Wood Trait Preferences of Neotropical Xylophagous Beetles (Coleoptera: Cerambycidae)

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Wood trait preferences of Neotropical xylophagous beetles (Coleoptera: Cerambycidae)

Christina Torres CUNY City College

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Wood trait preferences of Neotropical xylophagous beetles (Coleoptera: Cerambycidae).

Christina Ann Torres

Thesis Advisor: Dr. Amy Berkov

A thesis submitted in partial fulfillment of the requirements for the Degree of Bachelor of Science/Master of Science in Biology

THE CITY COLLEGE OF NEW YORK The City University of New York 2018

ABSTRACT

Tree life history strategies are correlated with functional plant traits, such as wood density

(WD), moisture content (MC), bark thickness (BT), and nitrogen content (N); these traits affect the nutrients available to xylophagous insects. Cerambycid beetles feed on substrates that vary in these traits, but little is known about how they affect community composition. The goal of this project is to document the abundance of two cerambycid subfamilies (Cerambycinae and

Lamiinae) and the WD, MC, BT, and N content in the wood they eat. In a salvage project conducted adjacent to the Panama Canal, trees were felled and exposed to Cerambycidae for oviposition. Disks from branches of differing thickness from the same plant individuals were used to calculate WD, MC, and BT in the field; nitrogen content was determined using mass spectrometry. Thick and thin branches tended to differ in wood trait values; therefore, data were analyzed separately in subsequent analyses. In thin branches, cerambycid abundance and species richness were higher in samples with less dense, moister wood, and thicker bark. Thick branches showed similar trends, but the wood traits accounted for little variability in beetle abundance or species richness. There were no significant correlations between beetle data and nitrogen.

Cerambycines emerged more slowly, and from denser, drier wood, than lamiines. Cerambycines might be more drought-tolerant than lamiines, and therefore more resistant to the longer, more severe dry seasons that are predicted to occur due to climate change.

Key Words: longhorn beetles, ecology, dry season, wood traits, early colonists, insect-plant interactions

Insect feeding strategies influence their life histories, community composition, and the ecosystem services they provide - including pollination, pest control, and decomposition. Most cerambycid larvae (Coleoptera: Cerambycidae) consume wood, starting at the cambial region, and then typically continuing to consume the sapwood (Haack 2017). Cerambycids promote wood decomposition, but the magnitude of their contribution is unknown (Calderón-Cortés et al. 2011;

Ulyshen and Wagner 2013; Ulyshen 2016). Tree life history strategies are correlated with their functional traits, such as wood density (WD) and moisture content (MC), bark thickness (BT), and nitrogen content (N); these traits affect the nutrients available to xylophagous insects (Filipiak and

Weiner 2017; Martius 1992; Müller and Müller 2016; Poorter et al. 2014). Little is known about the trait preferences of Neotropical cerambycids. If cerambycids taxa preferentially colonize wood with particular traits, those patterns could provide insight into beetle host preferences and the ecosystem services they provide (Filipiak and Weiner 2017; Martius 1992; Nahrung et al. 2014).

Wood is a nutritionally poor food substrate because of its low nitrogen and high lignin content (Haack and Slansky et al. 1987; Portillo-Quintero et al. 2014; Saint-Germain et al. 2007).

In the Neotropics, wood nitrogen content ranges between 0.03% - 0.59% (Martin et al. 2014;

Martius 1992). When cerambycid larvae feed on undecayed dead wood, nitrogen is the most limiting factor, while phosphorus is the limiting factor for larvae feeding on highly decayed wood

(Filipiak and Weiner 2017). Cerambycids may compensate for the poor wood nutritional quality by consuming wood-rot fungi, ingesting fungal enzymes, or harboring gut symbionts (Ayayee et al. 2015; Filipiak and Weiner 2017).

Wood density, like nitrogen, differs between plant taxa. Trees vary in the diameters of their conducting cells and the fortification of the fibers that contribute to their structural support, affecting the weight of a given volume of wood (basic WD). In trees, WD affects structural stability

4 and the efficiency of water transport (Gleason et al. 2016). Because density reflects the proportion of heavily lignified tissues, it directly affects substrate digestibility—especially for cerambycids that colonize living or moribund trees. Globally, WD varies by more than an order of magnitude

(0.1 g cm-3 – 1.5 g cm-3) (Chave et al. 2009). Wood density appears to increase across Amazonia as soil fertility declines (Ter Steege et al. 2006). Forest successional status also impacts local WD: in a lowland rainforest in Australia, trees in mature forests had significantly higher WD than pioneer trees (Apgaua et al. 2017). Considerable variation can be captured at a single site: on Barro

Colorado Island, Panama, WD ranges from 0.19 g cm-3 – 0.89 g cm-3 (Hietz et al. 2017).

Within trees, wood density and moisture content are inherently linked due to vessel diameters. Wood MC is influenced by environmental conditions and soil water content, but in both

Amazonian tree species and temperate pines, denser wood has less moisture (Dias and Marenco

2015; Eberhardt et al. 2017). Dense wood is not very porous and has small vessel diameters (Hacke et al. 2001). Less dense wood is more porous, with more metabolically parenchyma cells, has larger vessel diameters, and can store more water (Borchert and Pockman 2005; Nogueira et al.

2008). Higher MC allows a tree to be more resistant to physical stressors (more mechanically stable), whereas higher WD allows a tree to be more resistant to drought, because small vessels increase cavitation resistance (Hacke et al. 2001; Markesteijn et al. 2011; McCulloh et al. 2011).

Cerambycid genera tend to prefer either dry or moist wood (Haack 2017), and in some cases species select upper or lower surfaces of a tree based on MC (Iwata et al. 2007).

Bark is the corky barrier that reduces wood moisture loss (Haack 2017), and protects internal plant tissues from disturbances such as fire, pathogens, and herbivory (Paine et al. 2010;

Rosell 2016; Wiedenhoeft and Miller 2005). When 640 different tree species across 18 different ecological sites were studied, BT variation was mostly due to variation in stem size (Rosell 2016).

Bark protects larvae feeding in the cambial region, but can also pose a barrier to neonate larvae.

In a study of four traits potentially influencing cerambycid species associated with pines, BT was the most important factor affecting beetle community composition (Foit 2010).

The goal of this study is to explore the impact of these plant traits on the abundance and species richness of Neotropical cerambycid beetles. Dense wood is expected to have lower MC and nitrogen content due to the tradeoff between WD and MC, and for moister wood containing more parenchyma cells (Borchert and Pockman 2005; Hacke et al. 2001; Nogueira et al. 2008).

Cerambycids are expected to emerge in greater numbers and have a higher species richness from branches that have a high MC, low WD, thicker bark, and high nitrogen content due to cerambycid occurrence data being independently correlated with these traits in previous research (Foit 2010;

Haack 2017; Saint-Germain et al. 2007). Cerambycids are also expected to emerge the earliest from branches with these characteristics.

Furthermore, we investigate the plant traits associated with two cerambycid subfamilies

(Cerambycinae and Lamiinae). In our previous rearing experiments cerambycine beetles emerged preferentially from warmer, drier, microhabitats (canopy stratum, especially in the dry season;

Berkov 2018). Cerambycines are therefore expected to colonize trees with dense wood, and to emerge later than lamiines, because dense wood is expected to have lower MC and nitrogen content. Lamiines are expected to colonize trees with less dense wood, and to emerge earlier than cerambycines, because less dense wood is expected to have higher MC and nitrogen content.

MATERIAL AND METHODS

Study Area and Sampling Design. We reared cerambycids from freshly cut trees on a 1 ha headland on the W bank of the Panama Canal (Prov. de Colon, approximately 20 km from Barro

Colorado Island, 9º07’N, 79º45’W). The site, supporting tall secondary forest approximately 80 years old (A. Hernandez, pers. comm.) was inundated after the Canal was enlarged. Prior to inundation, we felled 67 trees and lianas (identified by A. Hernandez; 36 species in 22 families) during the dry season, (17–24 Mar 2010) (Appendix 1). Sixty branches segments (approximately

8 cm x 85 cm) were suspended in three large trees. The felled trees and these canopy baits were exposed to beetles for two months. Near the beginning of the rainy season (17–21 May 2010) we enclosed branch sections (when available: the canopy bait, three equivalent segments from a ground stratum branch, and six segments of thinner ground branches, approximately 2 cm x 85 cm; Appendix 3) separately in bags constructed from No-seeum netting and transported them to a rearing facility in Gamboa. The cages were monitored at least once a week for emerged adult beetles from 22 May 2010 through 22 May 2011. Adult cerambycids were stored in 95% EtOH.

Plant and insect vouchers were deposited at the Smithsonian Tropical Research Institute (STRI) and the University of Panama (UP). Remaining cerambycid specimens were exported to the

Berkov Lab at The City College of New York.

When the plants were initially felled, we collected thin and thick wood disks from each plant specimen (approximately 2 cm x 1 cm and 8 cm x 1cm, respectively), for analyses of branch traits (WD, MC, BT, and N). Nitrogen content was measured only for plants that produced beetles, and these plants were used in all subsequent analyses (28 species in 17 families; Appendix 2).

Each disk was immediately placed into an individual Ziploc bag until its fresh mass was weighed.

Volume was calculated using the water displacement method (Chave et al. 2005), with large disks split into quarters (or halves) before being submerged. Disks were then placed into individual paper bags and oven-dried to constant weight at 60º C. Bark thickness was calculated from fresh thick and thin disks as the mean of four measurements taken in mm around the circumference. Basic

7 density was calculated, for each disk, as oven-dry weight divided by wet volume (Fearnside 1997).

Moisture content was calculated as (fresh weight – dry weight) divided by dry weight x 100% (L.

Boddy; pers. comm.).

To analyze nitrogen content, we removed the bark from one disk per plant individual from thick and thin branches, and ground sections of the outermost sapwood into fine powder using a metal file. We placed 8-9 mg of each sapwood sample into a tin capsule and sent the samples to the Stable Isotope Paleo Environments Research Group (SIPERG, Ames, IA) for total nitrogen analysis. The SIPERG lab used a ThermoFinnigan Delta Plus XL mass spectrometer in conjunction with a Gas Bench II and CombiPal autosampler to obtain % N and d15N (the ratio of heavy to light nitrogen, relative to that found in the atmosphere). This study focused on % N, which was converted to mg/g prior to statistical analysis (Whigham et al. 2013).

Data Analyses. All analyses were performed in JMP-SAS (11.0.0). We first conducted T- tests to determine if there were trait differences between thick and thin branches. In subsequent tests, we analyzed data from thick and thin branches separately. We first tested whether dense wood was correlated with lower MC and nitrogen content. We next tested to see whether wood traits were correlated with cerambycid abundance or species richness; count data were square-root transformed prior to analysis. We calculated mean emergence week for all cerambycids emerging from a particular plant individual; these data were used in linear regressions woth wood triat data.

Mean emergence week was then calculated per cerambycid species; a t-test was conducted to test is emergemce week differed between cerambycid subfamilies with these data. Finally, we conducted T-tests to determine if there were any significant differences in the branch traits of plants colonized by cerambycine and lamiine species.

RESULTS

A total of 1,984 adult beetles were reared from 63 species; all within the Cerambycinae and Lamiinae subfamilies. More cerambycines emerged than lamiines (Appendix 1); the most abundant beetle species in this study (Haenkea thoracica Chevrolat; 785 individuals) is a member of the Cerambycinae subfamily (data not shown). More lamiine species emerged than cerambycines, with 35 Lamiinae species and 28 Cerambycinae species (Berkov 2018).

All wood traits varied among plant individuals (Table 1; Appendix 2). Relative to thick branches, thin branches had significantly thinner bark and more nitrogen (Table 2).

Table 1. Ranges of calculated wood traits. Wood Trait Range WD (g cm-3) 26.12 – 84.05

MC (%) 43.44 – 224.48 BT (mm) 0.50 – 7.25 N (mg/g) 1.10 – 11.20

Table 2. Wood traits of thick and thin branches. T-tests assume unequal variances. Thick mean = mean from thick branches. Thin mean = mean from thin branches. Std deviation = standard deviation. Significance value: p < 0.05. Wood Trait (df) t- p-value Thick mean ± Thin mean ± statistic Std deviation Std deviation WD (g cm-3) 81 -0.72 0.475 54.41 ± 12.47 52.44 ± 13.36 No Significance

MC (%) 64 1.96 0.054 80.29 ± 25.55 94.43 ± 39.44 No Significance BT (mm) 71 -5.09 p < 2.60 ± 1.26 1.56 ± 0.59 Thick > Thin 0.0001 N (mg/g) 60 3.79 p < 3.08 ± 1.58 4.86 ± 2.64 Thick < Thin 0.0004

Due to significant differences in BT and N, data from thick and thin branches were analyzed separately in subsequent analyses.

Fig. 1. WD vs. MC in thick and thin branches. Thick branches = red: R2 = 0.78, p < 0.0001. Thin branches = blue: R2 = 0.59, p < 0.0001. Significance value: p < 0.05.

Dense wood from thick and thin branches were low in moisture (Fig. 1). There were no significant correlations with nitrogen (WD vs. N (mg/g): thick branches, R2 = 0.01, p = 0.836; thin branches, R2 = 0.10, p = 0.141. MC vs. N (m/mg): thick branches, R2 = 0.01, p = 0.607; thin branches, R2 = 0.03, p = 0.389).

A B

Figs. 2A-C. Wood trait data vs. cerambycid abundance. Abundance data were square root transformed; thick branches = red, thin branches = blue. 2A: WD vs. cerambycid abundance: Red: R2 = 0.02, p = 0.377. Blue: R2 = 0.12, p = 0.028. 2B: MC vs. cerambycid abundance: Red: R2 = 0.00, p = 0.660. Blue: R2 = 0.10, p = 0.044. 2C: BT vs. cerambycid abundance: Red: R2 = 0.02, p = 0.345. Blue: R2 = 0.15, p = 0.015. Significance value: p < 0.05.

More cerambycids emerged from thin branches with lower WD, higher MC, and thicker bark (Figs. 2A-C). There was no significant correlation with nitrogen (N (mg/g) vs. cerambycid abundance: thick branches, R2 = 0.00, p = 0.953; thin branches, R2 = 0.01, p = 0.546). In thick branches, there were no significant correlations with any wood traits.

A B

Figs. 3A-B. Wood trait data vs. cerambycid species richness. Abundance data were square root transformed; thick branches = red, thin branches = blue. 3A: MC vs. cerambycid species richness: Red: R2 = 0.02, p = 0.294. Blue: R2 = 0.15, p = 0.015. 3B: BT vs. cerambycid species richness: Red: R2 = 0.02, p = 0.337. Blue: R2 = 0.23, p = 0.002. Significance value: p < 0.05.

Thin branches with higher MC and thicker bark yielded more cerambycid species (Figs.

3A-B). There were no significant correlations with wood density and nitrogen with species richness (WD vs. cerambycid species richness: thick branches, R2 = 0.00, p = 0.652; thin branches,

R2 = 0.02, p = 0.428. N (mg/g) vs. cerambycid species richness: thick branches, R2 = 0.09, p =

0.358; thin branches, R2 = 0.06, p = 0.126). In thick branches, there were no significant correlations with any wood traits.

A B

Figs. 4A-B. Wood trait data vs. estimated cerambycid mean emergence week. Thick branches = red, thin branches = blue. 4A: WD vs. cerambycid mean emergence week: Red: R2 = 0.15, p = 0.006. Blue: R2 = 0.29, p < 0.0003. 4B: MC vs. cerambycid mean emergence week: Red: R2 = 0.11, p = 0.017. Blue: R2 = 0.12, p = 0.027. Significance value: p < 0.05.

Cerambycids emerged from more rapidly from branches with low WD and high MC (Figs.

4A-B). There were no significant correlations with nitrogen and bark thickness, and mean emergence week (N (mg/g) vs. estimated cerambycid mean emergence week: thick branches, R2

= 0.05, p = 0.103; thin branches, R2 = 0.00, p = 0.881. BT vs. estimated cerambycid mean emergence week: thick branches, R2 = 0.00, p = 0.970; thin branches, R2 = 0.06, p = 0.144).

Table 3. T-tests with wood traits and emergence week, and cerambycid subfamilies. Ceram. = Cerambycinae mean. Lam. = Lamiinae mean. Std deviation = standard deviation. Significance value: p < 0.05. (df) t-statistic p-value Ceram. ± Std Lam. ± Std deviation deviation WD (g cm-3) 602 -7.30 p < 0.0001 53.99 ± 11.33 47.39 ± 10.87

MC (%) 599 6.40 p < 0.0001 87.05 ± 29.62 102.81 ± 30.87

Mean Emergence 34 -3.65 p < 0.0004 23.39 ± 14.79 12.51 ± 6.06 Week

Cerambycine host plants were significantly higher in WD and lower in MC than lamiine host plants (Table 3). Cerambycine species also emerged significantly later than the lamiine species.

DISCUSSION

We report a negative correlation between WD and MC – a result consistent with previous findings that dense wood has comparatively few metabolically active parenchyma cells, and that less dense wood has more parenchyma, increased moisture capacity, and fewer lignified cells (Fig.

1; Borchert and Pockman 2005; Dias and Marenco 2015; Nogueira et al. 2008). However, some studies in tropical forests have detected higher concentrations of parenchyma in denser wood

(Martin et al. 2014). These conflicting results might be explained by different volumes and functions of ray (lateral) and axial parenchyma (Martínez-Cabrera et al. 2009); these differences in the ratios of these metabolically active cells in our wood samples might also explain the lack of significant correlations with N. Although, our most productive wood samples tended to higher in

N (data not shown), Further work needs to be conducted to analyze the different ratios of parenchyma in our wood samples to better understand potential relationships between wood anatomy and N content.

The dry conditions that prevailed when we felled and exposed the trees might explain why more cerambycids (in abundance and species richness) emerged from branches that were relatively moist and had thicker bark (Fig. 2A-C, Fig. 3A-B). These dry conditions might have been harsh enough that the need for protection was prioritized over a preference for N, even though, in undecayed dead wood, nitrogen is considered the most limiting factor for cerambycid larvae

(Filipiak and Weiner 2017). Further research on the influence of wood traits on cerambycid communities in different climactic conditions needs to be conducted to support to this idea.

In addition to potentially protecting cerambycid larvae from desiccation, wood moisture content appears to affect life cycle duration. Increased MC and decreased WD were both associated with earlier adult emergence (Figs. 4A-B). We cannot calculate the duration of cerambycid life cycles because the branches were exposed for two months and we do not know the dates of oviposition. However, all trees were felled within one week, and cerambycids were visiting the baits within a few days (Berkov, pers. obs.). Because wood density is related to the proportion of heavily lignified cells, these data are consistent with the inherent challenges of lignin digestion.

Cerambycine species emerged later, and out of denser, drier wood, than lamiine species (Table 3).

In our previous rearing experiments, cerambycines emerged preferentially from warmer, drier, microhabitats (Berkov 2018); this study suggests that a preference for dry conditions may extend to the feeding substrate. Cerambycines could therefore be more resistant to the longer, more severe dry seasons that are predicted to occur in moist neotropical forests due to climate change (Allen et al. 2015; Greenwood et al. 2017). Also, since some cerambycine adults feed and mate on flowers

(Millar and Hanks 2017), later emergence could have a selective advantage—if it ensures synchrony with flower blooming during the dry season (Noguera 2017). However, more research would need to be conducted to support this.

Overall, the wood traits that we measured explained relatively little of the abundance or species richness of cerambycids. Bark thickness seemed to have the most predictive power, as noted in the community of primary saproxylic beetles associated with Scots pine (Foit 2010).

Wood density and moisture content appeared to play fundamental roles in explaining community composition, probably through a direct role in substrate digestibility. We might therefore expect

15 that there would be differences in the gut microbiomes of Neotropical cerambycines and lamiines due to their preferential associations with host plants of differing wood density. Due to the importance of cerambycids as early colonists of moribund wood, continuing to investigate their wood trait preferences is vital—especially given the increasing occurrence of drought events as a result of climate change (Allen et al. 2015; Greenwood et al. 2017). Trees with denser wood have lower mortality rates during drought events (Greenwood et al. 2017). Cerambycid host plants with varying wood traits, along with the cerambycids that depend on them, could therefore be affected differently by these events.

ACKNOWLEDGMENTS

We would like to thank the National Science Foundation (NSF-REU program), PSC

CUNY Research Foundation (Award #41), an anonymous donor, and the MARC/RISE fellowship

(Rise grant: 5R25GM056833-15) for providing funding to complete this project. We would also like to thank the Smithsonian Tropical Research Institute (STRI) for aiding in the 2010-2011 rearing study conducted by Dr. Berkov in Panama, and the entire team that helped in the field during that project (permits: Dr. Hector Barrios (UP, STRI), Hortensia Broce (ACP), Lil Camacho

(STRI); logistics and field support: Hector Barrios, Joyce Fassbender, Teofilio Gordon, Basilio

Gordon, Chris Roddick, Sara Pinzon, Marleny Rivera, and additional staff at STRI). We would like to thank Andres Hernandez (STRI) and Dr. Miguel Monné (MNRJ) for their help with determinations of plants and cerambycids, respectively. I would like to thank Dr. Amy Berkov,

Dr. David Lohman, Dr. Ana Carnaval, and the entire Berkov lab for their support and guidance during this project.

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APPENDICES Appendix 1: Beetle and host plant occurrence data from plant individuals. Productive and non-productive plant individuals shown. Code = plant individual code. Diameter = plant sample branch diameter. Ceram. = # Cerambycinae individuals. Lam. = # Lamiinae individuals. Plant Family Plant Species Code Diameter Ceram. Lam. Anacardiaceae Anacardium excelsum L. 8 Thick 5 34

Spondias radkoferi Donn.Sm. 57 Thick 1 27 Spondias radkoferi Donn.Sm. 57 Thin - 2 Spondias radkoferi Donn.Sm. 58 Thick - 2 Araliaceae Dendropanax arboreus (L.) 65 Thin - 7 Decne. & Planch. Arecaceae Attalea butyracea (Mutis ex L. 1 (photo - - - f.) Wess. Boer only) Bignoniaceae Tababuia ochracea (Cham.) 66 Thick 2 - Standl. Tababuia ochracea (Cham.) 66 Thin 1 - Standl. Boraginaceae Cordia bicolor A. DC. Ex DC. 52 - - - Burseraceae Bursera simaruba (L.) Sarg. 63 - - - Protium tenuifolium (Engl.) 45 Thick - 4 Engl. Combretaceae Terminalia oblonga (Ruiz & 2 Thick 4 - Pav.) Steud. Terminalia oblonga (Ruiz & 56 Thick 2 4 Pav.) Steud. Terminalia oblonga (Ruiz & 56 Thin - 1 Pav.) Steud. Dilleniaceae Davila nitida (Vahl) Kubitzki 49 Thick 22 - Fabaceae Dahlbergia retusa Hemsl. 64 Thick 1 - Dahlbergia retusa Hemsl. 64 Thin 2 - Dipteryx oleifera Benth. 59 Thick 1 1 Dipteryx oleifera Benth. 59 Thin 2 - Dipteryx oleifera Benth. 60 Thick 2 66 Dipteryx oleifera Benth. 60 Thin 5 4 Inga laurina (Sw.) Willd. 39 Thick 43 15 Inga laurina (Sw.) Willd. 39 Thin 7 4 Lonchocarpus sp. 40 Thick 4 6 Lonchocarpus sp. 40 Thin 1 - Lonchocarpus sp. 50 Thick 6 39 Lonchocarpus sp. 50 Thin - 5 Swartzia simplex (Sw.) Spreng. 34 Thick 5 - Swartzia simplex (Sw.) Spreng. 34 Thin 3 - Swartzia simplex (Sw.) Spreng. 35 Thick 23 - Swartzia simplex (Sw.) Spreng. 35 Thin 3 - Swartzia simplex (Sw.) Spreng. 36 Thick 6 -

Swartzia simplex (Sw.) Spreng. 36 Thin 3 - Tachigali versicolor Standl. & 28B Thick 4 - L.O. Williams Tachigali versicolor Standl. & 28B Thin 2 1 L.O. Williams Lecythidaceae Gustavia superba (Kunth) O. 13 Thick 16 8 Berg Gustavia superba (Kunth) O. 13 Thin 15 6 Berg Gustavia superba (Kunth) O. 14 Thick 34 25 Berg Gustavia superba (Kunth) O. 14 Thin 34 72 Berg Gustavia superba (Kunth) O. 15 Thick 18 37 Berg Gustavia superba (Kunth) O. 15 Thin 47 12 Berg Gustavia superba (Kunth) O. 16 Thick 7 38 Berg Gustavia superba (Kunth) O. 16 Thin 12 7 Berg Malpighiaceae Heteropsis laurifolia Marshall 47 Thick 1 - Malvaceae Cavanillesia platanifolia 0 - - - (Humb. & Bonpl.) Kunth Luehea seemannii Triana & 44 Thick 47 5 Planch. Luehea seemannii Triana & 44 Thin 2 2 Planch. Luehea seemannii Triana & 53 Thick 1 - Planch. Luehea seemannii Triana & 53 Thin 34 12 Planch. Luehea speciosa Willd. 24 Thin 3 - Luehea speciosa Willd. 32 Thick 7 - Luehea speciosa Willd. 32 Thin 2 - Pachira sessilis Benth. 29 Thick - 3 Pachira sessilis Benth. 29 Thin - 1 Pachira sessilis Benth. 31 Thick - 7 Pachira sessilis Benth. 31 Thin - 8

Sterculia apetala (Jacq.) H. 51 Thick - 26 Karst Sterculia apetala (Jacq.) H. 51 Thin - 44 Karst Melastomataceae Miconia argentea (Sw.) DC. 12 - - - Meliaceae Trichilia tuberculata (Triana & 37 Thin 1 - Planch.) C.DC. Trichilia tuberculata (Triana & 41 Thick 4 - Planch.) C.DC. Trichilia tuberculata (Triana & 42 Thick 4 - Planch.) C.DC. Moraceae Castilia elastica Cerv. 19 Thick - 2 Castilia elastica Cerv. 19 Thin - 1 Castilia elastica Cerv. 20 Thick - 24 Castilia elastica Cerv. 20 Thin - 5 Castilia elastica Cerv. 21 Thick 3 8 Castilia elastica Cerv. 21 Thin - 17 Maquira costaricana (Standl.) 26 Thick 3 2 C.C. Berg Maquira costaricana (Standl.) 26 Thin - 25 C.C. Berg Maquira costaricana (Standl.) 54 Thick - 14 C.C. Berg Maquira costaricana (Standl.) 54 Thin - 8 C.C. Berg Maquira costaricana (Standl.) 7 Thick - 7 C.C. Berg Maquira costaricana (Standl.) 7 Thin - 15 C.C. Berg Trophis racemosa (L.) Urb. 62 Thick - 7 Trophis racemosa (L.) Urb. 62 Thin - 3 Myristacaceae Virola sebifera Aubl. 22 Thick 7 - Virola sebifera Aubl. 5 Thick 1 - Virola sebifera Aubl. 6 Thick - 5 Virola sebifera Aubl. 9 Thick 3 - Nyctaginaceae Guapira standleyana Woodson 38 - - - Rhizophoraceae Cassipourea elliptica (Sw.) 17 Thick 4 - Poir. Cassipourea elliptica (Sw.) 18 Thick - 13 Poir.

Cassipourea elliptica (Sw.) 18 Thin 1 - Poir. Cassipourea elliptica (Sw.) 30 Thick - 1 Poir. Cassipourea elliptica (Sw.) 30 Thin - 2 Poir. Rubiaceae Alseis blackiana Hemsl. 10 Thick 45 - Alseis blackiana Hemsl. 10 Thin 62 - Alseis blackiana Hemsl. 11 Thick 152 1 Alseis blackiana Hemsl. 11 Thin 18 - Alseis blackiana Hemsl. 27 Thick 158 1 Alseis blackiana Hemsl. 27 Thin 69 - Alseis blackiana Hemsl. 28A Thick 221 - Alseis blackiana Hemsl. 28A Thin 51 - Macrocnemum roseum (Ruiz & 23 - - - Pav.) Wedd. Salicaceae Tetrathylacium johansenii 25 Thick - 1 Standl. Sapotaceae Chrysophyllum cainito L. 3 Thick 35 - Chrysophyllum cainito L. 3 Thin 5 - Verbenaceae Petrea aspera Turcz. 48 - - - Total of beetle 1287 697 individuals N = 1984

Appendix 2: Plant trait data from productive plant individuals. Only plants that yielded beetles are included. Code = plant individual code. Diameter = plant sample diameter. WD = wood density (g cm-3). MC = moisture content (%). BT = bark thickness (mm). N = nitrogen (mg/g). Plant Family Plant Species Code Diameter WD MC BT N Anacardiaceae Anacardium 8 Thick 39.84 97.65 2.00 4.00 excelsum L. Spondias radkoferi 57 Thick 37.94 104.37 1.56 3.30 Donn.Sm. Spondias radkoferi 57 Thin 38.87 104.49 2.00 1.60 Donn.Sm. Spondias radkoferi 58 Thick 38.80 104.49 2.00 2.60 Donn.Sm. Araliaceae Dendropanax 65 Thin 39.39 108.92 1.56 2.00 arboreus (L.) Decne. & Planch.

Bignoniaceae Tababuia ochracea 66 Thick 62.01 67.14 2.31 8.90 (Cham.) Standl. Tababuia ochracea 66 Thin 64.76 62.55 3.06 6.20 (Cham.) Standl. Protium tenuifolium 45 Thick 57.95 48.70 2.06 1.50 (Engl.) Engl. Combretaceae Terminalia oblonga 2 Thick 56.84 71.42 1.44 2.60 (Ruiz & Pav.) Steud. Terminalia oblonga 56 Thick 58.10 70.59 1.38 4.50 (Ruiz & Pav.) Steud. Terminalia oblonga 56 Thin 61.29 60.49 1.44 2.00 (Ruiz & Pav.) Steud. Dilleniaceae Davila nitida 49 Thick 53.15 102.37 7.25 2.60 (Vahl) Kubitzki Fabaceae Dahlbergia retusa 64 Thick 65.80 60.15 1.69 3.30 Hemsl. Dahlbergia retusa 64 Thin 71.65 57.24 3.88 1.60 Hemsl. Dipteryx oleifera 59 Thick 65.92 69.15 1.06 2.70 Benth. Dipteryx oleifera 59 Thin 74.54 49.97 1.50 1.10 Benth. Dipteryx oleifera 60 Thick 72.45 63.95 1.63 10.30 Benth. Dipteryx oleifera 60 Thin 77.60 49.05 1.81 1.80 Benth. Inga laurina (Sw.) 39 Thick 65.63 67.73 3.38 3.40 Willd. Inga laurina (Sw.) 39 Thin 68.59 62.50 2.50 7.60 Willd. Lonchocarpus sp. 40 Thick 54.11 88.61 1.75 4.20 Lonchocarpus sp. 40 Thin 54.19 84.72 2.38 3.90 Lonchocarpus sp. 50 Thick 53.46 98.08 1.88 15.40 Lonchocarpus sp. 50 Thin 57.84 76.64 2.69 3.40 Swartzia simplex 34 Thick 76.07 46.85 0.75 5.20 (Sw.) Spreng.

Swartzia simplex 34 Thin 84.05 43.44 1.25 2.70 (Sw.) Spreng. Swartzia simplex 35 Thick 78.53 46.20 2.19 4.80 (Sw.) Spreng. Swartzia simplex 35 Thin 79.26 45.13 1.44 4.80 (Sw.) Spreng. Swartzia simplex 36 Thick 76.16 48.79 0.94 4.30 (Sw.) Spreng. Swartzia simplex 36 Thin 78.48 44.49 1.56 2.60 (Sw.) Spreng. Tachigali 28B Thick 63.06 70.41 1.94 3.80 versicolor Standl. & L.O. Williams Tachigali 28B Thin 66.13 57.36 2.88 1.70 versicolor Standl. & L.O. Williams Lecythidaceae Gustavia superba 13 Thick 45.01 127.81 2.13 8.10 (Kunth) O. Berg Gustavia superba 13 Thin 51.53 101.85 3.56 2.90 (Kunth) O. Berg Gustavia superba 14 Thick 39.38 160.98 2.56 4.80 (Kunth) O. Berg Gustavia superba 14 Thin 50.18 110.46 4.25 3.50 (Kunth) O. Berg Gustavia superba 15 Thick 46.35 124.07 2.56 6.10 (Kunth) O. Berg Gustavia superba 15 Thin 51.76 104.67 4.44 3.80 (Kunth) O. Berg Gustavia superba 16 Thick 43.26 124.09 3.81 3.50 (Kunth) O. Berg Gustavia superba 16 Thin 44.07 136.32 1.81 10.80 (Kunth) O. Berg Malpighiaceae Heteropsis 47 Thick 54.70 99.60 2.30 11.20 laurifolia Marshall Luehea seemannii 44 Thick 41.05 86.92 1.56 3.80 Triana & Planch. Luehea seemannii 44 Thin 45.35 77.81 2.38 2.00 Triana & Planch. Luehea seemannii 53 Thick 31.46 154.05 1.63 2.50 Triana & Planch.

Luehea seemannii 53 Thin 38.80 110.87 1.94 1.80 Triana & Planch. Luehea speciosa 24 Thin 48.97 99.00 1.69 2.20 Willd. Luehea speciosa 32 Thick 52.95 84.62 1.94 4.40 Willd. Luehea speciosa 32 Thin 55.35 71.04 3.50 4.10 Willd. Pachira sessilis 29 Thick 48.80 66.62 1.13 4.60 Benth. Pachira sessilis 29 Thin 49.89 59.56 1.38 4.90 Benth. Pachira sessilis 31 Thick 51.23 98.33 3.06 2.00 Benth. Pachira sessilis 31 Thin 52.13 91.95 2.44 3.00 Benth. Sterculia apetala 51 Thick 31.32 160.06 2.38 3.90 (Jacq.) H. Karst Sterculia apetala 51 Thin 35.61 138.93 5.38 2.30 (Jacq.) H. Karst Meliaceae Trichilia 37 Thin 65.92 51.88 0.94 3.40 tuberculata (Triana & Planch.) C.DC. Trichilia 41 Thick 64.52 51.71 3.31 2.50 tuberculata (Triana & Planch.) C.DC. Trichilia 42 Thick 66.72 50.27 3.88 2.10 tuberculata (Triana & Planch.) C.DC. Moraceae Castilia elastica 19 Thick 26.12 224.48 1.75 4.80 Cerv. Castilia elastica 19 Thin 33.47 141.58 2.56 3.50 Cerv. Castilia elastica 20 Thick 31.49 173.58 1.88 6.30 Cerv. Castilia elastica 20 Thin 38.51 95.11 2.94 1.60 Cerv. Castilia elastica 21 Thick 37.64 137.36 1.81 5.90 Cerv.

Castilia elastica 21 Thin 41.49 100.05 2.88 1.90 Cerv. Maquira 26 Thick 47.79 95.09 1.25 10.10 costaricana (Standl.) C.C. Berg Maquira 26 Thin 52.70 77.04 1.81 2.70 costaricana (Standl.) C.C. Berg Maquira 54 Thick 47.86 86.01 0.88 3.60 costaricana (Standl.) C.C. Berg Maquira 54 Thin 50.81 79.99 1.75 4.40 costaricana (Standl.) C.C. Berg Maquira 7 Thick 47.44 80.84 0.69 6.40 costaricana (Standl.) C.C. Berg Maquira 7 Thin 51.03 73.58 1.19 4.50 costaricana (Standl.) C.C. Berg Trophis racemosa 62 Thick 43.13 104.48 0.50 1.70 (L.) Urb. Trophis racemosa 62 Thin 43.13 104.48 0.50 2.50 (L.) Urb. Myristacaceae Virola sebifera 22 Thick 55.06 61.47 1.38 2.10 Aubl. Virola sebifera 5 Thick 43.52 84.00 2.13 3.40 Aubl. Virola sebifera 6 Thick 36.64 96.07 2.94 2.50 Aubl. Virola sebifera 9 Thick 40.20 91.12 1.44 3.90 Aubl. Rhizophoraceae Cassipourea 17 Thick 60.71 76.19 1.75 2.80 elliptica (Sw.) Poir. Cassipourea 18 Thick 60.44 70.71 1.63 3.00 elliptica (Sw.) Poir. Cassipourea 18 Thin 60.52 77.54 0.75 6.20 elliptica (Sw.) Poir. Cassipourea 30 Thick 60.77 72.88 1.63 3.20 elliptica (Sw.) Poir. Cassipourea 30 Thin 67.32 68.83 1.13 2.60 elliptica (Sw.) Poir.

Rubiaceae Alseis blackiana 10 Thick 53.83 51.87 1.13 3.80 Hemsl. Alseis blackiana 10 Thin 55.80 60.22 1.19 5.70 Hemsl. Alseis blackiana 11 Thick 51.39 56.43 0.81 2.80 Hemsl. Alseis blackiana 11 Thin 55.56 67.69 0.75 4.50 Hemsl. Alseis blackiana 27 Thick 56.38 66.96 1.44 5.10 Hemsl. Alseis blackiana 27 Thin 57.01 65.25 1.75 4.80 Hemsl. Alseis blackiana 28A Thick 34.63 109.67 4.06 3.00 Hemsl. Alseis blackiana 28A Thin 44.00 100.26 2.56 4.10 Hemsl. Salicaceae Tetrathylacium 25 Thick 44.61 127.65 4.81 2.20 johansenii Standl. Sapotaceae Chrysophyllum 3 Thick 56.83 84.81 0.75 6.30 cainito L. Chrysophyllum 3 Thin 59.01 79.67 2.75 2.30 cainito L.

Appendix 3: Sampling of branch sections collected. Only plants that yielded beetles are included. Code = plant individual code. Branch section type = thickness of branches (thick or thin) and stratum (canopy or ground) from where branch sections were taken from. Plant Family Plant Species Code Branch Section Type Anacardiaceae Anacardium 8 Thick ground excelsum L. Spondias radkoferi 57 Thick canopy, thick ground, thin Donn.Sm. ground Spondias radkoferi 58 Thick ground Donn.Sm. Araliaceae Dendropanax 65 Thin ground arboreus (L.) Decne. & Planch. Bignoniaceae Tababuia ochracea 66 Thick ground, thin ground (Cham.) Standl. Protium tenuifolium 45 Thick ground (Engl.) Engl.

Combretaceae Terminalia oblonga 2 Thick canopy, thick ground (Ruiz & Pav.) Steud. Terminalia oblonga 56 Thick ground, thin ground (Ruiz & Pav.) Steud. Dilleniaceae Davila nitida 49 Thick canopy, thick ground (Vahl) Kubitzki Fabaceae Dahlbergia retusa 64 Thick ground, thin ground Hemsl. Dipteryx oleifera 59 Thick ground, thin ground Benth. Dipteryx oleifera 60 Thick ground, thin ground Benth. Inga laurina (Sw.) 39 Thick canopy, thick ground, thin Willd. ground Lonchocarpus sp. 40 Thick canopy, thick ground, thin ground Lonchocarpus sp. 50 Thick canopy, thick ground, thin ground Swartzia simplex 34 Thick ground, thin ground (Sw.) Spreng. Swartzia simplex 35 Thick canopy, thick ground, thin (Sw.) Spreng. ground Swartzia simplex 36 Thick canopy, thin ground (Sw.) Spreng. Tachigali 28B Thick canopy, thin ground versicolor Standl. & L.O. Williams Lecythidaceae Gustavia superba 13 Thick ground, thin ground (Kunth) O. Berg Gustavia superba 14 Thick canopy, thick ground, thin (Kunth) O. Berg ground Gustavia superba 15 Thick canopy, thick ground, thin (Kunth) O. Berg ground Gustavia superba 16 Thick canopy, thick ground, thin (Kunth) O. Berg ground Malpighiaceae Heteropsis 47 Thick ground laurifolia Marshall

Luehea seemannii 44 Thick canopy, thick ground, thin Triana & Planch. ground Luehea seemannii 53 Thick ground, thin ground Triana & Planch. Luehea speciosa 24 Thin ground Willd. Luehea speciosa 32 Thick ground, thin ground Willd. Pachira sessilis 29 Thick ground, thin ground Benth. Pachira sessilis 31 Thick canopy, thick ground, thin Benth. ground Sterculia apetala 51 Thick ground, thin ground (Jacq.) H. Karst Meliaceae Trichilia 37 Thin ground tuberculata (Triana & Planch.) C.DC. Trichilia 41 Thick ground tuberculata (Triana & Planch.) C.DC. Trichilia 42 Thick ground tuberculata (Triana & Planch.) C.DC. Moraceae Castilia elastica 19 Thick canopy, thin ground Cerv. Castilia elastica 20 Thick canopy, thick ground, thin Cerv. ground Castilia elastica 21 Thick canopy, thick ground, thin Cerv. ground Maquira 26 Thick canopy, thick ground, thin costaricana ground (Standl.) C.C. Berg Maquira 54 Thick grounf, thin ground costaricana (Standl.) C.C. Berg Maquira 7 Thick canopy, thick ground, thin costaricana ground (Standl.) C.C. Berg Trophis racemosa 62 Thick ground, thin ground (L.) Urb.

Myristacaceae Virola sebifera 22 Thick ground Aubl. Virola sebifera 5 Thick ground Aubl. Virola sebifera 6 Thick ground Aubl. Virola sebifera 9 Thick ground Aubl. Rhizophoraceae Cassipourea 17 Thick ground elliptica (Sw.) Poir. Cassipourea 18 Thick canopy elliptica (Sw.) Poir. Cassipourea 30 Thick ground, thin ground elliptica (Sw.) Poir. Rubiaceae Alseis blackiana 10 Thick canopy, thick ground, thin Hemsl. ground Alseis blackiana 11 Thick canopy, thick ground, thin Hemsl. ground Alseis blackiana 27 Thick canopy, thick ground, thin Hemsl. ground Alseis blackiana 28A Thick canopy, thick ground, thin Hemsl. ground Salicaceae Tetrathylacium 25 Thick canopy johansenii Standl. Sapotaceae Chrysophyllum 3 Thick canopy, thick ground, thin cainito L. ground