0 0 0 178.22 79 12788.43 49.92 147.33 329.97 0 0 527.22 80 2782.21 0 37.75 0 0 0 37.75 81 2278.13 0 479.73 0 0 376.24 855.97 82 9415.69 76.71 196.03 0 0 0 272.74 83 11064 44 0 0 0 0 44 84 1465 0 0 0 0 0 0 85 1610 0 0 0 0 0 0 86 3181 0 0 0 0 0 0 87 8789 35 2534 0 0 0 2569 88 17084.08 0 630.86 0 0 0 630.86 89 9280 0 483.27 0 0 0 483.27 90 12748.05 0 1308.24 0 0 0 1308.24 91 2992.03 230.35 0 0 0 230.35 92 7775.52 101.4 22.53 0 0 0 123.93 93 4147.53 55.08 102.66 0 0 0 157.74 94 5910 0 540 0 0 0 540 95 10958 391 2288 0 0 0 2679 96 4299 13 207 0 53 0 273 97 834 144 21 0 0 0 165 98 2398 0 112 0 0 0 112 99 28339.5 306.17 3683.95 1323.46 0 0 5313.58 100 22868 367 1627 0 0 0 1994 101 7287.65 91.36 0 0 0 0 91.36 102 6686.42 561.72 461.73 0 0 0 1023.45 103 13119.76 1364.22 1501.24 0 0 0 2865.46 104 4756.79 0 0 0 0 0 0 105 7318.35 618.98 201.22 409.01 0 0 1229.21 106 5014.14 0 187 0 0 0 187 107 4539.52 0 0 0 0 0 0 108 1921.45 0 890.18 0 0 0 890.18 109 9434 793 822 0 0 0 1615 no 4018 100 877 0 0 0 977 111 3859 273 102 0 0 0 375 112 4609 0 127 0 0 0 127 113 2638 0 239 0 0 0 239 114 2580 44 393 0 0 0 437 115 1669 0 162 0 0 0 162 116 1594 0 0 0 0 0 0 117 6758 221 108 0 0 0 329 118 6809 0 0 181 0 0 181 119 3882 0 0 0 0 0 0 120 1917 0 369 0 0 0 369 121 6816.39 59.67 545.76 0 0 0 605.43 122 4090.08 41.03 683.75 358.04 0 0 1082.82 123 7050.11 70.86 543.28 0 0 0 614.14 124 3599.02 0 207.61 0 0 0 207.61 125 8622.74 0 466.19 0 0 0 466.19 126 13671.57 1420.19 5434.86 0 0 0 6855.05 127 4592.42 0 25.68 0 0 0 25.68 128 4608.22 0 1377.73 0 0 0 1377.73 129 2171.77 0 0 0 0 0 0 130 1592.04 49.38 0 0 0 0 49.38 131 1036.01 0 59.26 342.7 0 118.51 520.47 132 3188 33 I 0 0 0 34 133 2285 54 123 0 0 0 177 134 3982 11 163 0 68 157 399 135 1048 0 0 0 0 0 0 136 1309 0 0 0 0 0 0 137 1288 0 0 0 0 0 0 138 1765 0 0 0 0 0 0 139 1842 0 0 0 0 0 0 140 1016 0 0 0 0 0 0 141 1674 0 0 0 0 0 0 142 1342 0 28 0 0 0 28 143 1600.95 0 0 0 0 0 0 144 3757.83 0 0 0 0 0 0 145 1873.34 42.53 0 0 0 0 42.53 146 2741.16 0 33.42 0 0 0 33.42 147 3550.42 339.23 0 0 0 0 339.23 148 1587.79 0 0 0 0 0 0 149 1720.44 0 71.9 0 0 0 71.9 150 1195.9 0 0 0 0 0 0 151 16559.5 414.88 278.51 0 0 0 693.39 152 3233.06 0 0 0 0 0 0 153 1585.12 72.73 28.1 0 0 102.48 203.31 154 2062.81 202.48 0 0 0 0 202.48 155 8882.05 93.87 187.05 0 0 0 280.92 156 3373.11 44.88 0 0 0 0 44.88 157 1637.24 23.81 0 0 0 0 23.81 158 4748.48 0 0 0 0 0 0 159 7336.87 0 0 0 0 0 0 160 3346.34 23.08 0 0 0 0 23.08 161 8354.92 114.99 108.23 0 0 0 223.22 162 3262.88 0 0 0 0 128.84 128.84 163 3054.05 0 0 449.2 0 0 449.2 164 9924.6 0 39.47 0 0 0 39.47 165 4769.54 0 321.28 0 0 0 321.28 166 5021.05 66.09 16.52 0 0 0 82.61 167 2782.23 0 98.22 0 0 0 98.22 168 4674.99 37.63 62.42 0 0 0 100.05 169 7523.61 77.78 0 69.45 0 0 147.23 170 2054.17 16.67 0 0 0 0 16.67 171 2665.28 50 24.31 0 0 0 74.31 172 1066.67 27.78 58.33 0 0 0 86.11 173 4165.28 141.67 59.03 0 0 0 200.7 174 3556.94 47.22 121.53 0 0 0 168.75 175 10426.45 105.78 0 0 0 0 105.78 176 1951.24 I! 4.88 0 0 0 0 114.88 177 1322.31 0 54.55 0 0 0 54.55 178 1936.36 0 38.84 0 0 0 38.84 179 1131.4 34.72 38.02 0 375.21 0 447.95 180 6157.02 49.59 0 0 0 0 49.59 181 4184.3 0 251.24 0 0 0 251.24 182 8148.76 0 123.97 0 0 0 123.97 183 3306.61 19.01 57.03 0 0 0 76.04 184 6489.26 0 0 0 0 0 0 185 2602.48 0 306.62 0 0 0 306.62 186 10013.98 53.94 0 0 0 0 53.94 187 1418.55 172.41 321.95 0 0 98.87 593.23 188 5056.43 10.62 278.64 0 0 0 289.26 189 8012.82 23.7 1480.65 0 0 0 1504.35 190 9145 71 152 0 0 0 223 191 2578 0 0 331 413 0 744 192 2513 0 0 0 0 0 0 193 9131 405 636 0 0 0 1041 194 2046 0 0 0 0 0 0 195 1662 0 0 0 0 0 0 196 12413 137 523 0 0 0 660 197 1943 0 0 0 0 0 0 198 2082 28 61 0 0 0 89 199 987 0 0 37 0 0 37 200 10345 110 234 0 0 0 344 201 14585 282 946 0 0 0 1228 202 3363 0 0 0 0 0 0 203 2161 0 161 0 0 0 161 204 2148 73 0 0 0 0 73 205 1653 0 0 0 0 0 0 206 7159 0 67 0 0 0 67 207 12424.21 373.32 552.08 0 0 0 925.4 208 9105.82 136.29 542.2 0 0 0 678.49 209 4623.03 42.47 0 0 0 0 42.47 210 5159.21 120.96 215.58 564.79 0 198.02 1099.35 211 8454.32 40 0 0 0 0 40 212 8992.78 0 252.64 0 0 0 252.64 213 2605.46 0 0 0 0 245.82 245.82 214 1858.25 147.3 170.7 0 0 0 318 215 14059 130 1577 0 0 0 1707 216 3519 0 53 0 0 0 53 217 1603 0 91 0 0 0 91 218 3109 138 1821 0 0 0 1959 219 6853 78 287 0 0 0 365 220 9295.44 124.45 0 0 0 0 124.45 221 2928.28 28.64 54.32 0 0 0 82.96 222 3845.78 27.65 42.47 0 0 0 70.12 223 9012 0 0 0 0 0 0 224 980.7 0 0 0 0 0 0 225 11700.78 246.91 358.52 0 0 0 605.43 226 11924.85 56.65 0 0 0 0 56.65 227 10186.94 0 0 0 0 0 0 228 2767.49 0 0 0 0 947.82 947.82 229 2537.62 0 0 0 0 262.28 262.28 230 7201.74 105.31 0 0 0 0 105.31 231 4441.34 0 862.75 0 0 207.59 1070.34 232 4851.87 124.47 227.18 0 0 0 351.65 233 1265.85 46.25 62.58 0 0 0 108.83 234 812.16 0 0 44.89 0 0 44.89 235 1383.75 0 57 86.75 0 0 143.75 236 7250.93 160.52 525.79 0 0 0 686.31 237 6893.83 508.08 3064.98 0 0 0 3573.06 238 10451.98 207.29 0 0 0 0 207.29 239 1264.7 27.58 0 0 0 0 27.58 240 1365.85 0 0 0 0 0 0 241 2134.87 0 0 0 0 0 0 242 8424.95 212.32 112.85 0 0 0 325.17 243 2438.29 0 102.81 0 0 208.13 310.94 244 1585.69 0 88.6 0 0 0 88.6 245 11177.38 303.56 291.72 0 0 0 595.28 246 5696.5 25.37 0 0 0 0 25.37 247 1831.47 18.6 0 175.03 0 0 193.63 248 6658.74 0 0 0 0 0 0 249 7719.01 93.39 0 0 0 0 93.39 250 9307.43 0 4448.76 0 0 0 4448.76 251 7684.3 25.62 96.69 0 0 0 122.31 252 2707.44 0 29.75 0 0 0 29.75 253 9737.61 190.59 902.79 0 0 0 1093.38 254 2395.74 0 0 0 0 365.3 365.3 255 4754.7 0 0 0 0 0 0 256 7020.88 41.8 2170.88 0 0 0 2212.68 257 8089.18 17.55 0 0 0 0 17.55 258 2304.63 0 0 0 0 0 0 259 1826.48 72.72 0 465.61 0 0 538.33 260 7616.89 71.06 0 0 0 0 71.06 261 514.93 0 0 0 0 0 0 262 2237.75 0 0 229.04 0 0 229.04 263 10514.18 0 0 0 0 0 0 264 1275.61 0 0 0 0 0 0 265 1726.17 0 122.88 0 0 0 122.88 266 6076.29 58.5 0 0 0 0 58.5 267 3315.25 19.23 96.13 0 0 0 115.36 268 1423.57 0 0 0 0 0 0 269 32102.48 62.81 0 0 0 0 62.81 270 7851.24 0 542.15 0 0 0 542.15 271 10575.21 0 3798.34 0 0 0 3798.34 272 21742.15 99.17 0 0 0 0 99.17 273 8277.69 0 0 499.17 0 0 499.17 274 6981.82 0 0 0 0 829.76 829.76 275 11281.38 0 438.26 0 0 0 438.26 276 5167.65 30.81 0 0 0 0 30.81 277 3465.31 29.81 0 94.41 0 0 124.22 278 7402.66 0 0 0 0 0 0 279 5282.93 61.61 45.71 0 0 0 107.32 280 14217 358 2685 199 0 0 3242 281 5488 0 261 0 0 0 261 282 3452 0 32 0 0 0 32 283 8355 95 675 0 0 0 770 284 948 40 0 0 0 0 40 285 6787.6 42.15 140.5 0 0 0 182.65 286 2709.92 55.37 358.68 0 0 0 414.05 287 2061.98 0 0 0 0 0 0 288 10830.58 38.85 1152.07 0 0 0 1190.92 289 1867.77 0 616.53 0 0 0 616.53 290 1996.69 0 400.83 0 0 0 400.83 291 1788.43 0 0 0 0 0 0 292 111572 226 2281 0 0 0 2507 293 767 0 0 0 0 0 0 294 352 0 0 0 0 0 0 295 8113 66 210 0 0 0 276 296 2964 0 51 298 0 0 349 297 2089 0 0 127 0 0 127 298 1344 0 0 0 0 0 0 299 1578 0 0 212 0 0 212 300 2147 0 0 268 0 167 435 Herbivory data from California.

Leaf Original Hole Feed Margin Feed Skeletonizing Galling Leaf Mining Total Number Leaf Area Area Area Area Area Area Damage Area 1 3521.68 328.06 0 0 7.4 0 335.46 2 4794.38 35.94 219.08 0 43.64 0 298.66 3 1540.09 0 30.33 49.99 0 0 80.32 4 1270.12 0 0 12.87 0 66.72 79.59 5 1324.41 0 200.74 0 0 0 200.74 6 3976.33 0 0 54.28 0 0 54.28 7 1194.52 0 56.38 0 0 0 56.38 8 2626.91 0 19.52 82.01 0 184.19 285.72 9 2098 0 0 86.25 0 32.75 119 10 4241.75 0 0 60 0 0 60 11 1553.75 40.25 0 0 0 0 35.94 12 2545.5 11.75 0 10.5 0 0 22.25 13 900.75 0 0 0 7.25 0 7.25 14 1073.25 10 31.75 0 0 0 41.75 15 1514.75 0 0 0 70.25 0 70.25 16 1365.75 0 0 42.25 0 27 69.25 17 1094 0 0 0 0 49.25 49.25 18 1706.61 0 0 0 0 0 0 19 2148.35 0 0 0 0 0 0 20 1814.51 0 0 0 0 0 0 21 2900 0 0 0 0 0 0 22 2047.73 0 0 0 0 13.02 13.02 23 1730.79 0 0 0 76.65 0 76.65 24 7488.19 71.54 0 0 0 0 71.54 25 2031.2 0 64.26 0 0 0 64.26 26 3042.98 0 0 0 0 0 0 27 2269.75 0 0 0 89.75 0 89.75 28 2183.75 0 0 0 0 0 0 29 2062.5 11.35 0 0 16.45 0 27.8 30 1641.33 0 0 0 44.52 0 44.52 31 2199.48 0 0 65.26 0 0 65.26 32 5782.64 0 125 0 0 667.01 792.01 33 3654.17 0 0 0 62.67 0 62.67 34 1809.03 0 35.76 0 0 0 35.76 35 2959.25 0 0 115.57 0 0 115.57 36 3914 0 0 0 0 718 718 37 5294.75 0 0 0 0 918.25 918.25 38 2629.25 0 0 68.25 0 83.5 151.75 39 996.68 0 55.11 96.19 0 0 151.3 40 651.28 2.29 0 0 0 0 2.29 41 2679.72 27.55 151.54 51.15 0 0 230.24 42 1422.93 15.39 56.66 0 0 0 72.05 43 2314.94 0 211.54 55.19 0 27.51 294.24 44 488.9 0 264.54 0 0 0 264.54 45 3122.7 0 104.46 335.85 0 0 440.31 46 2869.5 27 0 0 0 0 27 47 2021 0 0 0 11.75 0 11.75 48 349.69 0 25.31 0 0 0 25.31 49 3506.5 0 0 0 0 0 0 50 2135 11.75 0 0 5 0 16.75 51 8141.36 0 0 0 0 0 0 52 4383.75 0 88.5 0 0 0 88.5 53 3137.75 0 75.25 0 99.75 0 175 54 2108.75 0 0 8.75 8.75 0 17.5 55 6535.74 0 941.53 0 0 0 941.53 56 2447.25 0 0 0 0 0 0 57 1537.16 0 0 2.78 14.41 0 17.19 58 1494.45 0 0 2.08 44.1 0 46.18 59 2867.35 0 0 11.16 0 0 11.16 60 3822.31 6.82 629.96 0 0 0 636.78 61 4820.75 641.25 0 0 14.5 0 655.75 62 603.75 15.25 50.5 0 0 0 65.75 63 1061.5 16.25 0 19.5 0 0 35.75 64 632.75 0 28 14.5 0 0 42.5 65 915.5 16.5 0 0 0 0 16.5 66 900.75 0 17.25 0 0 0 17.25 67 1644.25 0 0 26.25 0 59.25 85.5 68 2261.25 0 0 0 0 0 0 69 2444 0 384.75 11 0 32 427.75 70 1322.5 0 0 8 0 27.25 35.25 71 2215.25 0 0 0 0 0 0 72 1805.75 0 0 38 0 0 38 73 3043.75 0 0 0 0 0 0 74 866.5 8 93.25 0 19.23 0 120.48 75 2418.75 0 0 0 0 0 0 76 2484.5 0 0 0 0 0 0 77 1762.25 0 119 34.5 0 0 153.5 78 3894 42.75 0 0 0 0 42.75 79 1152.5 0 0 0 0 0 0 80 ^ 5579.34 0 0 19.27 0 0 19.27 81 2102 9.75 57.5 0 0 0 67.25 82 3019 0 0 8 0 0 8 83 2027.25 0 0 51.25 0 0 51.25 84 1776.75 0 0 0 0 0 0 85 3039 0 0 16.25 0 0 16.25 86 2441.12 0 431.82 0 0 0 431.82 87 4784.71 146.28 0 0 0 0 146.28 88 2428.51 0 0 0 0 0 0 89 1383 0 26.5 0 59.25 0 85.75 90 1398 42.75 0 34.75 0 0 77.5 91 1302.89 0 0 0 0 0 0 92 3509.3 0 10.12 0 0 0 10.12 93 3431.5 0 0 0 0 78 78 94 4602.5 0 173.75 0 0 100.5 274.25 95 1829 0 0 0 0 0 0 96 3891 0 0 13 0 32.25 45.25 97 3093 0 0 0 0 0 0 98 3602.25 0 136.75 0 0 0 136.75 99 12107.81 0 753.99 0 0 31.08 785.07 100 2705.56 0 0 0 0 0 0 101 3041.15 0 0 53.13 0 160.59 213.72 102 6604.17 259.55 0 0 0 210.94 470.49 103 4137.5 274.67 0 0 0 0 274.67 104 3067.88 167.19 0 0 0 0 167.19 105 2395.31 10.25 5.03 0 0 0 15.28 106 2602.15 37.33 836.8 0 0 0 874.13 107 6933.5 104.25 0 37 0 0 141.25 108 4337.5 0 0 25.5 0 0 25.5 109 3094.75 0 395.75 0 0 0 395.75 110 7451 26.75 0 0 0 0 26.75 Ill 3778.25 0 0 0 0 0 0 112 2703.5 0 0 0 69.25 0 69.25 113 3012.25 41.75 61.25 29.5 0 0 132.5 114 1616.5 2.5 0 27.25 0 0 29.75 115 2843 37.25 55 0 0 0 92.25 116 9525.25 374.25 1466.5 53.75 0 0 1894.5 117 3577.25 0 0 0 61.5 0 61.5 118 603.5 0 202.75 0 0 0 202.75 119 3416.7 0 93.6 0 0 0 93.6 120 2574.25 0 0 108.75 0 0 108.75 121 5505.28 0 0 0 45.24 0 45.24 122 2598.5 0 0 0 0 0 0 123 5463.49 256.32 592.31 0 0 0 848.63 124 3626.73 0 67.65 0 0 0 67.65 125 1787.46 0 0 0 0 0 0 126 2141.87 0 0 0 0 0 0 127 5413 0 0 0 58.25 0 58.25 128 4674.25 0 0 0 0 0 0 129 3427.25 14.5 0 0 0 0 14.5 130 3813.25 0 70.25 0 0 0 70.25 131 4926.08 0 17.21 5.32 0 0 22.53 132 4800.25 0 0 0 0 0 0 133 3410 4.5 0 0 0 0 4.5 134 3926.16 28.55 14.59 0 0 0 43.14 135 3353.75 0 0 4.5 0 0 4.5 136 5921 0 0 0 0 0 0 137 2648 33.25 0 0 0 0 33.25 138 5263.5 0 292.5 0 0 0 292.5 139 3259 0 0 84.75 0 0 84.75 140 4876.75 10.5 119.5 34.25 0 0 164.25 141 1573 0 71.5 8.75 22 0 102.25 142 1298.5 0 9 0 0 0 9 143 1506.5 0 0 0 0 0 0 144 2323.5 0 0 0 0 181.75 181.75 145 2812.25 6.75 0 0 0 0 6.75 146 3099.75 0 0 0 0 0 0 147 1523.5 0 0 30.25 0 0 30.25 148 2913.75 0 0 31 0 0 31 149 4135 0 0 0 0 0 0 150 2646 17 0 0 0 0 17

00 Herbivory data from Cerro dc la Miiertc.

Leaf Original Hole Feed Margin Feed Skeletonizing Galling Leaf Mining Total Number Leaf Area Area Area Area Area Area Damage Area 1 2504.5 0 0 0 0 0 0 2 2623.5 0 2 0 0 0 2 3 1415 7.75 0 0 0 0 7.75 4 3613.75 15.75 69 0 0 0 84.75 5 3565.75 0 0 13.75 0 0 13.75 6 4247.25 0 42.25 0 0 0 42.25 7 3835.25 14.75 85 0 0 0 99.75 8 2359.75 9.75 16 0 0 0 25.75 9 1483.25 101 23.5 0 0 0 124.5 10 1622.5 7.25 0 0 0 0 7.25 II 2400.89 0 0 0 0 0 0 12 2535.95 51.48 410.8 0 0 0 462.28 13 3066.86 28.39 46.89 0 0 0 75.28 14 3396.75 39.06 87.57 0 0 0 126.63 15 2259.76 0 25.59 0 0 0 25.59 16 495.71 0 0 0 0 0 0 17 4383.88 13.64 55.58 0 0 0 69.22 18 3029.13 28.31 72.31 0 0 0 100.62 19 641.32 9.09 0 0 0 0 9.09 20 916.12 10.74 0 0 0 0 10.74 21 772.73 17.56 43.6 0 0 0 61.16 22 2219.1 0 0 74.13 0 0 74.13 23 1589.24 0 56.94 0 0 0 56.94 24 3829.75 42.98 372.94 0 0 0 415.92 25 4445.87 330.36 0 0 0 0 330.36 26 1025 0 0 0 0 0 0 27 1885.59 3.3 0 0 0 0 3.3 28 966.67 0 11.81 6.42 0 0 18.23 29 1289.76 5.9 0 0 0 0 5.9 30 4397.92 0 5.21 11.11 0 0 16.32 31 1386.36 0 31.4 0 0 0 31.4 32 3301.03 5.99 0 0 0 0 5.99 33 1414.58 0 53.99 0 0 0 53.99 34 2533.68 61.98 168.39 0 0 0 230.37 35 2718.64 0 245.27 0 0 0 245.27 36 2225 24.85 361.98 0 0 0 386.83 37 1064.64 16.57 0 0 0 0 16.57 38 1729.73 0 135.8 0 0 0 135.8 39 1414.64 0 295.27 0 0 0 295.27 40 2077.51 11.68 73.82 0 0 0 85.5 41 10296.25 0 0 254.75 0 0 254.75 42 4542.25 0 125.25 0 0 0 125.25 43 11932.75 382.25 0 0 0 0 382.25 44 2118.75 0 247 0 0 0 247 45 2518.75 5.25 0 0 0 0 5.25 46 1637.28 0 0 0 0 0 0 47 2965.38 0 6.07 0 0 0 6.07 48 3141.72 0 37.43 0 0 0 37.43 49 6886.25 9.5 0 0 0 0 9.5 50 23410.5 302 0 0 0 0 302 51 2234.75 0 24.5 0 0 0 24.5 52 1559.75 0 16.25 0 0 0 16.25 53 5036 40.75 0 0 0 0 40.75 54 1843.5 8.5 0 0 0 0 8.5 55 2328 0 0 0 0 0 0 56 2463.5 85.75 21.25 0 0 0 107 57 1202.75 0 0 0 0 25.5 25.5 58 1386.25 0 317.75 0 0 0 317.75 59 1665.75 0 0 0 0 0 0 60 2268.25 0 0 0 0 0 0 61 2525.5 0 0 0 0 0 0 62 4119.97 35.66 58.58 0 0 0 94.24 63 5742.46 51.48 307.1 0 0 0 358.58 64 2461.83 0 165.97 0 0 0 165.97 65 2413 17.75 33.25 0 0 0 51 66 2635 17.75 26.5 0 0 0 44.25 67 3538.5 4.25 0 0 0 0 4.25 68 2110 0 78 0 0 0 78 69 2177.5 17.5 33 0 0 0 50.5 70 3060 0 65.25 0 33.75 0 99 71 2713.5 0 148.5 0 0 0 148.5 72 2784.25 10 0 0 0 0 10 73 2163.25 0 0 0 0 0 0 74 1163.25 0 34.5 0 0 0 34.5 75 1868.75 0 10.75 0 0 0 10.75 76 2694.63 0 0 0 0 0 0 77 3846.9 0 0 0 0 0 0 78 1512.81 134.3 0 0 0 0 134.3 79 3120.45 0 29.13 0 0 0 29.13 80 2352.48 12.19 193.18 0 0 0 205.37 81 3623.35 33.88 129.13 0 0 0 163.01 82 4731.82 9.92 377.69 0 0 0 387.61 83 1708.26 49.79 20.46 0 0 0 70.25 84 952.07 0 16.94 0 0 0 16.94 85 1024.79 0 16.94 0 0 0 16.94 86 2611.98 36.78 0 0 0 0 36.78 87 3435.12 36.36 1257.23 0 0 0 1293.59 88 1440.7 0 13.64 0 0 0 13.64 89 7488.22 0 163.64 0 0 0 163.64 90 772.93 0 67.98 0 0 0 67.98 91 1317.77 0 62.19 0 0 0 62.19 92 4756.8 25.74 32.99 0 0 0 58.73 93 1376.25 0 12 0 0 0 12 94 3036.5 152.25 710 0 0 0 862.25 95 1778.25 0 32.25 0 0 0 32.25 96 5666 0 27 0 0 0 27 97 2304.75 0 0 0 0 0 0 98 5670.25 31.25 16.75 0 0 0 48 99 3895 69 85.75 0 0 0 154.75 100 8147 372 0 0 0 0 372 101 2824.85 97.62 97.64 0 0 0 195.26 102 866.86 0 0 0 0 0 0 103 1675.5 0 0 0 0 0 12.75 104 4607.75 26 93.25 0 0 0 119.25 105 4090.25 26.25 573.5 0 0 0 599.75 106 2234 16.75 101.25 0 0 0 118 107 2046 25.5 47 0 0 0 72.5 108 954 0 0 0 0 0 0 109 2372.5 0 9 0 0 0 9 110 2589.5 0 0 0 0 364.75 364.75 111 1217.36 0 212.36 0 0 0 212.36 112 5167.5 24.75 119.5 0 0 0 144.25 113 1369.21 0 137.19 0 0 0 137.19 114 1611.98 0 0 0 0 0 0 115 1052.27 0 43.39 0 0 0 43.39 116 4006 0 753.25 0 0 0 753.25 117 2350 17.25 101 0 0 0 118.25 118 2234.92 53.31 29.75 0 0 0 83.06 119 2132.02 0 0 0 0 0 0 120 1513.64 0 52.89 0 0 0 52.89 121 1442.77 0 152.9 0 0 0 152.9 122 1633.47 14.27 93.8 0 0 0 108.07 123 3576.25 0 115 0 0 0 115 124 4995.25 0 255.99 0 0 0 255.99 125 1809.75 0 0 0 0 0 0 126 4294 0 385.75 0 0 0 385.75 127 1152.26 4.17 3.3 0 0 0 7.47 128 623.61 0 68.57 0 0 0 68.57 129 1488.89 0 197.57 0 0 0 197.57 130 1605.03 5.56 0 0 0 0 5.56 131 2798.96 0 295.66 0 0 0 295.66 132 3257.99 0 191.6 0 0 0 191.6 133 2153.47 0 14.41 0 0 0 14.41 134 1493.58 0 170.14 0 0 31.42 201.56 135 1975.69 0 0 0 0 0 0 136 1004.34 0 30.03 0 0 0 30.03 137 568.75 3.99 0 0 0 0 3.99 138 3201.22 22.57 0 0 0 0 22.57 139 1660.24 45.66 0 0 0 0 45.66 140 2457.81 0 16.32 0 0 0 16.32 141 3589.76 26.39 29 0 0 0 55.39 142 1009.03 0 81.08 0 0 0 81.08 143 1867.71 0 35.24 0 0 0 35.24 144 2093.23 0 0 0 0 0 0 145 1245.49 0 28.13 0 0 0 28.13 146 2212.33 9.03 177.08 0 0 0 186.11 147 1656.94 0 0 0 0 0 0 148 2727.26 0 0 0 0 0 0 149 5283.85 0 0 0 0 0 0 150 5159.38 0 464.06 0 0 0 464.06 151 8253.13 90.28 1519.65 0 0 215.1 1825.03 152 1262.15 8.86 0 0 0 0 8.86 153 337.67 0 0 0 0 0 0 154 1602.95 0 0 0 0 0 0 155 4228.13 9.9 721.25 0 0 0 731.15 156 2037.15 0 0 0 0 0 0 157 10695.75 139.75 721.25 0 0 0 861 158 2653.47 0 0 0 0 0 0 159 1068.4 0 0 0 0 0 0 160 1984 0 0 0 0 0 0 161 4093.5 77.75 22.5 134.25 0 0 234.5 162 1949 0 0 0 0 0 0 163 3248.25 0 0 0 0 0 0 164 1821.75 0 19.5 0 0 0 19.5 165 1559 0 0 0 0 0 0 166 11308.88 166.94 951.44 0 0 0 1118.38 167 3020.25 0 37.19 0 0 0 37.19 168 2117.77 12.81 9.09 0 0 0 21.9 169 1761.78 0 29.13 0 0 0 29.13 170 2035.12 0 162.4 0 0 0 162.4 171 2518.18 0 36.98 0 0 0 36.98 172 10167.15 0 0 0 0 0 0 173 2652.89 0 43.39 0 0 0 43.39 174 1520.45 7.64 6.4 0 0 0 14.04 175 2429.75 0 8.47 0 9.09 0 17.56 176 1730.79 0 0 0 0 0 0 177 4240.91 0 74.79 0 0 0 74.79 178 2124.85 0 0 0 0 0 0 179 3366.72 0 114.35 0 0 0 114.35 180 2850.15 22.34 27.07 0 0 0 49.41 181 4176.65 0 0 0 0 0 0 182 3639.79 11.24 185.36 0 0 0 196.6 183 3852.07 17.75 0 41.86 0 0 59.61 184 2254.59 0 14.64 8.58 0 0 23.22 185 5569.21 0 295.03 0 0 0 295.03 186 1686.98 16.95 127.9 0 0 0 144.85 187 3446.9 0 526.74 0 0 0 526.74 188 4909.71 3.1 290.28 0 0 0 293.38 189 4335.74 41.53 603.09 0 0 0 644.62 190 2496.49 0 304.54 0 13.43 0 317.97 191 2693.6 0 38.64 0 0 0 38.64 192 2451.92 2.81 198.38 0 0 0 201.19 193 1935.06 35.94 0 0 0 0 35.94 194 3038.14 10.97 82.4 10.95 0 0 104.32 195 2314.64 0 86.98 0 0 0 86.98 196 1592.56 0 0 0 0 0 0 197 1628.25 0 0 0 0 0 0 198 1064.11 0 98.54 0 0 0 98.54 199 1827.22 5.47 69.82 0 0 0 75.29 200 1626.04 17.9 106.8 0 0 0 124.7 201 1884.92 10.74 69.21 10.12 0 0 90.07 202 2461.57 0 603.71 0 0 0 603.71 203 1918.39 8.68 13.02 0 0 0 21.7 204 1053.1 0 0 0 0 0 0 205 3116.75 0 0 0 0 0 0 206 2487 5.75 171.5 0 0 0 177.25 207 4060.25 14.25 401.5 0 0 0 415.75 208 1451.65 0 34.92 0 0 0 34.92 209 2524.11 10.08 5.61 14.29 0 0 29.98 210 2830.1 0 9.44 0 0 0 9.44 211 2334.25 0 24 0 0 0 24 212 8821 1254.5 115 0 0 0 1369.5 213 3706.5 9 133.25 0 0 0 142.25 214 1461.5 0 0 0 0 0 0 215 2826.25 14.25 157.75 0 0 0 172 216 1258 0 0 0 0 0 0 217 1077 0 0 0 0 0 0 218 2139.75 0 19.5 0 0 0 19.5 219 1810 25 91 0 0 0 116 220 2042.28 0 43.83 0 0 0 43.83 221 2639.54 10.33 77.29 0 0 0 87.62 222 1895.03 58.16 15.18 0 0 0 73.34 223 839.5 0 29.5 0 0 0 29.5 224 2272 6.5 0 0 0 0 6.5 225 1723.5 7.75 4.75 0 0 0 12.5 226 818 10.5 32.5 0 0 0 43 227 1316.25 17.75 131 0 0 0 148.75 228 1025.75 16.5 80.75 0 0 5 102.25 229 2021 0 44.75 0 0 0 44.75 230 2578.75 0 79 0 0 0 79 231 2354.5 0 110.75 0 0 0 110.75 232 3665.25 0 125.75 0 0 0 125.75 233 2326.75 13.75 120.25 0 0 0 134 234 2084.5 6.75 92.75 0 0 0 99.5 235 1626.25 0 0 0 0 0 0 236 1154.5 0 13.75 0 0 0 13.75 237 1776.5 0 25.25 0 0 0 25.25 238 2342.25 0 42 0 0 0 42 239 2607.75 0 285.75 24.5 0 0 310.25 240 3937.25 57.5 12.75 35.75 0 0 106 241 3082 0 0 0 0 0 0 242 3238.25 0 0 0 0 0 0 243 4130.5 7 20 0 0 0 27 244 1658.5 0 23 0 0 0 23 245 3452.75 0 121.75 0 0 0 121.75 246 2149 0 24 0 0 0 24 247 702 0 0 40.25 0 0 40.25 248 2441.5 0 137 0 0 0 137 249 1935.25 0 25.25 0 0 0 25.25 250 1122.25 0 25 0 0 0 25 251 2580.03 0 0 0 0 0 0 252 3385.24 0 7.64 0 0 0 7.64 253 1506.42 0 0 0 0 50.67 50.67 254 2247.4 0 0 0 0 0 0 255 2813.37 0 0 0 0 0 0 256 3086.28 0 42.19 14.92 0 0 57.11 257 2899.31 28.47 48.27 0 0 0 76.74 258 2147.25 0 0 0 0 0 0 259 1898.96 0 0 28.99 0 0 28.99 260 1837 0 43.5 0 0 0 43.5 261 3306.25 32.75 0 0 0 0 32.75 262 1586.25 0 130.75 0 0 0 130.75 263 1620 30.5 80.75 0 0 21 132.25 264 3613.25 0 0 34.75 0 0 34.75 265 2167.5 0 0 0 0 0 0 266 940.5 13.25 26.75 22.25 0 0 62.25 icn 1394.75 10.75 70.75 0 0 0 81.5 268 1767.25 48 0 0 0 0 48 269 2960.5 0 0 0 0 0 0 270 1912.25 8 154 0 0 0 162 271 2632.25 0 59.75 43.5 0 0 103.25 111 3685.76 0 0 0 0 0 0 273 2098.61 0 0 0 0 0 0 274 3214.06 0 25.35 0 0 0 25.35 275 493.06 0 0 0 0 0 0 276 1515.8 0 0 0 0 0 0 111 761.28 0 41.49 0 0 0 41.49 278 1089.93 0 10.07 0 0 0 10.07 279 1030.9 0 32.99 0 0 0 32.99 280 1076.22 4.69 0 0 0 0 4.69 281 2701.74 0 0 0 0 0 0 282 3415.97 0 0 85.59 0 0 85.59 283 6558.75 237 817.77 0 0 0 1054.77 284 2953.47 17.01 137.16 0 0 0 154.17 285 3616.75 0 32.25 0 0 0 32.25 286 1530.5 9 0 0 0 0 9 287 1789 15.75 0 0 0 0 15.75 288 1542.53 0 196 0 0 0 196 289 3683.33 0 293.4 0 0 66.84 360.24 290 3052 0 154.5 0 0 0 154.5 291 4072.92 154.86 493.79 0 0 0 648.65 292 3080.25 0 26 0 0 0 26 293 5543.5 17.75 0 0 0 0 17.75 294 2313.25 15.75 0 0 0 0 15.75 295 2784.72 0 0 0 0 0 0 296 1813.02 127.6 295.32 0 0 0 422.92 297 4152.3 0 0 0 0 0 0 298 3730.74 17.72 5.74 0 0 0 23.46 299 4495.66 13.14 31.76 0 0 0 44.9 300 1359.31 59.82 0 0 0 0 59.82 Herbivory data from Corcovado.

Leaf Original Hole Feed Margin Feed Skeletonizing Galling Leaf Mining Total Number Leaf Area Area Area Area Area Area Damage Area 1 12858.92 0 0 54.8 0 0 54.8 2 3719.36 0 0 0 0 0 0 3 2023.95 88.17 68.36 0 0 81.57 238.1 4 2436.69 149.4 494.6 0 0 0 644 5 1593.8 0 0 0 0 0 0 6 2943.52 0 0 0 0 0 0 7 2692.62 43.99 413.21 0 0 0 457.2 8 4572.43 21.89 0 0 0 0 21.89 9 1976.66 0 0 0 0 0 0 10 5247.8 21.05 0 0 0 0 21.05 11 10049.2 0 0 0 0 0 0 12 5654.22 239.96 324.97 0 0 0 564.93 13 713.06 0 0 0 0 0 0 14 8626.25 0 0 0 0 0 0 15 3719.92 0 0 0 0 0 0 16 2991.98 49.99 0 0 0 0 49.99 17 3520.05 41.17 0 0 0 0 41.17 18 8791.67 0 0 0 0 0 0 19 8846.88 0 0 0 0 0 0 20 5452.22 0 1274.7 0 0 0 1274.7 21 6044.23 17.31 2082.69 0 0 0 2100 22 1704.29 0 0 0 0 0 0 23 2288.16 130.64 0 0 0 381.2 511.84 24 1148.13 0 0 0 0 0 0 25 1934.32 81.14 0 0 0 0 81.14 26 1913.38 0 0 0 0 0 0 27 1679.24 25.7 0 0 0 0 25.7 28 5845.97 0 14.28 0 0 0 14.28 29 14204.65 805.41 0 0 0 0 805.41 30 4102.33 0 0 0 0 0 0 31 3499 0 1168.63 0 0 0 1168.63 32 1336.51 62.32 255.06 0 0 0 317.38 33 8301.02 447.58 1382.78 0 0 0 1830.36 34 5774.62 0 1078.57 0 0 0 1078.57 35 19816.35 0 0 0 0 0 0 36 16599.46 0 0 0 0 0 0 37 3073.89 75.82 0 0 0 0 75.82 38 1357.32 195.62 110.11 0 0 0 305.73 39 1236.76 0 0 0 0 0 0 40 29013.12 0 0 0 0 0 0 41 1760.14 0 0 0 0 0 0 42 5555.13 148.34 16.87 0 0 0 165.21 43 2344.94 0 0 0 0 0 0 44 1110.82 0 0 0 0 0 0 45 16660.42 660.22 83.06 0 0 0 743.28 46 7517.88 0 0 0 0 0 0 47 5803.12 52.6 52.26 0 0 0 104.86 48 2940.62 0 0 0 0 0 0 49 3042.88 23.44 134.03 0 0 0 157.47 50 6528.75 243.44 1206.7 0 0 0 1450.14 51 1493.4 43.57 106.94 0 0 0 150.51 52 17740.74 507.78 1961.26 0 0 0 2469.04 53 11299.54 0 0 0 0 0 0 54 12244.35 98.33 544.11 0 0 0 642.44 55 1528.3 0 0 0 0 0 0 56 6266.32 52.19 26.61 0 0 364.66 443.46 57 36000.59 0 0 0 0 0 0 58 11977.49 214.57 0 0 0 0 214.57 59 1294.05 55.59 38.01 0 0 0 93.6 60 1552.53 0 0 0 0 0 0 61 2100.11 0 0 0 0 0 0 62 1843.7 0 0 0 0 0 0 63 1873.72 0 46.81 0 0 0 46.81 64 4798.76 114.46 0 0 0 0 114.46 65 4200.21 14.26 0 0 0 0 14.26 66 4054.75 13.64 0 0 0 0 13.64 67 2074.17 62.4 871.08 62.4 0 0 995.88 68 4656.48 11.9 0 0 0 0 11.9 69 7550.95 172.53 602.35 0 0 0 774.88 70 2151.66 0 23.3 88.77 0 0 112.07 71 8728.49 0 0 0 0 0 0 72 2858.56 12.42 288.64 140.31 0 0 441.37 73 6361.06 0 19.22 0 0 0 19.22 74 2888.13 87.86 395.37 0 0 0 483.23 75 11397.2 21 707.56 53.12 0 0 781.68 76 11010.18 715.77 180.38 0 0 0 896.15 77 2104.16 0 0 0 0 0 0 78 1660.66 0 0 0 0 0 0 79 5064.38 0 0 0 0 0 0 80 3768.6 0 0 0 0 0 0 81 7324.14 0 0 0 0 0 0 82 7273.59 0 0 0 0 0 0 83 12635.8 116.13 1595.34 0 0 0 1711.47 84 3357.05 0 372.81 0 0 0 372.81 85 3242.73 146.78 36.64 0 0 0 183.42 86 2688.25 0 130.99 0 0 0 130.99 87 2558.05 0 87.33 0 0 0 87.33 88 5648.27 29.17 0 0 0 0 29.17 89 2974.24 0 56.93 0 0 0 56.93 90 1495.43 0 0 0 0 24.38 24.38 91 552.15 0 36.84 0 0 0 36.84 92 2696.9 0 0 0 0 0 0 93 11083.81 282.59 138.13 0 0 0 420.72 94 2032.29 34.39 0 0 0 65.74 100.13 95 4153.64 0 434.16 0 0 0 434.16 96 1026.29 20.72 47.14 0 0 0 67.86 97 13959.85 0 0 0 0 0 0 98 19726.72 1501.89 2343.41 0 0 0 3845.3 99 5084.05 74.97 27.97 0 0 0 102.94 100 1875.39 22.66 0 0 0 0 22.66 101 10826.27 112.31 113.57 0 0 0 225.88 102 9171.39 0 0 0 0 0 0 103 15773.12 1265.28 1199.68 0 0 0 2464.96 104 1696.29 0 0 0 0 0 0 105 11764.41 0 0 0 0 0 0 106 12665.29 1458.64 1320.62 0 0 0 2779.26 107 6459.09 0 575.81 0 0 0 575.81 108 4612.67 0 0 0 0 0 0 109 4312.26 0 0 0 0 0 0 110 5167.58 0 0 0 0 0 0 Ill 7260.3 0 535.5 0 0 0 535.5 112 7828.19 59.06 293.44 0 0 0 352.5 113 8765.93 78.06 0 0 0 0 78.06 114 8299.15 0 171.09 0 0 0 171.09 115 3684.38 0 280.97 0 0 0 280.97 116 6611.09 0 37.27 0 0 0 37.27 117 8650.74 0 0 0 0 0 0 118 14141.11 0 0 0 0 0 0 119 2722.04 77.07 0 0 0 0 77.07 120 7639.02 0 0 0 0 0 0 121 1943.03 72.33 0 0 0 0 72.33 122 8237.18 30.02 511.56 116.47 0 0 658.05 123 13124.56 0 0 93.35 0 0 93.35 124 5062.33 0 0 374.37 0 0 374.37 125 8619.37 0 0 0 0 0 0 126 2150.88 51.57 0 0 0 0 51.57 127 3134.17 269.53 0 0 0 0 269.53 128 3851.84 0 0 0 0 0 0 129 2954.19 0 0 0 0 0 0 130 3784.17 0 0 0 0 0 0 131 1585.66 0 0 0 0 0 0 132 2469.59 0 0 188.26 0 0 188.26 133 2055.5 0 0 0 0 0 0 134 5263.58 22.42 0 0 0 0 22.42 135 2906.37 0 0 0 0 0 0 136 2808.33 0 0 0 0 0 0 137 3097 0 0 0 0 0 0 138 3214.52 0 0 0 0 20.7 20,7 139 1060.08 0 64.01 0 0 0 64.01 140 1172.87 0 87.57 0 0 0 87.57 141 3442.39 18.58 52.72 0 0 0 71.3 142 3170.51 0 11.31 0 0 0 11.31 143 2919.28 102.8 109.95 0 0 0 212.75 144 3167.35 0 81.13 0 0 0 81.13 145 2640.19 0 67.42 0 0 0 67.42 146 1368.73 0 347.44 0 0 0 347.44 147 4049.65 145.1 86.63 0 0 0 231.73 148 1167.68 0 0 0 0 0 0 149 1441.02 0 0 87.87 0 0 87.87 150 686.8 0 104.04 0 0 0 104.04 151 15442.98 42.97 696.28 0 0 0 739.25 152 12082.44 22.73 0 0 0 0 22.73 153 5548.96 677.22 255.33 0 0 44.97 977.52 154 3830.18 0 0 0 0 0 0 155 4112.13 0 0 0 0 0 0 156 1418.05 0 0 0 0 0 0 157 14200.62 0 1111.77 0 0 0 1111.77 158 7016.12 206.2 99.59 510.33 0 0 816.12 159 10258.85 27.77 15.97 0 0 0 43.74 160 4725.35 48.26 0 0 0 0 48.26 161 3204.86 0 0 0 0 0 0 162 1995.14 0 0 0 0 0 0 163 12704 0 1216.5 0 0 0 1216.5 164 7951.25 0 0 0 0 0 0 165 3249 0 0 0 0 0 0 166 2817 0 0 0 0 0 0 167 4660.08 0 111.35 0 16.8 504.47 632.66 168 1778.57 34.06 0 0 0 0 34.06 169 2191.96 0 0 0 0 0 0 170 3143.24 0 0 0 0 0 0 171 1425.13 0 0 0 0 0 0 172 13772.06 44.62 0 0 0 0 44.62 173 11440.24 105.46 53.25 0 0 0 158.71 174 8207.69 359.03 0 0 0 0 359.03 175 4250.26 79.75 21.61 132.98 0 0 234.34 176 2486.32 0 0 0 0 0 0 177 1839.94 0 0 0 0 0 0 178 1588.61 0 0 0 0 0 0 179 9434.27 1263.8 0 0 0 0 1263.8 180 12990.97 59.55 208.51 0 0 0 268.06 181 10448 0 22.91 0 0 0 22.91 182 5425.17 0 1076.04 0 18.1 0 1094.1 183 1295.21 0 0 0 50.8 0 50.77 184 14418.34 998.21 955.92 0 0 0 1954.13 185 15797.14 0 0 0 0 0 0 186 3411.16 0 39.11 0 0 0 39.11 187 2987.57 0 132.54 0 0 0 132.54 188 2097.38 0 36.94 0 0 0 36.94 189 1428.35 57.68 0 0 0 0 57.68 190 7398.56 0 0 0 0 0 0 191 4710.24 0 1203.2 0 0 0 1203.2 192 7911.73 0 0 0 0 0 0 193 3000.32 172.16 42.88 0 0 0 215.04 194 3044 0 0 0 0 0 0 195 3115.77 0 0 0 0 0 0 196 5869.97 0 0 0 0 0 0 197 2983.66 0 0 0 0 0 0 198 2012.68 0 0 0 0 0 0 199 1142.48 0 0 0 0 0 0 200 1531.73 0 0 0 0 0 0 201 7659.31 63.53 61.6 0 0 33.67 158.8 202 10040.43 29.72 134.44 0 0 0 164.16 203 26552.07 179.56 83.57 0 0 46.9 310.03 204 1473.35 0 0 0 0 0 0 205 946.28 0 0 0 0 0 0 206 1161.98 10.33 7.44 0 0 0 17.77 207 13077.53 791.32 33.18 0 0 0 824.5 208 7664.76 0 1581.78 0 0 0 1581.78 209 2766.81 0 0 0 0 0 0 210 2298.43 0 188.74 0 0 0 188.74 211 4017.24 0 0 0 0 0 0 212 1884.64 0 32.79 0 0 0 32.79 213 1881.68 0 0 0 0 0 0 214 2493.04 0 115.16 0 0 0 115.16 215 2149.53 6.72 0 0 0 0 6.72 216 1152.99 0 110.23 0 0 0 110.23 217 1227.26 0 0 0 0 0 0 218 1393.38 0 0 0 0 0 0 219 9306.5 223.41 74.94 53.33 58.4 0 410.06 220 7413.41 510.02 123.21 0 0 0 633.23 221 5577.81 471.26 1646.66 0 0 0 2117.92 222 3688.22 0 0 0 0 0 0 223 1682.62 0 0 15.35 0 0 15.35 224 16502.07 383.15 436.69 0 0 0 819.84 225 2836.83 58.44 267.6 0 0 0 326.04 226 12786.39 234.4 9.45 0 0 0 243.85 227 1182.99 0 0 0 0 0 0 228 4498.11 129.29 6.05 0 0 0 135.34 229 1549.53 0 0 0 0 0 0 230 1111.91 0 36.48 0 0 0 36.48 231 14735.19 67.14 1949.35 0 0 56.14 2072.63 232 3851.73 0 719.53 0 0 0 719.53 233 1677.5 17.2 48.96 41.59 0 0 107.75 234 26345.56 407.42 0 0 23 0 430.45 235 8302.37 0 0 0 68.8 0 68.8 236 5601 0 0 0 0 100.79 100.79 237 1188.54 0 0 0 0 0 0 238 4057.14 0 0 0 0 0 0 239 1250.15 0 36.76 0 0 0 36.76 240 17576.56 563.01 744.28 0 0 0 1307.29 241 5717.77 61.15 1299.8 0 0 0 1360.95 242 1525.41 78.3 0 0 0 0 78.3 243 1408.26 0 0 0 0 92.56 92.56 244 8298.26 155.2 976.04 0 0 0 1131.24 245 2028.31 147.73 0 0 0 0 147.73 246 3406.61 0 0 0 0 0 0 247 2415.08 0 0 12.81 0 0 12.81 248 1384.71 10.74 0 0 0 0 10.74 249 16941.49 138.91 1858.33 0 0 0 1997.24 250 3676.39 0 0 0 0 0 0 251 7457.78 283.05 0 0 0 0 283.05 252 7218.77 337.58 21.14 0 0 0 358.72 253 3280.99 0 0 0 0 0 0 254 3814.52 183.11 851.95 0 0 0 1035.06 255 736.11 0 0 0 0 0 0 256 1439.6 0 0 35.95 0 0 35.95 257 15661.63 262.91 1459.75 0 0 0 1722.66 258 4224.4 0 0 0 0 0 0 259 2343.89 37.67 0 0 0 0 37.67 260 3433 94.78 0 0 0 0 94.78 261 2233.28 0 0 0 0 0 0 262 2277.89 0 0 0 0 0 0 263 10031.01 8.49 1443.56 0 0 0 1452.05 264 3611.22 0 0 0 0 0 0 265 15929.34 169.8 179.51 0 0 0 349.31 266 13264.58 665.45 1206.43 0 0 0 1871.88 267 6196.18 289.58 1268.4 0 0 0 1557.98 268 1328.82 99.65 0 0 0 0 99.65 269 1078.65 19.79 65.62 0 29 0 114.4 270 1367.19 21.35 0 0 0 0 21.35 271 2656.25 0 0 0 0 0 0 272 1101.22 0 0 0 0 0 0 273 1622.57 33.68 31.94 0 0 0 65.62 274 1066.67 0 0 0 0 0 0 275 1528.65 0 0 0 0 0 0 276 1062.85 0 0 0 0 0 0 277 24466.53 127.48 6305.58 0 0 0 6433.06 278 3060.95 0 139.05 0 0 0 139.05 279 1515.7 84.71 0 0 0 0 84.71 280 6741.02 0 0 0 95.7 0 95.65 281 1730.62 0 0 0 0 0 0 282 2241.4 29.68 0 0 0 0 29.68 283 1963.33 16.26 0 0 0 0 16.26 284 3796.6 164.08 242.35 0 0 0 406.43 285 1433.84 0 0 0 0 0 0 286 2083.36 84.31 0 0 0 0 84.31 287 1131.19 0 0 0 0 0 0 288 3415.95 0 222.77 0 0 0 222.77 289 1386.67 0 0 0 19 0 19.01 290 1427.45 38.16 0 0 0 0 38.16 291 5438.69 0 485.59 0 0 0 485.59 292 2202.85 0 0 0 0 0 0 293 1812.16 0 0 0 36.2 0 36.16 294 1753.4 0 0 0 0 0 0 295 1738.58 0 0 0 0 0 0 296 1893.52 0 0 0 97.8 0 97.78 297 1228.71 0 0 0 0 0 0 298 2526.94 22.52 0 536.61 0 0 559.13 299 1576.31 0 0 0 196 0 195.58 300 2033.71 0 0 0 0 0 0 301 17448 0 0 0 0 0 0 302 2397.07 0 0 0 0 0 0 303 3455.51 0 0 0 0 0 0 304 3492.09 0 0 0 0 0 0 305 2794.77 20.16 52.04 0 0 0 72.2 306 2028.7 0 0 0 0 0 0 307 3883.37 14.95 5.61 0 12.5 0 33.07 308 2125.93 46.3 0 10.21 0 0 56.51 309 2405.13 0 0 0 10.8 0 10.78 310 3704.95 0 0 0 0 0 0 311 2827.81 24.87 210.34 0 0 0 235.21 312 2627.25 67.13 8.48 0 0 0 75.61 313 7132.28 125.56 0 0 0 0 125.56 314 6895.03 52.09 0 0 0 0 52.09 315 9136.27 144.58 330.68 407.04 0 0 882.3 316 10899.07 91.14 0 0 0 0 91.14 317 5837.12 0 0 65.52 7.78 0 73.3 318 11114.51 76.57 0 0 0 0 76.57 319 5569.73 0 0 0 0 0 0 320 2295 18.42 9.56 0 0 0 27.98 321 3361.84 0 18.47 0 0 12.1 30.57 322 2185.03 29.12 0 0 0 0 29.12 323 1429.6 38 0 0 0 0 38 324 1084.46 0 0 0 0 105.28 105.28 325 2846.46 18.04 0 0 0 0 18.04 326 2583.04 0 0 76.85 0 62.19 139.04 327 14527.15 0 0 0 0 0 0 328 12886.03 0 32.86 5.43 0 8.14 46.43 329 15145.03 0 0 0 0 0 0 330 8000.58 220.85 0 0 0 0 220.85 331 3195.39 0 0 0 0 0 0 332 23728.6 21.82 0 0 0 0 21.82 333 3209.76 53.66 429.07 0 0 0 482.73 334 1492.52 0 0 0 0 0 0 335 15487.46 28.04 630.61 17.37 12.8 0 688.85 336 6644.93 0 26.22 34.88 0 55.53 116.63 337 10307.89 0 0 0 0 0 0 338 2462,98 0 0 0 0 0 0 339 2445.61 0 0 0 0 45.8 45.8 340 630.41 0 7.22 0 0 0 7.22 341 2921.73 0 0 0 0 0 0 342 2011.35 3.62 0 0 0 0 3.62 343 5620.58 122.69 0 0 0 0 122.69 344 2909.48 0 0 0 0 0 0 345 11213.08 14.01 0 0 0 82.58 96.59 346 2008.59 30.64 88.18 0 0 0 118.82 347 2941.99 6.54 0 0 0 0 6.54 348 1534.61 0 5.98 0 0 0 5.98 349 889.86 0 0 0 0 0 0 350 4085.32 0 119.98 0 0 0 119.98 351 7353.44 0 0 0 0 0 0 352 6387.55 0 0 0 0 0 0 353 4438.39 142.23 708.43 0 0 0 850.66 354 5171.63 83.31 41.65 0 0 0 124.96 355 2671.02 39.1 108.39 0 0 0 147.49 356 5675.8 0 12.39 0 0 0 12.39 357 22693.09 169.22 755.69 0 39.8 0 964.74 358 5931.23 9.87 0 0 0 0 9.87 359 8291.31 30.53 1.45 0 0 0 31.98 360 14050.66 137.93 1420.69 0 15.5 0 1574.14 361 2519.5 40.59 179.71 0 0 0 220.3 362 7020.13 0 26.42 0 0 0 26.42 363 6096.86 19.92 0 0 0 0 19.92 364 3969.18 98.1 820.96 0 0 0 919.06 365 3599.58 4.2 0 0 0 0 4.2 366 1690.19 28.12 0 0 0 0 28.12 367 3955.35 14.68 0 0 0 0 14.68 368 1286.37 0 0 0 0 0 0 369 27465.43 80.78 563.99 0 293 0 937.3 370 2745.02 98.64 119.46 0 0 0 218.1 37! 3291.18 17.2 138.46 0 0 0 155.66 372 527.21 0 0 0 0 0 0 373 926.61 0 250.65 0 0 0 250.65 374 2889.37 0 6.33 0 0 0 6.33 375 3109.73 85.75 33.71 0 0 0 119.46 376 15710.67 19.43 0 0 0 0 19.43 377 4635.33 47.29 0 47.53 0 0 94.82 378 5360.9 0 0 0 0 0 0 379 8377.45 206.74 7.02 0 0 0 213.76 380 3321.64 0 475.62 0 0 0 475.62 381 5093.52 0 25.05 0 0 0 25.05 382 633.56 0 5.62 0 0 0 5.62 383 11417.47 0 0 0 0 0 0 384 16702.13 10.26 0 0 0 0 10.26 385 7170.29 141.26 0 0 0 0 141.26 386 3811.49 0 0 0 0 0 0 387 16003.61 43.68 0 0 0 0 43.68 388 14396.91 49.22 18.86 0 0 0 68.08 389 5240.79 0 27.62 0 0 0 27.62 390 4314.57 0 0 0 0 0 0 391 8426.97 8.25 0 0 0 0 8.25 392 6478.56 73.39 485.08 0 0 0 558.47 393 1739.99 0 0 0 0 0 0 394 2421.71 19.76 0 33.8 0 0 53.56 395 3437.01 0 0 7.83 0 0 7.83 396 5484.7 16.38 17.62 0 16.4 0 50.37 397 2108.72 0 0 0 0 0 0 398 1200.89 0 38.61 0 0 0 38.61 399 12430.42 14.97 202.47 0 32.1 73.58 323.09 400 2160.97 0 107.78 0 0 0 107.78 401 7514.54 0 0 296.07 0 0 296.07 402 4950.88 68.76 354.22 0 0 0 422.98 403 8720.63 0 0 0 0 0 0 404 3767.98 0 0 0 0 0 0 405 2096.27 21.22 0 0 0 0 21.22 406 1853.24 0 0 0 0 0 0 407 18096.78 91.68 859.87 0 0 0 951.55 408 10388.24 0 491.57 0 0 0 491.57 409 9168.52 0 0 0 0 0 0 410 1415.42 15.89 0 0 3.34 56.12 75.35 411 9326.94 163.76 0 0 0 0 163.76 412 3204.63 0 0 0 0 0 0 413 7418.03 0 0 2523.5 86.5 19.31 2629.33 414 3135.65 2.35 0 0 0 0 2.35 415 11637.89 0 2082.7 0 0 0 2082.7 416 4110.93 251.54 605.27 0 0 0 856.81 417 8673.43 0 0 0 0 177.98 177.98 418 4339.84 4.41 0 0 0 0 4.41 419 5656.41 58.23 0 16.53 0 0 74.76 420 3450.86 10.29 0 0 0 0 10.29 421 11328.72 76.57 171.53 0 0 0 248.1 422 2809.32 3.15 0 0 0 0 3.15 423 2043.7 23.8 3.02 34.77 0 0 61.59 424 1026.57 0 0 0 0 0 0 425 15674.64 83.4 710.22 0 0 0 793.62 426 2118.63 0 0 0 0 0 0 427 4932.63 12.45 52.66 0 0 29.45 94.56 428 3667.2 41.15 242.17 0 0 0 283.32 429 11046.33 3.02 7.55 0 0 0 10.57 430 2727.94 0 28.12 0 0 0 28.12 431 22869.32 134.88 148.85 0 0 0 283.73 432 7084.92 626.83 588.61 0 0 0 1215.44 433 3336.16 0 227.74 0 0 0 227.74 434 5751.04 0 0 0 0 0 0 435 6408.38 23.65 18.27 0 0 0 41.92 436 9984.51 0 0 15.49 0 0 15.49 437 4836.65 12.01 0 0 0 0 12.01 438 7828.91 0 250.52 0 0 0 250.52 439 3884.83 0 0 0 0 0 0 440 4475.82 3.94 218.55 45.14 0 0 267.63 441 3275.85 150.95 243.26 0 0 0 394.21 442 3492.95 0 27.65 19.43 0 0 47.08 443 2397.59 5.85 57.57 0 0 0 63.42 444 3989.91 118.26 7.85 0 0 0 126.11 445 1951.8 0 253.89 0 0 0 253.89 446 2416.63 12.89 0 0 0 378.89 391.78 447 2633.35 0 0 0 0 0 0 448 2206.23 140.29 434.97 0 0 0 575.26 449 2401.72 0 9.88 0 0 0 9.88 450 2645.69 12.34 203.24 0 0 0 215.58 451 5295.19 0 0 62.79 0 0 62.79 452 3004.67 51.08 7.23 0 0 0 58.31 453 2219.84 0 32.39 0 0 0 32.39 454 1138.89 0 274.02 0 0 0 274.02 455 1552.5 23.53 3.67 0 0 0 27.2 456 2888.72 12.51 11.25 0 0 0 23.76 457 1845.11 10.21 284.33 0 0 0 294.54 458 1587.05 0 49.36 0 0 0 49.36 459 5322.72 81.39 185.23 0 0 0 266.62 460 6474.45 142.23 136.41 0 0 0 278.64 461 2332.43 57.4 0 0 0 0 57.4 462 905.65 29.3 3.43 0 0 0 32.73 463 1064.51 22.78 8.05 0 0 0 30.83 464 1127.23 0 0 0 0 0 0 465 10073.56 32.22 0 0 0 0 32.22 466 8923.63 3.45 0 0 0 0 3.45 467 3718.21 11.59 168.49 0 0 0 180.08 468 1738.28 8.41 0 0 0 0 8.41 469 2687.94 0 346.86 0 0 0 346.86 470 5026.96 32.66 0 0 0 0 32.66 471 3285.35 0 0 0 0 0 0 472 1940.74 38.56 96.15 0 0 0 134.71 473 984.1 4.6 0 0 0 0 4.6 474 899.05 0 0 0 0 0 0 475 4761.62 28.08 91.48 0 0 0 119.56 476 2473.52 0 4.11 0 0 0 4.11 477 4074.64 111.02 6.9 0 0 0 117.92 478 1845.49 36.3 87.6 0 0 0 123.9 479 1447.88 0 0 0 0 0 0 480 1698.18 35.15 25.78 0 0 0 60.93 481 12417.98 173.85 219.52 0 0 0 393.37 482 1203.5 0 103.63 0 0 0 103.63 483 1321.26 11.82 15.6 0 0 0 27.42 484 990.43 0 0 0 0 0 0 485 2121.68 33.16 87.37 0 0 0 120.53 486 4950 0 5.38 0 0 0 5.38 487 4685.46 23.94 8.63 0 0 0 32.57 488 7353.43 299.35 1035.33 0 0 0 1334.68 489 4782.75 46.24 22.38 0 0 0 68.62 490 4292.26 67.66 147.31 0 0 0 214.97 491 3500.87 319.1 0 0 0 0 319.1 492 3537.38 8.76 111.88 0 0 0 120.64 493 5054.42 0 0 0 0 0 0 494 1806.62 0 63.25 0 0 0 63.25 495 4877.14 79.29 64.97 0 0 0 144.26 496 2895.79 0 24.04 0 0 0 24.04 497 3348.57 59.28 56.71 0 0 0 115.99 498 2050.88 4.04 15.6 0 0 0 19.64 499 1986.28 31.01 17.62 0 0 0 48.63 500 1909.38 0 176.55 0 0 0 176.55 501 2048.31 30.46 33.95 0 0 0 64.41 502 2351.37 0 90.54 0 0 0 90.54 503 1436.64 16.79 9.86 0 0 0 26.65 504 2057.86 0 402.06 0 0 0 402.06 505 1598.24 4.26 9.98 0 0 0 14.24 506 1307.42 6.6 0 0 0 0 6.6 507 1185.56 0 10.28 0 0 0 10.28 508 1004.77 14.61 60.95 0 0 0 75.56 509 922.76 0 17.99 0 0 0 17.99 510 1315.68 0 223.35 0 0 0 223.35 511 13324.34 0 13.82 0 0 0 13.82 512 5427.04 135.59 111.45 0 0 0 247.04 513 3058.43 16.32 250.92 0 0 0 267.24 514 4309.94 18.06 242.86 0 0 0 260.92 515 3326.41 28.51 0 0 0 0 28.51 516 6897.82 0 78.56 0 0 0 78.56 517 1362.85 36.95 0 0 0 0 36.95 518 1617.3 0 34.8 0 0 0 34.8 519 2109.86 0 346.53 0 0 0 346.53 520 1337.28 9.5 134.12 0 0 0 143.62 521 2970.37 0 86.51 0 0 0 86.51 522 1866.49 11.57 0 0 0 0 11.57 523 2964.49 28.29 581.41 0 0 0 609.7 524 3487.22 39.68 66.67 0 0 0 106.35 525 3190.02 0 44.25 0 0 0 44.25 526 3359.33 14.51 54.84 0 0 0 69.35 527 3348.21 0 0 0 0 0 0 528 4025.39 15.98 55.84 0 0 0 71.82 529 2697.06 106.53 0 0 0 0 106.53 530 2270.01 7.1 209.44 0 0 0 216.54 531 3236.69 6.8 0 0 0 0 6.8 532 2332.8 7.94 115.84 0 0 0 123.78 533 2091.26 65.85 43.76 0 0 0 109.61 534 2232.66 11.77 0 0 0 0 11.77 535 2767.5 18.37 22.08 0 0 0 40.45 536 1580.37 139.96 39.01 0 0 0 178.97 537 2080.78 10.1 109.86 0 0 0 119.96 538 1813 0 207.66 0 0 0 207.66 539 1457.55 18.37 95.16 0 0 0 113.53 540 1359.91 0 0 0 0 0 0 541 3504.01 0 90.41 0 0 0 90.41 542 3989.52 4.95 814.75 0 0 0 819.7 543 3569.24 0 29.11 0 0 0 29.11 544 3563.59 0 0 160.59 0 105.66 266.25 545 4699.95 22.94 596.49 0 0 0 619.43 546 4661.02 0 25.68 0 0 0 25.68 547 2058.26 21.68 0 0 0 0 21.68 548 5511.63 75.42 627.05 0 0 0 702.47 549 5131.3 393.05 93.15 0 0 0 486.2 550 3583.52 0 614.75 0 0 0 614.75 551 3250.51 40.26 281.82 86.52 0 0 408.6 552 3121.8 89.55 32.32 0 0 0 121.87 553 2856.24 39.32 100.45 0 0 0 139.77 554 2542.39 27.41 0 0 0 0 27.41 555 3174.84 0 369.48 0 0 0 369.48 556 937.98 8.28 34.55 0 0 0 42.83 557 1837.47 83.79 279.96 0 0 0 363.75 558 2544.17 0 0 0 0 0 0 559 2876.62 8.23 0 0 0 28.82 37.05 560 1822.23 17.09 0 0 0 0 17.09 561 1478.15 46.57 57.3 0 0 0 103.87 562 1056.64 0 4.67 0 0 0 4.67 563 1965.54 8.39 12.12 0 0 0 20.51 564 1987.16 19 105.88 0 0 0 124.88 565 1315.74 32.23 45.78 0 0 0 78.01 566 2129.95 6.38 180.78 0 0 0 187.16 567 1492.94 6.85 9.81 0 0 0 16.66 568 1312.79 0 0 0 0 0 0 569 654.29 0 9.97 0 0 0 9.97 570 1028.62 0 23.36 0 0 0 23.36 571 3452.7 0 66.96 0 0 0 66.96 572 3026.19 6.76 118.8 0 0 0 125.56 573 2685.45 0 11.19 0 0 0 11.19 574 1764.09 0 0 0 0 0 0 575 2175.1 42.81 36.44 0 0 0 79.25 576 2257.94 61.5 63.53 0 0 0 125.03 577 1738.21 12.12 50.48 0 0 0 62.6 578 1192.42 0 107.75 0 0 0 107.75 579 1636.79 0 392.17 0 0 0 392.17 580 3777.42 0 0 0 0 0 0 581 1711.63 11.98 96.88 0 0 0 108.86 582 2287.47 11.98 96.88 0 0 0 108.86 583 2905.56 0 65.8 0 0 0 65.8 584 1760.76 0 14.06 0 0 0 14.06 585 2552.78 47.61 185.76 0 0 0 233.37 586 1788.07 0 0 0 0 0 0 587 1583.51 0 282.12 0 0 0 282.12 588 1501.91 45.14 15.8 0 0 0 60.94 589 1406.06 0 0 0 0 0 0 590 1704.92 9.91 0 0 0 0 9.91 591 1402.94 22.38 0 0 0 0 22.38 592 1355.9 33.33 13.89 0 0 0 47.22 593 1405.32 48.07 26.06 0 0 0 74.13 594 1153.94 0 0 0 0 0 0 595 985.69 0 17.61 0 0 0 17.61 596 1123.78 0 0 0 0 0 0 597 1343.4 48.26 0 0 0 0 48.26 598 1242.71 0 15.98 0 0 0 15.98 599 1023.44 26.91 16.84 0 0 0 43.75 600 900.18 11.74 143.67 0 0 0 155.41 Herbivory data from La Sclva.

Leaf Original Hole Feed Margin Feed Skeletonizing Galling Leaf Mining Total Number Leaf Area Area Area Area Area Area Damage Area 1 4814.84 0 516.55 0 0 0 516.55 2 4340.74 255.82 641.58 0 0 0 897.4 3 1493.3 64.66 0 0 0 0 64.66 4 7640.8 233.3 396.12 537.09 0 0 1166.51 5 1929.02 34.37 0 0 0 0 34.37 6 3058.42 24.32 399.58 0 0 0 423.9 7 5727.65 77.98 18.3 0 0 0 96.28 8 6371.82 16.13 693.22 0 0 0 709.35 9 2090.92 n.i 0 0 0 0 11.1 10 4594.37 0 1049.71 0 0 69.91 1119.62 II 4145.12 0 0 0 0 0 0 12 6663.41 243.9 0 0 0 0 243.9 13 6026.52 41.75 106.24 0 0 0 147.99 14 7358.98 14.34 136.62 0 0 0 150.96 15 5648.4 1143.52 0 0 0 0 1143.52 16 3470.54 0 0 0 0 0 0 17 3402.88 0 0 0 0 0 0 18 3274.98 72.33 163 0 0 0 235.33 19 1988.7 0 0 0 0 345.36 345.36 20 2774.29 0 0 0 0 0 0 21 2407.37 35.97 0 0 0 0 35.97 22 2204.1 0 0 0 0 0 0 23 4577.9 74.08 2761.56 0 0 0 2835.64 24 2604.06 13.37 0 0 0 107.83 121.2 25 1332.17 0 0 0 0 0 0 26 3683.74 11.62 0 0 0 0 11.62 27 9830.17 357.46 0 0 0 0 357.46 28 2883.35 0 0 0 0 0 0 29 3246.35 0 0 0 0 0 0 30 2874.01 0 0 0 0 0 0 31 5189.67 1407.09 8.68 0 0 0 1415.77 32 5028.1 121.91 62.4 0 0 0 184.31 33 8038.2 42.24 2245.62 0 0 0 2287.86 34 4470.56 95.73 857.53 0 0 0 953.26 35 5656.18 314.37 105.62 0 0 0 419.99 36 4606.23 222.64 27.55 0 0 0 250.19 37 5005.28 110.75 14.52 0 0 0 125.27 38 9380.19 77.93 1639.62 53.41 0 0 1770.96 39 4263.43 40.09 482.24 0 0 0 522.33 40 6680.87 328.36 109.21 0 0 0 437.57 41 14968.44 82.37 591.38 0 0 0 673.75 42 3798.74 137.55 363 0 0 0 500.55 43 13891.17 1094.57 384.15 88.34 0 0 1567.06 44 6176.04 196.44 19.43 0 0 0 215.87 45 6251.7 163.22 229.24 0 0 0 392.46 46 5312.26 141.18 169.92 47.13 0 0 358.23 47 7063.6 123.38 0 0 0 0 123.38 48 5683.01 338.84 410.37 0 0 0 749.21 49 1873.14 0 38.91 722.22 0 0 761.13 50 7431.22 414.79 513.58 127.38 0 0 1055.75 51 5115.69 80.7 50.18 0 0 0 130.88 52 9347.32 428.22 0 0 0 0 428.22 53 7755.4 567.06 1220.68 0 0 0 1787.74 54 7135.87 410.56 196.53 0 0 0 607.09 55 5635.77 0 0 2743.46 0 0 2743.46 56 9294.36 343.17 2769.18 0 0 0 3112.35 57 5496.35 81.34 0 72.7 0 0 154.04 58 6666.01 0 264.32 33.14 0 0 297.46 59 5675.23 155.21 351.4 0 0 0 506.61 60 5711.35 71.62 42.35 0 0 0 113.97 61 5300.3 18.82 90.39 385.61 0 0 494.82 62 6652.24 98.15 50.62 0 26.18 0 174.95 63 6064.67 208.95 0 0 0 0 208.95 64 2908.3 0 254.22 0 0 0 254.22 65 2749.88 24.39 0 0 0 114.92 139.31 66 3266.67 0 332.89 385.61 0 0 718.5 67 3930.61 33.08 0 25.45 0 0 58.53 68 3497.79 57.03 46.19 396.99 0 0 500.21 69 4851.84 42.21 591.39 972.55 0 0 1606.15 70 5391.39 0 0 0 0 0 0 71 5878.2 372.17 57.68 0 0 0 429.85 72 3406.56 191.8 19.67 52.45 0 0 263.92 73 4116.21 46.19 15.14 0 0 184.29 245.62 74 5933.4 112.2 0 677.24 0 0 789.44 75 5843.21 475.51 38.54 0 0 0 514.05 76 6580.24 40.01 0 0 0 0 40.01 77 2354.06 0 375.31 0 0 0 375.31 78 6427.64 84.5 88.64 595.86 0 0 769 79 3534.8 63.7 0 0 114.66 0 178.36 80 3177.49 0 0 0 0 0 0 81 2066.54 17.57 0 179.66 0 0 197.23 82 1966.17 0 205.71 0 0 0 205.71 83 5214.72 0 0 0 0 0 0 84 5927.88 164.42 68.6 0 0 0 233.02 85 2906 79.78 216.98 0 0 0 296.76 86 2566.6 0 0 48.76 0 0 48.76 87 3327.46 0 394.81 0 21.75 0 416.56 88 1839.68 0 0 0 0 0 0 89 6180.35 47.93 0 0 0 0 47.93 90 2954.61 0 0 297.72 0 0 297.72 91 3038.88 43.1 0 0 0 0 43.1 92 3843.59 0 13.64 0 0 0 13.64 93 3299.7 143.54 487.88 0 0 0 631.42 94 3442.82 77.89 154.81 0 0 0 232.7 95 3969.33 0 0 0 0 0 0 96 2256.85 40.62 132.58 375.84 0 0 549.04 97 2965.62 0 0 0 6.39 0 6.39 98 14662.27 33.76 850.43 0 0 0 884.19 99 2062.6 27.4 175 8 0 0 210.4 100 2228.2 166 34.8 90.6 0 0 291.4 101 6522 384 197.2 240.4 0 0 821.6 102 4335.97 28.31 0 0 0 0 28.31 103 13051.33 367.14 168.6 0 0 0 535.74 104 4320.37 749.69 0 25.08 0 0 774.77 105 4264.67 57.35 101.25 0 0 0 158.6 106 3541.13 0 0 0 0 0 0 107 2450.8 0 21.91 0 0 0 21.91 108 2002.31 32.99 16.58 757.04 0 0 806.61 109 5589.57 0 111.54 0 0 0 111.54 110 6924.65 0 218.18 0 0 0 218.18 III 3563.25 0 527.36 0 0 0 527.36 112 5933.3 134.76 302.19 0 0 0 436.95 113 7978.78 0 97.63 0 0 0 97.63 114 4787.02 0 0 0 0 0 0 115 6958.8 0 605.75 0 0 0 605.75 116 4438.48 0 0 251.72 0 0 251.72 117 4045.19 0 0 0 0 0 0 118 998.26 0 307.97 0 0 0 307.97 119 1410.82 0 0 11.43 0 0 11.43 120 1804.96 0 9.33 0 0 0 9.33 121 4507.07 208.79 111.55 0 44.13 0 364.47 122 7802.78 196.58 438.25 0 0 0 634.83 123 3989 50.86 933.79 0 0 0 984.65 124 6421.15 142.95 147.01 26.28 0 0 316.24 125 3301.38 0 0 37.59 0 0 37.59 126 3921.37 50.21 0 36.97 0 0 87.18 127 7898.85 229.06 479.71 0 28.79 0 737.56 128 4335.98 0 0 52.88 0 0 52.88 129 2651.34 0 20.1 411.97 0 0 432.07 130 2986.25 0 0 203.73 0 0 203.73 131 2129.81 0 0 73.19 0 0 73.19 132 5119.34 74.65 31.68 0 0 0 106.33 133 5501.556 64.05 0 0 0 0 64.05 134 3177.52 0 75.96 0 0 0 75.96 135 1833.73 0 0 0 0 0 0 136 2944.64 0 57.37 0 0 0 57.37 137 4629.38 0 111.85 0 0 0 111.85 138 2881.24 0 165.5 0 0 0 165.5 139 3317.73 8.49 0 0 0 0 8.49 140 4188.64 0 105.48 100.93 0 0 206.41 141 5989.33 188.43 274.76 125.04 0 0 588.23 142 3613.49 0 25.33 0 0 0 25.33 143 3518.11 20.12 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0 459.46 170 3769.89 18.17 289.61 0 0 0 307.78 171 6998.74 60.28 267.53 356.69 0 0 684.5 172 2715.89 18.58 0 253.9 0 0 272.48 173 5137.44 210.52 281.4 0 0 0 491.92 174 7433.41 99.37 275.54 165.26 0 0 540.17 175 6143.19 0 236.56 361.98 0 0 598.54 176 5122.83 137.69 18.38 113.55 0 0 269.62 177 4234.4 37.82 0 323.96 0 0 361.78 178 8979.32 186.38 361 219.89 0 0 767.27 179 8368.22 279.13 108.86 0 0 0 387.99 180 2648.29 38.78 198.34 0 0 0 237.12 181 5176.59 70.82 451.52 426.15 0 0 948.49 182 5726.05 0 0 598.83 0 0 598.83 183 6364.26 91.02 0 0 0 0 91.02 184 6703.46 0 0 1126.56 0 0 1126.56 185 3904.73 25.33 138 242.15 0 0 405.48 186 3696.79 34.97 12.85 203.4 0 0 251.22 187 4201.51 7.37 0 362.76 0 0 370.13 188 4968.81 0 0 457.28 0 0 457.28 189 3215.57 84.5 647.3 0 0 0 731.8 190 3994.78 206.71 459.09 0 0 0 665.8 191 2489.84 106.06 22.55 0 0 0 128.61 192 6118.36 96.37 29.64 102.14 0 0 228.15 193 5248.78 264.41 40.1 0 0 0 304.51 194 4794.27 461.46 278.12 0 0 0 739.58 195 3560.45 0 0 674.56 0 0 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477.25 518 6718 89.5 453.5 0 0 0 543 519 4352.73 62.25 1437.17 0 0 0 1499.42 520 2520.41 11.64 0 0 0 0 11.64 52! 5431.73 581.65 88.38 0 0 0 670.03 522 6754.68 0 796.73 0 0 0 796.73 523 16805.66 762.49 925.32 130.82 0 0 1818.63 524 1052.59 82.57 0 10.66 0 0 93.23 525 668.73 13.94 16.95 0 0 0 30.89 526 1678.67 13.12 47.84 0 0 0 60.96 527 2626 23.79 0 0 0 0 23.79 528 1557.55 0 0 0 0 0 0 529 5805.9 411.21 1123.95 0 0 0 1535.16 530 4725.7 450 0 0 0 0 450 531 3636.48 41.56 410.1 0 0 0 451.66 532 4048.77 0 0 0 0 0 0 533 1802.65 0 613.06 0 0 0 613.06 534 5267.38 26.69 0 0 0 0 26.69 535 2327.07 0 444.42 0 0 0 444.42 536 4457.36 64.45 340.18 0 0 0 404.63 537 4311.19 361.72 0 0 0 0 361.72 538 5901.1 163.02 106.93 0 0 0 269.95 539 3777.62 36.65 41.47 0 0 0 78.12 540 5210.91 48.39 592.82 0 0 0 641.21 541 3471.72 156.29 0 0 0 0 156.29 542 1475.19 0 63.56 0 0 0 63.56 543 2789.15 45 53.99 0 0 0 98.99 544 2994.36 0 82.87 0 0 0 82.87 545 3030.64 324.64 242.78 0 0 0 567.42 546 1941.5 0 1941.5 0 0 0 1941.5 547 6168.82 87.08 46.3 0 87.37 0 220.75 548 4672.99 217.5 290.84 0 0 0 508.34 549 2675.58 16.32 0 0 0 0 16.32 550 3727.96 114.31 197.86 0 0 0 312.17 551 9433.75 679.47 136.21 661.9 0 0 1477.58 552 2594.08 129.15 9.62 74.56 0 0 213.33 553 5391.27 390.54 631.66 877.07 0 0 1899.27 554 3958.73 230.92 27.52 0 0 0 258.44 555 8798.97 487.66 101.37 0 0 0 589.03 556 11041.48 821.28 69.09 0 0 0 890.37 557 2843.4 0 0 232.87 0 0 232.87 558 6469.1 16.13 105.63 0 0 0 121.76 559 2542.71 25.52 269.62 0 0 0 295.14 560 3586.46 0 529.68 0 0 0 529.68 561 9559.9 12.15 538.02 0 0 0 550.17 562 5077.07 135.66 540.48 0 0 0 676.14 563 6937.11 128.79 655.24 0 0 491.5 1275.53 564 5144.49 112 0 0 0 0 112 565 5626.91 158.52 226.76 0 0 0 385.28 566 2739.56 0 0 0 0 0 0 567 8282.12 1083.84 968.06 0 0 0 2051.9 568 6035.76 91.31 639.93 0 0 0 731.24 569 4145.54 0 27.52 0 0 0 27.52 570 6723.09 173.96 78.29 0 0 0 252.25 571 5284.29 457.37 154.93 0 0 0 612.3 572 6844.92 72.65 0 0 0 1036.11 1108.76 573 4885.74 277.24 88.89 0 0 0 366.13 574 6067.83 20.18 24.37 0 0 89.29 133.84 575 8084.21 0 0 0 0 0 0 576 6220.63 188.55 193.96 0 0 0 382.51 577 6829.65 210.33 1423.58 0 0 0 1633.91 578 10063.43 320.78 1149.58 0 0 0 1470.36 579 1905.85 97.1 19.48 0 0 0 116.58 580 3177.63 438.34 700.12 0 0 0 1138.46 581 2464.25 0 14.75 0 0 0 14.75 582 5394.26 0 0 0 0 0 0 583 9831.25 185 141 0 0 0 326 584 7356.42 216.42 12.18 0 0 0 228.6 585 7281.85 296.79 0 0 29.83 0 326.62 586 5384.42 58.34 0 0 0 0 58.34 587 3252.16 127.15 592.9 0 0 0 720.05 588 4953.98 236.27 197.23 0 0 0 433.5 589 6807.72 300.93 371.29 0 0 0 672.22 590 8048.77 408.32 136.11 0 0 0 544.43 591 2740.74 0 87.96 0 100 0 187.96 592 3916.48 22.06 45.14 0 0 0 67.2 593 7066.36 191.66 42.28 0 0 0 233.94 594 3555.31 90.96 315.07 735.74 0 0 1141.77 595 10525.47 350.68 2877.61 18.05 0 0 3246.34 596 7533.51 330.82 199.03 0 0 0 529.85 597 3684.8 254.84 0 0 0 0 254.84 598 6137.81 383.32 898.39 0 0 0 1281.71 599 7603.48 66.33 0 75.35 0 0 141.68 600 2461.09 0 52.19 0 0 0 52.19 601 4285.69 69.5 34.63 0 0 0 104.13 602 9709.25 1538.5 51 0 0 0 1589.5 603 5079 32 369 0 0 0 401 604 3800 40.25 23.5 0 0 0 63.75 605 1645.75 33 141.75 0 0 0 174.75 606 3655 87 133.25 0 0 0 220.25 607 4108 57.25 391.5 0 0 0 448.75 608 1899.22 0 33.65 0 0 0 33.65 609 9233.35 516.87 87.36 0 0 0 604.23 610 9457.98 785.44 479.79 0 0 0 1265.23 611 3706.67 0 261.17 0 0 0 261.17 612 5715.71 28.61 94.36 0 0 0 122.97 613 2328.35 0 0 0 0 0 0 614 2704.59 0 431.33 0 0 0 431.33 615 2432.58 184.05 27.03 0 0 0 211.08 616 5724.45 29.12 1618.6 0 0 0 1647.72 617 1582.41 0 0 0 0 0 0 618 1521.42 20.25 220.23 0 0 0 240.48 619 2450.77 56.05 28.89 0 0 0 84.94 620 1451.8 0 0 0 0 0 0 621 4077.62 7.16 747.14 99.25 0 0 853.55 622 3035.69 0 0 0 0 0 0 623 1934.25 0 314.81 0 0 0 314.81 624 4096.39 224.68 0 0 0 0 224.68 625 3574.92 144.44 165.17 0 0 0 309.61 626 8261.75 52.75 867.5 0 0 0 920.25 627 4459.33 0 30.86 0 0 0 30.86 628 3158.25 28.75 347 0 0 0 375.75 629 11177.95 1260.95 1736.11 0 0 0 2997.06 630 5462.25 435.75 94 0 0 0 529.75 631 10487 133.75 20 0 0 0 153.75 632 7073.75 123.75 30.25 0 0 0 154 633 6145.88 148.79 108.07 0 0 0 256.86 634 6155.16 23.2 0 89.17 0 0 112.37 635 5884.55 680.24 875.93 0 0 0 1556.17 636 3118.64 0 195.18 0 0 0 195.18 637 1153.75 0 0 0 0 0 0 638 6583,51 524.67 54.17 0 0 0 578.84 639 3510.07 10.41 989.93 0 0 0 1000.34 640 7702.43 144.1 2530.56 0 0 0 2674.66 641 6066.91 96.13 583.2 0 0 0 679.33 642 10442.95 1008.32 610.36 85.61 0 0 1704.29 643 2317.9 0 217.97 0 0 0 217.97 644 3319.8 24.84 43.83 0 0 0 68.67 645 3610.5 115.24 0 0 0 0 115.24 646 2842.16 94.08 141.72 0 0 0 235.8 647 2690.09 15.53 330.32 0 0 0 345.85 648 6837.88 0 468.11 0 0 0 468.11 649 3851.98 0 0 0 0 0 0 650 4471.12 298.49 197.1 0 0 0 495.59 651 3691.41 333.14 60.95 0 0 0 394.09 652 11986.34 57.04 48.94 0 0 0 105.98 653 7579.96 259.71 571.36 0 0 0 831.07 654 2174.67 310.77 0 0 0 0 310.77 655 1756.73 224.9 555.75 0 0 0 780.65 656 2454.49 0 0 0 0 0 0 657 3443.06 0 117.8 0 0 0 117.8 658 3970.28 109.29 274.57 0 0 0 383.86 659 4674.97 294.96 156.78 0 0 0 451.74 660 2326.83 81.89 61.49 0 0 0 143.38 661 8469.27 344.65 0 0 0 0 344.65 662 6247.19 325.71 251.74 0 0 0 577.45 663 2352.58 0 40.07 0 0 0 40.07 664 4636.31 52.05 334.84 0 0 0 386.89 665 3788.27 341.98 119.76 0 0 0 461.74 666 6006.17 439.82 534.57 0 0 0 974.39 667 7322.53 310.17 154.32 0 0 0 464.49 668 2766.67 0 0 0 0 0 0 669 2252.47 0 0 0 0 0 0 670 4970.37 0 245.68 0 0 0 245.68 671 3321.17 249.36 574.77 0 0 0 824.13 672 4823.68 208.39 16.29 0 0 0 224.68 673 2953.15 0 0 0 0 0 0 674 14397.25 453.86 33.82 0 0 0 487.68 675 5422.16 0 393.07 0 0 0 393.07 676 7134.07 218.27 60.11 0 0 0 278.38 677 4956.23 0 0 0 0 0 0 678 3026.59 19.67 1152.35 0 0 0 1172.02 679 8443.8 897.51 725.2 0 0 0 1622.71 680 5823.35 53.52 24.59 0 0 0 78.11 681 1878.31 0 74.58 0 0 0 74.58 682 3790.74 439.66 477.06 0 0 0 916.72 683 3803.51 0 0 0 0 0 0 684 2673.55 55.79 49.79 0 0 0 105.58 685 2298.56 108.66 144.89 0 0 0 253.55 686 1926.22 17.89 0 0 0 0 17.89 687 34122.93 1621.32 866.19 0 0 0 2487.51 688 11703.88 0 3100 0 0 0 3100 689 1612.19 10.53 56.23 0 0 0 66.76 690 2034.88 74.69 96.61 0 0 0 171.3 691 2136.73 150.31 320.67 0 0 0 470.98 692 2639.2 0 194.44 0 0 0 194.44 693 2135.19 0 0 0 0 0 0 694 3871.91 0 20.68 0 0 0 20.68 695 3654.32 0 0 0 0 0 0 696 5723.82 0 206.37 0 0 0 206.37 697 3075.55 262.73 672.05 0 0 0 934.78 698 1448.4 33.15 0 0 0 0 33.15 699 2291.88 0 0 0 0 0 0 700 1916.37 0 0 0 0 0 0 701 3099.38 0 0 0 0 0 0 702 4301.71 0 277.56 0 0 0 277.56 703 4029.49 137.95 243.9 0 0 0 381.85 704 2289.69 0 270.31 0 0 0 270.31 705 3936.19 48.1 975.6 0 0 0 1023.7 706 6415.22 60.88 712.6 0 0 0 773.48 707 4491.42 272.74 428.3 0 0 0 701.04 708 3727.47 88.58 146.61 0 0 0 235.19 709 1159.38 0 0 0 0 0 0 710 1496.01 0 0 0 0 0 0 711 2630.9 0 477.61 76.91 0 0 554.52 712 2013.72 99.31 255.38 0 0 0 354.69 713 7936.3 87.07 35.06 0 0 0 122.13 714 1632.81 0 506.26 0 0 0 506.26 715 3757.64 38.37 137.15 0 0 0 175.52 716 3191.49 0 23.09 0 45.49 0 68.58 717 6534.11 370.79 408.76 0 0 0 779.55 718 6595.18 141.42 204.24 0 0 0 345.66 719 1900.17 85.92 21.48 0 0 0 107.4 720 1454.51 0 392.08 0 0 0 392.08 721 3609.49 0 334.55 0 0 0 334.55 722 4211.94 0 1080.95 0 0 308.63 1389.58 723 1187.61 0 176.3 0 0 0 176.3 724 3508.26 0 1087.12 0 0 0 1087.12 725 1532.9 0 50.86 0 0 0 50.86 726 2840.63 0 45.92 0 0 0 45.92 727 8393.01 0 50.86 0 0 0 50.86 728 4143.05 0 280.98 0 0 0 280.98 729 2418.76 0 77.37 0 0 0 77.37 730 2958.73 0 132.87 0 0 0 132.87 731 3098.98 0 313.04 0 0 0 313.04 732 3377.85 414.48 48.39 0 0 0 462.87 733 1567.94 147.32 0 0 0 0 147.32 734 5577.34 0 65.35 0 0 0 65.35 735 7342.95 0 159.67 0 0 0 159.67 736 9225.3 0 0 0 0 0 0 737 4722.73 0 0 0 0 0 0 738 3268.18 0 0 0 0 0 0 739 4299.17 114.4 401.67 0 0 0 516.07 740 7916.34 0 93.35 0 0 0 93.35 741 3596.95 56.79 837.95 0 0 0 894.74 742 12459.56 1531.02 2405.82 0 0 0 3936.84 743 1439.99 0 67.8 0 0 0 67.8 744 3393.47 0 0 0 0 0 0 745 2226.4 0 76.26 0 0 0 76.26 746 3211.25 408.72 73.74 0 0 0 482.46 747 3836.15 93.99 172.22 0 0 0 266.21 748 15694.2 135.33 343.39 0 0 0 478.72 749 8556.57 56.87 1445.47 0 0 0 1502.34 750 10930.98 70.57 526.99 0 0 0 597.56 751 5770.81 170.23 163.42 0 0 0 333.65 752 3044.4 73.47 46.76 0 0 0 120.23 753 4691 0 0 0 0 0 0 754 3633.8 0 0 0 0 0 0 755 6646.91 352.84 235.17 0 0 0 588.01 756 4129.15 0 293.17 0 0 0 293.17 757 7096.97 87.69 430.37 0 0 0 518.06 758 8560.95 181 459.86 0 0 0 640.86 759 5960.92 0 75.46 0 0 0 75.46 760 5790.05 1929.64 312.77 0 0 0 2242.41 761 4921.46 93.11 410.36 0 0 0 503.47 762 6943.81 363.54 92.91 0 0 0 456.45 763 20331.58 3422.22 2355.06 0 0 0 5777.28 764 3538.34 0 0 0 0 0 0 765 4835.41 97.09 382.11 0 0 0 479.2 766 5183.71 479.78 779.76 0 0 0 1259.54 767 7216.47 400.26 355.13 0 0 0 755.39 768 11041.11 336.28 498.94 0 0 5076.21 5911.43 769 4999.98 7.84 191.82 0 0 175.51 375.17 770 13294.89 63.22 1053.2 0 0 0 1116.42 771 6433.08 0 316.28 0 0 0 316.28 772 3167.53 0 276.28 0 0 0 276.28 773 6998.99 12.35 127.4 0 0 0 139.75 774 4524.52 314.05 404.68 0 0 0 718.73 775 9670.56 0 35.43 0 0 0 35.43 776 7004.39 165.73 247.1 51.04 0 0 463.87 Ill 4497.95 740.99 384.61 0 0 0 1125.6 778 8583.63 746.2 109.05 0 0 0 855.25 779 4228.94 0 27.02 0 0 0 27.02 780 14736.27 0 445.26 0 0 0 445.26 781 18178.57 265.72 471.99 0 0 634.11 1371.82 782 3781.27 426.36 191.26 0 0 0 617.62 783 5974.19 150.81 128.16 0 0 0 278.97 784 27709.81 993.82 0 0 0 0 993.82 785 4808.28 427.56 1665.86 0 0 0 2093.42 786 11583.67 6.12 45.66 0 0 0 51.78 787 8140.74 64.19 369.14 0 0 0 433.33 788 5833.02 347.55 91.05 0 0 0 438.6 789 5585.19 112.66 122.22 0 0 0 234.88 790 3700.62 30.25 472.84 0 0 0 503.09 791 25686.5 0 1885.25 0 0 305.5 2190.75 792 3658.06 12.28 30.76 0 0 0 43.04 793 3464.25 107 14.25 0 0 0 121.25 794 4071 0 0 0 0 0 0 795 12515.97 0 1535.96 0 0 0 1535.96 796 7896.56 71.97 582.57 0 0 0 654.54 797 4987.85 54.2 212.06 0 0 0 266.26 798 4320.87 0 0 0 0 0 0 799 5982.39 178.72 0 0 0 0 178.72 800 2736.85 202.69 87.47 0 0 0 290.16 801 1387.65 0 37.7 13.57 0 0 51.27 802 8651.49 0 0 0 0 0 0 803 4984.92 0 18.04 0 0 73.02 91.06 804 6794.44 0 204.01 0 0 0 204.01 805 8562.96 130.56 70.06 0 0 0 200.62 806 2162.66 0 107.21 0 0 0 107.21 807 6413.89 73.15 36.42 0 0 0 109.57 808 3774.07 250.62 0 0 48.77 179.63 479.02 809 10585.8 0 44.44 0 0 0 44.44 810 7802.3 31.76 579.83 302.55 0 0 914.14 811 4979.37 42.31 1141.8 0 0 0 1184.11 812 1895.5 0 46.88 0 0 0 46.88 813 12261.21 481.86 163.77 0 0 0 645.63 814 3216.27 0 892.59 0 0 0 892.59 815 6206.08 188.59 165.77 0 0 0 354.36 816 8539.78 29.99 221.24 0 0 0 251.23 817 3861.56 5.16 431.99 0 0 0 437.15 818 2752.35 410.34 110.79 0 0 0 521.13 819 4635.9 131.95 197.42 0 0 0 329.37 820 3393.18 0 490.43 0 0 0 490.43 821 3106.47 0 163.08 0 0 0 163.08 822 2403.32 138.08 20.23 0 0 0 158.31 823 3950.77 85.88 1050.92 0 0 0 1136.8 824 9082.71 45.58 2303.32 0 0 0 2348.9 825 16080.17 0 964.08 0 0 0 964.08 826 5173.26 183.82 64.44 0 0 0 248.26 827 5346.74 94.28 310.76 0 0 0 405.04 828 5465.83 30.03 1056.51 0 0 0 1086.54 829 4373.82 0 61.98 0 0 0 61.98 830 3475.89 146.15 122.63 0 0 0 268.78 831 3922.78 184.61 74.7 0 0 0 259.31 832 2906.85 348.6 177.04 35.7 0 0 561.34 833 4015.6 222.03 163.69 0 0 0 385.72 834 3425.38 0 381.1 0 0 0 381.1 835 4790.42 304.92 205.5 0 0 0 510.42 836 5952.75 14.5 1146.5 0 0 0 1161 837 6789.45 442.86 366.78 0 0 0 809.64 838 3973.3 0 188.51 0 0 0 188.51 839 3825.03 221.43 31.21 0 0 0 252.64 840 7647.5 664.25 544.5 0 0 0 1208.75 841 3729.92 0 0 0 0 0 0 842 3116.35 137.06 143.14 34.88 0 0 315.08 843 10323.37 27.66 96.13 0 0 0 123.79 844 7776.57 161.41 134.98 0 0 0 296.39 845 8467.47 702.55 120.12 0 0 0 822.67 846 4150.59 0 103.51 0 0 0 103.51 847 2434.54 241.68 36.68 0 0 0 278.36 848 2678.93 0 98.12 0 0 0 98.12 849 1728.84 0 610.78 0 0 0 610.78 850 1459.72 0 26.26 0 0 0 26.26 851 7133 235.11 503.44 0 0 0 738.55 852 3562.66 0 0 0 0 0 0 853 3235.53 0 338.68 0 0 0 338.68 854 7825.5 165.68 0 0 0 0 165.68 855 4794.28 0 69.13 0 0 0 69.13 856 2701.73 0 26.52 0 0 0 26.52 857 6151.75 400.57 123.31 0 0 0 523.88 858 2636.46 31.07 329.77 0 54.68 0 415.52 859 6666.06 382.38 3156.35 0 0 0 3538.73 860 6966.01 112.91 537.24 0 0 0 650.15 861 6523.64 751.58 1046.67 0 0 0 1798.25 862 3003.2 19.75 852.56 0 0 0 872.31 863 4483.08 92.23 50.38 0 0 0 142.61 864 3156.1 40.48 740.13 0 0 0 780.61 865 2673.7 69.28 108.63 0 0 0 177.91 866 7884.88 364.2 819.76 0 0 0 1183.96 867 5289.19 507.24 494.32 0 0 0 1001.56 868 2199.07 44.14 0 0 0 0 44.14 869 8154.32 961.11 542.89 0 0 0 1504 870 5649.38 57.06 494.07 0 0 0 551.13 871 6387.35 435.18 791.37 0 0 0 1226.55 872 2859.88 22.84 711.73 0 0 0 734.57 873 3709.43 0 0 0 0 0 0 874 2874.75 0 0 0 0 0 0 875 3988.55 0 77.32 0 0 0 77.32 876 9714 385.38 134.55 0 0 0 519.93 877 4355.35 0 283.4 0 0 0 283.4 878 4898.09 76.1 513.22 0 0 0 589.32 879 2901.39 0 0 0 0 0 0 880 2591.9 23.47 0 0 0 0 23.47 881 3921.18 146.22 355.56 0 0 0 501.78 882 3417.15 132.9 0 0 0 0 132.9 883 5916.92 76.34 445.28 0 0 0 521.62 884 5965.43 163.4 818.47 0 0 0 981.87 885 5858.64 389.21 790.34 131.19 0 0 1310.74 886 2755.82 0 37.55 0 0 0 37.55 887 5157.02 57.55 1394.42 0 0 0 1451.97 888 2434.51 367.51 267.76 0 0 0 635.27 889 4746.66 0 71.45 0 0 0 71.45 890 2334.19 0 713.28 0 0 0 713.28 891 8380.51 424.63 828.89 0 0 0 1253.52 892 3883.55 661.03 134.29 0 0 0 795.32 893 3493.6 79.25 225.66 0 0 0 304.9! 894 2149.55 0 103.75 0 0 0 103.75 895 5987.46 63.26 0 0 0 0 63.26 896 2632.54 162.4 0 0 0 0 162.4 897 6322.8 547.2 201.67 0 0 0 748.87 898 5245.49 0 550.39 0 0 0 550.39 899 2106.69 0 0 0 0 0 0

lo SO Herbivory data from Palo Verde.

Leaf Original Hole Feed IVfargin Feed Skeletonizing Galling Leaf Mining Total Number Leaf Area Area Area Area Area Area Damage Area I 16393.78 0 1213.14 0 0 0 1213.14 2 17198.27 0 2860.05 0 0 0 2860.05 3 2424.26 0 359.21 0 0 0 359.21 4 2684.47 20.48 0 0 0 0 20.48 5 2018.23 0 35.12 0 0 0 35.12 6 6386 333 952.5 0 0 0 1285.5 7 7257.25 201 174 72.75 0 0 447.75 8 5034.75 0 30.5 0 252.25 0 282.75 9 4220.25 0 0 0 0 0 0 10 2695 47.25 0 41.75 0 0 89 11 3045.25 0 22.25 20.5 0 0 42.75 12 9077 0 1796.75 0 0 0 1796.75 13 9801.5 298.8 733.5 0 0 0 1032.25 14 10372.5 0 309.25 0 0 0 309.25 15 2955.12 0 0 0 0 0 0 16 6107 0 0 0 59.5 0 59.5 17 2200 0 0 0 0 0 0 18 2059.83 9.7 271.19 0 0 0 280.89 19 2721.5 0 0 0 0 0 0 20 2768.42 0 885.87 0 28.81 0 914.68 21 12324.8 189.1 2530.67 0 0 0 2719.8 22 9724.65 75.35 2335.76 0 0 0 2411.11 23 4998.38 0 1285.7 0 0 0 1285.7 24 3242.22 58.71 0 0 0 0 58.71 25 3221.99 0 0 0 0 0 0 26 2889.19 0 297.06 0 0 0 297.06 27 1504.17 0 40.63 0 0 0 40.63 28 13.149.38 327.5 5343.82 0 0 0 5671.29 29 3072.93 140.4 320.77 0 0 0 461.17 30 4122.22 0 0 0 0 0 0 31 2777.87 0 19.04 0 0 0 19.04 32 8927.78 105.6 0 0 0 0 105.56 33 7034.26 64.51 184.26 94.14 0 0 342.91 34 3207.38 0 24.98 0 0 0 24.98 35 10879.14 0 3383.87 0 0 0 3383.87 36 6165.53 0 0 0 0 0 0 37 2849.56 0 90.38 0 0 0 90.38 38 1859.17 0 0 0 0 0 0 39 7235.06 0 120.69 0 392.76 427.2 940.67 40 6964.65 0 0 0 0 0 0 41 2896.01 0 59.47 0 0 0 59.47 42 5069.32 0 0 0 177.14 0 177.14 43 6068.53 33.03 0 0 0 0 33.03 44 5635.28 188.3 0 0 0 0 188.25 45 3879.45 83.47 146.82 0 0 0 230.29 46 2156.65 0 188.85 0 0 0 188.85 47 2893.45 32.12 0 25.53 0 0 57.65 48 6374.95 23.76 1518.51 0 0 0 1542.27 49 2769.38 0 0 0 0 0 0 50 4763.03 14.38 0 0 0 0 14.38 51 2812.21 0 848.79 0 139.15 0 987.94 52 6500.42 53.78 1300.89 0 0 0 1354.67 53 3357.78 0 0 0 0 0 0 54 2738.93 0 0 0 0 0 0 55 4461.87 0 47.83 0 0 55.2 103.03 56 2969.9 0 60.91 0 0 0 60.91 57 6266.18 16.36 0 0 0 0 16.36 58 3817.49 0 42.83 0 0 0 42.83 59 2782.18 0 0 0 0 0 0 60 1361.29 0 0 0 0 0 0 61 3083.4 0 772.16 0 0 0 772.16 62 5883.45 24.11 0 0 0 0 24.11 63 5490.8 71.26 1138.93 0 0 0 1210.19 64 6521.43 0 36.16 0 0 0 36.16 65 2492.89 0 7.45 0 0 0 7.45 66 4501.48 0 0 0 0 0 0 67 2844.05 0 0 0 0 0 0 68 1646.27 0 895.55 0 0 0 895.55 69 2199.11 12.13 0 0 0 0 12.13 70 3052.22 46.6 0 0 0 0 46.6 71 3508.73 30.62 0 0 0 0 30.62 72 2684.91 0 15.53 0 0 0 15.53 73 7197.07 lll.l 0 0 0 0 111.1 74 2915.18 0 0 0 0 0 0 75 4536.16 199.1 673.64 0 0 0 872.69 76 11735.7 116 0 0 0 0 116.02 77 5210.35 0 0 0 0 0 0 78 5016.72 52.96 1352.96 0 0 0 1405.92 79 3283.28 81.8 595.12 0 0 0 676.92 80 7782.34 0 2314.7 31.79 0 0 2346.49 81 7162.62 209.6 1274.94 0 0 0 1484.49 82 3305.4 0 673.42 0 0 0 673.42 83 4379.94 77.47 95.21 0 0 0 172.68 84 3322.06 0 31.17 0 0 0 31.17 85 3755.06 0 0 0 0 0 0 86 6239.6 65.58 2470.71 0 0 0 2536.29 87 1774.81 0 0 0 0 0 0 88 2154.67 0 246.91 58.87 0 0 305.78 89 2056.3 0 0 0 0 0 0 90 1800.49 0 61.43 0 0 0 61.43 91 11351.45 0 885.03 0 0 0 885.03 92 4068.05 0 0 0 0 0 0 93 3614.52 0 0 0 0 0 0 94 3310 128.9 0 0 0 0 128.87 95 5001.32 0 0 0 0 0 0 96 2302.19 0 0 73.92 0 0 73.92 97 2039.92 0 0 0 0 0 0 98 2202.5 0 0 0 0 0 0 99 2056.88 24.81 31.64 0 0 0 56.45 100 1583.99 0 0 0 0 0 0 101 3298.76 0 362.6 0 0 0 362.6 102 2077.72 0 0 0 0 0 0 103 1790.5 0 121.84 0 0 0 121.84 104 8858.43 0 0 0 0 0 0 105 1604.67 0 0 0 0 0 0 106 1186.32 6.31 0 28.67 0 0 34.98 107 2374.58 0 0 0 0 0 0 108 5593.16 132.3 414.72 0 0 0 547.01 109 3986.32 0 184.09 0 0 0 184.09 110 3231.82 12.89 720.05 0 0 0 732.94 111 1611.69 0 35.94 0 0 0 68.08 112 1930.75 0 427.47 0 0 0 427.47 113 2616.22 0 0 6.14 0 0 6.14 114 3203.84 0 132.11 0 158.58 0 290.69 115 2622.92 0 0 0 0 0 0 116 2727.83 10.11 28.18 0 0 0 38.29 117 3437.72 17.94 0 0 0 0 17.94 118 1531.67 0 0 0 0 0 0 119 1647.69 0 0 0 0 0 0 120 2244.84 254.2 63.92 0 0 0 318.15 121 2378.55 24.31 0 0 0 0 24.31 122 5841.27 255.9 1151.02 0 0 0 1406.87 123 2248.44 0 0 0 0 0 0 124 1415.75 0 15.43 0 0 0 15.43 125 1008.92 0 73.08 0 0 0 73.08 126 2065.67 7.48 0 0 0 0 7.48 127 3541.32 0 0 0 0 0 0 128 6743.06 188.2 499.31 120.8 0 0 808.35 129 2522.05 66.32 448.61 0 0 0 514.93 130 3878.47 0 0 0 0 0 0 131 2959.2 26.57 0 0 0 0 26.57 132 2581.12 0 0 0 0 0 0 133 1325.92 0 305.44 0 0 0 305.44 134 6423.84 0 236.64 0 208 0 444.64 135 2115.68 0 28.64 0 0 32 60.64 136 3120.48 0 82.72 0 0 0 82.72 137 5454.24 46.24 0 0 0 0 46.24 138 9358.36 62.53 0 0 0 0 62.53 139 2803.8 0 69.54 0 0 0 69.54 140 4111.32 0 0 0 0 0 0 141 1240.32 0 0 0 0 0 0 142 1633.02 0 10.23 0 0 0 10.23 143 8145.83 381.3 0 0 0 0 381.25 144 3029.17 37.68 159.2 0 0 0 196.88 145 1620.14 0 0 0 0 0 0 146 1403.99 0 0 0 0 70.66 70.66 147 893.92 4.17 112.5 0 0 30.38 147.05 148 1367.36 0 72.39 0 0 0 72.39 149 972.22 0 0 0 0 0 0 150 745.66 0 0 31.42 0 0 31.42 151 8958.75 41.25 870.25 0 0 0 911.5 152 6032.72 677.2 170.99 0 0 0 848.15 153 4044.14 0 500 0 0 0 500 154 2545.68 132.4 0 0 0 0 132.4 155 6516.27 0 117.75 0 0 0 117.75 156 8871.3 63.58 0 0 0 0 63.58 157 2745.68 0 0 0 0 39.51 39.51 158 14339.35 37.28 422.04 0 0 0 459.32 159 2950.5 0 23.72 0 0 0 23.72 160 2358.88 0 0 0 0 0 0 161 5514.58 0 243.2 0 0 0 243.2 162 5746.07 0 0 0 0 0 0 163 5589.64 0 0 0 0 0 0 164 3021.66 0 0 0 0 0 0 165 2041.96 30.92 73.26 0 0 0 104.18 166 5739.37 0 147.05 0 0 0 147.05 167 2890.05 0 891.33 0 0 0 891.33 168 8075.35 0 0 496.5 0 0 496.51 169 4320.98 0 0 0 0 0 0 170 3653.87 0 274.86 0 0 0 274.86 171 4374.7 0 0 0 0 0 0 172 4670.32 0 0 0 0 28.09 28.09 173 3398.02 49.5 0 0 0 0 49.5 174 11648.99 364.9 554.68 0 0 0 919.53 175 6628.01 0 839.38 0 0 0 839.38 176 9218.87 389.4 0 0 0 0 389.38 177 3220.48 0 0 0 0 0 0 178 2392.81 28.06 0 0 0 0 28.06 179 17681.27 174.6 67.77 0 0 0 242.36 180 12066.25 0 0 564.5 0 0 564.54 181 2187.61 0 56.89 0 0 40.2 97.09 182 1828.14 0 53.84 0 0 168.1 221.89 183 2672.76 0 0 0 0 236.3 236.27 184 3963.22 0 0 0 0 0 0 185 7407.32 524.5 39.63 0 0 0 564.16 186 5665.6 0 27.62 1161 0 0 1188.38 187 4811.11 28.22 458.77 0 0 0 486.99 188 4222.03 0 0 0 0 0 0 189 4015.81 0 0 0 0 0 0 190 6269.13 0 0 0 0 0 0 191 3854.75 0 345.46 0 0 0 345.46 192 7809.63 66.76 0 0 0 0 66.76 193 5515.5 322.2 154.61 0 0 0 476.77 194 3510.93 80.6 0 0 0 0 80.6 195 3140.5 0 552.35 0 0 0 552.35 196 3220.73 15.85 0 24.87 0 0 40.72 197 4352.44 0 0 0 0 0 0 198 5540.2 0 0 0 0 0 0 199 3043.36 270.6 432.67 0 0 0 703.3 200 2447.68 0 0 0 0 0 0 201 1422.91 107.1 0 0 0 7.49 114.55 202 1150.63 0 0 0 0 0 0 203 864.66 7.36 0 0 0 0 7.36 204 4064.12 59.89 369.84 0 0 0 429.73 205 1390.06 0 9.17 0 0 0 9.17 206 5227.97 43.82 36.52 0 0 0 80.34 207 1046.27 76.57 105.92 0 0 0 182.49 208 3341.1 26.88 168.26 0 0 0 195.14 209 4237.36 0 768.6 126.8 0 0 895.38 210 405.78 0 0 0 0 0 0 211 3205.79 163.7 842.21 0 0 0 1005.9 212 6157.45 367.4 0 0 0 0 367.42 213 7334.54 0 266.96 521.1 0 0 788.07 214 5282.42 7.11 0 0 0 0 7.11 215 6211.98 0 163.55 0 0 0 163.55 216 3354.45 0 395.09 0 0 0 395.09 217 2811.45 0 0 0 0 0 0 218 3851.38 0 0 0 0 0 0 219 1950.44 0 105.24 0 0 0 105.24 220 2402.99 0 27.02 0 0 0 27.02 221 4100.07 0 82.79 0 0 0 82.79 222 2219.27 20.7 199.01 0 0 0 219.71 223 2222.32 0 258.42 0 0 0 258.42 224 1330.16 58.75 64.23 0 0 0 122.98 225 1858.88 71.22 0 0 0 0 71.22 226 3037.99 0 798 0 0 0 798 227 4341.75 104.4 243.2 0 0 0 347.6 228 4798.93 0 62.4 0 0 0 62.4 229 4640.77 0 1502.8 0 0 0 1502.8 230 3163.59 0 0 0 0 0 0 231 4009.36 0 1732.56 0 0 0 1732.56 232 1850.88 0 0 0 0 0 0 233 2953.75 0 0 0 0 0 0 234 244.4 0 52.15 0 0 0 52.15 235 179.29 0 27.45 0 0 0 27.45 236 268.47 0 69.43 0 0 0 69.43 237 180.8 0 25.87 0 0 0 25.87 238 2724.41 0 0 0 0 0 0 239 3179.25 0 959.34 0 0 0 959.34 240 2017.65 20.14 281.42 0 0 0 301.56 241 2149.59 0 15.8 0 0 0 15.8 242 2422.85 0 0 0 0 0 0 243 1351.7 0 98.78 0 0 0 98.78 244 3877.15 111.5 0 28.64 0 0 140.09 245 3074.48 0 0 0 0 0 0 246 1994.74 608.7 0 0 0 0 608.67 247 3634.2 0 0 0 0 0 0 248 3237.33 220.1 195.83 0 0 0 415.97 249 1110.94 0 407.81 0 0 0 407.81 250 1688.72 0 9.2 0 0 0 9.2 251 2713.72 0 0 0 0 0 0 252 2442.53 0 0 0 0 0 0 253 1985.24 0 166.49 0 0 0 166.49 254 245.49 0 0 0 0 0 0 255 263.89 0 24.13 0 0 0 24.13 256 6425.52 0 0 0 0 0 0 257 2036.11 16.67 0 155.6 0 0 172.23 258 3576.91 0 0 0 0 0 0 259 2998.09 0 41.32 0 0 0 41.32 260 4209.9 0 17.19 0 0 0 17.19 261 2639.52 0 30.72 0 0 0 30.72 262 4905.12 54.88 104 0 0 0 158.88 263 3153.12 51.84 0 0 0 0 51.84 264 2733.28 0 0 0 0 0 0 265 2960.32 20.96 378.56 0 0 0 399.52 266 1560.96 0 0 0 0 0 0 267 1156.16 0 258.56 0 0 0 258.56 268 2340.36 0 0 0 0 35.82 35.82 269 1961.18 0 247.23 0 0 0 247.23 270 3119.2 0 466.81 0 0 0 466.81 271 2999.92 0 26.82 0 0 20.16 46.98 272 2944.44 0 0 0 0 0 0 273 1660.97 0 14.5 0 0 0 14.5 274 1262.64 0 0 22.66 0 13.99 36.65 275 1826.74 107.8 319.87 0 0 0 427.66 276 1778.76 0 0 0 0 0 0 277 4088.24 0 36.08 0 0 0 36.08 278 888.4 0 94.03 0 0 0 94.03 279 881.22 0 0 0 0 0 0 280 1671.5 80.33 0 0 0 0 80.33 281 1326.72 0 0 0 0 0 0 282 1406.22 0 0 0 0 0 0 283 1551.68 0 0 0 0 0 0 284 3292.07 0 0 0 0 0 0 285 323.78 0 9.63 0 0 0 9.63 286 354.25 0 10.44 0 0 0 10.44 287 9022.12 0 1666.59 0 0 0 1666.59 288 6058.5 0 413.79 0 0 0 413.79 289 4767.18 0 81.33 0 0 93.22 174.55 290 8957.91 178.4 0 0 0 0 178.36 291 4782.88 0 42.81 0 0 0 42.81 292 3746.49 0 0 0 0 0 0 293 7327.47 54.7 251.13 0 0 0 305.83 294 6452.32 62.3 3330.32 0 0 0 3392.62 295 3828.3 0 0 0 0 0 0 296 5535.31 0 1196.67 0 0 0 1196.67 297 2442.81 0 0 0 0 0 0 298 3300.83 0 1157.19 0 0 0 1157.19 299 5628.54 0 101.3! 0 0 0 101.31 300 4070.87 0 0 0 0 0 0

C\ o Herbivory data from Green River.

Leaf Original Hole Feed Margin Feed Skeletonizing Galling Leaf Mining Total Number Leaf Area Area Area Area Area Area Damage Area 1 611.55 0 0 0 0 0 0 2 435.82 0 0 Ml 0 0 1.11 3 337.82 0 9.5 0 0 0 9.5 4 273.3 0 0 0 15.78 0 15.78 5 363.81 0 0 0 0 0 0 6 454.41 0 0 0 0 0 0 7 499.73 0 0 0 0 0 0 8 317.28 3.79 0 0 0 0 3.79 9 622.33 0 0 0 0 0 0 10 214.64 0 0 0 0 0 0 II 320.9 0 0 0 0 0 0 12 336.16 0 0 0 0 0 0 13 318.25 0 0 0 0 0 0 14 272.46 0 0 0 0 0 0 15 321.21 0 0 0 0 0 0 16 447.56 11.59 0 0 0 0 11.59 17 143.31 0 0 0 0 0 0 18 314.37 0 0 0 0 0 0 19 333.44 0 0 0 0 0 0 20 729.23 0 26.14 0 0 51.2 77.34 21 205.21 0 0 0 0 0 0 22 190.83 0 0 0 0 0 0 23 588.14 0 0 0 0 0 0 24 224.95 0 0 0 0 0 0 25 477.45 0 0 0 0 0 0 26 229.54 0 0 0 0 0 0 27 223.91 0 0 0 0 0 0 28 401.86 0 0 0 0 0 0 29 417.61 0 0 0 0 0 0 30 493.66 0 0 0 0 0 0 31 333 0 0 0 0 0 0 32 218.42 0 0 0 0 0 0 33 166.13 0 0 0 0 0 0 34 286.36 0 0 0 0 0 0 35 515.04 0 0 0 0 0 0 36 276.07 3.96 0 0 0 0 3.96 37 655.93 0 0 0 0 0 0 38 537.53 0 0 0 0 0 0 39 238.35 0 0 0 0 0 0 40 283.15 0 32.45 0 0 0 32.45 41 432.54 0 0 0 0 0 0 42 377.21 0 0 0 0 0 0 43 312.31 0 0 0 0 0 0 44 86.31 0 0 0 0 0 0 45 333.91 0 0 0 0 0 0 46 74.52 0 0 0 0 0 0 47 103.25 0 0 0 0 0 0 48 122.63 0 0 0 0 0 0 49 146.18 0 0 0 0 0 0 50 460.13 0 0 0 0 0 0 51 296.25 0 0 0 0 0 0 52 352.13 0 0 0 0 0 0 53 898.38 0 0 0 0 0 0 54 570.21 0 0 0 0 0 0 55 1201.92 0 0 0 0 0 0 56 601.81 0 0 0 0 0 0 57 1309.8 0 0 0 0 0 0 58 586.32 0 6.03 0 0 0 6.03 59 2283.26 129.72 0 81.77 0 0 211.49 60 620.95 0 0 0 0 0 0 61 290.16 0 0 0 0 0 0 62 50.15 0 0 0 0 0 0 63 47.03 0.79 0 0 0 0 0.79 64 254.58 0 49.74 0 0 0 49.74 65 33.49 0 0 0 0 0 0 66 52.82 0 0 0 0 0 0 67 154.71 0 0 0 0 0 0 68 52.41 0 0 0 0 0 0 69 156.57 0 0 0 0 0 0 70 193.49 0 0 0 0 0 0 71 30.7 0 0 0 0 0 0 72 364.94 0 0 0 0 0 0 73 169.59 0 0 0 0 0 0 74 62.67 0 0 0 0 0 0 75 535 0 72.93 0 0 0 72.93 76 144.08 0 0 0 0 0 0 77 111.87 0 0 0 0 0 0 78 168.01 0 0 0 0 0 0 79 95 0 0 0 0 0 0 80 110.12 0 3.94 0 0 0 3.94 81 93.15 0 0 0 0 0 0 82 146.82 0 0 0 0 0 0 83 102.93 3.46 11.55 0 0 0 15.01 84 119.13 0 0 0 0 0 0 85 364.94 0 0 0 0 0 0 86 271.42 0 0 0 0 0 0 87 162.76 0 0 0 0 0 0 88 69.97 0 0 0 0 0 0 89 157.47 0 0 0 0 0 0 90 34.15 0 0 0 0 0 0 91 176.71 0 0 0 0 0 0 92 243.92 7.05 0 0 0 0 7.05 93 89.26 0 0 0 0 0 0 94 57.74 0 0 0 0 0 0 95 81.13 0 0 0 0 0 0 96 107.27 0 0 47.21 0 0 47.21 97 120.92 0 0 0 0 0 0 98 307.38 0 0 0 0 0 0 99 110.2 0 0 0 0 0 0 100 378.18 ^ 0 0 0 0 0 0 101 166.45 0 0.84 0 0 0 0.84 102 337.67 0 0 0 0 0 0 103 351.41 _J 0 20.93 0 0 0 20.93 104 111.54 0 0 0 0 0 0 105 119.05 0 0 0 0 0 0 106 113.32 0 0 0 0 0 0 107 76.01 0 0 0 0 0 0 108 219.32 0 0 0 0 24.44 24.44 109 113.98 0 0 0 0 0 0 110 130.73 0 0 0 0 0 0 111 381.91 0 0 0 0 0 0 112 171.53 0 0 0 0 0 0 113 144.24 0 0 0 0 0 0 114 116.34 0 0 0 0 0 0 115 133.44 0 0 0 0 0 0 116 36.74 0 0 0 0 0 0 117 141.77 0 0 0 0 0 0 118 162.82 0 0 0 0 0 0 119 132.74 0 0 0 0 0 0 120 92.95 0 0 0 0 0 0 121 235.98 0 0 0 0 0 0 122 155.36 0 0 0 0 0 0 123 469.05 27.14 0 0 0 0 27.14 124 67.59 0 0 0 0 0 0 125 110.49 0 0 0 0 0 0 126 162.43 0 0 0 0 0 0 127 376.52 0 31.27 0 0 0 31.27 128 93.94 0 0 0 0 0 0 129 52.38 0 0 0 0 0 0 130 376.21 0 0 0 0 0 0 131 240.93 0 2.97 0 0 0 2.97 132 533.6 0 0 0 0 0 0 133 48.53 0 0 0 0 0 0 134 99.21 0 0 0 0 0 0 135 131.3 0 0 0 0 0 0 136 476.87 0 0 0 0 0 0 137 189.45 0 0 0 0 0 0 138 2774.54 31.06 220.88 0 0 0 251.94 139 755.03 0 6.26 0 0 23.43 29.69 140 667.21 0 0 0 0 0 0 141 1242.73 7 0 0 0 0 7 142 1318.27 5.95 0 0 2.48 0 8.43 143 475.86 0 0 0 3.3 0 3.3 144 328.29 0 0 0 0 0 0 145 213.07 2.05 0 0 0 0 2.05 146 1117.81 0 0 0 1.25 0 1.25 147 505.91 8.89 0 0 0 0 8.89 148 1142.77 0 0 0 0 0 0 149 727.02 0 0 0 0 0 0 150 246.4 0 0 0 0 0 0 151 633.4 9.16 0 0 0 0 9.16 152 416.17 1.02 0 0 0 0 1.02 153 205.6 0 0 27.98 0 0 27.98 154 756.47 15.51 0 0 0 0 15.51 155 954.61 0 0 0 0 0 0 156 348.82 0 0 0 0 0 0 157 1279.76 0 0 0 0 0 0 158 681.76 3.66 43.68 0 0 0 47.34 159 105.13 0.78 0 0 0 0 0.78 160 91.81 0 0 0 0 0 0 161 793.69 0 0 0 0 0 0 162 235.4 0 4.56 0 0 0 4.56 163 1576.86 0 0 0 0 0 0 164 1892,89 0 0 0 0 0 0 165 276.35 0 0 119.29 0 0 119.29 166 758.92 0 0 0 0 0 0 167 540.55 0 0 0 0 0 0 168 242.36 6.42 0 0 4.13 0 10.55 169 461.87 0 0 0 0 0 0 170 1111.95 0 0 0 0 0 0 171 319.18 17.11 0 0 0 0 17.11 172 1592.34 0 0 0 0 0 0 173 1503.43 0 0 0 0 0 0 174 802.31 0 0 0 0 0 0 175 1319.26 4.61 0 0 0 0 4.61 176 1107.26 12.68 0 0 0 0 12.68 177 627.88 18.47 0 0 0 0 18.47 178 670.08 0 42.23 0 0 0 42.23 179 59.3 0 0 0 0 0 0 180 167.63 0 0 31.64 0 0 31.64 181 594.27 0 0 0 0 0 0 182 1139.61 61.01 0 0 0 0 61.01 183 238.25 56.13 0.54 0 0 0 56.67 184 1526.39 55.41 34.72 404.77 0 0 494.9 185 1168.98 0 0 0 0 0 0 186 132.94 0 1.46 0 0 0 1.46 187 707.06 0 0 0 0 0 0 188 1140.64 7.06 164.1 0 0 0 171.16 189 1178.73 3.18 21.7 0 0 0 24.88 190 392.88 0 0 0 0 0 0 191 1451.07 27.56 0 0 0 0 27.56 192 1689.29 36.27 0 0 0 0 36.27 193 370.03 2.27 11.49 0 0 0 13.76 194 290.86 0 1.03 0 0 0 1.03 195 2100.75 46.98 0 0 0 0 46.98 196 943.57 77.47 54.36 0 0 0 131.83 197 919.05 0 0 0 0 0 0 198 627.36 3.99 87.85 0 0 0 91.84 199 684.04 0 6.12 0 0 0 6.12 200 294.42 0 0 0 0 0 0 201 236.12 10.44 11.07 0 0 0 21.51 202 176.97 0 0 0 0 0 0 203 2932.27 0 0 0 0 0 0 204 373.97 0.86 0 0 0 0 0.86 205 225.48 0 0 0 0 0 0 206 5034.9 0 0 0 0 0 0 207 3397.02 0 0 0 0 0 0 208 1875.12 0 0 0 0 0 0 209 1057.81 0 0 0 0 0 0 210 5102.15 151.84 0 0 0 0 151.84 211 462.75 5.81 0 0 0 0 5.81 212 2234.18 0 0 0 0 0 0 213 746.25 2.28 0 0 0 0 2.28 214 1097.46 0 0 0 0 0 0 215 242.88 0 0 0 0 0 0 216 103.75 0 0 0 0 0 0 217 80.8 0 0 0 0 0 0 218 434.04 0 0 0 0 0 0 219 563.6 0 0 0 0 0 0 220 653.87 0 0 0 0 0 0 221 738.9 0 0 0 0 0 0 222 320.57 2.57 0 0 0 0 2.57 223 583.58 0 0 0 0 0 0 224 2062.14 3.73 0 0 0 0 3.73 225 1714.84 0 165.2 0 0 0 165.2 226 821.87 0 0 0 0 0 0 221 468.98 0 0 0 0 0 0 228 896.98 27.66 87.87 153.56 0 0 269.09 229 331.99 7.58 0 0 0 0 7.58 230 541.7 27.84 0 0 0 0 27.84 231 90.29 0 0 0 0 0 0 232 940.85 0 65.08 0 0 0 65.08 233 337.24 0 0 0 0 0 0 234 833.12 0 15.51 0 0 0 15.51 235 746.53 0 97.17 0 0 0 97.17 236 742.69 0 0 0 0 0 0 237 819.77 0 0 0 10 0 10 238 4202.35 3.74 0 0 0 0 3.74 239 760.86 0 0 85.48 0 0 85.48 240 592.1 0 0 0 0 0 0 241 921.18 0 0 0 0 0 0 242 384.93 0 0 0 0 0 0 243 491.21 0 0 93.25 0 0 93.25 244 718.44 0 0 0 0 0 0 245 245.74 0.82 10.4 0 0 0 11.22 246 405.23 0 75.93 0 0 0 75.93 247 235.82 15.88 11.2 0 0 0 27.08 248 253.62 0 0 0 0 0 0 249 3295.92 26.84 0 0 0 0 26.84 250 1583.84 17.71 0 0 0 0 17.71 251 374.42 0 6.11 0 0 0 6.11 252 244.68 0 0 0 0 0 0 253 231.03 0 0 0 0 0 0 254 1099.67 0 168.77 0 0 0 168.77 255 118.71 0 0 0 0 0 0 256 670.26 0 0 0 0 0 0 257 446.11 0 0 0 13.58 0 13.58 258 190.85 11.57 0 0 0 0 11.57 259 242.41 0 4.09 0 0 0 4.09 260 218.5 0 0 0 0 0 0 261 937.44 0 0 0 0 0 0 262 677.19 0 0 0 10.81 0 10.81 263 319.57 0 0 0 0 0 0 264 726.88 0 0 0 0 0 0 265 526.3 0 0 0 0 0 0 266 404.3 0 0 0 0 0 0 267 124.26 2.04 0 0 0 0 2.04 268 141.13 0 0 0 0 0 0 269 624.04 12.48 0 0 0 0 12.48 270 141.76 0 0 0 0 0 0 271 380.56 0 0 16.32 0 0 16.32 272 320.24 1.8 22.6 0 0 0 24.4 273 213.65 0 0 0 0 0 0 274 388.91 0 0 0 0 0 0 275 1097.7 0 0 16.94 0 0 16.94 276 306.32 0 5.54 0 0 0 5.54 277 356.06 0.68 0 0 0 0 0.68 278 1039.16 0 0 0 14.42 0 14.42 279 324.47 0 27.38 7.7 0 0 35.08 280 859.74 0 72.02 0 0 0 72.02 281 294.9 0 0 0 0 0 0 282 1128.48 5.82 0 0 0 0 5.82 283 1251.41 0 21.84 0 0 0 21.84 284 2411.66 0 0 0 0 0 0 285 2859.66 0 0 88.8 0 0 88.8 286 2249.03 0 0 0 0 0 0 287 834.5 0 0 0 0 0 0 288 708.98 0 0 0 0 0 0 289 1833.9 0 0 0 0 0 0 290 790.81 0 175.05 0 0 0 175.05 291 2645.08 0 0 0 0 0 0 292 107 0 0 0 0 0 0 293 107.65 0 0 0 0 0 0 294 68.56 0 0 0 0 0 0 295 318.22 0 0 0 0 0 0 296 292.56 8.17 0 0 0 0 8.17 297 360.33 0 0 0 0 0 0 298 57.55 4.64 0.33 0 0 0 4.97 299 283.36 0 0 0 0 0 0 300 307.16 0 0 0 0 0 0 301 320.61 0 0 0 0 0 0 302 266.77 0 0 0 0 0 0 303 495.59 0 0 0 0 0 0 304 260.43 1.65 0 0 0 0 1.65 305 390.29 3.72 0 0 0 0 3.72 306 493.54 0 0 0 0 0 0 307 555.24 11.06 0 0 0 0 11.06 308 247.72 0 0 0 0 0 0 309 615.89 36.28 68.68 0 0 0 104.96 310 207.87 0 0 0 0 0 0 311 199.7 0 5.7 0 0 0 5.7 312 249.01 0 1.18 0 0 0 1.18 313 71.71 0 1.94 0 0 0 1.94 314 336.52 27.35 0 0 0 0 27.35 315 212.92 0 0 0 0 0 0 316 555.65 0 0 0 0 0 0 317 200.87 8.13 43.96 0 0 0 52.09 318 342.83 0 0 0 0 0 0 319 328.26 0 0 0 0 0 0 320 311.78 10.56 0 0 0 0 10.56 321 123.88 1.56 14.94 0 0 0 16.5 322 219.6 0 0 0 0 0 0 323 121.35 0 26.79 0 0 0 26.79 324 182.14 0 1.61 0 0 0 1.61 325 313.87 0 0 0 0 0 0 326 93.63 0 0 0 0 0 0 327 54.27 0 0 0 0 0 0 328 211.06 0 0 0 0 0 0 329 386 0 0 31.56 0 0 31.56 330 1637.44 24.71 0 0 0 0 24.71 331 216.73 0 0 0 0 0 0 332 159.22 1.63 8.92 0 0 0 10.55 333 152.69 0 0 0 0 0 0 334 186.37 0 31.95 0 0 0 31.95 335 254.44 0 0 0 0 0 0 336 155.24 0 0 0 0 0 0 337 222.94 0 0 0 0 0 0 338 147.72 23.35 0 0 0 0 23.35 339 442.78 12.4 0 0 0 0 12.4 340 284.29 0 0 0 0 0 0 341 421.92 0 0 0 0 0 0 342 293.81 3.32 25.05 0 0 0 28.37 343 68.65 0 0 0 0 0 0 344 372.42 0 0 0 0 0 0 345 232.37 0 51.75 0 0 0 51.75 346 261.09 7.87 2.7 0 0 0 10.57 347 231.06 7.85 4.85 0 0 0 12.7 348 167.25 0 0 0 0 0 0 349 202.86 0 0 0 0 0 0 350 308.59 0 0 0 0 0 0 351 303.82 0 0 0 0 0 0 352 83.15 0 0 0 0 0 0 353 179.99 1.27 0 0 0 0 1.27 354 220.1 0 0 0 0 0 0 355 330.11 0 0 0 0 0 0 356 184.28 0 0 0 0 0 0 357 328.71 0 0 0 0 0 0 358 133.82 0 0 0 0 0 0 359 171.66 0 35.21 0 0 0 35.21 360 183.15 0 0 0 0 0 0 361 106.29 0 0 0 0 0 0 362 158.45 0 0 0 0 0 0 363 244.33 0 0 0 0 0 0 364 131.26 0 0 0 0 0 0 365 185.59 0 0 0 0 0 0 366 270.61 0 0 0 0 0 0 367 225.58 0 0 0 0 0 0 368 325.98 0 0 0 0 0 0 369 327.2 0 0 0 0 0 0 370 124.56 0 0 0 0 0 0 371 123.53 10.77 0 0 0 0 10.77 372 282.72 0 15.95 0 0 0 15.95 373 287.18 0 23.1 0 0 0 23.1 374 308.22 0 14.79 0 0 0 14.79 375 290.16 0 0 0 0 0 0 376 124.13 0 0 0 0 0 0 377 126.05 0 5.36 0 0 0 5.36 378 140.8 0 0 0 0 0 0 379 132.01 0 11.93 0 0 0 11.93 380 568.4 0 0 0 0 0 0 381 227.4 0 0 0 0 0 0 382 164.84 0 0 0 0 0 0 383 217.91 0 0 0 0 0 0 384 205.27 0 0 0 0 0 0 385 78.74 1.45 0 0 0 0 1.45 386 106.28 8.91 0 0 0 0 8.91 387 151.57 0 0 0 0 0 0 388 131.47 0 10.02 0 0 0 10.02 389 168.21 0 0 0 0 0 0 390 119.86 0 0 0 0 0 0 391 381.23 0 9.69 0 0 0 9.69 392 171.7 0 0 0 0 0 0 393 115.68 0 0 0 0 0 0 394 167.26 0 0 0 0 0 0 395 151.5 2.1 0 0 0 0 2.1 396 301.01 0 0 0 0 0 0 397 235.94 9.59 0 0 0 0 9.59 398 222.22 0 0 0 0 0 0 399 98.39 0 0 0 0 0 0 400 424.01 0 0 0 0 0 0 401 124.69 0 0 0 0 0 0 402 302.03 0 0 0 0 0 0 403 748.78 0 0 0 0 0 0 404 219.84 0 0 0 0 0 0 405 240.37 0 0 0 0 0 0 406 93.33 0 0 0 0 0 0 407 240.79 0 0 0 0 0 0 408 66.28 0 0 0 0 0 0 409 71.93 0 0 0 0 0 0 410 127.23 2.92 0 0 0 0 2.92 411 85.18 0 0 0 0 0 0 412 251.6 0 0 0 0 0 0 413 187.58 0 0 0 0 0 0 414 70.81 0 0 0 0 0 0 415 70.67 0 4.36 0 0 0 4.36 416 95.32 0 0 0 0 0 0 417 135.11 0 0 0 0 0 0 418 210.48 0 0 0 0 0 0 419 71 0 0 0 0 0 0 420 173.81 3.69 0 0 0 0 3.69 421 506.9 0 0 0 0 0 0 422 633.72 5.8 0 0 0 0 5.8 423 37.44 0 0 0 0 0 0 424 573.38 0 0 0 0 0 0 425 697.75 0 0 0 0 0 0 426 65.45 0 0 0 0 0 0 427 316.96 0 22.13 0 0 0 22.13 428 256.45 0 0 0 0 0 0 429 45.67 0 0 0 0 0 0 430 190.26 0 12.04 0 0 0 12.04 431 235.59 0 0 0 0 0 0 432 212.84 0 0 0 0 0 0 433 261.6 0 102.61 0 0 0 102.61 434 47.14 0 1.67 0 0 0 1.67 435 127.92 0 0 0 0 0 0 436 150.53 0 0 0 0 0 0 437 185.29 0 0 0 0 0 0 438 100.56 0 0 0 0 0 0 439 580.06 0 29.28 0 0 0 29.28 440 270.66 0 0 0 0 0 0 441 282.8 0 0 0 0 0 0 442 360.87 3.49 0 0 0 0 3.49 443 142.11 0 0 0 0 0 0 444 165.01 0 7.5 0 0 0 7.5 445 155.26 0 0 0 0 0 0 446 138.72 0 22.77 0 0 0 22.77 447 350.79 3.26 32.27 0 0 0 35.53 448 362.7 0 0 0 0 0 0 449 356.32 0 6.78 0 0 0 6.78 450 170.68 0 0 0 0 0 0 451 1077.01 0 0 0 0 0 0 452 178.42 0 0 0 0 0 0 453 486.61 5.01 0 0 0 0 5.01 454 506.22 0 0 0 0 0 0 455 365.41 40.29 0 0 0 0 40.29 456 380.64 4.57 0 0 0 0 4.57 457 771.17 0 0 0 0 0 0 458 192.71 0 0 0 0 0 0 459 106.45 0 0 0 0 0 0 460 402.55 0 0 0 0 0 0 461 252.5 0 51.26 0 0 0 51.26 462 137.79 0 0 0 0 0 0 463 31.72 0 0 0 0 0 0 464 111.09 0 0 0 0 0 0 465 94.16 0 0 0 0 0 0 466 131.06 0 0 0 0 0 0 467 294.79 0 0 0 0 0 0 468 170.93 0 0 0 0 0 0 469 365.58 0 0 0 0 0 0 470 100.16 0 0 0 0 0 0 471 457.29 0 0 0 0 0 0 472 184.04 0 0 28.07 0 0 28.07 473 377.37 0 0 0 0 0 0 474 128.84 0 0 0 0 0 0 475 610.73 6.39 32.1 0 0 0 38.49 476 202.35 0 23.14 0 0 0 23.14 477 72.71 0 0 0 0 0 0 478 634.46 0 37 0 0 0 37 479 208.74 0 0 0 0 0 0 480 290.43 0 19.38 0 0 0 19.38 481 275.54 0 0 0 0 0 0 482 259.16 0 0 0 0 0 0 483 101.72 0 0 0 0 0 0 484 532.44 0 0 0 0 0 0 485 216.58 0 0 0 0 0 0 486 122.61 0 0 0 0 0 0 487 169.6 0 0 0 0 0 0 488 142.15 0 0 91.23 0 0 91.23 489 131.95 0 0 0 0 0 0 490 575.63 0 0 0 0 0 0 491 179.31 0 0 0 0 0 0 492 198.24 0 0 0 0 0 0 493 360.18 0 0 0 0 0 0 494 144.43 0 0 0 0 0 0 495 291.49 0 0 0 0 0 0 496 90.44 0 0 0 0 0 0 497 100.79 0 0 8.44 0 0 8.44 498 204.59 0 0 0 0 0 0 499 569.23 0 0 0 0 0 0 500 253.03 1.2 0 0 0 0 1.2 501 205.42 0 0 0 0 0 0 502 204.19 0 0 0 0 0 0 503 38.26 0 0 0 0 0 0 504 156.27 0 0 0 0 0 0 505 321.29 0 0 0 0 0 0 506 262.46 0 39.61 0 0 0 39.61 507 144.68 0 0 0 0 0 0 508 359.5 0 0 0 0 0 0 509 238.47 0 0 0 0 0 0 510 428 0 9.25 0 0 0 9.25 511 543.11 0 0 0 0 0 0 512 119.95 0 0 0 0 0 0 513 360.58 0 4.35 0 0 0 4.35 514 464.7 7.37 0 0 0 0 7.37 515 408.73 12.92 56.5 0 0 0 69.42 516 105.85 0 0 0 0 0 0 517 116.33 0 0 0 0 0 0 518 211.77 0 0 0 0 0 0 519 198.96 0 0 0 0 0 0 520 290.24 13.61 19.95 0 0 0 33.56 521 238.79 0 0 0 0 0 0 522 163.91 0 0 0 0 0 0 523 280.47 0 0 0 0 0 0 524 349.2 5.94 0 0 0 0 5.94 525 488.07 0 0 0 0 0 0 526 132.2 0 6.24 0 0 0 6.24 527 375.85 0 0 0 0 0 0 528 167.41 7.72 6.81 0 0 0 14.53 529 82.86 0 0 0 0 0 0 530 85.38 0 0 0 0 0 0 531 ^ 92.81 0.83 0 0 0 0 0.83 532 273.3 28.09 0 0 0 0 28.09 533 105.62 0 0 0 0 0 0 534 91.34 0 0 0 0 0 0 535 477.24 6.44 0 0 0 0 6.44 536 271.04 6.86 0 0 0 0 6.86 537 142.66 0 0 0 0 0 0 538 377.91 0 30.59 0 0 0 30.59 539 36.82 0 0 0 0 0 0 540 636.96 0 30.8 0 0 0 30.8 541 121.25 0 0 0 0 0 0 542 181.32 0 0 0 0 0 0 543 113.76 0 10.23 0 0 0 10.23 544 67.18 0 0 0 0 0 0 545 136.57 0 0 0 0 0 0 546 71.58 0 0 0 0 0 0 547 124.87 0 0 0 0 0 0 548 215.52 0 0 0 0 0 0 549 831.45 0 0 0 0 0 0 550 202.47 0 0 0 0 0 0 551 676.58 0 0 0 0 0 0 552 133.57 0 0 0 0 0 0 553 177.21 0 0 0 0 0 0 554 110.12 7.16 0 0 0 0 7.16 555 281.62 8.35 1.62 0 0 0 9.97 556 65.51 0 0 0 0 0 0 557 402.7 0 4.69 0 0 0 4.69 558 348.25 0 0 0 0 0 0 559 462.64 0 22.88 0 0 0 22.88 560 636.74 0 0 0 0 0 0 561 151.59 0 0 0 0 0 0 562 108.25 0 0 0 0 0 0 563 89.84 0 10.93 0 0 0 10.93 564 631.96 2.84 0 0 0 0 2.84 565 389.79 9.25 51.18 0 0 0 60.43 566 69.92 0 0 0 0 0 0 567 325.81 0 0 0 0 0 0 568 324.2 0 0 0 0 0 0 569 439.77 0 0 0 0 0 0 570 883.39 0 0 0 0 0 0 571 236.07 0 0 0 0 0 0 572 252.29 0 0 0 0 0 0 573 147.27 0 0 0 0 0 0 574 331.44 0 0 0 0 0 0 575 1097.26 14.27 0 0 0 0 14.27 576 668.54 0 0 225.22 0 0 225.22 577 110.69 0 4.28 0 0 0 4.28 578 82.31 0 5.12 0 0 0 5.12 579 289.03 0 0 0 0 0 0 580 1943.23 0 0 0 0 0 0 581 231.86 0 26.84 0 0 0 26.84 582 39.5 0 0 0 0 0 0 583 977.42 0 0 0 0 0 0 584 1054.11 0 0 0 0 0 0 Herbivory data from Florissant.

Leaf Original Hole Feed Margin Feed Skclctoni/ing Galling Leaf Mining Total Number Leaf Area Area Area Area Area Area Damage Area 1 1865.22 0 0 0 0 0 0 2 456.62 0 0 0 0 0 0 3 828.07 0 0 0 0 0 0 4 145.4 0 8.83 0 0 0 8.83 5 77.11 3.14 0 0 0 0 3.14 6 110.54 0 0 0 0 0 0 7 56.87 0 0 0 0 0 0 8 283.65 0 0 0 0 0 0 9 541.93 4.84 0 0 0 0 4.84 10 403.4 0 0 0 0 0 0 II 854.37 0 0 0 0 0 0 12 91.61 1.02 0 0 0 0 1.02 13 18.54 1.16 0 0 0 0 1.16 14 311.87 0 0 0 0 0 0 15 529.72 0 0 0 0 0 0 16 130.76 0 47.74 0 0 0 47.74 17 268.89 0 0 0 0 0 0 18 1187.95 0 0 0 0 0 0 19 124.97 0 0 0 0 0 0 20 1199.78 0 0 0 0 0 0 21 573.43 0 0 0 0 0 0 22 411.05 0 0 0 0 0 0 23 380.98 0 0 0 0 0 0 24 365.15 27.76 3.67 0 0 0 31.43 25 258.66 0 0 0 0 0 0 26 231.04 0 0 0 0 0 0 27 124.16 0 0 0 0 0 0 28 1098.7 7.54 0 0 0 0 7.54 29 561.03 0 0 0 0 0 0 30 90.54 0 0 0 0 0 0 31 278 0 8.86 0 9.75 0 18.61 32 59.27 0 0 0 0 0 0 33 234.38 0 0 0 0 0 0 34 279.45 0 0 0 0 0 0 35 164.24 0 0 0 0 0 0 36 751.28 70.4 0 0 0 0 70.4 37 197.1 0 0 0 0 0 0 38 88.26 0 0 0 0 0 0 39 238.89 0 0 0 0 0 0 40 236.93 0 0 0 0 0 0 41 206.39 0 10.11 0 0 0 10.11 42 374.5 14.79 0 0 0 0 14.79 43 154.43 0 0 0 0 0 0 44 150.15 1.13 0 0 0 0 1.13 45 1245.65 157.32 0 0 0 0 157.32 46 517.16 0 12.75 0 0 0 12.75 47 269.05 0 0 0 0 0 0 48 127.66 0 0 0 0 0 0 49 564.76 3.96 45.45 0 0 0 49.41 50 709.25 0 0 0 0 0 0 51 171.82 0 5.45 0 0 0 5.45 52 119.37 0.95 0 0 0 0 0.95 53 48.33 0 0 0 0 0 0 54 370.99 0 0 0 0 0 0 55 190.63 2.05 0 0 0 0 2.05 56 208 0 0.95 0 0 0 0.95 57 233.8 0 0 0 0 0 0 58 1127.25 0 0 0 0 0 0 59 1392.41 0 0 0 0 0 0 60 29.01 0 0 0 0 0 0 61 198.82 2.48 0 0 0 0 2.48 62 139.41 0 0 0 0 0 0 63 162.29 48.82 0 0 0 0 48.82 64 1086.61 7.75 0 0 0 0 7.75 65 347.68 2.95 0 0 0 0 2.95 66 270.79 3.82 13.21 0 0 0 17.03 67 657.66 7.07 0 0 0 0 7.07 68 254.3 2.44 0 0 0 0 2.44 69 288.67 26.31 0 0 0 0 26.31 70 190.06 0 5.97 0 0 0 5.97 71 60.02 2.7 0 0 0 0 2.7 72 314.99 8.54 13.23 0 0 0 21.77 73 175.67 10.2 0.96 0 0 0 11.16 74 175.4 0.93 0 0 0 0 0.93 75 163.65 0 0 0 0 0 0 76 309.07 0 14.46 0 0 0 14.46 77 79.6 0 0 0 0 0 0 78 164 0 4.89 0 0 0 4.89 79 54.5 0 0 0 0 0 0 80 170.58 0.79 0 0 0 0 0.79 81 244.29 0 0 0 0 0 0 82 114.38 0 0 0 0 0 0 83 197.15 0 0 0 0 0 0 84 126.03 3.78 0 0 0 0 3.78 85 99.56 0 0 0 0 0 0 86 161.62 0 0 0 0 0 0 87 31.99 0 0 0 0 0 0 88 182.74 0 0 0 0 0 0 89 1337.73 0 0 0 29.76 0 29.76 90 266.75 6.21 0 0 0 0 6.21 91 147.11 0 0 0 0 0 0 92 85.61 0 0 0 0 0 0 93 146.08 1.33 0 0 0 0 1.33 94 112.91 0 14.34 0 0 0 14.34 95 213.55 0 0 0 0 0 0 96 78.22 0 0 0 0 0 0 97 93.21 0 0 0 0 0 0 98 414.66 0 48.63 0 0 0 48.63 99 130.91 0 0 0 0 0 0 100 601.79 0 0 0 0 0 0 101 712.71 0.65 0 0 14.61 0 15.26 102 234.91 0 0 0 0 0 0 103 803.28 0 4.27 0 0 0 4.27 104 410.5 0 0 0 0 0 0 105 86.2 0 0 0 0 0 0 106 514.86 0 0 0 0 0 0 107 92.97 0 0 11.24 0 0 11.24 108 164.16 0 0 0 0 0 0 109 152.02 0 0 0 0 0 0 no 214.17 0 0 0 0 0 0 III 212.05 0 0 0 0 0 0 112 168.32 0 0 0 0 0 0 113 192.02 0 0 0 0 0 0 114 101.06 0 0 0 0 0 0 115 187.12 0 0 0 0 0 0 116 102.18 0 0 0 0 0 0 117 452.45 0 0 0 0 0 0 118 98.26 0 0 0 0 0 0 119 76.08 0 0 0 0 0 0 120 194.25 0 0 0 0 0 0 121 164.81 0 0 0 0 0 0 122 189.68 0 0 0 0 0 0 123 336.44 0 0 0 0 0 0 124 153.37 0 9.34 0 0 0 9.34 125 201.85 24.08 20.11 0 0 0 44.19 126 44.66 0 0 0 0 0 0 127 275.33 0 0 0 0 0 0 128 163.97 0 0 0 0 0 0 129 224.13 0 0 0 0 0 0 130 196.55 0 0 0 0 0 0 131 333.04 0 0 0 0 0 0 132 178.96 0 0 0 0 0 0 133 428.92 0 0 0 0 0 0 134 175.85 0 0 0 0 0 0 135 139.47 1.56 0 0 0 0 1.56 136 544.52 0 0 0 0 0 0 137 232.18 6.29 0 0 0 0 6.29 138 446.43 0 0 0 0 0 0 139 298.81 0 0 0 0 0 0 140 283.29 0 0 0 0 0 0 141 250.04 0 0 0 0 0 0 142 106.17 0 0 0 0 0 0 143 201.88 0 0 0 0 0 0 144 162.04 0.87 0 0 0 0 0.87 145 211.22 0 0 0 0 0 0 146 244.69 0 0 0 0 0 0 147 201.71 0 0 0 0 0 0 148 69.15 0 0 0 0 0 0 149 269.34 0 0 0 0 0 0 150 183.29 0 0 0 0 0 0 151 489.82 0 0 0 0 0 0 152 521.81 0 0 0 0 0 0 153 250.02 0 0 0 0 0 0 154 327.5 0 0 0 0 0 0 155 484.17 0 0 0 0 0 0 156 630.56 0 0 0 0 0 0 157 531.69 0 0 0 0 0 0 158 392.57 0 17.64 0 0 0 17.64 159 243.57 0 0 0 0 0 0 160 67.32 0 0 0 0 0 0 161 488.07 0 0 0 0 0 0 162 674.64 0 0 0 0 0 0 163 300.24 0 11.6 0 0 0 11.6 164 294.78 0 0 0 0 0 0 165 104.56 0 0 9.67 0 0 9.67 166 245.93 0 0 0 21.73 0 21.73 167 163.09 0 0 0 0 0 0 168 237.77 0 0 0 0 0 0 169 707.84 0 0 0 0 0 0 170 443 0 0 17.6 0 0 17.6 171 74.8 0 0 0 0 0 0 172 458.06 0 0 0 0 0 0 173 363.42 0 0 0 0 0 0 174 200.31 0 4.45 0 0 0 4.45 175 382.64 2.34 0 0 0 0 2.34 176 268.5 0 3.26 0 0 0 3.26 177 323.13 0 0 0 0 0 0 178 200.06 1.14 0 0 0 0 1.14 179 555.81 0 0 0 0 0 0 180 131.24 0 0 0 0 0 0 181 156.78 0 1.06 0 0 0 1.06 182 524.43 0 0 0 0 0 0 183 326.35 0 0 0 12.7 0 12.7 184 305.47 0 0 50.9 0 0 50.9 185 518.82 0 0 0 0 0 0 186 188.63 0 0 0 0 0 0 187 78.4 0 0 0 0 0 0 188 210.76 0.37 0 0 0 0 0.37 189 474.39 0 11.17 0 0 0 11.17 190 431.08 0 0 0 0 0 0 191 138.61 0 0 0 0 0 0 192 571.71 0.46 0 0 0 0 0.46 193 27.03 0 0 0 0 0 0 194 334.82 0 0 0 0 0 0 195 104.9 0 0 0 0 0 0 196 1092.51 0 0 0 0 0 0 197 340.24 0 0 0 0 0 0 198 332.79 0 0 0 0 0 0 199 281.05 0 0 0 0 0 0 200 706.7 0 0 0 0 0 0 201 138.86 0 0 0 0 0 0 202 143.45 0 0 0 0 0 0 203 225.68 12.34 0 0 0 0 12.34 204 418.46 0 0 0 0 0 0 205 402.24 0 10.13 0 0 0 10.13 206 406.58 1.23 0 0 0 0 1.23 207 140.07 1.64 0 0 0 0 1.64 208 384.48 52.41 0 0 0 0 52.41 209 192.91 0.98 0 0 0 0 0.98 210 383.94 0 0 0 0 0 0 211 583.39 0 0 0 15.94 0 15.94 212 256.56 0.64 0 0 0 0 0.64 213 350.91 0 0 0 0 0 0 214 113.83 0 0 0 0 0 0 215 243.51 2.98 6.17 0 0 0 9.15 216 148.36 0 0 0 0 0 0 217 241.37 0 0 0 0 0 0 218 161.26 0 0 0 0 0 0 219 127.18 0 0 0 0 0 0 220 125.06 0 0 0 0 0 0 221 65.23 0 0 0 0 0 0 222 345.18 0 0 0 0 0 0 223 99.24 0 0 0 0 0 0 224 218.47 0 0 0 19.97 0 19.97 225 116.74 0 5.53 0 0 0 5.53 226 220.92 0 0 0 2.41 0 2.41 227 383.13 0 0 0 0 0 0 228 177.3 0 2.16 0 0 0 2.16 229 199.92 0 0 0 0 0 0 230 261.87 0 0 0 0 0 0 231 218.37 0 0 0 0 0 0 232 67.68 0 0 0 0 0 0 233 43.82 0 0 0 0 0 0 234 43.73 0 0 0 0 0 0 235 1043.77 0 0 0 0 0 0 236 1472.98 0 0 0 0 0 0 237 201.71 0 0 0 0 0 0 238 361.69 0 0 0 0 0 0 239 1053.99 13.16 0 0 0 0 13.16 240 618.6 0.76 0 0 0 0 0.76 24! 1064.06 1.77 0 0 0 0 1.77 242 1449.53 0 0 0 0 0 0 243 1277.06 2.46 6.34 0 0 0 8.8 244 550.36 0 0 0 0 0 0 245 374.06 2.35 0 0 0 0 2.35 246 704.99 0 0 0 0 0 0 247 1064.68 57.02 0 0 0 0 57.02 248 665.22 0 0 0 0 0 0 249 1415.76 16.88 0 0 0 0 16.88 250 1498.45 7.28 0 0 0 0 7.28 251 1636.79 148.82 221.23 0 0 0 370.05 252 1193.29 81.68 0 0 0 0 81.68 253 1309.7 0 0 0 0 0 0 254 3137.6 189.34 0 0 10.82 0 200.16 255 2578.42 3.03 0 0 0 0 3.03 256 865.9 l.I 0 0 0 0 1.1 257 1476.38 0 99.3 0 0 0 99.3 258 1382.83 0 0 0 0 0 0 259 1363.55 129.87 0 0 0 0 129.87 260 527.88 89.36 0 0 0 0 89.36 261 617.16 47.49 0 0 0 0 47.49 262 574.55 0 0 0 6.44 0 6.44 263 418.03 27.96 0 0 0 0 27.96 264 1269.32 0 0 188.45 0 0 188.45 265 1002.35 0 0 0 0 0 0 266 775.85 0 0 0 0 0 0 267 114.49 0 0 0 0 0 0 268 290.48 0 0 0 0 0 0 269 124.25 0 0 0 0 0 0 270 417.33 0 0 0 0 0 0 271 538.1 0 0 0 0 0 0 212 1044.82 0 0 0 0 0 0 273 1345.11 0 0 0 0 0 0 274 307.5 0 0 0 0 0 0 275 314.3 0 0 0 0 0 0 276 139.11 0 0 0 0 0 0 277 818.88 0 0 0 0 0 0 278 1107.16 0 0 0 0 0 0 279 408.95 0 0 0 0 0 0 280 181.74 0 0 0 0 0 0 281 335.32 0 3.87 0 0 0 3.87 282 179.65 0 0 0 0 0 0 283 194.31 0 0 0 0 0 0 284 411.1 0 0 0 0 0 0 285 241.62 0 0 0 0 0 0 286 316.56 0 0 0 0 0 0 287 105.19 2.55 0 0 0 0 2.55 288 307.11 0 0 0 0 0 0 289 474.89 3.65 0 0 0 0 3.65 290 181.36 2.12 0 0 0 0 2.12 291 1031.34 4.62 0 0 0 0 4.62 292 299.99 14.79 0 0 0 0 14.79 293 201.18 0 0 0 0 0 0 294 100.76 0 0 0 0 0 0 295 222.02 0 0 0 0 0 0 296 163.61 0 0 0 0 0 0 297 235.72 0 0 0 0 0 0 298 133.53 0 0 0 0 0 0 299 233.79 0 0 0 0 0 0 300 534.77 0 0 0 0 0 0 301 133.38 0 0 0 0 0 0 302 146.43 0 0 0 0 0 0 303 102.32 0 0 0 0 0 0 304 133.05 0 0 0 0 0 0 305 242.19 0 0 0 0 0 0 306 118.36 0 0 0 0 0 0 307 1487.81 0 0 0 0 0 0 308 201.26 0 0 0 0 0 0 309 1061.83 0 0 0 0 0 0 310 905.99 0 0 0 0 0 0 311 1185 0 0 0 0 0 0 312 2057.24 0 0 0 0 0 0 313 1924.78 0 0 0 0 0 0 314 1139.2 0 0 0 0 0 0 315 840.05 0 0 0 0 0 0 316 1407.92 0 0 45.21 0 0 45.21 317 3811.16 0 0 0 0 0 0 318 1665.85 0 0 0 0 0 0 319 1765.65 0 0 0 0 0 0 320 83.61 0 0 0 0 0 0 321 128.78 0.46 0 0 0 0 0.46 322 113.55 0 0 3.5! 0 0 3.51 323 60.9 0 0 0 0 0 0 324 134.44 0 0 0 0 0 0 325 350.67 0 0 0 0 0 0 326 44.24 0 0 0 0 0 0 111 61.09 0 0 0 0 0 0 328 138.49 0 0 0 0 0 0 329 79.62 0 0 0 0 0 0 330 296.53 0 0 0 0 0 0 331 229.06 0 0 0 0 0 0 332 395.86 1.94 0 0 0 0 1.94 333 68.35 0 0 0 0 0 0 334 48.87 0 0 0 0 0 0 335 123.17 0 4.96 0 0 0 4.96 336 45.21 0 0 0 0 0 0 337 23.2 0 0 0 0 0 0 338 15.78 0 0 0 0 0 0 339 740.81 0 0 0 0 0 0 340 95.01 0 0 0 0 0 0 341 404.3 0 0 0 0 0 0 342 1158.61 0 0 0 0 0 0 343 588.8 0 0 0 0 0 0 344 133.1 0 0 0 0 0 0 345 215.71 0 0 0 0 0 0 346 284.01 0 0 0 0 0 0 347 346.28 0 0 0 0 0 0 348 111.43 0 0 0 0 0 0 349 105.08 0 0 0 0 0 0 350 92.8 0 0 0 0 0 0 351 538.02 0 0 0 0 0 0 352 210.54 0 14.86 0 0 0 14.86 353 341.4 0 0 0 0 0 0 354 213.46 0 0 0 0 0 0 355 158.82 0 0 0 0 0 0 356 135.78 1.42 0 0 0 0 1.42 357 107.05 0 0 0 0 0 0 358 223.91 0 0 0 0 0 0 359 272.8 0 0 0 0 0 0 360 95.79 0 0 0 0 0 0 361 314.62 0 0 0 0 0 0 362 277.06 0 0 0 0 0 0 363 115.57 0 0 0 0 0 0 364 86.35 0 0 0 0 0 0 365 55.03 0 0 0 0 0 0 366 93.24 0 0 0 0 0 0 367 103.74 0 0 0 0 0 0 368 11.6 0 0 0 0 0 0 369 56.74 0 0 0 0 0 0 370 55.52 0 0 0 0 0 0 371 48.08 0 0 0 0 0 0 372 48.98 0 0 0 0 0 0 373 36.1 0 0 0 0 0 0 374 1271.36 0 0 0 0 0 0 375 107.18 0 0 0 0 0 0 376 418.7 0 0 0 0 0 0 'ill 555.33 0 0 0 0 0 0 378 125.16 0 0 0 0 0 0 379 188.58 0 0 0 0 0 0 380 146.6 0 0 0 0 0 0 381 692.08 0 0 0 0 0 0 382 832.01 0 22.49 0 0 0 22.49 383 2407.61 38.24 0 0 0 0 38.24 384 326.89 19.72 0 0 0 0 19.72 385 293.63 0 0 0 0 0 0 386 363.38 0 0 0 0 0 0 387 104.54 0 0 0 0 0 0 388 1286.18 0 198.44 0 0 0 198.44 389 711.91 0 0 0 0 0 0 390 405.57 0 0 0 0 0 0 391 119.56 0 0 0 0 0 0 392 379.63 0 56.78 0 0 0 56.78 393 41.74 0 0 0 0 0 0 394 125.91 0 0 0 0 0 0 395 103.22 0 0 0 0 0 0 396 510.32 1.62 0 0 0 0 1.62 397 806.16 0 0 0 0 0 0 398 1084.05 0 0 0 0 0 0 399 1606.79 0 144.25 0 60.16 0 204.41 400 260.24 0 14.99 0 0 0 14.99 401 497.83 5.16 38.74 0 0 0 43.9 402 1257.18 15.26 85.06 0 0 0 100.32 403 127.47 0 0 0 0 0 0 404 174.13 0 0 0 0 0 0 405 186.64 0 0 0 0 0 0 406 324.78 0 0 0 0 0 0 407 64.3 0 0 0 0 0 0 408 31.8 0 1.02 0 0 0 1.02 409 21.87 0 0 0 0 0 0 410 173.9 0 13.62 0 0 0 13.62 411 123.23 0 0 0 8.17 0 8.17 412 302.26 0 0 0 0 0 0 413 574.63 0 0 0 0 0 0 414 35.65 0 0 0 0 0 0 415 16.61 0 0 0 0 0 0 416 104.13 0 0 0 0 0 0 417 44.66 0 0 0 0 0 0 418 46.67 0 1.08 0 0 0 1.08 419 103.69 0 0 0 0 0 0 420 131.3 0 0 0 0 0 0 421 121.2 0 1.89 0 0 0 1.89 422 606.52 0 0 0 0 0 0 423 405.57 1.68 0 0 0 0 1.68 424 114.74 0 0 0 0 0 0 425 141.14 0 0 0 0 0 0 426 111.03 0 0 0 0 0 0 427 842.67 0 0 0 3.09 0 3.09 428 464.17 0 0 0 0 0 0 429 27.24 0 0 0 0 0 0 430 509.14 0 0 0 0 0 0 431 111.97 0 0 0 0 0 0 432 646.06 0 0 0 0 0 0 433 225.08 0 0 0 0 0 0 434 903.95 0 0 0 0 0 0 435 847.01 2.47 14.77 0 0 0 17.24 436 201.15 0 0 0 0 0 0 437 443.57 0 0 0 0 0 0 438 229.43 0 0 0 0 0 0 439 63.29 0 0 0 0 0 0 440 141.4 0 0 0 0 0 0 441 21.64 0 2.07 0 0 0 2.07 442 325.49 0 0 0 0 0 0 443 463.72 3.13 22.49 0 0 0 25.62 444 206.88 0 0 0 0 0 0 445 109.53 0 0 0 0 0 0 446 194.96 0 0 0 0 0 0 447 45.16 0 0 0 0 0 0 448 57.04 0 0 0 0 0 0 449 358.52 0 0 0 0 0 0 450 826.6 0 0 0 0 0 0 451 728.84 0 0 0 0 0 0 452 72.07 0 0 0 0 0 0 453 78.99 0 0 0 0 0 0 454 39.53 0 0 0 0 0 0 455 384.62 0 33.71 0 0 0 33.71 456 310.83 0 0 0 0 0 0 457 261.7 0 0 0 0 0 0 458 386.36 0 0 0 0 0 0 459 93.73 0 0 0 0 0 0 460 215.45 0 0 0 0 0 0 461 125.13 0 0 0 0 0 0 462 721.2 0 0 0 0 0 0 463 152.33 0 0 0 0 0 0 464 122.5 0 0 0 0 0 0 465 531.11 0 0 0 0 0 0 466 1290.75 0 0 0 0 0 0 467 455.99 0 0 0 0 0 0 468 379.47 0 0 0 0 0 0 469 1676.03 11.13 0 0 0 0 11.13 470 159.31 0 0 0 0 0 0 471 200.97 0 0 0 0 0 0 472 199.83 0 0 0 0 0 0 473 328.7 0 0 0 0 0 0 474 408.07 0 0 0 0 0 0 475 350.81 0 0 0 0 0 0 476 216.77 0 0 0 0 0 0 477 258.97 0 0 0 0 0 0 478 459.64 0 0 0 0 0 0 479 408.07 0 0 0 0 0 0 480 931.88 0 0 0 0 0 0 481 560.87 0 0 0 0 0 0 482 617.39 0 0 0 0 0 0 483 542.68 0 0 0 0 0 0 484 464.17 0 0 0 0 0 0 485 113.24 0 0 0 0 0 0 486 271.83 0 0 0 0 0 0 487 506.63 0 0 0 0 0 0 488 348.37 0 0 0 0 0 0 489 242.34 0 0 0 0 0 0 490 1887.53 0 0 0 0 0 0 491 896.06 0 0 0 0 0 0 492 534.77 0 0 0 0 0 0 493 171.11 0 0 0 0 0 0 494 1383.55 0 0 0 0 0 0 495 467.08 0 0 0 0 0 0 496 257.92 0 0 0 0 0 0 497 247.26 0 0 0 0 0 0 498 596.68 0 0 0 0 0 0 499 323.24 0 0 0 0 0 0 500 282.07 0 0 0 0 0 0 501 208.96 0 0 0 0 0 0 502 76.42 0 0 0 0 0 0 503 94.98 0 0 0 0 0 0 504 188.45 0 0 0 0 0 0 505 1576.41 0 0 0 0 0 0 506 123.55 0 0 0 0 0 0 507 179.17 0 0 0 0 0 0 508 140.6 0 0 0 0 0 0 509 367.7 0 0 0 0 0 0 510 34.88 0 0 0 0 0 0 511 508.72 0 0 0 0 0 0 512 521.9 0 0 0 0 0 0 513 392.32 0 0 0 0 0 0 514 330.05 3.92 0 0 0 0 3.92 515 238.96 0 0 0 0 0 0 516 369 0 0 0 0 0 0 517 578.15 0 0 0 0 0 0 518 711.02 0 0 0 0 0 0 519 384.49 0 0 0 0 0 0 520 51.79 0 0 0 0 0 0 521 167.3 0 0 0 0 0 0 522 192.4 0 0 0 0 0 0 523 423 0 0 0 0 0 0 524 593.78 0 0 0 0 0 0 525 570.87 0 0 0 0 0 0 526 102.56 0 0 0 0 0 0 527 183.58 0 0 0 0 0 0 528 393.19 0 0 0 0 0 0 529 544.99 0 0 0 0 0 0 530 395.81 0 0 0 0 0 0 531 416.02 0 0 0 0 0 0 532 380.15 0 0 0 0 0 0 533 280.46 0 0 0 0 0 0 534 219.58 0 0 0 0 0 0 535 270.47 0 0 0 0 0 0 536 178.85 0 0 0 0 0 0 537 869.64 0 0 0 0 0 0 538 419.32 0 0 0 0 0 0 539 195.51 0 0 0 0 0 0 540 499.61 0 0 0 0 0 0 541 931.53 0 0 0 0 0 0 542 211.7 0 0 0 0 0 0 543 814.94 0 0 0 0 0 0 544 368.37 0 0 0 0 0 0 545 74.94 0 0 0 0 0 0 546 649.23 0 0 0 0 0 0 547 126.85 0 0 0 0 0 0 548 297.64 0 0 0 0 0 0 549 1018.3 0 0 0 0 0 0 550 295.51 0 0 0 0 0 0 551 1549.99 0 0 0 0 0 0 552 468.63 0 0 0 0 0 0 553 805.46 0 0 0 0 0 0 554 606.21 0 0 0 0 0 0 555 294.4 14.43 23.95 0 0 0 38.38 556 247.56 0 0 0 0 0 0 557 126.69 0 0 0 0 0 0 558 719.47 0 0 0 0 0 0 559 419.98 0 0 0 0 0 0 560 442.33 0 0 0 0 0 0 561 409.84 0 0 0 0 0 0 562 218.1 0 0 0 0 0 0 563 107.52 0 0 0 0 0 0 564 191.32 0 0 0 0 0 0 565 767.53 0 0 0 0 0 0 566 93 0 0 0 0 0 0 567 62.9 0 0 0 0 0 0 568 398.47 0 0 0 0 0 0 569 112.28 0 0 0 0 0 0 570 174.93 0 9.01 0 0 0 9.01 571 144.31 0 0 0 0 0 0 572 248.53 0 0 0 0 0 0 573 518.59 0 0 0 0 0 0 574 135.85 0 0 0 0 0 0 575 869.87 0 0 0 0 0 0 576 175.78 0 0 0 0 0 0 577 422.17 0 0 0 0 0 0 578 659.19 0 0 0 0 0 0 579 153.33 2.16 0 0 0 0 2.16 580 385.24 0 0 0 0 0 0 581 968.21 0 0 0 0 0 0 582 267.43 0 0 0 0 0 0 583 733.87 26.2 0 0 0 0 26.2 584 532.23 2.55 0 0 0 0 2.55 585 1461.19 0 0 0 0 0 0 586 1357.44 0 0 0 0 0 0 587 200.98 0 0 0 0 0 0 588 562.48 0 0 0 0 0 0 589 265.34 0 0 0 0 0 0 590 407.83 0 0 0 0 0 0 591 509.85 0 0 0 0 0 0 592 489.58 24.7 0 0 0 0 24.7 593 179.76 0 0 0 0 0 0 594 564.06 0 0 0 0 0 0 595 168.44 0 0 0 0 0 0 596 219.91 0 0 0 0 0 0 597 207.56 0 0 0 0 0 0 598 288.97 0 0 0 0 0 0 599 212.69 0 0 0 0 0 0 600 221.24 0 0 0 0 0 0 601 84.41 0 0 0 0 0 0 602 123.05 0 0 0 0 0 0 603 128.89 0 0 0 0.95 0 0.95 604 295.06 0 0 0 0 0 0 605 295.05 0 0 0 0 0 0 606 127.03 0 0 0 0 0 0 607 88.65 0 2.66 0 0 0 2.66 608 172.79 0 3.93 0 0 0 3.93 609 149.68 0 0 0 0 0 0 610 128.26 l.I 0 0 0 0 1.1 611 297.16 0 0 0 0 0 0 612 756.56 0 0 0 0 0 0 613 182.25 0 0 0 0 0 0 614 114.7 0 0 0 0 0 0 615 155.39 0 0 0 0 0 0 616 286.72 0 0 0 0 0 0 617 127.35 0 0 0 0 0 0 618 151.97 0 0 0 0 0 0 619 122.57 0 0 0 0 0 0 620 107.65 0 0 0 0 0 0 621 186.32 0 0 0 0 0 0 622 453.76 0 0 0 0 0 0 623 1331.2 0 0 0 0 0 0 624 292.43 0 0 0 0 0 0

OJ ou> 304

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