Crown Structure & the Canopy Arthropod Biodiversity of 100 Year Old and Old Growth Tasmanian Eucalyptus obliqua

by

Yoav Daniel Bar-Ness B.Sc.(Wildlife Sciences)

Submitted in fulfilment of the requirements for the degree of Masters (Research) of Environmental Studies, University of Tasmania, January 2005

Table of Contents Page Section 5 …………Statement of Originality 6 …………Abstract: 7 …………Acknowledgments 11 …………1) Introduction: 11 …………1.1 Biodiversity and forest structure 17 …………1.2 Research questions 1) What are the structural differences between 100 year old and old-growth E. obliqua, and how can they be measured and displayed? 2) How is the canopy arthropod biodiversity different in 100 year old and old-growth E. obliqua? 3) In what ways does crown structure influence arthropod biodiversity in E. obliqua? 18 …………1.3 Limitations of the present study 19 ………… 2) Aims and Background 19 …………2.1 Addressing the research questions 24 …………2.2 Background Research 48 …………2.3 Synthesis of previous research 49 …………3) Study Design and Site 49 …………3.1 Study Subject and Site 55 …………3.3 Study design 64 …………4) Crown Structure – Methods 64 …………4.1 Definition of tree elements 75 …………4.2 Structural data collection 84 …………4.3 Field Recordings 90 …………4.4 Vector derived variable generation 93 …………4.5 Tree Structural Descriptor Generation 95 …………4.6 Analysis of structural descriptors 96 …………4.7 Arborograph generation and analysis 98 …………4.8 Image file size analysis 101 …………4.9 Structural features 103 …………5) Crown Structure – Results 105 …………5.1 Total numbers 105 …………5.2 Summary and range of structural descriptors. 106 …………5.3 Univariate T-tests 117 …………5.4 Multivariate classifications 119 …………5.5 File Size comparison 120 …………5.6 Structural features 134 …………5.7 Other untabulated features 137 …………6) Crown Structure – Discussion 137 …………6.1 Distinctness of crowns 151 …………6.2 Variability in tree crowns 155 …………6.3 Incidence of structural features 157 …………6.4 More complexity of information Table of Contents Page Section 161 …………7) Canopy Arthropods - Methods 161 …………7.1 Trap design types 172 …………7.2 Trap processing 173 …………7.3 Data Analysis 175 …………7.4 Analysis Approaches 176 …………7.5 Biodiversity aspects 182 …………7.6 Scaling of results 190 …………8) Canopy Arthropods –Results 190 …………8.1 Trap survival 192 …………8.2 Total collection overview 192 …………8.3 Trap type comparison 194 …………8.4 Abundance results 199 …………8.5 Richness results 204 …………8.6 Community structure results 206 …………8.7 Diversity results 211 …………8.8 Species accumulation and estimated richness results

214 …………8.9 Distinctness results 234 …………8.10 RTU level morphospecific results 236 …………9) Canopy Arthropods – Discussion 236 …………9.1 Abundance of faunal activity 238 …………9.2 Richness of fauna 242 …………9.3 Diversity of fauna 245 …………9.4 Distinctness of fauna 250 …………10) Correlation – Methods 250 …………10.1 Exploring correlations between structure and arthropod biodiversity 251 …………10.2 Bivariate Correlations 254 …………10.3 Multivariate correlations 255 …………11) Correlation – Results 255 …………11.1 Bivariate results 264 …………11.2 Multivariate Correlation results 268 …………12) Correlation – Discussion 268 …………12.1 The influence of crown structure on canopy arthropods of E. obliqua 269 …………12.2 Bivariate 271 …………12.3 Multivariate compositional predictors 272 …………12.4 Surrogate measures 272 …………12.5 Applicability Table of Contents Page Section 274 …………13) Conclusions 274 …………13.1 Mapping Aim 1: Quantitatively assess the differences in crown structure and size between 100yr and old E. obliqua 275 …………13.2 Mapping Aim 2: Investigate the presence of structural features in the crowns of 100yr and old E. obliqua 276 …………13.3 Mapping Aim 3: Modify the conifer mapping technique of Van Pelt et al. (2004b) for Eucalyptus trees 278 …………13.4 Mapping Aim 4: Develop a technique for displaying crown structure of forest trees using computer models based on 3-dimensional spherical coordinates 279 …………13.5 Mapping Aim 5: Expand Jacobs 1955 theory of intra- branch competition in Eucalyptus saplings to mature and old-growth E. obliqua 280 …………13.6 Mapping Aim 6: Generate a predictor data set to explore the influence of crown structure on canopy arthropods 280 …………13.7 Trapping Aim 1: Determine what differences exist in arthropod biodiversity between 100 year old and old growth Eucalyptus obliqua 281 …………13.8 Trapping Aim 2:Contribute to the knowledge of Eucalyptus canopy arthropods 281 …………13.9 Trapping Aim 3: Develop robust, inexpensive trap designs suitable for transport to and use in E. obliqua 282 …………13.10 Trapping Aim 4: Generate a response data set to explore the influence of crown structure on canopy arthropods 283 …………13.11 Conclusions: Answering the Research Questions 1) What are the structural differences between 100 year old and old-growth E. obliqua, and how can they be measured and displayed? 2) How is the canopy arthropod biodiversity different in 100 year old and old-growth E. obliqua? 3) In what ways does crown structure influence arthropod biodiversity in E. obliqua? 285 …………13.12 Future directions 287 …………14) References (311) …………15) Appendix 1. Arborograph Line Up (312) …………16) Appendix 2 Arborograph CAD Script Generation (313) …………17) Appendix 3. Sample Arborograph Script (353) …………18) Appendix 4. Blank Data Sheet (354) …………19) Appendix 5. Sample Data Sheet (355) …………20) Appendix 6. RTU listing (359) …………21) Appendix 7. A Review of Canopy Arthropod Biodiversity Research in Australian Eucalyptus (389) …………22) Appendix 8. Additional Spider Taxonomy (391) …………23) Appendix 9. Arthropod Voucher Photograph Collection

Please note that PDF computer files are electronically bookmarked Crown Structure & the Canopy Arthropod Biodiversity of 100 Year Old and Old Growth Tasmanian Eucalyptus obliqua

by

Yoav Daniel Bar-Ness B.Sc.(Wildlife Sciences)

Submitted in fulfilment of the requirements for the degree of Masters (Research) of Environmental Studies, University of Tasmania, January 2005

This thesis contains no material which has been accepted for a degree or diploma by the University or any other institution, except by way of background information and duly acknowledged in the thesis, and to the best of the my knowledge and belief, contains no material previously published or written by another person except where due acknowledgement is made in the text of the thesis.

Yoav Daniel Bar-Ness

This thesis may be made available for loan and limited copying in accordance with the Copyright Act 1968. Abstract:

The crown structure and canopy arthropods of Eucalyptus obliqua were studied at the Warra Long Term Ecological Research Site in the wet sclerophyll forests of Southern Tasmania. Eight 100 year old and eight old-growth (between 300-500 years old) trees were studied in pairs. Despite their critical role in the economy and ecology of Tasmania, few studies have addressed the canopy arthropod biodiversity of these trees. The crown structure was mapped by recording a vector in spherical space for every branch. 3- dimensional computer models were generated to aid in illustrating the tree maps. Sticky traps, flight intercept traps, and funnel crawl traps were placed in the crown to sample mobile arthropods.

The age classes were quantifiably different in crown structure. 100 year old trees had young mature crowns of original branches. Old trees had either senescent original crowns or resprouted epicormic secondary crowns. Old trees expressed a greater variability in crown structure, and were more complex as measured by the amount of information required to display the computer image. Structural attributes such as hollows, dead tops, dead flanks, trunk fissures, burls, vascular epiphytes, snapped trunks, and litter collections were all present more often in the old trees than the 100 year old trees.

Old trees showed a more diverse community structure and the evidence suggested a more rich fauna in these trees. The age classes showed differences in composition that were often masked by environmental effects.

The structural descriptors were tested against the arthropod biodiversity variables to explore the influence of crown structure on canopy arthropod communities. In the context of the study trees, several rank correlations were identified. Old trees with a higher portion of their total wood volume in the trunk had a more diverse total arthropod fauna. Trees with less crown depth had a richer Diptera fauna. Trees with a narrower range of dead branch starting diameters had a more diverse sticky trap catch. Trees with a wider range of live branch starting diameters had a richer hangtrap catch. Old trees with greater mean upwards arc had a richer hangtrap catch. Trees with a lower mean branch height as % of total height had a richer hangtrap catch. No other study is known to have investigated the influence of crown structure on canopy arthropod biodiversity in a forest tree.

Dr. Jamie Kirkpatrick & Dr. Peter McQuillan provided academic supervision in complementary ways, and were integral in making this project a reality.

The Australian Geographic Society provided material support and coverage of this project.

Vertical Pty Ltd, a West Australian manufacturer of climbing & industrial access equipment, provided material sponsorship.

Special thanks to Simon Grove Dick Bashford, Natalie Kelly, Tim Wardlaw, John Hickey, Belinda Yaxley FT Huon District: Geeveston Warra LTER FT Conservation Branch FT Aerial Photos Division

Kate Harrison, Anna Hopkins, Caroline Mohammed, Marie Yee

Field Crew: It is insufficient to merely list names. All of these people literally put their life on the line for this project. Their participation was absolutely integral. Thank you very much!

-Abby Barrows moved to Hobart from Maine and spent two months climbing trees in the field and three months in the lab sorting bugs. No other volunteer put in such a sterling effort.

-Robert Junker came out from Germany and assisted for three months of rigging, climbing, and trapsetting. With his help, all 16 study trees were scouted, rigged, climbed, and trapped. He was actively involved in the development of our own unique style of rigging.

-Melissa Whitman took a break from her research in Madagascar to spend two months in the field. She constantly developed new techniques with a huge supply of creativity. She also provided several of the photos in this document, which are denoted by her initials “MW”.

-Matt Cracknell is a constant inspiration through his safe ropework ethics and positive attitudes in the treetops and underground. Several adventures later, he still maintains the same care and calm no matter how remarkable the circumstances.

-Gene Miller -Staumm,Sophie, The Tahune Forest Airwalk incongruously helped tackle Kyatt, Joe H., & provided the luxuries of a cafe, a ramp into the virtual tree Rina G. aided in stringybark treetops, and a picnic shelter with rendering & trap placement and barbeques only a ten minute drive from camp assisted with trees mapping tree 3 stoves, ropes, and rain. 1, 2, and 4 Josh and Dylan provided excellent company at all hours. -Katy Dika and -Craig Airey Paul Ossimo helped with helped place the crossbow rigging of hangtraps tree 5 & 6 The Geography Dept's Les Wood, Paulene efficiently & Harrowby, & Moya Kilpatrick tackled effectively Occupational Health & Safety issues effectively and discretely, as well as providing assistance in -Elaine Vale countless of administrative tasks -Jerry Romanski helped with offered constructive horizontal rigging criticism and rigged of trees 12 & 16 John Stacpoole offered excellent input on tree 4 ropework, and provided an excellent deal on a motor vehicle with exquisite timing

-Wally and Eva -Michael Sumner Herrmann helped map tree 2 enthusiastically and climb tree 3 -Peter Cornish enthusiastically and creatively facilitated research and aided laser & GPS development of trap designs and rigging weaponry, and graciously surveys of study constructed several trap components trees -Denis Charlesworth assisted with processing dozens of receipts for sundry exciting items including cordage, duct tape, dowel rods, cableties, wire, -Angus Munro- several kilometres of cord, rope and climbing equipment, fishing reels and helped climb and -Sue Baker helped line, dowel rods, and duct tape, and more cord. rig trees 2 and 4, map tree 4 and -Southern Tasmanian Caverneers enthusiastically shared their rope work and contributed to offered suggestions styles during several instances underground and at South Hobart Quarry. development of on handling beetle -South Hobart Recycling Centre never even spared us a glance as we dug horizontal rigging collections through a mountain of recycled bottles. They provided hundreds of plastic skills bottles at no cost. -Australian Chainsaw in Launceston provided a large amount of climbing equipment at an excellent price -Brendan Keyes- -Ben Meadows Forestry Suppliers provided 2 60m ropes for the price of 1 assisted in mapping -Malcolm Grant suggested several design ideas for funnel traps tree 3 and provided -Bob Cotgrove graciously housed Melissa Whitman the crossbow used -Scott Sharpe -Erin Buttermore kindly hosted Robert Junker for rigging all of the helped rig tree 4 -Jilly Middleton assisted with insect curation trees and is a -Dan Bickel, Liz Turner, David Green, and Nicky Meeson expressed treeclimbing interest and helped identify animals from their favourite taxonomic groups, inspiration! flies, spiders, mites, and ants. These four taxa were the most difficult, by far. -Mike Driessen assisted in identifying the arboreal Kinemania cave cricket. -Phillip Grosse- -Tom Dunlap advised Melissa to get involved with this research project. helped rig tree 12, -Jon Majer provided useful input following a pilot study and showed trap helped flail at tree -Damien and laboratory techniques in use in W.A. 14, and pointed out Catchpole- helped -Peter Lillywhite provided access to microscopes & other resources at the that the awl on a in collecting the Melbourne Museum Swiss Army Knife traps,and drank much -Tim Sloan at Mountain Designs Hobart- provided many pieces of could be used as a peppermint tea. climbing equipment for an excellent price wood drill -Malcolm Grant, Kirra Enders, Lydia Marino, and Craig Airey actively offered hospitality, conversation about things besides trees and several much appreciated cups of tea. -Lucie Whitten -Kelly Moore helped develop early mapping skills in a Tassie Blue Gum on helped map and -Robert Shaw- Waterworks Road derig tree 13, and accompanied a -Wenda Matthews provided an excellent work environment. provided warm and scouting trip up the -Walsh Optics, Hobart provided a laser rangefinder and assisted in digital patient slopes of Mt camera repair companionship Frederick - UTAS Environmental Studies Student Society provided material support throughout a very to attend the International Congress of Entomology in Brisbane July '05. busy season. Thanks. This project was directly inspired by experiences & contact with the all of the following people: Several more people could not be listed. Thanks! -Dave James -Cara McGary- · Robert Van Pelt, the Lorax · Nalini Nadkarni assisted in mapping assisted in · Stephen Sillett · Tom Greenwood tree 15 explorations near · Jerry Franklin · Dave Shaw tree 3 · Brendan Kelly · Matt Dunlap · Kelly Gleason · Richard Gwosz · John Miller · Megan Wilson Stromberg · Bruce Rinker · Brett Nichols · Jim Spickler · Carl Gillick · Alex Zeravla· Andrew Steckler

-Jack Burns and Laurie Fox were “Ultimately, there would be only a integral supporters of this project. single tree, incorporating all living things without exception...” E.C. Pielou, Biogeography 1) Introduction and Aims

1) Introduction:

1.1 Biodiversity and forest structure

The conservation of biological diversity is an overriding concern in a world well supplied with urgent environmental issues (Hunter, 1999). Arthropods form a vast portion of the Earth’s eukaryotic biodiversity (Erwin, 2004; Gaston & Spicer, 1998). Despite their critical importance, they are under-represented and under-valued by both the general public and the decision makers (Kim, 1993; New, 1999). The range of arthropod ecosystem services and ecological niches underpins the survival of humans on this planet, and actively working towards their conservation will benefit us greatly in the future (Beattie, 1994; Grove & Stork, 2000; Kim, 1993; New, 1999). Any loss of arthropod biological diversity in the forest canopy can impact humans in many ways, including monetarily, ecologically and spiritually (Beattie, 1994; Kim, 1993; New, 1999; Yen, 1993).

Arthropod biodiversity reaches its greatest levels in forest ecosystems (Erwin, 1995). Australian Eucalyptus forests harbour a globally significant number of species (Majer et al., 1994). Like other animals in these ecosystems, their biodiversity is dependant on forest structure (Spies, 1998).

Forest management drastically alters both the structure and demographics of Eucalyptus forests (Floren et al., 2001; Lindenmayer & McCarthy, 2002; Lindenmayer et al., 2000a). Studies in other forest regions have addressed the impact of harvesting on the canopy arthropod community (Chey et al., 1998; Floren & Linsenmair, 2001; Winchester & Ring, 1996).

Very little is known about the arthropod biodiversity or the habitat structures in ancient or mature Eucalyptus trees (Majer et al., 1997). Forest management can only be considered sustainable if it maintains biodiversity (Lindenmayer, 1995). If old-growth trees are reservoirs of arthropod biodiversity, then the loss of these irreplaceable trees would be an unsustainable practice.

Alteration of Tasmanian forests: The wet sclerophyll Eucalyptus forests of Tasmania are ecologically and economically critical to the island (Dovers et al., 1994; Whiteley, 1999; Williams & Brooker, 1997; Williams & Potts, 1996), and contain the tallest and largest flowering plants in the world (Hickey et al., 2000; Kostoglou, 2000;

1) Introduction and Aims

Mifsud, 2003). Eucalyptus trees in wet sclerophyll forests grow and decay at high speeds, with an estimated lifespan of 400-500 years. (Alcorn et al., 2001; Gibbons & Lindenmayer, 1996; Lindenmayer et al., 2000a)

Forest management in Tasmania is based on an 80-100 year rotation of clearfelling, burning, and replanting of wet Eucalyptus forests (Figure 1.1). It is a politically sensitive issue and the source of a limitless quantity of public debate and spite (ForestryTasmania, 2003; Green, 2003).

Figure 1.1. Clearfelling alters the structure of Eucalyptus forests.

Forest management impacts arthropod biodiversity: A critical element of this controversy is the impact of forest management on the native fauna. Studies on the arthropods associated with E. obliqua forests play an important role in assessing this impact. Arthropods biodiversity is linked to structures (Spies, 1998). Like all arthropod biodiversity research, these must deal with an overwhelming number of animals and the lack of comprehensive taxonomic knowledge. Targeted studies investigating the impact of land management on these animals can offer solutions to mitigate the loss of biodiversity (Bashford et al., 2001; Brown et al., 2001; Grove, 2003; Michaels & McQuillan, 1995).

Under current forest harvest regimes, older trees are removed from the Tasmanian Eucalyptus forest landscape, and replaced with young trees destined for

1) Introduction and Aims

harvesting before formation of old-growth characteristics. The tree population, and subsequently the forest environment, will change dramatically (Lindenmayer et al., 2000a). Research comparing different tree ages offers insight into the results of changing the forest tree demographics (Harrison et al., 2003; Yee et al., 2001).

Forest biodiversity is managed through structures: Management of arboreal biodiversity requires knowledge of arboreal habitats. Despite humans’ great ability to destroy natural forest structures, our ability to generate them is very limited. To properly manage forest structural attributes to conserve biodiversity, more knowledge is needed on their occurrence and formation.

A critical component of that knowledge is information on how and where structures form, and consequently how to best plan for their conservation (Ball et al., 1999; Gibbons & Lindenmayer, 1996; Hallett et al., 2001; Lindenmayer, 1995). Using natural disturbance regimes as a model for forest management can most effectively conserve these structures (Franklin et al., 2002).

Modern forest management aspires to mimic natural disturbances (Franklin et al., 2002; Hickey & Wilkinson, 1999; Lindenmayer & McCarthy, 2002). In Eucalyptus forests, the major disturbance factor is fire. In practice, the proximate goal is to protect biodiversity from the adverse effects of harvest by preserving specific, identifiable forest structures that would survive a fire. The ultimate goal is to foster similar dynamics such as regrowth and decay in the disturbed forest (Grove et al., 2002). Measures to conserve features of forest stands such as standing dead trees, large old trees, and coarse woody debris are considered part of a “new forestry” and have been adopted with varying degrees of success (Franklin, 1989). In Tasmania, the Silvicultural Systems Trial has been testing these measures (Hickey et al., 2001).

Targeted preservation of structure is the best approach: Taking a conservative approach to conserving unknown arthropod biodiversity (Bickel & Tasker, 2004) would require the preservation of all forest structures. This is incompatible with modern forestry. Research on the use and formation of habitat structures can maximize the value of the efforts being made by the industry for wildlife conservation.

Forest managers protect biodiversity by retaining trees: The most effective method of conserving forest structures is to not destroy them. Forest managers have

1) Introduction and Aims

the mandate of preserving native animal biodiversity to be factored into planning for the primary requirement of extracting forest products for economic value (Norton & May, 1994; Orr & Gerrand, 1998). One method to achieve this goal is the retention of old, living trees in clusters or singly (Neyland et al., 1999).

Legal requirements to manage forests for endangered species in Australia and the United States have prompted research into their habitat requirements. For example, large, older trees are required for the survival of animals such as the red- cockaded woodpecker (Hanula et al., 2000), marbled murrelets (USFS, 2004), and Leadbetter’s possums (Lindenmayer et al., 1990). The conservation of ancient oak trees required by saproxylic beetles had been addressed in Sweden (Ranius, 2000). Because these trees take so long to develop, preservation of forest trees in harvested areas is the basic tool of foresters in biodiversity conservation (Franklin, 1989; Lindenmayer et al., 1993). Replacement is not an option in the scale of a human lifetime.

Retained tree selection guided by knowledge of habitat use and formation: The selection of retained trees requires knowledge of which attributes contribute to animal biodiversity. When these attributes are not readily visible, assessing the likelihood of their presence is critical (Whitford, 2002). Retained trees in Eucalyptus forestry are usually old trees with visible hollows. By examining trees felled for harvest, Koch (2004) has been studying marsupial occupancy of the tree hollow habitat in E. obliqua. Using similar techniques Harrison et al. (2003) have been studying the beetle communities in these trees and their association with fungus.

Lindenmayer et al. (1993) and Gibbons & Lindenmayer (2002) discuss factors in hollow formation and their application to retaining trees in the landscape. Gibbons & Lindenmayer (2002) esti mate that 303 Australian native vertebrate species utilize hollows in live or dead trees, and that each one of these animals has its own requirements at to what makes a suitable hollow. Larger animals require larger hollows only found in larger, older trees (Mackowski, 1984). Forest birds for example, may seek out exceptionally large trees of a particular species (Hansell, 2000), and forest marsupials may seek out trees with a secondary epicormic crowns (Mackowski, 1984). However, management of tree hollows for vertebrate fauna is one element of a complex problem. Other habitats exist besides hollows, and myriad other organisms exist besides vertebrates. In addition, these vertebrate animals may only be

1) Introduction and Aims

present at a specific tree because of the availability of arthropods as food (Abbott & Van Heurck, 1985; Hanula & Franzreb, 1998; Recher et al., 1996; Scarff et al., 1998).

Knowledge of habitat utilisation aids management of habitats: Creation or fostering of animal habitat is an option when evidence exists for its effectiveness. A very familiar example is the common artificial birdhouse. Gibbons & Lindenmayer (2002) and Mackowski (1984) discuss the use of artificial nest boxes in creating marsupial habitat in the absence of tree hollows. The killing of trees can be a simple way to generate standing dead wood habitat in managed landscapes where it is in short supply (Hallett et al., 2001; Moorman et al., 1999). Killing branches within a tree crown may create similar habitat in microcosm. The same knowledge that guides tree retention can be applied to habitat creation.

Keystone structures support biodiversity: Forest structure is the template for the picture of biodiversity (Spies, 1998). Tews et al. (2004) introduce the concept of keystone structures. These structures are a resource, or combination of resources required by certain taxa to exist in a region, and the removal or lack of these resources will result in their extirpation. The trees in the forest are prominent keystone structures.

A keystone structure may be an aggregate of other keystone structures. It may be abundant or scarce. It may seem irrelevant to humans..

Different animals may require contradictory or mutually exclusive structures. A tree with many branches may be better than a tree with few branches to some animals. In Victorian Eucalyptus forests, for example, understorey Acacia trees and old decayed eucalypts have been identified as a keystone structures for marsupials (Lindenmayer, 1995).

However, location of these structures may be critically important as well. McCune et al. (2000) investigate the microhabitats of arboreal bryophytes and report that similar structures in different locations within the tree are distinct habitats. The same Acacia trees from the Victorian forests can be irrelevant to the animals if in the wrong context, such as steep slopes (Lindenmayer, 1995).

Not all trees have all keystone structures: Trees in the forest may be keystone structures in that they contain, or have the potential to form, specific structures. Trees at different life stages, and even pieces of the tree at different stages,

1) Introduction and Aims

can potentially fulfil different requirements for different animals (Mackowski, 1984; Whitford, 2002). For example, leaves are more likely to be found on a given tree than burls. Fruit is a temporary resource (Andersen & New, 1987).

Some keystone features are age linked: For example, several marsupial and bird species require hollows formed in old trees (Abbott & Whitford, 2001; Ambrose, 1981; Gibbons, 1994; Hanula et al., 2000; Haseler & Taylor, 1993; Lindenmayer et al., 1997). While some elements of a tree may be similar to those in other trees (e.g. foliage), the presence of a hollow increases the biodiversity value of the tree and the forest stand. Conversely, the loss of these features by natural or human disturbance can destroy animal biodiversity (Franklin et al., 2002).

Elements such as flowers, leaf litter, fruit, pollen, shedding bark, and fallen branches are all structures within a tree that change as the tree are grow, ages, and dies. Some structures, such as charcoal on tree trunks, are created during disturbances that may change the abundance of several other structures. The same fire that left charcoal on the tree trunks may have destroyed all of the nonvascular epiphytes. Research on the chance formation of these structures over time and their relation to disturbance regimes can aid prediction of their presence in the landscape (Lertzman et al., 1996; Lindenmayer et al., 1990; Mackowski, 1984; McCarthy & Lindenmayer, 1998; Mushinsky & Gibson, 1991; Nilsson et al., 2002; Pausas et al., 1997).

Management of structures: The perfect forest management plan would optimally conserve cryptic taxa by conserving all types of structures (Bickel & Tasker, 2004). This also means preserving the chance rhythms of disturbance that may stochastically generate these structures.

The presence of a structure does not necessarily mean it is a keystone structure. However, a conservative approach to management would assume, and plan accordingly, that all of these elements are utilized and required at some stage by some animal. Human knowledge of forest invertebrate biodiversity is very poor in regards to the number of species in the forest, let alone the ecological requirements of these unknown animals (Basset et al., 2003; Erwin, 1995; Grove & Stork, 2000; Stork et al., 1997).

Research on structures in Eucalyptus forests, such as large, old living and dead trees, and coarse woody debris, has emphasized that forest industry land managers

1) Introduction and Aims

must plan for the loss and recruitment of these features throughout the landscape over time (Grove et al., 2002; Lindenmayer et al., 1997). This requires detailed knowledge of the abundance, formation, and loss of these structures in relation to the planned forest harvest regime. Unfortunately, most studies of long term sustainability of old- growth structures in Eucalyptus forests harvested by total clearing predict that these goals are not being met (Gibbons, 1994; Gibbons & Lindenmayer, 1996; Kirkpatrick, 1994; Lindenmayer et al., 2000b). Little is known about the canopy arthropods of old Eucalyptus trees, and consequently little is known about the impact of forest management on them.

1.2 Research questions

Gaps in the knowledge:, three major research questions were identified and engaged. The overarching goals are to aid forest managers in conservation of arboreal animal biodiversity, and to fill some of the gaps in human knowledge of these ecosystems

1) What are the structural differences between 100 year old and old- growth E. obliqua, and how can they be measured and displayed?

Eight trees in each age class were mapped and compared. Spatial measurements of every branch within the tree crown were collected and summarized. The presence of structural features, such as hollows and vascular epiphytes, was noted. Computer visual models of each tree were generated to illustrate the shape of each tree.

2) How is the canopy arthropod biodiversity different in 100 year old and old-growth E. obliqua?

Three types of traps were set in the crowns of the study trees and the collected arthropods analysed for differences in abundance, richness, diversity, community structure, and distinctness.

3) In what ways does crown structure influence arthropod biodiversity in E. obliqua?

1) Introduction and Aims

The arthropod biodiversity responses to tree structural descriptors was explored. Several correlations between structural predictors and arthropod responses of composition, abundance, richness, and diversity are identified and explored. These are presented as promising avenues of future investigation.

1.3 Limitations of the present study

The effort involved in accessing the study trees limited all aspects of fieldwork. Days lost to windy conditions had to be made up later in the season. Without the assistance of several enthusiastic field helpers, studying sixteen trees would have been impossible.

The study trees were biased towards safe climbing trees. Most of the trees in the study site were deemed too dangerous for climbing.

Mapping of branches required careful consideration of resolution. It was impossible to map every leaf and branch on trees reaching 75 metres in height, so a precise method of scaling back the mapping resolution was required. However detailed any tree map may be, the structural measures of gross branch morphology that seemed important to a human climber may actually be irrelevant to some arthropods.

Like all arthropod biodiversity studies, time and resources limited trap placement and processing. For some arthropod taxa, taxonomic identification was aided by local experts (please see section 7.2). Consequently other taxa were not sorted to the same degree. Fieldwork was only done in summer, therefore missing out on seasonal differences in fauna.

E. obliqua also grows in dry sclerophyllous forests. It is present in a wide range of environments in Tasmania and mainland Australia. The wet sclerophyll forest site in the study has a strong Gondwana rainforest floristic element, and contains representatives of some of the tallest and biggest trees in Australia. The environmental conditions at the study location are significantly different from other E. obliqua localities. It may therefore not adequately represent Eucalyptus in other circumstances.

2) Aims and Background

2) Aims and Background

In this section, the study aims are defined. These aims were designed to answer the research questions and to maximize the value of the time and effort spent in the Eucalyptus trees. A review of the relevant literature is presented. Previous crown structural mapping research is cited, and current models of crown development in Eucalyptus are presented. Next, a brief review of trends in canopy arthropod research is followed by a more detailed treatment of what is known about Eucalyptus trunk and canopy arthropods. Finally, the literature on the link between tree crown structure and arthropod biodiversity is discussed. 2.1 Addressing the research questions

2.1.1 Study Design

Comparison of neighbouring old and young trees: By studying trees of different ages, a picture can be formed of their structural development through time (Van Pelt & Nadkarni, 2004). Tasmanian Eucalyptus forests are difficult to age by ring counts and high intra-cohort variability can mask differences in age (Alcorn et al., 2001). Therefore, the study site was selected where cohorts of Eucalyptus were sufficiently distinguishable by size. Comparison of two ages illuminates only two stages of the 450 year lifespan of E. obliqua (Hickey et al., 1998), but allows for control of spatial confounds (Burgman & Williams, 1995; Richardson et al., 1999).

Schowalter (1995), Schowalter & Ganio (1998), and Jukes et al. (2002) compared the canopy arthropods of different tree species simultaneously with forest age, and reported that most variation in forest arthropod communities was linked to the tree species sampled. To control the confounding effects of tree species and site, nearby old and young study trees of a single species were paired, controlling for site- specific effects (Burgman & Williams, 1995; Richardson et al., 1999).

Habitats are studied in situ in live trees: Study of the arboreal environment within a tree is best done by accessing the trees (Nadkarni, 1995). Several habitats in mature Eucalyptus trees cannot be studied after the tree has been felled. Characteristics, such as the distinctive arrangement of branches in the crown, are modified greatly as the tree lands. Animals living in the airspace of the tree are likely

2) Aims and Background to have flown away. Modern canopy access techniques allow observations and measurements of undisturbed tree crowns in situ (Moffett & Lowman, 1995).

2.1.2 Study Aims and Applicability Few researchers have climbed Eucalyptus trees for study purposes (Hickey et al., 2000; Larson, 2004; Lowman et al., 1987; Mifsud, 2003; Ohmart et al., 1983b; Palzer, 1983; Van Pelt et al., 2004). The present project was therefore exploratory in nature. The project aims were developed to address the research problems. The aims are divided into structural “mapping” aims, and arthropod “trapping” aims. Not all aims were built around hypothetical tests. Embedded in the study aims are eight hypothesized quantifiable differences between the age classes. In the presentation of this thesis, the mapping aims will be addressed first, followed by the trapping aims, and then an exploration of their correlation.

Throughout the rest of this document, the 100 year old E. obliqua are referred to as “100yr” trees, and the old-growth E. obliqua are referred to as “old trees.”

The ten study aims are:

1. Mapping Aim 1: Quantitatively assess the differences in crown structure and size between 100yr and old E. obliqua

2. Mapping Aim 2: Investigate the presence of structural features in the crowns of 100yr and old E. obliqua

3. Mapping Aim 3: Modify the conifer mapping technique of Van Pelt et al. (2004b) for Eucalyptus trees

4. Mapping Aim 4: Develop a technique for displaying crown structure of forest trees using computer models based on 3-dimensional spherical coordinates

5. Mapping Aim 5: Expand Jacob’s (1955) theories of intra-branch competition in Eucalyptus saplings to mature and old-growth E. obliqua

6. Mapping Aim 6: Generate a predictor data set to explore the influence of crown structure on canopy arthropods

2) Aims and Background

7. Trapping Aim 1: Determine what differences exist in arthropod biodiversity between 100 year old and old growth Eucalyptus obliqua

8. Trapping Aim 2: Contribute to the knowledge of Eucalyptus anopy arthropods

9. Trapping Aim 3: Develop robust, inexpensive trap designs suitable for transport to and use in E. obliqua

10. Trapping Aim 4: Generate a response data set to explore the influence of crown structure on canopy arthropods

These aims are described in greater detail:

Mapping Aim 1: Quantitatively assess the differences in crown structure and size between 100yr and old E. obliqua: Identifying age-related changes in crown structure can aid in understanding the growth of these trees, and aid in assessing what forest structures are lost with harvested old-growth trees.

• Hypothesis 1) Old E. obliqua are quantifiably and objectively distinct from 100yr trees, i.e. old trees are not scaled-up versions of 100yr trees. • Hypothesis 2) Old E. obliqua have a more variable crown structure than 100yr E. obliqua.

Mapping Aim 2: Investigate the presence of structural features in the crowns of 100yr and old E. obliqua. The opportunity to observe and record these features in situ allows for discussion of their formation and location. A photographic record can aid future researchers in selecting arboreal habitats to investigate.

• Hypothesis 3) Structural features such as epiphytes, hollows, dead tops, snapped trunks and burls are more commonly present in old E. obliqua than 100yr E. obliqua.

Mapping Aim 3: Modify the conifer mapping technique of Van Pelt et al. (2004b) for Eucalyptus trees. The mapping methods used in the present project are derived from techniques of Van Pelt et al. (2004b). Their methods were developed in coniferous trees. Eucalyptus has a distinct growth pattern which required

2) Aims and Background

modifications to their technique. Researchers wishing to map Eucalyptus or other trees in a similar fashion can utilize these adopted changes if appropriate.

Mapping Aim 4: Develop a technique for displaying crown structure of forest trees using computer models based on 3-dimensional spherical coordinates. The rendering of the study trees using 3-dimensional computer models allows for illustration of crown dynamics and offers a faithful visual representation of the collected data.

• Hypothesis 4) Old E. obliqua will have a greater complexity than 100yr trees, as measured in surrogate by the amount of information required to describe them.

Mapping Aim 5: Expand Jacob’s (1955) theories of intra-branch competition in Eucalyptus saplings to mature and old-growth E. obliqua. The crown structure of E. obliqua is a product of competitive interactions between individual branch units (Halle, 1995). Jacobs (1955) introduced the concept for saplings, but did not extend it to older trees. Applying the principles of intra-branch competition to mature and old-growth E. obliqua can clarify differences in crown structural measurements.

Mapping Aim 6: Generate a predictor data set to explore the influence of crown structure on canopy arthropods. When combined with measures of arthropod biodiversity, structural attributes of Eucalyptus trees can be identified that are important in determining the abundance, richness, diversity, and composition of canopy arthropods.

Trapping Aim 1: Determine what differences exist in arthropod biodiversity between 100 year old and old growth Eucalyptus obliqua. The loss of old, senescent Eucalyptus trees and the spread of regrowth forests has been identified as a threat to conservation of marsupial biodiversity (Lindenmayer et al., 1999). It is unknown if arthropod communities are similarly at risk. While the literature suggests that old-growth trees have a more diverse arthropod fauna than younger trees (Martikainen et al., 2000; Schowalter, 1995), this has not been addressed in Eucalyptus trees.

2) Aims and Background

• Hypothesis 5) Old-growth E. obliqua trees have a more abundant arthropod fauna than 100 year old trees, i.e more individual arthropod animals. • Hypothesis 6) Old-growth E. obliqua trees have a richer arthropod fauna than 100 year old trees, i.e. more recognizable types of arthropods. • Hypothesis 7) Old-growth E. obliqua trees have a more diverse arthropod fauna than 100 year old trees, i.e. a greater quantitative index implying both a richer community and more even abundances between morphospecies. • Hypothesis 8) Old-growth E. obliqua trees have a distinct arthropod fauna from 100 year old trees, i.e. the taxonomic composition of animals differs.

Trapping Aim 2: Contribute to the knowledge of Eucalyptus canopy arthropods. There is a gap in the knowledge of the arthropod biodiversity in wet sclerophyllous Eucalyptus trees (Majer et al., 1997). Because Eucalyptus is the dominant forest tree in Australia (Lindenmayer et al., 1997), and because arthropods are a critical part of the global ecosystem (Kim, 1993), it is important to study their presence and diversity. Listing, photographing, and curation of collected specimens will add to the resources available for arthropod biodiversity research.

Trapping Aim 3: Develop robust, inexpensive trap designs suitable for transport to and use in E. obliqua. Because of the effort involved in accessing the crowns of tall forest trees, passive traps allow greater flexibility in placement schedules than active collecting methods. Traps are needed that are durable and inexpensive. Recycled bottles and compact disk cases were found to fulfil both of these requirements.

Trapping Aim 4: Generate a response data set to explore the influence of crown structure on canopy arthropods. In tandem with Mapping Aim 6, the results from the arthropod trapping can be paired with the structural data. The combination of detailed structural measurements and the arthropod biodiversity results allows for exploration into their correlation. It is assumed that crown structure predicts arthropod biodiversity, rather than the other way around (Spies, 1998; Tews et al., 2004).

2) Aims and Background

Arthropods responses of abundance, richness, diversity, and composition can be related to structural variables for each tree.

2.2 Background Research

2.2.1 Crown Structure and development

Vertical structure: The structural qualities of individual tree crowns and forest canopies is often referred to as vertical structure, whereas the landscape pattern is referred to as habitat heterogeneity (Brokaw & Lent, 1999; Tews et al., 2004; Tscharntke & Brandl, 2004). In the present study, only vertical structure is addressed. Vertical structure includes all of the aerial portions of the forest (Brokaw & Lent, 1999). The vertical structure of a forest location can be a determining factor in the climate and light on the forest floor (Holbrook & Lund, 1995; Nunez, 1985; Parker, 1995; Van Pelt & Franklin, 2000). The forest industry is based on utilizing the trunk component of vertical structure.

Tree crowns as opposed to forest canopies: The terms “crown” and “canopy” have different meanings and are frequently confused (Moffett, 2000). An individual tree has a crown, while forests have a canopy composed of combined crowns (Moffett, 2000; Parker, 1995). The distinction can blur (Nadkarni et al., 2004). Conifers with reiterated trunks can be viewed as an aerial collection of smaller crowns forming a canopy (Sillett & Van Pelt, 2000). In savannah ecosystems, or Eucalyptus woodlands, tree crowns form “islands in the sky” that usually do not overlap (Dean et al., 1999; Dial et al., 2004; Hnatiuk et al., 2003; Kruger & McGavin, 1997). Herbs, shrubs and grasslands may be considered canopies by smaller animals (Nadkarni et al., 2004; Parmenter & MacMahon, 1984). Furthermore, the subject of interest may be the gaps, or airspace between tree crowns (Dial et al., 2004; Van Pelt & Franklin, 2000). In the context of the present study, the unit of interest is the individual tree crown. In the wet sclerophyll forests, mature Eucalyptus crowns form an emergent island over the closed canopy of the rainforest below.

Quantification of plant structure: Two major approaches to quantification of plant physical structure exist. Descriptive empirical mapping of a crown structure,

2) Aims and Background used in this study, is distinct from modelling hypothetical tree architecture (Godin, 2000).

Halle (1995) defined the architectural unit as an individual plant’s unique expression of an architectural model dictated by its genotype. Events in the life of a forest tree can dictate the exact form of the architectural unit, such as:

1. decay (Ball et al., 1999; Manion, 1981; May & Simpson, 1997; Palzer, 1983; Parks & Shaw, 1996; Wardlaw & Neilsen, 1999; White & Kile, 1994; Whitford, 2002),

2. insect herbivory (Landsberg, 1990; Lowman et al., 1987; Lowman & Heatwole, 1987; Mopper et al., 1991),

3. competition from other plants(Aiba & Kohyama, 1997; Anten & Hirose, 2001; Dial et al., 2004), and

4. fire (Franklin et al., 2002; Jacobs, 1955).

Even the loss of one bud at a branching point can change the future shape of the plant (Hadlington & Johnston, 1988; Halle, 1995). For example, the foliage scorching of E. obliqua saplings can completely kill the dominant treetop and create an entirely new shape, with several smaller branches vying for apical leadership (Wilkinson & Jennins, 1993). This familiar “candelabra” shaped pattern of reiterated branches is one of the more obvious outcomes of individual tree life experiences, and can be witnessed in ancient forest trees such as Sequoia sempervirens (Sillett & Van Pelt, 2000) and Thuja plicata (Van Pelt, 2002). The stochastic event responsible for the pattern, and the tree’s growth responses to it, are the focus of modelling, whereas those mapping the tree focus on the observed results of these factors.

Although the actual distinction between these two approaches is flexible, the scientific literature suggests that studies of plant architecture using the modelling approach appear in genotypic, predictive and mechanistic studies (Attiwill, 1962; Ball et al., 1999; Coder, 2000; Godin et al., 2004; Hanan & Room, 1997; Maguire et al., 1998; Parker, 1995). By contrast, descriptive mapping lends itself towards phenotypic, experimental and biogeographic analysis (Clement & Shaw, 1999; Ishii et al., 2004; Sillett, 1999; Sillett & Rambo, 2000; Van Pelt & Nadkarni, 2004).

2) Aims and Background

Previous plant mapping research: The subjects of descriptive plant mapping studies can be divided into four types:

1. Features of plants 2. Individual plants 3. Similar plants 4. Different plants Examples of these include:

1) Features of plants: The subject of interest is the physical location of a component within the plant. Mackowski (1984) dissected two E. pilularis and reported on their position and formation. Whitford (2002) concentrated on tree hollows in West Australian E. marginata and Corymbia calophylla. Peeters (2002) examined leaf structural features and their impact on herbivore communities.

2) Individual plants: No comparisons are made within the framework of a study design. When trees are studied individually, they are usually of exceptional size or age. The popular literature has several qualitative portraits of individual trees (Lewington & Parker, 1999; Mortimer & Mortimer, 2003; Pakenham, 1996), but these are rarely linked to any quantifiable measurements beyond height, girth, and crown spread. The crown of a Sequoia sempervirens tree with a complex network of secondary trunks was described and illustrated by Sillett & Van Pelt (2000). The crown of a giantSequoiadendron giganteum was described by Sillett et al. (2000a). The largest and tallest representatives of Western North American species have been measured and illustrated by Van Pelt (2002).

3) Similar plants: Several similar plants are used as representatives of a larger group. Ishii & Wilson (2001) measured the branching architecture of six old Pseudotsuga menziesii and report on the processes of branch death and epicormic resprouting in these trees. Takenaka et al. (1998) used a simple angular and distance measuring devices to quantify and illustrate several Japanese forest shrubs, and to quantify the light capture of each plant. Clement & Shaw (1999) quantified the crown shape and branching characteristics of P. menziesii and its influence on epiphyte biomass. They discuss the importance of limb size and vertical height. Sillett (1999) measured the crown shape of old-growth Sequoia sempervirens and detailed

2) Aims and Background epiphyte habitats at a branch scale. Sillett & Bailey (2003) compared crown-level structural variables with the biomass of epiphytic fern in Picea sitchensis.

4) Different plants: The structure of plants representing multiple groups is compared. They are not necessarily from different species. The present study is a comparison of different ages of trees. Schmid & Bazzaz (1994) compared the leaf mortality and height growth of the perrenial flowering plants Aster lanceolatus and Solidago canadensis throughout a single growing season. Van Pelt & Nadkarni (2004) and Ishii & McDowell (2001) measured the branching architecture of P. menziesii of different ages to investigate trends in crown development. Clement et al. (2001) studied the crown structure and foliage arthropods of the Chilean conifer Fitzroya cupressoides. They compared trees ~500 years old with those ~2000 years old, and discussed the differences in crown structure, epiphytes, and arthropod communities.

2.2.2 Growth and developmental of Eucalyptus

Fifty years ago, M.R. Jacobs summarized a career’s worth of knowledge of Eucalyptus trees in a single book. The Growth Habits of the Eucalypts (1955) begins with a discussion on the dynamics of buds, and moves up the spatial scale to leaves, branches, crowns, and silvicultural stands. The concepts are illustrated in clear and appealing line drawings. Jacobs’ (1955) interest extended primarily to silviculturally useful specimens, in contrast to the modern scientific interest in the growth and structure of old-growth trees (Ishii & Wilson, 2001; Van Pelt & Nadkarni, 2004). Decadent, gnarled old trees that were considered “useless veterans” (Jacobs 1955, figure 137) are now considered to be an invaluable and effectively irreplaceable resource for wildlife (Lindenmayer et al., 1997; Mackowski, 1984). In the context of the current study, Jacobs’ (1955) treatments of Eucalyptus crown and branch development are utilized.

Eucalyptus crowns: (Jacobs, 1955) distinguished four crown stages, or “types”, in Eucalyptus (Figure 2.1) These are:

A. Vigorous young trees, which develop into B. Early mature trees, which develop into either C. Trees with fire-induced epicormic crowns or

2) Aims and Background

D. Mature trees of high quality

All 100 year old trees in the present research possessed early mature crowns. Old trees had either an epicormic crown or a mature, high quality crown. If a crown fire had burnt through the crown, most original branches were lost and epicormic branches sprouted. If a fire had not burnt the crown, original branches were lost slowly to crown senescence, and epicormic branches sprouted to replace them.

Figure 2.1: Crown types in Eucalyptus. Clockwise from upper left, A, B, D, C. Figure 58 in Jacobs (1955)

2) Aims and Background

Eucalyptus branch dynamics: Jacobs (1955) illustrated a progression of ages downwards through four years of branches in the crown of a Eucalyptus sapling (Figure 2.2). The youngest branches are uppermost in the crown. These branches are held upright, and descend to a horizontal attitude over time. The lowest branches, older than four years, are moribund and will be dropped as the tree ages. Competition between branches was discussed only in regards to the terminal branch. This model was not extended to old-growth Eucalyptus.

Curtin (1970) addressed the crown shape and branch allometry of even-aged E. obliqua between 10 and 93 years old. This range of ages may be less than one quarter of the lifespan of E. obliqua at Warra LTER (Hickey et al., 1998). He discussed the “non-plasticity” of branch sizes. It is, however, unclear if the dynamics as illustrated by Curtin (1970) and Jacobs (1955) hold true for Eucalyptus older than 100 years of age.

2) Aims and Background

Figure 2.2: Intra- crown competition of branch cohorts in E. sieberana sapling crown. The youngest branches are the most vertical, the oldest live branches are more horizontal, and the oldest live branches are lowest in the crown. Figure 55b from Jacobs (1955)

The competition of branches within the crown of young and old-growth forest trees is addressed by Ishii & McDowell (2001). Following Halle’s (1995) description of branches as competing individuals, they treated branches as microcosms of whole trees (Figure 2.3). They present a model of Pseudotsuga menziesii development in which branch competition is compared to stand dynamics (Franklin et al., 2002). Lower original branches are older branches. Branch cohorts of decreasing altitude and increasing age show similar dynamics to the development of forest stands (Halle, 1995). The young upper crown of old trees resembles the young upper crown of younger trees, but the older lower crown shows its age in lower

2) Aims and Background

branch density and higher numbers of dead branches (Ishii & Wilson, 2001). Ishii and McDowell (2001) likened the death of branches in the lower crown of P. menziesii to the mortality of trees in a forest stand, and the sprouting of epicormic buds to the recruitment of young trees.

The concepts introduced by Jacobs (1955) for Eucalyptus saplings, by Halle (1995) for rainforest trees, and by Ishii & McDowell (2001) for P. menziesii development are applied in the present study to aid in understanding the crown structure of 100yr old and old-growth E. obliqua.

Figure 2.3: Conceptual model of crown development comparing branch cohorts to stand development. From Ishii & McDowell (2001)

Empirical crown structural mapping of Eucalyptus crowns: Current research on Eucalyptus crowns has generally moved away from Jacobs’ (1955) focus on tree ontogeny. The forest industry continually refines its models of forest stand growth, but published literature on Eucalyptus crown structure at a tree scale is less common. Recent studies have addressed light dynamics, disease responses, and the measurement of exceptional specimens of the genus.

Kelly et al. (2004) quantified light levels and leaf numbers in a sapling E. regnans. They applied computer technology to assign 3-dimensional coordinates to every branching point on the young trees.

2) Aims and Background

Rather than measure the position and size of branches, another approach to quantification of structure is the mass of tree elements. Lowman et al. (1987a) measured the aerial and subterranean biomass of two E. nova-anglica trees, one of which was suffering from rural Eucalyptus dieback syndrome. They quantify the loss of root biomass and increase in wood-borer damage in the dieback tree.

Published literature showing the crown structure of individual trees is rare for Eucalyptus. Exceptionally tall Eucalyptus trees have measured in one dimension in efforts to find the tallest living specimen (Hickey et al., 2000; Kostoglou, 2000; Mifsud, 2003). Van Pelt et al. (2004b) detailed a method of quantifying forest structure at branch-scale and stand-scale, and compare tree crown attributes of old- growth E. regnans and P. menziesii. They link a hand-drawn profile illustration of an old-growth wet sclerophyll Eucalyptus forest stand to 3-dimensional computer profiles of a forest tree. Mackowski (1984) presents a detailed hand-drawn rendering of two E. pilularis trees to depict the location of hollows.

2.2.3 Canopy Arthropods

The study of treetop arthropods: Sampling canopy arthropods from trees is inherently difficult (Basset et al., 2003c). Animals must be either brought down into reach, or the researcher must be brought up to them (Mitchell, 1982; Nadkarni, 1995). Despite these difficulties, canopy arthropods are the subject of a growing interest (Basset et al., 2003b; Stork et al., 1997).

Current concentrations in canopy arthropod research include:

• The total number of species in the world forest canopy, e.g. (Erwin, 2004)

• the host specificity of insects in trees, e.g. (Kitching et al., 2003; Odegaard, 2000),

• the function of arthropods as herbivores, e.g. (Basset, 2001; Landsberg & Cork, 1997; Lowman, 1995; Lowman & Rinker, 2004; Rinker et al., 2001; Wotherspoon, 1998)

• the effects of forest management, e.g. (Chey et al., 1998; Chung et al., 2000; Floren & Linsenmair, 2001, 2003)

2) Aims and Background

• vertical stratification, e.g. (Basset et al., 2003a; Le Corff & Marquis, 1999; Shaw, 2004)

New techniques have been developed in the last thirty years to collect arthropods living in regions out of reach. Most important among these are chemical knockdown (Basset et al., 1997; Erwin, 1995; Majer & Recher, 1988; Stork & Hammond, 1997), canopy cranes (Basset et al., 2003c; Davis, 2001; Odegaard, 2000), and rope access techniques (Dial & Tobin, 1994; Moffett & Lowman, 1995; Perry, 1978). While a thriving field of study, the difficulties in access and the vast numbers of animals present a difficult challenge. It is apparent that much remains to be learned (Basset et al., 2003b, c; Stork et al., 1997).

Chemical knockdown, or canopy fogging by insecticide mist, is considered to be less biased in that it collects several animal taxa, but the ability to link animals to the habitat they were collected from is limited (Majer et al., 1990; Richardson et al., 1999; Stork et al., 2001). As an active collecting technique, chemical knockdown vulnerable to temporal and climatic (such as wind) conditions that can render it useless (Majer et al., 1996).

Canopy cranes are used in several research sites around the world. While offering safe and comfortable access to all parts of the outer tree crown, cranes are prohibitively expensive to build, operate, and maintain, and are fixed in a single location (Basset et al., 2003c; Moffett & Lowman, 1995; Wind River Canopy Crane Research Facility, 2002).

Rope techniques offer the most inexpensive and portable method of direct access to the canopy. The initial investment of equipment and training allows study of canopy arthropods in their habitat by direct observation, active collecting or passive trapping methods. Drawbacks to the methods include risks associated with working at heights, and difficulties in movement away from sturdy branches (Nadkarni, 1995; Smith & Padgett, 1996). However, to sample and measure canopy biodiversity directly, rope techniques allow exact placement of traps and the ability to observe habitats in situ.

2.2.4 Comparing ages of trees

It is an ecological truism that old, complex forest trees sustain an animal biodiversity both richer and distinct than that of younger, simpler trees. In Australian

2) Aims and Background

Eucalyptus forests, and other forests of the world, several studies support this claim for vertebrates animals (Gibbons, 1994; Jackson, 2000; Lindenmayer et al., 2000; Lindenmayer et al., 1991; Mackowski, 1984; McCarthy & Lindenmayer, 1998; Pausas et al., 1997; Spies, 1998). While it is perhaps a logical next step to conclude the same is true for invertebrates, less evidence is available.

Studies comparing the arthropod biodiversity of different forest ages (Floren & Linsenmair, 2001; Schowalter, 1995) are not designed to compare different ages of individual trees of the same species. Studying different sites may confound results even when comparing trees of the same species.

Published evidence of age-related arthropod differences: Differences in arthropod biodiversity associated with plant age have been identified in several plant types around the world. These investigations have generally found a richer fauna in older plants, forests, or structural elements. It is sometimes unclear if this is due to differences in size- older plants are usually larger than younger plants. A confounding element in comparing results is the measure of scale, i.e. did the researcher choose to standardize arthropod results by some measure of scale. This confound is discussed in detail in section 7.6.

Within a single plant, Waltz & Whitham (1997) studied juvenile ramets vegetatively propagating clonally from the base of mature adult Populus trees. The fauna of the adult was both different and richer than that of the juvenile foliage. Between plants, Banerjee (1981) found more herbivore species in older individuals of plantation tea (Camellia sinensis).

Working from a canopy crane, Basset (2001) reported a greater abundance and richness of phytophages collected per unit of leaf area in mature specimens of Panamanian tree Pourouma bicolor than in saplings. Differences in herbivore composition were attributed to the greater resource of young foliage, microclimate differences, increased toughness of leaves in mature trees, and the presence of enemy- free space. Ants were far more abundant on saplings, suggesting a different trophic web in the crowns of P. bicolor at different ages. At the same site, Barrios (2003) compared the herbivores on mature and sapling Castilla elastica and found a greater richness and diversity in the mature specimens.

2) Aims and Background

Thunes et al. (2003) found a greater richness of arthropods collected by canopy fogging in older Norwegian Pinus, but not when quantitatively controlling for their greater volume.

Clement et al. (2001) studied the foliage of Fitzroya cupressoides (Cupressaceae), the giant alerce of Chile, and found more psocids and aphids, but fewer cercopids and staphylinids in the larger, older trees ( >2,000 years old) than the smaller, younger (~500 years old) trees.

Schowalter (1995) studied age effects on Pseudotsuga menziesii forests in Oregon, USA, and found a higher diversity and abundance of predators and detritivores in older forests. In Finland, Martikainen et al. (2000) found a richer beetle fauna in old-growth Picea forests of than in younger regrowth forests.

Applicability to biodiversity conservation: The management implications of these results to conserving biodiversity are related to the changing demographics of forested landscapes. Pinus, Picea and Pseudotsuga are important forest resources that are harvested today. Like Eucalyptus forests, harvested old trees in their native environment are not replaceable within the time scale of a human life.

In Tasmania, other concurrent projects at Warra LTER (described below) are comparing the biodiversity associated with 100 year old E. obliqua with that of large old growth trees. Harrison et al. (2004) is investigating the invertebrate and fungal biodiversity of interior habitats in E. obliqua by dissecting felled trees, and preliminary results show a distinct and richer beetle fauna in old trees. Yee et al. (2001) have found a distinct fauna between small and large naturally fallen logs. These projects, as well as the canopy fogging by Grove et al. (2002) were all performed within a 10km radius at Warra LTER (Bashford et al., 2001).

2.2.5 Eucalyptus Canopy arthropods

Knowledge of Eucalyptus canopy arthropod biodiversity: When the co- dominance between Acacia and Eucalyptus of the entire Australian continent is considered, very little is known about the arthropod biodiversity associated with their crowns (Majer et al., 1997; Williams & Brooker, 1997). A chart outlining the

2) Aims and Background scientific literature addressing the canopy, foliage, and trunk arthropod biodiversity of adult Eucalyptus in Australia is presented as Table 2.1.

Most research on Eucalyptus canopy arthropods has addressed three broadly overlapping themes:

1) herbivore and foliage arthropods (Abbott et al., 1992; Abbott & Wills, 2001; Elliot et al., 2002; Fensham, 1994; Fox & Morrow, 1983; Landsberg & Cork, 1997; Lowman et al., 1987),

2) comparisons between species or subgenera (Morrow, 1977a, b; Ohmart et al., 1983a, b; Woinarski & Cullen, 1984; Wotherspoon, 1998), or

3) biodiversity assessments of the Eucalyptus forest canopy in Western Australia, New South Wales, and Tasmania (Elliot et al., 2002; Grove et al., 2002a; Majer et al., 1994; Majer et al., 2000; Recher et al., 1996b; Recher et al., 1993).

Some of the significant findings of these studies are quantitative evidence that:

1) arthropod biodiversity in Eucalyptus forest canopy may be richer than in other temperate forest ecosystems (Majer et al., 1994),

2) the arthropods associated with Eucalyptus differ throughout the seasons (Bell, 1985; Lowman & Heatwole, 1992; Recher et al., 1996b),

3) herbivory is an important factor in the ecosystem (Landsberg & Cork, 1997; Ohmart, 1984; Ohmart & Edwards, 1991),

4) arthropods respond to drought (Bell, 1985),

5) some arthropods respond to fire (Radho-Toly et al., 2001; Steinbauer et al., 1998),

6) arthropod and bird communites are interlinked (Abbott & Heurck, 1985b; Evelegh et al., 2001; Recher et al., 1996a),

2) Aims and Background

7) there are differences in communities associated with different subgenera (Burdon & Chilvers, 1974a; Morrow, 1977a; Ohmart et al., 1983a; Wotherspoon, 1998), and

8) communities vary with altitude (Burdon & Chilvers, 1974b) , geographic region (Burgman & Williams, 1995; Majer et al., 1990; Richardson et al., 1999) and within the tree (Majer et al., 1990).

The present research fits into the third theme of biodiversity assessment. Very little research has been done on the biodiversity of Eucalyptus without limiting the scope to phytophagous of foliage insects.

There are three outstanding gaps in the knowledge: Very little is known about old-growth trees (Bashford et al., 2001; Majer et al., 1997), the effect of tree age on the arthropod community (Abbott & Whitford, 2001), or the habitats utilized by arthropods in the crowns of mature trees. These three subjects are critically important to forest management in Australia. The clearfelling and replanting of native forests is reducing the number of very old tree with young trees (Lindenmayer et al., 1990). Planned forestry rotations will not allow sufficient time for the regeneration of this resource (Lindenmayer, 1995). It is unknown if old-growth Eucalyptus are as important as reservoirs for arthropod biodiversity as they are for vertebrates (Gibbons & Lindenmayer, 2002). Consequently, little is known about the impacts of forest harvesting on Eucalyptus canopy arthropods.

Research on Eucalyptus trunk and canopy arthropods: In addition to the foliage, the surface and airspace within the tree crown are inhabited by arthropods. The invertebrate biodiversity associated with the trunks of live, standing Eucalyptus trees has been studied on Western Australia (hereafter “WA”) by Majer et al. (2003) using flight intercept and funnel traps, by Bickel & Tasker (2004) in NSW using sticky traps, by Scarff et al. (1998) in WA by hand-collecting, and by Harrison et al. (2004) on recently fallen E. obliqua using emergence traps on pieces of bark. Larson (2004) climbed E. grandis in subtropical rainforest and trapped flying insects with flight intercept and sticky traps.

2) Aims and Background

Table 2.1 (following pages) Citations in the scientific literature on native Eucalyptus trunk and canopy biodiversity

2) Aims and Background

2) Aims and Background

2) Aims and Background

Table 2.1 (previous pages) Citations in the scientific literature on native Eucalyptus trunk and canopy biodiversity

Trunk and canopy biodiversity research in Eucalyptus has generally focused on the “active” collecting rather than “passive” trapping. Only two published trunk studies have used trapping techniques (Bickel, 2003; Majer et al., 2003). Bagging, beating, or sweep-netting foliage has been used to compare differences in the invertebrate fauna between tree species (Abbott et al., 1992; Abbott & Wills, 2001; Evelegh et al., 2001; Majer et al., 1996; Majer & Recher, 1988; Morrow, 1977a; Ohmart et al., 1983b; Yen, 1989) and to test ecological effects such as fire, predation or drought (Bell, 1985; Evelegh et al., 2001; Radho-Toly et al., 2001; Yen, 1989). Wotherspoon (1998) reported on the phytophagous foliage insects of E. obliqua in comparison to other Eucalyptus species.

Canopy fogging, or insecticide knockdown, has been performed on Eucalyptus in W.A. and New South Wales by Recher et al. (1996b) and Majer et al. (1990). In Tasmania, canopy fogging collections have been taken from mature specimens of E. obliqua within 5 km of the trees studied by the present research (Bashford et al., 2001; Grove et al., 2002a). Yen & Lillywhite (1990) performed a preliminary fogging project in Tasmanian rainforest plants, which are often found growing near or with Eucalyptus (Kirkpatrick & Backhouse, 1981).

The foliage and trunk biodiversity of the southwestern WA dry sclerophyll trees, E. marginata and Corymbia callophylla have been researched by several different projects and techniques, mostly in comparison to different Eucalyptus species (Abbott et al., 2000; Majer et al., 1990; Majer et al., 2003; Radho-Toly et al., 2001). No comparable body of research exists for Eucalyptus in the rest of the continent.

In sclerophyllous forests throughout the world, fire regularly disturbs the ecological community (Pielou, 1979). Radho-Toly et al. (2001) investigated the responses to fire of arthropods on native and exotic Eucalyptus in W.A. They linked a higher abundance on burnt trees to higher nutrient levels in the leaves. Steinbauer et al. (1998) examined the preference of a Hemipteran pest to fire coppiced Eucalyptus trunks over intact stems.

2) Aims and Background

2.2.6 Previous research on the link between structure and arthropod biodiversity

Branches in crown are analogous to trees in forest: Halle (1995) considers branches of a tree as independent pieces sprouting from similar pieces. Ishii and McDowell (2001) analysed the branch distribution within a Pseudotsuga crown as corresponding to trees in a forest stand. In a forest management setting, older, mixed- age forests containing dead trees are considered to be more desirable than younger single-cohort plantations for biodiversity conservation (Seymour & Hunter, 1999). A parallel situation may exist within the crown of individual trees, with those trees containing dead branches fostering more biodiversity than a simpler tree (Paviour- Smith & Elbourn, 1993). Standing dead trees (McComb & Lindenmayer, 1999) and dead branches (Schiegg, 2001) may be comparable, similar habitats.

This analogy can be taken further: shed branches may be like falling trees, branches in poor health like dying trees, old healthy branches like older trees, and epicormic resprouts like seedlings. Sillett &Van Pelt (2000b) found the outstanding number of reiterated secondary trunks in a Sequoia crown to be best described as a “forest canopy” of its own. The relationships between forest structures and forest biodiversity may therefore apply at the finer scale of the tree crown.

Surrogate measures of complexity: Habitat complexity is often presented as a predictor of biodiversity (Tews et al., 2004), but is often inadequately defined (Loehle, 2004). Quantification of the crown structure and the presence of distinct structural attributes can be surrogate measures of geometric, physical complexity (Loehle, 2004). (Anand & Orloci, 1996) comment that complex things require more effort to describe than simple things. In the context of the current structure, the amount of information need to describe different trees is quantifiably measured (in surrogate) by the computer file sizes of 3-d virtual illustrations.

The abundance or presence of a certain feature, or attribute, in a landscape may be considered to add complexity to it. For example, the abundance of trees, and the presence of volcanos, islands, mountains, and valleys on one island might make it more appealing to the geologist and the biologist than an island with none of these features. Similarly, the crown of a tree with abundant branches, epiphytes, litter collection, and shards of broken wood may be considered to be more complex than a

2) Aims and Background

tree with few branches and none of these features. For the purposes of this study, complexity is measured in surrogate by characteristics of branch architecture, and the presence of structural features.

Linking structure to biodiversity: Lawton (1983) highlights five key components of plant architecture that determine insect diversity: size, growth form, seasonal development, variety of above-ground elements, and persistence of individual components. In this study, the size, growth form, and variety of elements is quantified.

In the framework of the current study, the effects of age and crown structure may be indistinguishable. Like all organisms, Eucalyptus follows a distinct ontogeny (Jacobs, 1955). Different structural attributes may develop together as the tree ages. Research on their influence may be confounded at a tree level by their interaction.

Lawton (1983) presents two approaches to explaining the higher diversity detected in larger, more complex plant types. First, more complex plants have more areas to be colonized by insects. Second, more complex plants offer larger numbers and types of spatial situations and resources. The former can be addressed by looking at tree size and shape through its crown structure, and the latter can be addressed by looking at the presence or absence of different features.

Lawton’s (1983) approach 1 – Complex trees offer more area for arthropods: Strong et al. (1984) present a comparison of broad growth forms and show a descending trend in phytophagous insect diversity from trees through shrubs, perennial herbs, annual herbs, and monocotyledons. They describe these growth forms as a descending trend of architectural complexity.

A further refinement of the concept could be used to compare different forms of the same species, or even within the same trees. Fowler (1985) investigated the phytophage fauna on Betula seedlings, shrubs, and trees. His study found little significant difference in composition and richness between the three, except for a winter sampling period in which seedlings had the poorest fauna. He postulates that the proximity of the three size classes to each other could foster recruitment of the fauna from the trees to the smaller classes, and that a lack of overwintering sites on seedlings could be responsible for difference in winter.

2) Aims and Background

Experiments on plant structure and its effect on spider communities has shown that structural complexity supports spider abundance (Rypstra et al., 1999). Sundberg & Gunnarsson (1994) and Gunnarsson (1990) investigated the relationship of spider abundance to foliage needle structure Picea branches. Densely needled branches contained more spiders, and experimental removal of needles resulted in a decrease in numbers.

Similarly, Halaj et al. (2000) experimentally modified the foliage habitats of Pseudotsuga by removing needles, removing branches, and tying branches together. They found several differences in predator and prey relationships, and conclude that the habitat structure within the tree may influence the animal food web in a “bottom- up,” fashion, in which changes in prey affect the predators.

Structures at a scale finer than the leaf may influence arthropod composition. Peeters (2002) examines the presence of specific leaf attributes and their relationship with the function feeding group community. Plants with leaf defences such as spines or thickened cuticles had lower levels of herbivores.

Research in conifers of Western North America has identified several relationships between crown structure and epiphyte biodiversity. McCune et al. (2000) examine the lichen biodiversity in several arboreal microhabitats. Clement & Shaw (1999) and Sillett (1999) report on the effect of branch-level descriptor variables such as branch size and crown depth on arboreal resources such as epiphytes and soil collections. Arthropod communities utilize these resources, and subsequently are affected by the tree structure (Nadkarni & Longino, 1990; Prinzing, 1997).

Larger trees have more habitat area: A larger tree may represent a larger “island” to arthropods (MacArthur & Wilson, 1967). Southwood & Kennedy (1983) compare the geographic range of trees to the size of an island. The title under which they publish their theory is “Trees as islands.” An alternative analogy views individual trees as islands. An appropriate title for this view would be “A tree as an island."

A basic principle of island biogeography is that larger islands support more species (MacArthur & Wilson, 1967). In the present study, the old trees were significantly larger that the 100yr trees in several measures. Height, crown volume, total foliage, surface area, and wood volume were significantly greater in the old

2) Aims and Background

trees. If these measures are analogous to the 2-dimensional land area of an island, then richness should therefore be greater in the old trees. Other possible measures of size include the abundance of fruits (Andersen & New, 1987) or volume of dead wood (Schiegg, 2001). The choice of a measure of scale is discussed in detail in the canopy arthropods methods section.

The relationship between the size of an island and the number of species of a taxa is presented by MacArthur &Wilson (1967) as

# of species = C * AreaZ

where

C is a constant which varies between taxa,

Z is a constant which varies with habitat heterogeneity from the theoretical expected value of 0.27 for a population with a lognormal distribution, and

Area is a two-dimensional measure of habitat size.

Other measures of size could substitute for island land area. For trees, it may be crown volume, foliage mass or surface area. As discussed above, none of these measures are satisfactory.

An important observation is that usually Z < 1. If Z = 1, then species richness would scale linearly with size. In the context of trees, a tree twice as big as another (with no differences other than size) would therefore have twice the number of species within in. The definition of “twice as big” is problematic and dependent on the organisms of interest. An appropriate measure of scale is difficult to choose (see section 7.6). Abundance might (but not necessarily) scale linearly with the available quantity of the appropriate habitat, but because Z < 1, species richness increases more slowly than area.

Lawton’s (1983) approach 2 – Complex trees have a variety of spatial situations and resources: Carey (1996) writes that, for arboreal mammals, “all attributes of canopies could have biological importance.” The same statement applies to arboreal arthropods.

The invertebrate biodiversity associated with several different arboreal habitats has been investigated in several studies. Majer et al. (1997) review the literature on habitats within Eucalyptus forests. When an animal community is found

2) Aims and Background to be associated with a certain feature, it can be assumed that the loss of that feature would affect that community. These include, but are not limited to:

1. foliage and live branches ((Abbott et al., 2000; Basset, 2001; Clement et al., 2001; Lowman et al., 1993; Peeters, 2002; Schowalter, 1995; Winchester, 1997; Woinarski & Cullen, 1984; Wotherspoon, 1998) 2. dead wood (Hanula & Franzreb, 1998; Paviour-Smith & Elbourn, 1993; Ranius, 2000; Ranius & Hedin, 2000; Schiegg, 2001) 3. hollows in the trunk and branches (Harrison et al., 2004; Ranius & Wilander, 2000) 4. flowers, fruit and cones (Abbott & Heurck, 1985a; Andersen & New, 1987; Jaffe et al., 2003; Kirmse et al., 2003; Majer et al., 1997; Roubik et al., 2003; Shea, 1989; Turgeon et al., 1994) 5. bark and branch surfaces (Bickel & Tasker, 2004; Buchs, 1990; Hanula & Franzreb, 1998; Heterick et al., 2001; Majer et al., 2003; Moeed & Meads, 1983; Nicolai, 1986; Nicolai, 1995; Proctor et al., 2002) 6. fungal hyphae and fruiting bodies, which may actually owe its presence in the tree to a saproxylic animal (Harrison et al., 2004; Komonen 2001; Sippola, 2001) 7. individual leaf structural features (Lill & Marquis, 2004; Peeters, 2002; Shaw & Walter, 2003; Walter & O'Dowd, 1995) 8. suspended soils (Nadkarni & Longino, 1990; Paoletti et al., 1991) 9. vascular epiphytes (Benzing, 1983; Coxson & Nadkarni, 1995) 10. non-vascular epiphytes (Prinzing & Wirtz, 1997; Rhoades, 1995) 11. vines (Odegaard, 2000) 12. charcoal on tree trunks (McQuillan, personal communication) 13. water-filled tree hollows, or phytotolmata ((Kitching & Callaghan, 1982)) 14. leaves bound by caterpillar-silk (Lill & Marquis, 2004) Other features with the potential to foster animal biodiversity include sap flows, cambium wounds, cracked wood, suspended bark ribbons, branch fusions, and rubbing points between branches.

Bickel & Tasker (2004) argue for consideration of unknown fauna in land management. The presence of structural attributes in a tree suggests the potential to

2) Aims and Background

harbour more animal diversity than their absence would. Cryptic ecological associations, chance probability, and the time scale involved in the formation of these structural attributes are factors to be considered in forest management.

Age of elements: The age of elements within a tree may alter its role for animals in the arboreal ecosystem. Furthermore, structures within a tree may grow, shrink, or stay the same size as the tree ages. Branches sprout, grow and die in a microcosm of the entire tree (Attiwill, 1962; Jacobs, 1955; Lowman, 1992). Dead branches in a tree, like dead trees on the forest floor, change as they age. As logs and coarse woody debris age, they are utilized by different animal communities (Grove et al., 2002b; Irmler et al., 1996; Yee et al., 2001).

Young leaves in a tree are chemically and physically distinct from old leaves (Landsberg, 1990). Studies by Lowman (1985) and Peeters (2002) found higher abundances of herbivores on softer, greener young foliage than harder, lignified mature foliage. Waltz & Whitham (1997) compared resprout suckers and mature foliage within Populus trees, and found significantly more insect species on mature foliage. They relate their findings to a larger scale view, and comment on the influence of developmental heterogeneity in single trees may contribute to the biodiversity in a stand of trees. Within a tree crown, the presence of both younger and older foliage together may foster animal diversity.

Tree trunks in older Eucalyptus trees are often decayed and harbour fungal hyphae, whereas younger tree trunks may not (Barry et al., 2000; Harrison et al., 2004; Wardlaw & Neilsen, 1999). Fungal biodiversity is interlinked with animal biodiversity (Komonen 2001).

Hollows develop and grow over time, and are suitable for different animals at different stages (Gibbons & Lindenmayer, 2002; Mackowski, 1984). Fungal hyphae can develop fruiting bodies (May & Simpson, 1997). Water filled tree-hollows can dry out or overflow (Kitching & Callaghan, 1982). Buds can flower, fruit, and disperse (Andersen & New, 1987). The tree can be burnt (Radho-Toly et al., 2001). Jukes et al. (2002) found an increase in saproxylic beetles in older British Pinus plantations. Pinus may carry more dead wood in its crown as it ages, and similarly, older branches in trees may contain more dead branchlets.

2) Aims and Background

2.3 Synthesis of previous research

Crown structural mapping addresses the empirical shape of individual trees.

Mapping measures the actual expression of the trees innate architectural model

(Section 2.2.1). Eucalyptus crown structure has been studied from various aspects.

Jacobs (1955) addressed stages of crown development and senescence; a theoretical

branch competition framework utilising his stages and the analogy of branches to

trees presented by Ishii & McDowell (2001) is presented.

Empirical mapping of Eucalyptus trees has been done at the level of biomass

(Lowman et al. 1987a), leaf position (Kelly et al. 2004), tree height (Mifsud, 2003),

hollow locations (Mackowski 1984), and the branch structural level (Van Pelt et al.

2004b).

Canopy arthropods have been studied by several methods, but few studies

have compared the arthropod fauna associated with different aged forests or trees.

Eucalyptus canopy arthropods have been studied in respects to herbivores, faunal

composition differences between species, and biodiversity assessments comparing

eastern and Western Australia.

The link between structure and arthropod biodiversity is discussed in reference

to the analogy of branches in a tree as trees in a forest. Structural complexity, tree

size, habitat presence, and the age of tree elements are all proposed as factors

determining arthropod composition and biodiversity in trees.

3) Study Design & Site

3) Study Design and Site

The primary objectives of the research were to compare the differences in crown structure and canopy arthropod biodiversity of 100 year old and old growth wet sclerophyll Eucalyptus obliqua trees. The study site and design were selected to most effectively approach this objective.

The Warra Long Term Ecological Research Site (hereafter “LTER”) was chosen to utilize and integrate with the body of previous scientific research performed there (Alcorn et al., 2001; Allen, 2001; Beaulieu, 2004; Brown et al., 2001; Coops, 2001; Hickey et al., 2001). In a multi-aged forest, old-growth trees and 100 year old trees were selected in nearby pairs. The age classes were coexisting and readily distinguishable by basal girth (Alcorn et al., 2001). A confounding intermediate age class was not present. Selecting in pairs controlled for any potential environmental differences in fauna within the study area (Kirkpatrick, 2004; Richardson et al., 1999; Schowalter, 1995). 3.1 Study Subject and Site

3.1.1 Tasmanian wet sclerophyll forests:

The wet sclerophyll forests of Tasmania are largely dominated by E. obliqua, E. regnans, E. delegatensis, and E. globulus (Forest_Resources, 1979). These are the tallest and most massive angiospermous forests on Earth (Hickey et al., 2000; Kostoglou, 2000; Mifsud, 2003; Van Pelt et al., 2004), and are second in height only to the coniferous forests of western North America (Van Pelt, 2002). Age cohorts of Eucalyptus originate from fire events (Ashton, 2000; Gill, 1997; Jacobs, 1955). Fires of insufficient intensity to cause complete mortality generate a cohort of Eucalypts seedlings (Lindenmayer et al., 2000). Successive fires will therefore create multiple aged forests. In the absence of regular fires, high fuel loads in the wet sclerophyll forests can enable uncontrollable conflagrations, a fact well noted by residents of forested areas of Hobart (Gill, 1997; Marsden-Smedley & Slijepcevic, 2001).

In the absence of fire, temperate rainforest vegetation develops beneath the Eucalyptus canopy (Gilbert, 1959). The rainforest is floristically linked to the ancient supercontinent of Gondwanaland (White, 1986), and supports an herbivorous

3) Study Design & Site

arthropod fauna distinct from that of the more recent Eucalyptus trees (McQuillan, 1993; Pielou, 1979; Yaxley, 2000).

Clearfelling, burning, and seeding of Eucalyptus forests is the most widely applied silvicultural method, although other methods are being trialled (Hickey et al., 2001). The management of these forests for economic gain is the source of heated, ongoing controversy and is considered a matter of national significance (Beale, 2003; ForestryTasmania, 2003; Green, 2003; Lindenmayer et al., 1990).

3.1.2 Eucalyptus obliqua

E. obliqua is the type species for the Eucalyptus genus. The holotype was collected by Captain Cook’s 1777 expedition to Bruny Island, 50 km east of the study site, and described by Charles-Louis L’Heritier de Brutelle in 1788 (Costermans, 1981).

E. obliqua is one of the tallest and largest trees flowering plant species in the world, after the closely related and co-existing E. regnans, and is the most widespread eucalypt in Tasmania (Figure 3.1) (Kirkpatrick & Backhouse, 1981; Williams & Potts, 1996). E. obliqua is known as “brown-top stringybark” in Tasmania, and as “messmate” in Victoria (Forest_Resources, 1979). It is more flexible in its environmental requirements than E. regnans, and will dominate it on less fertile sites (Ashton, 2000; Kirkpatrick, 2004) .

It is a non-lignotuberous member of the Monocalyptus “ash” subgenus (Costermans, 1981). Characteristic flakes of flammable dead phloem cells cover the trunk and larger branches (Jacobs, 1955).

Figure 3.1: Distribution of E. obliqua in Tasmania, from Kirkpatrick & Backhouse (1981)

3) Study Design & Site

Figure 3.2 : Location of Warra LTER in southern Tasmania. Hobart is located beneath the “E” in “LTER”. Graphic from www. warra.com 3.1.3 Warra Long Term Ecological Research Site

Warra LTER is a 16,000 hectare site (Figure 3.2) straddling the border of the Southwest World Heritage Area 60 km southwest of Hobart (Brown et al., 2001). It is primarily covered by virgin and managed E. obliqua forests, with additional areas of temperate rainforests and buttongrass moorland (Corbett & Balmer, 2001). The south and east boundaries follow the Huon and Weld rivers. At the confluence of the Huon and Picton rivers in the southeast corner, the Tahune Airwalk ( offers tourists the opportunity to view E. obliqua trees from a canopy walkway (Figures 3.3, 3.4). Forestry Tasmania maintains a web site with information, metadata, and species lists from the LTER at http://www.warra.com.

Figure 3.3(left): The Tahune Airwalk offers a canopy-level view of E. obliqua similar to the study trees. Several illustrative photographs were taken from this platform. Figure 3.4 (right): Robert. Junker and Elaine Vale on the Tahune Airwalk.

3) Study Design & Site

3.1.4 West end of Manuka Road

Manuka Road runs upstream on the north banks of the Huon River and ends 10km west of the Airwalk (Figure 3.5). This site is the only road-accessible region of contiguous virgin forest in Warra. Several other studies have been conducted along Manuka Road, including investigations of: a) sustainable silvicultural methods (Hickey et al., 2001; Neyland et al., 1999) b) the association of beetles and fungus in E. obliqua (Harrison, 2004; Harrison et al., 2004),c) chemical knockdown of arboreal arthropods (Bashford et al., 2001; Grove et al., 2002), and d) the arthropod biodiversity of fallen logs (Grove & Bashford, 2003; Yee et al., 2001).

Southwest World Heritage Area Approx. Boundary e t i B S

y d u t S C

Ma nuk a Rd A

Hu on R iv er

A Canopy fogging tree True Tahune North Airwalk B Log decay study 10 km C 1996 harvest regen 500m

Figure 3.5: Aerial photograph of the west end of Manuka Road, Warra LTER, Tasmania. Aerial photograph supplied by Forestry Tasmania. Note the large crowns of old-growth trees in the northern section, and the finer grain of the younger trees in the southern section.

3) Study Design & Site

3.1.5 Topography and Geology

The trees used in the study were located within a 2 km2 region immediately west of the road’s western extent, on the southern slopes of Mt. Frederick Plateau. Study trees were between 100 and 300 m altitude. The Huon River flows eastward 1 km south at ~75 m altitude. The area is well drained, steep, and southern facing (250 m altitude gained in 1 km). All creeks drain southwards to the Huon River. All creeks were flowing throughout the year. The geology is composed of igneous Jurassic (~170 million years old) dolerite overlaying the Parmeener supergroup of Permian mudstone sediments (~300 million years) (Laffan, 2001). Pieces of mudstone with fossils of brachiopods and bryozoan Fenestella were visible where forest soil had been disturbed by landslide or uprooted trees (White, 1986). A small limestone outflow cave was found 1km west of the study site.

3.1.6 Vegetation

The uppermost forest stratum consists of very tall wet sclerophyllous E. obliqua, with E. regnans present in the northern section (Figures 3.7 and 3.8). Eucalyptus tree crowns do not overlap, forming an open canopy from 30-80 m in height.

Figure 3.7 (left): 100yr E. obliqua cohort as seen from Manuka road (MW) Figure 3.8 (right): Old-growth E. regnans present in the northern section of the study site, with R. Junker descending.

Beneath the Eucalyptus trees, a middle stratum of rainforest trees 10-30 m tall forms a closed canopy (Figures 3.9 and 3.10). Nothofagus cunninghamii, Atherosperma moschatum and Pomaderris apetala were the most abundant trees, followed by Acacia melanoxylon, Eucryphia lucida, Phyllocladus aspleniifolius, and Acacia dealbata. Immediately beneath this stratum were Dicksonia antarctica treeferns 1-4 m tall. Patches of the study site were pure rainforest consisting of these trees without Eucalyptus. Fallen logs of large Eucalyptus and other smaller trees were abundant on the forest floor (Figure 3.11).

3) Study Design & Site

Figure 3.9 (left): Acacia melanoxylon was often encountered growing immediately below E. obliqua. M. Whitman shown examining epiphytic bryophytes

Figure 3.10 (right): The dead tops of old E. obliqua tower above the pointed crowns of Atherosperma moschatum. (MW)

Understorey vegetation included Tasmannia lanceolata, Monotoca glauca, Acacia verticillata, Anodopetalum biglandulosum, Olearia argophylla, Cenarrhenes nitida, Anopterus glandulosus, Pimelea drupacea, Gahnia grandis, Urtica incisa, Polystichum proliferum, Hymenophyllum spp. and numerous bryophytes (Figures 3.12, 3.13, 3.14) (Collier et al., 1992; Corbett & Balmer, 2001; Costermans, 1981; Forest_Resources, 1979; Jarman & Kantvilas, 2001a, b; Kirkpatrick, 2004; Kirkpatrick & Backhouse, 1981).

Figure 3.11 (left): Burnt Eucalyptus stumps are frequently encountered as vertical shards. (MW) Figure 3.12 (right): 100yr E. obliqua trunk and Dicksonia antarctica

3) Study Design & Site

Figure 3.13 (left): Nothofagus cunninghamii (left edge) and Phyllocladus aspleniifolius (right edge, dark foliage) near the base of tree 4 Figure 3.14 (right): Dicksonia antarctica near old tree 12, with Robert Junker on a fallen log

3.3 Study design

3.2.1 Age comparison

Trees originating from fires around 1898-1906 (“100yr”) were compared to veteran older neighbouring trees that survived that fire event (“old”.)

E. obliqua establishes after fire events (Kirkpatrick et al., 1988). In Tasmania, the repetition of small-scale burning heterogeneity and the survivorship of individual trees creates multi-aged forests with cohorts originating from each fire event (Alcorn et al., 2001). Older eucalypts were invariable rotten through the centre, masking dendrochronology through ring counts (Allen, 2001; Hickey et al., 1998). No seedlings of Eucalyptus were ever seen in the forest. Nothofagus seedlings were common. In the absence of fire, the rainforest vegetation was regenerating and the sclerophyll vegetation was abiding for the next burn.

The age of the E. obliqua trees was inferred from the regional fire history compiled by Hickey et al. (1998). Old trees were inferred to range between 300 and 450 years of age, originating from a fire before recorded history (Figure 3.15). The 100yr trees germinated after fires in the period of 1898 to 1906 (Hickey et al., 1998). The natural fire rotation has been determined to be 448 years long (Hickey et al., 1998).

3) Study Design & Site

Alcorn et al. (2001) warns of inferring wet sclerophyll E. obliqua age by diameter, but at this site size differences between the two age cohorts were sufficient for ready identification. Unlike other regions at Warra, the west end of Manuka road does not appear to have burnt in 1936 or at any other time since 1898-1906.

This survivorship of old trees contrasts with certain areas of E. regnans forest in Victoria, in which a single intense fire, followed by a long exclusion of fire, causes total mortality of a stand. This leads to even-aged stands of Eucalyptus, such as seen at Wallaby Creek, Victoria (Figure 3.16) (Ashton & Chinner, 1999; Mifsud, 2003).

Figure 3.15 (left): Matt Cracknell standing next to an old tree showing charcoal at its base. This indicates that the tree had survived a fire. Most old trees were observed with these markings Figure 3.16 (right): Even-aged stand of E. regnans at Wallaby Creek, Victoria. Note the intra-cohort variation of stem diameters.

3.2.2 Tree selection methods

Eight pairs of E. obliqua trees were studied from August 2003 to April 2004 (Figures 3.17, 3.18, 3.19, 3.21). Each pair consisted of a 100yr old tree and an old tree. The two trees were always less than 50 m from each other, and all eight pairs were within a 2 km2 region. Climber safety and tree health were the primary selection criteria. Because of the greater abundance of 100yr trees and the decayed nature of old E. obliqua (Jacobs, 1955), location of appropriate old trees guided selection of 100yr trees. By definition, the old trees had survived or avoided the fire events in the years 1898-1906 that gave rise to the 100yr cohort (Alcorn et al., 2001; Allen, 2001;

3) Study Design & Site

Gill, 1997). The fire records compiled by Alcorn et al. (2001) found no evidence of fire in the location since then.

Figure 3.17 (left): The crown of 100yr tree 1 Figure 3.18 (right): The crown of old tree 8 (MW)

3.2.3 Study tree description

The old trees in the study were commercially overmature veterans (Figure 3.20) (Jacobs, 1955). They were no longer alive at the top and had lost most of their original branches, but still retained a number of original branches in addition to more recent epicormic growth. Many of the living branches in the old trees were not the original branches, but rather were mature epicormic shoots arising from the trunk or the basal portions of original branches (Figures 3.24, 3.25). These successful resprouts, especially when lower on the tree trunk, probably grew immediately after a fire that the tree survived. The old trees were very likely to have a hollow rotten centre (Gibbons & Lindenmayer, 2002). The branches on old trees were often decayed or hollow.

The 100yr trees possessed their original crowns with a live terminal shoot. They were vigorously growing upwards, but were often shaded by the old trees. The 100yr trees were always taller than the understorey trees (Acacia melanoxylon, Nothofagus cunninghamii, Pomaderris apetala, Atherosperma moschatum) standing nearby. Dominant trees possess an abundance of sturdy branches suitable for access.

3) Study Design & Site

Subdominant trees generally had no suitable branches for climbing, and exhibited more kinks and irregularities in their trunk. All trees used in the study represented the dominant, least decayed members of their cohort.

Figure 3.19: The study pair of 100yr tree 7 and old tree 8. Note the difference in crown shapes. Tree 7 is rounded and vigorous, whereas tree 8 is snag-topped and in decline. Mt. Picton is in the background across the Huon River, in the World Heritage Area. (MW) .

3) Study Design & Site

Figure 3.20 (left): Closer view of dead top of old tree 8 with an epicormic crown beneath. The “V” shape of the funnel drift fence is just visible on the trunk at the bottom of the photograph (MW) Figure 3.21 (right): looking downwards from tree 1, both age cohorts are visible. An old tree is in the upper right corner, and the rest of the stems are 100yr trees. Note cave at base of old tree (R. Junker)

Judging by the lack of stringy bark in the upper crown of some study trees (especially old trees 3 and 4), it is possible that these are hybrids with E. regnans (Figures 3.22, 3.23) (Kirkpatrick & Backhouse, 1981; Williams & Potts, 1996).

Figure 3.22 (left): The upper trunk of old tree 4 is smooth barked. This may indicate hybridization with E. regnans. Sue Baker is shown descending. Figure 3.23 (right): The smooth bark on the upper trunk of old tree 3 may indicate hybridization with E. regnans. Brendan Kayes is shown recording data .

3) Study Design & Site

Figure 3.24 (left): Old E. obliqua with a secondary epicormic crown (MW) Figure 3.25 (right): Old E. obliqua with snapped trunk, and a single large epicormic branch

3.2.4 Study tree identification

Each study tree was assigned a number (Table 3.1). With two exceptions, study pairs were made of consecutively numbered trees. The 100yr trees were given an odd number and the old trees given the even number immediately following. The first four trees were named in a pilot study and contradict this numbering scheme. Trees 1 and 2 are 100yr trees, and 3 and 4 are old trees. Renumbering these trees for clarity’s sake was deemed to be too much of a risk to data integrity, and therefore the contradictory numbers were retained.

Study Pair ABCDEFGH 100yr Tree Number 1 2 5 7 9 11 13 15 Old Tree Number 3 4 6 8 10121416

Table 3.1: Tree identification numbers for each study pair

3.2.5 Rope techniques

Access to the trees was gained by launching a crossbow bolt trailed by fishing line over a sturdy branch. (Figures 3.26, 3.27, 3.28) A cord 2.5 mm in diameter was tied to the fishing line and the line reeled in (Mitchell, 1982). A climbing rope was then installed by tying it to the rigging cord installing a rope for climbing entry.

Caver-style Single Rope Techniques (SRT) were used to climb to the branch (Figures 3.29, 3.30). In SRT, a climber’s weight is alternated between a pair of ratcheting ascenders or knots to move up the rope (Perry, 1978; Smith & Padgett, 1996; Warild, 2004)

3) Study Design & Site

Figure 3.26 (left): Phillip Grosse using a compound bow for access to old tree 14 Figure 3.27 (right): Gene Miller using crossbow for access to old tree 4, the rightmost stem.

Figure 3.28 (left): Lucie Whitten using crossbow to access 100yr tree 11. Figure 3.29 (right): Matt Cracknell ascending into tree 12 using SRT rope technique

Once in the crown, arborist doubled rope techniques were used to move upwards from the access branch until the highest safe point was reached (Figures 3.31, 3.32). In arborist rope technique, a rope is thrown over a branch, anchored to the climbers harness, and the standing end controlled by a ratcheting friction hitch knot

3) Study Design & Site

(Figure 3.32)(Budworth, 1999; Dial & Tobin, 1994; Moffett & Lowman, 1995; Smith & Padgett, 1996).

The time required to access and rig each tree varied widely. For example, tree 1 was shot, climbed, and rigged in ~5 hours, whereas tree 14 took ~6 days of full-time work. Some time was wasted rigging trees that were considered, but not actually studied. Not all trees had easily targeted branches or slopes nearby to offer higher crossbow shots. Availability of field assistance and the proper equipment controlled whether rigging trees would be successful. Once in the tree, the time to reach the highest point varied with crown structure (branch anchor abundance), weather, field assistance, and myriad other concerns.

Figure 3.30 (left): Melissa Whitman ascending into old tree 8 using SRT rope technique Figure 3.31 (right) Jerry Romanski secured with arborist rope techniques in old tree 4.

3) Study Design & Site

Figure 3.32 (left): R Junker utilizing arborist rope techniques in E. regnans ~250 years old. Figure 3.33 (right): Arborist doubled ropes: a schwabish friction knot and a barrel termination knot

4) Crown Structure – Methods

4) Crown Structure – Methods

In this section, the methods used in mapping the crown structure of the study trees are presented. A definition of elements is established first. The concept of “a branch” is discussed in detail. The concept of branch resolution and generation is discussed. A framework for measuring the decomposition, geometry, and topology of a tree using stacked, relative, spherical pocometric coordinates is presented. The information actually collected in the field is listed, and the process of generating and analysing tree structural descriptor variables from this information is described. The layers used in generating a 3-dimensional computer arborograph illustration are shown, followed by a novel method of measuring structural complexity based on the amount of information required to display the tree map. Finally, the structural features recorded in the field are introduced. 4.1 Definition of tree elements

Working definitions of several terms are presented to assist in understanding the data. Moffett (2000) organizes several definitions of concepts in canopy biology. He specifically highlights the divergent perceptions of what is “a branch” to remind authors to “clearly specify they are establishing a novel usage.” To that end, detailed definitions are presented for the tree trunk, and the branches.

Trunk definition: The identification of the trunk and branches of a given tree may seem obvious, but their measurement in studies of structure requires careful definition. The trunk is the main vertical stem of a tree, usually descended from successive terminal buds and dominant over other vertical stems in the tree (Moffett, 2000). In wet sclerophyll tall Eucalyptus forests, the distinction of a trunk is much more apparent than in multiple stemmed trees such as the dry mallee eucalypts. While not always vertical (Ishii & Higashi, 1998; Jacobs, 1937), the trunk is usually more vertical than any of its branches (Figures 4.1, 4.2). Leaning trees were not considered safe for access and therefore tree selection was biased towards completely upright trees.

4) Crown Structure – Methods

Figure 4.1 (left): M. Cracknell at a steeply tilted old E. obliqua Figure 4.2 (right): Old E. obliqua leaning over stream. Some of the branches are more vertical than the trunk.

The trunk may split into several contender branches vying for crown dominance (Figures 4.3, 4.4, 4.5, 4.6). The trunk may be considered to end where no single definitive leader can be distinguished, or where the trunk has broken or lost dominance, and the definitive treetop leader is a branch that arises laterally and grows upward (Hadlington & Johnston, 1988; Sillett & Van Pelt, 2000).

Figure 4.3 (left): E. obliqua ~100 years old visible from the Airwalk showing contenders Figure 4.4 (right): Old E. obliqua with several resprout contenders from two large original contenders, visible from tree 16.

4) Crown Structure – Methods

Figure 4.5 (left): Several failing contenders in the largest known E. obliqua, “Gothmog” Styx Valley Tasmania Figure 4.6 (right): Several failing contenders in Thuja plicata “Quinault Giant,” the largest known non- Sequoia tree, Olympic Peninsula, USA. Note similarity with previous figure.

Branches and resolution levels: A central concept in the measurement of any “branch” is the definition of the resolution level (Godin, 2000). Only two truly fixed resolution levels exist: the entire plant and the individual buds. At the most basic level, an entire tree is viewed a single unit with no branching (Coder, 2000; Van Pelt & North, 1996). At the most advanced scale, every single leaf and bud on a plant is addressed (Figure 4.7) (Hanan & Room, 1997; Kelly et al., 2004).

Halle (1995) identifies an intermediate level of resolution, the reiterated architectural unit. These are repeated structures that are a microcosm of the entire plant. In Eucalyptus, these are the “crown units” (Jacobs, 1955) or “foliar units” (Van Pelt et al., 2004) (Figures 4.8, 4.9, 4.10).

4) Crown Structure – Methods

Figure 4 .7: Every leaf of an E. astringens sapling digitally fixed in space, demonstrating maximum resolution at a bud level. Image generated and supplied by Natalie Kelly, University of Melbourne- Creswick Forestry School. .

4) Crown Structure – Methods

Figures 4.8, 4.9, 4.10: Physically one of the smallest eucalypts, the Western Australian E. sepulcralis (top, left) shows a superficially similar structure to the vigorous reiterated foliage units growing from parent branches the much larger E. obliqua crown (right).

Inevitably, one must compromise between the two extremes of resolution. Practical considerations of time, effort, and organization will force a researcher to summarize the finest branches. This decision delineates the resolution level. In the context of this study, “branches” are above the resolution level, and “branchlets” are below the resolution level. In analogy, a table of contents does not list every word, or every sentence, in a book. Listing every sentence would be impractical. There are three distinct approaches to setting a resolution level (Figure 4.11):

4) Crown Structure – Methods

1) Setting resolution by size: branches greater than a certain diameter or length are measured, as used in Van Pelt et al. (2004b); or choosing the larger branch at each branching node,

2) Setting resolution by branch characteristic: foliar units, branches extending from the trunk or a vertically reiterated trunk, as used in Clement & Shaw (1999) and Ishii &Wilson (2001).

3) Setting resolution by branching generation: lateral branches sprouting from primary branches sprouting from the trunk are second generation branches (Attiwill, 1962; Godin, 2000); or in the other direction, where leaf buds are the base unit and parent branches are counted inwards to the root or trunk (Hanan & Room, 1997). The observed branching generation is not necessarily equivalent to the actual bud generation. Bud mortality and epicormic branching can create a current structure that deviates from the original growth model. If identification of Halle’s (1995) reiterations is possible with precision, these units can define the resolution level.

Figure 4.11: Generation, resolution, and terminal concepts illustrated. Branching generation is counted from the base outwards instead of from the terminations inward. Note that not all branching points are bifurcations. Also note that branches of the same generation can be of different lengths. While not illustrated, terminal branches may be of different generations.

4) Crown Structure – Methods

A branch originates from a single parent branch or trunk location, and may have multiple children branches growing from it. These children branches are sister branches to each other. The exponential growth in the number of branches as resolution level becomes finer precludes field measurements of every one for even small trees (with Kelly et al. (2004) a notable exception). Branchlets are the unmeasured branches below the level of resolution. A practical working definition of a “branch” summarizes and encompasses all of the children branchlets from a given parent location.

Terminal branch measurements have no children branch measurements. Moffett (2000) defines a terminal branch as the smallest shoots on a tree. In the context of the present study, terminal refers to the smallest measured branches. Segment branch measurements are associated with further measurements of its children branches. These children branches may be segments with children themselves, or terminal.

Branching generation does not imply a fixed periodicity (Romanski, 2004). The physical length or time between branching points may differ between generations. The assigned generations may not be equivalent to the actual bud generation. In the present study, branching generation is based on observed wood structure.

When referring to branching patterns, an analogy with human ancestral lineage is used for descriptive purposes. The terminal branches are the current generation- you and your sisters. The branchlets below the resolution line are your unborn children (born by parthenogenesis). The branch you sprout from is your parent, and its parent is your grandparent. The branching path that leads to you is your lineage. The base of the entire tree is the ancestor of your whole family.

To study E. obliqua, a large tree with multi-generational branching pattern, branch characteristics and branch size were used simultaneously to generate two resolution levels. The measured branching system was based on the observed wood structure, rather than the actual bud and vascular developmental structure.

All branches originating from the trunk (a characteristic) were measured as terminal branches, defining the first, coarser resolution levels. These were the simple branches.

4) Crown Structure – Methods

Subsequent branchings below this level were measured again, down to an attempted size of ~15 cm starting diameter (a size). This was the second, finer resolution level. These were the complex branches. Branchlets smaller than ~15 cm resolution were summarized into terminal measurements, and those larger were measured in more detail by segments and their associated children measurements. However, practical difficulties of access and time can generate variability in this resolution. The base of a complex branch is the segment connected to the trunk.

Care in analysis must be taken when comparing parent branches with children branches, as they are not independent sampling units. Even comparing branches within the same tree can be problematic, because they are linked entities within the same system. Looking at branches as individual units (Halle, 1995; Ishii et al., 2000) is a valid approach, but one slightly less stable than looking at them as elements placed along a continuum from the base of the tree to the smallest branchlets, or a continuum along the trunk the crown (Ishii et al., 2002; Ishii & McDowell, 2001).

In coniferous trees, where reiterated stems grow upwards from limbs, and branches grow horizontally from stems, distinguishing branches from secondary trunks is simple. The upright stems allow for a fixed resolution using a characteristic rule (Van Pelt & Nadkarni, 2004). In wet sclerophyll Eucalyptus, the distinction between “limbs” and “branches” as used in Van Pelt et al. (2004b) is problematic due to the absence of true reiterations (Kelly, 2004). They define a limb as leading to a reiteration, which is “architecturally indistinguishable from freestanding trees.” The reiterated foliar crown unit in Eucalyptus is actually below the resolution level of their data, as well as that of the present research (Jacobs, 1955).

A further bias in resolution comes from the selection of the highest climbable branch, estimated in an exceedingly conservative manner by combining instinct, safety considerations, and climber caution.

The absolute treetop summit is intrinsically unreachable by rope techniques, a situation exacerbated by the dead tops of the old trees. At the highest safe climbing anchor, the crown above was measured as best as possible, and the uppermost branches summarized in the same manner as a whole branch.

Furthermore, anyone observing tree branching patterns should remember that the fields of crown structure and canopy physical shape are only examining the aerial

4) Crown Structure – Methods portions of plants (Figure 4.12). An equally complex, understudied branching system exists underground in the form of roots (Lowman et al., 1987). Roots may respond similarly to factors such as herbivory and resource availability (Mopper et al., 1991).When looking up in the forest, one should remember the description of the forest canopy as “roots to the sky” (Alvarez, 2002).

Figure 4.12: Looking downwards in the crown of old E. regnans in Wallaby Creek, Victoria

Original and Epicormic Branches: Connected to the trunk and generally directed laterally are original and resprouted epicormic branch systems. (Ishii & Ford, 2001; Jacobs, 1955)

Original branches begin as lateral buds at the growing tip of the tree. Consequently, as the tree ages, original branches can be traced centre of the tree and have grown radially from and proportionally with the trunk.

By contrast, epicormic branches sprout from hormonally suppressed buds beneath the bark of the trunk or branches (Figure 4.13) (Jacobs, 1955; Romanski, 2004). These buds are released in response to fire, herbivory or branch decline. Epicormic branches are connected to the wood of the tree only to the depth corresponding to the living cambium at the time of sprouting, and frequently grow on

4) Crown Structure – Methods a tangent from its surface. Eucalyptus trees defoliated by fire can grow an epicormic secondary crown (Figures 4.14, 4.17) (Jacobs, 1955).

Mature epicormics may be difficult to distinguish from original branches (Jacobs, 1955). Original branches are radially connected, and proportional in size to, their parent branch or trunk, and often arise opposite to another original branch, while epicormic branches are often tangentially connected to and significantly smaller than their parent, and may appear at irregular locations. The distinctive upward elbowing shape of Eucalyptus (Figures 4.15, 4.16) is caused by an original branch snapping off and epicormics shooting from the upper surface taking over as the dominant branch tip (Jacobs, 1955; Mackowski, 1984).

Figure 4.13 (left): Two original branches in old tree 4, with Damien Catchpole descending. Knocking on these branches produced a hollow noise.. Figure 4.14 (right): Epicormic branches on old tree 8 are visible in the upper centre. The author is just barely visible beneath those branches. (MW)

4) Crown Structure – Methods

Figure 4.15: Elbow in original branch crown of 100yr tree 13.The original leader branch pointing horizontally has been lost and the branch pointing upwards has become the new leader.

Figure 4.16 (left): Upwards elbowing exhibited in large broken originals on the left side of old tree 12, with R. Junker visible. Figure 4.17 (right): E. regnans in the Styx Valley, Tasmania, exhibiting a secondary epicormic crown

4) Crown Structure – Methods

4.2 Structural data collection

Godin (2000) distinguishes three types of information used in representing plant architecture. The field collection techniques developed to fulfil these three types for E. obliqua are inspired by and directly descended from participation in the conifer mappings published in Van Pelt et al. (2004b). These three information categories are:

1) Decomposition describes the pieces composing the representation. In this study, all pieces, or elements, are line segments, with extra information to “flesh out” the shape added. When combined with circumferences, lines form truncated cones used for describing branch segments, and complete cones used for terminal branches. The trunk of the tree is quantified as a sequential, vertical arrangement of truncated cones.

2) Geometry places these elements within the universe. Vectors in 3- dimensional space are used in this study, defining the end point of a line in relation to a starting point with angular measurements of slope and azimuth, and linear measurements of distance.

3) Topology is the connection between these elements, or the branching logic. Topology as a mathematical science has its roots in the familiar concept of plants branching, and a hierarchical branching tree form is seen in the taxonomic branches of life (Adelson, 2001). The name of an element traces its lineage back through its parent vectors (father, grandfather, great-grandfather....) to the origin point.

Decomposition & Geometry: Defined by Vectors: All field measurements were built around vectors in a 3-dimensional spherical space system, from the base of the branch to the end of the branch (Figures 4.18, 4.19, 4.20) (Takenaka et al., 1998). In spherical space, measurements of slope angle, azimuth angle, and linear distance define an end point in space from a given origin point. By contrast, in cylindrical space, vector end points are defined by linear height, angular azimuth, and linear distance. In Cartesian, or XYZ space, three linear measurements are used: change in X axis (east-west), change in Y axis (north-south), and change in Z axis (up-down).

4) Crown Structure – Methods

The selection of a space system is most important in determining field techniques. The fixing of a point in space can be mathematically converted between the three formats.

Figure 4.18:Iillustration of vectors as seen from profile view of 100yr tree 2. (Mike Sumner)

4) Crown Structure – Methods

Figure 4.19: The author as vector detector in old tree 4. (Sue Baker)

Figure 4.20: Old E. obliqua visible from Airwalk, with overlay showing a vector depiction.

A spherical coordinate system was selected for its ease of application to Eucalyptus trees, in which the trunk is the only reliably vertical element. Studies in coniferous trees of Western North America have generally used cylindrical

4) Crown Structure – Methods coordinate systems, due to the use of laser rangefinders with inbuilt trigonometric functions, and the strong distinction between horizontal branches and vertical (reiterated, secondary trunks within these tree crowns (Clement et al., 2001; Clement & Shaw, 1999; Ishii & Wilson, 2001; Sillett et al., 2000; Van Pelt, 2002).

The branch vector starting point is the point at which the centre of the branch connects to the surface of the trunk, as opposed to the centre of the tree (Van Pelt et al., 2004). The branch vector end point is defined as the farthest point of the branch from the base, generally aligned with the branch shaft. Branches with strong curves in any direction were therefore modelled less accurately than neutral branches. When multiple points fit this definition, a midpoint measurement was used.

Slope was measured to the nearest 5 degrees upwards (+) or downwards (-) from the horizon using a clinometer (Figure 4.21). Azimuth was measured in to the nearest 5 degrees using a compass (Figure 4.22). Distance was measured to the nearest 0.25 meters using a laser rangefinder or tape measure (Figure 4.23). The combination of these three values is defined as “pocometry” (polar coordinate metrics) by Takenaka et al. (1998).

Note that the distance measurement describes the length of the branch in a direction, rather than the extension from a central axis as used in the cylindrical coordinate systems in Van Pelt et al. (2004b).

4) Crown Structure – Methods

Figure 4.21: Vector slope illustrated in 100yr tree 2 at 42.8metres. Note the inherent difficulty expressing 3-d concepts in 2-d images.

4) Crown Structure – Methods

Figure 4.22:Vector azimuth illustrated in 100yr tree 2 at 42.8metres. Note the inherent difficulty expressing 3-d concepts in 2-d images

. Figure 4.23: Vector distance illustrated in 100yr tree 2 at 42.8metres. Note the inherent difficulty expressing 3-d concepts in 2-d images.

Topology: Defined in the vector name: All vectors were named so as to distinguish between them and to reconnect them into a hierarchical branching system. The number of digits in its lineage name is equivalent to its branching from the trunk (Figures 4.24, 4.25).

The lineage of a vector is encoded within its name. Vector names take the form:

t[Tree Number]_[height]b[branch number sequence]

where

“t” is a prefix character indicating “tree number”,

[Tree number] is an identifier of which tree the branch is located in,

“_” is a divider character,

4) Crown Structure – Methods

[height] is the height of the branch above the ground at the point it is attached to the trunk, to one decimal place,

“b” is a divider character, and

[branch number sequence] is a listing of single digits to record the lineage of each branch. The first digit identifies which branch it is connected to the trunk through. The following digits identify the following generations.

Thus, t02_42.8.2b1 is the first branch arising from a height of 42.8 metres. Another sister branch arising at the same height would be named t02_42.8b2. If three branches arose from this second branch, they would be named t02_42.8b21, t02_42.8b22, and t02_42.8b23.

If t02_42.8b22 had children they would be identified according to the same pattern by adding another digit to their name: t02_42.8b221, t02_42.8b222, t02_42.8b223, and so on. This last branch is not “two hundred and twenty three,” but rather is “two two three.” Note that this system could use any name or letter instead of numbers. While slightly intimidating at first look, the branch names hold the information about its lineage from the base of the tree.

Simple summaries of complex branching systems use the name of the basal segment, with the suffix “sys” added to distinguish it from the data recorded for the basal segment. If 10 sister branches exist, another divider character “b” can be added.

The simple, coarse resolution branch summary associated with a complex branch was labelled bSys, for “system”.

4) Crown Structure – Methods

>

=

3

b

8 < . = 2 4 4 2 .8 b 2

Trunk at <=42.8b1 42.8m

Figure 4.24: Simple branches. In tree 2, branch 42.8 and its children branches described at a lower resolution as a whole branch with a single terminal vector. The elements identified as 42.8b21, b22, b23, and b24 in the preceding figure are now branchlets below the resolution level. In the data archive, this “system summary” of a branch would be labeled as t02_b42.8bsys.

4) Crown Structure – Methods

< = b < 2 = 3 b < 3 =b 1 => 22 b32 > 4= b2 <=b21

< = 4 42.8b3=> 2 .8 b 2

Trunk at <=42.8b1 42.8m

. Figure 4.25: Complex branches. Branch 42.8 and its children branches described at a higher resolution. Branch naming system illustrated in100yr tree 2 at highest climbed point of 42.8m. The resolution applied to branch 42.8b2 results in the descendants of this branch anchor point described with one basal segment and four terminal vectors. In the data archive, these would be recorded as nine vectors named separately. .

By using the height of the branch, branch names are more parsimonious than if a new branching point was considered at each branch encountered along the trunk. This system would be more accurate if applied only to original branches, but the loss of buds, loss of branches, and the presence of resprouts at different points along vectors confuses the issue. If plants faithfully followed their architectural growth model (Halle, 1995), without losing buds, losing branches, or resprouting, each branching node along the could be treated as a new parent location.

Van Pelt et al. (2004b) use a slightly different naming system for branch lineages, developed from experiences in conifers. They identify basal branch segments with the prefix M, and use a different divider character for segments and terminal branches. 4.3 Field Recordings

4) Crown Structure – Methods

Vectors: Working downwards in a spiral from the uppermost climbed branch, every branch originating from the trunk was measured as a simple branch, i.e. one vector. Larger, more complex branches were further represented by branch segments leading to terminal branches. Resolution, naming and vector definition were as described above. With the help of a data recorder, each vector took approximately 90 seconds to record, not counting the time required in climbing to each piece.

Circumferences: Branch segments were represented by an orthogonally truncated cone built around the branch vector. Start and end circumferences were measured around the vector by tape measure (Figure 4.26). Measurements avoided the swelling of wood at a branching node. For terminal branches, only a start circumference was recorded, forming a cone. Small, dead terminal branch stubs were recorded with an end circumference when they had no children branches summarized by the vector.

Figure 4.26: Start and end circumference locations on an original branch basal segment in old tree 4, (with D. Catchpole.)

The term megabranch was subjectively applied to describe the largest branches on an old tree (Junker, 2004). These were invariably larger in girth than the arm span of an average human (1.5m circumference), and were often contender leaders (Figure 4.27).

4) Crown Structure – Methods

Trunk rings: Trunk sections were special cases of branch segments, in which the end circumference, or trunk ring, of one section was also the start circumference of another section. The shape of the tree trunk was quantified by measuring circumference at five metre intervals and at 1.34 metres (chest height), forming a sequence of stacked orthogonally truncated cones (Figure 4.28).

Figure 4.27 (right): Two original megabranches in old tree 12, with R. Junker placing an insect trap. Figure 4.28 (left): A. Barrows measuring the circumference of 100yr tree 11 at 10 metres height.

Anchor azimuth: A second azimuth was recorded for non-radial branches, in which the vector azimuth is different from the azimuth between the trunk centre and the attachment point (Figure 4.29). Epicormic branches frequently begin at one point and point in different directions, and are misrepresented by vectors originating from the trunk centre (Figure 4.30). Because the branches were measured from the point at which they connect to the trunk surface, a separate trunk anchor azimuth was recorded for non-radial branches. This number was not recorded for radial branches, for which trunk anchor azimuth equals branch vector azimuth.

4) Crown Structure – Methods

Figure 4.29 (left): Radial and non-radial branches illustrated. Figure 4.30 (right):The epicormic crown of this old E. obliqua one km south of the study site would be misrepresented by radial vectors.

Distance along parent: The distance along the parent branch was recorded for branches that did not originate from the end of its parent vector (Figure 4.31). The branches were numbered and named sequentially from the base outwards. When branches originated from the end of the vector, this distance value was not recorded. Measuring height along the trunk is a special case of this technique.

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An ass

4) Crown Structure – Methods

Figure 4.31: Distance along parent vector illustrated. The first three children branches, as counted from right to left, are recorded with a distance along. The three leftmost children originate from the vector end. All but the leftmost vector are epicormic resprouts from the upper surface. The dead vector is the original terminal bud.

Airspace Boundary Fields: To further describe the branching structure below the resolution at which terminal branches were summarized, four field vectors were defined for each branch encompassing the branchlets and foliage (Figures 4.32, 4.33). Angular measurements of the lower, upper, left, and right extent of the foliage were recorded and used to create vectors in spherical space with the same starting point and equivalent distance as the branch vector. These four measurements were easily obtained and accommodate asymmetrical branches. Vertical and horizontal arcs were calculated by summing down with up, and left with right, respectively.

Van Pelt et al. (2004b) used linear spread and linear depth to encompass the foliage along its widest and tallest axes. These linear measurements were much less vulnerable to error from the distance of the branch, but mask asymmetry.

Figure 4.32: Vertical boundary airspaces illustrated on vertical and horizontal planes. Note the inherent difficulty expressing 3-d concepts in 2-d images

4) Crown Structure – Methods

Figures 4.33: Horizontal boundary airspaces illustrated on vertical and horizontal planes. Note the inherent difficulty expressing 3-d concepts in 2-d images.

Foliage: A calibrated estimate of foliar units was recorded for each branch vector. Foliar units in E. obliqua are the “globs” or “balls” of leaves (Figure 4.34). Jacobs (1955) refers to them as “crown units”. Van Pelt et al. (2004b) refers to “repeating clusters of stems and foliage” and adopts the term “foliar units.”

A foliar unit is a branchlet because it is below the resolution level. It is simultaneously defined by its size and its penultimate terminal branching generation. Unlike branches, the foliar unit branchlets were not fixed in space or in the topology. They were counted as a continuous variable associated with a vector.

In the study trees these units were approximately 0.75m in diameter. Inherent differences between the size of these units in different trees, situations, and heights created errors in these estimates. Van Pelt et al. (2004b) and Ishii et al. (2002) discussed the concept in detail and highlight difficulties in unit measurements, and Kelly et al. (2004) digitally quantified the foliar units of E. globulus. The foliar units used in the present research are provided as basic estimate only for comparison between branches within a tree and total foliage between trees.

4) Crown Structure – Methods

Figure 4.34: Approximate size and appearance of foliar units in 100yr tree 13, with R. Junker aloft. The actual volume filled by foliar unit was dependent on leaf density.

Percent living and percent gum-barked: The percentage of live wood was assessed and recorded for each vector. In addition, percentage of branching surface with “gummy” smooth bark was recorded (Forest_Resources, 1979; Jackson, 2000; Mortimer & Mortimer, 2003). On young branches, encountered high in the tree, branches of E. obliqua are smooth barked (Figure 4.35). They resemble branches of gum trees such as E. regnans and may indicate hybridization (Kirkpatrick & Backhouse, 1981).

4) Crown Structure – Methods

Figure 4.35: The gummy smooth barked upper trunk of old tree 4, with Sue Baker recording structural data.

Tree height and diameter. The height of each tree was measured from the ground using a laser rangefinder and clinometer using trigonometric techniques described by Goodwin (2004), Herrmann (2003), and Van Pelt (2002). This value may be different than the height generated by a computer model summing all of the vectors, and was considered less vulnerable to resolution effects. The diameter at breast height (1.3 metres) was taken using a tape measure.

Traps: The location of each trap was recorded within the framework of the mapping numbers. CD sticky and funnel traps were associated with a branch, or given a height and azimuth along the trunk. The height of each hangtrap was fixed in a cylindrical coordinate space by recording an azimuth and distance from the trunk.

4.4 Vector derived variable generation

Using the field data generated for whole branches, additional variables were calculated to describe each vector and the entire tree. These were the XYZ vector end points, trunk diameter at height, crown depth, trunk and branch cone volumes and surface area, crown and branch polyhedra volumes and surface area, and a scaling ratio between the branch and trunk diameters.

4) Crown Structure – Methods

XYZ conversion: For ease of analysis, the vectors collected for spherical coordinate vectors were converted into Cartesian XYZ coordinates vectors (www.wikipedia.org, 2004). Change in X, Y, and Z are each listed separately:

∆X-axis = distance * sine (90-slope) * cosine (1-azimuth) ∆Y-axis = distance * sine (90-slope) * sine (1-azimuth) ∆Z-axis = distance * cosine (90-slope)

Note the conversion of slopes by subtracting from 90, and the conversion of azimuths by adding a negative sign. This reflects the “right hand rule” in vector mathematics, a situation opposite to the clockwise azimuth readings. In the mathematical system, slope is measured from the overhead zenith down to the point, rather than upwards from the horizon. This conversion could be ignored, but the resulting points would be a rotated mirror image of the intended construction (Bentley Systems, 1998).

The benefit of conversion to XYZ Cartesian coordinates (hereafter XYZ) is the ease with which vectors can be added. Summing the figures for dX, dY, and dZ separately generate the combined vector.

Trunk diameter functions: The measurements of trunk girth at different heights were regressed linearly for each tree to generate a simplified function defining radius (y) as a function of height along trunk (x) in the form y=mx +b. where m and b are variables for each tree. Field data was collected in circumferences, analysis in diameters, and computer graphic in radii.

Crown depth: The crown depth was generated for each tree by subtracting the height of the lowest branch from the total tree height.

Cone Volume: The wood volume of a branch was measured in surrogate by the vector cone volume. For terminal branches, the vector volume was assumed to represent that of the summarized branchlets below the resolution level. Volume for the trunk and vector cones was calculated using the formula in Van Pelt et al. (2004b): Volume in m3= (π/3)*distance*((0.5diamA)2+((0.5diamB)2)+((diamA*diamB)/4))

4) Crown Structure – Methods

where distance= length of cone segment diamA = diameter of ring in metres diamB= diameter of ring in metres

Total trunk wood volume was generated by adding the cone volumes for each trunk segment.

Cone Surface area: Similarly, the surface area was calculated:

Surface area in m2 =

(((π*diamA)*(π*diamB))/2) * distance

where distance= length of fustrum diamA = diameter of start ring in metres diamB= diameter of end ring in metres

Minimum polyhedra: The crown airspace volume encompassed by the entire tree crown was calculated by fitting a minimum convex polyhedra around the outermost XYZ points using the computer program Mathematica 4.1 (Kelly, 2004; Wolfram Research, 2001). These polyhedra are computer-generated envelopes of joined triangle planes. They are computed to encompass the smallest possible volume that encloses all branches. The branch vector ends form the vertex at which these planes are connected. Minimum polyhedra can be visualized as “shrink-wrapping in crystal.” The trunk and the base of the tree were excluded. Crown surface areas of the polygonal shape were also generated.

The branch airspace and surface area encompassed by each terminal branch’s branchlet and foliage airspace field was calculated using minimum polyhedra in a similar fashion. Six points were entered: branch vector start, branch vector end, and the four airspace boundary vector ends.

Scaling factors: The ratio in size between a branch and its parent was calculated and labelled the scaling factor (Figure 4.36).

4) Crown Structure – Methods

For simple branches, this is the starting diameter divided by the diameter of the trunk at height of attachment (calculated by linear regression as described above). For children branches, the diameter of the child branch was divided by the diameter of its parent vector at the start point was generated. If this start point was the end of the parent branch, then the ending diameter of the parent was the denominator in this calculation. When children arose from a point along the parent branch before the end, the diameter at that point was calculated arithmetically between the start and end circumferences at a portion of the distance.

Scaling factors: Diam b1 Diam b2

÷÷ Diam Trunk Diam Trunk 1

= = 0.69 0.76 Diam b1=> <= Diam b2 scaling scaling factor factor

Diam Trunk -> Figure 4.36: Scaling factors illustrated in 100yr tree 2 for two original branches.

4.5 Tree Structural Descriptor Generation

107 descriptor variables for each tree were used in analysis. These descriptor variables were organized for clarity in ten categories: 1. Tree Size 2. Branch Counts 3. Crown Depth 4. Cones 5. Polyhedra 6. Foliage

4) Crown Structure – Methods

7. Start Diameters 8. Distance 9. Vertical plane 10. Horizontal plane

Tree-level variables, such as trunk volume, height, and DBH, were used unmodified. Branch-level variables were summarized for each tree using arithmetic means, standard deviations (“Std Dev”) sums, ranges (maximum value-minimums), maximums (“Max”, and proportions (described individually). Only simple branch data were used.

Due to small sample sizes, the presence of features was analysed separately by comparison and exposition rather than using a statistical technique.

Most statistics were run in triplicate for three branch types: all whole branches, living whole branches only, and completely dead branches only. Because a whole branch with any foliage was considered living, the small dead branches below the resolution level are under represented. Therefore, “dead only” should be interpreted as “completely dead simple branches only.”

When calculating crown depth statistics, the secondary trunks arising from the base of trees 6 and 7 were not counted.

The tree descriptor statistics generated are listed in Table 4.1.

4) Crown Structure – Methods

T Crow P Tre Si MeanDt Branchh Height (m) all Sumlh Polyhedrad Volume (m3) all Totalli e Height (m) DBH branches Mean Branch branches Sum Polyhedra Volume (m3) Foliage Units Mean Foliage Units Diamat13(m)Trunk Height (m) live Mean Branch Height live only Sum Polyhedra Volume (m3) live only Mean % of branch living all Volume (m3) Wood Volume (m3) (Trunk & (m) dead Mean branch ht as % of dead only Mean Polyhedra Volume (m3) all branches Mean % of branch gummy al Vector Total) TreePolyhedra tree height Mean live branch ht as % of treebr anches Mean Polyhedra Volume (m3) branches Volume (m3) TreePolyhedra Surface height Mean dead branch ht as % of treelive only Mean Polyhedra Volume (m3) D Area (m2) Ratio Polyhedra height Total Crown dead only Max Polyhedra Volume (m3) all itMean Distance (m) all m3:Trunk m3 % of Wood Depth (m) Range Branch Height (m) branches Max Polyhedra Volume (m3) branches Mean Distance Volume in Trunk% of Vector Volume in Dead live only Range Branch Height (m) live only Max Polyhedra Volume (m3) (m) live only Mean Distance (m) Branches dead only Std Dev Branch Height (m) all dead only Std Dev Polyhedra Volume (m3) all dead only Max Distance (m) all Branch branches Std Dev Branch Height (m) branches Std Dev Polyhedra Volume (m3) branches Max Distance Ct# Branches live only Std Dev Branch Height (m) live only Std Dev Polyhedra Volume (m3) (m) live only Max Distance (m) all branches # dead only Lowest Branch Height (m) all dead only Sum Polyhedra Surface Area (m2) all dead only Branches live on#ly branches Highest Branch Height (m) all branches Sum Polyhedra Surface Area (m2) Horizont Branches dead onl% yof Total Branches branches live only Sum Polyhedra Surface Area (m2) llMean Horizontal Arc (deg) al are Dead # Measured Vectors all dead only Mean Polyhedra Surface Area (m2) allbr anches Mean Horizontal Arc (deg) branches Sum Cone Volume (m3) all branches Mean Polyhedra Surface Area (m2) live only Mean Horizontal Arc (deg) Start branches Sum Cone Volume (m3) live only Mean Polyhedra Surface Area (m2) dead only Di Meant Start Diameter (m) all live only Sum Cone Volume (m3) dead only branches Mean Start Diameter (m) dead only Mean Cone Volume (m3) all Vertic live only Mean Start Diameter (m) branches Mean Cone Volume (m3) Meanll Slope (deg) all dead only Range Start Diameter (m) all live only Mean Cone Volume (m3) branches Mean Slope branches Range Start Diameter (m) dead only Max Cone Volume (m3) all (deg) live only Mean Slope (deg) live only Range Start Diameter (m) branches Max Cone Volume (m3) dead only Mean Vertical Arc (deg) all dead only Std Dev Start Diameter (m) allliv e only Max Cone Volume (m3) branches Mean Vertical Arc (deg) branches Std Dev Start Diameter (m) dead only Std Dev Cone Volume (m3) all live only Mean Vertical Arc (deg) live only Std Dev Start Diameter (m) branches Std Dev Cone Volume (m3) dead only Mean Down Arc (deg) all dead only Mean Scaling Ratio all live only Std Dev Cone Volume (m3) branches Mean Down Arc branches Mean Scaling dead only Sum Cone Surface Area (m2) all( deg) live only Mean Down Arc (deg) Ratio live only Mean Scaling Ratio branches Sum Cone Surface Area (m2) dead only Mean Up Arc (deg) all dead only Std Dev Scaling Ratio all live only Sum Cone Surface Area (m2) branches Mean Up Arc branches Std Dev Scaling dead only Mean Cone Surface Area (m2) all(deg) live only Mean Up Arc (deg) Ratio live only St Dev Scaling Ratio branches Mean Cone Surface Area (m2) dead only (Tree dead only Range Scaling Ratio all live only Mean Cone Surface Area (m2) 107 descriptorillustrat ion) branches Range Scaling dead only variables in 10 categories Ratio live only Range Scaling Ratio generated for dead only ach of 16 obliqua study trees

Table 4.1: 107 tree structural descriptor variables in ten categories 4.6 Analysis of structural descriptors

Univariate t-tests: T-tests were used to compare mean values of each descriptor variable between eight 100yr trees and the eight old-growth trees using JMP software ((SAS Institute, 2003). The null hypothesis that the descriptor was not different between age classes was tested. An assumption was made that the structural descriptors are normally distributed throughout the forest, and extended to the sampled trees. The goal of these tests was to identify quantitative aspects in which the crown structure was different between the two age classes, and to test Hypothesis 1 (Old E. obliqua are quantifiably and objectively distinct from 100yr trees, i.e. old trees are not scaled-up versions of 100yr trees. )

NMS ordination: Non-metric multidimensional scaling ordination of the 16 trees was performed using PC-ORD software, incorporating all 107 descriptors (McCune, 1999). This was used to show how the descriptors organised the trees by structural similarity. Data was square root transformed. The Sorensen distance measure between each combination of two trees was calculated in a computer n- dimensional hyperspace, with each of 107 dimensional axes ranking the trees by a

4) Crown Structure – Methods specific descriptor variable. Computing power is applied in depicting their hyper- dimensional spatial relationship in a 2 or 3-dimensional picture (Clarke & Warwick, 2001; McCune, 1999). The reported stress levels offer a comparative measure of how accurate this low-dimensional picture represents the hyperspace picture. This tested Hypotheses 1 (Old E. obliqua are quantifiably and objectively distinct from 100yr trees, i.e. old trees are not scaled-up versions of 100yr trees) and 2 (Old E. obliqua have a more variable crown structure than 100yr E. obliqua).

Dendrogram: Cluster analysis was performed by generating a dendrogram using Wards linkage method with PC-ORD. This branching diagram groups the trees by their relationships in the descriptor variables (Clarke & Warwick, 2001; McCune, 1999). Like the NMS, this was used to assess the distinction between age classes based on the structural descriptors, and to test Hypothesis 1 (Old E. obliqua are quantifiably and objectively distinct from 100yr trees, i.e. old trees are not scaled-up versions of 100yr trees)

Assumptions in multivariate analysis: The use of these multivariate techniques (NMS and cluster analysis) with the 107 structural descriptors assumes that all of these variables contribute equally to the definition of the tree. Despite intrinsic correlations between variables (e.g. cone volume and surface area), these multivariate techniques offer a useful method of visualization. However, measures of cones and polyhedra will contribute disproportionately because of the number of descriptors generated using them. Furthermore, some measures are perfect correlated. For example, the maximum cone volume of all branches is always equivalent to the maximum cone volume of live branches. Interpretation of the resulting graphical displays must bear this assumption in mind. 4.7 Arborograph generation and analysis

A virtual computer model was generated for each tree. Microstation Computer Aided Design software (CAD) version 05.07.00.41 was used to create a virtual 3-d computer arborograph (Bentley Systems, 1998). CAD engineering drafting software can create 3-d models using scripted text files. Once the arborographs are scripted, the possibilities for illustration are limitless. Foliage can be rendered on to the branches to add to the verisimilitude of the models (Figures

4) Crown Structure – Methods

4.36, 4.37). A photo of the stringy bark of E. obliqua was rendered as the “wallpaper” around the branch and trunk vectors (Figures 4.38, 4.39).

Figure 4.36 (far left): Rendering of E. obliqua foliage onto the branches of 100yr tree 1 Figure 4.37 (mid left): Rendering of E. obliqua foliage onto the branches of old tree 3 Figure 4.38 (mid right): Stringybark rendered onto side view arborograph of old tree 3, illustrating branch t03_32.2b1 Figure 4.39 (far right): Cropped photo of stringybark surface used for rendering

Six layers were created for each tree (Figure 4.40, overlead): 1. Trunk 2. Cone 3. Line 4. Cross 5. Kite 6. Extras

The full lineup of arborographs is in Appendix 1. Instructions for generating Microstation CAD scripts from the vector-based structural data are given in Appendix 2. In addition, the sample text scripts used in creating Figure 4.40 are provided in Appendix 3.

4) Crown Structure – Methods

Figure 4.40: Layers used in CAD arborographics. The scripts used in generating these images can be found in Appendix 3.

4) Crown Structure – Methods

4.8 Image file size analysis

A novel, simple method for comparing visual complexity of the generated tree models was developed. The mean file size in kilobytes of the computer graphic file depicting eight 100yr and eight old tree arborographs was compared using t-tests (Figure 4.41). This was used to test Hypothesis 4 (Old E. obliqua have a greater complexity than 100yr trees, as measured in surrogate by the amount of information required to describe them).

Graphical output from Microstation CAD software was in .PNG (Portable Networks Graphic) format, a format analogous to .GIF (Graphical Interchange Format). These vector based image recording formats summarize lines precisely and efficiently, and are more appropriate than .BMP bitmaps or .JPG (Joint Photo Experts Group) images. Bitmaps are raster-based, and record every pixel independently. Bitmaps are unsuitable because they record an equivalent amount of information for empty white background pixels as for filled content pixels. Photo JPG format uses a complex algorithm to summarize shades of colour too similar for the human eye to distinguish.

Implicit in this technique is the designation of complex things taking more effort to describe than simple things (Anand & Orloci, 1996; Roche, 2001). The practical outcome of this can be seen in the download times required for elaborate web pages, the length of a scientific article describing complicated field work, or the increased price of long distance phone calls when one has many stories to tell.

Comparing the file size of graphical output from Microstation CAD is feasible because all images were standardized. The repetition of eight camera angles allows for comparison between trees. The actual resolution level used in the different age classes may be a further source of bias, as complex branches were used rather than their whole branch summaries. The image file size is undoubtedly correlated with the measured vector count. This resolution level, in analogy with a long distance telephone call, is the judgement of the speaker as to what stories are worth telling.

Once computer models are generated, generating these variables is a simple process aided by the CAD software image output functions. File sizes were

4) Crown Structure – Methods calculated using Microsoft Windows NT computer operating system (Microsoft, 2000).

Figure 4.41: The camera angles used in image output and analysis, with tree 10 cones and trunk layers as an example. In addition, analysis was done with these layers plus the kite, cross, and line layers.

4) Crown Structure – Methods

4.9 Structural features

During field data collection, notes on selected structural habitat features were recorded. The location of features was recorded as accurately as possible in the framework of the vectors. The general types of features are presented below. Photographs and notes on specific occurrences are presented with the results.

Three distinct biases exist in this data collection: 1. Recognition bias, the researcher’s recognition of what constitutes a feature of interest (analogous to an indicator species), 2. Observation bias, the possibility of missing features in the inaccessible outer or upper crown (analogous to cryptic taxa), and 3. Distinction bias, the lumping of features with different ontogeny by their similar appearance (analogous to polyphyletic taxa).

Several of the recorded features are described as or related to hollows. Hollow formation in Eucalyptus is a complex, stochastic process (Ball et al., 1999). Distinction, observation, and recognition biases add difficulty to field recordings. Further information on animal and fungal associations with E. obliqua hollows is available through Harrison et al. (2004) and for the genus Eucalyptus through Ambrose (1981), Gibbons & Lindenmayer (2002), and Mackowski (1984).

Following is a listing of features recorded. A further description of each is presented in the results section. The presence of the distinction bias made distinguishing between the features both a method and a result.

1. Trough branches 2. Elbow hollows 3. Litter collections 4. Vascular epiphytes 5. Burls 6. Dead tops 7. Snapped main stems 8. Secondary trunks 9. Gallery branches 10. Dead flanks

4) Crown Structure – Methods

11. Collar hollows 12. Trunk fissures

5) Crown Structure - Results

5) Crown Structure – Results

Figure 5.1 Isometric view arborographs of the sixteen study trees.

5) Crown Structure - Results

Figure 5.2 Front view arborographs of the sixteen study trees

5) Crown Structure - Results

The results of the analysis of structural descriptors is presented in this chapter. T-test results comparing the mean values of descriptors are listed. The ten categories are: tree size, branch counts, crown depth, cones, polyhedra, foliage, start diameters, distance, vertical plane, and horizontal plane. Several differences between age classes were identified, and the meaning of the descriptors discussed. Following, the multivariate classification of study trees is presented, showing a strong distinction between age classes. Next, file size analysis of image complexity is reported. Old trees had significantly more complex visual arborographs. Lastly, occurrences of tree structural features are detailed. All structural features were present more often in old trees than 100yr trees. 5.1 Total numbers

Approximately 10,800 numbers describing the crown structure of the sixteen trees were recorded. These data form ~1,000 vectors representing 538 simple branches and 16 trunks for the 16 trees. Combining measurements from all trees, the arborographs represent organisms totalling 0.842 kilometres of vertical height, ~ 980 cubic metres of wood volume, and encompassing ~49,024 cubic metres of airspace. Isometric and front views of the arborographs are presented in Figures 5.1 and 5.2. 5.2 Summary and range of structural descriptors.

Basic summaries of size are given for each tree in Table 5.1.

At 72.4 m, old tree 4 was the tallest of the study trees, and 100yr tree 15 was the shortest at 32.8 m. There was a height difference of 39.8 m between these two trees. Old tree 10 had the most trunk volume, at 117 m3, and 100yr tree 15 was the smallest, with one-twentieth the volume at 6.4 m3. The largest crown airspace was occupied by old tree 16, encompassing 6588 m3, and the smallest airspace was again 100yr tree 15, with one-seventh the airspace at 891 m3.

5) Crown Structure - Results

Height in Diameter at 1.34 Trunk Wood Airspace Tree Number Name Age class Pair metres metres Volume m3 Volume m3 1 Crowofine 100yr A 51.9 1.05 16.7 1442.0 2 Rocky 100yr B 46.1 1.27 21.7 2837.2 5 Wigachu 100yr C 46.1 1.07 13.4 2587.7 7 Blaze Tree 100yr D 47.8 0.88 10.2 1472.1 9 Cmdr Keen 100yr E 37.2 0.99 11.7 3313.3 11 Oxylobl 100yr F 44.8 0.92 15.0 992.2 13 Peasybaum 100yr G 49.4 1.35 18.1 2039.5 15 KeepyLefty 100yr H 32.8 0.80 6.4 891.4 100year Mean 44.5 1.0 14.1 1946.9 100yr Std Dev 6.4 0.2 4.8 893.2

3 Lord Niles Old A 66.8 2.16 79.5 4540.9 4 Gemini B Old B 72.4 2.31 73.2 2870.8 6 Deedlum Old C 61.1 2.64 89.8 3619.1 8 Vorticon Old D 54.0 3.49 92.8 5541.6 10 McMire Old E 53.4 3.04 116.6 3656.4 12 Megadendra Old F 59.8 2.88 103.3 4596.1 14 Bogelhart Old G 62.0 2.86 164.0 2050.6 16 Pepperberry Old H 56.9 2.96 59.5 6587.8 Old Mean 60.8 2.8 97.3 4182.9 Old St Dev 6.4 0.4 32.2 1453.7

p<0.05? * * * *

Table 5.1. Basic size measures for each tree, compared with t-tests between age classes. 5.3 Univariate T-tests

Of the 107 descriptor variables, 53 exhibited significant differences between 100yr and old E. obliqua trees. Note that a mean and standard deviation is used to summarize descriptor variables that are themselves means, standard deviations, ranges, or arithmetic relationships of the branch population within a tree.

5.3.2 Tree Size

Tree Size: 100yr Mean Std Dev Old Mean Std Dev p<0.05?

Tree Height (m) 44.51 6.38 60.80 6.43 <.005 DBH Diam at 1.3 (m) 1.04 0.19 2.79 0.42 <.005 Trunk Volume (m3) 14.14 4.80 97.32 32.19 <.005

Wood Volume (m3) (Trunk and Vector Total) 15.60 5.43 107.12 32.91 <.005

TreePolyhedra Volume (m3) 1946.91 893.23 4182.92 1453.66 <.005 TreePolyhedra Surface Area (m2) 892.23 249.96 1569.13 317.47 <.005 Ratio Polyhedra m3:Trunk m3 144.56 68.15 49.44 28.86 <.005 % of Wood Volume in Trunk 90.88 2.90 90.49 3.89 ns % of Vector Volume in Dead Branches 12.79 15.63 6.37 4.17 ns

Table 5.2: Comparison of tree size descriptors The old trees were significantly taller and bigger than their 100yr counterparts (Table 5.2). The difference in diameter at 1.34 metres was expected, as this was the primary selection criterion to distinguish between the age classes. On average, the old trees were 16 metres taller, and more than 7 times the wood volume. The crown

5) Crown Structure - Results

airspace volume occupied by an old large tree was only twice that of a 100yr old tree, implying substantial wood invested in the trunk rather than the outer foliage. Relative to the trunk size, the crowns of 100yr old trees were three times larger, due to the narrow trunks and long horizontal branches of these trees.

No significant differences were detected in the mean proportion of total wood volume invested in the trunk instead of branches. In both age classes, 90% of the wood volume was in the trunk. This value was between 85% and 95% for all trees; even though branches and trunks tended to be larger in the old trees (see below), their proportions were the same.

In 100yr old trees, the proportion of branch cone volume in dead whole branches was twice that of old trees, but this difference was not significant.

5.3.3 Branch Counts

Branch Counts: 100yr Mean Std Dev Old Mean Std Dev p<0.05?

# Branches all branches 31.5 6.5 35.9 16.7 ns # Branches live only 21.5 5.3 22.1 9.9 ns # Branches dead only 10.0 5.4 13.8 7.7 ns % of Total Branches are Dead dead only 30.86 15.01 35.96 10.72 ns # Measured Vectors all branches 35.5 5.9 77.1 33.0 <.005

Table 5.3: Comparison of branch count descriptors The number of whole branches originating from the trunk was not significantly different between ages (Table 5.3). On average, 33 branches arose from the trunk of each study tree. No significant difference was detected in the number of live or completely dead whole branches, or the proportion of whole branches that were dead. For all of these descriptors, the mean was slightly higher in old trees.

The mean number of vectors measured in each old tree was twice that of the 100yr old trees. This reflects the opinion of the measurer on what constituted a branch segment, and was aimed at a ~15 cm diameter resolution level. Another measure related to vector count is the size of the computer image files (see below).

5.3.4 Crown Depth

5) Crown Structure - Results

Crown Depth: 100yr Mean Std Dev Old Mean Std Dev p<0.05?

Mean Branch Height (m) all branches 31.24 4.24 33.69 9.67 ns Mean Branch Height (m) live 32.92 4.92 35.94 8.81 ns Mean Branch Height (m) dead 28.38 3.57 29.64 11.37 ns Mean branch ht as % of tree height 70.341 3.451 54.972 12.019 <.005 Mean live branch ht as % of tree height 74.117 5.749 58.734 10.242 <.005 Mean dead branch ht as % of tree height 63.952 3.510 48.229 15.342 <.05 Total Crown Depth (m) 11.60 3.09 24.86 5.77 <.005 Range Branch Height (m) live only 17.26 5.23 32.41 7.15 <.005 Range Branch Height (m) dead only 18.89 10.31 25.11 8.35 ns Std Dev Branch Height (m) all branches 6.35 1.06 10.09 2.08 <.005 Std Dev Branch Height (m) live only 5.21 1.56 9.71 2.25 <.005 Std Dev Branch Height (m) dead only 6.67 3.13 8.98 1.60 ns Lowest Branch Height (m) all branches 16.41 6.62 15.59 8.89 ns Highest Branch Height (m) all branches 39.36 5.40 48.93 8.00 <.05

Table 5.4: Comparison of crown depth descriptors

Tree 16 (old) Tree 9 (100yr) Arborographs of 100yr and old trees with horizontal axis stretched three times. Flattening the image helps to show the regular, original crown of the 100yr trees, and the irregular resprouted crown of the old tree. Gray cones are completely dead branches. Note the large number of failed epicormics in tree 16, especially beneath large, successful resprouts.

Figure 5.3: Crown depths visually compared by horizontally stretched arborographs The average height of branch attachment to the trunks of all study trees was 32 metres (Table 5.4). No significant differences was detected in the average height of branches. This could be linked to related to the canopy height of nearby Nothofagus cunninghamii, Acacia melanoxylon, Acacia dealbata, and Pomaderris apetala. The standard deviation of mean branch height for old trees was twice that of 100yr trees, showing a less centralized crown.

The mean height of the dead branches was almost always lower than the mean height of the live branches. However, distinct processes in the different age classes may have caused this (Figure 5.3). In 100yr trees, a live crown was situated over a dead crown (Figures 5.4, 5.5). The lower dead crown is made of previous original branches that were being abandoned as the trees grew upward competing for light (Jacobs, 1955). By contrast, in old trees, the lowest original branches had been occluded in the past, and dead branches arising from the lower trunk were more recent epicormic resprouts that had failed to succeed (Figure 5.6).

5) Crown Structure - Results

Figure 5.4 (left): Dead original branches in lower crown of 100yr tree 1, with R. Junker ascending Figure 5.5 (centre): The dead original branches in the lower crown of 100yr tree 7 are visible below the upper live crown of original branches. R. Junker is nearby a snapped dead branch Figure 5.6 (right): The lower crown of old tree 8 visible from 100yr tree 7, with failed epicormic branches visible. A contradictory situation occurred when comparing the mean branch height as a percentage of the total tree height (Figure 5.7). For all, live only, and dead branches, this measure was significantly higher in the 100yr trees. 100yr old trees are more “top heavy” than the old trees. Old trees are already in a dominant position and have low epicormic branches that can take advantage of light conditions caused by canopy gaps caused by tree fall or fire (Franklin et al., 2002; Ishii & Ford, 2001; Ishii & Wilson, 2001; Jacobs, 1955). By contrast, the 100yr trees are investing in growing upwards in competition with their cohort, and are abandoning the original branches in the lower crown (Falster & Westoby, 2003; Jacobs, 1955).

Mean height of tree (column) and mean height of branches (diamond) 80

70

60

50

40

30 Height in metres Height 20

10

0 12579111315346810121416 100yr Old Tree Number

Figure 5.7: Mean height of branches (diamond mark) plotted against tree height (column)

5) Crown Structure - Results

For similar reasons, the depth of the crown, and the range in heights for live branches, was greater in the old trees. The height range of dead branches was insignificantly greater in old trees.

A related measure of crown depth is the standard deviation of branch heights. These followed the same pattern as the range in heights: significantly greater in old trees for all branches and live branches but not the dead branches. Old trees had more dead branches in the upper crown than the 100yr trees.

For both age classes, the mean lowest branch height was approximately 16 metres, perhaps in response to the height of the other tree species nearby. A significant difference in the height of the mean highest branch was found. This measure, however, reflects both the objective difference in tree heights between age classes, and the subjective ability of the climber to access the uppermost crown of some trees.

5.3.5 Cones

Cones: 100yr Mean Std Dev Old Mean Std Dev p<0.05?

Sum Cone Volume (m3) all branches 1.46 0.82 9.80 3.81 <.005 Sum Cone Volume (m3) live only 1.32 0.88 9.24 3.86 <.005 Sum Cone Volume (m3) dead only 0.14 0.17 0.55 0.40 <.05 Mean Cone Volume (m3) all branches 0.05 0.02 0.39 0.31 <.01 Mean Cone Volume (m3) live only 0.06 0.03 0.55 0.41 <.01 Mean Cone Volume (m3) dead only 0.01 0.01 0.07 0.10 ns Max Cone Volume (m3) all branches 0.27 0.17 4.20 3.94 <.05 Max Cone Volume (m3) live only 0.23 0.17 4.20 3.94 <.05 Max Cone Volume (m3) dead only 0.08 0.14 0.26 0.22 ns Std Dev Cone Volume (m3) all branches 0.06 0.03 0.92 0.95 <.05 Std Dev Cone Volume (m3) live only 0.05 0.03 1.10 1.18 <.05 Std Dev Cone Volume (m3) dead only 0.02 0.04 0.09 0.08 ns Sum Cone Surface Area (m2) all branches 46.64 17.15 110.54 29.48 <.005 Sum Cone Surface Area (m2) live only 40.90 18.86 99.39 26.32 <.005 Sum Cone Surface Area (m2) dead only 5.74 4.82 11.16 6.51 ns Mean Cone Surface Area (m2) all branches 1.48 0.48 3.71 1.70 <.005 Mean Cone Surface Area (m2) live only 1.86 0.62 5.14 1.97 <.005 Mean Cone Surface Area (m2) dead only 0.52 0.28 1.05 0.82 ns

Table 5.5: Comparison of cone descriptors Descriptors of the volumes and surface areas associated with the vector cones representing branches show similar patterns to that expressed before: significant differences for all branches and live branches, but not for dead branches (Table 5.5).

The total of wood volume in branches (all and live only) was ~6 times greater in the old trees. The proportion of this volume in dead branches was not significantly different between ages, as described above. However, the total volume of completely dead branches was ~4 times greater in the older trees. This number is undoubtedly an underestimate, due to dead branchlets below the resolution level.

5) Crown Structure - Results

The mean cone volume followed the described pattern. The average volume of dead whole branches in old trees was 7 times greater than in 100yr trees, but a large standard deviation makes this a statistically insignificant result. Dead branches in 100yr trees were abandoned originals being self-pruned, whereas those in old trees were a combination of small failed epicormics and large broken stubs of originals. This variability in dead branch size is expressed in a higher standard deviation of cone volume.

The mean of the standard deviation of cone volume illustrates the much greater range of live branch sizes in the old trees, with this measure of variability twice as great as in the 100yr trees. Old branches have huge original “megabranches” (Junker, 2004), large old epicormics, and recent small epicormics.

The maximum cone volume again followed the described pattern. Results for the sum and mean surface area mirrored the cone volume results, as they are calculated similarly.

5.3.6 Polyhedra

Polyhedra: 100yr Mean Std Dev Old Mean Std Dev p<0.05?

Sum Polyhedra Volume (m3) all branches 1128.59 666.68 1923.88 915.97 ns Sum Polyhedra Volume (m3) live only 232.11 156.27 973.48 367.14 <.005 Sum Polyhedra Volume (m3) dead only 42.36 50.11 106.25 74.21 ns Mean Polyhedra Volume (m3) all branches 34.86 18.39 54.55 10.94 <.05 Mean Polyhedra Volume (m3) live only 47.14 20.28 82.97 11.01 <.005 Mean Polyhedra Volume (m3) dead only 3.96 2.90 3.49 3.24 ns Max Polyhedra Volume (m3) all branches 140.71 55.89 415.19 202.24 <.005 Max Polyhedra Volume (m3) live only 140.71 55.89 415.19 202.24 <.005 Max Polyhedra Volume (m3) dead only 17.72 12.36 37.02 59.53 ns Std Dev Polyhedra Volume (m3) all branches 35.71 14.23 90.94 31.22 <.005 Std Dev Polyhedra Volume (m3) live only 35.80 13.24 102.51 35.79 <.005 Std Dev Polyhedra Volume (m3) dead only 5.85 3.90 9.58 12.74 ns Sum Polyhedra Surface Area (m2) all branches 467.86 370.80 1171.53 632.78 <.05 Sum Polyhedra Surface Area (m2) live only 464.43 371.67 1151.53 629.77 <.05 Sum Polyhedra Surface Area (m2) dead only 3.43 2.35 20.00 46.15 ns Mean Polyhedra Surface Area (m2) all branches 14.25 10.86 32.57 14.67 <.05 Mean Polyhedra Surface Area (m2) live only 19.43 12.57 50.21 19.70 <.005 Mean Polyhedra Surface Area (m2) dead only 0.39 0.33 1.01 2.18 ns

Table 5.6: Comparison of polyhedra descriptors T-test results for the minimum convex polyhedra describing the airspace occupied by branchlets and foliage were similar to those for the cone volumes, with two exceptions (Table 5.6).

The mean sum of airspace cubic metres for all branches, and for dead branches only, was not significantly different between age classes. This contrasts with the results for total crown airspace, which was twice as great in the old trees. This suggests a sparser canopy in the larger, more irregular trees than in the more regular 100yr trees.

5) Crown Structure - Results

For all other comparison of polyhedra volumes- mean per branch, maximum, standard deviation- all branches and live only again exhibited significant differences, whereas dead only did not.

Mean polyhedra surface area results the same as the mean volume results. For old trees, a significantly higher surface area was found for all branches and live branches only. However, the total polyhedra surface area results were different. The sum of all branches surface area was significantly different, but this was not the case for the total of the volumes. If the surface area is greater in old trees, but the volume is not, this would imply a less efficient “packing” of these contents in an equivalent airspace (Kelly et al., 2004). A more efficient “packing” system would be a more regular airspace with symmetrical boundaries, as encountered in the 100yr trees.

5.3.7 Foliage

Foliage: 100yr Mean Std Dev Old Mean Std Dev p<0.05?

Total Foliage Units 134.5 52.8 232.0 62.4 <.005 Mean Foliage Units live only 4.21 1.20 6.75 3.82 ns Mean % of branch living all branches 55.92 15.76 45.09 9.72 ns Mean % of branch gummy all branches 32.01 17.23 13.57 8.52 <.05

Table 5.7: Comparison of foliage descriptors Old trees carried nearly twice as much foliage as 100yr trees, a value which corresponds neatly to other measures of crown size: total branch polyhedra volume is also double in old trees, as is crown polyhedra volume (Table 5.7).

The mean number of foliage units per branch was not significantly different between age classes, nor was the percentage of branches that were dead.

Branches in 100yr trees showed a significantly higher amount of gumminess, especially in the upper crown. Gumminess in old trees was confined to smaller branchlets and young epicormics.

5.3.8 Start Diameters

5) Crown Structure - Results

100yr Old Start Diameters: Std Std p<0.05? Mean Dev Mean Dev Mean Start Diameter (m) all branches 0.14 0.03 0.27 0.09 <.005 Mean Start Diameter (m) live only 0.15 0.03 0.32 0.07 <.005 Mean Start Diameter (m) dead only 0.09 0.01 0.20 0.14 <.05 Range Start Diameter (m) all branches 0.24 0.06 0.84 0.26 <.005 Range Start Diameter (m) live only 0.22 0.07 0.82 0.25 <.005 Range Start Diameter (m) dead only 0.12 0.07 0.35 0.19 <.01 Std Dev Start Diameter (m) all branches 0.06 0.02 0.21 0.07 <.005 Std Dev Start Diameter (m) live only 0.05 0.01 0.23 0.09 <.005 Std Dev Start Diameter (m) dead only 0.04 0.02 0.10 0.04 <.005 Mean Scaling Ratio all branches 0.35 0.07 0.28 0.07 ns Mean Scaling Ratio live only 0.42 0.08 0.34 0.08 ns Mean Scaling Ratio dead only 0.19 0.04 0.19 0.09 ns Std Dev Scaling Ratio all branches 0.18 0.06 0.26 0.08 <.05 Std Dev Scaling Ratio live only 0.18 0.07 0.29 0.10 <.05 St Dev Scaling Ratio dead only 0.06 0.05 0.15 0.06 <.01 Range Scaling Ratio all branches 0.75 0.24 1.22 0.62 ns Range Scaling Ratio live only 0.70 0.26 1.20 0.62 ns Range Scaling Ratio dead only 0.24 0.24 0.49 0.25 ns

Table 5.8: Comparison of branch diameter descriptors The average diameter at the base of branches in old trees was twice that of branches in 100yr trees, a statistically significant result (Table 5.8). In addition to the mean, the range and standard deviation of starting diameters was always higher in the old trees. As with the cone volumes, this is reflecting the similarity of the original branches in 100yr trees to each other, and the presence in old trees of large originals, large old epicormics and small recent epicormics.

Comparing the girth of the branch to the girth of the trunk, the average scaling factor for the branches exhibited no statistically significant differences between ages. Values for all branches and live only tended to be slightly higher in 100yr trees, a situation reflecting original branches that have been growing for the same time as the trunk at that height. This value is a quantitative measure of how much the larger bud branch lineage is dominating its lateral sister branches.

Surprisingly, the mean scaling factor for dead branches was the same between tree ages. This may be a coincidental relationship between the girth of the remaining lower crown stubs in 100yr trees and the occasional failed epicormic resprout in the upper or lower crown in old trees. This value may also reflect that the average size at death of an epicormic branch is fixed. This is the girth at which the branch is no competitive, i.e. a failed investment.

Different measures of variability in scaling ratio showed different patterns. For all branch types, the standard deviation of branch diameters in each tree showed significant differences between ages. However, the range of branch diameters was not significantly different.

5) Crown Structure - Results

The presence of mostly original branches in 100yr trees may be responsible for this divergence. Differences in significance between range and standard deviation could be caused by polymodal distributions of mean scaling factor in the old trees. A polymodal distribution could result in a higher standard deviation than a unimodal one, even if the range was the same. The similar results for the mean could reflect a coincidental balance in the central tendency.

If one assumes that there is a constant rate of growth that is somewhat similar between the original branches, and their big sister- the trunk upwards- then scaling factor should be unimodal for a tree with an original crown. When small, new epicormic branches sprout from the lower trunk, a polymodal distribution of scaling factors may result.

5.3.9 Distance

Distance: 100yr Mean Std Dev Old Mean Std Dev p<0.05?

Mean Distance (m) all branches 5.90 1.06 5.89 1.14 ns Mean Distance (m) live only 7.08 0.90 7.79 1.04 ns Mean Distance (m) dead only 2.95 0.93 2.54 0.86 ns Max Distance (m) all branches 13.44 3.60 18.13 4.82 <.05 Max Distance (m) live only 12.44 3.25 18.13 4.82 <.05 Max Distance (m) dead only 7.34 4.70 7.69 4.01 ns

Table 5.9: Comparison of distance descriptors The mean distance of branches on the tree was not different between tree ages, but the maximum distance of live branches was higher in old trees (Table 5.9). The situations associated with these longest branches were dissimilar, however.

Several notably long branches were encountered. However, they were of different types in the different age classes. In old trees the longest branches were contender leaders heading upwards. In the 100yr trees, some branches of exceptional length were encountered with a very horizontal attitude. This is mechanically stressful. Catenary forces of leverage create incredible loads on the anchor point when the load is held horizontally, perpendicular to gravity (Smith & Padgett, 1996).

A truly exceptional branch was encountered in tree 9. An original branch 18 meters long was found held at an only 20 degrees above horizontal. A gap in the branches above is show in Figure 5.8. This length of this branch was almost half the height of the entire tree, making it an ideal candidate for snapping in the near future.

5) Crown Structure - Results

Figure 5.8: Isometric arborograph of 100yr tree 9. Arrow points to a branch 18 m long. Note gap in branches directly above the long branch. 5.3.10 Vertical plane

Vertical plane: 100yr Mean Std Dev Old Mean Std Dev p<0.05?

Mean Slope (deg) all branches 33.44 9.69 38.14 6.69 ns Mean Slope (deg) live only 33.65 9.79 39.81 6.28 ns Mean Slope (deg) dead only 30.18 12.46 35.95 10.75 ns Mean Vertical Arc (deg) all branches 23.97 10.31 30.04 17.19 ns Mean Vertical Arc (deg) live only 30.15 7.85 43.54 22.03 ns Mean Vertical Arc (deg) dead only 16.48 29.24 5.04 5.52 ns Mean Down Arc (deg) all branches 5.80 2.33 7.75 3.89 ns Mean Down Arc (deg) live only 7.51 2.83 11.34 5.40 ns Mean Down Arc (deg) dead only 2.03 1.89 1.27 1.23 ns Mean Up Arc (deg) all branches 18.17 8.92 22.29 13.94 ns Mean Up Arc (deg) live only 22.65 6.62 32.20 17.88 ns Mean Up Arc (deg) dead only 14.45 29.29 3.77 4.90 ns

Table 5.10: Comparison of vertical plane descriptors No significant differences were found in the boundary airspace along the vertical plane (Table 5.10). The mean angular slope above the horizon, mean downwards boundary angle, mean upwards boundary angle, and the mean total boundary arc were all compared between ages.

On average, the branches in the study trees were held at a 36 degree angle above horizontal. In older branches, this can be caused by repeated resprouting

5) Crown Structure - Results

upwards from parent branches with a dead tip. Contending leader megabranches in old trees were more vertical than non- contenders. This contributed to the slightly higher mean slope old trees.

In all instances, the mean up arc was higher than the mean down arc. Foliage arises from the top surface of the branch (Jacobs, 1955).

5.3.11 Horizontal plane

Horizontal plane: 100yr Mean Std Dev Old Mean Std Dev p<0.05?

Mean Horizontal Arc (deg) all branches 25.78 7.24 32.91 9.17 ns Mean Horizontal Arc (deg) live only 34.24 5.09 48.37 10.10 <.005 Mean Horizontal Arc (deg) dead only 8.84 9.11 4.69 3.76 ns

Table 5.11: Comparison of horizontal plane descriptors The horizontal plane was compared for total symmetrical angular arc only (Table 5.11). Mean left and right arcs were considered irrelevant to describing the tree. A significant higher value was exhibited in this angular spread measure in the live branches of old trees.

For dead branches, an insignificantly higher horizontal arc occurred in the 100yr trees. The dead whole branches in these trees were formerly original branches. These tend have a horizontal spread greater than young, failed epicormics, which died before attaining any lateral spread.

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5.4 Multivariate classifications

5.4.1 NMS ordinations

NMS by Structural Descriptors With SqRt Transform t16 t08 t03 t12 t10 t09

t04 t02 t05 Axis 2 Axis

t06 t13 t14 t01 t07

t11 t15

Axis 1

Figure 5.9: NMS ordination of trees by structural descriptors standardized by square root transformation 2-d stress 4.63. (McCune, 1999) A distinct organization was seen in the rendering of the placements of the 16 trees in NMS hyper-dimensional descriptor-space (Figure 5.9). Unlike techniques such as PCA, the axes shown are arbitrary and are equally scaled (McCune, 1999). Older trees (green squares) tended strongly to the upper left corner, and 100yr trees (red circles) tended to the lower right.

An unexpected outcome of this ordination is the two distinct clusters of 100yr trees (1,7,11,13,15 against 2,5,9). Observation of the arborograph lineup shows that the second group appear to have denser, larger rounder crowns than the first. These

5) Crown Structure - Results are subsequently clustered closer to the older of trees. 100yr tree 15, the smallest and subjectively the simplest, was farthest away from old trees 8, 16, and 12.

5.4.2 Cluster analysis dendrograms

Figure 5.10: Dendrogram of trees classified by descriptors performed on PC-ORD using Euclidean distances, Wards linkage method. Old trees are underlined.

Structural Variables Distance (Objective Function) 6.7E+03 2E+07 4E+07 5.9E+07 7.9E+07 Information Remaining (%) 100 75 50 25 0

t01 t07 age t11 t15 1 2 t13 t14 t02 t05 t04 t09 t06 t03 t10 t12 t08 t16

Figure 5.11: Dendrogram of trees classified by descriptors performed on PC-ORD using Euclidean distances, Wards linkage method. 100yr trees are red circles, and old trees are green squares

5) Crown Structure - Results

A cluster analysis dendrogram (Figures 5.10, 5.11) showed an initial division demarcating the old trees 16, 8, 12, 10, and 3. The sister group included all of the old trees, with the crossover exceptions of old trees 4, 6, and 14.

These three old trees were indeed different than the other old trees, and this kinship with the 100yr trees can also be seen in their NMS ordination positions. With the exception of the secondary trunk on tree 6, these trees also had shallower crowns, less whole branches, and more trunk exposed, than the other old trees. The 100yr trees 2, 5, and 9, identified as an outlying cluster in the NMS ordination, were in a cluster with old trees 4 and 6.

5.5 File Size comparison

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Microstation .PNG image outputsize in Kilobytes (software v. 05.07.00.41 Bentley Systems, Incorporated, 1998)

Layers Camera 100yr Old T-test Mean Std Dev Mean Std Dev p<.05?

down 53 5.4 76 12.1 <.005 front 54 5.5 72 8.8 <.005 isometric 59 5.9 75 8.2 <.005 right 48 4.4 62 9.2 <.005 Cones & trunk right iso 59 5.1 75 7.9 <.005 only top 64 11.0 82 16.9 <.05 up A 72 12.1 99 26.2 <.05 up B 285 38.2 367 51.5 <.005 Sum 694 82.1 907 124.1 <.005

down 84 16.7 140 48.4 <.01 front 211 32.8 279 47.1 <.005 isometric 221 42.2 286 44.8 <.01 Cones, trunk, right 76 13.7 104 23.1 <.05 kite, cross, & right iso 89 13.1 120 20.3 <.005 boundary lines top 122 31.5 154 42.5 ns up A 134 41.4 185 62.0 <.05 up B 187 42.8 256 58.7 <.05 Sum 1124 224.3 1523 302.7 <.01

Table 5.12: File sizes of arborograph images All but one combination of camera angle and image layers showed significantly higher file size for older trees (Table 5.12). The applicability of this measure to the reality of the study tree is linked to the precision in resolution level. These results indicate that the computer images representing older trees consistently contain more information than the images representing 100yr trees.

The standard deviation of file size was invariably higher in the old trees, indicating a greater variability in crown complexity as measured by file size. 5.6 Structural features

5) Crown Structure - Results

X = Present in Tree 100yr Old 100yr Old Total Total Tree # 1 2 5 7 9 11 13 15 3 4 6 8 10 12 14 16 Trough branches X X XXXX X 1 6 Elbow hollows XXX XX 14 Litter collections XX X03 Vascular epiphytes XX 02 Burl X XXX 0 4 Dead tops XXXXXXX 0 7 Snapped main stem XX02 Secondary trunk XX 11 Gallery Branch XX 11 Dead flanks XX X 0 3 Collar hollow XXXX X X 0 6 Trunk Fissures X01

Table 5.13: Presence of features in each tree

All structural features tabulated were encountered more often in old trees than 100yr trees (Table 5.13). Eight of the twelve features were only in old trees.

Trough branches: Large branches, often originals, were decayed on their upper surface, forming a canoe-shaped trough. These branches appeared healthy from below and from the side, and often continued to support several children branches and large quantities of foliage. A brown, dusty layer of stringy bark fragments was found in the hollow. Trough branches were classified as three types.

a. Dead wood forming a pit on the upper surface of living branches (100yr tree 13, old trees 3,8,16). Trough depth was less than width (Figures 5.12, 5.15, 5.16).

b. Original megabranches hollowed into a canoe shape perhaps a more advanced stage of trough type “a” (old trees 6, 10, 12, 16). Trough depth was greater than width (Figure 5.18).

c. Fragmenting dead branches, perhaps pecked by marsupials or cockatoos such as the Black Cockatoo, Calyptorhynchus funereus (Simpson et al., 1999) (old trees 3, 6, 8, 10, 12, 16) (Figures 5.13, 5.14).

Trough branches were extremely variable in shape and depth. Large canoe shaped megabranches were found only in very large, old branches. Canoe troughs encountered in tree 12 were large enough to encompass at least five average-sized human. All trough branches were orientated in such a way that they would collect

5) Crown Structure - Results rainwater (Mackowski, 1984). Types a) and b) had a positive slope and are likely to channel water into the trunk.

No fragmenting dead branch troughs were recorded in 100yr old trees, despite the presence of branches similar in size.

Figure 5.12 (left): Trough branch in 100yr tree 13 Figure 5.13 (right): Trough branch in old tree 8

Figures 5.14 (left), 5.15 (centre), 5.16 (right): Trough branches in old tree 3

Elbow hollows: When the original terminal apex of a branch died, an epicormic bud on the upper surface gained dominance and the branch grew upwards in an elbow shape. The dead apex may include a woody branch that stays attached as a stub. As the living branch grew around that stub, a collar of living wood formed around the stub with the resprout arising from the top. When the stub fell off, a circular hollow was left. These were often found in conjunction with troughs.

Similar to the troughs, elbow hollows were encountered in both old large branches and smaller epicormics. 100yr tree 13 and old trees 3, 4, 12, and 14 had elbow hollows present. In old trees 12 and 3, the elbow hollows were contiguous with a trough branch (Figure 5.18). The elbow hollow found in 100yr tree 13 seems to have formed at an earlier stage than those in the old trees (Figure 5.17). The live wood

5) Crown Structure - Results

has grown around a snapped dead branch end and formed a shallow hollow, but appears to be solid wood throughout. By contrast, those in old trees seem to have formed when the branch end of an already hollow live branch snapped off.

Figure 5.17 (left): Elbow hollow in 100yr tree 13 Figure 5.18 (right): Elbow hollow in old tree 12

Litter collections: Leaf, fruit, twig, and bark litter collected in tight branching junctions, in troughs, or in branch collar hollows. Broken “widow maker” branches hanging in the tree crown were not counted as litter. Only collections deeper than 5cm were recorded. Shallower collections occur frequently at the base of live branches.

Collections greater than 5 cm deep consisted mostly of stringy bark fibre. Other debris included twigs, leaves, fruits, charcoal, and dead wood. Litter collections were only encountered in old trees:

a. in a trough branch or elbow hollow (four old branches in tree 12, tree 8) (Figures 5.20, 5.21),

b. in a tight branching point, such as an epicormic fan (old trees 8,16) (Figure 5.23), or

c. in a collar hollow (old tree 8) (Figure 5.19).

5) Crown Structure - Results

Figure 5.19 (left): Litter collection in a collar hollow in old tree 8 Figure 5.20 (centre): Litter collections in a trough branch in old tree 8 Figure 5.21 (right): Litter collections in an trough branch in old tree 3

Vascular epiphytes: Unlike the nearby Nothofagus and Acacia trees, epiphyte cover was scarce in E. obliqua, perhaps due to constantly shedding bark (Jacobs, 1955). Vascular epiphytes were only found in trough branches and litter collections.

Two vascular epiphyte occurrences were encountered, only in trough branches in old trees. Their presence was attributed to seed deposition by birds (McQuillan, 2004). A pair of unidentified seedlings, perhaps Phyllocladus aspleniifolius (Kirkpatrick, 2004), in a trough megabranch in old tree 12. The stems were approximately 3 cm in length (Figure 5.22). A vigorous specimen of Mountain Pepper, Tasmannia lanceolata (Winteraceae) in a branching crotch litter collection in old tree 16. (Figures 5.23, 5.24)

Figure 5-22 (left): Unidentified seedlings in tree 12 Figures 5-23 (centre), 5.24 right): Tasmannia lanceolata in tree 16

Burls: Large swellings on the tree trunk formed in response to an irritation or infection. While small swellings were numerous, perhaps epicormic buds

5) Crown Structure - Results immediately before sprouting, burls larger than 0.5 metres diameter were recorded as special features.

Large burls were encountered only in old trees 3, 10, 12 and 14. (Figures 5.25, 5.26) The largest, in tree 10, was 1.4 in diameter (Figure 5.27). This burl had a hard, knobbly surface distinct from the normal bark on the others.

Figures 5.25 (left), 5.26 (centre): Burl in old tree 3, with stringy bark surface Figure 5.27 (right) Burl in old tree 10 with a distinctly knobbly surface. R. Junker placing traps.

Dead tops: The original terminal shoot of some trees was no longer living. Other branches may be actually higher than these dead tops, but originate from the side of the trunk. All old trees except tree 3 had dead tops (Figure 5.28, 5.29). No 100yr trees had dead tops. The dead top of tree 6 was the only one to retain fine branchlets, indicating a more recent dieback of the summit (Figure 5.30).

Figure 5.28 (left): Dead top in old tree 4, from below Figure 5.29 (centre): View of old tree 8, from tree 7. Figure 5.30 (right): The dead top of old tree 6 retains smaller branchlets

5) Crown Structure - Results

Snapped main stem: A distinct break in the tree trunk, leaving a shard or stub section on top, was encountered. While dead tops could lose their leader and snap off, snapped stems were large enough that one could imagine a substantial portion of the living tree crown falling with it. Unless burnt, the snapped section is nearby on the forest floor.

Two old trees had snapped main stems. In both trees a lateral branch gained an upright position and became the treetop. These upright lateral branches resembled younger trees, and were the only branches encountered which could be described as reiterations (Halle, 1995). A shard of wood was often remaining (Figure 5.32).

Snapped stems were common in surrounding trees, but were often considered too hazardous trees and rejected for study (Figure 5.31).

Figure 5.31 (left): Snapped stem on E. obliqua visible from the Airwalk. Figure 5.32 (right): The snapped stem of old tree 10 is evidenced by a large shard of dead wood. A lateral branch is now the treetop leader. CD sticky trap visible.

Secondary trunk: These arioe from the base of the tree, well below the rest of the tree crown. It is unclear if these are resprouts from the low trunk, or a overgrown neighbouring tree. As E. obliqua is not lignotuberous, it is unlikely a normal resprout (Jacobs, 1955).

5) Crown Structure - Results

Secondary trunks were present in old tree 6 and 100yr tree 7. In surrounding forests, secondary stems were observed more frequently in old trees than younger trees.

Doubt remains as to whether these stems were neighbouring trees or basal sprouts. It is possible that a swallowed neighbouring tree could have fused or interlocking roots with the older swallowing tree (Kelly, 2004).

The secondary trunk in old tree 6 was of comparable size to surrounding 100yr trees, but shorter and wider (Figures 5.33, 5.35). It had a recently sprouted epicormic crown, and leaned away from the main stem. The main trunk bends abruptly over the top of the secondary trunk (Figure 5.34). The stems split at 3m above the ground. There was evidence of decay and charcoal at the basal fusion. A bracket fungal fruiting body was present below the fusion.

The secondary trunk in tree 7 was smaller than any of the surrounding trees and was completely dead. (Figure 5.36, 5.37) The stem split at 1m above the ground.

Figure 5.33 (left): Main trunk and secondary trunk of old tree 6 Figure 5.34 (centre): The upper crown of old tree 6 main trunk bends over the secondary trunk Figure 5.35 (right): Arborograph of old tree 6, secondary trunk visible

5) Crown Structure - Results

Figure 5.36 (left):: Looking down the main trunk of 100yr tree 7. Dead secondary trunk is visible in centre of photo in front of red climbing rope (MW) Figure 5.37 (right): Arborographs of 100yr tree 7, secondary trunk visible

Gallery branch: These were branches with several openings on the upper surface of a live branch. One gallery branch was found in each age class.

A live original branch in the upper crown of 100yr tree 11 was identified as a gallery branch. A series of holes into a hollow interior approximately 10mm in diameter were in a line on the upper surface of the branch.

A gallery branch encountered in old tree 4 snapped during the course of the study (Figure 5.37, 5.38, 5.39). The branch was living and carrying a large amount of foliage. It was the lowest branch on the crown and there were no branches near it on the trunk. An ant colony (unidentified species) inhabited the complex cavities in the branch. Ants were active on the branch surface before and after the break. It is unclear if the pattern of holes in the wood was caused by fungal decay, ant colony activity, or previous insect activity such as termites. Several small holes approximately 10mm in diameter on the upper surface were observed before breakage.

Figures 5.37 (left), 5.38 (centre), 5.39 (right) The snapped gallery branch in old tree 4

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Dead flanks: A substantial proportion of the upper trunk was dead and decayed, and with dry dead wood exposed an charcoal marks surrounding. Branches from the live side wrapped around the tree to replace this side of the crown. Bark flakes overhanging the side of the dead flanks offer a habitat distinct from the sheets of stringybark that peel off of other trees. Huntsman spiders, Delena sp., and skinks, Niveoskinkus, were regularly seen sheltering in the flakes on the edge of the dead flanks.

Dead flanks were only observed in old trees. In old trees 10, and 16, a large dead flank extended downward from the stem snap to cover a portion of the upper trunk (Figures 5.39, 5.40, 5.41). In tree 16, the dead flanks was approximately 30 metres in vertical length, and occupied ~40% of the girth. This could have resulted from a strip of wood staying attached and peeling off when the upper trunk fell. In tree 12, a 4m section of trunk approximately 2m wide was exposed dead wood (Figure 5.42). Unlike the others, live wood continued above this patch.

Figures 5.39 (left), 5.40 (right): Dead flank of old tree 16, with smooth dead wood exposed

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Figure 5.41:Detail of dead flank of old tree 16, with charcoal visible. Dark portion is approximately 15 cm across.

Figure 5.42: Dead flank on old tree 12,in the upper left of the photograph. (MW)

Collar hollow: Similar to an elbow hollow, but found on the trunk. When a branch died but did not fall off, trunk wood continued to grow around the stub. After the branch fell, the live wood formed a cavity (Figure 4.42). Collar hollows may eventually heal over. While open, they maintain a vector for further decay (Hadlington & Johnston, 1988).

Several collar hollows were found in each old tree except 10 and 14. None were found in 100yr trees. Several stages were observed:

a. shard of branch retained (all old trees save 10 and 14) (Figure 5.43)

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a. without shard retained (all old trees save 10 and 14) (Figure 5.44)

b. healing over (tree 4 only) (Figure 5.45)

Different effect of original and epicormics due to different depths of anchor pathway to heart (Hadlington & Johnston, 1988).

Figure 5.43 (left): Collar hollow in old tree 4 with branch shard Figure 5.44 (centre): Collar hollow in old tree 4 without branch shard Figure 5.45 (right): Collar hollows healing over in old tree 4

Trunk fissures: Vertical fissures on the lower tree trunk offered a window into a completely hollow stem. These fissures formed “legs” upon which crown is supported. The legs were sometimes slightly splayed, creating a bulge in the trunk, perhaps due to the weight pressing directly down. Another theory for the formation of the fissures and the splayed legs is that the lower trunk is composed of more rigid wood than the upper living crown, and the fissures are caused by mechanical stress as the tree sways in the wind (Romanski 2004).

Old tree 3 was the only study tree with trunk fissures (Figure 5.46). A similar phenomenon was observed on other old trees in the region. Four vertical fissures averaging 6m long and 10cm wide opened into a hollow interior. By placing a camera through the fissure, photographs could be taken of the trees interior (Figure 5.47, 5.48). Charcoal, lichen, and spider webs were observed in the interior. Brown dust coated several of the upper spider webs.

5) Crown Structure - Results

The burl in tree 3 (Figure 5.25) was above the fissures. At the upper extent of the fissures, the trunk bulged outwards (i.e. reverse taper).

Most old trees in the vicinity exhibited basal sinuses and buttresses (Figure 5.49, 5.50, 5.51). The fissuring in tree 3 may result from mechanical stress from wind or crown weight causing the buttresses to splay.

Figure 5.46: Trunk fissure, visible in the bottom centre, in old tree 3 viewed externally

Figure 5.47 (left): Hollow interior of old tree 3 with Brendan Keyes’ hand for scale Figure 5.48 (right): Hollow interior of old tree 3 with author’s boot for scale

5) Crown Structure - Results

Figure 5.49 (left): Basal sinuses visible on old tree 8 (MW. Figures 5.50 (centre), 5.51 (right : An old E. obliqua at the study site shows a possible future for trees with fissures. The hollow interior is exposed and the entire weight of the crown is resting on three buttress legs, one of which is buckling out. The upper crown of the tree appears healthy, and from one side of the tree it appears intact save for a daylit fissure. 5.7 Other untabulated features

Large branch breakages: Two large branches from old trees were found broken on the ground between November 2002 and August 2004. One was the gallery branch in old tree 4 described above (Figure 5.37). The other was the trough branch in old tree 6.

It is likely that other branches in study trees failed but not noticed. These two megabranches were immediately apparent from the ground because of their size.

Figure 5.52 (left): Large original trough branch failure on old tree 6 Figure 5.53 (right): Lichen on dead branch in old tree 4 Lichen on dead branch: A light green lichen, perhaps an Usnea sp., was frequently encountered on the upper surface of dead branches (Figure 5.53). The individual tufts were about 20mm tall.

5) Crown Structure - Results

Branch base indentation: An indentation indicating branch weakness was observed at the base of a living branch in old tree 4 (Figure 5.54). The discontinuity between bark pattern on the trunk and on the branch suggests that this branch is on the verge of breaking.

Figure 5.54 (left): Branch base indentation in old tree 4. Figures 5.55 (centre), 5.56 (right): Birds nest in lowest branch of old tree 8

Bird nest: An unoccupied bird’s nest was observed in the lowest dead branch of old tree 8 (Figures 5.55, 5.56)

Berry-like foliage gall: A red and yellow gall resembling a berry approximately 5 mm in diameter was found attached to a leaf in the top of 100yr tree 5 (Figure 5.57). When cut open, it appeared to be filled with white material, but was unoccupied (Figure 5.58).

Figure 5.57 (left), 5.58 (centre): Berry-like foliage gall in tree 5

5) Crown Structure - Results

Figure 5.59 (right): Scribbly tracks in old tree 3

Scribble tracks: Symmetrical markings tracks were observed on a live gum- barked branch in the upper crown of old tree 3 (Figure 5.59). This was attributed to the larvae of an Yponomeutid scribbler moth (CSIRO, 1967). No specimens of this taxon were found in the arthropod trapping.

Bark strip flakes: Long, flaky strips of bark up to 4 m in length were frequently encountered hanging along the trunk (Figure 5.60). Unlike the long, sturdy, bottom-anchored ribbons found hanging in the gummy E. regnans (Jacobs, 1955) , the stringy bark strips were loosely constructed. Bark strips were found hanging as a loose flake anchored at the top, or as smaller flakes anchored at the bottom. Large hanging strips were fragile and often fell off when disturbed.

Large hanging strips were easily created artificially when investigating the bark. By digging one’s fingers under the bark, a vertical flake could be pulled off to reveal animals living beneath. Large social huntsman spiders, Delena sp., were often found living beneath these flakes (Figure 5.61).

5) Crown Structure - Results

Figure 5.60 (left): Short bark strips visible above R. Junker along the trunk in 100yr tree 5. Figure 5.61 (right): Seven-legged Delena sp. huntsman spider in old tree 16. Blue rope is 9 mm in diameter.

Flaky armpit bark: Stringy bark was often seen forming flaky bridges between the branch and trunk on the underside of branches (Figure 5.62). They were not sampled for arthropods, but represent a habitat found only in stringy barked Eucalyptus.

Figure 5.62 (left): Flaky armpit bark in 100yr tree 2 Figure 5.63 (right): Branch armpit ridging visible in old tree 4, with R. Junker.

Branch armpit ridging: A pattern of parallel ridges was observed in many branches with a diameter greater than 15 cm (Figure 5.63). This may be an evolved mechanism to support the weight of heavy branches, or may be a deformation caused as a branch gains mass and dips downwards.

6) Crown Structure - Discussion

6) Crown Structure – Discussion

The crowns of the old trees were found to be quantifiably different from the 100yr trees. The old trees showed more variability in structure. The dynamics of intra- crown competition between branches is discussed. All structural features were more common in the older trees, following the literature that indicates time is required for their formation. The measure of complexity is discussed in regards to studying crown structural complexity. 6.1 Distinctness of crowns

Ha: Old trees are quantifiably and objectively distinct from 100yr trees, i.e. old trees are not scaled-up versions of 100yr trees.

Ho: Old E. obliqua trees differ only in size to 100yr trees.

6.1.1 Summary of evidence

Several descriptor variables show quantitative, significant distinctions between the tree ages. Descriptors of scaling ratios, proportion of gum bark, horizontal arcs, ratio of crown polyhedra: trunk wood volume, and proportional mean branch height highlight differences in crown structure beyond that of scale.

Other allometric relationships can be viewed in the results. For example, the ratio of tree height between 100yr and old trees was about 1:1.3, while the ratio of wood volume was about 1:7. This clearly shows a strong difference in allometry between the age classes. 100yr trees were slender compared to the thick old trees.

Structural features found in old trees were absent from most 100yr trees. A scaled up version of the 100yr trees would not have these features.

Old trees showed either a secondary epicormic crown, indicating a crown fire, or a senescing primary crown. These trees were distinguished in the descriptor dendrogram and NMS analyses (Figures 5.9, 5.10).

The evidence supports the rejection of the null hypothesis that old E. obliqua trees differ only in size to 100yr trees.

6.1.2 Crown developmental stages of E. obliqua in wet sclerophyll forests

6) Crown Structure - Discussion

The sixteen study trees represented the mature, primary, and secondary crowns types described by Jacobs (1955). All study trees had at one point been vigourous young trees, and all old trees had at one point been early mature trees (Figure 6.1).

The 100yr trees were all early, dominant mature trees. The old trees were either trees with epicormic crowns that had survived a crown fire, or mature trees with senescing primary crowns.

Figure 6.1: Crown structural development in wet sclerophyll E. obliqua. All trees eventually die. All eight of the 100yr study trees had primary crowns. Three old study trees had experienced a crown burn. Five old study trees retained primary crowns in various stages of senescence. A crown fire changes the crown structural path by killing the original branches of the primary crown and triggering a secondary epicormic crown. The trunk may snap. If a crown fire does not occur, original branches will die over time as crown senescence progresses. Note that the tree can die at any stage.. Trees end up as logs on the forest floor or are harvested.

All 100yr study trees were mature dominant trees: The 100yr age cohort showed no signs of crown scorch or crown senescence. The primary crown was still present. The upper crown was healthy, and the top had not yet died. Branch thinning

6) Crown Structure - Discussion on the lower crown was responsible for the presence of dead branches, rather than senescence. Most epicormic activity was on the branches, rather than from the trunk.

The differences in crown structure were caused by intra-cohort variability in vigour. While these trees were dominant over others of their age class, they still competed with trees on higher ground and the taller old trees.

The arborographs of the 100yr trees displayed rounded, live crowns (Figure 6.2, 6.3). The crown was significantly shallower than in the old trees. The uppermost branches were living and competed with the lower branches. The lower original branches longer than the higher branches and at a lower angle (Ishii & McDowell, 2001; Jacobs, 1955).

Small branches from the lower trunk were not present. The tree was investing its resources in vertical height. In its lifetime, the trunk has been shaded. No gaps had formed from trees falling, and the tree had been adjacent to fire in which the understorey was cleared. If these events did occur, epicormic branches in the lower crown would be present (Mifsud, 2003).

Figures 6.2 (left), 6.3 (right): 100yr trees 5 (left) & 9 (right) shows rounded primary crowns and clean lower trunks with no epicormic activity.

6) Crown Structure - Discussion

Features such as dead tops, trough branches, elbow hollows, and dead flanks are symptoms of crown damage caused by senescence or fire. These features were uncommon or absent in the 100yr trees.

Crown Fire event: A divergence in crown structure development is caused by a crown fire event. If it is not fatal, a crown fire will kill the original branches and trigger a secondary crown of resprout (Figure 6.4).

Figure 6.4. After loss of foliage and original branches of the primary crown to a fire, dormant epicormic buds are released. From Figure 44c in Jacobs (1955)

Old trees that experienced crown fire had a secondary epicormic crown: Trees 8, 10, and 16 showed evidence of a crown burn in the dense epicormic crowns, lack of original branches and low epicormic branches. Trees 10 and 16 also have dead flanks and snapped trunks.

Trees 8, 10, and 16 all showed deep, dense epicormic crowns (Figure 6.5, 6.6, 6.7). In comparison to the other old trees, branches were dense along the trunk. Original branches were mostly absent. Large epicormic branches dominated smaller epicormic branches.

Trees 10 and 16 were significantly shorter than the other old trees. Their trunks have snapped and the uppermost foliage was from an epicormic branch. Epicormic branches wrapped around the trunk over a dead flank to access light.

6) Crown Structure - Discussion

Figures 6.5 (left), 6.6 (centre), 6.7 (right): After the trunk snap in old tree 10, a vertical epicormic branch contends for leadership with a much larger branch growing at a slope.

Tree 10 and 12 both possessed dead flanks and snapped trunks (Figure 6.8). The dead flanks were covered in charcoal and signalled massive damage to the tree crown. Tree 8 may have burnt more recently, or suffered less damage in the crown fire. The top was dead. A dead flank was not observed and the trunk had not snapped.

Dead flank Dead flank

Old tree 10 Old tree 16

Figure 6.8: The dead flanks on trees 10 and 16 were evidence of massive crown damage during a fire

6) Crown Structure - Discussion

Old trees that did not experience crown fire had senescent primary crowns: Old trees 3, 4, 6, 12, and 14 retained primary crowns in varying degrees of senescence. The progression of senescence through time may be accelerated by stress on the tree.

Judging by the degree of resprouting and the number of original branches remaining, an assessment of the trees’ decreasing vigour and/or increasing age may be formed. Trees 6, 14, 3, 12, and 4 show declining abundances and condition of original branches, and an increasing amount of epicormic activity on the trunk of on the branches (Figures 6.9, 6.10).

The absence of a crown fire is shown in the arborographs by a taller, sparser crown than that of the secondary crowned old trees. Epicormic activity on the lower trunk is less pronounced. Original branches are retained with epicormic foliage on the upper surface (Figure 6.10).

Figures 6.9 (left), 6.10 (centre), 6.11 (right): In the upper crown of old tree 4 several large original branches remain. Epicormic branches arise from the upper surfaces. Damien Catchpole descending past large original branches in the upper crown of old tree 4.

Most of these trees had dead tops and recent collar hollows where original branches were failing. None of these trees had dead flanks or snapped tops. These trees appear to have experienced burning at some stage, but not of sufficient intensity

6) Crown Structure - Discussion

to reach the crown. The charcoal in the hollow interior of tree 3 suggests interior burning that the tree somehow survived. A crown fire could still alter the crown structure of these trees and trigger a secondary crown.

6.1.3 Crown development as competition between cohorts

100yr trees follow Jacobs (1955) sapling model: The 100yr old trees in the present study show a similar crown structure and dynamic to Jacobs’ (1955) illustration of branch competition in a Eucalyptus sapling (Figures 2.2, 6.12). Curtin (1970) discussed the “nonplasticity” of E. obliqua after studying the allometry of trees between 10 and 93 years of age. Similarly, the 100yr old trees in the study appear to be scaled up versions of younger trees.

6) Crown Structure - Discussion

Figure 6.12: Crown of 100yr tree 1

Older branches compete with younger branches by being lower and longer: Branches within a tree compete by producing auxin (Raven et al., 1986). This hormonal growth suppressant inhibits the growth of other buds lower in the tree or branch. Vigorous branches will suppress other branches to a greater extent than decadent branches.

In the dominant 100yr trees, the younger upper branches were more vertical than the lower older branches. Unlike the tapered, bottom heavy crown of P. menziesii, the younger, vigorous branches appeared to dominate over the older branches beneath. As the older branches adopted a lower angle, they moved out from

6) Crown Structure - Discussion under the influence of the upper branches. When competing upwards for light, the younger branches have the advantage of a higher origin point. The additional length of the older branches allows them to gain a high position at an angle away from the youngest branches. Foliage grows upwards from the branch surfaces towards the light. The rounded crown shape of the 100yr trees forms from this phenomenon of longer, older branches lowering from vertical. Jacobs (1955) attributes this to the increased weight of the older branches

Older, lower branches were sometimes of sufficient length or in suitable circumstances to retain an upright attitude and compete with the younger branches of the upper crown (Figure 6.13, 6.14).

Figures 6.13 (left), 6.14 (right): In 100yr tree 2, a lower branch visible on the leftside is competing with the upper crown. Its length and steep slope allow it to compete with younger branches with a higher anchor point.

By contrast, older branches in Douglas-fir compete by being longer: In P. menziesji, most branches maintain a horizontal, geotropic attitude throughout life

6) Crown Structure - Discussion

(Ishii & McDowell, 2001; Van Pelt et al., 2004). Older branches compete with the upper branches by virtue of their longer axes which bring foliage out from under the shading of the upper tree (Figure 2.3). Reiteration of branches in these coniferous trees may be an attempt by the older branches to exploit any advantage they may have over the younger branches above.

The original branches lowest in the primary crown are the oldest and the least competitive: In Jacobs’ (1955) illustration of crown growth in Eucalyptus (Figure 2.2), the lower dead crown in a sapling encompasses the oldest branches at four years of age. Similarly, in the 100yr study trees, the oldest branches remaining on the tree are in the lower crown (Figures 6.15, 6.16). Even if they had been exceptionally vigorous in the past, young branches were in a stronger position by virtue of their height. These branches were rarely observed at their length at death. Lower dead branches had usually snapped and remained as shards. Dead branches were rare in the upper crown of 100yr trees. This suggests that most branch death in this age class was due to the interlinked effects of old age and the lack of competitiveness in the lower crown. The visible effects of mortality from herbivory may be confined to branchlets and buds beneath the resolution level (Lowman, 1987; Mopper et al., 1991).

Figures 6.15 (left), 6.16 (right): The lower dead crown in 100yr tree 1 contains the oldest branches in the tree. Some are retained only as stubs.

Competition of branches in old trees: In the old E. obliqua, branches have a lifespan shorter than that of the tree. The analogy of branches in the crown to trees in a forest can be extended to the old trees. The old trees in the study are of sufficient age that most original branches are dead or dying. The catastrophic loss of branches to

6) Crown Structure - Discussion

a crown fire or the gradual loss of branches to age releases epicormic buds from dormancy (Jacobs, 1955). Similarly, in forest systems, shade-intolerant seedlings cannot survive unless a catastrophic disturbance or the death of an established dominant tree releases them from suppression by established trees.

A crown fire is analogous to a stand level fire: Crown fires, such as those that affected old trees 8, 10, and 16, can cause complete or partial mortality of the original branches. A complete crown burn, like the E. regnans forest in Wallaby Creek, Victoria, generates an even-aged cohort of branches on the tree (Ashton & Chinner, 1999). There may be some surviving original branches. If the foliage was scorched, then epicormic buds will be released along the branch surface. These epicormic buds are analogous to seedlings.

Ishii et al. (2002) notes the lack of increased epicormic activity in the harsh conditions of the upper crowns of P. menziesii. They infer that epicormic production in these trees is a constant process independent of external stimuli. By contrast, epicormic activity in E. obliqua is caused by external factors such as fires and crown damage, and by internal factors such as the loss of the tree top and branches to decay (Jacobs, 1955).

Sillett & Van Pelt (2000) records the structure of reiterated secondary trunk in the crown of ancient redwood tree that survived a crown fire. Due to the long time period since the fire, it is unclear if these reiterations arose from surviving original branches, or epicormic resprouts.

Epicormic buds compete to survive: Not all of these epicormic branches will survive. Like the competitive exclusion phase of stand development following a disturbance such as harvest or wildlife, density-dependant mortality results in the loss of less vigorous epicormic branches (Franklin et al., 2002). An epicormic branch in a lower position is generally less competitive than one above it. The upper branch shades the lower one and is in a stronger position to compete with neighbouring trees. The upper branch produces auxin that will suppress the lower branches (Jacobs, 1955) (Figures 6.17, 6.18).

Franklin et al. (2002) describes a stage of rapid biomass accumulation concurrent with competitive exclusion. Similarly, epicormic branches that suppress or

6) Crown Structure - Discussion outlive other branches will have less competition for the resources and will grow quickly.

Figure 6.17 (left): Low epicormic branches in old tree 6 have failed when higher ones have thrived. Figure 6.18 (right): The end result of competition between epicormic resprouts is visible in this arborograph of old tree 16.On the left side of the picture, several small dead epicormic stubs (grey) have been overcome by the large branch above (brown).

In absence of crown fire, branches will die of old age: Epicormic branches can also be released by the death of original branches from decay, mechanical disturbance, or herbivory (Jacobs, 1955). The primary crowns of the old E. obliqua are in dieback (Figure 6.19). The tree is no longer gaining height and is commercially overmature. The top is usually dead.

In a microcosm of the entire tree, original branches decay over time (Mackowski, 1984). The dead tip of a branch is analogous to the dead top of the entire trunk. When the top of a E. obliqua dies, the lower branches released from suppression contend for leadership.

The death of the branch tip can trigger the release of previously dominated epicormic buds on the upper surface of the branch (Jacobs, 1955). These branches will contend for domination of the entire branch. Elbow hollows can result when the branch end falls off (Mackowski, 1984).

6) Crown Structure - Discussion

Figure 6.19: The senescing upper crown of tree 3 has several original branches with epicormic activity on their upper surface.

Epicormic branches on the trunk are released by the death of originals: Epicormic branches on the trunk can result from decay or damage to the upper crown. Through time, the primary crown of E. obliqua senesces. When the original branches fail, dormant buds are released from their suppression and can replace the lost branches. Jacobs (1955) notes that they are not as efficient as the originals they replace, and that several cohorts of epicormic branches may sprout and die before the death of the entire tree. Contender branches may have arisen from epicormic buds, rather than original branches.

In P. menziesii, epicormic branches sustain foliage in the lower crown, where original branches are in decline (Ishii & Ford, 2001) (Figure 2-3).

Epicormics on the lower trunk can form megabranches: Epicormic branches on the lower trunk may be competitive because of favourable conditions

6) Crown Structure - Discussion surrounding the tree (Figures 6.20, 6.21, 6.22, 6.23). In old trees 3, 8, 12, and 16, megabranches arose from the lower trunk and were the largest branches on the tree.

Mifsud (2003) reports on the bursting of epicormic buds on wet sclerophyllous Eucalyptus in response to the opening of gaps as nearby trees fall. Jacobs (1955) (Figure 6.4) illustrates epicormic branches low on the trunk in response to fire.

All old study trees survived the fire events of 1898, although some may have experienced a crown burn . The fire would have caused a decrease in the abundance of neighbouring trees. Fire damage to the trunk, combined with the increased light levels, may trigger the growth of epicormic branches which were in the proper conditions to grow to tremendous sizes.

Competition from other branches would be minimal the middle trunk below the crown. In this zone, low original branches had been occluded in the past. The future megabranch would be able to grow steeply upwards without competition from established originals. If fire damage did not extend further up the trunk, the epicormic activity above would be absent.

Figures 6.20 (left), 6.21 (right): A megabranch low on the trunk of old tree 12 (at 13m) may have originated immediately after damage from a fire that did not reach the tree crown. Large

6) Crown Structure - Discussion

original megabranches can compete with branches in the upper crown. Competition obstacles for a newly sprouted epicormic on the lower crown would have been minimal after a fire. Original branches in the zone immediately above would have been self-pruned in the past, and neighbouring trees would be less abundant. There are no branches on the trunk in the shade of this megabranch. M. Cracknell shown placing traps.

Figures 6.22 (left), 6.23 (right): A large, low epicormic megabranch in tree 14 at 22m. R. Junker shown in the tree crown. Note the lack of epicormic branches in the zone shaded by the large branch. The next branch upwards on the trunk (31m), seen immediately above the climber does not compete with the low megabranch but bifurcates and invests in another location. The megabranch is probably the older of the two. 6.2 Variability in tree crowns

Ha: Old trees have a more variable crown structure than 100yr E. obliqua.

Ho: No difference exists in variability of crown structure between 100yr and old E. obliqua.

6.2.1 Summary of evidence

Observations in the field strongly indicated the higher variability of tree crowns in older E. obliqua.

Visual assessment of the arborograph lineups (Figures 5.1, 5.2) showed a higher variability in the crown structure of old trees. As the most extreme example,

6) Crown Structure - Discussion

old trees 4 and 6 are visually distinct from the 100yr trees and the other old trees. By contrast, the 100yr old trees show a smaller range of size and shape.

NMS ordinations (Figure 5.9) showed 2 distinct clusters of 100yr trees against a more evenly spread of old trees. However, there was a relatively similar spread of trees overall in both age categories.

PNG file size analysis (Table 5.12) showed a consistently higher standard deviation for the older trees. The amount of information, used as a surrogate for complexity, required to display the arborographs was therefore less variable for the 100yr trees.

The occurrence of structural features was much higher in the old trees. Each individual feature encountered was unique. When the higher rate of incidence is combined with a presence of a distinction bias adds further evidence that old trees are more variable in structure.

The discussion of intra-crown inter-branch competition highlights the regular pattern of branch growth in 100yr trees and the disruption of this pattern by stochastic events such as crown burn and branch senescence. All of the 100yr trees were identified as mature primary crowned Eucalyptus, whereas old trees were divided into primary- and secondary- crowned trees

The evidence supports the rejection of the null hypothesis that no difference exists in variability of crown structure between 100yr and old E. obliqua.

6.2.2 Comparison with previous research

The findings of the present study support the contention of Richardson et al. (2001) that older trees have a higher plasticity than younger trees. \Plasticity is meant in this context as the range of expression in physical form, rather than as the ability to tolerate change. Richardson et al. (2001) showed that within the crown of older Tsuga heterophylla, the range of leaf shape expression is greater than the younger trees. The present findings show that this holds true for the branching structure of E. obliqua.

Life experiences change trees and create unique, idiosyncratic crown structures (Halle, 1995; Ishii & McDowell, 2001; Sillett & Van Pelt, 2000). Young organisms faithfully express their genetic growth pattern, or architectural model. As time progresses, deviations from that model accumulate and exacerbate. Small

6) Crown Structure - Discussion

deviations can magnify. Older E. obliqua, like older people, older cars, and older buildings, have had a longer time to collect injuries and experiences. Those injuries and experiences continue to create deviations in the architectural model over time. Green (2002) and Fay (2002) discuss the arboriculture industry’s recognition of an ancient tree’s accumulation of experience.

The old study trees have had about 300 years more experience than the 100yr trees (Hickey et al., 1998). Fires in the crown caused the most apparent changes in the studied E. obliqua tree crowns (Wilkinson & Jennins, 1993). Experiences such as herbivory (Mopper et al., 1991), wind damage (Mifsud, 2003)) and fungal infection (May & Simpson, 1997; Wardlaw & Neilsen, 1999) impact each tree in a unique way.

A potential confounding factor is the increased range in age of the old trees. Using the fire history of Hickey et al. (1998), the ages determined for the 100yr trees is more precise than the lumping of the old trees into one age class. Differences in crown structure of 100yr tree are probably caused by environmental and competition factors rather than age. The old trees are all likely older than 300 years, and younger than 500 years, but determining their age is impossible (Hickey et al., 1998). The increased range of crown structures in the old trees is interlinked with the increased range of tree ages.

Like disturbances or competition in a forest stand (Franklin et al., 2002), branch mortality can change the population of branches in the tree crown. Ishii & Wilson (2001) discuss epicormic activity and branch death in different strata of the crown. Sprouting of epicormics and the death of original branches, in analogy with a forest stand, leads to an increase in heterogeneity of horizontal and vertical structure, albeit rotated ninety degrees (Brokaw & Lent, 1999; Song et al., 1997; Van Pelt & Nadkarni, 2004).

The old trees in the study had been present for three centuries before the 100yr trees. They had experienced the drastically different conditions following the fires in 1898 (Hickey et al., 1998). During this time period a difference collection of stresses acted upon the surviving trees (Figure 6.24). These stresses have never been experienced by the younger trees that germinated after that period. With the100yr cohort, the trees survived the gradual changes of the post-burn environment into the rainforest understorey seen today.

6) Crown Structure - Discussion

The old trees show plasticity in their observed crown structure. The evidence collected on the old E. obliqua in this project strongly contradicts his findings. The crown structure of the old trees shows strong differences from both the 100yr trees and from each other. The plasticity in structure of the old trees is apparent in different patterns of epicormic activity (Ishii et al., 2000), different levels of retention of original branches (Jacobs, 1955), and different incidences of structural features (Gibbons & Lindenmayer, 2002). Similarly, Clement et al. (2001) discuss the variability in the open, asymmetrical crowns of ancient Fitzroya cupressoides in comparison to the conical crowns of younger specimens. Sillett (1999) details the variation in crown structure of eight old-growth Sequoia sempervirens.

Figure 6.24: Eucalyptus globulus growing amidst coastal shrublands at Point Reyes, California, has been able to express its architectural model without disturbance from native insect or fungal stresses. The dead branch visible in the foreground illustrates how this vigour may be more than the tree can mechanically handle. It is likely that these exotic Eucalypts would show less variability in crown structure than they would in Australia. Hollows may never form in the absence of co-evolved agents such as fungus and termites. Note the clean healing of occluded branch visible in the front right and far left of the photo. Although other environmental factors will impact them, the lack of these stresses means fewer deviations from the model. Eric and Gary shown climbing ~2m from the ground.

6) Crown Structure - Discussion

6.3 Incidence of structural features

Ha: Structural features such as epiphytes, hollows, dead tops, snapped trunks and burls are more commonly present in old E. obliqua than 100yr E. obliqua.

Ho: Structural features are equally common in 100yr and old E. obliqua.

6.3.1 Summary of evidence

All structural features recorded were present in more old trees than 100yr trees (Table 5.13). Any and all of the old trees had more structural features than the 100yr trees.

The recording of structural features was subject to both recognition and observational biases. The four incidences recorded in 100yr trees (a trough branch, elbow hollow, secondary trunk, and gallery branch) all appeared to originate from different processes than those same features recorded in old trees. This also evidences of a distinction bias.

The evidence supports the rejection of the null hypothesis that structural features are equally common in 100yr and old E. obliqua.

6.3.2 Comparison with previous research

The higher numbers of structural features in the old E. obliqua is a product of the increased variability in the older crowns, and the distinct patterns of crown structure that are not present in the 100yr trees.

The presence of these features is due to more than just the increased size of the old trees. Increased age and decreased vigour foster the formation of several structural features. Subdominant trees may resemble older trees because of their decreased vigour (Hickey, personal communication).

The death of the uppermost portions appear to be an invariable feature of age in E. obliqua. Dead tops and trunk snaps of Eucalyptus may result from decay following crown fire, or from wind damage (Mifsud, 2003). The old trees have

6) Crown Structure - Discussion probably survived more storms than the younger trees. Dead tops can also be symptomatic of stress and internal decay unrelated to fire or wind. The dead flanks in old trees 10 and 16 were undoubtedly caused by a crown fire.

Tree hollows develop with time (Gibbons & Lindenmayer, 2002; Gibbons & Lindenmayer, 1996; Mackowski, 1984). The process of decay in trunk and branches is gradual and the 100yr trees are not old enough to form them. Whitford (2002) reports that hollows in Eucalyptus are bigger in more senescent trees. Also, trees may need to be of sufficient size to enclose hollows. Lindenmayer et al. (1993) reports that taller, larger Eucalyptus have more hollow branches than smaller ones.

The collar and elbow hollows form when the entire, or the end of a branch breaks (Gibbons & Lindenmayer, 2002). In the older trees, more branches have grown and failed than in the 100yr trees. Furthermore, these branches are usually in worse condition.

Trough branches and litter collections were often found in decaying original branches, or large old epicormics. The branches in the 100yr tree were often not of sufficient size to create a canoe trough. Litter collections were sometimes found in tight branching points where several epicormic branches were found. Sillett (1999) reports on the collection of debris at the base of a dense collection of reiterated trunks in an ancient Sequoia sempervirens. Similarly, dense branches in Eucalyptus foster the collection of litter.

The rare presence of vascular epiphytes was dependent on an appropriate litter collection to support it. They were probably deposited through bird deposition. Vascular epiphytes appear to be completely absent from the crown of the younger trees. Sillett (1999) links the presence of ericacious shrubs in a redwood crown to wood decay. Unlike epiphyte communities in tropical forests (Benzing, 1995; Hietz, 1997), vascular epiphytes in Eucalyptus trees are not canopy specialists, but appear to be chance incidences of understorey trees.

The large burls found in old trees were not found in the 100yr trees. The diameter of the hemispherical burl in old tree 10 was of comparable size to the basal trunk diameter of the 100yr trees. No burls of any size were noted on the trunk of the 100yr trees. Burls may form over time as a response to wounds or infections, and the 100yr trees may not be old enough.

6) Crown Structure - Discussion

The gallery branches and secondary trunks were likely to have been lumped in error due to a distinction bias. The formation of the two incidences of gallery branches is unclear. Further mystery remains in determining whether the secondary trunks are basal epicormic resprouts, or neighbouring trees being swallowed.

The trunk fissures of old tree 3 appear to be an advanced symptom of senescence and trunk hollowing. A similar feature was observed in a nearby old tree. Nothing resembling this feature was found in any younger trees. 6.4 More complexity of information

Ha: Old trees will have a greater complexity than 100yr trees, as measured in surrogate by the amount of information required to describe them.

Ho: Virtual 3-d models of 100yr and old E. obliqua do not differ in complexity, as measured by the amount of information required to describe them.

6.4.1 Summary of evidence

The mean file size of the arborographs was significantly higher in old trees for all but one of 15 camera angles. The accuracy of this measure was subject to the faithfulness in resolution when branches were measured in the field (Table 5-12).

A higher incidence of structural features was found in the old trees. Each feature requires more information to describe, and therefore add complexity to the trees (Table 5-13).

The evidence supports rejection of the null hypothesis that 100yr and old E. obliqua do not differ in complexity, as measured by the amount of information required to describe them.

6.4.2 Comparison with previous research

Structural complexity measured by information: The quantification of complexity is a difficult task. Loehle (2004) specifically identifies the measurement of geometric complexity of tree crowns as an outstanding challenge. Mathematical measures of complexity often approach the subject by measuring the information required to describe a system (Anand & Orloci, 1996). Roche (2001) appeals for

6) Crown Structure - Discussion

consideration of biocomplexity, a measure beyond mathematical summaries of communities.

Sillett & Van Pelt (2000a) describe the crown of an ancient Sequoia tree as potentially the most complex on Earth. They present a crown diagram illustrating the several generations of reiterated secondary trunks and the numerous fusions of wood in the crown. This diagram can be compared to those of other, simpler redwood trees (Sillett, 1999). Several additional measures of its higher complexity can be generated. More ink was used to illustrate its crown, and more words were used to discuss it. Sumida (1995) generates arborographs of trees using interlinked lines. The number and length of these lines can be used as a measure of complexity.

Complexity is more than just branches: The structural complexity in the crown of a tree may be more than measures of branch structures. The incidence of structural features adds complexity. Each attribute may contribute to the overall complexity more than a given branch. Live branches can be relatively similar, whereas tree hollows, bark flakes, and are individually distinct and have their own ontogeny.

Composite measures used as surrogate for overall complexity: Sillett & Bailey (2003) used a composite measure for structural complexity. Several features (e.g. volume of main trunk, number of branches, number of reiterated trunks) were delegated to represent complexity. They were combined using Principal Components Analysis (PCA) (McCune & Grace, 2002). The first PCA axis, representing most of the variation between the trees, was delegated as a surrogate for “overall complexity.” This composite measure allows for several contributing variables. A subjective bias exists in the choice of variables, and a decision is made that these variables contribute equally to complexity (Sillett & Bailey, 2003).

PNG file analysis allows: No other use of this measure is known. Computer visualization images are easily quantifiable using computer operating systems. When controlled for software and formatting variables, computer images provide a simple index of visual complexity.

Other tree maps could be analysed simply in this methods. The chronosequence of the P. menziesii illustrations from Van Pelt & Nadkarni (2004)

6) Crown Structure - Discussion were analysed for information content (Figure 6.25). The computer bitmap graphic of the chronosequence was cropped for each age class and converted to PNG graphics files. While PNG vector-based graphics are not perfectly suited for analysing ink drawings, they offer a standardized method. Like the present study, older trees show a trend to have larger file sizes. This measure is linked to the amount of ink used in the drawing, and perhaps to the number of measurements made in the structural measurements used as a drawing framework. The height of each tree illustration predicts the file size.

90

80

70

60

50

40

30 Height in metres 20

5010

50y 100 157 280 500 500 650 950 Age in years 10kb 17 21 27 24 33 59 31 PNG computer file size in kilobytes

Portraits of Pseudotsuga menziesii (Pinaceae) in Washington State, USA. Hand drawn by Robert Van Pelt using in situ structural measurements. All graphics and data used for PNG generation are from Van Pelt and Nadkarni (2004).

Figure 6.25 (left): Portraits of Pseudotsuga menziesii from Van Pelt & Nadkarni (2004). Hand drawings by Robert Van Pelt.

6) Crown Structure - Discussion

Figure 6.26 (right): Relationship of graphic computer file size of P. menziesii portrait plotted against increasing tree age. Note that the height of the tree predicts file size.

7) Canopy Arthropods - Methods

7) Canopy Arthropods - Methods

In this chapter, the traps collection and analysis of the canopy arthropods are detailed. The canopy arthropod biodiversity of the sixteen study E. obliqua trees was sampled using traps of three designs. Active collecting methods, such as fogging, branch clipping, and foliage beating, were rejected because of the unpredictable time investment required in accessing each tree. Mobile, rather than sessile, arthropods were targeted. Placing traps allowed flexibility for inevitable complications, as the trapping time (~60 days) was much greater than the placement time (~7 days). Analysis compared seven aspects of biodiversity between ages. The aspects were abundance, richness, community structure, species accumulation, diversity, distinctness, and individual taxon responses. The concept of scaling arthropods to tree size is discussed in detail to justify the rejection of scaling methods. 7.1 Trap design types

Fourteen traps were placed within the crown of each tree (Figure 7.1). Traps were active in January-March 2004.Three trap types were selected for their relative advantages: broad collection of taxa, low cost, durability in the conditions of the treetops, and portability.

CD sticky traps were placed on the trunk in the lower and upper crown to target local fliers, and animals intending to land on or jump along the trunk (Bickel, 2003). The trap collection was inferred to represent the community associated with the airspace close to the trunk.

Funnel crawl traps were placed on a live branch, dead branch, and trunk position in the upper and lower crown. Branch traps were positioned horizontally to capture animals moving inwards toward the trunk, and trunk traps were positioned vertically for animals heading upwards. The trap collection was inferred to represent the bark associated fauna.

Three hangtraps, omni-directional flight intercept traps were hung from branches on a cord approximately 2 metres from the trunk, in the upper, lower, and middle crown. Each trap had a top and a bottom collecting bottle filled with ethylene glycol. The trap collection was inferred to represent the community in the airspace within the tree crown.

7) Canopy Arthropods - Methods

Figure 7.1: Illustration of generalized trapping positions

7) Canopy Arthropods - Methods

7.1.1 Sticky CD traps

Goals : Sticky traps are relatively inexpensive and simple to construct, but are messy and generally yield poor quality specimens (Basset et al., 1997). Sticky traps target animals landing on tree surfaces (Figure 7.2), as do the flight intercept traps used by Majer et al. (2003) (Majer et al., 2003). A compact casing was required to carry a surface painted with sticky glue into the tree crown without allowing the glue to come in contact with other objects, while also allowing protection of the collected specimens during transport to the ground.

Figure 7.2: Arthropods susceptible to sticky traps, from Bickel & Tasker (2004)

7) Canopy Arthropods - Methods

Construction: A standard compact disk case served as both the sticky surface and carrying case (Figure 7.3). When the interior CD mounting crown is removed, they form a transparent plastic box 125mm x 140mm x 10mm. The inside front panel was coated with Tanglefoot ® glue to form a trapping area of 17,500 sq mm at a cost of ~$0.05 (Australian). CD cases are available from office or music suppliers at about $0.35 each. Note that most of the CD cases currently produced are the “slimline” variety, in which the mounting crown and the back panel are a single piece. These are not well suited for sticky traps, as there is very little space in the enclosed box.

A soldering iron was used to melt a hole through the centre of the trap (Figure 7.4), and a nail ($0.01) or cord (variable price) was used to mount to the tree trunk or branches. Construction of each trap took about 3 minutes.

Figure 7.3 (left): Components for CD sticky traps: Capture glue, CD case with coated half surface protected by uncoated half, mounting crowns to discard or reuse. Figure 7.4 (right): R. Junker melting anchor hole through CD cases with soldering iron

Trap Setting : The coated cases were carried into the tree crown, and handled by first labelling the outside of the sticky half, opening the lid and nailing the trap to the desired location. Trap labelling was done in situ to avoid complications if traps were dropped. The uncoated half of the case was retained to use during retrieval, in order to protect the fragile animals and to isolate the sticky surface. Placement of each trap took about 1 minute.

7) Canopy Arthropods - Methods

Traps could be placed in any accessible position. (Figure 7.5) Traps in foliage can be tied into position with string, and could also be placed in hollows or any other feature of interest, facing in any direction.

The collection results presented are from one upper and one lower trap mounted directly on the tree trunk for 60 days (Figure 7.6). Traps were also placed and collected on two live branches, two dead branches, and in foliage, but were not analysed due to time constraints. Almost a quarter (60 of 256) of the exposed traps failed, probably due to strong winds.

Figure 7.5 (left): CD sticky trap on dead branch in old tree 4 Figure 7.6 (right): CD sticky traps on trunk in 100yr tree 7 . A funnel crawl trap is partially visible.

Retrieval and processing : Traps were retrieved by bringing the retained CD lids up into the sampled tree, removing the exposed trap from its retaining nail, and recombining the case sections. Labels were checked while at the trap location. Once in the laboratory, animals were removed by soaking the entire trap in a kerosene bath for two hours then rinsing the animals off with water from a squeeze bottle.

Care must be taken not to let the traps soak for too long. When allowed to soak for 24 hours in mineral turpentine or citrus-based solvent, the plastic case chemically decomposed and irretrievably entombed the animals. It is possible that these solvents would work well if soaking time was less. Citrus oil-based solvents are generally superior to turpentine or kerosene, especially for removing specific individual animals.

7) Canopy Arthropods - Methods

Intact animals were hand sorted from the material. A large proportion of collected material was arthropod fragments, leading to underestimates of the true collection totals. Wings were often damaged. Specimens were generally in much worse condition than those in the liquid collection chambers of the other traps.

7.1.2 Funnel Crawl traps

Goals: Funnel crawl traps target animals walking on the tree surface and are based on the designs of Majer et al. (2003) and Hanula & New (1996). Trunk mounted funnels have primarily been used for capturing animals walking upwards by Majer et al. (2003), but can also be used to capture animals moving downwards (Moeed & Meads, 1983). A similar trap design is sometimes referred to as a “photo- eclector” (Nicolai, 1993). The design described here can also be used for animals walking horizontally on branches. Hanula & New (1996) found that drift fences leading into the funnels increased trap yield. Majer et al. (2003) used a portable angle grinder to create a mounting groove for a plastic fence. A technique was required for mounting a drift fence that could be anchored to the irregular stringy bark of E. obliqua, with a minimum of equipment carried into the tree crown.

Construction: Funnel traps were constructed from the bottlenecks cut from plastic drink bottles and configured to lead into intact collection bottles. A visit to a recycling centre netted 600 1.5 litre plastic (HDPE) drink bottles and caps at no monetary cost (Figures 7.7, 7.8). These bottles were rinsed in soapy water (~30 seconds per bottle) prior to use.

7) Canopy Arthropods - Methods

Figures 7.7 (left), 7.8 (right): Paul Osimo and Katy Dika acquiring plastic bottles at Hobart Recycling Centre

A funnel for each trap was made by cutting a bottle (~15 seconds) and pairing it with a similar but intact collection bottle. The collection bottles were filled with 200ml of 50% ethylene glycol /50% water and capped for transport (~$0.20). Several 10 mm x 10 mm x 1 m strips of closed cell foam (such as used for sleeping mats) served as drift fences (~$0.20). Several nails were used to mount the trap assembly on the tree. (~$0.06)

For funnel traps leading upwards, an extra bottle was used as a elbow chamber before the collection bottle. A motorized drill bit was used to make a 15mm hole in the side of this bottle.

To join the funnels to the collecting bottles, gaffers duct tape was used in the field (~$0.20). The care invested in creating this join strongly determined whether the collecting bottle would stay in place. If possible, a plastic female-female screw socket could be constructed to create a more robust binding, and is strongly recommended.

About ten minutes were required to construct each trap.

Trap setting: In the tree, the bottleneck funnel was nailed to the trunk or branch in the desired place and direction (Figure 7.9). A foam fence was then nailed into position with two strips leading into the funnel. When traps targeting insect movement in opposite directions were adjacent, a foam strip could serve as a fence for both. Labelling and bottleneck joining were performed in the tree. Joining the necks during initial construction would create a stronger trap, but not allow the collecting bottles to be transported while capped. Inferior bottle joining was the major source of trap failure.

7) Canopy Arthropods - Methods

For upwards traps, the elbow bottle was fitted securely onto the funnel bottleneck and the collecting bottle joined to it (Figure 7.10, 7.11). Downwards traps were vulnerable to dilution and obstruction falling debris. Basset et al. (1997) suggest salt water as a collecting fluid relatively resistant to dilution.

Installation required about 10 minutes per trap. Funnels can be placed on any surface of the tree and could be combined with mesh to create an emergence trap ((Grove & Bashford, 2003).

Data are from funnel traps set for 60 days targeting upward walking animals on the upper and lower trunk, and from horizontal traps targeting animals walking inwards on 4 branches coded as: upper crown live, lower live, upper dead, and lower dead. Traps were also placed and collected targeting outwards and downward walking animals, but were not analysed due to time constraints. More than a quarter (28 of 96) of the placed funnel traps failed, overwhelmingly due to failure of the duct tape bottle join.

Figure 7.9 (left): R. Junker placing live branch funnel traps on a megabranch in old tree 12. Figure 7.10 (centre): Upper trunk funnel traps in 100yr tree 7. Drift fences target upwards and downwards travelling animals. Trap loss for downwards traps was high due to extra stress on adhesive tape join from rainwater collection. Only upwards traps were processed. Figure 7.11 (right): A Hemipteran bug, Pentatomidae YDB sp.3, after entering the elbow bottle. These were the most commonly observed animals on the trunks of E. obliqua during field work

Retrieval and processing: Traps were retrieved by climbing to the trap location with the retained bottle caps, cutting through the duct tape to remove the trap, and recapping the bottle. Traps were relabelled in the field to safeguard against confusion.

7) Canopy Arthropods - Methods

Collecting bottles served as transport and storage devices until processing. Animals were generally in excellent condition and required very little sorting from debris.

7.1.3 Hangtraps

Goals: Omni-directional trap designs were based on the hanging traps illustrated by Wilkening et al. (1981) and the Malaise traps used by Basset (1991). Martikainen et al. (2000) used a similar design to sample flying beetles. Unlike sticky traps, or flight-intercept panels mounted parallel to the trunk surface (Majer et al., 2003), which are designed to capture animals landing on tree surfaces, these traps sample animals moving through the airspace within the crown. Unlike the flight intercept traps used by Chenier & Philogene (1989) the panels were not coated with sticky paint, and unlike the traps used by Yee et al. (2001), there was no open pan of collecting fluid.

Construction: The funnel and collecting components are similar to those described above for the funnel crawl traps, with 2 litre bottle funnels used for upwards and downwards bottlenecks (Figure 7.12). Bottle joins were of superior quality to the funnel traps due to easier working conditions on the ground. An omnidirectional panel assembly was used as the intercept surface (~$0.50) This was constructed by using plastic electrician’s ratcheting cable-ties to connect three plastic panels approx. 200 mm x 60 mm x 4 mm together in a triple- cross section shape through holes drilled in them (Figure 7.14). Alternatively, two panels twice the size could be slotted together in four-cross section. These were subsequently cable-tied to the funnels through holes melted or drilled in the bottlenecks. The panels could be constructed of any rigid transparent material. A total of 8 cable-ties were used for each trap (~$0.05), but could be replaced by wire or constrictor knots in cord (Budworth, 1999).

The plastic panels used were recycled from the flight intercept traps used by Yee et al. (2001), and required about 5 minutes of cutting to size. When all material were gathered, construction time was 15 minutes for each trap.

Trap setting: A cord was set in the uppermost branches of the tree crown by a climber and the traps were connected from the ground (Figure 7.13). The two legs of

7) Canopy Arthropods - Methods

the cord were tied together to form a closed loop and traps raised and lowered by pulling on the other side of the cord. Three traps were placed on each line using a clove hitch (Budworth, 1999) around the bottleneck. The uppermost trap was installed on the line first. Care must be taken when raising traps past understorey trees. Once the line was in place, trap installation required about 30 minutes for 3 traps.

Trap placement is limited only by the ability to place cordage and haul traps up through the forest canopy. Winchester (2004) placed hanging traps on strings across canopy gaps, and Basset (1991) placed them within each tree

Three traps were hung in the tree crown for 60 days. Traps were spaced equidistantly on the cord with the uppermost trap ~1m below the highest accessible branch, and the lowermost at the bottom of the tree crown. Each trap had two collecting bottles. Only 5% of trap bottles failed (5 of 96), mostly upon retrieval when the upper elbow chamber bottle caught on understorey foliage.

Retrieval and processing: Retrieval of the traps was performed from the ground by lowering them on the cord. The bottle join was removed and bottlecaps replaced and relabelled.

Animals were in excellent condition and free from debris.

Figure 7.12 (left): Paul Osimo melting panel attachment holes in bottlenecks using soldering iron. Figure 7.13 (centre): Upper and lower chambers of a omni directional flight intercept hangtrap Figure 7.14 (right): R. Junker assembling tri-panel intercept trap with plastic electricians cable-ties 7.1.4 Comparison charts

A diagram of trap construction and materials is presented in Figure 7.15 and a table comparing construction and placement of each design is presented in Table 7.1.

7) Canopy Arthropods - Methods

Figure 7.15: Trap construction diagrams

7) Canopy Arthropods - Methods

Hanging Omidirectional Sticky CD Case Funnel Crawl Flight Intercepts Fliers in tree airspace, Bark walkers, hoppers, skimmers, Target Fliers in tree airspace jumpers close fliers Ranking of 5 Diptera, Hymenoptera, Diptera, Coleoptera, most abundant Diptera, Hemiptera, Coleoptera, Coleoptera, Hemiptera, Lepidoptera, Hymenoptera, taxa (First is Hymenoptera, Blattodea Isoptera Araneae most abundant) Funnel not fixed closely enough to Potential Wind blowing in aerial trap surface- animals walk right Trap hanging in canopy of confounds plankton, Bark debris under, down traps are vulnerable neighbouring plants to rainfall and debris Specimen Medium, wings often lost Fine Fine conditions Bulky, but light. Most difficult Bulky, but light. Most difficult Portability into Compact, can use component is pre-filled collecting component is pre-filled field commercial music CD carriers bottle collecting bottle Total Time Effort (minutes:second 7:25 15:30 15:00 s per trap)

Cutting and combining of intercept panels, Washing From intact CD case: Washing and cutting funnels; and cutting funnels and elbow Construction Removing crown and painting Washing and filling collecting chambers, Washing and filling Time Effort glue ~20; Melting anchor hole bottles; Cutting of foam drift collecting bottles, Attachment ~5 seconds fences (optional); ~ 30 seconds of two funnels to intercept trap ~ 7 minutes

Once at branch, nailing drift fence Placement Time Once at branch, nailing into Once line is in place,~3 ~ 5 minutes; nail funnels and join Effort branch or trunk ~2 minutes minutes to clove hitch to cord bottles ~5 minutes Trap dissolving and specimen Trap Processing washing, ~ 20 minutes; Sorting animals from debris, ~ 5 Sorting animals from debris, Time Effort Sorting animals from debris, ~ minutes ~ 5 minutes 5 minutes Can label plastic directly with Can place final label in collecting Can place final label in Label permanent marker or scratch, bottle. Best done after placement collecting bottle. Best done considerations at risk of being lost in glue to avoid difficulty if a trap is after placement to avoid solvent dropped. difficulty if a trap is dropped.

Access to treetop, carrying bag, Cord rigged in canopy, Access to treetop, carrying hammer with nail pouch, marker or knowledge of clove hitch., bag, hammer with nail pouch, pencil and paper to label trap. marker or pencil and paper to Tool required marker or nailscratch to label Gaffer's tape for bottlejoins. Knife label trap. Gaffer's tape for trap. Must retain lid. or hands to cut foam to size. Must bottlejoins. Must retain retain bottlecap bottlecap. Total cost per $0.41 $0.67 $0.95 plus cordage trap

Table 7.1: Comparison of effort for each trap type

7.2 Trap processing

In April 2004, all traps were collected. Adult arthropods were identified to Recognizable Taxonomic Units (hereafter “RTU,” similar to morphospecies (Baldi, 2003; Basset et al., 2000; Oliver & Beattie, 1996; Pik et al., 1999)) and counted. Voucher specimens were photographed, assigned to family, and archived (Figure

7) Canopy Arthropods - Methods

7.16, 7.17). Araneae, Acarina, Hymenoptera-Formicidae, and larvae were counted but not sorted.

Coleoptera were pinned and identified to named species at the Tasmanian Forest Insect Collection (“TFIC”, Forestry Tasmania, Hobart, Australia). The species binomial of a RTU matched to a holotype in the TFIC contains the code “TFIC.”

RTU matched to Coleoptera in the Warra LTER collections of Kate Harrison or Marie Yee (Harrison et al., 2004; Yee et al., 2001) contain the code “KH” or “YEE.” All other RTU contain the prefix “YDB” to indicate their collection in the present study.

Araneae will be sent to E. Turner at the Queen Victoria Museum (Launceston, Australia) and Diptera to D. Bickel at the Australian Museum (Sydney, Australia). All other voucher specimens, photo catalogues, and computer CD archives of raw and processed data, will be lodged at the TFIC.

(left)Figure 7.16: A. Barrows using digital voucher photograph collection for arthropod sorting (right) Figure 7.17: Collection from upper trunk funnel crawl trap of 100yr tree 2

7.3 Data Analysis

All statistical tests and analyses were directed at comparing 100yr and old trees. Data analysis of the fauna collection from several types of traps was facilitated by identification of four approaches to studying seven aspects of biodiversity.

7) Canopy Arthropods - Methods

Four distinct approaches to analysis were used:

1. Age level 2. Tree level 3. Trap level 4. Placement level

The collected fauna of 100yr and old trees was compared for differences in seven aspects of biodiversity:

1. Animal abundance 2. Species richness 3. Community structure (plot of rank and abundance) 4. Resampled species accumulation curves and estimated true richness 5. Species diversity (evenness) 6. Community composition distinctness 7. Incidence/abundance trends for each RTU When reference is made to “all traps,” this means the pooling of all three trap types. When reference is made to “all traps+3,” this means analysis was done four times in total: first on all traps types pooled, then on each trap type separately.

In several instances, only the five most prominent arthropod orders were investigated: Coleoptera, Diptera, Hemiptera, Hymenoptera, and Lepidoptera. For these five taxa, the traps collected more than 20 RTU, and were deemed sufficiently abundant for statistical analysis. Other taxa were either not resolved to a sufficient taxonomic level, or were considered too scarce for analysis. When reference is made to “all taxa,” this includes these five and all other RTU encountered. When reference is made to “all taxa +5,” this refers to the repetition of the analysis six times: first for all collected RTU, and then separately for each of the five orders. “All taxa” does not mean that each RTU was tested separately and that they were all in agreement, but rather that these were all pooled together.

Tabulation and graphical representations were generated by Microsoft Excel (Microsoft, 2000). Univariate and bivariate analyses were performed using both JMP and Microsoft Excel (Microsoft, 2000; SAS Institute, 2003) . Multivariate analyses utilized PC-ORD software (McCune, 1999). Species accumulation curves and

7) Canopy Arthropods - Methods associated estimators were generated with EstimateS (Colwell, 2004). Analysis followed the advice of Clarke and Warwick (2001), McCune and Grace (2002), and Magurran (2004). 7.4 Analysis Approaches

Approach 1- Age level : At an age level, the fauna from all trees from each age class were pooled, with no reference to what tree they were collected in. There are only two entities contributing: the 100yr and the old age classes. This approach is only used for illustration and summarization. Statistical analysis is not possible because there is no replication at this level. The age level is the lumping of the tree level statistics.

Approach 2- Tree level : At a tree level, the independent sampling entity is considered to be a single tree. The entire collections from each of eight 100yr trees are compared to that of eight old trees. An assumption is made that the loss of traps was random and unbiased across the two age classes. When a trap type in a specific tree had one or no surviving traps, the entire tree was excluded from the analysis. This occurred with CD traps of trees 7 (100yr) and 16 (old), and with funnel traps of trees 5 (100yr) and 6 (old). In theory, sixteen entities contributed: eight in each age.

The consideration of a tree as the sampling unit can be seen in the equivalent treatment of fauna collected from trees of differing sizes (Grove et al., 2002; Majer et al., 2000), as opposed to the standardization by volume or mass encountered in techniques such as branch clipping (Abbott et al., 2000).

The tree level is the lumping of the trap level statistics.

Approach 3- Trap level : At a trap level, each trap is considered as a sampling unit and the pool of traps is divided into traps from 100yr trees, and traps from old trees. The pools are of similar sizes, supporting the previous assumption that trap loss was random across age classes. 93 traps from 100yr trees and 91 traps from old trees contributed to the trap pool, for a total of 184.

Because the individual tree is the subject of consideration, this is a pseudo- replicated approach. The lumping of trap results from different situations (upper with lower, or funnel with CD) is a detrimental consequence of this approach.

7) Canopy Arthropods - Methods

Trap level analysis was only used for multi-variate community analyses in which the tree may not be the fundamental unit of distinction between faunas. In these analyses, both age and individual tree identifier were treated as distinct factors.

Further pseudo-replication exists in trap-level statistics in the treatment of the upper and lower chambers of hangtraps as separate entities. The trap level is the lumping of the placement level statistics.

Approach 4- Placement level: At a placement level, pseudoreplication and lumping of trap results is avoided by only comparing traps of the same type and position. The effect of missing traps is minimized by calculating values based on means. This approach is the most statistically robust and requires the least assumptions, but offers the least immediately interpretable results. Different trap positions may exhibit conflicting trends. At a placement level, sixteen trees theoretically contributed, and this was repeated fourteen times to address each placement. Tabulation of statistics at a placement level also allows comparison of different positions within the tree, with a robust replicated design.

7.5 Biodiversity aspects

Aspect 1- Abundance: Abundance is the number of animal individuals present. Abundance is a simple measure of how many animals were collected, and is of limited applicability when lumping several taxonomic units. This measure is used to test Hypothesis 5 (Old-growth E. obliqua trees have a more abundant arthropod fauna than 100 year old trees, i.e more individual arthropod animals). Abundance is most useful when describing different taxa and their relation to another. This measure is the most vulnerable to differences in sampling effort due by missing traps.

By analogy, this is like counting the number of words in a book, without reference to their classifications or whether they had been encountered before. By counting the words in two shelves (i.e. age classes) of books, comparisons can be made between the shelves. Comparisons can be made between two books (i.e. age classes) by counting the words in each. Studying the abundance of one group, for example, the insect order Coleoptera, is analogous to studying only one chapter of the book.

7) Canopy Arthropods - Methods

Radar charts are presented at an age level illustrating abundances of taxa with >20 individuals for alltraps+3. Radar charts assign each taxon an axis that radiates outwards from a centre point. The distance from the centre to a mark is proportional to the value. Along each axis, a mark is made for each age class. Lines connecting the marks for each age class assist in highlighting differences in abundances of different taxa. Because the ordering of the axes, and therefore the points on the polygon are arbitrary (alphabetical), the shape is for visual emphasis only. Natural logarithmic Ln() transformations on the abundance data were used to preserve relationships but shrink the effect of outliers.

At a tree level, the mean total abundances per tree for alltraps +3 was tabulated for each age and compared with a nonparametric Kruskal-Wallis test (hereafter “K-W test”). This compares the two ages using their ranks, and produces output similar to a T-test (SAS Institute, 2003). T-tests were rejected because abundance data was not normally distributed.

At a placement level, the mean and total abundance of all taxa per trap placement is tabulated and compared with by K-W tests. A second table presents similar information subdivided into different taxonomic units.

Aspect 2- Richness: Species richness (hereafter “RTU count”, or “richness”), is measured in surrogate by the number of recognizable taxonomic units present in a given situation. This measure is used to test the Hypothesis 6 (Old-growth E. obliqua trees have a richer arthropod fauna than 100 year old trees, i.e. more recognizable types of arthropods.) Oliver & Beattie (1996) test the use of RTU by non-expert taxonomists in determining true species richness and conclude that the two errors of splitting and lumping taxa do not significantly alter the end results. Richness is the aspect of communities most often equated with diversity in a system (Magurran, 2004). Gaston & Spicer (1998) explore the use of richness as the most readily applicable measure of biodiversity.

Returning to the analogy of a book, species richness of a given community would be the number of different words encountered in the entire book, counting each distinct word only once. The richness of an order would be the number of different words in a chapter, and the richness of a family would be analogous to the number of different words in a chapter section.

7) Canopy Arthropods - Methods

Richness was compared in a similar fashion to abundance, but only using the five most prominent orders. Radar charts were generated at an age level, and mean abundances are tabulated at a tree level and placement level.

Aspect 3- Community structure: Community structure examines the rarity and dominance associated with a group of organisms. Graphical representations are generated relating abundance of an RTU to its relative abundance rank, thereby incorporating a measure of evenness (Magurran, 2004). The taxonomic identity of an RTU is ignored. A community with an even distribution of abundances for all taxa would be considered more diverse than a community with several very common and several rare species. This was used to test hypothesis 7 (Old-growth E. obliqua trees have a more diverse arthropod fauna than 100 year old trees).

By analogy, a human society in which several ethnic groups have an equal voice is considered diverse and desirable. Cultures with many groups present in which one group has a voice that dominates all others are considered less diverse. Cultures in which everyone has an equal voice, but where few ethnicities are present, are also considered less diverse.

A Whittaker rank abundance plot was generated with 100yr and old curves plotted together (age level) for all traps +3. Each RTU was ranked from most abundant to least abundance on the X-axis, and relative abundance was plotted on a log-scaleY-axis. The shape of the resulting curves were visually assessed to assign to a community structure model as guided by Magurran (2004). A steeper curve was interpreted as representing a less even community, in which the most abundant RTUs are more abundant and dominant than other taxa.

Aspect 4- Diversity: Diversity indices are quantifications incorporating richness and evenness, used for comparison between communities. A highly diverse community has a higher richness and is more even than a less diverse one. Magurran (2004) suggests the use of the negative ln() transformations of Simpson’s “D” Diversity Index (Simpson, 1949). After this transformation, a higher value indicates a more diverse community. This was used to test hypothesis 7 (Old-growth E. obliqua trees have a more diverse arthropod fauna than 100 year old trees).

Simpsons D is calculated using the formula:

7) Canopy Arthropods - Methods

D=Σ(( ni(ni-1)) ÷ (N(N-1)))

Where ni is the number of individuals in the ith RTU, and N= the total number of individuals (Magurran, 2004; Simpson, 1949)

-ln(D) was analysed in a similar fashion to abundance and richness. Age level radar charts were graphed. Tabulations of tree level and placement level diversity comparisons for alltraps +3 and alltaxa+5 were generated.

Aspect 5- Species accumulation curves & estimators: By computer randomization of the sequence in which samples were sorted, a species (i.e. RTU) accumulation curve can be plotted showing the cumulative number of species encountered (Y axis) as sampling effort increases (X axis) (Colwell, 2004; Gotelli & Colwell, 2001; Longino, 1994). As the rate in which new species are encountered decreases, one can surmise that one is getting closer to having encountered most of the species present. Because resources are limited, biodiversity researchers can rarely “get them all” and must “sample and estimate” the true richness (Longino, 1994). The true richness can be interpreted as the asymptote hinted at by the curve.

Species accumulation curves for old and 100yr trees were plotted together and the shape and behaviour of these curves compared. For this analysis, “species” means “RTU.” This was used to test 6 Hypothesis 6 (Old-growth E. obliqua trees have a richer arthropod fauna than 100 year old trees) and Hypothesis 7 (Old-growth E. obliqua trees have a more diverse arthropod fauna than 100 year old trees).

A useful analogy can be drawn with the determination of how many restaurants (i.e. species) are in a region. By questioning of guests (i.e. samples) at a party as to what restaurants they have dined at in the last week, an estimate can be formed. As one enters the room, no questions have been asked, and one has not heard of any restaurants. Sampling effort is zero, and restaurants accumulated are zero. The first guest approached has been to a group of restaurants, and the next guest has been to a different group of restaurants. Only restaurants not yet encountered are added to the list as more guests are approached. At a certain sampling effort, by which time the list has grown to a certain length, it will seem guests have fewer or no new restaurants for the list. This decline in rate is the asymptote, and the list can be considered to be approach completeness.

7) Canopy Arthropods - Methods

At parties with people who dine out often (traps collect a wide range of animals), this point is reached more quickly than at parties with people who most often eat at home (less effective traps), who only go to popular restaurants (less likely to add to the list) or who never eat out at all (traps that did not catch any animals).In regions without many restaurants (low species richness), the list is shorter than regions with many (high richness). Parties with only vegetarians, recent immigrants, or poor students will generate a bias in the list.

Software such as EstimateS (Colwell, 2004) randomizes the order in which samples are encountered (i.e. in which guests are approached) and generates a curve representing the average species accumulation.

In addition to a curve describing the accumulation of observed species, the number of animals encountered once (“singletons”) and animals encountered twice (“doubletons”) can be utilized to generate a true richness estimator. Following the recommendation of Magurran (2004), the Chao 1 true richness estimator was calculated. The Chao 1 abundance based richness estimator is introduced by Chao (1984) and presented by Magurran (2004) in the form:

2 Chao 1 True Richness Estimator Schao1 = Sobs + ((F1 ) ÷ (2F2))

Where

Sobs= number of species actually observed in sample

F1 = number of RTU encountered in a single individual

F2 = number of RTU encountered by two individuals (Chao, 1984; Magurran, 2004)

Using data entered at a tree level, species observed accumulation curves and Chao 1 true richness estimator curves were produced using 50 resampling runs in EstimateS software (Colwell, 2004). This was repeated for each age class, and each trap type. Both age classes were plotted simultaneously on a scatterplot for all traps +3. Cumulative species encountered or estimated was plotted on the Y-axis, and individuals sampled on the X-axis. The steepness and tendency towards asymptote of each curve was compared visually. This was performed for alltraps+3.

7) Canopy Arthropods - Methods

Sampling adequacy was calculated for each combination of age class and all traps+3. This was expressed by the percent proportion of the estimated true richness actually encountered.

Sampling adequacy =Sobs ÷ Schao1 x100

Aspect 6- Distinctness: A different approach than measuring biodiversity is to compare the taxonomic composition between the age classes. Communities may be composed of completely different elements, but summary measures of richness, diversity, and community structure may be the same. By analogy, two books may have a similar abundance, richness, and diversity of words, but the actual words used were different. Several techniques were used to test Hypothesis 8 (Old-growth E. obliqua trees have a distinct arthropod fauna from 100 year old trees, i.e. the taxonomic composition of animals differs.)

At an age level, Venn interlocking circles illustrate the overlap in RTU between ages for all traps +3. The size of plotted circles was proportional to the RTU richness.

Multi Response Permutation Procedures (MRPP, (McCune & Grace, 2002)) were run on PC-ORD software (McCune, 1999) to test the null hypothesis of no difference in community composition between the age classes. This was performed at a tree level, a trap level, and a placement level. MRPP tests only for a difference (like a 2-tailed t-test) but are not associated with a direction or an indicator species.

At a tree level, non-metric multidimensional scaling ordinations (NMS, sometimes referred to as MDS in other research) were generated using Sorenson/Bray-Curtis distance matrices on PC-ORD to display relative community similarities (Clarke & Warwick, 2001).. NMS was selected as a non-parametric test that has no assumption of normality and is robust to sampling irregularities, following the suggestion of McCune & Grace (2002). Abundances were transformed using the formula (log x+1) to mitigate the effect of overabundant taxa. A decision was made to include uncommon taxa (less than five individuals) in the ordinations. Because of the large number of uncommon taxa, removing them from analysis would result in an

7) Canopy Arthropods - Methods

undesirable loss of information. Study pairs are graphically linked by a vector from the 100yr tree to the old tree. This was performed for alltraps+3 for all taxa+5.

A tree level dendrogram showing a hierarchical clustering (by Ward’s method) of trees based on the Pythagorean distance matrix of their fauna was generated (Clarke & Warwick, 2001; McCune & Grace, 2002). Again, the data was log- transformed.This was performed for alltraps+3 for all taxa.

Aspect 7- Incidence/Abundance of each RTU: The response of each RTU was examined to determine if any taxa showed strong bias of occurrence or abundance to one age class. This was used to test Hypothesis 8 (Old-growth E. obliqua trees have a distinct arthropod fauna from 100 year old trees, i.e. the taxonomic composition of animals differs.)

This is analogous to performing an analysis on each word encountered in two books, to see if a specific word is encountered more often in one book or the other.

The age level abundance was investigated by counting the RTU that were more abundant in one age class. The tree level presence absence was investigated by counting the RTU that were present in more trees of one age class. The mean abundance of the 116 RTU with greater than 5 individuals collected was compared with t-tests at a tree level.

7.6 Scaling of results

Arthropods results were not scaled: Care must be taken in interpreting results. An equivalent number of traps were placed in trees of vastly differing size. Consideration is required before interpreting these as comparable, equivalent samples. A decision was made not to scale results to a measure of tree size.

Scaling up or down can reconcile differences between the scale of the investigation and the scale of the sampling (Figure 7.18). The choice to scale up or down can change the interpretation of the results. Researchers must consider their

7) Canopy Arthropods - Methods

object of concern, and the scale of their sample. Two critical questions must be asked: 1) Which is the unit of interest, a piece of the tree or the whole tree? 2) How much of the tree was sampled, a piece or the whole?

Four combinations of answers exist:

A) Number per tree is desired, and the entire tree is sampled. No scaling is required. (Majer et al., 1994; Tassone & Majer, 1997)

B) Number per tree is desired and quantifiable pieces are sampled. Scaling up produces densities per tree. It addresses the entire tree when the piece is sampled. (Hooper, 1995)

C) Number per unit is desired and the entire tree is sampled. Scaling down produces densities per unit of scale. It addresses the piece when the entire tree is sampled. (Thunes et al., 2003)

D) Number per unit is desired, and units are sampled. pieces of the tree are sampled. No scaling is required. (Abbott et al., 1992)

In the present study, the scale of interest is the entire tree (A or B). The difficulty lies in judging the scale of sampling when using passive traps. Did the traps sample a finite piece of the tree, or did they sample the entire plant (or proportion of the plant)?

For example, Majer et al. (1990) do not scale results from chemical knockdown in Eucalyptus trees. They fogged four species of Eucalyptus trees and collected the falling animals in funnels suspended in the tree. The funnels were of a standard size and were placed to collect below an equalized amount of foliage. It is unknown if this represents a sample of an entire plant or just a piece of the plant. Similarly, they do not scale their results. Scaling requires a measure of size. Crown area, foliage volume, crown volume are all measures of size, and to some extent, they may all be inappropriate.

7) Canopy Arthropods - Methods

Scale of Sampling

Sampling the Sampling a entire plant quantifiable piece

A. No scaling B. Scale up

Numbers per tree

Scale of

interest C. Scale down D. No scaling

Numbers per unit of scale

Figure 7.18: Scaling considerations. The researchers are looking through their sampling hand lens at either the entire tree or a piece of the tree. They are considering either the entire tree or the piece of the tree.

Relationship of abundance and richness to scale: Abundance and biomass is assumed to be linearly related to the unit of scale (Hooper, 1995). When scaling, biomass or abundance can be arithmetically calculated up or down by multiplication or division (Hooper, 1995; Schowalter, 1995).

Species richness cannot be calculated in this fashion. A basic principle of island biogeography is that richness increases non-linearly with increased 2- dimensional island study area (MacArthur & Wilson, 1967). Species accumulation curves can combine several samples in a tree to scale up to the size of an entire tree using a measure of sampling effort (Gotelli & Colwell, 2001). Coleman rarefaction curves can be used to scale down larger samples (Colwell et al., 2004; Gotelli & Colwell, 2001). Species accumulation curves and rarefaction curves can also be used to scale species richness by the number of individuals. However, in the present context the unit of scale is a physical measurement of size. Both represent used as measure survey of effort.

7) Canopy Arthropods - Methods

Scaling up to address the tree: Studies of foliage arthropods frequently present results as densities per kilogram (Abbott et al., 1992; Majer & Recher, 1988; Schowalter, 1995; Wotherspoon, 1998). Small pieces of a tree were sampled, with the effort measured by number of small pieces.

When combined with the amount of foliage per tree or per hectare, studies can be scaled up. In the absence of foliage measures for each tree or hectare, the reported numbers do not relate to the biodiversity of a tree, but instead relate to an amount of foliage. Research sampling small pieces of a tree is made much more useful by the inclusion of the relative size of each tree, to allow for scaling upwards.

When not scaled up, it is important to remember that results are for densities per unit rather than for densities per tree. Schowalter (1995) compares intensities of taxa per kg of foliage, and reports them using phrases such as “more abundant in old- growth trees than in younger trees.” The use of the word “trees” implies that the values presented are at the scale of a “tree”. However, these are actually densities linked to a unit of scale within the tree, rather than a density linked to the entire tree. What Schowalter (1995) is actually reporting is that these taxa were “more dense” rather than “more abundant”. Scaling up was judged inappropriate for the present study. It is unclear what the appropriate unit of scale is, and unclear what portion of that unit the traps sampled.

Scaling down to address the piece: When the scale of interest is the piece of the tree, samples from different sized trees can be by scaling down abundances arithmetically and/or richnesses using rarefaction curves (Gotelli & Colwell, 2001). For example, Mody et al. (2003) and Thunes et al. (2003) sampled entire trees using canopy fogging, and then scaled down to unit of volume. Rather than use the captured numbers as densities by tree, their sample of interest was the densities by piece. They scaled down to generate densities per unit of volume.

However, scaling down does not address the tree as the actual unit of consideration. Because trees are counted by the stem, scaling down does not address resource management concerns. Scaling down may be appropriate for studies of ecophysiology and herbivory, but in the context of the present study it was judged unnecessary. The subject of interest is the entire tree.

7) Canopy Arthropods - Methods

Assumptions in scaling: Two large assumptions are required when scaling. Scaling up requires an assumption that the sampled fauna is homogenous throughout the entire plant. Scaling down requires an assumption that the compared plants are congruent in all aspects save size. There is evidence against both of these assumptions.

Majer et al. (2001) report on high intra-crown variability in fogged invertebrates in Eucalyptus trees when compared to inter-crown variability. This suggests that fauna is not homogenously distributed through the crown of Eucalyptus trees. In the present research, evidence was collected that emphasizes that the smaller and larger trees differ in more than just size.

Hooper (1995) measured the biomass of arthropods on the bark Pinus palustris in South Carolina. He compared trees at representing ten year intervals in a chronosquence from 20-130 years. Data was collected per unit of bark area. Hooper (1995) presents the raw results per m2 simultaneously with scaled results per tree. When biomass was scaled up multiplicatively by surface area, total biomass per tree was highest in old trees. Before scaling up, results standardized to area showed the younger trees to have a higher biomass per m2. Despite arthropod biomass being more sparsely distributed on the surface of larger trees, the greater surface area meant higher biomass for the entire tree. The context of his study was to assess the food available for the red-cockaded woodpecker Picoides borealis. To the bird, a high density of arthropod biomass per unit of surface area in the younger, smaller Pinus means less foraging time, The lower density of biomass in the entire tree means less food available in the entire tree.

The most critical decision in scaling down or up is the unit of scale. Foliage is easily quantifiable by mass and is assumed to be homogenous throughout a tree when scaling up (Fowler, 1985). In studies of other habitats, the unit of scale is not so easily defined.

Choosing a measure of scale: Choosing any measure of scale is based on assumptions. For example, Majer et al. (1990) collected arthropods in funnels of standard diameter in fogged Eucalyptus crowns. Had they chosen to scale up, it would be unclear if these funnels had sampled a quantifiable area of crown spread, or a quantifiable volume of crown volume, or a quantifiable mass of foliage.

7) Canopy Arthropods - Methods

Thunes et al. (2003) do not find a correlation between tree height and chemical knockdown species richness. Before scaling down, they found a significantly richer fauna in old, larger volumed Pinus sylvestris than in younger, smaller volumed trees. Their decision to scale down by crown volume is not supported by this lack of correlation. Other habitats besides foliage within the tree may not scale down in step with volume. For example, in the present study, total crown polyhedra volume in 100yr E. obliqua was approximately half that of old trees, whereas the total live branch polyhedra volume was approximately one quarter.

Therefore, scaling down requires the assumption that smaller and larger trees are compositionally similar in all aspects except size. In Eucalyptus trees, most foliage is in the outer crown (Jacobs, 1955). For animals associated with foliage, the empty space is irrelevant.

A further problem with using values per kilogram of foliage is that the samples may not have actually been exactly one kilogram of foliage. When standardizing these varying masses of foliage by kilogram, a similar assumption is made that richness and abundance are both scaling linearly with mass. Few studies that present values per kg of foliage actually report the amount of foliage per sample. Schowalter (1995) and Clement (2001) report on the length of branches sampled, but then present results as arthropod numbers standardized by kg of foliage. It is unclear if each sample was greater than a kilogram, or less than a kilogram. When standardizing by mass, it is unclear if they multiply their samples and thereby “create” animals that never existed, or did they divide them and therefore “split” the animals they did collect? A potential solution is to present the size of the sample in the units of scale used for standardization (Majer & Recher, 1988), or, preferably, to present the size of the sample and leave the results unscaled.

It is possible that, for a given taxa, the unit of scale is cryptic. For example, Andersen & New (1987) studied the arthropods living in Eucalyptus fruits. For these animals, the most appropriate unit of scale would be the number of fruits in the tree. Using crown volume assumes that fruits were evenly distributed spatially throughout the crown, i.e. fruit habitat abundance scales linearly with crown volume. .

It is possible that cryptic organisms require cryptic habitats not normally considered by humans in land management (Bickel & Tasker, 2004), such as

7) Canopy Arthropods - Methods mistletoes (Watson, 2004), branch armpit ridging, or even fungal fruiting bodies growing in stringybark litter in small elbow hollows on dead epicormic branches on the sunny side of the tree. In the absence of a comprehensive knowledge of the arthropod communities in old eucalypts (Majer et al., 1997), these examples cannot be disregarded. The appropriate unit of scale for a cryptic taxon may be unrecognizable to human researchers.

Other measures of scale: The most common units for scaling are crown volume, bark surface area or leaf biomass. Other units have been used, including profile area, leaf area, and height.

Martin (1966) found abundance densities of arthropods per unit of crown profile area to be lowest in older Pinus plantations. Crown profile area was calculated by multiplying crown depth by crown diameter. Results were not scaled up, and crown sizes were not provided to allow the reader to scale up arithmetically. They do, however, comment that the highest abundances per tree were taken in the older plantations (25 years), in which trees were largest. The adoption of a crown size unit of scale reflects a silvicultural interest in phytophagous pests. The chosen measurement of scale is a 2-dimensional simplification of a 3-d space. The choice of a vertical area unit is unique in the literature.

Leaf surface area can be used as a measure of foliage scale instead of mass. Fowler (1985) scales down phytophage richness to Betula leaf surface area by using a linear approximation of an observed species accumulation curve (before the development of computer bootstrapping resampling techniques used in Colwell (2004). Scaling down was not performed from samples of an entire plant, but from the larger branches clipped in the larger plants. Fowler (1985) performs this scaling down in the context of comparing the architecture of seedlings, saplings, and trees, and discusses the greater variety of habitats in the larger, more complex trees. The assumption that sampled leaves on branch trips were equivalent to leaves elsewhere in the tree is stated. Examination of the species accumulation by cumulative leaf area plots presented by Fowler (1985, Figure 1) shows a steeper accumulation of phytophagous insects on trees than seedlings. This suggests a different, richer community structure on the larger, older trees. However, when scaled down, this differences is negated.

7) Canopy Arthropods - Methods

Tree height offers a one-dimensional unit of scale. Richardson et al. (1999) collected arthropods using chemical knockdown in Melaleuca and found an correlation between tree height and species richness. They did not, however, find a similar correlation for crown volume and species richness or abundance. This suggests crown volume is not an appropriate unit of scale.

Scaling up or down were both rejected: Scaling up or down was not used in the present research. Doubt remains as to the appropriate measure of scale, and as to whether traps sampled a fixed portion of that unit.

In data analysis, the assumption that traps sample a fixed volume (B in Figure 7.18) was judged to be less stable than the assumption that traps sample a fixed proportion of the tree crown (A in Figure 7.18), and was therefore rejected. The reality is certainly somewhere between these two assumptions.

The important failings of the “fixed volume” assumption (situation B) are twofold: choosing a unit of scale is problematic, and measuring how much of that unit was sampled is questionable. For phytophage studies, foliage is easily quantified and is a discrete, repeating resource, whereas the units sampled by the traps the present study are not so clearly defined.

In the current context, evidence from the current research shows that 100yr and old E. obliqua differ in more than just size. Old trees are not merely scaled up version of 100yr trees. Consequently, the catch from old trees was not scaled up in proportion using any calculated measure of size.

The values presented represent relative measures per trap. Results should be interpreted as relative levels standardized by trap numbers, rather than as abundance per tree or abundance per unit of size, volume, or foliage.

8) Canopy Arthropods - Results

8) Canopy Arthropods –Results

The results from the traps for each aspect of biodiversity are presented. For abundance, richness, and diversity, differences were detected more often at a placement level than at a tree level. Abundance was even between age classes for CD sticky and hangtraps but more higher in 100yr trees for the funnel traps. Richness was higher in the old traps for the CD sticky and hangtraps, but lower in the funnels. Diversity was higher in the old trees for most taxa. Community structure rank/abundance plots showed a more dominated community in the 100yr trees. Chao 1 estimators showed a higher species richness in the old trees for the CD sticky traps, about equal in the hangtraps and higher in the 100yr trees for the funnels. Distinction in fauna were detected at a trap level. NMS ordinations showed several strong age class distinctions that were often masked by environmental effects. 8.1 Trap survival

The survival of traps varied between trap types. CD sticky traps and funnel traps failed more often than the hangtraps. All lost hangtraps were upper chambers that entangled during raising or lowering along the cord.

In total, 93 traps were collected from 100yr trees, and 91 from old trees (Table 8.1). In each of 100yr tree 5 and old tree 6, only one funnel trap survived. In 100yr tree 7 and old tree 16, no CD sticky traps survived. These trees were subsequently removed from tree level analysis for that trap type.

A chi-squared analysis of variance was performed to test the assumption that trap loss was randomly distributed. No significant difference from the expected values 2 was detected (14 placement rows x 2 age columns; χ 13 d.f. =1.779, p=<0.9999). 8) Canopy Arthropods - Results

Trap survival by placement and trap 100yr Old 1 = trap processed Pair: A B C D E F G H 100yr A B C D E F G H Old Tree: 12579111315Total 3 4 6 810121416Total CDSticky Upper crownTrunk CD 111-1 - 1 1 6 111- 1 1 1 - 6 Lower crownTrunk CD 111-1 1 1 - 6 1111 1 1 1 - 7 Total 12 Total 13 Funnel: Upper crown Live Funnel 1--11111 6 11-1111- 6 Upper crownDead Funnel 11--11-1 5 ----1111 4 Upper crownTrunk Funnel 11-11111 7 11-11111 7 Lower crownLive Funnel 11---111 5 11111111 8 Lower crown Dead Funnel 111--111 6 11--1-11 5 Lower crown Trunk Funnel -1-111-1 5 -1-1--11 4 Total 34 Total 34 HangTrap: Upper crownUp bottleHangtrap 11111 1 1 1 8 -111 1 - 1 1 6 Upper crownLow bottleHangtrap 11111 1 1 1 8 1111 1111 8 Mid crown Up bottleHangtrap -1111 1 1 1 7 1111 1 1 - 1 7 Mid crown Low bottleHangtrap 11111 1 1 1 8 1111 1111 8 Lower crownUp bottleHangtrap 11111 1 1 1 8 -111 1111 7 Lower crownLow bottleHangtrap 11111 1 1 1 8 1111 1111 8 Total 47 Total 44 100yr Grand Total 93 Old Grand Total 91

Table 8.1: The survival of traps in different placements compared across trees and age classes; only trap positions actually studied are listed. “1”= processed; “-“ = lost trap. 8.2 Total collection overview

6490 arthropods were collected and classified into 312 RTU (Table 8.2). Four of the 312 RTU were larval hexapods. The hangtraps caught the most animals (2628), followed by the funnels (2724). The sticky trapping caught the least animals (1240).

Overall, the collection was dominated by Diptera, comprising almost half of all collected animals (47%) (Table 8.3). Diptera, Coleoptera, Hymenoptera, Hemiptera, and Lepidoptera combined were 85% of the entire collection. Funnels and hangtraps caught a wider range of animals than the sticky traps.

Abundance Richness 100yr Old 100yr Old CD Sticky 637 589 67 81 Funnel 1564 1115 137 110 Hangtrap 1210 1227 155 170

Table 8.2: Comparison of abundance, richness compared between ages and trap types

8.3 Trap type comparison

For comparison, the ordinal level results from the canopy fogging of five E. obliqua at Warra are presented (Bashford et al., 2001; Grove et al., 2002). The fogged 8) Canopy Arthropods - Results trees were less than 10 km from the sixteen study trees. The total abundance of collection from the fogging, funnels, and hangtraps was coincidentally similar (Tables 8.2, 8.3). The total CD sticky collection had about half as many individuals as these other techniques. Proportionally, fogging caught proportionally more Coleoptera and substantially less Diptera. Collembola, Thysanura, Thysanoptera and Dermaptera were present in the fogging but rare in the traps. Lepidoptera were much more common in the traps. Plecoptera and Orthoptera were found mostly in funnel traps but absent in the other techniques except for a few specimens in the fogging.

Abundance (#), Percent of Trap Total Catch (%) and Mean Number collected per trap (mju) Project Total Sticky Funnel HangTrap Fogging Identified Taxa # % # % µ # % µ # % µ # % Acarina 100 1.54 5 0.40 0.20 78 2.86 1.10 17 0.67 0.35 71 2.91 Araneae 321 4.95 5 0.40 0.20 180 6.61 2.54 136 5.38 2.83 54 2.21 Blattodea 225 3.47 1 0.08 0.04 198 7.27 2.79 26 1.03 0.54 43 1.76 Coleoptera 792 12.20 116 9.35 4.64 376 13.80 5.30 300 11.88 6.25 1070 43.78 Collembola 15 0.23 0 -- 9 0.33 0.13 6 0.24 0.13 159 6.51 Dermaptera 3 0.05 0 -- 3 0.11 0.04 0 -- 18 0.74 Diptera 3071 47.32 831 67.02 33.24 792 29.07 11.15 1448 57.32 30.17 185 7.57 Hemiptera 645 9.94 53 4.27 2.12 465 17.07 6.55 127 5.03 2.65 523 21.40 Hymenoptera 655 10.09 185 14.92 7.40 331 12.15 4.66 139 5.50 2.90 251 10.27 Isoptera 48 0.74 22 1.77 0.88 13 0.48 0.18 13 0.51 0.27 1 0.04 Lepidoptera 388 5.98 7 0.56 0.28 166 6.09 2.34 215 8.51 4.48 34 1.39 Neuroptera 2 0.03 0 -- 1 0.04 0.01 1 0.04 0.02 3 0.12 Orthoptera 17 0.26 0 -- 17 0.62 0.24 0 -- 1 0.04 Plecoptera 48 0.74 0 -- 48 1.76 0.68 0 -- 2 0.08 Pseudoscorpionidae 2 0.03 0 -- 2 0.07 0.03 0 -- 4 0.16 Psocoptera 7 0.11 1 0.08 0.04 0 -- 6 0.24 0.13 0 - Thysanoptera 0 - 0 -- 0 -- 0 -- 8 0.33 Thysanura 0 - 0 -- 0 -- 0 -- 17 0.70 Trichoptera 3 0.05 0 -- 0 -- 3 0.12 0.06 0 - Total 6490 animals 1240 animals 2724 animals 2526 animals 2444 animals (Excludes fogging) from from from from 25 traps 71 traps 48 traps 16 trees 16 trees 16 trees 5 trees

Table 8.3: Summary of total arthropod catch by trap type, with Warra canopy fogging data presented for comparison (Grove et al., 2002)

The five most abundant taxa were also the five most RTU-rich taxa (Figure 8.1). Overall, the hangtrap collection was the richest (226), followed by the funnels (162) and the CD sticky traps (102). 8) Canopy Arthropods - Results

RTUs collected by Trap Type

100

80

60

40 RTU count 20

0 Cole Dipt Hyme Hemi Lepi Blat Psoc Isop Plec Tric Coll Derm Neur Ortho Pseud Acar Aran

CDSticky 27 28 27 13 4 1 1 2 0 0 0 0 0 0 0 1 1 Funnel 60 21 27 30 12 5 0 1 2 0 1 1 1 1 1 1 1 HangTrap 79 46 36 32 20 5 4 1 0 2 1 0 1 0 0 1 1 Order x Trap Type

Figure 8.1: Richness of each order by trap type. From left to right: Coleoptera, Diptera, Hymenoptera, Hemiptera, Lepidoptera, Blattodae, Psocoptera, Isoptera, Plecoptera, Trichoptera, Collembola, Dermaptera, Neuroptera, Orthoptera, Pseudoscorpionidae, Acarina, Araneae. The overlap between trap types is illustrated in Figure 8.2. Hangtraps had the most RTU that were not found in any other trap type. Funnel traps and hangtraps had the most RTU in common, and the CD sticky and funnel traps had the least in common. Because the CD sticky traps were always placed immediately next to a funnel trap, this indicates that these trap types sampled different communities of arthropods.

306 total RTU

CD Funnel Sticky 23 6 51 Crawl 52 21 53

100

Hangtraps

Figure 8.2: RTU overlap between trap types.

8) Canopy Arthropods - Results

8.4 Abundance results

The total collection of arthropods in hangtraps and CD sticky was similar between ages (Figure 8.3). The total collection in funnel traps was insignificantly higher from the 100yr trees than the old trees (KW test, p>0.05, d.f.= 16).

Total abundances of trap type for 100yr and Old trees

2000 100yr Old 1500 1564 1210 1227 1000 1115

Abundance 500 637 589 0 Cdsticky Funnel Hangtraps Trap Type

Figure 8.3: Total abundances for each trap type compared by age. Mean total abundances per tree were not significantly significant between ages.

Overall, taxonomic abundances viewed at an ordinal level were similar between ages (Figure 8.4). Acarina, Coleoptera, and Lepidoptera were more abundant in 100yr trees, and Araneae, Blattodea, Hymenoptera: Formicidae, and Isoptera were more abundant in old trees. The actual shape of the radar plot line links is arbitrarily based on the alphabetical ordering of the axes. Therefore, the lines linking points are only useful in visualizing the position of a given point. 8) Canopy Arthropods - Results

Abundance for taxa with >20 individuals, All traps 100yr Old

1800 1710 1600 1400 1361 1200 1000 800 600 412380 327318 400 229 158163 163 188208 159 85 113112 96 200 15 18 30 0 Diptera Acarina Isoptera Araneae Blattodea Hemiptera Coleoptera Lepidoptera Formicidae Hymenoptera

(Hymenoptera)

Figure 8.4: Radar chart comparing abundances at an age level of taxa with >20 individuals, for all trap types In the CD sticky trap collections (Figure 8.5), Acarina were more abundant in 100yr trees. Blattodea and Formicidae were absent in the 100yr tree collections, but were common in the old trees. The lines moving inward along the axes are irrelevant. They are caused by the absence of Blattodea and Formicidae in the CD sticky traps.

In the funnel collections (Figure 8.6), Acarina were again more abundant in 100yr trees, and Formicidae were again more abundant in the old trees. Lepidoptera and Diptera were slightly more abundant in the 100yr trees, and the reverse was true for Coleoptera and Araneae.

In the hangtrap collections (Figure 8.7), Acarina were again more abundant in 100yr trees, although the trend for Formicidae to be less abundant was not observed. Old trees collected more Blattodea, Coleoptera, Hymenoptera, and Isoptera than 100yr trees. 8) Canopy Arthropods - Results

Abundance for taxa with >20 individuals, CD Sticky traps 100yr Old

500 457 450 400 374 350 300 250 200

150 93 68 71 100 48 21 23 30 50 4 1 2 3 1 7 15 2 5 0 Diptera Acarina Isoptera Araneae Blattodea Hemiptera Coleoptera Lepidoptera Formicidae Hymenoptera

(Hymenoptera)

Abundance for taxa with >20 individuals Funnel traps 100yr Old

600 512 500 400 300 280 230 238227

200 146 120 97 103 95 109 83 77 100 69 70 64 57 9 7 6 0 Diptera Acarina Isoptera Araneae Blattodea Hemiptera Coleoptera Lepidoptera Formicidae Hymenoptera

(Hymenoptera)

Abundance for taxa with >20 individuals Hang traps 100yr Old

800 741 707 700 600 500 400 300 166 200 134 118 97 59 77 66 61 60 100 26 31 12 5 10 16 22 4 9 0 Diptera Acarina Isoptera Araneae Blattodea Hemiptera Coleoptera Lepidoptera Formicidae Hymenoptera (Hymenoptera)

Figures 8.5 (top), 8.6 (middle), 8.7 (bottom): Radar charts comparing abundances at an age level of taxa with >20 individuals, for each trap type. Note the absence of Blattodea and Formicidae in 100yr CD sticky traps.

Tree level comparisons of age using K-W tests (Table 8.4) detected no significant differences in abundance between the age classes for any of the 24 combination of alltraps+3 and alltaxa+5. 8) Canopy Arthropods - Results

Abundance Comparing total catch from 8 100yr and 8 old trees 100yr Old Kruskal Wallis tes mean std min max mean std min max p<0.05? Alltraps All taxa 435.63 210.8716 229 857 375.63 165.4785 213 636 ns Coleoptera 51.5 27.08189 26 108 47.5 19.82062 21 82 ns Diptera 213.75 126.8664 77 401 170.13 69.43535 103 299 ns Hemiptera 40.875 29.396 12 100 39.75 30.94581 7 90 ns Hymenoptera 35.5 20.07842 7 60 46.375 29.56319 12 99 ns Lepidoptera 28.625 15.38958 12 56 19.875 18.03518 7 61 ns

Cdsticky All taxa 92 33.1562 55 156 85.143 34.80627 46 130 ns Coleoptera 6.8571 4.947342 3 17 9.7143 7.134757 4 23 ns Diptera 65.286 29.69127 17 116 53.429 22.18 28 78 ns Hemiptera 4.6 4.827007 1 13 4.2857 2.811541 1 9 ns Hymenoptera 13.286 11.61485 1 35 13.143 7.425824 3 26 ns Lepidoptera 1 011 1.25 0.5 1 2 ns

Funnel All taxa 199.5 144.3765 6 489 141 127.5125 6 339 ns Coleoptera 32.857 19.94516 17 73 18.25 20.65188 1 62 ns Diptera 64 55.44882 1 169 35 37.20983 1 107 ns Hemiptera 34 25.16611 5 80 32.429 27.77803 5 70 ns Hymenoptera 16.75 8.940278 2 34 24.625 22.8969 1 74 ns Lepidoptera 18.167 10.53407 8 32 9.5 10.40673 1 26 ns

Hangtraps All taxa 155.63 68.95534 59 281 160.13 80.55245 63 309 ns Coleoptera 16.75 7.778175 6 28 20.75 9.452891 10 34 ns Diptera 92.625 64.47134 22 225 88.375 53.85944 26 192 ns Hemiptera 8.25 3.84522 3 15 7.625 4.10357 3 14 ns Hymenoptera 8.1429 7.559289 2 24 9.2857 7.696629 2 25 ns Lepidoptera 14.75 6.250714 6 24 12.125 10.00625 3 35 ns

Table 8.4: Abundance between ages compared at a tree level for all traps +3 and all taxa +5

At a placement level, no significant differences were detected in the total abundance collected in any placement (Table 8.5).

Abundances of different trap Traps processed Total collection Mean arthropods per trap Kruskal Wallis tes placements compared between ages 100yr Old Total 100yr Old Total 100yr Old p<0.05?

CDSticky : Total 12 13 1238 643 595 49.5 53.6 45.8 ns Lower crown Trunk 6 7 649 382 267 49.9 63.7 38.1 ns Upper crown Trunk 6 6 589 261 328 49.1 43.5 54.7 ns

Funnel: Total 35 35 2700 1576 1124 38.6 45.0 32.1 ns Lower crown Dead 6 5 104 46 58 9.5 7.7 11.6 ns Lower crown Live 5 8 171 51 120 13.2 10.2 15.0 ns Lower crown Trunk 6 4 1123 817 306 112.3 136.2 76.5 ns Upper crown Dead 5 4 50 26 24 5.6 5.2 6.0 ns Upper crown Live 6 7 132 45 87 10.2 7.5 12.4 ns Upper crown Trunk 7 7 1112 583 529 79.4 83.3 75.6 ns

HangTrap: Total 47 44 2526 1245 1281 27.8 26.5 29.1 ns Lower crown Lower chamber 8 8 426 220 206 26.6 27.5 25.8 ns Lower crown Upper chamber 8 7 516 194 322 34.4 24.3 46.0 ns Mid crown Lower chamber 8 8 492 212 280 30.8 26.5 35.0 ns Mid crown Upper chamber 7 7 388 211 177 27.7 30.1 25.3 ns Upper crown Lower chamber 8 8 391 222 169 24.4 27.8 21.1 ns Upper crown Upper chamber 8 6 313 186 127 22.4 23.3 21.2 ns

Grand Total Total 94 92 6464 3464 3000 34.8 36.9 32.6 ns

Table 8.5: All taxa total abundances between ages compared at a placement level for each trap type.

A more detailed comparison of abundances at a placement level was performed on each major taxonomic group (Table 8.6). Several significant differences were detected, and several trends identified. The collections of Acarina, Diptera, and Plecoptera were more abundant in 100yr trees. Araneae, Blattodea, Coleoptera, Formicidae, and Hymenoptera (excluding Formicidae) were more abundant in older trees. 8) Canopy Arthropods - Results

Mean abundances of different trap placements and orders compared between ages Traps processed Acarina Araneae Blattodea Coleoptera Collembola Dermaptera P-values reported when p<0.20, * label when p<0.05 100yr Old 100yr Old p 100yr Old p 100yr Old p 100yr Old p 100yr Old p 100yr Old p Note that Hym=Hymenoptera & Form=Formicidae CDSticky : Total 12 13 0.33 0.08 0.17 0.23 - 0.08 4.00 5.23 ---- Lower crown Trunk 6 7 0.50 - 0.11 0.17 0.14 -- 5.00 6.71 ---- Upper crown Trunk 6 6 0.17 0.17 0.17 0.33 - 0.17 3.00 3.50 ----

Funnel: Total 35 35 1.97 0.26 2.77 2.37 2.94 2.71 6.57 4.17 0.20 0.06 - 0.09 Lower crown Dead branch 6 5 0.17 - 1.50 1.86 0.33 0.71 3.00 1.57 0.33 - - 0.14 Lower crown Live branch 5 8 -- 1.20 0.50 0.60 0.75 0.80 1.00 - - - 0.25 Lower crown Trunk 6 4 4.86 0.14 0.09 4.29 3.14 2.57 8.00 0.15 17.14 9.86 - 0.14 - - Upper crown Dead branch 5 4 2.00 0.13 0.06 0.60 3.63 *<.01 0.40 0.75 2.40 1.50 ---- Upper crown Live branch 6 7 0.17 0.20 1.50 1.00 0.33 0.60 1.00 2.40 - 0.20 - 0.20 Upper crown Trunk 7 7 3.50 1.50 6.50 3.00 12.67 5.50 12.67 10.00 0.83 - - -

HangTrap: Total 47 44 0.26 0.11 1.26 1.75 0.21 0.36 2.85 3.77 0.06 0.07 - - Lower crown Lower chamber 8 8 - 0.17 0.50 0.67 0.50 0.83 2.88 1.50 - 0.17 - - Lower crown Upper chamber 8 7 0.13 0.13 1.13 1.13 - 0.25 0.17 5.13 6.00 ---- Mid crown Lower chamber 8 8 0.29 - 0.17 1.14 1.57 0.14 0.71 1.86 1.71 ---- Mid crown Upper chamber 7 7 0.25 0.13 1.88 3.00 - 0.13 4.25 6.00 ---- Upper crown Lower chamber 8 8 0.13 0.14 0.88 1.57 *0.05 0.38 0.29 0.75 2.57 0.09 -0.14 - - Upper crown Upper chamber 8 6 0.75 0.13 0.11 2.00 2.25 0.25 0.13 2.50 4.25 0.16 0.38 0.13 - -

Grand Total Total 94 92 0.90 0.16 1.68 1.77 1.20 1.22 4.38 4.13 0.11 0.05 - 0.03 Traps processed Diptera Hemiptera Hym-Form. Hym ex Form Isoptera Lepidoptera 100yr Old 100yr Old p 100yr Old p 100yr Old p 100yr Old p 100yr Old p 100yr Old p

CDSticky : Total 12 13 38.08 28.77 1.92 2.31 - 0.46 7.75 6.62 0.58 1.15 0.17 0.38 Lower crown Trunk 6 7 27.33 27.86 2.50 2.29 - 3.98 *0.04 6.00 7.00 0.83 1.43 0.17 0.29 Upper crown Trunk 6 6 48.83 29.83 0.17 1.33 2.33 - 2.00 0.18 9.50 6.17 0.33 0.83 0.17 0.50

Funnel: Total 35 35 14.63 8.00 6.80 6.49 0.77 1.51 3.06 4.11 0.20 0.17 3.11 1.63 Lower crown Dead 6 5 1.17 2.57 0.33 1.00 0.33 2.75 0.05 0.33 1.86 - - - 0.14 Lower crown Live 5 8 0.20 0.50 - 1.00 *0.04 0.20 0.500.13 0.20 0.50 0.19 0.20 0.25 - - Lower crown Trunk 6 4 20.86 18.43 16.29 17.57 1.71 8.32 4.57 10.43 0.14 0.14 6.29 4.57 Upper crown Dead 5 4 1.60 2.75 1.60 1.63 0.20 3.51 0.15 0.40 2.88 - 0.25 - 0.13 Upper crown Live 6 7 1.33 4.20 0.50 1.00 0.83 0.71 1.17 0.60 - 0.20 0.17 - Upper crown Trunk 7 7 56.83 22.00 0.09 18.50 18.75 1.00 3.86 10.17 7.50 0.11 0.83 0.25 10.67 5.75

HangTrap: Total 47 44 15.77 16.07 1.40 1.39 0.23 0.32 0.98 1.55 0.09 0.20 2.51 2.20 Lower crown Lower chamber 8 8 12.38 10.50 1.63 1.17 0.50 1.75 1.00 3.83 *0.03 0.13 0.33 3.75 1.33 Lower crown Upper chamber 8 7 15.00 9.38 0.17 1.50 1.00 0.13 0.52 1.25 0.250.12 0.25 0.63 2.00 0.50 *0.04 Mid crown Lower chamber 8 8 18.71 15.71 1.86 1.29 0.14 0.38 1.00 0.57 - - 4.57 3.00 Mid crown Upper chamber 7 7 14.88 18.13 1.13 1.38 0.13 1.77 0.63 2.13 0.07 0.13 0.25 1.50 1.75 Upper crown Lower chamber 8 8 15.63 28.86 0.10 1.75 3.43 0.13 0.38 1.50 1.29 - - 2.63 6.71 Upper crown Upper chamber 8 6 18.38 14.00 0.63 0.25 0.38 0.35 0.50 1.63 - - 0.88 0.38

Grand Total Total 94 92 18.19 14.79 3.48 3.46 0.40 0.79 2.62 3.24 0.19 0.33 2.44 1.73 Traps processed Neuroptera Orthoptera Plecoptera Pseudo- Psocoptera Trichoptera 100yr Old 100yr Old p 100yr Old p 100yr Old p 100yr Old p 100yr Old p 100yr Old p

CDSticky : Total 1213------0.08- -- Lower crown Trunk 6 7 ------0.17 - - - Upper crownTrunk 6 6 ------

Funnel: Total 35 35 0.03 - 0.29 0.20 1.29 0.09 0.06 - ---- Lower crown Dead 6 5 -- - 0.29 ------Lower crownLive 5 8 - - 0.60------Lower crown Trunk 6 4 -- 0.14 0.43 4.43 0.43 0.08 ------Upper crown Dead 5 4 -- 0.20 - -- 0.40 - ---- Upper crownLive 6 7 - - 0.50------Upper crown Trunk 7 7 0.17 - 0.33 0.50 2.33 - *0.03 ------

HangTrap: Total 47 44 0.02 ------0.11 0.02 - 0.07 Lower crown Lower chamber 8 8 ------Lower crown Upper chamber 8 7 ------0.25 - - 0.13 Mid crown Lower chamber 8 8 ------0.14 Mid crown Upper chamber 7 7 ------0.13 - - - Upper crown Lower chamber 8 8 0.13 ------0.14 Upper crown Upper chamber 8 6 ------0.25 0.13 - - Grand Total Total 94 92 0.02 - 0.11 0.08 0.48 0.03 0.02 - 0.06 0.01 - 0.03

Table 8.6: Abundances compared between ages at a placement level for each major taxonomic grouping. “Hym- Form” = Hymenoptera: Formicidae; “Hym ex Form” = Hymenoptera excluding Formicidae; and “Pseudo”= Pseudoscorpionidae. 8.5 Richness results

A total of 233 RTU were collected in the old trees, and 226 in the 100yr trees. For the CD sticky traps and hangtraps, richness was higher in the old trees (Figure 8.8). The reverse was true for the funnels. 8) Canopy Arthropods - Results

Figure 8.8: Richness in each age class, by trap type Coleoptera were the most speciose taxa. One third (113) of all encountered RTU were beetles. Diptera, Hemiptera and Hymenoptera each comprised approximately one sixth of the RTU catalogue, with 55, 47, and 44 RTU. 20 lepidopteran RTU were collected.

When pooling all trap types, more Diptera, Coleoptera, and Hymenoptera species were found in old trees, and more Hemiptera in 100yr trees (Figure 8.9). The contribution of each taxa to the total RTU catalogue for each tree was similar.

8) Canopy Arthropods - Results

RTU richness for the five largest taxa 100yr Old All traps

90 79 80 77 70 60 49 50 41 39 35 35 40 32 30 20 18 18 10 0

ra ra e era tera e pt p pt o o o le Dipt id p Hemipteramen e Co y L H

Figure 8.9: Radar chart comparing richness between ages of the five most prominent taxa, for all trap types Comparison of the radar charts for each trap types shows that CD sticky and hangtrap fauna were richer in old trees, but the opposite tendency was seen in the funnel traps.

In the CD sticky trap collections, all of the five most prominent taxa were richer in the old trees, except Hymenoptera (Figure 8.10). The differences in richness of Coleoptera, Diptera, and Hemiptera were the most pronounced.

Funnel traps exhibited the opposite trend: all five taxa were more species rich in the 100yr trees (Figure 8.11).

The hangtrap collection (Figure 8.12) showed a similar pattern to the CD sticky traps. All taxa except Lepidoptera were more species rich in the old trees than the 100yr trees. RTU richness for Lepidoptera was equal. 8) Canopy Arthropods - Results

RTU richness for the five largest taxa CD Sticky traps 100yr Old

30 25 25 21 20 19 19 17 17 15 11 10 7 4 5 3

0

ra te tera tera p p p o leoptera Di id p Hemi e Co L Hymenoptera

RTU richness for the five largest taxa Funnel traps 100yr Old

50 45 45 40 35 35

30 26 25 20 20 20 19 15 16 15 10 10 8 5 0

ra ra a e tera tera ter ipte p Dip no op em e pid H e Coleopt ym L H

RTU richness for the five largest taxa Hangtraps 100yr Old

60 53 54 50 40 40 31 30 24 25 24 19 20 17 17

10

0

a ra ra e e pt opter Diptera miptera o e e id H Col Lep Hymenopt

Figures 8.10 (top), 8.11 (middle), 8.12 (bottom): Radar charts comparing richness at an age level for the five most prominent taxa for each trap type

8) Canopy Arthropods - Results

Analysis at a tree level (Table 8.7) showed no significant difference in the total RTU count for all traps +3 and all taxa +5.

RTU count Comparing total catch from 8 100yr and 8 old trees 100yr Old Kruskal Wallis test mean std min max mean std min max p<0.05? Alltraps All taxa 88.50 22.20 52 124 83.38 18.06289 59 113 ns Coleoptera 21.63 6.63 13 32 21.13 5.139136 13 28 ns Diptera 17.75 5.04 11 26 17.75 4.399675 11 24 ns Hemiptera 13.25 4.59 5 20 11.75 4.399675 6 17 ns Hymenoptera 12.50 4.17 6 17 14.00 5.398412 5 22 ns Lepidoptera 8.25 2.60 4 13 6.13 2.167124 3 10 ns

Cdsticky All taxa 20.86 5.21 14 31 24.00 6.855655 16 35 ns Coleoptera 3.86 1.77 2 6 5.29 2.13809 3 9 ns Diptera 8.00 1.63 5 10 8.57 2.225395 6 12 ns Hemiptera 1.43 1.13 0 3 2.43 1.511858 0 4 ns Hymenoptera 4.57 2.44 1 8 4.57 1.618347 2 7 ns Lepidoptera 0.14 0.38 0 1 0.14 0.377964 0 1 ns

Funnel All taxa 46.50 20.58 4 67 35.13 20.53873 4 62 ns Coleoptera 12.14 4.38 5 17 7.75 5.338539 1 16 ns Diptera 6.63 3.25 1 11 4.88 2.799872 1 10 ns Hemiptera 9.57 3.60 5 14 6.71 3.817254 2 12 ns Hymenoptera 6.50 2.93 1 11 6.50 4.242641 1 13 ns Lepidoptera 2.50 2.14 0 6 1.50 1.069045 0 3 ns

Hangtraps All taxa 44.13 12.14 29 69 47.50 10.87592 33 62 ns Coleoptera 11.25 3.92 6 17 12.25 4.803273 8 22 ns Diptera 11.75 5.09 8 21 12.25 3.535534 7 17 ns Hemiptera 5.13 1.64 3 8 5.63 2.386719 3 9 ns Hymenoptera 4.86 2.34 2 8 6.38 2.924649 1 10 ns Lepidoptera 5.88 2.47 4 11 4.75 2.12132 1 7 ns

Table 8.7: RTU richness compared between ages at a tree level for all traps +3 and all taxa +5

At a placement level, significant (p<0.05) differences were detected for some combinations of all taxa +5 and placement (Table 8.8). In the upper live branch funnels, the total RTU count was significantly higher in old trees. The collection from old trees was significantly higher in upper dead branch funnels for Hemiptera, and in the upper bottle of the uppermost hangtrap for Hymenoptera. Results for Lepidoptera were contradictory, with one hangtrap placement significantly richer in 100yr trees, and one hangtrap placement richer in old trees.

When viewing results where p<0.20, additional trends can be identified. Total RTU count was higher in old trees in two hangtrap and two funnel placements. Coleoptera and Hymenoptera both tended to be richer in old trees. 8) Canopy Arthropods - Results

Mean richness of different trap RTU count- All Animals Coleoptera Diptera placements and orders compared between ages 100yr std Old std p 100yr std Old std p 100yr std Old std p P-values reported when p<0.20, * label when p<0.05 CDSticky Upper crown Trunk 13.33 4.97 15.71 6.47 3.00 2.37 3.71 2.14 4.83 0.41 6.14 2.27 0.12 Lower crown Trunk 12.67 3.78 12.67 3.56 1.83 0.41 2.50 0.84 0.11 6.33 1.75 5.17 2.32

Funnel: Upper crown Live 4.33 2.34 7.00 2.83 *0.03 1.50 1.05 1.57 1.13 0.67 0.82 1.14 1.46 Upper crown Dead 3.20 0.45 5.00 2.16 0.11 0.80 0.84 1.00 0.82 0.20 0.45 0.50 0.58 Upper crown Trunk 23.14 #### 20.86 9.99 5.00 3.65 3.71 2.50 4.57 2.30 3.29 0.76 Lower crown Live 7.20 4.44 7.25 4.86 1.60 1.82 1.50 1.93 1.40 1.14 1.25 1.04 Lower crown Dead 5.17 3.31 7.80 7.53 0.67 0.82 1.80 3.03 1.17 0.98 2.20 1.92 Lower crown Trunk 34.00 #### 21.25 13.30 7.17 4.31 3.25 2.75 0.16 4.50 1.05 4.00 1.83

HangTrap: Upper crown Up bottle 8.75 7.13 10.17 4.26 0.88 1.46 1.50 1.87 3.38 3.11 2.83 1.17 Upper crown Low bottle 14.13 5.99 11.50 3.66 3.88 2.75 3.50 1.07 5.25 2.43 4.50 1.20 Mid crown Up bottle 9.86 5.08 9.57 3.55 1.71 1.70 1.57 1.51 3.00 2.16 2.86 1.35 Mid crown Low bottle 11.75 4.13 14.63 5.01 0.19 3.50 1.51 4.13 2.42 4.63 2.00 4.63 1.69 Lower crown Up bottle 9.00 4.81 13.57 5.00 0.09 0.75 0.71 2.00 1.63 0.09 3.38 1.85 3.86 2.04 Lower crown Low bottle 10.50 6.91 11.88 5.14 2.13 2.85 3.63 3.07 0.12 4.25 3.24 4.75 1.83 Hemiptera Hymenoptera Lepidoptera 100yr std Old std p 100yr std Old std p 100yr std Old std p

CDSticky Upper crown Trunk 0.83 0.98 1.86 0.90 0.10 2.83 1.83 2.86 2.19 0.17 0.41 0.14 0.38 Lower crown Trunk 0.83 0.75 1.00 0.00 2.83 1.83 2.33 1.03 0.17 0.41 0.50 0.55

Funnel: Upper crown Live 0.33 0.52 0.57 0.79 0.67 0.52 1.57 1.13 0.14 0.00 0.00 0.14 0.38 Upper crown Dead 0.00 0.00 1.00 0.82 *0.04 0.40 0.55 1.00 0.00 0.10 0.00 0.00 0.00 0.00 Upper crown Trunk 5.14 3.29 4.43 3.05 2.43 1.62 3.86 2.54 0.86 0.69 1.00 0.82 Lower crown Live 1.20 1.10 0.88 1.13 0.60 0.89 1.38 1.19 0.00 0.00 0.13 0.35 Lower crown Dead 0.33 0.52 0.80 0.84 1.33 1.37 1.40 1.14 0.17 0.41 0.00 0.00 Lower crown Trunk 6.83 4.75 4.50 2.65 5.33 2.73 4.00 3.74 2.50 1.87 1.00 1.15

HangTrap: Upper crown Up bottle 0.75 0.71 1.00 1.10 0.75 0.89 2.00 0.89 *0.04 2.25 1.83 1.33 1.03 Upper crown Low bottle 1.25 1.04 0.88 0.64 1.00 1.07 0.63 0.74 1.38 1.30 0.25 0.71 *0.03 Mid crown Up bottle 1.14 0.69 1.14 1.35 0.71 0.76 0.71 0.76 2.14 1.77 1.43 0.98 Mid crown Low bottle 1.00 0.93 1.25 1.28 0.63 0.74 1.88 1.73 0.09 0.75 0.89 1.00 1.07 Lower crown Up bottle 1.50 1.77 2.43 1.90 1.13 1.25 1.29 0.76 1.13 0.64 2.29 1.11 *0.04 Lower crown Low bottle 0.63 1.06 0.25 0.46 0.75 0.89 1.25 1.49 0.75 0.89 0.38 0.52

Table 8.8: RTU richness compared between ages at a placement level for all taxa +5 8.6 Community structure results

Visual assessment of the rank abundance plots generated for all traps +3 point to a lognormal species abundance model (Magurran, 2004) (See section 7.5, aspect 3).

A consistent trend for the 100yr collections to exhibit a steeper slope can be seen for all four plots (Figures 8.13, 8.14, 8.15, 8.16). This trend was most obvious for the sticky traps (Figure 8.14), and least obvious for the funnels (Figure 8.15).

A steeper slope indicates a higher dominance of the most abundant taxa, and therefore a less evenly abundant community structure. Consequently, the rank/abundance plots exhibit a less diverse community structure in the 100yr old collections.

The length of the curve along the x-axis indicates the total richness of the community. The flattened level of the tail at the right end of the curve represents the animals only encountered once. The next flat level above represents the animals encountered exactly twice. Because the Chao 1 true richness estimator is derived from these number on the ratio, the relative length of the singletons level is a sign of high 8) Canopy Arthropods - Results richness. For all traps pooled, CD sticky traps, and funnel traps, the length of the singletons tail was longer in old trees.

Similarly, the old curves are slightly above the 100yr curves, visually suggesting a potentially longer tail, and thus true richness, along the right axis for the old curve.

Whittaker Rank/Abundance Plot

1 Rel Abun 100yr Rel Abun Old 0.1

0.01 `

0.001 Old

Relative Abundance Relative 100yr 0.0001 0 50 100 150 200 250 Species Rank

Figure 8.13: Whittaker rank/ abundance plot for all traps

8) Canopy Arthropods - Results

Rank/Abundance Plot for CD Sticky traps

1 Rel Abun 100yr Rel Abun Old 0.1

0.01 100yr line ends (hidden)

Relative Abundance 0.001 0 20 40 60 80 100 Species Rank

Rank/Abundance Plot for Funnel traps

1 Rel Abun 100yr 0.1 Rel Abun Old

0.01 Old 0.001

Relative Abundance 0.0001 100yr 0 20 40 60 80 100 120 140 160 Species Rank

Rank/Abundance Plot for Hangtraps

1 Rel Abun 100yr 0.1 Rel Abun Old

0.01

0.001

!00yr line ends (hidden)

Relative Abundance 0.0001 0 50 100 150 200 Species Rank

Figures 8.14 (top), 8.15 (middle), 8.16 (bottom): Whittaker rank/ abundance plots for each trap type

8) Canopy Arthropods - Results

8.7 Diversity results

Figure 8.17: -ln (Simpson’s diversity) in each age class, by trap type At an age level, –ln(D) was higher for old trees (2.3402) than 100yr trees (2.3331). Because diversity indices have no intrinsic scale, these values are meaningless outside of their comparisons.

The diversity index was greater in old trees for funnels and hangtraps, and greater in 100yr trees for CD sticky traps (Figure 8.17). Because richness was higher in the old trees for CD sticky traps, evenness in these trees must have been lower to result in a lower diversity index (a composite of richness and evenness) at the age level. However, at the tree and placement level, CD sticky traps had a higher diversity in the old trees. Conversely, because richness was lower in the old trees for the funnel traps, evenness must have been higher to results in a higher diversity index. 8) Canopy Arthropods - Results

Radar charts at an age level show a higher diversity in the old trees for all 24 combinations (Figure 8.19, 8.20, 8.21) of all traps+3 and the five most prominent taxa with three exceptions: Diptera in the CD sticky traps, and Lepidoptera in the pooled traps and the funnel traps. This consistent trend reflects a quantitative index of the observations made of the rank/abundance graphs, above. In all cases, the Coleoptera fauna was more diverse in the old trees (Figure 8.18).

-ln(D) for the five largest taxa 100yr Old All traps

2.34 2.32 2.312.31 2.302.30 2.29 2.3 2.28 2.28 2.26

2.26 2.24

2.24 2.22 2.22 2.21 2.2 2.18 2.16 2.14

a a ra e tera pter ipt optera leopter Di m n idop He Co Lep Hyme

Figure 8.18: Radar charts comparing –ln(Simpsons D) between ages of the five most prominent taxa for all traps pooled 8) Canopy Arthropods - Results

-ln(D) for the five largest taxa CD Sticky traps 100yr Old

2.28 2.3 2.26 2.242.25 2.24 2.25 2.23 2.2 2.15 2.12

2.1 2.05 2.05 2.02 2 1.98 1.95 1.9 1.85 1.8

ra ra ra ra tera te e pte Dip o opt leop en d Hemipte Co Lepi Hym

-ln(D) for the five largest taxa Funnel traps 100yr Old

2.3 2.272.27 2.252.26 2.25 2.21 2.21

2.2 2.17 2.16 2.15 2.15 2.10 2.1 2.05 2

a a ra ra ter ter pte o pte Dip mip o en id He m ep Coleoptera L Hy

-ln(D) for the five largest taxa Hangtrap 100yr Old

2.35 2.32 2.32 2.32 2.31 2.30 2.31 2.3 2.252.26 2.25 2.2

2.14 2.15 2.12 2.1 2.05 2

ra e ptera iptera o leopt Dipteram en pidoptera He m e Co L Hy

Figures 8.19 (top), 8.20 (middle), 8.21 (bottom): Radar charts comparing –ln(Simpsons D) at an age level for the five most prominent taxa for each trap type K-W test comparisons a tree level showed a significantly higher diversity in the old trees for all taxa caught in the CD sticky traps (Table 8.9). 8) Canopy Arthropods - Results

-ln Simpsons D Comparing total catch from 8 100yr and 8 old trees 100yr Old Kruskal Wallis test mean std min max mean std min max p<0.05? Alltraps All taxa 1.3384 0.023814 1.297923 1.36008 1.3465 0.010861 1.327124 1.357956 ns Coleoptera 1.2459 0.090678 1.102921 1.328181 1.29279 0.065557 1.135112 1.336362 ns Diptera 1.249 0.029716 1.205376 1.290527 1.24995 0.031412 1.193986 1.300258 ns Hemiptera 1.2489 0.046772 1.152101 1.288637 1.22367 0.069493 1.091314 1.295502 ns Hymenoptera 1.1765 0.084084 1.062368 1.297709 1.23817 0.050557 1.169064 1.301811 ns Lepidoptera 1.1747 0.062527 1.08825 1.263411 1.11007 0.108041 0.944462 1.217876 ns

Cdsticky All taxa 1.2397 0.025759 1.185687 1.260325 1.27166 0.031426 1.221786 1.309292 * Coleoptera 2.5372 0.114227 2.300395 2.647538 2.59204 0.07089 2.428025 2.64549 ns Diptera 2.4873 0.081852 2.300395 2.560212 2.5152 0.04627 2.428025 2.568687 ns Hemiptera 0.8368 0.152488 0.653926 0.980829 0.90647 0.188405 0.693147 1.163151 ns Hymenoptera 0.857 0.234313 0.544727 1.141172 1.07902 0.075007 0.944462 1.146403 ns Lepidoptera 0.3466 0.490129 0 0.693147 0.17329 0.346574 0 0.693147 ns

Funnel All taxa 1.282 0.069144 1.141172 1.343267 1.29335 0.081025 1.098612 1.346948 ns Coleoptera 2.62 0.112308 2.363981 2.703299 2.59799 0.113226 2.37331 2.710348 ns Diptera 2.4822 0.069849 2.363981 2.554505 2.45993 0.081476 2.340393 2.562153 ns Hemiptera 1.1825 0.034564 1.128181 1.221845 1.09714 0.13561 0.867501 1.240693 ns Hymenoptera 1.0441 0.169 0.693147 1.280934 0.98486 0.16787 0.693147 1.171183 ns Lepidoptera 0.6734 0.262699 0.41058 1.045368 0.48336 0.346126 0 0.998529 ns

Hangtraps All taxa 1.2647 0.074576 1.118483 1.324569 1.28548 0.039657 1.227777 1.3302 ns Coleoptera 1.7921 0.145937 1.433531 1.858033 1.83516 0.029739 1.77413 1.864999 ns Diptera 1.0981 0.082756 0.998032 1.218395 1.10837 0.077146 1.003289 1.192913 ns Hemiptera 1.1406 0.072905 1.019025 1.255266 1.10181 0.125863 0.820981 1.237874 ns Hymenoptera 1.0448 0.118751 0.900787 1.203973 1.08897 0.175985 0.693147 1.247586 ns Lepidoptera 0.9531 0.12804 0.71744 1.148384 0.87264 0.234495 0.470004 1.125641 ns

Table 8.9: -ln(d) compared between ages at a tree level for all traps +3 and all taxa +5

Comparing at a placement level, several significant differences were detected. At p<0.05, all statistically significant differences indicated a more diverse fauna in old trees (Table 8.10).

Within the upper crown, sticky trap, live branch funnel, and trunk funnel were significantly more diverse in old trees for all taxa.

The Coleoptera collection was significantly more diverse in old trees in several placements in the lower crown: the sticky trap, dead branch funnel, and both chambers of the hangtrap. Diptera in the upper sticky trap were more diverse in old trees. 8) Canopy Arthropods - Results

-ln (Simpsons D) for different -ln Simps D; All Taxa Coleoptera Diptera placements and orders compared between ages 100yr std Old std p 100yr std Old std p 100yr std Old std p P-values reported when p<0.20, * label when p<0.05 CDSticky Upper crown Trunk 1.55 0.36 2.00 0.25 *0.04 0.88 0.61 0.93 0.66 0.84 0.31 1.14 0.08 *0.02 Lower crown Trunk 1.53 0.36 1.78 0.41 0.49 0.26 0.80 0.18 *0.04 1.31 0.41 1.16 0.49

Funnel: Upper crown Live 1.20 0.21 1.53 0.33 0.06 0.42 0.38 0.48 0.55 0.23 0.40 0.46 0.58 Upper crown Dead 1.05 0.38 1.50 0.46 0.23 0.40 0.23 0.40 0.00 - 0.00 0.00 Upper crown Trunk 2.25 0.42 2.36 0.42 *0.04 0.75 0.49 0.88 0.54 1.12 0.52 0.97 0.16 Lower crown Live 1.59 0.70 1.36 0.44 0.75 0.65 0.60 0.82 0.42 0.50 0.34 0.38 Lower crown Dead 1.24 0.76 1.70 0.66 0.20 0.34 1.02 0.46 0.14 0.20 0.44 0.78 0.63 0.14 Lower crown Trunk 2.24 0.47 2.33 0.56 1.26 0.75 0.77 0.18 0.69 0.30 0.95 0.30

HangTrap: Upper crown Up bottle 1.35 0.47 1.63 0.30 0.66 0.70 0.77 0.81 0.60 0.39 0.61 0.34 Upper crown Low bottle 2.05 0.48 2.14 0.23 1.02 0.75 1.02 0.37 1.14 0.31 1.26 0.23 Mid crown Up bottle 1.48 0.58 1.43 0.49 0.69 0.69 0.61 0.63 0.72 0.31 0.60 0.34 Mid crown Low bottle 1.95 0.43 1.94 0.46 1.09 0.53 1.33 0.36 1.07 0.36 0.94 0.46 Lower crown Up bottle 1.54 0.65 1.45 0.57 0.14 0.31 0.55 0.44 0.13 0.67 0.53 0.47 0.42 Lower crown Low bottle 1.59 0.62 1.92 0.26 0.40 0.82 0.94 0.76 0.09 0.77 0.64 1.02 0.33 Hemiptera Hymenoptera Lepidoptera 100yr std Old std p 100yr std Old std p 100yr std Old std p

CDSticky Upper crown Trunk 0.22 0.24 0.45 0.45 0.70 0.42 1.00 0.40 0.00 - 0.00 - Lower crown Trunk 0.17 0.35 0.00 0.00 0.80 0.41 0.40 0.36 0.00 - 0.00 0.00

Funnel: Upper crown Live 0.00 0.00 0.20 0.34 0.00 0.00 0.23 0.40 - - 0.00 - Upper crown Dead - - 0.23 0.40 0.00 ------Upper crown Trunk 1.04 0.58 0.85 0.71 0.45 0.47 0.83 0.61 0.19 0.09 0.21 0.17 0.24 Lower crown Live 0.25 0.49 0.35 0.45 0.00 0.00 0.35 0.40 - - 0.00 - Lower crown Dead 0.00 0.00 0.20 0.34 0.35 0.49 0.35 0.49 0.00 - - - Lower crown Trunk 1.12 0.71 1.01 0.52 1.18 0.59 0.71 0.62 0.36 0.30 0.43 0.37

HangTrap: Upper crown Up bottle 0.14 0.31 0.25 0.49 0.00 0.00 0.31 0.36 0.70 0.58 0.36 0.51 Upper crown Low bottle 0.37 0.41 0.10 0.24 0.18 0.21 0.41 0.00 0.00 0.44 0.53 0.47 - Mid crown Up bottle 0.19 0.31 0.25 0.57 0.23 0.40 0.23 0.40 0.59 0.60 0.35 0.38 Mid crown Low bottle 0.16 0.40 0.55 0.53 0.23 0.40 0.46 0.52 0.24 0.33 0.25 0.43 Lower crown Up bottle 0.62 0.61 0.77 0.62 0.36 0.51 0.23 0.36 0.18 0.31 0.63 0.48 0.07 Lower crown Low bottle 0.37 0.63 0.00 0.00 0.00 0.00 0.30 0.42 0.32 0.37 0.00 0.00

Table 8.10: -ln(D) compared between ages at a placement level for all taxa +5

8.8 Species accumulation and estimated richness results

Figure 8.22: Sampling adequacy: Observed richness presented as a percentage of the true richness estimated by Chao 1, comparing ages for each trap type. Cryptic RTU are those estimated to exist but not collected. 8) Canopy Arthropods - Results

Pooling traps, true richness was estimated to be slightly higher for the old trees. True richness was estimated to be higher in the old trees for CD sticky, and higher in the 100yr trees for funnels. Hangtraps were estimated to have an equal richness between ages (Figure 8.22).

Comparing between traps, the estimated true richness was highest for the hangtraps, followed by the funnels and then the CD sticky traps.

CD sticky traps showed the poorest performance as measured by sampling adequacy. Due to the high estimated true richness in the old trees, sampling in them was only two thirds as adequate as in the 100yr trees. Adequacy inequality was less pronounced for the other trap types. Pooling all traps, sampling adequacy was greater in the 100yr trees than the old trees.

Species accumulation curves output from EstimateS (Colwell, 2004) showed different trends for different trap types. Similar to the results for richness, funnel results did not agree with that of the CD sticky traps and the hangtraps.

100yr Species observed 400 100yr Chao 1 Estimator 350 Old Species observed 300 250 Old Chao 1 Estimator 200 150 100 50 Cumulative Species 0 0 500 1000 1500 All trapped individuals

Figure 8.23: Species observed accumulation curves with Chao 1 estimator individuals for all traps When comparing the curves for all traps pooled, old trees showed a slightly higher curve than 100yr trees, indicating a higher observed richness and a higher estimate of true richness (Figure 8.23). Observation of the last two points of the Chao 1 estimator curves for each age class hints at a flattening of the 100yr curve relative to the old curve. This implies a higher overall richness in the old trees.

8) Canopy Arthropods - Results

100yr Species observed 250 100yr Chao 1 Estimator 200 Old Species observed Old Chao 1 Estimator 150

100

50 Cumulative Species 0 0 50 100 150 200 CD Sticky trapped Individuals

Figure 8.24: Species observed accumulation curves with Chao 1 estimator individuals for CD sticky traps Curves from the resampled CD sticky traps show a very strong divergence in the Chao 1 estimators, with the old trees adopting a much steeper slope than the 100yr trees (Figure 8.24). This indicates much higher species richness in the community of animals prone to being caught in sticky traps, and that the sampling effort was not adequate for the Chao 1 estimator to asymptote. The presence of a large number of animals only encountered once in the old sticky traps may be responsible for the abrupt rise. The curves for species observed show a slightly higher value for old trees, but the divergence is not visible.

250 100yr Species observed 100yr Chao 1 Estimator 200 Old Species observed Old Chao 1 Estimator 150

100

50 Cumulative Species Cumulative 0 0 200 400 600 800 Funnel trapped Individuals

Figure 8.25: Species observed accumulation curves with Chao 1 estimator individuals for funnel traps The funnel trap curves show an opposite pattern, with 100yr trees showing a higher curve for both the species observed and the Chao 1 estimator (Figure 8.25). This indicates a richer community of animals prone to being trapped in funnels in the 100yr trees. 8) Canopy Arthropods - Results

100yr Species observed 300 100yr Chao 1 Estimator 250 Old Species observed 200 Old Chao 1 Estimator 150 100 50

Cumulative Species Cumulative 0 0 200 400 600 800

Hangtrapped Individuals

Figure 8.26: Species observed accumulation curves with Chao 1 estimator individuals for hang traps Hangtrap species accumulation curves show a flattening of the estimator and species observed curves in comparison to the other trap types (Figure 8.26). The estimator appears to asymptote at around 250 RTU, suggesting that the trapping regime was adequate to estimate the true richness. This does not mean the trapping regime was adequate to actually collect the true richness.. The species observed curve does not flatten as distinctly as the estimator.

Comparison of ages using the hangtrap curves presents a less clear picture than the other trap types. For species observed, old trees are show a slightly higher, richer curve.

The Chao 1 estimator shows an unexpected result. For most of the curve, the old tree curve is higher and steeper than the 100yr curve. However, it appears to flatten out by dipping and rising again, with the final points showing the old curve below the 100yr curve.

To confirm this was not an anomalous result, EstimateS analysis was redone with 8000 resampling runs. No difference in output was detected.

It is possible that if further sampling added another point to the 100yr curve, it would also dip down. The Chao 1 estimator results for hangtraps are ambiguous. For the other trap types, the trend of Chao 1 mirrored the trends of species observed. This may be true for hangtraps. Regardless of whether the Chao 1 results are integrated or not, the overall inclination of hangtrap curve output indicates a richer fauna in the old trees.

8) Canopy Arthropods - Results

8.9 Distinctness results

8.9.1 RTU overlap

Approximately half of all encountered RTU were found only in one age class, and half were found in both age classes (Figure 8.27). Slightly more RTU were found in old trees only (82) than in 100yr trees only (75).

Venn diagram RTU Overlap illustrating RTU overlap between tree ages for all trap types pooled. 100 year Old-growth Size of circles is proportional to RTU richness. Overlap in circles is not proportional 75 155 82

Figure 8.27:Venn diagram showing overlap between tree ages for all traps pooled For both hangtraps and CD sticky traps, more RTU were collected in old trees only than in 100yr trees only (Figure 8.28). The results for the funnel traps were different. Almost twice as many RTU were found only in 100yr funnel traps (62) than were found only in old funnel traps (35).

Venn diagram illustrating RTU 100 year RTU Overlap Old-growth overlap between tree ages for three CD 12 27 42 41 13 trap types. Sticky traps traps Size of circles is proportional to RTU richness.

Funnel 35 62 81 36 35 Crawl traps traps

Hang 47 62 100 79 44 traps traps traps

Figure 8.28: Venn diagram showing overlap between tree age for each trap type. Trap count includes hangtrap chambers as separate traps.

8) Canopy Arthropods - Results

8.9.2 MRPP results

MRPP analysis at a tree level detected no significant differences in the composition of RTU between the age classes (Table 8.11).

MRPP comparison Comparing total catch from 8 100yr and 8 old trees

MRPP p-value P <0.05? Alltraps All taxa 0.806 ns Coleoptera 0.374 ns Diptera 0.604 ns Hemiptera 0.639 ns Hymenoptera 0.143 ns Lepidoptera 0.104 ns

Cdsticky All taxa 0.636 ns Coleoptera 1.000 ns Diptera 0.936 ns Hemiptera 0.907 ns Hymenoptera 0.767 ns Lepidoptera 0.587 ns

Funnel All taxa 0.267 ns Coleoptera 0.567 ns Diptera 0.248 ns Hemiptera 1.000 ns Hymenoptera 0.130 ns Lepidoptera 0.453 ns

Hangtrap All taxa 0.979 ns Coleoptera 0.652 ns Diptera 0.742 ns Hemiptera 0.411 ns Hymenoptera 0.273 ns Lepidoptera 0.131 ns

Table 8.11: MRPP comparison of composition between ages at a tree level At a trap level, several significant differences in RTU composition were identified (Table 8.12). The collection of all funnel traps on live branches was significantly different between ages. This trend appeared to be driven by the Hymenoptera, and to a lesser extent, the Diptera. The lower chambers of all hangtraps pooled collected a significantly different coleopteran fauna. No significant differences in composition were detected in the fauna of sticky or funnel traps on the trunks. Analysis of lepidopteran fauna was not always possible due to their lower abundances. 8) Canopy Arthropods - Results

MRPP comparing a defined pool of sorted traps between ages P-values reported when <0.20; * label when p<0.05

All Animals Coleoptera Diptera Hemiptera Hymenoptera Lepidoptera p<.20? p<.20? p<.20? p<.20? p<.20? p<.20? CDSticky : All traps pooled ns ns ns ns ns ns Funnel: All traps pooled ns ns ns ns ns ns Trunk traps only ns ns ns ns ns ns Live branch traps only * <.01 ns 0.20 ns * 0.01 (inv) Dead branch traps only ns 0.12 ns ns ns (inv) Live and Dead pooled * <.01 ns 0.08 ns * 0.01 (inv) HangTrap: Both chambers pooled ns ns ns ns ns ns Upper chambers only ns ns ns ns ns ns Lower chambers only 0.16 * 0.03 0.20 0.12 0.06 ns All Trap Types All Positions Pooled ns ns ns ns ns (inv) Pooled

Table 8.12: MRPP comparison of composition between ages compared at a trap level

At a placement level, several significant differences in composition were detected (Table 8.13). The total RTU collection for all taxa was significantly different in two placements, with two more significant at p<0.10. The dipteran community in the lower chamber of the uppermost hangtrap was different between ages.

MRPP comparing 8 100yr and 8 old trees by placement All Orders Coleoptera Diptera Hemiptera Hymenoptera Lepidoptera P-values reported when <0.20; * label when p<0.05 p<.20? p<.20? p<.20? p<.20? p<.20? p<.20? CDSticky : Total: ns ns ns ns ns ns Lower crown Trunk ns ns ns ns ns ns Upper crown Trunk ns ns ns ns ns (inv) Funnel: Total: ns 0.19 ns ns ns ns Lower crown Dead ns 0.07 ns (inv) ns (inv) Lower crown Live *0.01 ns ns ns 0.11 (inv) Lower crown Trunk 0.09 ns 0.13 ns ns ns Upper crown Dead 0.19 ns (inv) (inv) 0.13 (inv) Upper crown Live 0.08 ns ns (inv) 0.19 (inv) Upper crown Trunk ns ns ns ns ns ns HangTrap: Total: ns ns ns ns ns ns Lower crown Lower chamber ns ns ns ns ns 0.17 Lower crown Upper chamber ns ns 0.10 ns ns ns Mid crown Lower chamber ns 0.10 ns ns 0.19 ns Mid crown Upper chamber ns ns ns ns 0.06 ns Upper crown Lower chamber *0.03 ns *0.04 ns 0.19 ns Upper crown Upper chamber ns ns 0.10 ns ns ns All Trap Types All Positions ns ns ns ns ns ns Pooled Pooled

Table 8.13: MRPP comparison of composition between ages compared at a placement level

8) Canopy Arthropods - Results

8.9.3 NMS ordination results

Visual assessment of NMS ordinations showed several trends in the compositional relationships of the trees. Distinction between age classes was strongest for Coleoptera, Diptera, and Hymenoptera, and weakest for Hemiptera and Lepidoptera. Vectors with arrows connect study pairs from 100yr to old. Several ordinations showed distinct clustering and/or separation of the age classes. Results for Lepidoptera should be viewed with the knowledge that abundances were generally very low for this taxon.

The 24 combinations of all traps+3 and all taxa +5 are presented in the following order:

a) all taxa in all traps pooled, then repeated with only the 122 RTU with >5 individuals collected (2 ordinations)

b) all taxa from sticky CD traps, all taxa from funnels, and all taxa from hangtraps (3 ordinations)

c) each of the five most prominent orders for all traps pooled (5 ordinations)

d) each of the five most prominent orders for each trap type separately (15 ordinations).

When reference is made to a study pair, it is in the form 100yr-Old.

Trees plotted closer together have a more similar composition of fauna. (Clarke & Warwick, 2001; McCune & Grace, 2002) The axes in NMS ordinations are arbitrary. Unlike other ordination techniques (e.g. Principle Components Analysis, PCA) axes represent equal amounts of variation, and have no intrinsic meaning (Clarke & Warwick, 2001; McCune & Grace, 2002).

No CD traps survived in trees 7 or 16, and only a single funnel trap survived in each of trees 5 and 6. These were removed from analyses of that trap type. While not removed in ordinations pooling all trap types, these trees are artificially drawn closer to each other because of the corresponding under representation.

8) Canopy Arthropods - Results

All taxa in all traps:

NMS of All taxa in All traps t05 t06 Age 100yr Old t03

t04 t08 t10 t02 t12 Axis 3 Axis

t09 t01 t11 t15

t16 t14 t13 t07

Axis 2

Figure 8.29: NMS ordination of trees by all taxa in all traps; 3-d stress=12.3; 6490 individuals.

All taxa, all traps: Clustering was evident for both age classes, with several outliers outside of the clusters (Figure 8.29). All old trees except 14 and 16 were grouped together. Old trees 4, 10, 12, 8, 3, and 6 exhibited one major cluster, while 100yr trees 2, 15, 13, 9, 1, 11, and 7 exhibited another. Old trees 16 and 14 were outlying old trees, overlapping with the 100yr trees.

Trees 5 and 7 were separated from the other 100yr trees. Trees 5-6 were close to each other and isolated from the others. These anomalies were likely a result of the missing funnel traps. 8) Canopy Arthropods - Results

The pairs 13-14 and 15-16 clustered together away from the other trees. These pairs were topographically higher and physically farthest from the others. Other study pairs exhibiting relative proximity in the ordination were 5-6, and 2-4. This was evidence that small-scale regional effects were a factor in determining composition.

Vectors showed a general trend upward. The compositional change between 100yr trees and old trees was therefore similar for each study pair except 15 -16 and 5 -6. The missing traps in these pairs could cause this exception.

RTU >5 individuals

t11 Age 100yr Old

t14 t12 t01 t07 t04 t13

t15 t03

Axis 3 t16 t09 t08

t10 t05

t06

t02

Axis 1

Figure 8.30: NMS ordination of trees by taxa with greater than five individuals, all traps; 3-d stress 9.9; 5257 individuals

All taxa with >5 individuals, all traps: A similar structuring to Figure 8.29 was exhibited when rare animal (<5 individuals) were removed (Figure 8.30). Compositional changes were again consistent between ages for each study pair, as shown by vectors trending rightwards. Study pairs 15-16 and 5-6 were again 8) Canopy Arthropods - Results exceptions. A regional effect linking pairs was seen in the clustering of 13-14 and 15- 16. Trees 7 and 16, which lost all CD sticky traps, tended to the left hand side. All old trees except 14 and 16 clustered together.

All taxa in each trap type:

NMS of All taxa in CD Sticky Traps

t04 Age 100yr Old

t11

t01 t10 t12 t02 t09 Axis 2 t03 t08

t05 t15

t06

t13 t14

Axis 1

Figure 8.31: NMS ordination of trees by all taxa in CD sticky traps; 2-d stress=20.5;1240 individuals.

All taxa, CD sticky traps: These two groups overlapped. No strong trends were identified comparing between ages (Figure 8.31). A general trend to the right suggested similarity in compositional differences. The study pair of tree 13-14 appeared closer to each other than to the other trees, suggesting a distinct airspace fauna in their vicinity.

8) Canopy Arthropods - Results

NMS of All taxa in Funnel Traps t13 Age 100yr t02 Old t16 t09 t14 t04 t15 t07 t11 t12 t10 Axis 2 Axis

t01 t08

t03

Axis 1

Figure 8.32: NMS ordination of trees by all taxa in funnels; 2-d stress=15.4; 2724 individuals

All taxa, funnel traps: 100yr trees exhibited a consistent difference in composition from the old trees (Figure 8.32). The majority of vectors are parallel, showing a regular compositional difference between ages. This intra-pair compositional trend is overshadowed by inter-pair variability. The spread of points appears similar. Old trees 14 and 16 are again separate from the other old trees. Pairs 13-14 and 15-16 again are in proximity, as is 11-12. This again suggests a distinct fauna in their vicinity. These six trees were the highest along the study site slope, and the farthest to the northeast.

8) Canopy Arthropods - Results

NMS of All taxa in Hangtraps t02 Age 100yr Old

t13 t06 t10 t14 t04 t16 t08

t01 t15 t05 Axis 2 Axis t07 t09

t03

t11

t12

Axis 1

Figure 8.33: NMS ordination of trees by all taxa in hangtraps; 2-d stress=36.6; 2526 individuals

All taxa, hangtraps: Trees were weakly clustered with five old grouped together (Figure 8.33). Old trees 16, 3 and 12 were outliers. 100yr trees 11 and 2 were outliers from the other 100yr trees. Study pairs exhibiting proximity in composition were 15-16, 13-14, and 7- 8. These shorter vectors suggest regional effects influenced composition of the hangtrap catch.

8) Canopy Arthropods - Results

Each taxon in all traps:

All Traps Coleoptera

t02

t09 t16 t13 t06 t10 t07 t14 t08 t01

t15 Axis 2

t03 t12 t04 t05

t11

Axis 1

Figure 8.34: NMS ordination of trees by Coleoptera in all traps pooled; 2-d stress 14.5; 792 individuals

Coleoptera, all traps: Strong clustering of the beetle fauna was observed (Figure 8.34). Beetle communities were distinct beetle communities between the tree ages. Tree 11 was an outlying 100yr tree, closest to its paired tree 12. Vectors tended rightwards. 8) Canopy Arthropods - Results

All traps Diptera

t13 t03

t04 t14 t10 t16 t15 t01 t02

t08 Axis 2 Axis t06

t12 t09

t11 t05 t07

Axis 1

Figure 8.35: NMS ordination of trees by Diptera in all traps pooled; 2-d stress 22.1; 3071 individuals

Diptera, all traps: Clustering was observed in dipteran composition (Figure 8.35), with a distinct group of old trees in close proximity to each other (4,10, 14, 15). A weak trend in vector directions upward was present. Exceptions to this trend were pairs 5-6, 7-8, and 15-16, all of which had at least one tree suffering heavy loss of traps. 8) Canopy Arthropods - Results All Traps Hemiptera

t12

t03 t16 t05 t07 t01t14

2 t13 t15 t09 t08 t10 t02 Axis Axis

t04 t11

t06

Axis 1

Figure 8.36: NMS ordination of trees by Hemiptera in all traps pooled; 2-d stress 18.6; 645 individuals

Hemiptera, all traps: Clustering of the hemipteran communities by ages was not observed (Figure 8.36). Variability was higher in old trees than 100yr trees. Pairs 13-14 and 1-3 showed strong similarity in hemipteran fauna within and between the pairs. 8) Canopy Arthropods - Results

All Traps Hymenoptera t02 age t05 1 2

t08

t03

t07 t06 t10 Axis 2 Axis t12 t09 t04

t13 t16 t14 t11

t15 t01

Axis 1

Figure 8.37: NMS ordination of trees by Hymenoptera in all traps pooled; 2-d stress 22.4; 655 individuals

Hymenoptera, all traps: The hymenopteran fauna was distinct between tree ages (Figure 8.37). Old trees showed a strong clustering to the lower right, with some overlapping 100yr trees. Vectors all tended rightwards, with pair 1-3 showing an exception. Variability in the 100yr trees was greater than in old trees. 8) Canopy Arthropods - Results

All Traps Lepidoptera t14 age 1 t13 2 t11

t09 t15 t10

t08 t06 Axis 2 Axis t04 t05

t03 t07 t12

t16

t01 t02

Axis 1

Figure 8.38: NMS ordination of trees by Lepidoptera in all traps pooled; 2-d stress 23.5; 388 individuals

Lepidoptera, all traps: Again, clustering of the old trees with overlaps from the 100yr trees was observed (Figure 8.38). Strongly parallel vectors showed that changes in composition between ages were consistent for the study pairs. The variability in the 100yr trees was greater than that in the old trees.

8) Canopy Arthropods - Results

Each taxon for each trap type:

CD Sticky Coleoptera Funnel Coleoptera t05 t12

t06 t14 t09 t13 t10 t03 t15 t13 t16 t01 t03 t09 t08 t08 t12 Axis 2 Axis Axis 3 t15 t04 t01 t11 t07 t14 t10 t02 t11 t02 t04 Axis 1 Axis 2

Hangtrap Coleoptera t05

t09 t12 t15 t07 t01

t04 Axis 2 Axis t02 t14 t03 t08 t06 t16 t13 t10

t11 Axis 1

Figures 8.39 (left), 8.40 (right), 8.41 (bottom): NMS ordination of trees by Coleoptera, for each trap type. From left to right: CD sticky (3-d stress 11.9; 116 individuals), funnel (3-d stress 10.7; 376 individuals), and hangtrap (3-d stress 14.2; 300 individuals)

Coleoptera for each trap type: The Coleoptera fauna exhibited strong age- related trends in all three trap types (Figure 8.39, 8.40, 8.41). In all cases, the fauna of the old trees was clustered strongly and showed less variability than the 100yr trees. In the CD sticky traps, old trees were strongly clustered. Composition changes were inconsistent, due to the high variability of the 100yr trees. In the funnel traps, all but one vector were very strongly parallel. The study pair 2-4 was the exception. The isolation of old tree 4 from the other old trees appears to drive the change in vector direction. In the hangtraps, a similar pattern to the CD sticky traps was observed. Old trees were strongly clustered and old trees were highly variable. The similarity in patterns could reflect the overlap of sampling airspace

8) Canopy Arthropods - Results

CD Diptera t06 Funnel Diptera t03 t10 t13t11 t05 t15 t04 t14 t09 t02 t12 t16 t13 t10 t15 t14 t03 t02 t07

Axis 2 Axis t08 t01

t04 Axis 2 t12 t08

t01

t11 t09 Axis 1 Axis 1

Hangtrap Diptera t14 t13 t02

t01 t09 t08 t03t06

Axis 3 Axis t10 t15 t16

t05 t12 t04 t11 t07 Axis 2

Figures 8.42 (left), 8.43 (right), 8.44 (bottom): NMS ordination of trees by Diptera, for each trap type. From left to right: CD sticky (2-d stress 15.1; 831 individuals), funnel (2-d stress 15.2; 792 individuals), and hangtrap (3-d stress 12.0; 1448 individuals)

Diptera for each trap type: In CD sticky traps, the dipteran fauna showed little distinction between age classes (Figures 8.42, 8.43, 8.44). By contrast, clusters appear in funnel and hangtrap dipteran composition. For the funnel traps, age classes were segregated and compositional differences consistent, with vectors showing strong trends to the left. The physically isolated pairs of 13-15 and 15-16 show similarity within and between the pairs. In the hangtraps, old trees cluster in the lower right. Variability was greater in the 100yr trees. Trees 14 and 16 was outlying from the other old trees, as seen in the ordinations of all taxa in all traps.

8) Canopy Arthropods - Results

CDSticky Hemiptera Funnel Hemiptera t12 t04

t11 t01 t02 t10 t15t11 t03 t14 t04 t14 t09 Axis 3 Axis t10 3 Axis t08 t13 t13 t07

t15 t01& t16 t08 t03t09 t06 t02 t12 Axis 2 Axis 1

Hangtrap Hemiptera t15

t04 t03t14

t13 t11 t09 t05 Axis 3 Axis

t01 t06 t07 t16 t10 t02 t12 t08 Axis 1

Figures 8.45 (left), 8.46 (right), 8.47 (bottom): NMS ordination of trees by Hemiptera, for each trap type. From left to right: CD sticky (3-d stress 14.2; 53 individuals), funnel (3-d stress 10.2; 465 individuals), and hangtrap (3-d stress 15.7; 127 individuals) Note that trees 1 and 8 are overlapping in the CD sticky traps. No Hemiptera were found in sticky traps in Tree 5.

Hemiptera for each trap type: The hemipteran fauna was not strongly distinct between age classes (Figures 8.45, 8.45, 8.47). In sticky traps, pairs 11-12 and 13-14 were dissimilar to the other trees and showed a high intra-pair similarity. For funnels, trees were intermingled and showed no trends in vectors. Segregation was strongest in the hangtraps, in which two groups of old trees formed separate clusters. Vectors tended rightwards. A diametrically opposed vector was observed for trees 11- 12, which were also outliers in the sticky traps.

8) Canopy Arthropods - Results

CDSticky Hymenoptera Funnel Hymenoptera t15 t08

t04 t11 t01 t16 t10 t15 t02 t04 t13 t09 t12 t14 t13 t05t03 t11 Axis 2 Axis Axis 3 Axis t02 t12 t09

t01 t10 t07 t14 t06 t08 t03 Axis 1 Axis 1

Hangtrap Hymenoptera t02 t01

t14 t13 Axis 2 Axis t06 t12 t10 t08

t16 t05 t04 t11 t15 t09 Axis 1

Figures 8.48 (left), 8.49 (right), 8.50 (bottom): NMS ordination of trees by Hymenoptera, for each trap type. From left to right: CD sticky (3-d stress 12.8; 185 individuals), funnel (3-d stress 11.1; 331 individuals), and hangtrap (3-d stress 14.0; 139 individuals)

Hymenoptera for each trap type :The hymenopteran fauna was distinct between age classes (Figures 8.48, 8.49, 8.50). Age classes clustered together in all trap types. In CD sticky traps, vectors tended downwards. Variability in fauna appeared similar for each age class. For the funnel traps, variability was again similar. With the pair 1-3 as exceptions, 100yr trees grouped in the lower right, and old trees in the upper left. In the hangtraps, variability was higher in 100yr trees.

8) Canopy Arthropods - Results

CDSticky Lepidoptera

t03 Funnel Lepidoptera t14 t12& t14t12t14 t10 t08 t02 t12 Axis 3

Axis 3 Axis t07 t03 t15 t13 t15 t11 t16

t10& t05 t05t10 t09 Axis 2 Axis 1

Hangtrap Lepidoptera t04 t03

t12

t09t10

t07 t06 t11 Axis 2 Axis

t02 t14 t05 t13 t15 t01 t08 t16 Axis 1

Figures 8.51 (left), 8.52 (right), 8.53 (bottom): NMS ordination of trees by Lepidoptera, for each trap type. From left to right: CD sticky (3-d stress 0.003; 7 individuals), funnel (3-d stress 9,84; 166 individuals), and hangtrap (2-d stress 19.3; 215 individuals). No Lepidoptera were collected in sticky traps in trees 1,2,4,6,8,9,11, or 13. Note overlap of 12 &14, and 10&5. No Lepidoptera were collected in funnel traps in trees 1 or 4.

Lepidoptera for each trap type: NMS ordinations by lepidopteran fauna were performed on very low abundances (Figures 8.51, 8.52, 8.53) . Lepidopterans were not strongly segregated by age. The catch in CD sticky traps was too sparse to generate a useful ordination. In funnel traps old trees tended leftwards and 100yr trees rightward. Vectors tended to the upper left. Old tree 10 and 100yr tree 13 were outlying from their age class. The hangtrap collection was overlapping between age classes. Variability was higher in old trees than 100yr trees.

8) Canopy Arthropods - Results

8.9.4 Dendrograms

Alltraps Alltaxa Dendrogram Distance (Objective Function) 4.6E+02 2E+04 4E+04 6.1E+04 8.1E+04 Information Remaining (%) 100 75 50 25 0

t03 t08 age t11 t06 1 2 t07 t01 t05 t12 t04 t09 t10 t14 t02 t13 t15 t16

Figure 8.54: Dendrogram grouping trees by total arthropod composition

The dendrograms classifying trees by all taxa showed little distinct clustering by age (Figure 8.54). For all traps pooled, trees 13, 15, and 16 formed a sister group to all the rest, supporting the NMS evidence showing a distinct fauna for the upper northwest study pairs. The pair of 15 with 16 was the only study pair classified immediately together.

CDSticky Alltaxa Dendrogram Distance (Objective Function) 8.9E+01 2.5E+03 4.8E+03 7.2E+03 9.6E+03 Information Remaining (%) 100 75 50 25 0

t01 t09 age t12 t10 1 2 t02 t04 t03 t05 t06 t08 t14 t15 t11 t13

Figure 8.55: Dendrogram grouping trees by CD sticky arthropod composition In the CD sticky dendrogram, the study pair of 5 with 6 formed its own group (Figure 8.55). The pair of 9 with 10 formed a group with the physically nearby tree 12, but were categorically distant from its pair of tree 11. Tree 13 was classified as distinct from all other trees. Clustering by age was not apparent. 8) Canopy Arthropods - Results

Funnel Alltaxa Dendrogram Distance (Objective Function) 9.2E+01 6.6E+03 1.3E+04 2E+04 2.6E+04 Information Remaining (%) 100 75 50 25 0 t01 t03 age t08 t11 1 2 t10 t12 t04 t07 t02 t15 t09 t16 t14 t13

Figure 8.56: Dendrogram grouping trees by funnel arthropod composition

The funnel trap dendrogram again shows tree 13 distinguished from all other study trees (Figure 8.56). Little clustering by age was apparent.

Hangtrap Alltaxa Dendrogram Distance (Objective Function) 1.2E+02 8.1E+03 1.6E+04 2.4E+04 3.2E+04 Information Remaining (%) 100 75 50 25 0 t01 t08 age t14 t03 1 2 t09 t07 t12 t11 t04 t06 t13 t10 t05 t15 t02 t16

Figure 8.57: Dendrogram grouping trees by hangtrap arthropod composition The hangtrap dendrogram showed little structuring by age (Figure 8.57). 8.10 RTU level morphospecific results

At an age level, 22 RTU were equally abundant between ages, 146 were more abundant in the 100yr collection, and 140 more abundant in the old collection.

Comparisons of RTU presence at a tree level showed a similar balance. 50 RTU were equally abundant between ages, 131 were detected in more 100yr trees than old trees, and 127 were found in more old trees than 100yr trees.

T-tests comparing the mean abundance between eight 100yr and eight old trees were performed on the 128 RTU that were encountered in more than five individuals. To compensate for the expected number of significant results in a large number of comparisons, only results with p<0.03 were considered. Five RTU exhibited significant differences (Table 8.11):

8) Canopy Arthropods - Results

Total Number of T-test Notes on taxa ecological Order Family Species binomial abundance trees prob niche 100yr Old 100yr Old Defoliating pest of Coleoptera Chrysomelidae Sterromela subcostata 7 0 4 0 0.014 Eucalyptus Parasite of Blattodea, Hymenoptera Evaniidae Evaniidae YDB sp.2 3 14 3 6 0.020 often on tree trunks Dead wood and bark Coleoptera Anobiidae Anobiidae YDB sp.01 14 41 6 8 0.026 borers Predator on Coleoptera Coleoptera Coccinelidae Rodatus TFIC sp. 01 25 2 7 2 0.026 and Hemiptera Eusocial, xylophage, Isoptera Isoptera YDB sp.2 5 14 5 7 0.029 herbivore

Table 8.11: RTU exhibiting significant differences in mean abundance per tree. Ecological niche notes from (CSIRO, 1967) RTU significantly more abundant in 100yr trees were Sterromela subcostata, a leaf-eating forestry pest beetle (Figure 8.58), and Rodatus TFIC sp. 01, a predatory beetle (Figure 8.59).

Significantly more abundant in old trees were Anobiidae YDB sp.01, a saproxylic wood borer (Figure 8.60); Isoptera YDB sp. 2, a winged xylophagous termite (Figure 8.61); and Evaniidae YDB sp.02, a wasp parasite of cockroaches (Figure 8.62). Cockroaches were insignificantly more abundant in old trees.

8.58: 8.59: 8.60:

8.61: 8.62:

Figure 8.58, Sterromela subcostata; 8.59, Rodatus TFIC sp. 01; 8.60, Isoptera YDB sp. 2; 8.61, Evaniidae YDB sp.02; 8.62, Anobiidae YDB sp.01. Scale = 1mm between blue marks

9) Canopy Arthropods - Discussion

9) Canopy Arthropods – Discussion

The differences detected in abundance, richness, diversity, and composition of canopy arthropods between the two classes are discussed. Overall abundance was not higher in the older trees, but is the measure most directly affected by scaling considerations. At a more detailed level, different taxa and different placements showed significant differences. The weight of evidence indicated a more rich and diverse fauna in the old trees. Strong evidence of distinct compositions between the age classes was found. These findings are linked to possible determinants of biodiversity in older trees: greater size, habitat variety, variety of element ages, and dominant position in the canopy. The greater space available in the older trees may foster a more diverse fauna by allowing species to avoid competing with each other.. Several taxa showed a less variable composition in the old trees and a more variable composition in the 100yr trees. This supports the findings of Schowalter (1995) in coniferous forests. 9.1 Abundance of faunal activity

Ha: Old-growth E. obliqua trees have a more abundant arthropod activity than 100 year old trees, i.e more individual arthropod animals.

Ho: No difference exists in arthropod activity between 100yr old and old E. obliqua

9.1.1 Summary of evidence

Overall, no significant differences in the total arthropod activity were found when comparing ages, for any trap position or for all traps pooled. When investigating specific taxa, variation in abundances could be identified. At a placement level, several taxa exhibited significant differences in abundance. Five RTU showed significant differences.

Results would be different if scaled up. The old trees were much larger than the 100yr trees. If measures such as crown volume or bark surface area were used to scale up, abundances would be much greater in old trees.

The evidence does not support the rejection of the null hypothesis of no difference in abundance between the age classes.

9) Canopy Arthropods - Discussion

Sufficient evidence exists to suggest more detailed trends, however. Different taxa were found in greater abundances in one of the age classes. More individuals of mites, beetles, and moths were detected for the same trapping effort in 100yr trees, and more spiders, cockroaches, ants, and termites in the old trees.

Spider abundance has been linked to older forests and architecturally complex habitats (Gunnarsson, 1990; McNett & Rypstra, 2000; Pettersson, 1996; Rypstra et al., 1999; Sundberg & Gunnarsson, 1994). In agreement with these works, the placement level results from the present study of Eucalyptus consistently showed more spiders in the older, more complex trees.

Beetle abundance is higher in old-growth Picea forests than in younger ones (Martikainen et al., 2000). Similarly, hangtrap results show a slightly higher abundance beetles in the old E. obliqua.

The higher abundance of mites in the 100yr trees and ants in the old trees is unsupported by the encountered literature. It is possible that ant abundance is linked to the greater presence in old trees of internal colony sites, i.e. decay and hollows.

9.1.2 Arthropod activity in 100yr and old trees

Lawton (1983) identified the size of a tree as a determinant in the number of habitats within its crown. Similarly, a larger tree may simply represent a larger “island” for an insect species to spread throughout (MacArthur & Wilson, 1967). Abundance likely would increase linearly with increase in habitat presence. As discussed before, habitat presence might not increase concurrently with any other measure of size. Different habitats may increase at different rates. For example, trunk interior wood volume does not scale up in step with amount of foliage.

Overall abundance of arthropod activity was generally similar between age classes. However, almost all possible measures of scale determined from the crown structural mapping are many times higher than those for the 100yr trees. The relevant habitat for certain taxa is likely to be more abundant in these larger trees. Scaling up would generate much higher abundance in the old trees. A strong example of this situation is reported by Basset (2001). He found a 2.5x higher abundance of herbivores per m2 of foliage beaten on mature trees of Pourouma bicolor than on the much smaller saplings. While his results are not scaled up, higher density per unit on the larger trees indicates a higher total abundance on the mature trees.

9) Canopy Arthropods - Discussion

Like Hooper (1995), Basset (2001) concludes that arthropod activity would diverge further between the age classes if results were scaled up. Using biomass as a surrogate for abundance, his results showed that older P. palustra had more activity in the entire tree, but less activity per unit of surface area.

Similar to the present study, Hanuala et al. (2000) viewed the catch from funnel traps as a sample of the entire tree (scaling situation A), and therefore did not scale up. However, unlike the present study, they placed drift fences around the entire circumference of tree trunk. The congruence of their unscaled results with Hooper’s (1995) scaled up results indicates that the funnel crawl traps do indeed sample at a tree level. Despite the smaller angular coverage of the drift fences in the present study, this supports the decision not to scale up results for funnel traps.

Further difficulties arise in using abundance as a measure of biodiversity. Individual animals may utilize trees of different ages at different stages of their life. Thunes et al. (2003) found a higher abundance of adult Thysanoptera on old trees, whereas the seed feeding juveniles were more abundant in younger trees. It is possible that the same Thysanoptera individual utilizes both age classes.

9.2 Richness of fauna

Ha: Old-growth E. obliqua trees have a richer arthropod fauna than 100 year old trees, i.e. more recognizable types of arthropods.

Ho: No difference exists in arthropod species richness between 100yr old and old E. obliqua.

9.2.1 Summary of evidence

The observed and expected richness was greater in the old trees for hangtraps and CD sticky traps, but not for funnel traps. Significant differences in richness were not detected at a tree level, but were at a placement level. The majority of significant differences at a placement level evidence a richer fauna in old trees.

Analysis of community structure through rank/abundance plots and species accumulation curves suggest a richer community overall in the old trees.

9) Canopy Arthropods - Discussion

A longer singletons tail in the old trees, matched with a shorter doubletons tail, can be seen in the rank/abundance plots. The presence of many singletons implies that sampling has not been adequate to fully survey the community. Magurran (2004) commented that a study in which all species are represented by at least two individuals may be deemed complete. In the species accumulation curve, the relationship of singletons to doubletons is utilized in estimating the true species richness via the Chao 1 formula. Towards the end of the all traps Chao 1 curves, the 100yr curve appears to have flattened off while the old curve appears to be rising still. Furthermore, the curve representing species actually observed is slightly higher for the old trees.

The higher richness in the old CD sticky traps and hangtraps may be due to the targeting of aerial taxa, especially Diptera, by the trap type. Because the old trees are taller, the upper crown aerial intercept traps may sample an uppermost forest stratum that the upper crown traps in 100yr cannot reach.

Funnel traps present the most contradictory story to the CD sticky and hangtraps. The species accumulation curves show a higher potential richness in the 100yr old trees. One potential explanation is that the trunk and bark fauna of E. obliqua is indeed richer in 100yr trees, while the airspace in the crown is richer in old trees. The high levels of richness in the funnel traps were due to a few outstanding traps. Another explanation is that the sampled area in 100yrs was a larger angular proportion in the smaller branches and trunk of 100yr trees.

The evidence supports rejection of the null hypothesis of no difference in richness between the age classes. Analysis of community structure and expected richness shows a more divergent picture than the observed species richness.

9.2.2 Possible determinants of richness

Despite widespread public opinion that old trees harbour more richness than younger trees (Fay, 2002; Green, 2002), few studies have compared arthropod richness in trees of different ages while controlling for environmental factors (Thunes et al., 2003).

Simultaneously, Harrison (2004) has found a higher species richness in old E. obliqua at Warra than in younger trees of similar age to the 100yr cohort. These

9) Canopy Arthropods - Discussion complementary results have addressed forests within ten km of each other at Warra LTER.

Like the current study, Thunes et al. (2003) found a higher species richness, before scaling down, in old Pinus sylvestris when compared to mature trees. Their study design, using pairs of nearby trees, closely parallels that of the present study.

Several factors may be simultaneously responsible for the higher richness observed in the old trees. These factors also contribute to a distinction of fauna between age classes.

1. More habitats and structural features in larger trees 2. Variety of ages of elements 3. Less vigorous defence 4. Dominant position in canopy

More habitats and structural features in larger trees: Lawton (1983) discusses the size of a tree as a determinant in the number of habitats within its crown. Several structural features form with increasing time (Mackowski, 1984). When studying growing trees, it is not always possible to separate the effects of size and age.

Specific features or elements within a tree have been shown to have an associated arthropod fauna. Nadkarni & Longino (2001) investigated the fauna in suspended soils, Paviour-Smith & Elbourn (1993) that of dead wood in tree crowns, and Ranius (2000) that of tree hollows. Pettersson (1996) linked spider richness to the abundance of lichen in Picea. All of these structural features were found more often in old trees.

In addition to the greater variety of habitats caused by the presence of these features, several habitats may be more abundant in the older trees. For example, five times the volume of dead whole branches was measured in the old trees of this study. To animals that see dead wood as the only habitat worthy of consideration, these trees represent a more favourable environment.

Variety of ages of elements: Structural features and elements in older trees branches represent a greater range of ages. In the 100yr trees, the live and dead branches originate from a regular pattern over a shorter period of time. The oldest

9) Canopy Arthropods - Discussion

possible branch is a century old, but due to branch occlusion in the lower crown the oldest remaining branch is certainly younger (Ishii & McDowell, 2001). In the upper crown of the 100yr old tree, the presence of an 85 year old dead branch is unlikely. In the old trees, live and dead branches originate from different times in the life of the tree. In the old trees, the oldest branches in the crown have been present for several centuries (Hickey et al., 1998).

Some organism are associated with features of different ages. Irmler et al. (1996) and Rambo et al. (2001) linked older rotting logs on the forest floor to increased richness of beetles and bryophytes, respectively. Similarly, Basset (2001) found a greater proportion of young tender foliage in the crowns of adult Pourouma bicolor than on sapling, and a higher herbivore richness. In agreement with the findings of Peeters (2002), he surmises that the increased richness may be to a preference or requirement of some species for younger foliage. Waltz & Whitham (1997) controlled for environmental factors by investigating the arthropod community on Populus in mature and basal resprout foliage within the same tree. The differences in fauna suggest that the presence of different aged elements fosters greater species richness.

Research in Eucalyptus forests has shown that marsupials require habitats of a certain age. For example, several marsupials require hollows of sufficient age and size for them to enter and live (Lindenmayer et al., 1990; Lindenmayer et al., 1997). Hollows in the 100yr trees will not be of sufficient age or size for these animals. Similar requirements may apply to arthropods.

Previous utilization by other organisms may be more important than the actual age of these older elements. The hollows that the marsupials live in result from activity of termites and fungi (Mackowski, 1984; May & Simpson, 1997). In the absence of these internal organisms, the hollow may not form or grow over time. It is possible that Californian eucalypts will never form hollows in the absence of these agents. The mechanical disturbance of dead wood by parrots and cockatoos in the wet sclerophyll forests may allow other organisms to colonize.

Less vigorous defence: In the present study, the higher abundance of ants and termites in older trees may indicate more disease and rotten interior nesting sites. A wood borer, Anobiidae YDB sp. 1, was associated with the older trees. Younger trees

9) Canopy Arthropods - Discussion

may be more vigorous and therefore can mount a stronger immune defence to herbivores, infection and decay (Lowman et al., 1987; Lowman & Heatwole, 1987). Organisms such as saproxylic insects are dependant on features which are symptoms of poor tree health. The old E. obliqua are invariably rotten and may be riddled with decay. Harrison (2004) has identified a higher species richness in the rotten interior of older E. obliqua at Warra than the less rotten interiors of younger ones.

Dominant position in canopy: The old trees were, on average, 15 m taller than the 100yr trees and occupy a dominant, emergent position in the canopy (Figures 9.1, 9.2 ). Trap positions were set relative to proportional crown heights (upper, middle, lower crown), rather than to absolute heights in the canopy above the forest floor as in Winchester (1997). Shaw (2004), and Basset et al. (2003) discuss the vertical stratification in forest canopy arthropod communities. The higher species richness detected for in animals such as Diptera, Hymenoptera, and Lepidoptera in the upper placements in old trees could be linked to the sampling of a distinct strata above the 100yr trees.

Figures 9.1 (left), 9.2 (right): Old E. obliqua are emergent above the average canopy level. Traps in the upper crowns of these trees may be sampling from a distinct stratum.

9.3 Diversity of fauna

Ha: Old-growth E. obliqua trees have a more diverse arthropod fauna than 100 year old trees, i.e. a greater quantitative index implying both a richer community and more even abundances between morphospecies.

Ho: No difference exists in arthropod diversity between 100yr old and old E. obliqua.

9.3.1 Summary of evidence

9) Canopy Arthropods - Discussion

Diversity, incorporating both richness and evenness in Simpson’s index (Magurran, 2004; Simpson, 1949), was higher in the old trees. Visual examination of radar charts shows that for most combinations of alltraps+3 and alltaxa+5, diversity was higher in the old trees. At a tree level, CD sticky trap diversity was higher in old trees. At a placement level, all significant (p<0.05) differences indicate a higher diversity in the old trees.

For the funnel traps, diversity at an age level was higher in the old trees for all five prominent taxa except Lepidoptera. This was in contrast to the higher RTU richness observed in the 100yr funnel traps.

In the Whittaker rank/abundance plots for all traps, an increased dominance of common taxa can be seen in the steeper slope of the 100yr curve. Increased dominance means lower evenness, and this less diversity.

The evidence supports rejection of the null hypothesis of no difference in diversity between the age classes.

9.3.2 Community structure and size

Community structure of older trees: The results of the present study agree with the findings of other studies have also found older trees to have both a more even and more species rich community structure. In agreement with the present study, Schowalter (1995) found a higher Shannon-Wiener diversity per kg of foliage in Oregonian old-growth Pseudotsuga menziesii forest trees than in young plantations. Barrios (2003) found mature trees of the Panamanian rainforest tree Castilla elastica to have a higher Simpsons D of herbivores than saplings, standardized by area of leaves beaten. It is unclear if the results would be similar if scaled up.

In the funnel traps, the old trees showed a higher diversity despite a lower richness. This means that the evenness of the funnel arthropod catch was higher in old trees. The community structure may be more even in the larger because there was more physical space for animals to co-exist without interacting (Jeffries & Lawton, 1984). It is possible that trunk surface area is an appropriate measure of the habitat size.

Bigger may mean more diverse by virtue of space: The size of plants may foster a more even community structure (Floater, 2001). Haysom & Coulson (1998)

9) Canopy Arthropods - Discussion

found a positive correlation between the height of Scottish heath Calluna vulgaris stands and sweep netted diversity (1/Simpsons) of Lepidoptera. In the present study, the taller, older trees showed a more diverse fauna. Like a larger island, a larger plant has more space to be filled by species (MacArthur & Wilson, 1967). With more space to disperse, evenness could increase because of decreased species interactions. In smaller spaces, strong dominant species may have more contact with less dominant species. They would therefore suppress them more effectively than in a larger space, where less dominant species may be able to find a place with less competition stress.

Diversity and Chao 1 richness estimator are linked: True richness and diversity will generally be positively correlated. A community with a higher Chao 1 estimate of true richness will also show a higher diversity index. This was observed for both the CD sticky and hangtraps Chao 1 estimator of true species richness rises with increased abundance of singletones, animals caught only once (Chao, 1984; Colwell, 2004). A large quantity of singletons can raise a measure of evenness, as all of these species were observed by a similar amount of individuals. They will also lessen the dominance of a single species. The long singletons tail will lessen the slope of a Whittaker rank-abundance plot.

Diversity indexes vulnerable to hyper-abundant taxa: The abundance of sciarid Dipterans may suppress the diversity indexes from the present research. A community with higher species richness may not have a higher diversity. This was observed in the funnel traps of the present study. The evenness aspect of a diversity index is vulnerable to hyper-abundant taxa. The community structure of a very rich community may also be very strongly dominated by a few species. For example, Abbott et al. (1992) collected outbreak levels of the gum leaf skeletonizer pest, Uraba lugens (Lepidoptera) on Eucalyptus foliage. The presence of these extra-abundant animals would strongly decrease the evenness of their collection.

This mathematical decrease would also be exacerbated by the trophic competition from the animals with high abundance. Their high abundance may be suppressing the numbers of other taxa in reality, as well as mathematically. A single abundant species can therefore strongly leverage the diversity index downwards. On a Whittaker rank-abundance plot, this would be represented by a very high, steep slope on the left.

9) Canopy Arthropods - Discussion

9.4 Distinctness of fauna

Ha: Old-growth E. obliqua trees have a distinct arthropod fauna from 100 year old trees, i.e. the composition of animals differs.

Ho: No difference exists in arthropod composition between 100yr old and old E. obliqua.

9.4.1 Summary of evidence

The age classes showed distinctions arthropod communities when viewed at a placement level, or when viewed via NMS ordinations which allow identification of environmental factors. Dendrogram and MRPP techniques pooling all taxa showed little distinction between the ages at age level. At a trap level and placement level, MRPP detected differences in composition between ages.

NMS ordinations allow for a more detailed analysis. Several taxa and trap type combinations showed a clustering in NMS that was not tested or detected by MRPP or dendrograms. The five most prominent taxa were treated separately. Regional differences were apparent in the clustering of trees in physical proximity. Strong differences in composition were observed for Coleoptera, Diptera, and Hemiptera. Funnel collections showed the clearest distinctions in fauna, and the most.

For the funnel trapped Hymenoptera and Coleoptera, and the sticky trapped Hymenoptera and Coleoptera, a strong parallel shift in composition was noted between the tree ages. This effect would have been undetected except for the study design pairing the trees to control for environmental effects.

The evidence supports rejection of the null hypothesis of no distinction in fauna between 100yr and old tree. The composition of RTU communities is, for some aspects, distinct between the age classes. Examining different combinations of trap types and taxa exposes several strong distinctions. Environmental and spatial factors can mask the differences between age classes.

9) Canopy Arthropods - Discussion

9.4.2 Age related distinction of fauna is distinct from other sources of variability:

In concordance with the results from the present project, Harrison (2004) has found a distinct saproxylic beetle fauna associated with the interiors of old E. obliqua at Warra.

Several other studies have found distinct faunal communities between forests or trees of different ages. Schowalter (1995) distinguished a characteristic old-growth forest community in western American coniferous forests. Martikainen et al. (2000) found a distinct saproxylic beetle community in old spruce forests when compared to managed younger forests. Sippola (2001) found distinctions in species composition between old, unmanaged boreal forest and younger, managed forests.

The factors involved in determining differences in taxonomic composition mirror those that determine richness. Richness may be driven by specialist fauna, which will distinguish the total fauna of the entire tree. Conversely, species that may contribute to overall richness but were not confined to one age class will blur distinctions in fauna. For example, if a species is only found in association with the dead tops of old Eucalyptus, then this taxa would contribute to distinction between age classes.

Geographic effects will mask other effects: Environmental and geographic effects may mask other patterns in composition. For example, the results for funnel traps show a parallel difference in composition between ages. If study trees were not selected in pairs, this spatial effect would mask the observation of an age effect. When comparing canopy arthropods of different forest stands, careful study design and replication is necessary (e.g. Schowalter, 1995). Previous studies have pseudoreplicated forest age and it is impossible to tell whether differences are actually due to environmental effect (Martin, 1966; Neumann, 1978). Studies of very mobile taxa such as Diptera may be more subject to this geographic effect than less mobile taxa such as Isoptera.

Richardson et al. (1999) quantified this geographic change by assessing canopy faunal similarity along linear distance for Melaleuca. Similarly, Burgman & Williams (1995) reanalysed the foliage clipping collections from E. marginata of

9) Canopy Arthropods - Discussion

Abbott et al. (1992) on a 2-dimensional geographic matrix and find an effect at the kilometer scale.

In the NMS ordinations of the present study, trees 13, 14, 15 and 16 often showed that they were more similar to each other than to the other trees. These trees were closer to each other than to the other study trees and provide evidence of a strong spatial effect.

In contrast with the present findings, Thunes et al. (2003) found tree age and structural descriptors were not as strong in determining overall arthropod composition in Norwegian P. sylvestris as environmental variables such as soil nutrition and topographic position. Their studies, like the present study, controlled for environmental variation by comparing pairs of old and mature trees.

Thunes et al. (2003) attributed some of the changes in species composition between study regions to differences in tree vigour. In the E. obliqua in the present study, the 100yr trees were much more vigorous than the old trees. Reid & Robb (1999) link coniferous tree vigour to the presence of a bark beetle pest, and Stone & Bacon (1995) describe a cycle in which drought stressed E. camaldulensis are more susceptible and to and is further stressed by herbivory. The combination of geographic effects and stress can further overshadow the differences between tree ages.

In the present study, the understorey and mid-canopy rainforest trees were not measured as predictor variables for the composition of arthropods. It is possible that the neighbouring plant species determines faunal composition more strongly than the structural factors measured. The rainforest understorey, botanically linked to the ancient supercontinent of Gondwana, has distinct arthropod elements from the Eucalyptus trees. (McQuillan 1993).

Intra crown variability will mask inter crown difference: Variability of the fauna within the tree crown may also mask differences between trees (Majer et al., 1990). For example, in the present research, the funnel trap results pool trunk traps with both live and dead branches. Viewed as a regime, interpretations may differ from if they are viewed separately. If any systematic difference between these three trap locations exists, they will increase the variability detected within each tree and therefore lessen the ability to detect differences between trees. The high intra-crown

9) Canopy Arthropods - Discussion

variability of Eucalyptus canopy invertebrates found by Majer et al. (1990) may suppress the detectability of inter-crown differences.

Evidence in the MRPP analysis from the present research shows this masking was occurring. MRPP comparisons at a tree level showed little differences between ages. However, at a placement level, different habitats were more effectively partitioned. With the masking effect controlled, more differences between ages appear.

Foliage may be the least variable of habitats: Fowler (1985) found a high overlap in phytophagous fauna in Betula foliage between seedlings, saplings, and trees. Most species found on seedlings and saplings were detected in the tree foliage. He reported on the similarity of habitat, as viewed by herbivores, between the age classes. Further, the proximity of the trees to seedlings may result in animals living in the habitats of trees feeding on the saplings.

At the intra-tree scale, Waltz & Whitham (1997) found a distinct fauna on younger foliage. In larger trees, young foliage may be more abundant (Basset, 2001). Because young foliage is present in both age classes, this would mask other differences.

One age class may be less variable than the other: Variability in the Coleoptera, Hymenoptera, and Lepidoptera composition was higher in the 100yr trees. For these taxa, old trees showed a consistent community of arthropods with 100yr trees showing a wide range of partially related communities. The old trees were more similar to each other than the 100yr trees were to each other. This pattern can be observed most strongly in the CD sticky and hangtrap Coleoptera fauna. NMS ordination for these trap types shows that the differences between age classes were not parallel, and that 100yr trees were highly variable in different ways when compared to the similarity between old trees. The old trees clustered in the middle and the 100yr trees were arrayed around them.

These match the findings of Schowalter (1995). In coniferous Pseudotsuga menziesii/Tsuga heterophylla at the H.J. Andrews LTER in Oregon, he found a distinct, consistent community on the foliage of old-growth forest trees. The trees in younger forests harboured a subset of the old-growth forest community, but with more variability in composition. The older forest replicates were more similar to each

9) Canopy Arthropods - Discussion

other than the younger forests were to each other. This pattern also appears at a tree scale- old Eucalyptus trees in the same forest are more similar to each other, for some taxa, than the 100yr trees are to each other.

By extension, following Ishii & McDowell’s (2001) analogy of older branches with older forests, the hypothesis can be formed that old branches in the tree crown will have a consistent, less variable fauna relative to the younger branches. Waltz & Whitham (1997) have found the herbivore communities within Populus crowns to be different on mature and juvenile branch suckers, and Clement & Shaw (1999) found large, old branches to support the biodiversity of epiphytes in Pseudotsuga. It remains to be tested if younger, smaller branches support a subset of the biodiversity associated with larger, older branches.

The more structurally variable old tree crowns have a less variable fauna than the more structurally consistent old trees: This pattern is in diametric opposition to the findings that crown structure in the old trees is highly variable. It is counter-intuitive that trees with divergent crown structures have convergent animal communities. The reasons for this remain unclear. A possible explanation for the results of the present study could that all of these animals live on the old trees, and only visit the 100yr trees on occasion. In analogy, the possums that are often in the hollows of old eucalypts could sometimes be detected in younger trees (Gibbons & Lindenmayer, 2002).

10) Correlation - Methods

10) Correlation – Methods

Correlations between structural attributes and arthropod biodiversity were investigated. The results can shed light on the influence of structure and the response of the animals. Using bivariate techniques, the structural descriptors in ten categories were matched against summaries of abundance, richness, diversity, and the abundance of each RTU. NMS vector biplots highlight structural descriptors most strongly correlated with changes in arthropod composition. Mantel tests investigated the strength of the relationship of similarity between trees for structure against the similarity between trees for arthropods. 10.1 Exploring correlations between structure and arthropod biodiversity

The effect of crown structure on the trunk and canopy arthropod biodiversity was explored at a tree level. Only information describing each tree was used. No trap or placement level analyses were conducted. The structural descriptors of the tree were considered independent predictors, and tree level arthropod biodiversity descriptors were considered dependent responses.

Interpretation of the results must accommodate that several structural descriptors were strongly correlated. For example, mean cone volume corresponds with mean cone surface area. Some descriptors were perfectly correlated. For example, the largest branch airspace was always found in a large branch, and therefore the descriptor of maximum branch airspace for all branches will be the same as the maximum branch airspace for dead branches.

Correlation does not equal causation. The correlation of a structural descriptor with any aspect of arthropod biodiversity does necessarily mean the arthropod is responding to that measure. It is possible that the animals are actually responding to cryptic predictors. If these cryptic predictors are themselves correlated with the observed predictor variables, then a potentially misleading predictor-response will be observed.

It is also worth noting that a correlation between measures of structure and arthropods does not indicate which is truly a predictor or a response. If one

10) Correlation - Methods

hypothetically accepts that arthropods are responsible for shaping the structure of a tree, then the correlations are also evidence of arthropods predicting structural responses.

The results of this meta-analysis are preliminary and exploratory. With an immense number of explored hypotheses defined robotically, and a very small number of replicates, statistical power is extremely low. Furthermore, many of the predictor responses are correlated to each other. Statistical techniques such as the Bonferroni correction minimize the chance of false positives by increasing the chance of false negatives (Benjamini & Yekutieli, 2001; Garcia, 2004). In the present exploratory research, analysis is performed in a contrasting manner. False positives were considered a lesser evil than false negatives. Positive results will suggest new avenues of investigation, which future investigations can assess. This is in contrast to fields such as medical drug research, where a false positive could expose people to needless harm (Perneger, 1998). Therefore, a decision has been made to prefer false positives to false negatives. 10.2 Bivariate Correlations

Bivariate analyses of correlation were performed using JMP software (SAS Institute, 2003). The structural descriptors were considered predictors, and tree level summaries of arthropod total abundance, richness, and diversity were considered responses. The non-parametric correlation test statistic Spearman’s rho (SAS Institute, 2003) was generated for each combination. This test tests the relationship between two variables by comparing the ranks of trees rather than the actual values. The assumption of normality was considered unstable for the full range of variables. The reported p-value was used to test a null hypothesis of no relationship between the variables.

To remove the confounding factor of age, analysis was done simultaneously and separately on each age class. The predictor variables were the 107 structural descriptors generated to describe each trees crown structure.

Summary variables of arthropod biodiversity were used as response variables. These were the 24 combinations of alltaxa +3 and alltaxa +5 for abundances, RTU richness, and diversity as measured by -ln(Simpsons D). In addition, the abundances per tree for the 128 RTU with greater than five individuals collected were used as

10) Correlation - Methods

response variables. The statistics software automatically removed invalid comparisons. This was usually due to situations in which no specimens of a taxon were caught in an age class.

Interpretation of p-value output: With α = 0.05, one in twenty pairs of variables could be expected to show a significant result by chance alone (Benjamini & Yekutieli, 2001; Garcia, 2004; Perneger, 1998).

Using techniques summarized in Garcia (2004), the following procedures were applied to compensate for the false discovery rate inherent in the high number of comparisons and to filter a manageable number of correlations for presentation.

Standard Bonferroni correction: The standard Bonferroni corrected significance was tested. The Bonferroni correction, or “one-step Bonferroni” (Garcia, 2004), divides the significance level α by the number of tests performed. This test is considered overly conservative and is designed to eliminate the chances of false positives (Perneger, 1998).

Step Up FDR technique: Sharpened “step-up False Discovery Rate” (hereafter “FDR”) significance levels were calculated (Benjamini & Yekutieli, 2001; Benjamini et al., 2002). The FDR technique was designed to control the proportion of false positives to observed positives. FDR is more tolerant of false positives when there are many observed significant results, and less tolerant when there are few (Benjamini & Yekutieli, 2001). The process is reworded by Garcia (2004):

a) p-values are ranked in ascending order, j being the resulting

rank. b) Proceed from jn to j1, until finding a first p-value, ranked k, satisfying

pk≤/zk */ α /n. c) then reject Ho for j ≤ k and accept all the remaining null hypotheses

Observed: Expected ratio for different levels of α: The uncorrected number of expected and observed significant results at both α = 0.1, α = 0.05, α = 0.01, α = 0.005, and α = 0.001 are presented in graphical form and discussed.

P<0.001 for either age class: Pairs of correlations with a p<0.001 for either age class were tabulated. This is one fiftieth of the normal α = 0.05. The p-value of the other age class was also reported.

When combinations were similarly correlated in both age classes, this supports the significant results and is evidence against a false positive. If the combination was

10) Correlation - Methods

highly correlated one age class but not the other, this could be interpreted in two ways. It is either evidence of a false positive, or an indication of different dynamics within the crowns of only one age classes. Similarly, sign mismatches (+/-), can be viewed as evidence of either a false positive or an opposing trend in the age classes.

Combinations were grouped by response type (abundance, richness, diversity, RTU abundance) and by predictor descriptor category (Tree size, crown depth, etc.). Within each chart, predictor categories are divided by lines.

Sorting the combinations by predictor category rather than RTU response was not intended to express an interest in predictive structures over responsive arthropod taxa. This imbalance in presentation was noted and the discussion addresses responding taxa as well as effective predictors.

P<0.05 for both age classes: Bivariate combinations in which both age classes reported a rank correlation probability of p<0.05 were tabulated. The signs of the rank correlation coefficients were compared to test that age classes were not showing opposite relationships. Opposite correlations would suggest a false positive, in which contradictory relationships existed for the different age classes. Combinations were grouped as above. To aid in visually identifying arthropod responses, letters were used to code similar responses. These letters served to link auto-correlated predictors and to serve as identifiers for discussions. 10.3 Multivariate correlations

NMS ordination: Multi-variate techniques were used to compare the effect of the structural descriptors on the faunal composition of each tree. The abundances of the 128 RTU with greater than five five individuals collected were used for NMS ordinations (species matrix), and the 107 structural descriptors were related to the ordination (environmental matrix) (McCune & Grace, 2002).

This was performed for all trees pooled. The r correlation coefficient between each structural descriptors and the axes of Non-metric Multidimensional Scaling (NMS) ordination was output from PC-ORD (McCune, 1999). These offer descriptive comparisons of influence. The unsquared r-value is presented to retain the sign (+/-) of the correlation. P-values were not output from PC-ORD as samples violate the assumption of independence (McCune & Grace, 2002)

10) Correlation - Methods

Higher correlations are evidence of an influence of a descriptor on the composition of the tree fauna. Graphical output was generated overlaying structural variables onto an NMS plot. Numerical output was generated by ranking and tabulating the correlation coefficients.

Mantel test: A Mantel test was performed between the structural descriptors and the arthropod fauna for alltraps +3 using PC-ORD software (McCune, 1999). This was done for both age classes pooled, and then for each age class separately. This test compares the correlation between the (Sorenson/Bray-Curtis) distance matrices relating the trees by structural descriptors and by arthropod collections (McCune & Grace, 2002).

:

11) Correlation - Results

11) Correlation – Results

Several correlations between structural descriptors and arthropod biodiversity were identified. The Bonferroni correction and Step Up FDR techniques yielded no significant results. At several levels of alpha, the old trees showed more significant results than expected with summaries of richness and diversity. Correlations are tabulated when p<0.001 for either age class, and when p<0.05 for both age classes. Mismatched signs (+/-) between the ages classes for a correlation were marked. These indicate either opposing trends in the different age classes, or false results. NMS biplot analysis graphically illustrated and identified structural descriptors that were most correlated with changes arthropod composition. Mantel tests showed no correlation between the structural descriptor distance matrix and arthropod composition distance matrix. 11.1 Bivariate results

11.1.1 Standard Bonferroni correction results

When filtered through the Bonferroni correction, no significant results were detected using Pearson’s rho rank correlation test. For the Bonferroni correction, the required probability for a significant result was approximately 0.00002. Output from JMP statistics software output results to only four decimal places.

11.1.2 Step Up FDR technique results

Similarly, no significant rank correlations were detected using Step Up FDR techniques. Manual implementation of the algorithm using Excel (Microsoft, 2000) and automated implementation using FDRalgo software (Benjamini et al., 2002) vgenerated a required probability of 0.

11.1.3 Observed: Expected ratios for different levels of α

Structural descriptors were correlated to arthropod abundances summaries less often than expected by chance alone (Figure 11.1). 11) Correlation - Results

Bivariate ScatterPlots Rho Correlations Proportion of Expected Significant Results Y at Alpha X 3

Abundance 100yr 2.5 Abundance Old Expected 2 Ratio 1.5 Observed to Expected 1

0.5

0 0.1 0.05 0.01 0.005 0.001 Alpha

Figure 11.1: Ratio of observed to expected significant results at different levels of significance for bivariate rank correlations of 107 structural descriptors to 24 abundance summaries

In the old trees, more rank correlations were found between the structural descriptors and summaries of arthropod richness than expected by chance alone (Figure 11.2). In the 100yr trees, fewer than expected correlations were found.

Bivariate ScatterPlots Rho Correlations Proportion of Expected Significant Results Y at Alpha X

5 4.5 4 3.5 Richness 100yr 3 Ratio Richness Old 2.5 Observed to Expected 2 Expected 1.5 1 0.5 0 0.1 0.05 0.01 0.005 0.001 Alpha

Figure 11.2: Ratio of observed to expected significant results at different levels of significance for bivariate rank correlations of 107 structural descriptors to 24 richness summaries

Similar to the richness results, old trees always had more correlations for diversity than expected by chance (Figure 11.3), and 100yr trees always had less than expected 11) Correlation - Results

Bivariate ScatterPlots Rho Correlations Proportion of Expected Significant Results Y at Alpha X

3 Diversity 100yr 2.5 Diversity Old Expected 2 Ratio 1.5 Observed to Expected 1

0.5

0 0.1 0.05 0.01 0.005 0.001 Alpha

Figure 11.3: Ratio of observed to expected significant results at different levels of significance for bivariate rank correlations of 107 structural descriptors to 24 diversity summaries

For RTU abundances, slightly more correlations than expected by chance alone were detected in the 100yr trees, and slightly fewer in the old trees (Figure 11.4). At all surveyed levels of significance, 100yr trees had a higher ratio of observed: expected significant results than old trees.

Bivariate ScatterPlots Rho Correlations Proportion of Expected Significant Results Y at Alpha X

1.8 1.6 1.4 1.2 1 Ratio Observed to 0.8 Expected RTU 100yr 0.6 RTU Old 0.4 Expected 0.2 0 0.1 0.05 0.01 0.005 0.001 Alpha

Figure 11.4: Ratio of observed to expected significant results at different levels of significance for bivariate rank correlations of 107 structural descriptors to 128 abundances of RTU

11.1.4 P<0.001 for either age class

No combinations of variables were detected as significantly rank correlated at p<0.001 in both age classes. Of 37 bivariate combinations in which one age class showed significant rank correlation at p<0.001, six predicted abundance summaries, seven predicted richness summaries, six predicted diversity summaries, and eighteen 11) Correlation - Results predicted the abundance of a specific RTU. Combinations with contradictory signs are marked with an asterisk (*). 18 of the 37 had mismatched signs.

Abundance:

Table 11.1. Spearman’ s rho rank correlations between 107 structural descriptors and 24 summaries of abundance in which at least one age class exhibited a p-value<0.001

In the 100yr age class, all significant correlations were found for animals in the CD sticky traps (Table 11.1). Coleoptera abundance was negatively correlated to the ratio of crown airspace to trunk wood volume. The 100yr trees with the most spreading crowns and slender trunks had the fewest beetles. Hymenoptera in CD sticky traps responded positively four auto-correlated descriptors of branch cone size. The 100yr trees with the largest total and average sized cones had more hymenopterans than those with smaller.

In the old trees (Table 11.1), Hemiptera abundance in funnels was negatively rank correlated with the mean vertical arc of live branches. Old trees with live branches with less vertical arc had a greater abundance of bugs.

These results were generally not matched by a significant correspondence in the other age class. The CD sticky hymenopteran abundance showed an contradictory correlation for total branch volume in the two age classes (i.e. signs did not match). It is unclear if this is evidence of an opposing trend or of a spurious result.

Richness:

Table 11.2. Spearman’ s rho rank correlations between 107 structural descriptors and 24 summaries of richness in which at least one age class exhibited a p-value<0.001 11) Correlation - Results

As indicated by Table 11.2, significant correlations with summaries of richness were detected more often in the old trees. Like the correlations of Hemiptera abundance in funnels with a measure of vertical arc, their richness was also negatively correlated with the mean upward arc of branches in old trees. The mean branch upward arc also negatively predicted the total richness collected in the hangtraps.

For old trees, Lepidoptera richness in the hangtraps was positively rank correlated with the mean upwards slope of live branches. Diptera richness in CD sticky traps was negatively predicted by the total live branch airspace. The richness of Hemiptera in hangtraps was positively correlated with the mean surface area of dead branches.

Only one correlation was detected in 100yr trees. The average surface area of dead branch airspace as calculated by polyhedra was positively correlated to the richness of Hymenoptera in the funnel traps.

Diversity:

Table 11.3. Spearman’ s rho rank correlations between 107 structural descriptors and 24 summaries of diversity in which at least one age class exhibited a p-value<0.001

Like the richness summaries, old trees showed more significant correlations of diversity responses at p<0.001 than 100yr trees (Table 11.3).

In the old trees, the total diversity of the arthropod fauna was positively correlated with the portion of the total wood volume in the trunk. Old trees with more wood in the trunk than the branches supported a more diverse fauna. Positive correlations were detected for the total wood volume of live branches with Hymenoptera diversity in the CD sticky traps, and for average branch length with Lepidoptera diversity in the funnel traps. Negative correlations were detected for beetle diversity in CD sticky traps with total wood volume of live branches, and for hangtrapped Lepidoptera diversity with the mean vertical arc of live branches. 11) Correlation - Results

In the 100yr age class, the diversity of Hemiptera was positively correlated with the mean start diameter of the branches. Trees with thicker branch bases had a more diverse bug fauna.

RTU abundance:

Table 11.4. Spearman’ s rho rank correlations between 107 structural descriptors and 128 abundances of RTU in which at least one age class exhibited a p-value<0.001

The abundance of several RTU showed rank correlations with structural descriptors (Table 11.4). Three moths showed multiple correlations. The grouped ants, three flies, two bugs, two wasps, and a beetle each showed single correlations.

An oecophorid moth, Oecophoridae YDB sp. 2, was positively predicted by the total crown depth in 100yr trees and the average surface area of dead branches in old trees. It was negatively predicted by the average live branch height in the 100yr trees. Total crown depth and mean live branch height are auto-correlated structural descriptor. Deeper crowns usually had lower live crowns, and therefore these correlations correspond. A tortricid moth, Torticidae YDB sp. 2, was positively correlated in the 100yr trees with the standard deviation and range of the dead branch scaling ratio, and negatively correlated in the old trees with the average branch starting diameter. A cosmopterygid moth, Cosmopterygidae YDB sp. 1, showed significant rank correlations in the 100yr trees. Its abundance was positively correlated with diameter at breast height, and negatively correlated with lowest branch height.

11.1.5 P<0.05 for both age classes

Only half (32 of 61) of the bivariate combinations with rank correlation significances of p<0.05 for both age classes agreed in their signs (+/-). Five 11) Correlation - Results correlations were identified describing abundances, fifteen correlations for each of richness and diversity, and 26 for RTU abundances. Combinations which contradictory signs are marked with an asterisk (*). A letter is placed next to the arthropod response to aid in visual assessment of response trends.

Abundance:

Abundance +/- 100yr Old Predictor Abundance Response mismatch? Rho P-value Rho P-value Sum Cone Volume (m3) all branches Hymenoptera- CDsticky A * 0.955 0.0008 -0.7928 0.0334 Mean Up Arc (deg) all branches Hymenoptera- Funnel B -0.8829 0.0085 -0.7748 0.0408 Mean Vertical Arc (deg) all branches Hymenoptera- Funnel B -0.8829 0.0085 -0.7748 0.0408 Mean Horizontal Arc (deg) all branches Hemiptera- Funnel C * -0.7748 0.0408 0.8214 0.0234 Mean Horizontal Arc (deg) live only Hymenoptera- Funnel B * -0.7568 0.0489 0.8469 0.0162

Table 11.5. Spearman’ s rho rank correlations between 107 structural descriptors and 24 summaries of abundance in which both age classes exhibited a p-value<0.05

Three of five detected correlations with abundance responses had mismatched signs (Table 11.5). Summaries of funnel trap Hymenopteran abundance responded negatively to measures of upwards arc, vertical arc, and live branch horizontal arc.

Richness:

Richness +/- 100yr Old Predictor Richness Response mismatch? Rho P-value Rho P-value Total Crown Depth (m) Diptera- All traps -0.7306 0.0396 -0.7711 0.0251 Mean branch height as % of treetop height All taxa- Hangtrap I -0.8571 0.0065 -0.8333 0.0102 Mean live branch height as % of treetop heighAll taxa- Hangtrap I -0.7857 0.0208 -0.8571 0.0065 Mean Cone Volume (m3) all branches Lepidoptera- Funnel G 0.7783 0.0393 0.8669 0.0115 Max Cone Volume (m3) dead only Lepidoptera- Funnel G * -0.8524 0.0148 0.8078 0.028 Std Dev Cone Volume (m3) dead only Lepidoptera- Funnel G * -0.8524 0.0148 0.9063 0.0049 Mean Cone Surface Area (m2) all branches Lepidoptera- Funnel G 0.7783 0.0393 0.8669 0.0115 Max Polyhedra Volume (m3) all branches All taxa- Hangtrap I * 0.7619 0.028 -0.8095 0.0149 Max Polyhedra Volume (m3) live only All taxa- Hangtrap I * 0.7619 0.028 -0.8095 0.0149 Std Dev Polyhedra Volume (m3) dead only All taxa- Hangtrap I * -0.9048 0.002 0.7143 0.0465 Std Dev Polyhedra Volume (m3) live only Diptera- Hangtrap * 0.7357 0.0375 -0.7186 0.0446 Range Start Diameter (m) live only All taxa- Hangtrap I 0.7857 0.0208 0.8333 0.0102 Mean Distance (m) all branches Lepidoptera- Funnel G 0.8154 0.0254 0.9063 0.0049 Mean Slope (deg) live only Hemiptera- Funnel C * -0.8829 0.0085 0.7857 0.0362 Mean Slope (deg) live only Lepidoptera- All traps * -0.747 0.0332 0.8051 0.0159

Table 11.5. Spearman’ s rho rank correlations between 107 structural descriptors and 24 summaries of richness in which both age classes exhibited a p-value<0.05

Eight of fifteen correlations with richness responses had mismatched signs (Table 11.5). The total richness of the fauna collected in hangtraps was negatively correlated with relative average branch heights, and with the standard deviation of 11) Correlation - Results dead branch airspace volume. The hangtrap richness was positively correlated with the range in start diameters for live branches. Trees with a range of branch girths had the richest hangtrap catch. The richness of flies in all traps pooled was negatively correlated with total crown depth. Their richness in hangtraps was negatively correlated with the standard deviation of live branch airspace volume. Lepidopteran richness in funnels corresponded positively with the branch volume, surface area, and length.

Diversity:

Diversity +/- 100yr Old Predictor Diversity Response mismatch? Rho P-value Rho P-value % of Wood Volume in Trunk Hemiptera- Hangtrap -0.7665 0.0265 -0.7619 0.028 Highest Branch Height (m) all branches Hemiptera- Funnel C -0.8571 0.0137 -0.8571 0.0137 Total Crown Depth (m) All taxa- CD Sticky D * 0.9286 0.0025 -0.9286 0.0025 Mean Branch Height (m) all branches Coleoptera- Funnel E -0.7857 0.0362 -0.8214 0.0234 Mean Branch Height (m) all branches Diptera- Funnel F -0.7857 0.0362 -0.8214 0.0234 Mean Branch Height (m) live Coleoptera- Funnel E -0.8571 0.0137 -0.8214 0.0234 Mean Branch Height (m) live Diptera- Funnel F -0.8571 0.0137 -0.8214 0.0234 Max Cone Volume (m3) live only Lepidoptera- Funnel G 0.9429 0.0048 0.8857 0.0188 Sum Cone Surface Area (m2) all branches Hemiptera- Hangtrap H 0.8383 0.0093 0.7619 0.028 Sum Cone Surface Area (m2) live only Hemiptera- Hangtrap H 0.7186 0.0446 0.7619 0.028 Sum Polyhedra Volume (m3) live only Hemiptera- Hangtrap H 0.8144 0.0138 0.7857 0.0208 TreePolyhedra Surface Area (m2) Hemiptera- Hangtrap H 0.8982 0.0024 0.7143 0.0465 Mean Start Diameter (m) all branches Lepidoptera- Funnel G 0.8857 0.0188 0.8286 0.0416 Range Start Diameter (m) all branches Hemiptera- Funnel C * -0.8214 0.0234 0.9286 0.0025 Range Start Diameter (m) dead only All taxa- CD Sticky D -0.8214 0.0234 -0.8108 0.0269

Table 11.6. Spearman’ s rho rank correlations between 107 structural descriptors and 24 summaries of diversity in which both age classes exhibited a p-value<0.05

For diversity responses, only two of fifteen correlations had mismatched signs (Table 11.6). The diversity of Hemiptera responded to several structural descriptors. Hemipteran funnel diversity was negatively correlated with the height of the highest branch. Their hangtrap diversity was negatively correlated with the portion of wood volume in the trunk, in contrast to the positive correlation with this predictor for diversity of all taxa in old trees (Table 11.3). Hangtrap Hemipteran diversity was further positively correlated with total branch surface area, branch airspace volume, and total crown airspace surface area.

Funnel Lepidoptera responded positively to maximum live branch volume, and to the average branch starting diameter. Both funnel Coleoptera and Diptera were negatively correlated with mean branch height. The total diversity of arthropods 11) Correlation - Results collected in CD sticky traps was correlated negatively with the range of dead branch diameters.

RTU abundance:

RTU Abundance +/- 100yr Old Predictor RTU Abundance Response mismatch? Rho P-value Rho P-value Ratio Polyhedra m3:Trunk m3 Coleoptera:Salpingidae:Orphanotrophium frigidum J -0.779 0.0227 -0.8607 0.0061 Mean branch height as % of treetop height Coleoptera:Brentidae:Apion tasmanicum J * 0.7407 0.0356 -0.8045 0.016 Mean dead branch height as % of treetop heigDiptera:Psychodidae YDB sp.1 K * -0.7326 0.0387 0.7559 0.03 Range Branch Height (m) dead only Diptera:Muscidae YDB sp.1 K * -0.8556 0.0067 0.7698 0.0255 Range Branch Height (m) dead only Lepidoptera:Oecophoridae YDB sp. 5 L * 0.7326 0.0387 -0.8452 0.0082 Std Dev Branch Height (m) dead only Orthoptera:Gryllacrididae:Kinemania YDB sp. 1 -0.7991 0.0173 -0.7559 0.03 Std Dev Branch Height (m) live only Diptera:Tachinidae YDB sp.1 K * 0.7259 0.0415 -0.7201 0.044 Sum Cone Volume (m3) dead only Lepidoptera:Oecophoridae YDB sp. 2 L.a * -0.8001 0.0171 0.8118 0.0144 Max Cone Volume (m3) dead only Lepidoptera:Oecophoridae YDB sp. 2 L.a * -0.7638 0.0274 0.7991 0.0173 Std Dev Cone Volume (m3) dead only Lepidoptera:Oecophoridae YDB sp. 2 L.a * -0.7274 0.0409 0.8244 0.0118 Mean Polyhedra Volume (m3) live only Diptera:Tipulidae YDB sp. 3 K * 0.7559 0.03 -0.7173 0.0452 Mean Polyhedra Volume (m3) dead only Hemiptera:Fulgoridae YDB sp.3 * -0.8248 0.0117 0.7412 0.0353 Max Polyhedra Volume (m3) all branches Coleoptera:Scirtidae:Pryonocyphon TFIC sp.01 J.b * 0.7638 0.0274 -0.8085 0.0151 Max Polyhedra Volume (m3) all branches Diptera:Phoridae YDB sp.1 K.c 0.8648 0.0056 0.7326 0.0387 Max Polyhedra Volume (m3) live only Coleoptera:Scirtidae:Pryonocyphon TFIC sp.01 J.b * 0.7638 0.0274 -0.8085 0.0151 Max Polyhedra Volume (m3) live only Diptera:Phoridae YDB sp.1 K.c 0.8648 0.0056 0.7326 0.0387 Std Dev Polyhedra Volume (m3) dead only Hymenoptera: Chalcidoid? YDB sp.1 M -0.8783 0.0041 -0.7109 0.0481 Total Foliage Units Coleoptera:Mycteridae:Trichosalpingus TFIC sp.01 J.d -0.7173 0.0452 -0.7325 0.0388 Range Start Diameter (m) all branches Coleoptera:Coccinellidae:Rhyzobius TFIC sp.05 J * 0.7325 0.0388 -0.7325 0.0388 Mean Scaling Ratio all branches Hymenoptera:Formicidae:Hymenoptera YDB sp. 22 M 0.7559 0.03 0.7326 0.0387 Mean Vertical Arc (deg) live only Diptera:Phoridae YDB sp.1 K.c 0.8154 0.0136 0.7326 0.0387 Mean Down Arc (deg) live only Diptera:Phoridae YDB sp.1 K.c 0.803 0.0164 0.7638 0.0274 Mean Horizontal Arc (deg) all branches Coleoptera:Mycteridae:Trichosalpingus TFIC sp.01 J.d * -0.7303 0.0397 0.8452 0.0082 Mean Horizontal Arc (deg) all branches Diptera:Mycetophilidae YDB sp.2 K 0.7326 0.0387 0.7279 0.0406 Mean Horizontal Arc (deg) live only Araneae:Grouped * -0.8193 0.0128 0.8333 0.0102 Mean Horizontal Arc (deg) live only Coleoptera:Mycteridae:Trichosalpingus TFIC sp.01 J.d * -0.7825 0.0217 0.8452 0.0082

Table 11.7. Spearman’ s rho rank correlations between 107 structural descriptors and 128 abundances of RTU in which both age classes exhibited a p-value<0.05

The significant rank correlations detected for abundance of each RTU are tabulated Table 11.7. Sixteen of 26 correlations had mismatched signs.

Coleopteran RTU responded to measures of crown airspace relative to trunk airspace, relative branch height, maximum branch airspace volume, total foliage, and horizontal branch arc. Dipteran RTU responded to relative branch height and ranges of dead branches, the standard deviation of live branch heights, average and maximum branch airspace volume, and vertical and horizontal arcs of live branches. Only a single Hemiptera RTU exhibited significant responses for both age classes. However, the signs of the correlations were mismatched.

Fulgoridae YDB sp. 3 was correlated with the mean airspace volume of dead branches. Hymenoptera responded negatively to the standard deviation of dead branch airspace volume, and positively to the average scaling ratio of all branches. A lepidopteran, Oecophoridae YDB sp. 2, again showed significant but mismatched correlations, this time with measures of dead branch volume. Spiders exhibited 11) Correlation - Results mismatched responses. A gryllacrid cricket, Kinemania YDB sp. 1 responded negatively to the standard deviation of dead branch heights. 11.2 Multivariate Correlation results

11.2.1 NMS ordination results

Correlations a NMS ordination discriminated the strongest structural predictors of changes in the composition of the arthropod community (Figure 11.5). The unsquared correlation coefficient r is tabulated for correlations where r2 > 0.15 in Table 11.8. Structural descriptors with both positive and negative r-values for an axis are influencing the community in opposing fashions (Figure 11.6)

All RTU with >5 fauna and Structural Descriptors B t05 Age 100yr Old t03 C D t12 t04 A t06

t08 t10 t11 H t02 G t01 t15t09 Axis 2 Axis t14 t13 t16

E

F t07

Axis 1

A, Lowest branch height; B, Mean up arc of all branches; C, Mean vertical arc of live branches; D,Total foliage units; E, Max distance of dead branches; F, % of vector volume in dead branches; G,Mean slope of all branches; H, % total wood volume in trunk r2 cutoff = .250; length of rays is proportional to r2

Figure 11.5: NMS of all RTU with >5 individuals. Joint plot with axis- correlated structural descriptors where r2 >.250. Mean 3-d stress 9.094 11) Correlation - Results

Of the six most strongly correlated descriptors, four were measures of dead branch size and distribution. Other highly correlated descriptors included the average branch slope, the lowest branch height, the number of live branches, and the portion of wood volume in the trunk.

Influence of structural descriptors on taxonomic composition r correlation coefficient with NMS axis: where r2 >0.15; ranked by highest r2 Axis Structural Descriptor 1 2 3 Mean Distance (m) dead only 0.043 0.69 -0.029 Max Distance (m) dead only -0.126 0.673 -0.148 Lowest Branch Height (m) all branches 0.176 -0.559 0.147 Mean Slope (deg) all branches 0.023 0.157 0.545 Range Branch Height (m) dead only 0.061 0.54 0.11 % of Vector Volume in Dead Branches -0.326 0.51 0.176 Mean Slope (deg) live only 0.022 0.199 0.501 # Branches live only 0.471 0.128 -0.054 % of Wood Volume in Trunk -0.084 -0.463 0.384 Highest Branch Height (m) all branches 0.455 -0.142 0.317 Range Branch Height (m) live only 0.45 -0.109 0.002 Max Distance (m) live only 0.443 0.075 -0.055 Sum Polyhedra Volume (m3) all branches 0.439 -0.001 -0.171 Mean Up Arc (deg) live only 0.423 -0.34 0.212 Mean Up Arc (deg) all branches 0.418 -0.255 0.215 Sum Polyhedra Surface Area (m2) live only 0.417 -0.079 -0.076 Std Dev Cone Volume (m3) dead only -0.414 0.055 0.085 Std Dev Branch Height (m) dead only 0.035 0.375 0.412 Mean Polyhedra Surface Area (m2) dead only -0.407 -0.04 -0.323 Total Foliage Units 0.406 -0.204 0.001 Sum Polyhedra Volume (m3) live only 0.397 0.013 -0.009 Std Dev Polyhedra Volume (m3) dead only -0.395 0.131 -0.326 Sum Polyhedra Surface Area (m2) all branches 0.393 -0.081 -0.091 Max Polyhedra Volume (m3) dead only -0.387 0.069 -0.315 Mean Foliage Units live only 0.084 -0.387 0.174

Table 11.8: Structural descriptors correlated with NMS ordination of arthropod community for which r2>0.15; sorted by strength of correlation with any axis

Comparison of descriptors correlated along the same axis of arthropod fauna shows opposition and agreement in structural influence (Table 11.8) Organisms differentiated by axis 1 were influenced in one direction by auto-correlated 11) Correlation - Results descriptors of branch height, airspace size, foliage, vertical arc and live branch numbers, and influenced in the other direction by measures of dead branch airspace and size variation.

Dead branches were influential on another axis of composition. The length, range and portion of wood volume associated with dead branches influences taxonomic composition in opposition to the portion of wood volume in the trunk, lowest branch height, and average foliage units per branch. The standard deviation of dead branch height and the average branch slope were correlated with arthropod composition but were not opposed by any other measures.

# Branches live only Highest Branch Height (m) all branches Range Branch Height (m) live only Max Distance (m) live only Std Dev Cone Volume (m3) dead only Sum Polyhedra Volume (m3) all branches Mean Polyhedra Surface Area (m2) dead only Mean Up Arc (deg) live only Axis 1 Std Dev Polyhedra Volume (m3) dead only Mean Up Arc (deg) all branches Max Polyhedra Volume (m3) dead only Sum Polyhedra Surface Area (m2) live only Total Foliage Units Sum Polyhedra Volume (m3) live only Sum Polyhedra Surface Area (m2) all branches

Mean Distance (m) dead only Lowest Branch Height (m) all branches Max Distance (m) dead only Axis 2 % of Wood Volume in Trunk Range Branch Height (m) dead only Mean Foliage Units live only % of Vector Volume in Dead Branches

Mean Slope (deg) all branches Mean Slope (deg) live only Axis 3 Std Dev Branch Height (m) dead only

Figure 11.6: Structural descriptors correlated with NMS ordination of arthropod community for which r2>0.15.. arranged by axis and +/- direction. For each direction, structural descriptors are ranked by descending correlation.

11) Correlation - Results

11.2.2 Mantel test results

Table 11.9: Mantel test results for both age classes pooled and separately for each trap type+3

Mantel tests did not show a significant correlation in the structure and arthropod distance matrices for both age classes pooled, or for each age class separately (Table 11.9). The most probable correlation was for all traps in the 100yr trees (p=0.16), but its likelihood of true significance is contradicted by the least probable correlation for its parallel in the old trees or in both age classes pooled.. No pattern is visible when comparing trap types between age classes and pooled ages. For all comparisons between ages or trap types, mismatched signs and highly variable p- values were observed.

12) Correlation - Discussion

12) Correlation – Discussion

Observations from the correlation analysis were subjectively chosen for discussion. Summaries of entire taxa or entire trap collections were discussed. The possibility that structural traits are surrogate measures of other phenomena are is noted, and a possible causal link between the structural descriptors and arthropod response is proposed.

A table with a selection of correlations worthy of future investigation is presented. Amongst many correlations, it was found that trees with a higher portion of their total wood volume in the trunk rather than the branches had a higher diversity of arthropods. Trees with a larger range of dead branch girths had a less diverse arthropods fauna in the CD sticky traps. More species were collected in the hangtraps in trees with a larger range of live branch girths, a greater upwards arc, or a lower proportional average branch height (relative to total tree height). Trees with shallower crowns had richer Dipteran fauna. Arthropod composition was found to be correlated with measures of crown depth, dead branch size, number and foliage mass of living branches, total crown foliage.

12.1 The influence of crown structure on canopy arthropods of E. obliqua

Several correlations between structural descriptors and measures of arthropod biodiversity were identified. Future assessment of their accuracy and causal relationships will require further research. A distinction bias was present in the researcher’s choice of variables for analysis. Another subjective choice was made in determining what levels of significance to report.

With the number of correlations generated, it is impossible to address each one separately. If one measure of a structural category showed a significant result, it is beyond the scope of this analysis to check if related measures corroborate this correlation. It is possible that the decision to present results at p<0.001 or p<0.05 is hiding a strong pattern of correlations

12) Correlation - Discussion

12.2 Bivariate

Rank correlations used for comparing within each age class may have masked differences in measured structural traits. The old trees express greater variation in their observed structure than the 100yr trees. Ranking trees by a structural descriptor may have applied an even spacing between trees for both age classes, when in fact the old trees were spaced farther apart by the unranked measure. This may account for the greater than expected number of significant results in the old trees for summaries of richness and diversity.

It is more likely that arthropods within the different age classes will correspond to structural descriptors than to respond in opposition. When comparing a strong correlation in one age class to a weak correlation in the other, little weight should be given to the weaker correlation. A high p-value suggests a lack of pattern, rather than an opposite trend. Mismatched signs in combinations were p<0.001 in one age class are less important than mismatched signs when p<0.05 for both age classes.

However, more suspicion should be given to mismatched signs (+/-) with strong correlations in both age classes. These were considered to be most likely evidence of false positives, rather than of truly opposing trends.

12.2.1 All taxa, all trap types

Only one structural descriptor was identified as a predictor of a biodiversity for all taxa, in all traps. There is no known evidence of this relationship in the literature. In the old trees, a greater diversity was rank correlated with the portion of the total wood volume in the trunk. No pattern of correlation was observed in the 100yr trees. A possible explanation is that trees with larger trunks and smaller branches would exhibit a more diverse fauna could be the emphasis in trapping on the trunk. Trees with more trunk habitat than branch habitat may have space for animals that would otherwise be pushed out to the branches. They may have a more diverse fauna because there is more space for them to occupy. This would allow more habitats for a richer fauna, and more space for them to coexist without dominating with each other.

12) Correlation - Discussion

12.2.2 All taxa, one trap type

Correlations were identified for summaries of total diversity in the CD sticky traps, and for total richness in the hangtraps. No structural predictors were identified for the total taxa of funnel traps, or for the total abundance of any trap type.

The total diversity in CD sticky traps in both age classes was negatively correlated with the range of dead branch diameters. Significant, but sign mismatched correlations were found for total crown depth. The diameter ranges for dead branches show variation in branch size as well as the, and could also indicate a greater range of time since branch death. This contradicts Irmler et al.’s (1996) findings that age since death of coarse woody debris to influences the insect fauna. The causal relationship between the two variables is unclear.

The total richness in hangtraps was positively correlated with the mean angular arc upwards on branches in old trees, and with the range in live branch start diameter for both age classes. The average upwards arc could be a surrogate for branch elbowing or epicormic activity on the upper surface of the branch. The range in live branch diameters increases with the presence of large, generally rotten older branches and small, healthy epicormics. If these branch types support a distinct fauna, then the total richness in the tree would increase with variability in live branch sizes.

Hangtrap richness was also negatively correlated with the mean proportional branch height. All of the old trees had a lower mean proportional branch height (Figure 5.7). This was linked to an increased abundance of epicormic branches in the lower crown. This trend could be also driven by proximity to the arthropods associated with the middle strata of rainforest trees. Trees with lower proportional branch height would have lower hangtraps. These traps would be closer to nearby non-Eucalyptus rainforest trees, which support a distinct faunal community (McQuillan, 1993).

12.2.3 One order, all trap types

Ordinal level results for all trap types pooled resulted in few correlations. The diversity of Hemiptera rose with the average branch diameter in 100yr trees. The

12) Correlation - Discussion

phytophagous Hemiptera may be more associated with branches than trunks (CSIRO, 1967). Larger branches would allow more habitat space for these animals, and allow less interference between species (Floater, 2001).

Dipteran richness was negatively correlated with crown depth in both age classes. Trees with shallower crowns had richer fly fauna. It is possible that species of flies were more densely distributed in trees with shallower crowns, and that exposed traps were sampling this more crowded fly fauna. However, abundance of Diptera did not correspond with richness. Crown depth may represent a measure of scale which was not accounted for.

Spider abundance was negatively correlated with mean horizontal arc in the 100yr trees but positively correlated in the old trees. Horizontal arc may be a surrogate for complexity of branchlets. Spider abundance has been linked to structural complexity in several situations (Balfour & Rypstra, 1998; McNett & Rypstra, 2000; Rypstra et al., 1999). The cause of this conflicting trend is unclear. It is possible that the compressed values caused by rankings of the 100yr tree descriptor created a misrepresented trend in the 100yr trees.

12.3 Multivariate compositional predictors

The number, size, height, and total foliage of living branches was identified as factors in determining arthropod composition in opposition to measures of dead branch airspace and size. This could reflects changes in herbivore and saproxylic communities. Martikainen et al. (2000), Grove et al. (2002), and Schiegg (2001) have identified the presence of dead wood as a factor contributing to beetle species richness. These structural descriptors may also be surrogate for overall tree vigour.

The length and crown depth of dead branches, and the proportion of branch volume in dead branches, was also determinant factors in arthropod composition. This composition trend was opposed by the average amount of foliage on each live branch, the lowest branch height, and the portion of wood volume in the trunk. In the bivariate correlations, the portion of wood volume in the trunk was also linked to a higher diversity of arthropods. None of these factors were significantly different between age classes, and therefore these measures are unlikely to be a surrogate for age classes.

12) Correlation - Discussion

Like axis 1, the average foliar units per branch could be a measure of vigour in opposition to the amount of dead branches. In E. obliqua forests on the mainland, Oliver et al. (2000) identified the number of stems, altitude, and litter depth as major determinants of ant and beetle composition. It is possible that the number of stems in these forest is analogous to the number of branchlets, or foliar units, in E. obliqua crowns.

12.4 Surrogate measures

In the context of the current study, it is impossible to fully test the veracity of correlations and to determine the causal mechanisms behind them. It is possible that the compositional changes are actually being driven by cryptic factors that are correlated to the structural descriptors. For example, measures of foliage abundance could be a surrogate for vigour or soil nutrient quality. Wormington et al. (2002) found the height of the tallest trees to positively influence the richness and abundance of marsupials in Queensland eucalypt forests, but comment that these may be reflecting a greater site productivity. Similarly, the correlation with the NMS ordination may actually be surrogates for the age class distinctions.

12.5 Applicability

It must be emphasized that these results are exploratory only. Although the structural predictors and arthropod responses were as comprehensive as possible, replication may have been inadequate. In data mining situations where many variables are tested with few replicates, doubt will always remain. The combination of structural descriptors with arthropod responses was the natural outcome of the crown structure and canopy arthropod measurements. Future analysis of the results at a placement or trap-level may offer more insights into the relationships between the two data sets.

It is clear that some arthropods respond to some structural descriptors. A selection of bivariate correlation is given in Table 12.1. Summaries of arthropod biodiversity responds to structural descriptors such as the portion of wood volume in the trunk, crown depth, start diameters, and upwards arc, but the causal link is

12) Correlation - Discussion

unknown. The composition of fauna responded to several structural descriptors, and the direction and axis that they lined up on offers insight into their relationships. Results from the Mantel tests show that the suite of structural descriptors used in analysis, do not determine en masse the composition of arthropods. It is hoped that future researchers will be able to use these as suggestions for more detailed measurements of structure and more targeted taxa of interest.

Group (+/-) Structural Predictor Summary Arthropod response Old trees with a higher % of total wood volume in the trunk had a more diverse total arthropod fauna Trees with less crown depth had a richer Dipteran fauna Trees with a narrower range of dead branch starting diameters had a more diverse sticky trap catch Trees with a wider range of live branch starting diameters had a richer hangtrap catch Old trees with greater mean upwards arc had a richer hangtrap catch Trees with a lower mean branch height as % of total height had a richer hangtrap catch

Table 12.1: A selection of statements regarding some of the observed correlations between structural predictors and arthropod biodiversity in the sixteen studied E. obliqua.

13) Conclusions

13) Conclusions In this section, the study aims are listed and an assessment is made of their fulfilment. Suggested avenues for future research are identified. The conclusions and implications follow and close this document. 13.1 Mapping Aim 1: Quantitatively assess the differences in crown structure and size between 100yr and old E. obliqua The collected data is the most comprehensive data set currently known comparing the crown structure of mature and old-growth Eucalyptus, and the most comprehensive analysis known of tree level structural descriptors for any species.

The structural descriptor variables illuminate several differences in crown structure between these age classes. Tree level analysis is sufficient to distinguish them and offers a framework for understanding the branch-level development in crown structure of E. obliqua.

Branch mapping allows future researchers a structure to plan studies within the same trees. Just as a map of a city allows a researcher to study its development, or plan their field trip, so can a map of a tree.

Structural descriptor variables are only approximations of the detail within the collected data. Continued analysis at a tree level can continue to compare ratios of descriptors between ages. For example, Jacobs (1955) comments that old crowns have approximately twice the foliage of a mature crown. A similar ratio was observed in the total number of foliar units within the study trees. But how does that doubling compare to other measures of tree size and vigour? The crown airspace of old trees was also doubled between age classes, but the total wood volume increased seven times between ages. This “uncoupling” of growth rates sheds light on resource allocation and growth dynamics for E. obliqua (Lusk et al., 2003).

Future analysis at a branch level could further illuminate differences in crown dynamics, and provide quantitative material to model the competition of branches within the crown, after Ishii &McDowell (2001). Curtin (1970) and Attiwill (1962) formulated several allometric formulae for young E. obliqua, but they did so at a time when old trees were considered “useless veterans” (Jacobs, 1955), and little or no distinction was made between natural and silvicultural regeneration. They did not

13) Conclusions

address old-growth trees in their studies. The next step for future research on Eucalyptus is to study more than just two age classes, after Van Pelt &Nadkarni (2004a) and Ishii & McDowell (2001).

Future comparisons to the collected crown structure information can be performed in the same trees at the LTER. It is likely that the study trees will outlive both the author and the readers of this document. At a later date, observations of the change in branch structure can assist in creating models of branch occlusion, crown senescence, secondary crown development, and other processes of growth and decline in E. obliqua. Furthermore, the structural data offers a ready source of comparisons for other data sets quantifying the architecture of Eucalyptus (Curtin, 1970; Kelly et al., 2004; Van Pelt et al., 2004) or other forest trees (Clement et al., 2001; Clement & Shaw, 1999; Ishii & Wilson, 2001; Sillett, 1999). 13.2 Mapping Aim 2: Investigate the presence of structural features in the crowns of 100yr and old E. obliqua The descriptions of structural features in the present study can only offer the most basic hints to the occurrence and formation of these features. The current project was hindered by the selection, access, and distinction biases. It was impossible to photograph or record every feature which attracted the attention of the field workers, and difficult to fit them into simple categories. Each phenomenon is worth investigating on its own. Tree hollows, in particular, have been studied in great detail. Gibbons & Lindenmayer (2002) have written a book reviewing their value for wildlife conservation. Researchers in Tasmanian E. obliqua at Warra LTER have focused on vertebrate and invertebrate use of hollows (Harrison, 2004; Koch, 2004).

However, few, if any, projects have investigated the development and faunal associations in E. obliqua of features such as dead tops, snapped main stems, burls, bark strip flakes, or flaky armpit bark. It is possible that these features support a distinct animal, epiphyte, fungal, or micro- organism community. By photographing and recording the presence of these features, future researchers can both consider them in their studies, and know which trees they can be found in. A future research project to assess the animal communities associated with these features can use the information in this document to mitigate difficulties in selection and distinction.

13) Conclusions

13.3 Mapping Aim 3: Modify the conifer mapping technique of Van Pelt et al. (2004b) for Eucalyptus trees A crown- mapping technique developed for use in the coniferous forest of the Western United States is published by Van Pelt et al. (2004b). This protocol was trialled in Eucalyptus forests in Victoria. In addition to the crown-mapping methods, a technique for quantifying forest stand structure and foliage distribution is described. Their crown mapping methods, as described, were modified for mapping the crowns of E. obliqua in the present research. Pocometric principles (Takenaka et al., 1998) were used for 3-dimensional surveying of vectors. The techniques developed were directly descended and inspired from the conifer-mapping methods but were changed in several ways:

a) Using spherical vectors with relative anchor points eliminates the dependence on a fixed vertical axis from the origin for cylindrical coordinates. Vectors branches are measured in pocometric space from their starting point, independently from any other points. Cylindrical coordinate systems do not work for all trees. Trees with a strongly leaning trunk do not have a central axis. In trees with contending branches, trunk snaps, or suppressed leaders, the central trunk axis may not be present above a certain point. It is unclear in their methods if azimuths and distances are always fixed to the central axis, or if they are recalculated relative to their parent vector.

b) The adoption of a pocometric spherical coordinate system instead of a horizontal extent allows for a more intuitive measurement of a branch than a cylindrical coordinate system, and easier virtualization using CAD software. The length of a branch can be interpreted immediately, whereas the horizontal extent of a branch must be trigonometrically combined with slope to interpret the length of a branch.

c) Asymmetrical foliage distributions around the measured branch are addressed by separate angular measurements of arc for down, up, left and right, rather than by a symmetrical depth and spread.

13) Conclusions

d) Using spherical instead of cylindrical coordinates eliminates the need for an expensive clinometer and laser rangefinder package with internal trigonometric functions.

e) Trunk anchor azimuths are recorded to represent non-radial branches.

f) Branches are defined with reference to resolution levels, following the advice of Moffett (2000). The distinctions between limb segments, branch segments, and branches are removed, as truly reiterated secondary trunks do not exist in Eucalyptus. In these trees, the architecturally reiterated sub-unit of Halle (1995) is the foliar unit, which topologically is below the resolution level of both Van Pelt et al. (2004b) and the present research.

g) A system using two levels of resolution allows statistically valid comparison between branches- the definition of the trunk allows for a standard definition of a branch. In Van Pelt et al. (2004b)’s context of studying foliage and wood volume, counting parent and children branches as distinct units may be justified, but misrepresents the actual topology and confounds branching generations.

h) The limb segment-branch topology naming system is abandoned and replaced with a system based solely on observed branching generation and not by the characteristic of reiterated secondary trunks.

Researchers wishing to map angiospermous trees may benefit from these modifications. However the mapping technique used in the present project was not as effective as other researchers’ in some aspects.

i) The biomass, leaf area, and leaf count of foliar units was not calibrated to each tree.

j) The tools used for measuring slope and distance (clinometer, Leica Disto laser and tape measure) were less precise than the

13) Conclusions

combination clinometer/laser (Impulse) used by Van Pelt et al. (2004b).

k) In the present study, a separate data field was not utilised to record whether a branch was original or epicormic. 13.4 Mapping Aim 4: Develop a technique for displaying crown structure of forest trees using computer models based on 3-dimensional spherical coordinates The study trees are individual organisms that have survived a century or more in a far corner of the world. In mapping, photographing, and discussing these trees, the unique shape of each tree is recorded. The old trees are a record of half a millennium of growth, and a picture of their form offers a link to the past. The arborograph models are presented in an Appendix 1. In addition, throughout this document photographs, focused arborographics, and text descriptions are provided to assist in perception of the variability in branching architecture and crown dynamics. The use of a computerized measurement of visual information as a surrogate of tree complexity is unknown in ecological literature

The virtualization of plant structure is a vibrant field. Several strands of thought are being reconciled: mapping with modelling, plant physiologist with computer scientist, and ecologist with agriculturalist (Godin, 2000; Godin et al., 2004; Hanan & Room, 1997). The present work contributes to the field by refining existing techniques of crown mapping (Van Pelt et al., 2004) and offering detailed instructions for virtualization in a computer aided design environment. This method was applied in trees of exceptional size and the arborographs presented are currently the most advanced known virtual representations of trees of their size.

The 3-d computer arborographs represent a parallel technique to the established methods of photographs and hand-illustration (Pakenham, 1996; Van Pelt, 2002). All three techniques show a tree in a different, complementary, way. Drawing trees is an ancient practice, and computer graphics offer a modern approach to precisely displaying an actual tree. Future uses and improvements of the data set and virtualization could include:

a) the generation of a software tool to eliminate the need for manual data conversion

13) Conclusions

b) an improved depiction of Eucalyptus foliage units

c) graphic animations of the arborographs, such as spinning around the trunk

d) a satisfactory solution to the problem of depicting little traps in large trees

e) posting of the arborographs on the internet with 3-d view control, photographs, structural data, and arthropod trap collection data. Available at http://www.geog.utas.edu.au/yoav

f) submission of the structural data set to the International Canopy Networks Big Canopy database at http://canopy.evergreen.edu/bcd/

The data archive can be viewed at http://scidb.evergreen.edu/databank/studycenter/ydbtasmania

Work with the International Canopy Network in Olympia continues to further develop 3-d visualisation tools. 13.5 Mapping Aim 5: Expand Jacobs’ (1955) theory of intra- branch competition in Eucalyptus saplings to mature and old- growth E. obliqua Integrating Halle’s (1995) concept of competing branches with Ishii & McDowell’s (2001) analogy of branch age to tree age, Jacobs’ (1955) illustration of sapling branch dynamics was extended to mature and old-growth Eucalyptus. At 100 years of age, crown structure is formed by similar dynamics to a sapling less than ten years old. At some stage between 100 and 500 years of age, E. obliqua no longer follows a regular pattern of upper original branch growth and lower crown mortality . Branch mortality from catastrophic crown fires or gradual branch senescence is balanced by the release of epicormic buds within the crown.

The concepts outlined in the model of crown structural development in old trees will be recognized by those familiar with Eucalyptus trees. However, no previous study is known that has addressed these dynamics for this genus in writing since Jacobs (1955). In addition, the computer 3-d arborographs offer empirical, visual evidence of Jacobs’ (1955) theoretical line drawings. This knowledge could be useful to arborists seeking to preserve individual Eucalyptus tree. Also, foresters

13) Conclusions

seeking to conserve biodiversity by managing for old-growth crown structures can gain insights from the analogy of branches to competing trees of different ages. 13.6 Mapping Aim 6: Generate a predictor data set to explore the influence of crown structure on canopy arthropods In tandem with Trapping Aim 4, the descriptor variables used for analysis of crown structural development can be used as a predictor data set to explore the influence of crown structure on canopy arthropod biodiversity. Mopper et al. (1991) describe how herbivory can influence the structure of plants. While outside the scope of this study, the crown structural descriptors are also response variables to an unmeasured set of environmental and experiential predictors. 13.7 Trapping Aim 1: Determine what differences exist in arthropod biodiversity between 100 year old and old growth Eucalyptus obliqua The results of the present study are the first known comparisons of Eucalyptus canopy arthropod biodiversity associated with different aged trees. Several differences in abundances, richness, diversity, community structure, and composition have been described.

After extensive literature search, only a single publication was found addressing the arthropod community of native Eucalyptus forests, or trees of different ages. Abbott et al. (1992) looked at foliage arthropods of sapling and resprout E. marginata, but did not address results at a tree level. No value of foliage mass per tree was given to allow scaling up. In their paper, this study is one of four concurrently reported projects. In an article spanning eleven pages, it receives exactly two paragraphs and one table. No other research project has addressed age-related changed in Australian Eucalyptus.

Old-growth forests are being replaced at an accelerating rate with young eucalypt plantations. Studies addressing the changes in biodiversity associated with this practice are urgently required to assess and mitigate its impact.

Concurrently, at Warra LTER, other research projects have looked at E. obliqua of similar age classes to those of the present study. Harrison (2004) has investigated the interior habitats of live Warra E. obliqua immediately after cutting

13) Conclusions

them down, and Yee et al. (2001) has investigated rotting logs of different ages and sizes (Bashford et al., 2001). 13.8 Trapping Aim 2:Contribute to the knowledge of Eucalyptus canopy arthropods Very little is known about the canopy arthropods of Eucalyptus (Majer et al., 1997), despite it being one of the most prominent tree genera in the world. Ecologically, economically, socially, and spiritually, Eucalyptus typifies the Australian continent. Despite this, surprisingly few researchers have studied the arthropods in their crowns.

Furthermore, little work has been done on the arthropod communities of very old Eucalyptus, living or dead (Grove et al., 2002; Grove & Bashford, 2003; Majer et al., 1997; Yee et al., 2001). In the politically and economically prominent wet sclerophyllous Eucalyptus forests of Tasmania, only Grove et al. (2002) has sampled canopy arthropods from old-growth Eucalyptus.

Beyond the comparison between tree ages analysed in this document, the data collected can be used to address other questions about Eucalyptus arthropods, beyond the scope of the current thesis document. What differences exist between upper and lower crowns? Does the trap collection overlap with the canopy fogging at Warra (Bashford et al., 2001)? How much overlap is there with the beetle collections from felled trees (Harrison, 2004)? Do dead branches have a distinct fauna from live branches?

Voucher specimens of all animals will be archived in the Tasmanian Forest Insect Collection at Forestry Tasmania in Hobart, donated to the Australian National Insect Collection in Canberra, or sent to interested taxonomists. Voucher photographs will be packaged with raw and processed data and made available electronically. 13.9 Trapping Aim 3: Develop robust, inexpensive trap designs suitable for transport to and use in E. obliqua The present research is the first available data from passive trapping in the upper crown of Eucalyptus trees. Two other projects have used passive traps in eucalypt crowns, but unfortunately no results are available at the present time. Larson (2004) placed sticky and flight-intercept traps in the crown of E. grandis in

13) Conclusions

subtropical rainforest, and the author of this document placed sticky traps in the crowns of E. regnans in Wallaby Creek, Victoria.

The traps designs presented here are inexpensive, robust, and simple to make. The concepts behind the designs have been tried and tested in Eucalyptus trees and on ropes in the canopy of other forest trees (Basset, 1991; Basset et al., 1997; Bickel & Tasker, 2004; Majer et al., 2003; Wilkening et al., 1981). By presenting details of construction and placement, and by tabulation of the time, materials, and monetary effort involved, it is hoped that other researchers will find them useful and appropriate for their studies. In particular, the use of CD cases for sticky panels offers an easy way to transport these notoriously messy traps.

A word of caution to any research intending to place traps while climbing on ropes in trees: great care must be given to the sequence of events and the risk of dropping items. The placement and timing of labels, hammers, nails, tape, bottles, and any other elements must be planned meticulously. Lanyards need to be attached to any critical tools that must not be dropped. Important items, like carefully printed trap labels, are worse than useless when they are drifting through the canopy in the wind. Ink labels may dissolve in alcoholic trapping fluids. Small items such as bottlecaps and pens can make climbing into the tree useless. For example, in the pilot study, a staple gun was used to anchor cardboard sticky traps to the stringy bark. Reloading the gun required releasing a spring load staple cartridge. If this was not done carefully, it would have be very easy to launch the staple cartridge right out of the tree crown! 13.10 Trapping Aim 4: Generate a response data set to explore the influence of crown structure on canopy arthropods A response data set at a tree level was generated using three sets of summaries and the abundances of each RTU. For each of the biodiversity aspect summaries of abundance, richness, and diversity, the 24 values for alltraps +3 and alltaxa +5 were tabulated. The abundances of the 118 RTU with abundances greater than five were tabulated. These 190 variables were the response variables to the 107 descriptors of crown structure.

Carey (1996) points out that any attribute of the arboreal ecosystem could be considered of import. The sum total interactions involved in the Eucalyptus treetop

13) Conclusions

ecosystems are vastly more complicated, stochastic, and variable than we can imagine. Exploratory research will never provide satisfactory and clear answers as to what is going on up there, but can only offer a few tantalizing clues.

The results of the exploration can guide future researchers in planning studies targeting a narrower range of structures or taxa. Despite the limitations inherent in the automated comparison of the data sets, it offers useful insights into the influence of crown structure on the canopy arthropods in E. obliqua. No comparable work is known for arboreal invertebrates. 13.11 Conclusions: Answering the Research Questions

1) What are the structural differences between 100 year old and old- growth E. obliqua, and how can they be measured and displayed?

Quantifiable structural differences exist between the 100 year old and old- growth E. obliqua at Warra LTER. The old trees have more variable crowns and a greater abundance of recognized structural features. Different dynamics of branch competition determine their crown structure. Intra-crown branch competition in old- growth E. obliqua trees is distinct from the mature 100 year old trees. Old trees have either senescent primary crowns or resprouted epicormic crowns, whereas 100 year old trees only have mature young crowns.

2) How is the canopy arthropod biodiversity different in 100 year old and old-growth E. obliqua?

High levels of arthropod biodiversity were found in the crowns of both 100yr and old growth trees. More than 300 recognizable taxonomic units were collected in only 3 months in only sixteen trees. Because spider, mites, and ants were unsorted, this value is certainly an underestimate. Furthermore, the traps types undoubtedly failed to collect animals that are sessile, only in the outer crown, or living inside the tree.

Evidence was collected that suggest higher levels of arthropod biodiversity exist in the crowns of the old-growth trees. The composition of arthropods collected in the crowns was different between the age classes. Changing the demographics of

13) Conclusions

the forest by replacing multi-aged stands with even-aged ones will impact the arthropods associated with both of these age classes. While only sampling two points in the lifespan of E. obliqua, the results of this study indicate the old growth trees support a greater and different arthropod biodiversity than the 100 year old trees. This suggests, in agreement with the literature for vertebrates, that older Eucalyptus trees sustain a greater and different arthropod biodiversity than younger trees.

3) In what ways does crown structure influence arthropod biodiversity in E. obliqua?

The arthropod biodiversity responses to tree structural descriptors was explored. Several correlations between structural predictors and arthropod responses of composition, abundance, richness, and diversity are identified and explored. These are presented as promising avenues of future investigation.

The structural results show that trees one century in age have yet to develop characteristically old-growth crowns, and by extension are probably not supporting a characteristically old-growth fauna. It is important not to forget that the old-growth trees potentially represent a larger range of ages than the 100yr trees. It is possible that the crown fire event that differentiated the old-growth crowns also impacted the arthropod fauna. Future analysis of the data will test the hypothesis that senescent primary crowns and resprouted epicormic old-growth crowns have different arthropod biodiversity. Future research linking the presence of structural features to arthropod biodiversity will be invaluable in assessing the impact of losing these potential keystone structures. An important direction for future research in the canopy of Eucalyptus forests is to look at even younger trees (i.e. within the planned silvicultural rotation of 80-100 years) in comparison with these two age classes.

Rank correlations between crown structure and arthropod biodiversity identified potential structural predictors of arthropod responses. Trees with a higher portion of their total wood volume in the trunk rather than the branches had a higher diversity of arthropods. Trees with a larger range of dead branch girths had a less diverse arthropods fauna in the CD sticky traps. More species were collected in the hangtraps in trees with a larger range of live branch girths, a greater upwards arc, or a lower proportional average branch height (relative to total tree height). Trees with

13) Conclusions shallower crowns had richer Dipteran fauna. Arthropod composition was found to be correlated with measures of crown depth, dead branch size, number and foliage mass of living branches, total crown foliage. The causal relationships behind these correlations remains to be tested by future investigation.

13.12 Future directions Very little research has been performed in situ in the Eucalyptus forest canopy. Very few projects have mapped tree crowns in detail. Very little is known of the biodiversity living in the Eucalyptus canopy. No other research is known for any plant species linking detailed crown structural mapping with a survey of canopy arthropod biodiversity.

Promising avenues for scientific research include investigations of the detected correlations between crown structure and arthropod biodiversity, the linking of structural features to animal communities, the mapping of crown structures in forest trees, and the study of canopy invertebrates. Eucalyptus forests are of world significance and all people would benefit by learning more about these magnificent trees. By providing branch structural data, information about structural features, 3- dimensional tree maps, arthropod voucher collections, suggestions on trap designs and placement, and access cords in place for the study trees at Warra LTER, it is hoped that other researchers will climb into these magnificent trees and benefit from the results of this exploration.

13) Conclusions

Please visit the WWW for more resources, information, and access to raw datasheets:

Project web page:

http://www.geog.utas.edu.au/yoav

Data Archive at the International Canopy Network:

http://scidb.evergreen.edu/databank/studycenter/ydbtasmania

Future correspondence:

[email protected]

14) References

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R

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15) Arborograph Lineup

15) Appendix 1. Arborogaph Lineup (Placeholder for print version)

Figure A15.1: Old Tree 10, down view 16) Arborograph Script Instructions

16) Appendix 2. Instructions for Generating Arborograph Scripts Arborographs composed of six layers Six layers were created for each tree: 1. Trunk 2. Cone 3. Line 4. Cross 5. Kite 6. Extras

Figure A16.1: Layers used in CAD arborographics. The scripts used in generating these images can be found in Appendix 2. Data preparation Using Microsoft Notepad, Word and Excel (Microsoft 2000), the field data was organized into columns and transformed into the text file scripts by placing the text of commands as columns between variables. This data format is distinct for Microstation, but can guide generation of similar scripts for other 3-d CAD software. Generation of layers In the virtual space, the centre of the base of the tree was the origin, where x=0, y=0, and z=0. Text as put into the program is presented in bold, variables are presented in bold italics, and annotations describing are in italics. Each line of text is preceded by a number. Variable transform

16) Arborograph Script Instructions

First, all circumference measurements were converted to radii in metres. A negative sign was added to the azimuths, and slope was subtracted from 90 (see XYZ conversion, above). In the context of describing computer input, reference is made to these transformed values. Coordinate systems input Vector input was in two formats: 3-d Cartesian XYZ (prefixed in the software as and hereafter “xy=”) and spherical (“ax=”). XYZ coordinates are always relative to the origin. Spherical coordinates are relative to the active point. Microstation can also interpret cylindrical coordinates, but this ability was not used. To place an object or move the active drawing point, vectors are defined. XYZ vectors are entered: 1) xy= X, Y, Z Spherical vectors are entered as 1) ax=distance, azimuth, slope

Trunk segments The trunk segments were defined by using the height and radius of the bottom ring, and the height and radius of the top ring. Because the trunk stretches surrounds the Z axis, and distance along Z axis is known, absolute xy= terms were used. 1) place cone radius (defines a cone by its radius) 2) top radius 3) bottom radius 4) xy=0,0,top ring height 5) xy=0,0,bottom ring height

Branch cones Branch vectors are used to fixing a cone in space. The process by which the starting point was determined is described in the section. Input for a branch cone: 1) place cone radius 2) start radius 3) end radius 4) ax=0,0,0 (active point is at starting point) 5) ax=distance, azimuth, slope

Lines Kites, crosses, and boundary lines are created by entering: 1) place line

16) Arborograph Script Instructions

2) ax=start point distance, start point azimuth, start point slope (if the active point is already at the start point, this is ax=0,0,0) 3) ax=end point distance, end point azimuth, end point slope (alternately, xy could be used for absolute coordinates)

Trunk Sprout The trunk sprout point for each vector originating from the trunk was computed using the branch attachment height, trunk radius at branch height, and anchor azimuth. From the origin point, the active point was moved: 1) xy=0,0,height (move up the centre of the trunk) 2) ax=radius at height, anchor azimuth, 90 (move out to trunk surface) Topological trace Whole branches begin at the trunk sprout point. To fix the point at which children branches begin, the vectors of their parent lineage was followed in sequence. From the trunk sprout, the active point was moved: 1) ax=1st gen distance, 1st gen slope, 1st gen azimuth (move along first generation basal vector) 2) ax=2nd gen distance, 2nd gen slope, 2nd gen azimuth (move along next generation) 3) (continued until starting point of current branch is reached) When a distance along parent was recorded, that value replaced the distance term at of the last line. This topological vector sequence prefixed all children branches. An alternative method is to add the converted XYZ vectors of each parent, taking care to generate an alternate XYZ vector when a distance along occurs. Live/Dead Colours As a prefix to each of the four branch layers, the active colour was changed depending on whether the branch was living or wholly dead. Grey was used for dead branches, and brown for live branches. By separating these categories into colours, rendering different materials is facilitated. 1) active color colour (replace final word with desired colour code, American spelling required) Layers construction 1) The trunk is a series of vertically stacked truncated cones. Each segment has a start and end radius defined by the trunk rings. The trunk layer was generated separately to the other layers. The script was generated by sequentially listing the input for each trunk segment defined above: Base segment, next segment ... top segment

2) A truncated or complete cone represents each terminal and segment branches.

16) Arborograph Script Instructions

Each branch was entered sequentially in the script using the components described above: trunk sprout point, topological trace, branch cone.

3) Four boundary lines extend from the base of each branch to the end of the boundary airspaces; these are the down, up, left, and right vectors. When the angular arc in one of these four directions was zero, the boundary line is invisibly present along the cone vector. The line script follows the topological trace once and, staying there, sprouts four lines: trunk sprout point, topological trace, downline, upline, rightline, leftline.

4) Two cross lines connect the boundary line ends. Down to up forms the vertical line; left to right forms the horizontal line. trunk sprout point, topological trace, line up-down, line left-right

5) The kite is a sequence of four lines connecting the end of the boundary airspaces: trunk sprout point, topological trace, line up-left, line left-down, line down-right, line right-up.

6) Extra pieces include: a compass rose, a vertical scale, virtual lighting for graphical rendering, and experiments with depicting traps and structural features. Despite accurate representations of traps and features, screen and print resolution inhibits showing objects ~100 cm2 on a tree 70 m tall. Example scripts are included in the Appendices.

Figure A16.2: Screen capture of Microstation CAD. Use of text scripts eliminates the need for all of the manual tools cluttering the display.

Figure A 16.3: The computer images represent the data which represent the measurers vision of the tree. Tree idiosyncrasies such as fusions or contact between branches can result in seemingly impossible renderings. The arrborograph of old tree 14 shows a seemingly impossible fusion where a branch is in close contact with the tree trunk .

17) Sample Arborograph Script 17) Appendix 3. Sample Arborograph Script for Old Tree 3 Script derived from data sheets in Appendix 1. Suitable for computer interpretation. Read columns downwards, and then across. Trunk define acs points place cone radius xy=0,0,55 0.397899093 0.523635206 0.849912462 0.767149451 xy=0,0,10 xy=0,0,1.3 spherical 0.17985039 xy=0,0,50 xy=0,0,45 xy=0,0,35 xy=0,0,25 xy=0,0,20 xy=0,0,7.5 xy=0,0,40 xy=0,0,30 xy=0,0,23.4 xy=0,0,15 place cone radius xy=0,0,0 0.288078943 place cone radius place cone radius 1.082285532 xy=1,0,0 xy=0,0,57.4 place cone radius place cone radius place cone radius place cone radius 0.767149451 0.884927582 1.268502308 xy=0,1,0 xy=0,0,55 0.28489575 0.397899093 0.523635206 0.849912462 0.854687251 0.926309088 xy=0,0,1.3 0.331052045 move acs 0.455196562 0.635046952 0.913576317 xy=0,0,15 xy=0,0,7.5 xy=0,0,0 xy=0,0,50 xy=0,0,40 xy=0,0,30 xy=0,0,23.4 xy=0,0,10 xy=0,0,5 xy=0,0,0 place cone radius xy=0,0,45 xy=0,0,35 xy=0,0,25 xy=0,0,20 place cone radius active color 54 0.288078943 place cone radius 0.926309088 0.28489575 place cone radius place cone radius place cone radius place cone radius 0.854687251 1.082285532 0.331052045 0.455196562 0.635046952 0.913576317 0.884927582 xy=0,0,5

Cone define acs points move acs 0 move acs ax=6.5,-215,25 ax=0.7,-90,85 ax=0.7,-90,85 move acs xy=0,0,32.2 spherical ax=0.100942,-350,90 move acs ax=0,0,0 move acs xy=0,0,43.9 move acs active color 78 xy=0,0,35.5 move acs xy=0,0,0 xy=0,0,51.7 ax=4.25,-150,35 xy=0,0,46.4 move acs move acs ax=.35,-10,75 place cone radius move acs ax=0.283663,-180,90 xy=1,0,0 active color 78 move acs move acs ax=0.1890685,-10,90 xy=0,0,43.7 active color 78 0.140127389 ax=0.2569825,-320,90 move acs xy=0,1,0 place cone radius ax=0.1260055,-10,90 move acs ax=0.168856,-190,90 move acs move acs place cone radius 0.143312102 ax=1.45,-180,40 move acs 0.038216561 xy=0,0,48.9 move acs ax=0.6,-10,30 ax=0.1906855,-260,90 0.049363057 ax=0,0,0 active color 144 move acs xy=0,0,0 0 active color 78 move acs ax=1.05,-170,70 move acs move acs 0 ax=1.4,-10,75 place cone radius ax=1.2,-200,10 move acs ax=0,0,0 place cone radius ax=0.1486435,-320,90 active color 144 ax=1.1,-70,70 ax=3.2,-260,40 ax=0,0,0 0.054140127 active color 144 xy=0,0,57.4 ax=4.25,0,80 0.101910828 place cone radius active color 78 active color 78 ax=2.5,0,20 move acs 0.047770701 place cone radius move acs 0 active color 144 0.066878981 place cone radius place cone radius xy=0,0,40.8 ax=0,0,0 0.060509554 ax=0.079921,-250,90 move acs ax=0,0,0 place cone radius 0 0.087579618 0.108280255 move acs move acs ax=1.3,-340,65 0 xy=0,0,54.1 ax=5.5,-10,85 0.100318471 ax=0,0,0 0 0 xy=0,0,40.8 ax=0.214132,-90,90 ax=0,0,0 active color 144 move acs 0 ax=7.5,-100,95 ax=0,0,0 ax=0,0,0 move acs move acs ax=2.25,-110,75 place cone radius ax=0.1066015,-300,90 move acs ax=0,0,0 ax=5.5,-15,65 ax=9.5,-322,60 ax=0.214132,-90,90 active color 78 xy=0,0,35.5 0.128980892 xy=0,0,51.5 ax=2,-320,60 move acs move acs place cone radius move acs move acs 0 active color 144 move acs xy=0,0,46.4 move acs move acs ax=0.7,-90,85 0.170382166 ax=0.2569825,-320,90 xy=0,0,32.2 ax=0,0,0 place cone radius ax=0.1276225,-5,90 move acs move acs xy=0,0,43.9 xy=0,0,43.7 move acs 0.140127389 move acs ax=4.5,-250,40 0.041401274 xy=0,0,48.5 ax=0.168856,-190,90 move acs move acs ax=.8,-10,75 ax=0,0,0 active color 144 ax=0.283663,-180,90 0.007961783 active color 78 move acs move acs ax=0.1890685,-10,90 ax=0.1906855,-260,90 active color 144 ax=0.7,-90,85 place cone radius move acs move acs ax=0,0,0 place cone radius ax=0.1518775,-350,90 ax=1.05,-170,70 move acs move acs place cone radius 0.076433121 ax=1.45,-180,40 xy=0,0,57.4 ax=3.25,-170,90 0.055732484 active color 144 ax=0.6,-10,30 ax=3.2,-260,40 0.02866242 move acs 0 move acs move acs 0 active color 144 place cone radius move acs active color 78 0.015923567 xy=0,0,40.1 ax=0,0,0 ax=.7,-200,10 ax=0.079921,-15,90 move acs ax=0,0,0 place cone radius 0.074840764 ax=1.1,-70,70 place cone radius ax=0,0,0 move acs ax=1.6,-320,75 active color 144 xy=0,0,54.1 ax=3.5,-5,40 0.159235669 0 active color 78 0.111464968 ax=0.3,-340,5 ax=0.2197915,-330,90 place cone radius active color 78 move acs 0 ax=0,0,0 place cone radius 0 move acs 0.02866242 place cone radius ax=0.1066015,-300,90 move acs ax=0,0,0 ax=8.25,-140,60 0.127388535 ax=0,0,0 move acs active color 78 xy=0,0,34.5 0.00955414 0.036624204 xy=0,0,50.8 ax=5.5,-350,65 0 ax=10,-340,75 xy=0,0,40.8 place cone radius move acs ax=0,0,0 0 active color 78 move acs move acs ax=0,0,0 move acs 0.128980892 ax=0.2650675,-60,90 ax=1.4,-200,20 ax=0,0,0 place cone radius ax=0.133282,-100,90 move acs xy=0,0,46.4 ax=7,-90,50 move acs ax=0.214132,-90,90 0 ax=6,-15,80 0.116242038 xy=0,0,47.2 move acs xy=0,0,43.7 move acs ax=0,0,0 active color 78 move acs 0 active color 78 move acs ax=0.168856,-190,90 move acs move acs ax=0.7,-90,85 ax=8,-330,60 place cone radius xy=0,0,32.2 move acs ax=0,0,0 place cone radius ax=0.162388,-16,90 xy=0,0,43.9 ax=0.1906855,-260,90 move acs 0.132165605 move acs xy=0,0,57.4 ax=9.5,-220,60 0.073248408 move acs active color 144 move acs ax=.9,-10,75 move acs 0 ax=0.283663,-180,90 move acs 0 ax=.9,-16,80 place cone radius ax=0.1890685,-10,90 active color 78 active color 144 xy=0,0,37.9 ax=0,0,0 move acs ax=0.079921,-135,90 move acs ax=0,0,0 active color 144 0.116242038 move acs place cone radius place cone radius move acs ax=8.5,-60,70 ax=1.45,-180,40 xy=0,0,53.1 ax=5,-100,60 place cone radius 0.097133758 ax=0.6,-10,30 0.211783439 0.031847134 ax=0.2375785,-320,90 move acs active color 78 move acs 0.065286624 ax=0,0,0 active color 78 0.191082803 0.031847134 move acs ax=.55,-200,10 place cone radius ax=0.1146865,-300,90 move acs 0 ax=1.05,-170,70 place cone radius ax=0,0,0 ax=0,0,0 active color 78 xy=0,0,33.4 active color 144 0.175159236 xy=0,0,50.2 ax=0,0,0 0.140127389 ax=3.2,-260,40 ax=0.4,-160,10 place cone radius move acs place cone radius 0 active color 78 move acs ax=7,-40,50 move acs 0.130573248 0.092356688 ax=0.273961,-350,90 0.050955414 ax=0,0,0 place cone radius ax=0.138133,-50,90 xy=0,0,46.1 ax=0,0,0 move acs move acs 0 move acs 0.047770701 ax=15,-135,5 0.062101911 move acs move acs ax=1.1,-70,70 xy=0,0,43.6 xy=0,0,40.8 ax=0,0,0 ax=1.35,-280,80 ax=0,0,0 0 active color 78 xy=0,0,47.2 ax=0.1712815,-160,90 move acs move acs ax=5.75,-300,65 active color 78 ax=0.75,-160,15 move acs ax=0,0,0 place cone radius move acs move acs ax=0.191494,-120,90 ax=0.214132,-90,90 place cone radius xy=0,0,56.4 ax=5.5,-10,70 0.049363057 ax=0.162388,-16,90 active color 78 xy=0,0,43.9 move acs move acs 0.058917197 move acs move acs 0 move acs place cone radius move acs active color 78 ax=0.7,-90,85 xy=0,0,37.9 0 xy=0,0,32.2 ax=0.088006,-60,90 move acs ax=0,0,0 ax=.9,-16,80 0.095541401 ax=0.1890685,-10,90 place cone radius move acs move acs ax=0,0,0 move acs xy=0,0,52.9 ax=3.5,-50,75 active color 144 0 0.068471338 ax=1.1,-10,75 ax=0.2375785,-320,90 ax=4.75,-170,30 ax=0.283663,-180,90 active color 78 move acs place cone radius ax=0,0,0 active color 78 0 active color 78 move acs place cone radius ax=0.1163035,-160,90 move acs 0.079617834 ax=9,-160,80 place cone radius ax=0,0,0 place cone radius active color 78 move acs ax=1.45,-180,40 0.073248408 xy=0,0,50.1 0 0.178343949 ax=4.25,-140,55 0.035031847 place cone radius xy=0,0,33.4 move acs 0 active color 144 move acs ax=0,0,0 move acs 0.167197452 0 0.101910828 move acs ax=.4,-200,10 ax=0,0,0 place cone radius ax=0.1389415,-245,90 ax=7.5,-55,75 xy=0,0,44.8 ax=0,0,0 move acs ax=0,0,0 0 ax=0.273961,-350,90 active color 78 ax=5.5,-60,70 0.055732484 move acs ax=0.6,-10,30 xy=0,0,42.5 ax=1.2,-260,5 ax=0,0,0 move acs place cone radius 0.007961783 active color 144 move acs ax=0.181792,-320,90 move acs ax=4.75,-340,45 ax=1.35,-280,80 0.035031847 move acs ax=0,0,0 place cone radius xy=0,0,47.2 move acs ax=0.2003875,-320,90 move acs active color 78 0 xy=0,0,56.1 ax=0.45,-160,60 0.108280255 move acs active color 78 xy=0,0,43.7 xy=0,0,40.8 move acs place cone radius ax=0,0,0 move acs 0.047770701 ax=0.162388,-16,90 place cone radius move acs active color 144 move acs xy=0,0,37.5 0.103503185 ax=2.5,-180,45 ax=0.0904315,-170,90 move acs ax=0,0,0 0.030254777 ax=0.1906855,-260,90 place cone radius ax=0.214132,-90,90 move acs 0 xy=0,0,52.9 ax=4,-235,70 active color 144 0 move acs 0.057324841 move acs ax=0.2408125,0,90 ax=0,0,0 move acs active color 144 move acs place cone radius ax=0,0,0 ax=.85,-260,40 0.039808917 ax=0.7,-90,85 ax=6.25,-320,65 xy=0,0,32.2 place cone radius ax=0.1163035,-140,90 move acs 0.10186216775425752 ax=2.2,-320,30 active color 144 ax=0,0,0 move acs active color 144 move acs 0.125796178 xy=0,0,50.1 029285373229349 place cone radius ax=3.25,-320,65 ax=1.4,-10,75 place cone radius move acs ax=0.283663,-180,90 0.111464968 active color 78 move acs 0.08912939678497533 move acs 0.073248408 active color 144 0.070063694 xy=0,0,33.4 move acs ax=0,0,0 place cone radius ax=0.1389415,-245,90 0256247015756804 xy=0,0,44.4 0 move acs place cone radius 0 move acs ax=1.45,-180,40 ax=2.1,-170,40 0.132165605 ax=0,0,0 move acs ax=0,0,0 xy=0,0,42.1 0.02388535 ax=0,0,0 ax=0.273961,-350,90 move acs 0 active color 144 ax=.9,-16,80 ax=0.185026,-260,90 ax=4,-10,80 move acs 0 ax=3.2,0,75 ax=4.5,-200,10 move acs ax=0,0,0 place cone radius ax=0.2036215,-250,90 ax=0,0,0 active color 78 active color 78 xy=0,0,55.5 ax=8,-140,55 0.063694268 move acs active color 78 move acs ax=2.5,-340,15 move acs place cone radius place cone radius move acs 0 xy=0,0,46.7 place cone radius xy=0,0,43.7 active color 144 xy=0,0,36.6 0.111464968 0.124203822 ax=0.0952825,-310,90 move acs ax=0,0,0 move acs 0.02866242 move acs place cone radius move acs move acs 0.108280255 0 xy=0,0,52.2 ax=8,-230,70 ax=0.1664305,-350,90 0 ax=0.1906855,-260,90 0.046178344 xy=0,0,40.8 ax=0.248089,-90,90 ax=0,0,0 ax=0,0,0 active color 78 move acs ax=0,0,0 move acs 0 move acs move acs ax=1.35,-280,80 ax=7.5,-130,65 place cone radius ax=0.121963,-100,90 move acs active color 144 ax=3.5,-220,70 ax=2.1,-260,40 ax=0,0,0 ax=0.214132,-90,90 ax=0.7,-90,85 0.081210191 xy=0,0,50.1 place cone radius active color 144 ax=3.5,-180,50 move acs active color 144 move acs move acs 0 active color 78 move acs 0.078025478 move acs place cone radius ax=0.7,-90,85 place cone radius xy=0,0,32.2 xy=0,0,32.2 ax=0,0,0 place cone radius ax=0.1389415,-245,90 0.081210191 xy=0,0,43.9 0.111464968 move acs move acs 0.079617834 move acs move acs ax=6.5,-20,65 0.078025478 ax=0,0,0 move acs 0.095541401 xy=0,0,41.3 ax=1.4,-10,75 0.063694268 ax=0.283663,-180,90 ax=0.283663,-180,90 0 active color 144 ax=0.65,-350,70 ax=0.1890685,-10,90 ax=0,0,0 move acs active color 78 ax=0,0,0 move acs move acs move acs ax=0,0,0 place cone radius move acs ax=1.15,-240,85 ax=0.2100895,-260,90 place cone radius ax=0.9,-90,5 ax=1.45,-180,40 ax=1.45,-180,40 xy=0,0,55.5 ax=7,-50,40 0.044585987 move acs ax=0.6,-10,30 0.103503185 move acs active color 78 move acs 0 xy=0,0,46.4 move acs move acs active color 78 0 move acs ax=4.5,-200,10 place cone radius ax=0.0952825,-310,90 move acs ax=0,0,0 move acs ax= xy=0,0,43.7 place cone radius ax=0,0,0 xy=0,0,36.6 active color 78 0.242038217 xy=0,0,52.2 ax=6.5,-30,65 ax=0.168856,-190,90 active color 78 move acs 0.030254777 ax=5,-100,30 move acs place cone radius 0.219745223 active color 144 move acs move acs place cone radius ax=0.1906855,-260,90 0 ax=0.248089,-90,90 0.162420382 ax=0,0,0 place cone radius ax=0.121963,-100,90 move acs ax=.5,-170,70 0.031847134 move acs ax=0,0,0 move acs 0 ax=4.5,-200,10 0.085987261 xy=0,0,49.1 active color 144 0 ax=2.7,-260,40 ax=3.5,-230,25 xy=0,0,40.8 active color 144 ax=0,0,0 0.079617834 active color 144 move acs place cone radius ax=0,0,0 active color 78 move acs place cone radius ax=9,-160,50 move acs ax=0,0,0 place cone radius ax=0.1470265,-340,90 0.035031847 ax=3.5,-310,50 place cone radius move acs ax=0.214132,-90,90 0.082802548 xy=0,0,32.2 ax=2.2,-10,55 0.103503185 0.02388535 0.050955414 xy=0,0,41.3 move acs 0.092356688 move acs move acs 0.079617834 active color 144 ax=0,0,0 move acs 0 move acs ax=0.7,-90,85 ax=0,0,0 xy=0,0,32.2 ax=0.283663,-180,90 move acs ax=0,0,0 place cone radius ax=1.5,-180,75 xy=0,0,43.9 ax=0,0,0 ax=0.2100895,-260,90 move acs ax=0.7,-90,85 move acs xy=0,0,55.1 ax=0.85,-15,45 0.111464968 move acs ax=3.5,-20,20 ax=1.4,-10,75 ax=0.283663,-180,90 active color 78 move acs 0 move acs ax=0.1890685,-10,90 active color 144 active color 144 move acs move acs place cone radius ax=0.0985165,-90,90 move acs ax=0,0,0 xy=0,0,46.4 move acs move acs place cone radius place cone radius xy=0,0,35.5 ax=1.45,-180,40 0.313694268 xy=0,0,52.2 ax=3.5,-340,70 move acs ax=0.6,-10,30 xy=0,0,43.7 0.01910828 0.111464968 move acs move acs 0.246815287 active color 78 move acs ax=0.168856,-190,90 move acs move acs 0 0.047770701 ax=0.2569825,-320,90 ax=4.5,-200,10 ax=0,0,0 place cone radius ax=0.121963,-100,90 move acs move acs ax= ax=0.1906855,-260,90 ax=0,0,0 ax=0,0,0 active color 78 ax=1.45,-180,40 0.082802548 xy=0,0,48.9 ax=1.05,-170,70 active color 144 move acs ax=2.2,-300,75 ax=3.5,-30,60 active color 144 place cone radius 0 active color 78 move acs active color 144 place cone radius ax=3.2,-260,40 place cone radius 0.168789809 move acs ax=0,0,0 place cone radius ax=0.1486435,-150,90 place cone radius 0.092356688 active color 78 move acs move acs 0.057324841 0 xy=0,0,31.3 ax=7.25,-90,80 0.063694268 0.020700637 0.047770701 place cone radius xy=0,0,40.8 xy=0,0,40.8 0.02388535 ax=0,0,0 move acs 0 active color 144 0.001592357 ax=0,0,0 0.095541401 move acs move acs ax=0,0,0 ax=7.25,-200,20 ax=0.2909395,-330,90 move acs ax=0,0,0 place cone radius ax=0,0,0 ax=1.4,-20,65 0 ax=0.214132,-90,90 ax=0.214132,-90,90 ax=0.9,-30,85 move acs xy=0,0,54.8 ax=7.5,-350,50 0.044585987 ax=3,-170,45 ax=0,0,0 move acs move acs move acs ax=0.56,-330,80 17) Sample Arborograph Script active color 144 move acs 0.038216561 active color 144 move acs 0 active color 144 move acs place cone radius ax=0.2909395,-330,90 move acs ax=0,0,0 place cone radius ax=0.309535,-320,90 move acs ax=0,0,0 place cone radius ax=0.31762,-40,90 0.030254777 xy=0,0,31.3 ax=1.2,-290,85 0.047770701 xy=0,0,29.0 ax=2.5,-280,55 0.050955414 0.02388535 active color 144 move acs 0.036624204 active color 144 move acs 0.044585987 active color 144 ax=0,0,0 place cone radius ax=0.2909395,-330,90 move acs ax=0,0,0 place cone radius ax=0.309535,-320,90 move acs ax=0,0,0 place cone radius ax=0.7,-350,25 0.046178344 xy=0,0,31.0 ax=1.45,-70,70 0.036624204 xy=0,0,29.0 ax=1.05,-330,70 0.042993631 0.039808917 active color 144 move acs 0.035031847 active color 78 move acs 0.003184713 move acs ax=0,0,0 place cone radius ax=0.293365,-70,90 move acs ax=0,0,0 place cone radius ax=0.309535,-320,90 move acs ax=0,0,0 xy=0,0,31.3 ax=0.56,-330,80 0.031847134 xy=0,0,29.0 ax=0.9,-260,40 0.055732484 xy=0,0,28.0 ax=4.5,-100,50

Line define acs points move acs spherical xy=0,0,55.5 xy=0,0,0 move acs xy=1,0,0 ax=0.0952825,-310,90 xy=0,1,0 move acs active color 112 xy=0,0,0 place line move acs ax=0,0,0 xy=0,0,57.4 ax=2.2,-10,90 move acs ax=0.079921,-250,90 place line ax=0,0,0 active color 112 ax=2.2,-10,20 place line ax=0,0,0 place line ax=4.5,-250,90 ax=0,0,0 ax=2.2,-10,55 place line ax=0,0,0 place line ax=4.5,-250,-40 ax=0,0,0 ax=2.2,-10,55 place line move acs ax=0,0,0 xy=0,0,55.1 ax=4.5,-240,40 move acs ax=0.0985165,-90,90 place line ax=0,0,0 active color 80 ax=4.5,-265,40 place line move acs ax=0,0,0 xy=0,0,57.4 ax=7.25,-90,80 move acs ax=0.079921,-15,90 place line ax=0,0,0 active color 80 ax=7.25,-90,50 place line ax=0,0,0 place line ax=6,-15,90 ax=0,0,0 ax=7.25,-20,80 place line ax=0,0,0 place line ax=6,-15,65 ax=0,0,0 ax=7.25,-140,80 place line move acs ax=0,0,0 xy=0,0,54.8 ax=6,-10,80 move acs ax=0.100942,-350,90 place line ax=0,0,0 active color 80 ax=6,-20,80 place line move acs ax=0,0,0 xy=0,0,57.4 ax=4.25,0,85 move acs ax=0.079921,-135,90 place line ax=0,0,0 active color 80 ax=4.25,0,60 place line ax=0,0,0 place line ax=15,-135,70 ax=0,0,0 ax=4.25,5,80 place line ax=0,0,0 place line ax=15,-135,-145 ax=0,0,0 ax=4.25,-5,80 place line move acs ax=0,0,0 xy=0,0,54.1 ax=15,-45,5 move acs ax=0.1066015,-300,90 place line ax=0,0,0 active color 112 ax=15,-190,5 place line move acs ax=0,0,0 xy=0,0,56.4 ax=3.25,-170,90 move acs ax=0.088006,-60,90 place line ax=0,0,0 active color 80 ax=3.25,-170,90 place line ax=0,0,0 place line ax=5.5,-60,90 ax=0,0,0 ax=3.25,-170,90 place line ax=0,0,0 place line ax=5.5,-60,20 ax=0,0,0 ax=3.25,-170,90 place line move acs ax=0,0,0 xy=0,0,54.1 ax=5.5,-55,70 move acs ax=0.1066015,-300,90 place line ax=0,0,0 active color 80 ax=5.5,-70,70 place line move acs ax=0,0,0 xy=0,0,56.1 ax=9.5,-220,90 move acs ax=0.0904315,-170,90 place line ax=0,0,0 active color 112 ax=9.5,-220,5 place line ax=0,0,0 place line ax=2.1,-170,90 ax=0,0,0 ax=9.5,-200,60 place line ax=0,0,0 place line ax=2.1,-170,-10 ax=0,0,0 ax=9.5,-255,60 place line move acs ax=0,0,0 xy=0,0,53.1 ax=2.1,-170,40 move acs ax=0.1146865,-300,90 place line ax=0,0,0 active color 80 ax=2.1,-170,40 place line move acs ax=0,0,0 xy=0,0,55.5 ax=5.5,-10,90 move acs ax=0.0952825,-310,90 place line ax=0,0,0 active color 80 place line ax=0,0,0 ax=6.5,-20,90 place line ax=0,0,0 ax=6.5,-20,10 place line ax=0,0,0 ax=6.5,-15,65 place line ax=0,0,0 ax=6.5,-30,65 17) Sample Arborograph Script 17) Sample Arborograph Script 17) Sample Arborograph Script 17) Sample Arborograph Script 17) Sample Arborograph Script 17) Sample Arborograph Script 17) Sample Arborograph Script 17) Sample Arborograph Script 17) Sample Arborograph Script 17) Sample Arborograph Script 17) Sample Arborograph Script 17) Sample Arborograph Script 17) Sample Arborograph Script 17) Sample Arborograph Script 17) Sample Arborograph Script 17) Sample Arborograph Script 17) Sample Arborograph Script 17) Sample Arborograph Script 17) Sample Arborograph Script 17) Sample Arborograph Script 17) Sample Arborograph Script 17) Sample Arborograph Script 17) Sample Arborograph Script 17) Sample Arborograph Script 17) Sample Arborograph Script 17) Sample Arborograph Script 17) Sample Arborograph Script 17) Sample Arborograph Script 17) Sample Arborograph Script 17) Sample Arborograph Script 17) Sample Arborograph Script

ax=8,-170,55 move acs move acs place line ax=1.4,-20,90 ax=0,0,0 place line xy=0,0,40.1 move acs ax=0.1389415,-245,90 ax=.9,-16,80 ax=0,0,0 ax=1.15,-240,85 ax=0,0,0 place line move acs xy=0,0,52.2 ax=7.5,-90,95 place line ax=3.25,-320,90 ax=0,0,0 ax=0.2197915,-330,90 move acs active color 112 active color 112 ax=0,0,0 place line ax=0.4,-160,10 ax=0.121963,-100,90 place line place line place line ax=1.4,-20,40 ax=0,0,0 place line move acs active color 80 ax=0,0,0 ax=0,0,0 ax=0,0,0 ax=1.15,-240,85 ax=0,0,0 xy=0,0,40.8 place line active color 80 ax=4,-235,90 ax=7,-40,85 ax=7.5,-115,95 place line move acs ax=3.25,-320,40 move acs ax=0,0,0 place line move acs ax=0,0,0 xy=0,0,43.7 ax=0.214132,-90,90 ax=8,-330,85 ax=0,0,0 place line place line xy=0,0,46.4 ax=1.4,-20,65 move acs place line ax=7,-50,80 ax=0,0,0 ax=0,0,0 move acs ax=0.1906855,-260,90 ax=0,0,0 move acs place line ax=4,-235,50 ax=7,-40,-25 ax=0.168856,-190,90 place line ax=3.25,-320,65 ax=0.7,-90,85 ax=0,0,0 place line ax=0,0,0 move acs move acs ax=8,-330,5 ax=0,0,0 place line place line move acs ax=1.4,-20,65 ax=2.7,-260,40 place line ax=1.1,-10,75 ax=7,-50,-15 ax=0,0,0 ax=0,0,0 ax=1.05,-170,70 move acs ax=0,0,0 active color 80 place line ax=4,-235,70 ax=7,-25,50 xy=0,0,43.9 active color 80 ax=3.25,-320,65 place line ax=0,0,0 place line active color 112 move acs place line move acs ax=0,0,0 ax=8,-320,60 ax=0,0,0 place line place line place line ax=0.1890685,-10,90 ax=0,0,0 xy=0,0,42.1 ax=1.2,-260,70 ax=7,-45,40 ax=0,0,0 ax=0,0,0 ax=0,0,0 ax=3.5,-20,75 move acs place line ax=4,-235,70 ax=7,-50,50 ax=8.25,-140,85 move acs ax=0.2036215,-250,90 place line ax=0,0,0 place line move acs move acs ax=0.6,-10,30 place line ax=0,0,0 ax=8,-345,60 ax=0,0,0 xy=0,0,50.1 xy=0,0,47.2 place line move acs ax=0,0,0 active color 112 ax=1.2,-260,-100 move acs ax=7,-55,40 move acs move acs ax=0,0,0 ax=1.1,-70,70 ax=3.5,-20,-150 place line xy=0,0,37.9 move acs ax=0.1389415,-245,90 ax=0.162388,-16,90 ax=8.25,-140,-15 active color 80 ax=0,0,0 place line move acs xy=0,0,52.2 place line place line ax=3.5,-180,65 ax=0,0,0 ax=0.2375785,-320,90 move acs active color 112 move acs place line ax=0,0,0 ax=0,0,0 ax=1.2,-245,5 ax=0.121963,-100,90 place line ax=.9,-16,80 ax=0,0,0 ax=5.5,-15,80 ax=3.5,70,20 place line active color 80 ax=0,0,0 ax=8.25,-110,60 ax=0,0,0 place line place line active color 112 ax=8,-230,90 active color 112 place line place line ax=3.5,-180,-5 ax=0,0,0 ax=0,0,0 place line place line place line ax=0,0,0 ax=0,0,0 ax=1.2,-270,5 ax=5.75,-300,25 ax=0,0,0 place line ax=0,0,0 ax=0,0,0 ax=5.5,-15,35 ax=3.5,-75,20 place line move acs ax=0.85,-15,90 ax=0,0,0 ax=7.5,-55,90 ax=8.25,-180,60 move acs ax=0,0,0 xy=0,0,40.8 place line ax=8,-230,40 move acs place line xy=0,0,43.7 ax=3.5,-160,50 move acs ax=0,0,0 place line place line xy=0,0,46.4 ax=0,0,0 move acs ax=0.214132,-90,90 ax=5.75,-300,20 ax=0,0,0 place line ax=0,0,0 move acs ax=5.5,-10,65 ax=0.1906855,-260,90 place line ax=0.85,-15,0 ax=0,0,0 ax=7.5,-55,25 ax=0.168856,-190,90 ax=0,0,0 move acs place line ax=8,-225,70 place line move acs ax=3.5,-190,50 ax=0.7,-90,85 ax=0,0,0 place line place line active color 112 ax=0,0,0 ax=3.2,-260,40 move acs move acs ax=5.75,-275,65 ax=0,0,0 place line ax=0,0,0 place line ax=5.5,-20,65 xy=0,0,41.3 ax=1.4,-10,75 ax=0.85,-15,45 ax=0,0,0 ax=7.5,-55,75 ax=0,0,0 move acs active color 80 move acs active color 112 place line ax=8,-235,70 ax=1.05,-170,90 xy=0,0,43.9 place line ax=0.2100895,-260,90 place line ax=0,0,0 place line move acs place line move acs ax=0,0,0 ax=0,0,0 ax=5.75,-330,65 ax=0,0,0 xy=0,0,50.1 ax=0,0,0 place line ax=0.1890685,-10,90 ax=6.5,-215,60 active color 80 ax=2.5,-340,85 move acs ax=0.85,-15,45 move acs ax=7.5,-65,75 ax=0,0,0 place line xy=0,0,37.9 move acs ax=0.1389415,-245,90 move acs ax=1.05,-170,50 move acs place line ax=0,0,0 place line move acs xy=0,0,52.2 xy=0,0,47.2 ax=0.6,-10,30 ax=0,0,0 ax=3.5,-230,70 ax=0,0,0 ax=0.2375785,-320,90 move acs active color 112 move acs place line move acs ax=6.5,-215,-55 ax=2.5,-340,-65 ax=0.121963,-100,90 place line ax=0.162388,-16,90 ax=0,0,0 ax=1.1,-70,70 place line active color 80 ax=0,0,0 ax=1.05,-170,70 active color 80 place line ax=0,0,0 place line place line active color 80 ax=6.5,-30,90 active color 112 place line ax=0,0,0 ax=3.5,-230,-60 ax=0,0,0 ax=0,0,0 place line place line place line ax=0,0,0 ax=6.5,-195,25 ax=2.5,-338,15 ax=4.75,-340,70 ax=0,0,0 place line ax=0,0,0 ax=0,0,0 ax=7,-90,55 place line ax=7.5,-350,80 ax=0,0,0 ax=.9,-16,80 ax=1.05,-170,70 place line ax=0,0,0 place line place line ax=6.5,-30,25 move acs place line ax=0,0,0 ax=3.5,-220,25 ax=0,0,0 ax=0,0,0 place line place line xy=0,0,46.1 ax=0,0,0 ax=6.5,-310,25 ax=2.5,-342,15 ax=4.75,-340,-40 ax=0,0,0 place line ax=0,0,0 move acs ax=7,-90,-15 move acs place line move acs ax=7.5,-350,10 ax=0,0,0 ax=.9,-16,80 ax=0.1712815,-160,90 xy=0,0,43.7 ax=0,0,0 xy=0,0,40.8 place line ax=6.5,-15,65 place line move acs ax=3.5,-245,25 move acs ax=0,0,0 place line place line active color 80 ax=0,0,0 ax=0.1906855,-260,90 move acs ax=0.214132,-90,90 ax=4.75,-330,45 ax=0,0,0 place line ax=0,0,0 place line ax=7,-70,50 xy=0,0,41.3 ax=7.5,-345,50 ax=0,0,0 ax=.9,-16,80 ax=0,0,0 move acs move acs move acs place line ax=6.5,-40,65 ax=9,-160,90 place line ax=3.2,-260,40 ax=0.2100895,-260,90 ax=0.7,-90,85 ax=0,0,0 place line move acs place line ax=0,0,0 move acs ax=4.75,-350,45 ax=0,0,0 xy=0,0,49.1 ax=0,0,0 place line ax=7,-105,50 active color 80 active color 112 ax=1.4,-10,75 move acs ax=7.5,-350,50 move acs ax=.9,-16,80 ax=0,0,0 move acs place line place line active color 80 xy=0,0,37.5 move acs ax=0.1470265,-340,90 move acs ax=9,-160,20 xy=0,0,43.9 ax=0,0,0 ax=0,0,0 place line move acs xy=0,0,51.7 xy=0,0,46.7 move acs ax=9.5,-322,35 ax=2.2,-300,85 ax=0,0,0 ax=0.2408125,0,90 move acs active color 112 move acs place line ax=0.1890685,-10,90 ax=5,-100,70 ax=0.1260055,-10,90 place line ax=0.1664305,-350,90 ax=0,0,0 place line place line active color 112 ax=0,0,0 ax=9,-150,80 move acs ax=0,0,0 ax=0,0,0 place line place line active color 80 ax=3.5,-340,85 active color 112 ax=0.6,-10,30 ax=9.5,-322,5 ax=2.2,-300,55 ax=0,0,0 ax=0,0,0 place line place line place line ax=5,-100,-65 ax=3.2,0,60 ax=0,0,0 place line ax=0,0,0 ax=0,0,0 active color 80 place line place line ax=5.5,-10,90 ax=0,0,0 ax=0.65,-350,90 ax=9,-225,80 place line ax=0,0,0 ax=0,0,0 place line place line ax=3.5,-340,35 move acs ax=0,0,0 ax=9.5,-298,60 ax=2.2,-295,75 ax=0,0,0 ax=0,0,0 place line place line xy=0,0,44.8 ax=1.1,-70,90 ax=5,-60,30 ax=3.2,0,35 ax=0,0,0 place line ax=0,0,0 move acs place line place line ax=5.5,-10,60 ax=0,0,0 ax=0.65,-350,50 ax=0.181792,-320,90 place line ax=0,0,0 ax=0,0,0 place line place line ax=3.5,-335,70 ax=0,0,0 ax=9.5,-370,60 ax=2.2,-305,75 ax=0,0,0 ax=0,0,0 place line place line active color 80 ax=1.1,-70,50 move acs move acs ax=5,-120,30 ax=3.2,0,75 ax=0,0,0 place line ax=0,0,0 place line xy=0,0,43.7 xy=0,0,40.8 move acs ax=5.5,-5,85 ax=0,0,0 ax=0.65,-350,70 ax=0,0,0 place line move acs move acs xy=0,0,40.8 place line ax=3.5,-350,70 ax=2.2,-320,70 ax=0,0,0 ax=0.1906855,-260,90 ax=0.214132,-90,90 move acs ax=0,0,0 place line move acs place line ax=1.1,-70,70 ax=0.214132,-90,90 ax=3.2,-20,75 ax=0,0,0 xy=0,0,48.9 ax=0,0,0 place line move acs move acs move acs ax=5.5,-15,85 move acs ax=0.65,-350,70 ax=0,0,0 place line ax=3.2,-260,40 ax=0.7,-90,85 move acs xy=0,0,36.6 move acs ax=0.1486435,-150,90 move acs ax=2.2,-320,-60 ax=0,0,0 move acs ax=0.7,-90,85 move acs xy=0,0,51.5 xy=0,0,46.4 ax=1.1,-70,70 active color 80 ax=.35,-10,75 move acs ax=0.248089,-90,90 move acs active color 112 move acs place line move acs place line active color 80 ax=1.4,-10,75 ax=0.1276225,-5,90 place line ax=0.168856,-190,90 ax=0,0,0 xy=0,0,43.9 ax=0,0,0 place line active color 112 move acs ax=0,0,0 ax=2.2,-310,30 move acs ax=10,-340,75 ax=0,0,0 place line ax=0.7,-90,85 active color 80 ax=4.25,-150,70 move acs ax=0.1890685,-10,90 ax=2.5,0,60 ax=0,0,0 place line ax=.5,-170,70 place line place line ax=3.5,-30,90 active color 112 ax=0,0,0 place line ax=0,0,0 active color 80 ax=0,0,0 place line place line ax=3.5,-5,80 ax=0,0,0 active color 112 ax=2.2,-330,30 place line ax=10,-340,10 ax=0,0,0 place line ax=0,0,0 ax=4.25,-150,-45 place line move acs ax=0,0,0 ax=2.5,0,-55 ax=0,0,0 ax=0.9,-90,90 place line ax=0,0,0 xy=0,0,44.4 ax=0.6,-10,90 place line ax=3.5,-30,30 ax=0,0,0 place line ax=1.5,-180,90 move acs ax=0,0,0 place line place line ax=3.5,-5,-30 ax=0,0,0 ax=0.185026,-260,90 place line ax=10,-320,75 ax=0,0,0 place line ax=0,0,0 ax=4.25,-125,35 place line ax=0,0,0 ax=2.5,40,20 ax=0,0,0 ax=0.9,-90,-80 place line ax=0,0,0 active color 80 ax=0.6,-10,-30 place line ax=3.5,-30,60 ax=5.5,-10,40 ax=0,0,0 place line ax=1.5,-180,60 place line ax=0,0,0 place line place line ax=3.5,0,40 ax=0,0,0 ax=0,0,0 place line ax=10,-382,75 ax=0,0,0 place line ax=0,0,0 place line ax=4.25,-170,35 place line ax=3.5,-220,85 ax=0,0,0 move acs ax=2.5,-10,20 ax=0,0,0 ax=0.9,-90,5 ax=0,0,0 place line move acs ax=0,0,0 ax=0.6,-10,30 xy=0,0,43.7 move acs ax=3.5,-30,60 ax=5.5,-5,70 ax=0,0,0 xy=0,0,48.9 ax=1.5,-180,75 place line move acs xy=0,0,40.8 move acs place line ax=3.5,-10,40 move acs ax=0,0,0 place line ax=0.1906855,-260,90 move acs xy=0,0,40.8 ax=0,0,0 place line move acs ax=0.1486435,-320,90 place line ax=3.5,-220,40 ax=0,0,0 ax=0.214132,-90,90 move acs ax=0.9,-90,5 ax=0,0,0 xy=0,0,50.8 ax=0,0,0 ax=0.6,-10,30 active color 80 ax=0.214132,-90,90 move acs ax=5.5,-20,70 move acs active color 112 ax=1.5,-180,75 place line move acs place line move acs xy=0,0,36.6 move acs ax=0.133282,-100,90 place line move acs ax=0,0,0 xy=0,0,43.7 ax=0,0,0 ax=0.7,-90,85 move acs move acs xy=0,0,52.9 ax=0,0,0 xy=0,0,46.4 ax=3.5,-215,70 move acs ax=3.2,-260,90 move acs ax=0.7,-90,85 ax=0.248089,-90,90 move acs active color 80 ax=2,-320,90 move acs ax=0.1906855,-260,90 ax=.8,-10,75 ax=0.1163035,-160,90 place line ax=0.168856,-190,90 place line place line active color 112 active color 80 active color 112 ax=0,0,0 place line ax=0,0,0 move acs ax=0,0,0 place line place line place line active color 112 ax=5,-100,75 ax=0,0,0 move acs ax=3.5,-225,70 ax=.85,-260,40 ax=3.2,-260,-10 ax=0,0,0 ax=0,0,0 ax=0,0,0 place line ax=2,-320,-15 ax=1.05,-170,70 move acs ax=0.3,-340,90 ax=1.4,-10,90 ax=0.7,-90,90 ax=0,0,0 place line xy=0,0,43.9 active color 112 place line ax=0.45,-160,90 ax=0,0,0 place line active color 112 move acs place line ax=0,0,0 place line place line place line ax=5,-100,25 ax=0,0,0 place line ax=0.1890685,-10,90 ax=0,0,0 ax=3.2,-260,40 ax=0,0,0 ax=0,0,0 ax=0,0,0 place line ax=2,-320,60 ax=0,0,0 ax=4,-10,90 ax= ax=1.4,-10,60 ax=0.7,-90,80 ax=0,0,0 place line ax=3,-170,90 move acs place line ax=0.3,-340,-80 ax=0.45,-160,30 ax=0,0,0 place line ax=0.6,-10,30 place line ax=0,0,0 place line place line ax=5,-100,60 ax=0,0,0 place line move acs ax=0,0,0 ax=3.2,-260,40 place line ax=0,0,0 ax=0,0,0 place line ax=2,-360,60 ax=0,0,0 active color 80 ax=4,-10,55 move acs ax=0,0,0 ax=1.4,-10,75 ax=0.7,-90,85 ax=0,0,0 place line move acs ax=3,-170,0 place line xy=0,0,43.6 ax=0.3,-340,5 ax=0.45,-160,60 ax=0,0,0 xy=0,0,48.5 ax=0,0,0 place line move acs place line place line ax=5,-110,60 move acs place line ax=3.5,-310,75 ax=0,0,0 ax=0.191494,-120,90 place line ax=0,0,0 ax=0,0,0 place line move acs ax=0.1518775,-350,90 ax=0,0,0 ax=4,-5,80 ax=0,0,0 ax=1.4,-10,75 ax=0.7,-90,85 ax=0,0,0 xy=0,0,50.2 ax=3,-170,45 place line active color 80 ax=0.3,-340,5 move acs move acs ax=0.45,-160,60 move acs active color 112 ax=0,0,0 place line place line move acs xy=0,0,40.8 xy=0,0,35.5 move acs ax=0.138133,-50,90 place line place line ax=3.5,-310,-20 ax=0,0,0 ax=0,0,0 xy=0,0,40.8 move acs move acs xy=0,0,52.9 ax=0,0,0 ax=0,0,0 ax=4,-20,80 ax=4.25,-140,85 move acs ax=0.214132,-90,90 ax=0.2569825,-320,90 move acs active color 80 ax=5.5,-350,85 ax=3,-170,45 place line move acs ax=0.214132,-90,90 ax=0.1163035,-140,90 place line move acs ax=0,0,0 xy=0,0,43.7 place line active color 80 active color 112 ax=0,0,0 place line xy=0,0,46.4 ax=3.5,-305,50 move acs ax=0,0,0 move acs place line place line active color 80 ax=3.5,-50,80 ax=0,0,0 move acs ax=0.1906855,-260,90 ax=4.25,-140,0 ax=0.7,-90,85 ax=0,0,0 ax=0,0,0 place line ax=5.5,-350,20 ax=0.168856,-190,90 place line move acs ax=0.7,-90,120 ax=0.9,-30,90 ax=0,0,0 place line ax=0,0,0 move acs place line ax=.9,-10,75 ax=8,-140,85 ax=0,0,0 place line move acs ax=3.5,-330,50 ax=2.1,-260,40 ax=0,0,0 active color 112 place line place line ax=3.5,-50,55 ax=0,0,0 ax=1.05,-170,70 move acs ax=4.25,-130,55 place line ax=0,0,0 ax=0,0,0 place line ax=5.5,-346,65 xy=0,0,43.9 active color 112 ax=0,0,0 ax=0.7,-90,50 ax=0.9,-30,80 ax=0,0,0 place line active color 112 move acs place line place line ax=0.4,-160,90 ax=8,-140,5 ax=0,0,0 place line place line ax=0.1890685,-10,90 ax=0,0,0 ax=0,0,0 place line place line ax=3.5,-45,75 ax=0,0,0 ax=0,0,0 ax=1.15,-240,85 ax=4.25,-155,55 place line ax=0,0,0 ax=0,0,0 place line ax=5.5,-365,65 ax=7.5,-100,90 move acs move acs ax=0,0,0 ax=0.7,-65,85 ax=0.9,-30,85 ax=0,0,0 place line move acs ax=0.6,-10,30 place line xy=0,0,42.5 ax=0.4,-160,-70 ax=8,-115,55 ax=0,0,0 xy=0,0,47.2 place line move acs ax=0,0,0 move acs place line place line ax=3.5,-60,75 move acs ax=0,0,0 active color 112 ax=1.15,-240,85 ax=0.2003875,-320,90 place line ax=0,0,0 ax=0,0,0 place line move acs ax=0.162388,-16,90 ax=7.5,-100,75 place line ax=0,0,0 ax=0.7,-65,85 ax=0.9,-30,85 ax=0,0,0 xy=0,0,50.1 ax=0,0,0 place line active color 112 ax=0.4,-160,10 move acs move acs 17) Sample Arborograph Script xy=0,0,35.5 ax=0,0,0 ax=0,0,0 ax=0,0,0 active color 112 move acs place line ax=0,0,0 move acs move acs ax=8.5,-60,90 ax=6.25,-320,90 place line ax=2.25,-110,90 place line ax=4.5,-200,10 ax=0,0,0 place line ax=1.45,-70,70 xy=0,0,29.0 ax=0.2569825,-320,90 ax=0,0,0 ax=0,0,0 active color 80 ax=1.45,-180,90 ax=0,0,0 move acs place line place line ax=9,-160,-20 place line ax=0.75,-160,90 place line ax=0.56,-330,70 place line ax=0.309535,-320,90 active color 112 ax=0,0,0 ax=0,0,0 ax=0,0,0 ax=0,0,0 place line ax=0,0,0 place line ax=8.5,-60,25 ax=6.25,-320,20 place line ax=2.25,-110,50 place line ax=7.5,-130,90 ax=0,0,0 place line ax=1.45,-70,70 active color 112 ax=0,0,0 ax=0,0,0 ax=0,0,0 ax=1.45,-180,-10 ax=0,0,0 move acs place line ax=1.3,-340,90 place line place line ax=9,-160,50 place line ax=0.75,-160,-60 place line ax=0.56,-330,80 xy=0,0,29.0 ax=0,0,0 ax=0,0,0 ax=0,0,0 ax=0,0,0 ax=0,0,0 place line move acs ax=1.05,-330,90 place line ax=8.5,-30,70 ax=6.25,-295,65 place line ax=2.25,-110,75 place line ax=7.5,-130,-15 ax=0,0,0 place line ax=0.309535,-320,90 ax=0,0,0 ax=0,0,0 ax=0,0,0 ax=1.45,-180,40 ax=0,0,0 place line ax=1.3,-340,40 place line place line ax=9,-180,50 place line ax=0.75,-160,15 place line ax=0.56,-330,80 active color 112 ax=0,0,0 ax=0,0,0 ax=0,0,0 move acs ax=0,0,0 ax=0,0,0 place line move acs place line ax=1.05,-330,50 place line ax=8.5,-90,70 ax=6.25,-325,65 xy=0,0,32.2 ax=2.25,-115,75 place line ax=7.5,-125,65 ax=0,0,0 xy=0,0,31.3 ax=0,0,0 ax=0,0,0 move acs move acs move acs move acs ax=0,0,0 ax=1.45,-180,40 move acs ax=0.9,-260,90 place line ax=1.3,-340,65 xy=0,0,33.4 xy=0,0,33.4 ax=0.283663,-180,90 xy=0,0,32.2 ax=0.75,-160,15 place line move acs ax=0.2909395,-330,90 ax=0,0,0 move acs move acs move acs move acs ax=0,0,0 xy=0,0,31.3 place line ax=1.05,-330,70 place line ax=0.273961,-350,90 ax=0.273961,-350,90 move acs ax=0.283663,-180,90 xy=0,0,32.2 ax=7.5,-155,65 move acs active color 112 ax=0,0,0 ax=0,0,0 ax=1.45,-180,40 move acs move acs ax=0.2909395,-330,90 place line ax=0.9,-260,-10 place line ax=1.3,-340,65 move acs active color 80 move acs move acs ax=0.283663,-180,90 xy=0,0,32.2 ax=0,0,0 ax=0,0,0 move acs ax=1.35,-280,80 place line ax=4.5,-200,10 ax=1.45,-180,40 move acs move acs ax=1.2,-290,90 place line ax=1.05,-330,70 xy=0,0,35.5 ax=0,0,0 active color 80 move acs move acs ax=0.283663,-180,90 ax=0.56,-330,80 ax=0,0,0 move acs move acs active color 80 ax=1.35,-280,90 place line ax=.7,-200,10 ax=1.45,-180,40 place line ax=0.9,-260,40 xy=0,0,28.0 ax=0.2569825,-320,90 place line ax=0,0,0 active color 112 move acs move acs active color 112 ax=0,0,0 move acs ax=0,0,0 place line ax=7.25,-200,75 place line ax=.4,-200,10 ax=1.45,-180,40 place line ax=1.2,-290,80 place line ax=0.31762,-40,90 active color 112 ax=4.75,-170,65 ax=0,0,0 ax=0,0,0 active color 80 ax=0,0,0 ax=0,0,0 place line ax=1.35,-280,70 place line ax=1.4,-200,90 place line active color 80 ax=0.7,-350,90 place line ax=0.9,-260,40 active color 112 ax=0,0,0 place line ax=0,0,0 ax=0,0,0 place line ax=0,0,0 move acs place line ax=1.6,-320,85 ax=0,0,0 place line ax=7.25,-200,-90 place line ax=2.5,-180,90 ax=0,0,0 place line ax=1.2,-290,85 xy=0,0,29.0 ax=0,0,0 ax=4.75,-170,-60 ax=0,0,0 ax=0,0,0 ax=4.5,-200,90 ax=0,0,0 move acs ax=4.5,-100,90 place line ax=1.35,-280,80 place line ax=1.4,-200,-50 place line ax=0.7,-350,-40 place line ax=0.309535,-320,90 ax=0,0,0 place line ax=0,0,0 ax=0,0,0 place line ax=0,0,0 place line ax=1.6,-320,50 ax=0,0,0 place line ax=7.25,-140,20 place line ax=2.5,-180,-5 ax=0,0,0 place line ax=1.2,-290,85 active color 80 ax=0,0,0 ax=4.75,-140,30 ax=0,0,0 ax=0,0,0 ax=4.5,-200,-70 ax=0,0,0 move acs place line ax=4.5,-100,10 place line ax=1.35,-280,80 place line ax=1.4,-200,20 place line ax=0.7,-350,25 xy=0,0,31.0 ax=0,0,0 ax=0,0,0 place line move acs ax=0,0,0 ax=0,0,0 place line move acs ax=2.5,-280,90 place line ax=1.6,-305,75 ax=0,0,0 xy=0,0,32.2 ax=7.25,-220,20 place line ax=2.5,-175,45 ax=0,0,0 place line ax=0.293365,-70,90 ax=0,0,0 ax=4.75,-200,30 move acs move acs ax=0,0,0 ax=4.5,-200,10 ax=0,0,0 place line ax=4.5,-100,50 place line move acs ax=0.283663,-180,90 xy=0,0,32.2 ax=1.4,-200,20 place line ax=0.7,-350,25 active color 112 ax=0,0,0 ax=0,0,0 xy=0,0,33.4 move acs move acs ax=0,0,0 place line move acs place line ax=2.5,-280,-60 place line ax=1.6,-330,75 move acs move acs ax=0.283663,-180,90 xy=0,0,32.2 ax=2.5,-190,45 ax=0,0,0 xy=0,0,31.3 ax=0,0,0 ax=0,0,0 move acs ax=0.273961,-350,90 ax=1.45,-180,40 move acs move acs ax=4.5,-200,10 move acs ax=1.45,-70,90 place line ax=4.5,-100,50 xy=0,0,34.5 move acs move acs ax=0.283663,-180,90 xy=0,0,32.2 move acs ax=0.2909395,-330,90 ax=0,0,0 move acs move acs ax=4.5,-200,10 ax=1.45,-180,40 move acs xy=0,0,32.2 place line ax=2.5,-240,55 ax=0.2650675,-60,90 ax=1.35,-280,80 active color 80 move acs move acs ax=0.283663,-180,90 move acs active color 112 ax=0,0,0 place line ax=1.2,-200,10 ax=1.45,-180,40 ax=0.283663,-180,90 place line ax=1.45,-70,50 place line active color 80 active color 80 ax=0,0,0 active color 112 move acs move acs ax=0,0,0 ax=0,0,0 place line place line ax=9,-160,75 place line ax=.55,-200,10 ax=1.45,-180,40 active color 80 ax=0.56,-330,90 place line ax=2.5,-300,55

Cross define acs points ax=0.0904315,- place line ax=0.45,-160,60 ax=4.25,-150,35 move acs active color 96 ax=2.2,-320,-60 ax=5.5,-15,65 spherical 170,90 ax=4.25,0,80 active color 178 place line ax=4.25,-170,35 ax=0.162388,-16,90 place line ax=5.5,-15,80 xy=0,0,0 ax=4.25,0,60 place line place line ax=4,-235,70 move acs ax=7.5,-100,95 place line xy=1,0,0 active color 96 ax=0.45,-160,60 ax=5.5,-10,85 ax=4,-235,70 xy=0,0,48.9 active color 96 ax=7.5,-100,90 ax=2.2,-320,30 place line xy=0,1,0 place line place line ax=0.45,-160,60 ax=5.5,-10,90 move acs place line ax=2.2,-310,30 ax=5.5,-15,65 move acs ax=2.1,-170,40 ax=4.25,0,80 move acs place line ax=0.1486435,- ax=.9,-16,80 place line ax=5.5,-15,35 xy=0,0,0 ax=2.1,-170,90 ax=4.25,5,80 xy=0,0,52.9 place line ax=4,-235,70 320,90 ax=.9,-16,80 ax=7.5,-100,95 place line move acs move acs ax=5.5,-10,85 ax=4,-235,70 ax=7.5,-100,75 ax=2.2,-320,30 place line xy=0,0,57.4 place line place line ax=0.1163035,- ax=5.5,-10,60 move acs active color 96 place line ax=2.2,-330,30 ax=5.5,-15,65 move acs ax=2.1,-170,40 ax=4.25,0,80 140,90 xy=0,0,50.1 place line ax=.9,-16,80 place line move acs ax=5.5,-10,65 ax=0.079921,-250,90 ax=2.1,-170,-10 ax=4.25,-5,80 place line move acs ax=2,-320,60 ax=.9,-16,80 ax=7.5,-100,95 xy=0,0,44.4 move acs active color 178 ax=5.5,-10,85 ax=0.1389415,- ax=2,-320,90 ax=7.5,-90,95 move acs place line active color 96 place line xy=0,0,54.1 place line ax=5.5,-5,85 245,90 place line ax=0.185026,-260,90 ax=5.5,-15,65 place line ax=2.1,-170,40 move acs ax=8,-140,55 place line ax=.9,-16,80 place line ax=5.5,-20,65 ax=4.5,-250,40 ax=2.1,-170,40 ax=0.1066015,- ax=8,-140,85 place line active color 96 ax=2,-320,60 ax=.9,-16,80 ax=7.5,-100,95 active color 178 move acs ax=4.5,-250,90 300,90 ax=5.5,-10,85 place line ax=2,-320,-15 ax=7.5,-115,95 place line xy=0,0,43.9 place line place line ax=5.5,-15,85 ax=8,-230,70 place line move acs ax=3.5,-220,70 move acs place line ax=2.1,-170,40 active color 96 ax=8,-140,55 move acs ax=8,-230,90 place line ax=.9,-16,80 xy=0,0,46.4 ax=3.5,-220,85 ax=0.1890685,-10,90 ax=4.5,-250,40 ax=2.1,-170,40 place line ax=8,-140,5 xy=0,0,51.5 ax=2,-320,60 ax=.9,-16,80 move acs ax=4.5,-250,-40 move acs ax=3.25,-170,90 move acs place line ax=2,-320,60 move acs ax=0.168856,-190,90 place line move acs xy=0,0,55.5 ax=3.25,-170,90 place line ax=0.1276225,-5,90 ax=8,-230,70 xy=0,0,46.7 ax=3.5,-220,70 ax=0.6,-10,30 place line move acs ax=8,-140,55 ax=8,-230,40 place line move acs move acs ax=3.5,-220,40 move acs ax=4.5,-250,40 ax=0.0952825,- place line ax=8,-115,55 active color 178 ax=2,-320,60 ax=0.1664305,- ax=1.05,-170,70 ax=1.1,-70,70 ax=4.5,-240,40 310,90 ax=3.25,-170,90 place line place line ax=2,-360,60 350,90 active color 96 place line ax=3.25,-170,90 place line ax=3.5,-5,40 ax=8,-230,70 move acs place line ax=3.5,-220,70 active color 178 place line active color 178 ax=8,-140,55 ax=3.5,-5,80 ax=8,-225,70 xy=0,0,48.5 active color 96 ax=8.25,-140,60 ax=3.5,-215,70 place line ax=4.5,-250,40 place line place line ax=8,-170,55 move acs place line ax=8.25,-140,85 ax=7,-90,50 ax=4.5,-265,40 ax=6.5,-20,65 ax=3.25,-170,90 move acs place line place line ax=0.1518775,- ax=0.65,-350,70 place line ax=7,-90,55 move acs ax=6.5,-20,90 ax=3.25,-170,90 xy=0,0,52.2 ax=3.5,-5,40 ax=8,-230,70 350,90 ax=0.65,-350,90 place line ax=3.5,-220,70 xy=0,0,57.4 move acs ax=3.5,-5,-30 ax=8,-235,70 ax=8.25,-140,60 ax=3.5,-225,70 place line move acs place line place line ax=0.121963,-100,90 move acs active color 96 place line ax=8.25,-140,-15 move acs ax=7,-90,50 ax=0.079921,-15,90 ax=6.5,-20,65 ax=3.25,-170,90 place line xy=0,0,50.1 place line ax=0.65,-350,70 xy=0,0,43.9 ax=7,-90,-15 ax=6.5,-20,10 ax=3.25,-170,90 active color 178 ax=3.5,-5,40 move acs ax=5.5,-350,65 ax=0.65,-350,50 place line move acs active color 178 move acs place line ax=3.5,0,40 ax=0.1389415,- ax=5.5,-350,85 ax=8.25,-140,60 ax=0.1890685,-10,90 place line place line place line xy=0,0,54.1 ax=7,-50,40 245,90 place line ax=8.25,-110,60 ax=7,-90,50 ax=6,-15,80 ax=6.5,-20,65 move acs ax=7,-50,80 place line place line ax=0.65,-350,70 move acs ax=7,-70,50 ax=6,-15,90 ax=6.5,-15,65 ax=0.1066015,- ax=3.5,-5,40 active color 96 ax=5.5,-350,65 ax=0.65,-350,70 place line ax=0.6,-10,30 300,90 place line ax=3.5,-10,40 place line ax=5.5,-350,20 ax=8.25,-140,60 move acs place line place line place line ax=7,-50,40 move acs ax=6.5,-30,65 place line ax=8.25,-180,60 ax= ax=7,-90,50 ax=6,-15,80 ax=6.5,-20,65 active color 178 ax=7,-50,-15 xy=0,0,50.8 ax=6.5,-30,90 place line ax=0.65,-350,70 move acs ax=7,-105,50 ax=6,-15,65 ax=6.5,-30,65 place line move acs ax=5.5,-350,65 ax=0.65,-350,70 xy=0,0,46.4 active color 178 move acs move acs ax=9.5,-220,60 place line ax=0.133282,-100,90 place line ax=5.5,-346,65 move acs move acs place line xy=0,0,43.9 place line xy=0,0,55.5 ax=9.5,-220,90 ax=7,-50,40 ax=6.5,-30,65 xy=0,0,46.4 ax=0.168856,-190,90 ax=3.5,-310,50 move acs ax=6,-15,80 move acs ax=7,-45,40 active color 178 ax=6.5,-30,25 place line move acs ax=3.5,-310,75 ax=0.1890685,-10,90 ax=6,-10,80 ax=0.0952825,- place line place line ax=5.5,-350,65 ax=0.168856,-190,90 active color 96 310,90 ax=9.5,-220,60 place line ax=5,-100,60 place line ax=5.5,-365,65 place line place line move acs place line ax=9.5,-220,5 ax=7,-50,40 ax=5,-100,75 ax=6.5,-30,65 move acs move acs ax=1.05,-170,70 ax=3.5,-310,50 ax=0.6,-10,30 ax=6,-15,80 active color 96 ax=7,-55,40 ax=6.5,-15,65 xy=0,0,47.2 ax=.5,-170,70 ax=1.05,-170,90 ax=3.5,-310,-20 active color 178 ax=6,-20,80 place line place line move acs place line move acs active color 96 place line move acs ax=2.2,-10,55 ax=9.5,-220,60 xy=0,0,52.2 ax=5,-100,60 place line ax=0.162388,-16,90 place line place line place line ax=1.1,-70,70 xy=0,0,57.4 ax=2.2,-10,90 ax=9.5,-200,60 move acs ax=5,-100,25 ax=6.5,-30,65 ax=1.5,-180,75 ax=1.05,-170,70 ax=3.5,-310,50 ax=1.1,-70,90 move acs ax=0.121963,-100,90 ax=6.5,-40,65 move acs ax=1.5,-180,90 ax=1.05,-170,50 ax=3.5,-305,50 ax=0.079921,-135,90 place line place line place line move acs ax=.9,-16,80 place line ax=2.2,-10,55 ax=9.5,-220,60 active color 96 ax=5,-100,60 xy=0,0,49.1 active color 96 place line place line place line ax=1.1,-70,70 active color 178 ax=2.2,-10,20 ax=9.5,-255,60 place line ax=5,-100,60 move acs place line ax=1.5,-180,75 ax=1.05,-170,70 ax=3.5,-310,50 ax=1.1,-70,50 place line move acs ax=0.85,-15,45 ax=0.1470265,- ax=7,-40,50 ax=1.5,-180,60 ax=1.05,-170,70 ax=3.5,-330,50 ax=15,-135,5 place line xy=0,0,53.1 ax=0.85,-15,90 place line 340,90 ax=7,-40,85 move acs place line ax=15,-135,70 ax=2.2,-10,55 move acs ax=5,-100,60 place line place line xy=0,0,43.9 ax=1.1,-70,70 ax=2.2,-10,55 ax=0.1146865,- place line ax=5,-110,60 active color 96 place line ax=1.5,-180,75 ax=1.05,-170,70 move acs ax=1.1,-70,70 place line 300,90 ax=0.85,-15,45 move acs place line ax=7,-40,50 ax=1.5,-180,75 ax=1.05,-170,70 ax=0.1890685,-10,90 ax=15,-135,5 place line ax=0.85,-15,0 xy=0,0,50.2 ax=3.5,-340,70 ax=7,-40,-25 move acs place line ax=15,-135,-145 ax=2.2,-10,55 active color 178 move acs ax=3.5,-340,85 place line xy=0,0,46.1 move acs ax=1.1,-70,70 ax=2.2,-10,55 place line place line ax=0.138133,-50,90 place line ax=1.5,-180,75 move acs ax=0.6,-10,30 ax=1.1,-70,70 place line move acs ax=5.5,-10,70 ax=0.85,-15,45 place line ax=7,-40,50 ax=1.5,-180,75 ax=0.1712815,- move acs move acs ax=15,-135,5 xy=0,0,55.1 ax=5.5,-10,90 ax=0.85,-15,45 active color 178 ax=3.5,-340,70 ax=7,-25,50 move acs 160,90 ax= xy=0,0,43.9 ax=15,-45,5 move acs place line ax=3.5,-340,35 xy=0,0,46.4 move acs ax=0.0985165,-90,90 place line place line ax=3.5,-50,75 place line move acs active color 178 active color 96 ax=0.1890685,-10,90 place line ax=5.5,-10,70 ax=0.85,-15,45 ax=3.5,-50,80 place line ax=7,-40,50 ax=0.168856,-190,90 place line place line ax=15,-135,5 active color 178 ax=5.5,-10,40 ax=0.85,-15,45 ax=3.5,-340,70 ax=7,-50,50 ax=9,-160,80 ax=1.4,-20,65 active color 178 ax=15,-190,5 place line move acs place line ax=3.5,-335,70 move acs move acs ax=9,-160,90 ax=1.4,-20,90 place line move acs ax=7.25,-90,80 place line xy=0,0,52.2 ax=3.5,-50,75 xy=0,0,47.2 ax=1.05,-170,70 ax=0.6,-10,30 xy=0,0,56.4 ax=7.25,-90,80 ax=5.5,-10,70 move acs ax=3.5,-50,55 place line move acs active color 96 place line place line ax=0.6,-10,90 move acs ax=5.5,-5,70 ax=0.121963,-100,90 ax=3.5,-340,70 ax=0.162388,-16,90 place line ax=9,-160,80 ax=1.4,-20,65 ax=0.088006,-60,90 place line place line ax=3.5,-350,70 ax=3,-170,45 ax=9,-160,20 ax=1.4,-20,40 place line ax=7.25,-90,80 place line active color 178 ax=3.5,-50,75 move acs move acs ax=3,-170,90 ax=0.6,-10,30 active color 178 ax=7.25,-90,50 ax=5.5,-10,70 place line ax=3.5,-45,75 xy=0,0,48.9 ax=.9,-16,80 place line place line ax=0.6,-10,-30 place line ax=5.5,-20,70 ax=7.5,-350,50 move acs active color 96 place line ax=9,-160,80 ax=1.4,-20,65 ax=5.5,-60,70 place line move acs ax=7.5,-350,80 place line ax=0.1486435,- place line ax=3,-170,45 ax=9,-150,80 ax=1.4,-20,65 place line ax=5.5,-60,90 ax=7.25,-90,80 xy=0,0,52.9 ax=3.5,-50,75 150,90 ax=7.5,-55,75 ax=3,-170,0 ax=0.6,-10,30 ax=7.25,-20,80 move acs place line ax=3.5,-60,75 ax=7.5,-55,90 place line place line ax=0.6,-10,30 place line ax=0.1163035,- ax=7.5,-350,50 move acs active color 96 place line ax=9,-160,80 ax=1.4,-20,65 ax=5.5,-60,70 place line 160,90 ax=7.5,-350,10 xy=0,0,50.1 place line place line ax=3,-170,45 ax=9,-225,80 ax=1.4,-20,65 place line ax=5.5,-60,20 ax=7.25,-90,80 move acs ax=4.25,-150,35 ax=7.5,-55,75 ax=3,-170,45 move acs move acs ax=0.6,-10,30 ax=7.25,-140,80 active color 96 place line ax=0.1389415,- ax=4.25,-150,70 ax=7.5,-55,25 xy=0,0,44.8 xy=0,0,43.9 ax=0.6,-10,30 place line move acs place line ax=7.5,-350,50 245,90 place line move acs move acs move acs ax=5.5,-60,70 xy=0,0,54.8 ax=0.45,-160,60 ax=7.5,-345,50 place line place line ax=3,-170,45 ax=0.181792,-320,90 ax=0.1890685,-10,90 xy=0,0,43.7 ax=5.5,-55,70 move acs ax=0.45,-160,90 active color 96 ax=4.25,-150,35 ax=7.5,-55,75 ax=3,-170,45 move acs ax=0.100942,-350,90 place line place line ax=4.25,-150,-45 ax=7.5,-55,75 move acs active color 178 move acs ax=0.1906855,- place line place line ax=7.5,-350,50 ax=4,-235,70 xy=0,0,46.4 place line ax=0.6,-10,30 260,90 ax=5.5,-60,70 active color 178 ax=0.45,-160,60 ax=7.5,-350,50 ax=4,-235,90 place line place line move acs ax=2.2,-320,30 move acs ax=5.5,-70,70 place line ax=0.45,-160,30 move acs ax=4.25,-150,35 ax=7.5,-55,75 ax=0.168856,-190,90 ax=2.2,-320,70 ax=1.1,-70,70 move acs move acs ax=4.25,0,80 xy=0,0,51.7 place line ax=4.25,-125,35 ax=7.5,-65,75 ax=.85,-260,40 xy=0,0,56.1 ax=4.25,0,85 place line move acs ax=4,-235,70 move acs move acs place line active color 178 active color 96 move acs ax=0.45,-160,60 ax=0.1260055,-10,90 ax=4,-235,50 place line xy=0,0,47.2 ax=1.05,-170,70 ax=2.2,-320,30 place line place line 17) Sample Arborograph Script ax=4,-10,80 place line place line move acs move acs ax=0.9,-30,85 ax=6.25,-320,65 move acs move acs move acs ax=4,-10,90 ax=9.5,-322,60 ax=3.5,-180,50 move acs ax=1.4,-10,75 xy=0,0,37.9 ax=0.9,-30,85 ax=6.25,-295,65 ax=0.283663,-180,90 ax=1.45,-180,40 xy=0,0,31.0 ax=9.5,-370,60 ax=3.5,-160,50 ax=0.7,-90,85 move acs move acs active color 178 move acs place line move acs move acs active color 96 ax=0.2375785,- xy=0,0,35.5 place line move acs place line ax=0.293365,-70,90 ax=4,-10,80 xy=0,0,43.7 place line ax=.9,-10,75 place line 320,90 move acs ax=6.25,-320,65 ax=1.45,-180,40 ax=4.5,-200,10 ax=4,-10,55 move acs ax=3.5,-180,50 ax=3.5,-30,60 ax=0.2569825,- ax=6.25,-325,65 move acs ax=4.5,-200,90 active color 96 ax=0.1906855,- ax=3.5,-190,50 active color 96 ax=3.5,-30,90 active color 178 320,90 move acs ax=.7,-200,10 place line place line 260,90 move acs place line place line xy=0,0,33.4 place line ax=1.45,-70,70 ax=4,-10,80 xy=0,0,41.3 ax=0.4,-160,10 place line ax=4.75,-340,45 active color 96 move acs active color 96 ax=4.5,-200,10 ax=1.45,-70,90 ax=4,-5,80 move acs move acs ax=0.4,-160,90 ax=3.5,-30,60 ax=4.75,-340,70 place line ax=0.273961,-350,90 place line ax=4.5,-200,-70 ax=3.2,-260,40 ax=0.2100895,- ax=3.5,-30,30 ax=1.3,-340,65 ax=1.4,-200,20 place line place line active color 178 260,90 place line place line ax=1.3,-340,90 active color 178 ax=1.4,-200,90 place line ax=1.45,-70,70 ax=4,-10,80 place line ax=0.4,-160,10 place line ax=4.75,-340,45 place line ax=4.5,-200,10 ax=1.45,-70,50 ax=4,-20,80 ax=10,-340,75 active color 178 ax=0.4,-160,-70 ax=3.5,-30,60 ax=4.75,-340,-40 place line ax=1.35,-280,80 place line ax=4.5,-200,10 move acs ax=10,-340,75 place line ax=3.5,-30,60 ax=1.3,-340,65 ax=1.35,-280,90 ax=1.4,-200,20 place line xy=0,0,43.7 ax=3.5,-230,25 place line place line ax=1.3,-340,40 ax=1.4,-200,-50 place line ax=1.45,-70,70 move acs place line ax=3.5,-230,70 ax=0.4,-160,10 place line ax=4.75,-340,45 place line ax=4.5,-200,10 ax=1.45,-70,70 ax=0.1906855,- ax=10,-340,75 ax=0.4,-160,10 ax=3.5,-30,60 ax=4.75,-330,45 place line ax=1.35,-280,80 place line ax=4.5,-200,10 260,90 ax=10,-340,10 place line ax=3.5,-30,60 ax=1.3,-340,65 ax=1.35,-280,70 ax=1.4,-200,20 move acs place line ax=3.5,-230,25 place line move acs place line ax=1.3,-340,65 ax=1.4,-200,20 xy=0,0,32.2 ax=1.45,-70,70 move acs place line ax=3.5,-230,-60 ax=0.4,-160,10 xy=0,0,40.8 ax=4.75,-340,45 place line move acs ax=1.45,-70,70 ax=2.1,-260,40 ax=10,-340,75 ax=0.4,-160,10 move acs ax=4.75,-350,45 place line ax=1.35,-280,80 place line ax=0.283663,-180,90 move acs active color 96 ax=10,-320,75 place line move acs ax=0.214132,-90,90 move acs ax=1.3,-340,65 ax=1.35,-280,80 ax=1.4,-200,20 xy=0,0,29.0 place line ax=3.5,-230,25 xy=0,0,40.8 xy=0,0,37.5 ax=1.3,-340,65 ax=1.4,-200,20 active color 178 move acs ax=1.15,-240,85 place line ax=3.5,-220,25 move acs move acs move acs move acs place line move acs place line ax=0.309535,-320,90 ax=1.15,-240,85 ax=10,-340,75 ax=0.214132,-90,90 ax=0.7,-90,85 ax=0.2408125,0,90 xy=0,0,35.5 ax=1.35,-280,80 xy=0,0,32.2 ax=1.45,-180,40 ax=10,-382,75 place line active color 178 move acs ax=1.35,-280,80 move acs ax=1.45,-180,90 active color 96 place line move acs ax=3.5,-230,25 move acs place line active color 96 ax=0.2569825,- move acs ax=0.283663,-180,90 place line ax=1.15,-240,85 xy=0,0,43.7 ax=3.5,-245,25 ax=0.7,-90,85 ax=1.4,-10,75 place line 320,90 xy=0,0,32.2 place line ax=0.9,-260,40 ax=1.15,-240,85 move acs move acs move acs ax=1.4,-10,90 ax=3.2,0,75 move acs move acs ax=1.45,-180,40 ax=0.9,-260,90 ax=0.1906855,- xy=0,0,41.3 ax=1.1,-10,75 ax=3.2,0,60 active color 96 ax=0.283663,-180,90 ax=1.45,-180,40 ax=1.45,-180,-10 place line 260,90 move acs place line place line move acs place line ax=1.15,-240,85 ax=0.2100895,- active color 178 ax=1.4,-10,75 place line ax=1.6,-320,75 move acs ax=.55,-200,10 place line ax=0.9,-260,40 ax=1.15,-240,85 active color 178 260,90 place line ax=1.4,-10,60 ax=3.2,0,75 ax=1.6,-320,85 ax=1.45,-180,40 ax=1.45,-180,40 ax=0.9,-260,-10 place line ax=1.2,-260,5 ax=3.2,0,35 move acs active color 96 ax=1.45,-180,40 place line ax=3.2,-260,40 active color 96 ax=1.2,-260,70 place line place line ax=4.5,-200,10 place line place line ax=1.15,-240,85 ax=3.2,-260,90 place line ax=1.4,-10,75 place line ax=1.6,-320,75 ax=0.75,-160,15 place line ax=0.9,-260,40 ax=1.15,-240,85 ax=2.2,-300,75 place line ax=1.4,-10,75 ax=3.2,0,75 ax=1.6,-320,50 active color 178 ax=0.75,-160,90 ax=1.45,-180,40 ax=0.9,-260,40 move acs place line ax=2.2,-300,85 ax=1.2,-260,5 ax=3.2,0,75 place line ax=1.45,-180,40 xy=0,0,43.7 ax=3.2,-260,40 ax=1.2,-260,-100 place line place line ax=9,-160,50 place line move acs place line move acs ax=3.2,-260,-10 place line ax=1.4,-10,75 place line ax=1.6,-320,75 ax=9,-160,75 ax=0.75,-160,15 xy=0,0,31.3 ax=0.9,-260,40 ax=0.1906855,- ax=2.2,-300,75 place line ax=1.4,-10,75 ax=3.2,0,75 ax=1.6,-305,75 ax=0.75,-160,-60 move acs ax=0.9,-260,40 260,90 place line ax=2.2,-300,55 ax=1.2,-260,5 move acs ax=3.2,-20,75 place line ax=0.2909395,- move acs ax=3.2,-260,40 ax=1.2,-245,5 xy=0,0,40.8 move acs place line ax=9,-160,50 place line 330,90 xy=0,0,29.0 move acs ax=3.2,-260,40 place line move acs xy=0,0,36.6 ax=1.6,-320,75 ax=9,-160,-20 ax=0.75,-160,15 move acs ax=2.7,-260,40 ax=2.2,-300,75 place line ax=0.214132,-90,90 move acs ax=1.6,-330,75 ax=0.75,-160,15 move acs ax=0.309535,-320,90 active color 178 place line ax=2.2,-295,75 ax=1.2,-260,5 ax=0.248089,-90,90 move acs place line ax=0.56,-330,80 place line ax=3.2,-260,40 ax=1.2,-270,5 active color 178 xy=0,0,34.5 ax=9,-160,50 place line active color 96 active color 178 ax=3.5,-20,20 ax=3.2,-260,40 place line move acs place line move acs move acs ax=9,-160,50 ax=0.75,-160,15 place line place line ax=3.5,-20,75 move acs ax=2.2,-300,75 xy=0,0,40.8 ax=0.7,-90,85 ax=0.7,-90,85 ax=0.2650675,-60,90 ax=0.75,-160,15 ax=0.7,-350,25 ax=2.5,-280,55 xy=0,0,43.6 ax=2.2,-305,75 move acs ax=0.7,-90,120 active color 96 place line move acs ax=0.7,-350,90 ax=2.5,-280,90 place line move acs move acs ax=0.214132,-90,90 place line active color 178 ax=9,-160,50 xy=0,0,32.2 ax=3.5,-20,20 ax=0.191494,-120,90 xy=0,0,40.8 place line ax=0.9,-90,5 place line ax=9,-180,50 move acs place line place line ax=3.5,-20,-150 move acs move acs ax=0.7,-90,85 ax=0.9,-90,90 ax=8.5,-60,70 move acs ax=0.283663,-180,90 ax=0.7,-350,25 ax=2.5,-280,55 active color 178 ax=0.214132,-90,90 ax=0.7,-90,85 ax=0.7,-90,50 ax=8.5,-60,90 xy=0,0,32.2 ax=0.7,-350,-40 ax=2.5,-280,-60 place line place line move acs place line move acs move acs ax=3.5,-20,20 ax=4.25,-140,55 move acs ax=1.4,-10,75 place line ax=0.9,-90,5 place line ax=0.283663,-180,90 ax=1.45,-180,40 place line place line ax=3.5,70,20 ax=4.25,-140,85 ax=0.7,-90,85 ax=0.7,-90,85 ax=0.9,-90,-80 ax=8.5,-60,70 move acs ax=0.7,-350,25 ax=2.5,-280,55 move acs active color 96 ax=0.7,-65,85 ax=8.5,-60,25 move acs ax=.4,-200,10 ax=0.7,-350,25 ax=2.5,-240,55 place line place line ax=.35,-10,75 place line place line ax=1.45,-180,40 ax=3.5,-20,20 ax=4.25,-140,55 ax=2.5,-340,15 place line ax=0.9,-90,5 place line move acs active color 178 place line place line ax=3.5,-75,20 ax=4.25,-140,0 active color 178 ax=2.5,-340,85 ax=0.7,-90,85 ax=0.9,-90,5 ax=8.5,-60,70 ax=4.5,-200,10 place line ax=0.7,-350,25 ax=2.5,-280,55 move acs place line ax=0.7,-65,85 ax=8.5,-30,70 ax=2.5,-180,45 ax=0.7,-350,25 ax=2.5,-300,55 xy=0,0,43.7 place line ax=2.5,0,20 place line move acs place line active color 178 ax=2.5,-180,90 move acs move acs move acs ax=4.25,-140,55 ax=2.5,0,60 ax=2.5,-340,15 xy=0,0,40.1 ax=0.9,-90,5 place line place line xy=0,0,31.3 xy=0,0,29.0 ax=0.1906855,- ax=4.25,-130,55 ax=2.5,-340,-65 move acs ax=0.9,-90,5 ax=8.5,-60,70 ax=7.25,-200,20 place line move acs move acs 260,90 place line ax=0.2197915,- move acs ax=8.5,-90,70 ax=7.25,-200,75 ax=2.5,-180,45 ax=0.2909395,- ax=0.309535,-320,90 place line ax=2.5,0,20 place line 330,90 xy=0,0,36.6 move acs ax=2.5,-180,-5 330,90 move acs ax=4.25,-140,55 ax=2.5,0,-55 ax=2.5,-340,15 move acs xy=0,0,33.4 place line active color 96 ax=3.2,-260,40 ax=4.25,-155,55 ax=2.5,-338,15 active color 178 ax=0.248089,-90,90 move acs ax=7.25,-200,20 place line active color 96 place line active color 178 move acs place line place line ax=0.273961,-350,90 ax=7.25,-200,-90 ax=2.5,-180,45 place line ax=1.05,-330,70 place line xy=0,0,42.5 ax=2.5,0,20 place line ax=8,-330,60 active color 96 ax=2.5,-175,45 ax=0.56,-330,80 ax=1.05,-330,90 ax=6.5,-215,25 move acs ax=2.5,40,20 ax=2.5,-340,15 ax=8,-330,85 place line move acs place line ax=0.56,-330,90 ax=6.5,-215,60 ax=0.2003875,- ax=2.5,-342,15 ax=0.7,-90,85 ax=1.35,-280,80 ax=7.25,-200,20 place line place line 320,90 place line move acs place line ax=0.7,-90,90 active color 178 ax=7.25,-140,20 ax=2.5,-180,45 place line ax=1.05,-330,70 place line ax=2.5,0,20 xy=0,0,40.8 ax=8,-330,60 place line ax=2.5,-190,45 ax=0.56,-330,80 ax=1.05,-330,50 ax=6.5,-215,25 active color 96 ax=2.5,-10,20 move acs ax=8,-330,5 place line ax=4.75,-170,30 place line move acs ax=0.56,-330,70 ax=6.5,-215,-55 place line move acs ax=0.214132,-90,90 ax=0.7,-90,85 ax=4.75,-170,65 ax=7.25,-200,20 xy=0,0,32.2 place line ax=3.25,-320,65 xy=0,0,40.8 place line ax=0.7,-90,80 ax=7.25,-220,20 move acs place line ax=1.05,-330,70 place line ax=3.25,-320,90 move acs move acs ax=8,-330,60 place line move acs ax=0.283663,-180,90 ax=0.56,-330,80 ax=1.05,-330,70 ax=6.5,-215,25 ax=0.214132,-90,90 ax=0.7,-90,85 ax=8,-320,60 place line ax=4.75,-170,30 xy=0,0,32.2 ax=0.56,-330,80 ax=6.5,-195,25 place line move acs ax=0.7,-90,85 ax=4.75,-170,-60 move acs move acs place line ax=3.25,-320,65 move acs ax=1.4,-10,75 place line ax=0.7,-90,85 ax=0.283663,-180,90 ax=1.45,-180,40 place line ax=1.05,-330,70 place line ax=3.25,-320,40 ax=0.7,-90,85 ax=8,-330,60 place line move acs ax=0.56,-330,80 ax=1.05,-330,70 ax=6.5,-215,25 move acs active color 178 ax=8,-345,60 place line ax=4.75,-170,30 move acs ax=4.5,-200,10 ax=0.56,-330,80 move acs ax=6.5,-310,25 place line ax=.8,-10,75 place line move acs ax=0.7,-90,85 ax=4.75,-140,30 ax=1.45,-180,40 move acs xy=0,0,28.0 move acs ax=3.25,-320,65 ax=5,-100,30 xy=0,0,37.9 ax=0.7,-90,85 move acs active color 178 xy=0,0,31.3 move acs xy=0,0,43.7 ax=3.25,-320,65 active color 96 ax=5,-100,70 move acs move acs place line ax=1.2,-200,10 place line move acs ax=0.31762,-40,90 move acs place line ax=0.2375785,- xy=0,0,35.5 ax=4.75,-170,30 ax=7.5,-130,65 ax=0.2909395,- ax=0.1906855,- place line ax=0.3,-340,5 place line 320,90 move acs ax=4.75,-200,30 active color 96 ax=7.5,-130,90 330,90 active color 96 260,90 ax=3.25,-320,65 ax=0.3,-340,90 ax=5,-100,30 ax=0.2569825,- move acs place line place line ax=3.25,-320,65 ax=5,-100,-65 active color 178 320,90 xy=0,0,33.4 ax=2.25,-110,75 place line active color 96 ax=4.5,-100,50 move acs move acs place line place line move acs ax=2.25,-110,90 ax=7.5,-130,65 place line ax=4.5,-100,90 ax=3.2,-260,40 xy=0,0,42.1 ax=0.3,-340,5 place line ax=5.75,-300,65 active color 96 ax=0.273961,-350,90 ax=7.5,-130,-15 ax=1.2,-290,85 active color 178 move acs ax=0.3,-340,-80 ax=5,-100,30 ax=5.75,-300,25 place line place line ax=1.2,-290,90 place line place line ax=0.2036215,- ax=5,-60,30 ax=0.9,-30,85 move acs ax=2.25,-110,75 place line ax=4.5,-100,50 ax=9.5,-322,60 250,90 place line place line ax=0.9,-30,90 ax=1.35,-280,80 ax=2.25,-110,50 ax=7.5,-130,65 place line ax=4.5,-100,10 ax=9.5,-322,35 ax=0.3,-340,5 place line ax=5.75,-300,65 active color 178 ax=7.5,-125,65 ax=1.2,-290,85 active color 96 ax=0.3,-340,5 ax=5,-100,30 ax=5.75,-300,20 place line place line place line ax=1.2,-290,80 place line place line place line ax=5,-120,30 ax=0.9,-30,85 ax=6.25,-320,65 ax=2.25,-110,75 place line ax=4.5,-100,50 ax=9.5,-322,60 ax=3.5,-180,50 place line move acs place line ax=0.9,-30,80 ax=6.25,-320,90 ax=2.25,-110,75 ax=7.5,-130,65 place line ax=4.5,-100,50 ax=9.5,-322,5 ax=3.5,-180,65 ax=0.3,-340,5 xy=0,0,40.8 ax=5.75,-300,65 ax=7.5,-155,65 ax=1.2,-290,85 ax=0.3,-340,5 move acs ax=5.75,-275,65 place line place line place line move acs ax=1.2,-290,85 place line place line place line move acs ax=0.214132,-90,90 ax=0.9,-30,85 ax=6.25,-320,65 ax=2.25,-110,75 xy=0,0,32.2 ax=4.5,-100,50 ax=9.5,-322,60 ax=3.5,-180,50 xy=0,0,40.8 place line ax=0.9,-30,85 ax=6.25,-320,20 ax=2.25,-115,75 move acs place line ax=4.5,-100,50 ax=9.5,-298,60 ax=3.5,-180,-5 move acs move acs ax=5.75,-300,65 move acs ax=0.283663,-180,90 ax=1.2,-290,85 ax=0.214132,-90,90 ax=0.7,-90,85 ax=5.75,-330,65 place line place line xy=0,0,32.2 ax=1.2,-290,85

Kite define acs points xy=0,0,57.4 ax=2.1,-170,-10 move acs ax=3.25,-170,90 ax=0.1163035,- ax=7,-55,40 Place smartline xy=0,0,50.2 ax=8,-225,70 xy=0,0,48.9 spherical move acs ax=2.1,-170,40 ax=0.0985165,-90,90 ax=3.25,-170,90 160,90 ax=7,-50,80 ax=5.5,-10,90 move acs ax=8,-230,40 move acs xy=0,0,0 ax=0.079921,-135,90 ax=2.1,-170,90 active color 114 active color 96 ax=5.5,-5,85 ax=0.138133,-50,90 ax=8,-235,70 ax=0.1486435,- xy=1,0,0 active color 114 Place smartline move acs Place smartline move acs ax=5.5,-10,60 active color 114 ax=8,-230,90 150,90 xy=0,1,0 Place smartline move acs ax=7.25,-90,80 xy=0,0,54.1 ax=0.45,-160,90 xy=0,0,52.2 ax=5.5,-15,85 Place smartline active color 96 move acs ax=15,-135,70 xy=0,0,55.5 ax=7.25,-20,80 move acs ax=0.45,-160,60 move acs ax=5.5,-10,90 ax=3.5,-50,80 move acs Place smartline xy=0,0,0 ax=15,-45,5 move acs ax=7.25,-90,50 ax=0.1066015,- ax=0.45,-160,30 ax=0.121963,-100,90 ax=3.5,-45,75 xy=0,0,50.1 ax=4.25,-150,70 move acs ax=15,-135,-145 ax=0.0952825,- ax=7.25,-140,80 300,90 ax=0.45,-160,60 active color 96 move acs ax=3.5,-50,55 move acs ax=4.25,-125,35 xy=0,0,57.4 ax=15,-190,5 310,90 ax=7.25,-90,80 active color 114 ax=0.45,-160,90 Place smartline xy=0,0,51.5 ax=3.5,-60,75 ax=0.1389415,- ax=4.25,-150,-45 move acs ax=15,-135,70 active color 114 Place smartline ax=0.85,-15,90 move acs ax=3.5,-50,80 245,90 ax=4.25,-170,35 ax=0.079921,-250,90 Place smartline move acs ax=9.5,-220,90 move acs ax=0.85,-15,45 ax=0.1276225,-5,90 active color 96 ax=4.25,-150,70 active color 96 move acs ax=6.5,-20,90 xy=0,0,54.8 ax=9.5,-200,60 xy=0,0,52.9 ax=0.85,-15,0 active color 114 move acs Place smartline Place smartline xy=0,0,56.4 ax=6.5,-15,65 move acs ax=9.5,-220,5 move acs ax=0.85,-15,45 Place smartline xy=0,0,50.1 ax=6.5,-30,90 move acs ax=4.5,-250,90 move acs ax=6.5,-20,10 ax=0.100942,-350,90 ax=9.5,-255,60 ax=0.1163035,- ax=0.85,-15,90 ax=3.5,-5,80 move acs ax=6.5,-15,65 xy=0,0,48.9 ax=4.5,-240,40 ax=0.088006,-60,90 ax=6.5,-30,65 active color 114 ax=9.5,-220,90 140,90 ax=3.5,0,40 ax=0.1389415,- ax=6.5,-30,25 move acs ax=4.5,-250,-40 active color 114 ax=6.5,-20,90 Place smartline active color 114 move acs ax=3.5,-5,-30 245,90 ax=6.5,-40,65 ax=0.1486435,- ax=4.5,-265,40 Place smartline ax=4.25,0,85 move acs Place smartline xy=0,0,52.2 ax=3.5,-10,40 active color 96 ax=6.5,-30,90 320,90 ax=4.5,-250,90 ax=5.5,-60,90 move acs ax=4.25,5,80 xy=0,0,53.1 ax=8,-140,85 move acs ax=3.5,-5,80 Place smartline active color 96 ax=5.5,-55,70 xy=0,0,55.5 ax=4.25,0,60 move acs ax=8,-115,55 ax=0.121963,-100,90 ax=4,-235,90 move acs Place smartline move acs ax=5.5,-60,20 move acs ax=4.25,-5,80 ax=0.1146865,- ax=8,-140,5 active color 114 move acs ax=4,-235,70 xy=0,0,49.1 ax=2,-320,90 xy=0,0,57.4 ax=5.5,-70,70 ax=0.0952825,- ax=4.25,0,85 300,90 ax=8,-170,55 Place smartline xy=0,0,50.8 ax=4,-235,50 move acs ax=2,-320,60 move acs ax=5.5,-60,90 310,90 active color 114 ax=8,-140,85 ax=7.5,-350,80 move acs ax=4,-235,70 ax=0.1470265,- ax=2,-320,-15 ax=0.079921,-15,90 active color 96 move acs Place smartline ax=7.5,-345,50 ax=0.133282,-100,90 ax=4,-235,90 340,90 ax=2,-360,60 active color 114 move acs Place smartline xy=0,0,54.1 ax=5.5,-10,90 move acs ax=7.5,-350,10 active color 114 active color 96 ax=2,-320,90 Place smartline xy=0,0,56.1 ax=2.2,-10,90 move acs ax=5.5,-5,70 xy=0,0,52.2 ax=7.5,-350,50 Place smartline move acs Place smartline ax=6,-15,90 move acs ax=2.2,-10,55 ax=0.1066015,- ax=5.5,-10,40 move acs ax=7.5,-350,80 ax=5,-100,75 xy=0,0,50.1 ax=3.5,-340,85 move acs ax=6,-10,80 ax=0.0904315,- ax=2.2,-10,20 300,90 ax=5.5,-20,70 ax=0.121963,-100,90 ax=5,-100,60 move acs ax=3.5,-335,70 xy=0,0,48.5 ax=6,-15,65 170,90 ax=2.2,-10,55 active color 96 ax=5.5,-10,90 active color 114 move acs ax=5,-100,25 ax=0.1389415,- ax=3.5,-340,35 move acs ax=6,-20,80 active color 96 ax=2.2,-10,90 Place smartline Place smartline xy=0,0,51.7 ax=5,-110,60 245,90 ax=3.5,-350,70 ax=0.1518775,- ax=6,-15,90 Place smartline ax=3.25,-170,90 move acs ax=7,-50,80 move acs ax=5,-100,75 active color 96 ax=3.5,-340,85 350,90 ax=2.1,-170,90 move acs ax=3.25,-170,90 xy=0,0,52.9 ax=7,-45,40 ax=0.1260055,-10,90 Place smartline active color 96 move acs ax=2.1,-170,40 xy=0,0,55.1 ax=3.25,-170,90 move acs ax=7,-50,-15 active color 114 move acs ax=8,-230,90 move acs Place smartline 17) Sample Arborograph Script ax=5.5,-350,85 xy=0,0,46.4 ax=3.5,-310,-20 ax=4,-5,80 ax=3.2,-260,40 ax=0.214132,-90,90 ax=3.5,-30,60 active color 96 ax=0.273961,-350,90 xy=0,0,32.2 move acs ax=5.5,-346,65 move acs ax=3.5,-330,50 ax=4,-10,55 ax=3.2,-260,-10 move acs ax=3.5,-30,30 Place smartline move acs move acs xy=0,0,31.3 ax=5.5,-350,20 ax=0.168856,-190,90 ax=3.5,-310,75 ax=4,-20,80 ax=3.2,-260,40 ax=0.7,-90,85 ax=3.5,-30,60 ax=0.9,-90,90 ax=1.35,-280,80 ax=0.283663,-180,90 move acs ax=5.5,-365,65 move acs ax=4,-10,90 ax=3.2,-260,90 move acs ax=3.5,-30,90 ax=0.9,-90,5 active color 114 move acs ax=0.2909395,- ax=5.5,-350,85 ax=1.05,-170,70 move acs ax=.8,-10,75 ax=0.9,-90,-80 Place smartline ax=1.45,-180,40 330,90 active color 96 xy=0,0,43.9 move acs move acs active color 96 move acs ax=0.9,-90,5 ax=6.25,-320,90 move acs active color 96 move acs Place smartline move acs xy=0,0,43.7 xy=0,0,43.6 Place smartline xy=0,0,40.8 ax=0.9,-90,90 ax=6.25,-295,65 ax=.55,-200,10 Place smartline xy=0,0,47.2 ax=7.5,-100,90 ax=0.1890685,-10,90 move acs move acs ax=0.3,-340,90 move acs ax=6.25,-320,20 active color 96 ax=0.56,-330,90 move acs ax=7.5,-90,95 move acs ax=0.1906855,- ax=0.191494,-120,90 ax=0.3,-340,5 ax=0.214132,-90,90 move acs ax=6.25,-325,65 Place smartline ax=0.56,-330,80 ax=0.162388,-16,90 ax=7.5,-100,75 ax=0.6,-10,30 260,90 active color 114 ax=0.3,-340,-80 move acs xy=0,0,36.6 ax=6.25,-320,90 ax=0.75,-160,90 ax=0.56,-330,70 move acs ax=7.5,-115,95 move acs move acs Place smartline ax=0.3,-340,5 ax=0.7,-90,85 move acs ax=0.75,-160,15 ax=0.56,-330,80 ax=.9,-16,80 ax=7.5,-100,90 ax= ax=2.1,-260,40 ax=4.25,-140,85 ax=0.3,-340,90 active color 114 ax=0.248089,-90,90 move acs ax=0.75,-160,-60 ax=0.56,-330,90 active color 96 active color 96 active color 96 ax=4.25,-130,55 Place smartline active color 96 xy=0,0,33.4 ax=0.75,-160,15 Place smartline move acs Place smartline Place smartline ax=4.25,-140,0 move acs ax=1.4,-10,90 Place smartline move acs ax=0.75,-160,90 move acs ax=7,-40,85 xy=0,0,46.4 ax=1.4,-20,90 ax=1.15,-240,85 ax=4.25,-155,55 xy=0,0,40.8 ax=1.4,-10,75 ax=0.7,-90,90 ax=0.273961,-350,90 xy=0,0,31.3 ax=7,-25,50 move acs ax=1.4,-20,65 ax=1.15,-240,85 ax=4.25,-140,85 move acs ax=1.4,-10,60 ax=0.7,-90,85 active color 114 move acs move acs ax=7,-40,-25 ax=0.168856,-190,90 ax=1.4,-20,40 ax=1.15,-240,85 ax=0.214132,-90,90 ax=1.4,-10,75 ax=0.7,-90,80 Place smartline xy=0,0,32.2 ax=0.2909395,- ax=7,-50,50 move acs ax=1.4,-20,65 ax=1.15,-240,85 move acs move acs ax=1.4,-10,90 ax=0.7,-90,85 ax=1.35,-280,90 move acs 330,90 ax=7,-40,85 ax=1.05,-170,70 ax=1.4,-20,90 ax=1.15,-240,85 xy=0,0,42.5 ax=0.7,-90,85 ax=0.7,-90,90 ax=1.35,-280,80 ax=0.283663,-180,90 active color 96 active color 96 move acs move acs move acs ax=1.35,-280,70 move acs Place smartline move acs Place smartline move acs move acs ax=0.2003875,- ax=.9,-10,75 xy=0,0,40.8 move acs ax=1.35,-280,80 ax=1.45,-180,40 ax=1.2,-290,90 xy=0,0,47.2 ax=8.25,-140,85 xy=0,0,43.9 xy=0,0,43.7 320,90 active color 96 move acs xy=0,0,35.5 ax=1.35,-280,90 move acs ax=1.2,-290,85 move acs ax=8.25,-110,60 move acs move acs active color 96 Place smartline ax=0.214132,-90,90 move acs ax=.4,-200,10 ax=1.2,-290,80 ax=0.162388,-16,90 ax=8.25,-140,-15 ax=0.1890685,-10,90 ax=0.1906855,- Place smartline ax=0.4,-160,90 active color 114 ax=0.2569825,- move acs active color 114 ax=1.2,-290,85 move acs ax=8.25,-180,60 move acs 260,90 ax=3.25,-320,90 ax=0.4,-160,10 Place smartline 320,90 xy=0,0,32.2 Place smartline ax=1.2,-290,90 ax=.9,-16,80 ax=8.25,-140,85 ax=0.6,-10,30 move acs ax=3.25,-320,65 ax=0.4,-160,-70 ax=0.7,-90,120 active color 96 move acs ax=2.5,-180,90 active color 96 move acs ax=2.7,-260,40 ax=3.25,-320,40 ax=0.4,-160,10 ax=0.7,-65,85 Place smartline ax=0.283663,-180,90 ax=2.5,-175,45 move acs Place smartline move acs ax=1.1,-70,70 active color 114 ax=3.25,-320,65 ax=0.4,-160,90 ax=0.7,-90,50 ax=0.9,-30,90 move acs ax=2.5,-180,-5 xy=0,0,31.0 ax=7.5,-55,90 xy=0,0,46.4 active color 114 Place smartline ax=3.25,-320,90 ax=0.7,-65,85 ax=0.9,-30,85 ax=1.45,-180,40 ax=2.5,-190,45 move acs ax=7.5,-55,75 move acs Place smartline ax=3.5,-20,75 move acs ax=0.7,-90,120 ax=0.9,-30,80 move acs ax=2.5,-180,90 ax=0.293365,-70,90 ax=7.5,-55,25 ax=0.168856,-190,90 ax=5.5,-15,80 ax=3.5,70,20 move acs xy=0,0,40.8 ax=0.9,-30,85 ax=4.5,-200,10 active color 96 ax=7.5,-65,75 active color 96 ax=5.5,-10,65 ax=3.5,-20,-150 xy=0,0,42.1 move acs move acs ax=0.9,-30,90 active color 114 move acs Place smartline ax=7.5,-55,90 Place smartline ax=5.5,-15,35 ax=3.5,-75,20 move acs ax=0.214132,-90,90 xy=0,0,40.1 Place smartline xy=0,0,32.2 ax=1.45,-70,90 ax=1.05,-170,90 ax=5.5,-20,65 ax=3.5,-20,75 ax=0.2036215,- move acs move acs move acs ax=9,-160,75 move acs ax=1.45,-70,70 move acs ax=1.05,-170,70 ax=5.5,-15,80 250,90 ax=0.7,-90,85 ax=0.2197915,- xy=0,0,35.5 ax=9,-160,50 ax=0.283663,-180,90 ax=1.45,-70,50 xy=0,0,47.2 ax=1.05,-170,50 move acs active color 96 move acs 330,90 move acs ax=9,-160,-20 move acs ax=1.45,-70,70 move acs ax=1.05,-170,70 move acs xy=0,0,43.7 Place smartline ax=1.1,-10,75 active color 114 ax=0.2569825,- ax=9,-180,50 ax=1.45,-180,40 ax=1.45,-70,90 ax=0.162388,-16,90 ax=1.05,-170,90 xy=0,0,43.9 move acs ax=3.5,-180,65 active color 114 Place smartline 320,90 ax=9,-160,75 move acs active color 96 move acs ax=0.1906855,- ax=3.5,-160,50 Place smartline ax=8,-330,85 active color 96 ax=4.5,-200,10 move acs Place smartline move acs ax=0.1890685,-10,90 260,90 ax=3.5,-180,-5 ax=1.2,-260,70 ax=8,-320,60 Place smartline move acs active color 114 xy=0,0,29.0 ax=.9,-16,80 xy=0,0,46.1 move acs move acs ax=3.5,-190,50 ax=1.2,-245,5 ax=8,-330,5 ax=1.3,-340,90 xy=0,0,32.2 Place smartline move acs ax=.9,-16,80 move acs ax=0.6,-10,30 ax=3.2,-260,40 ax=3.5,-180,65 ax=1.2,-260,-100 ax=8,-345,60 ax=1.3,-340,65 move acs ax=7.5,-130,90 ax=0.309535,-320,90 ax=.9,-16,80 ax=0.1712815,- move acs active color 114 ax=1.2,-270,5 ax=8,-330,85 ax=1.3,-340,40 ax=0.283663,-180,90 ax=7.5,-125,65 active color 96 ax=.9,-16,80 160,90 ax=1.1,-70,70 Place smartline move acs ax=1.2,-260,70 ax=1.3,-340,65 move acs ax=7.5,-130,-15 Place smartline ax=.9,-16,80 active color 114 active color 114 ax=6.5,-215,60 xy=0,0,41.3 move acs ax=1.3,-340,90 ax=1.45,-180,40 ax=7.5,-155,65 ax=0.9,-260,90 Place smartline Place smartline ax=6.5,-195,25 move acs move acs xy=0,0,37.9 move acs ax=7.5,-130,90 ax=0.9,-260,40 move acs ax=9,-160,90 ax=7,-90,55 ax=6.5,-215,-55 ax=0.2100895,- xy=0,0,40.8 move acs move acs ax=4.5,-200,10 ax=0.9,-260,-10 xy=0,0,46.7 ax=9,-150,80 ax=7,-70,50 ax=6.5,-310,25 260,90 move acs ax=0.2375785,- xy=0,0,35.5 active color 114 move acs ax=0.9,-260,40 move acs ax=9,-160,20 ax=7,-90,-15 ax=6.5,-215,60 active color 114 ax=0.214132,-90,90 320,90 move acs Place smartline xy=0,0,32.2 ax=0.9,-260,90 ax=0.1664305,- ax=9,-225,80 ax=7,-105,50 Place smartline move acs active color 114 ax=0.2569825,- ax=7.25,-200,75 move acs 350,90 ax=9,-160,90 ax=7,-90,55 move acs ax=3.5,-230,70 ax=0.7,-90,85 Place smartline 320,90 ax=7.25,-140,20 ax=0.283663,-180,90 move acs active color 96 xy=0,0,43.7 ax=3.5,-220,25 move acs ax=5.75,-300,25 active color 96 ax=7.25,-200,-90 move acs xy=0,0,29.0 Place smartline move acs move acs move acs ax=3.5,-230,-60 ax=1.4,-10,75 ax=5.75,-275,65 Place smartline ax=7.25,-220,20 ax=1.45,-180,40 move acs ax=0.65,-350,90 xy=0,0,44.8 xy=0,0,43.9 ax=0.1906855,- ax=3.5,-245,25 active color 96 ax=5.75,-300,20 ax=1.6,-320,85 ax=7.25,-200,75 active color 114 ax=0.309535,-320,90 ax=0.65,-350,70 move acs move acs 260,90 ax=3.5,-230,70 Place smartline ax=5.75,-330,65 ax=1.6,-305,75 Place smartline active color 114 ax=0.65,-350,50 ax=0.181792,-320,90 ax=0.1890685,-10,90 move acs ax=2.5,-340,85 ax=5.75,-300,25 ax=1.6,-320,50 move acs ax=4.5,-200,90 Place smartline ax=0.65,-350,70 active color 114 move acs ax=3.2,-260,40 move acs ax=2.5,-338,15 ax=1.6,-330,75 xy=0,0,32.2 ax=4.5,-200,10 ax=2.5,-280,90 ax=0.65,-350,90 Place smartline ax=0.6,-10,30 active color 114 xy=0,0,41.3 ax=2.5,-340,-65 move acs ax=1.6,-320,85 move acs ax=4.5,-200,-70 ax=2.5,-240,55 ax=2.2,-320,70 active color 114 Place smartline move acs ax=2.5,-342,15 xy=0,0,37.9 ax=0.283663,-180,90 ax=4.5,-200,10 ax=2.5,-280,-60 move acs ax=2.2,-310,30 Place smartline ax=9.5,-322,35 ax=0.2100895,- ax=2.5,-340,85 move acs move acs move acs ax=4.5,-200,90 ax=2.5,-300,55 xy=0,0,46.4 ax=2.2,-320,-60 ax=1.1,-70,90 ax=9.5,-298,60 260,90 ax=0.2375785,- xy=0,0,34.5 ax=1.45,-180,40 ax=2.5,-280,90 move acs ax=2.2,-330,30 ax=1.1,-70,70 ax=9.5,-322,5 active color 96 move acs 320,90 move acs move acs move acs ax=0.168856,-190,90 ax=2.2,-320,70 ax=1.1,-70,50 ax=9.5,-370,60 Place smartline xy=0,0,40.8 active color 114 ax=0.2650675,-60,90 ax=1.2,-200,10 xy=0,0,32.2 move acs move acs ax=1.1,-70,70 ax=9.5,-322,35 ax=2.2,-300,85 move acs Place smartline active color 114 active color 96 move acs xy=0,0,29.0 ax=.5,-170,70 move acs ax=1.1,-70,90 ax=2.2,-295,75 ax=0.214132,-90,90 ax=4.75,-340,70 Place smartline Place smartline ax=0.283663,-180,90 move acs active color 96 xy=0,0,44.4 move acs ax=2.2,-300,55 move acs ax=4.75,-330,45 ax=8.5,-60,90 ax=2.25,-110,90 active color 114 ax=0.309535,-320,90 Place smartline move acs move acs xy=0,0,43.7 ax=2.2,-305,75 ax=0.7,-90,85 ax=4.75,-340,-40 ax=8.5,-30,70 ax=2.25,-110,75 Place smartline active color 96 ax=1.5,-180,90 ax=0.185026,-260,90 xy=0,0,43.9 move acs ax=2.2,-300,85 move acs ax=4.75,-350,45 ax=8.5,-60,25 ax=2.25,-110,50 ax=1.45,-180,90 Place smartline ax=1.5,-180,75 active color 114 move acs ax=0.1906855,- ax=1.4,-10,75 ax=4.75,-340,70 ax=8.5,-90,70 ax=2.25,-115,75 ax=1.45,-180,40 ax=1.05,-330,90 ax=1.5,-180,60 Place smartline ax=0.1890685,-10,90 260,90 move acs active color 114 ax=8.5,-60,90 ax=2.25,-110,90 ax=1.45,-180,-10 ax=1.05,-330,70 ax=1.5,-180,75 ax=3.5,-220,85 active color 114 move acs xy=0,0,40.8 Place smartline move acs ax=1.45,-180,40 ax=1.05,-330,50 ax=1.5,-180,90 ax=3.5,-215,70 Place smartline ax=3.2,-260,40 move acs ax=5,-100,70 xy=0,0,37.5 move acs move acs ax=1.45,-180,90 ax=1.05,-330,70 ax=3.5,-220,40 ax=0.6,-10,90 active color 114 ax=0.214132,-90,90 ax=5,-60,30 move acs xy=0,0,33.4 xy=0,0,32.2 ax=1.05,-330,90 move acs ax=3.5,-225,70 ax=0.6,-10,30 Place smartline move acs ax=5,-100,-65 ax=0.2408125,0,90 move acs move acs move acs xy=0,0,46.4 ax=3.5,-220,85 ax=0.6,-10,-30 ax=10,-340,75 ax=0.7,-90,85 ax=5,-120,30 active color 96 ax=0.273961,-350,90 ax=0.283663,-180,90 xy=0,0,31.3 move acs move acs ax=0.6,-10,30 ax=10,-320,75 move acs ax=5,-100,70 Place smartline move acs move acs move acs xy=0,0,28.0 ax=0.168856,-190,90 move acs ax=0.6,-10,90 ax=10,-340,10 ax=.35,-10,75 ax=3.2,0,60 ax=1.35,-280,80 ax=1.45,-180,40 ax=0.2909395,- move acs move acs xy=0,0,43.9 ax=10,-382,75 active color 114 move acs ax=3.2,0,75 active color 114 move acs 330,90 ax=0.31762,-40,90 ax=1.05,-170,70 move acs move acs ax=10,-340,75 Place smartline xy=0,0,40.8 ax=3.2,0,35 Place smartline ax=.7,-200,10 move acs active color 96 active color 96 ax=0.1890685,-10,90 xy=0,0,43.7 ax=2.5,0,60 move acs ax=3.2,-20,75 ax=4.75,-170,65 active color 96 ax=0.56,-330,80 Place smartline Place smartline move acs move acs move acs ax=2.5,40,20 ax=0.214132,-90,90 ax=3.2,0,60 ax=4.75,-140,30 Place smartline active color 96 ax=4.5,-100,90 ax=3,-170,90 ax=0.6,-10,30 ax=0.1906855,- xy=0,0,43.7 ax=2.5,0,-55 move acs ax=4.75,-170,-60 ax=1.4,-200,90 Place smartline ax=4.5,-100,50 ax=3,-170,45 move acs 260,90 move acs ax=2.5,-10,20 ax=0.7,-90,85 move acs ax=4.75,-200,30 ax=1.4,-200,20 ax=0.7,-350,90 ax=4.5,-100,10 ax=3,-170,0 ax= move acs ax=0.1906855,- ax=2.5,0,60 move acs xy=0,0,36.6 ax=4.75,-170,65 ax=1.4,-200,-50 ax=0.7,-350,25 ax=4.5,-100,50 ax=3,-170,45 active color 114 ax=.85,-260,40 260,90 ax=1.4,-10,75 move acs ax=1.4,-200,20 ax=0.7,-350,-40 ax=4.5,-100,90 ax=3,-170,90 Place smartline active color 96 active color 114 move acs active color 96 ax=0.248089,-90,90 move acs ax=1.4,-200,90 ax=0.7,-350,25 ax=3.5,-310,75 Place smartline Place smartline xy=0,0,40.8 Place smartline move acs xy=0,0,33.4 ax=0.7,-350,90 move acs ax=3.5,-305,50 ax=4,-10,90 ax=3.2,-260,90 move acs ax=3.5,-30,90 ax=0.7,-90,85 move acs move acs

Extras: Compass Rose

define acs points .5 xy=0,0,0 .05 ax=17,-270,90 spherical 0 ax=5,-45,90 active color 9 place cone radius .05 xy=0,0,0 ax=10,-0,90 active color 8 place cone radius .5 xy=0,0,0 active color 9 xy=1,0,0 ax=15,-0,90 place cone radius .05 0 ax=10,-270,90 place cone radius xy=0,1,0 active color 14 .05 .05 ax=10,-180,90 .05 move acs place text fvi .05 xy=0,0,0 ax=15,-180,90 place cone radius .05 xy=0,0,0 N xy=0,0,0 ax=5,-135,90 active color 9 .5 xy=0,0,0 active color 8 ax=15,0,90 ax=10,-90,90 place cone radius 0 ax=5,-315,90 place cone radius ax=17,0,90 active color 8 .05 ax=10,-270,90 .05 place cone radius place cone radius .05 ax=15,-270,90 .05 active color 9 .5 .05 xy=0,0,0 active color 14 xy=0,0,0 place cone radius 0 .05 ax=5,-215,90 place text fvi ax=10,-0,90 .05 ax=10,-90,90 xy=0,0,0 active color 8 W place cone radius .05 ax=15,-90,90 ax=10,-180,90 place cone radius ax=15,-270,90

Extras: 75m Vertical Scale Bar define acs points xy=15,0,75 .05 place text fvi place cone radius xy=16,0,30 xy=16,0,36 spherical xy=15,0,10 15 .05 place text fvi place cone radius xy=0,0,0 place cone radius xy=16,0,10 xy=16,0,14 .05 30 .05 xy=1,0,0 .05 place text fvi xy=16,0,16 xy=15,0,25 xy=16,0,29 .05 xy=0,1,0 .05 10 place cone radius xy=16,0,25 xy=16,0,31 xy=15,0,40 move acs xy=15,0,5 xy=16,0,9 .05 place text fvi place cone radius xy=16,0,40 xy=0,0,0 xy=16,0,5 xy=16,0,11 .05 25 .05 place text fvi place text fvi xy=15,0,20 xy=16,0,24 .05 40 active color 72 5m place cone radius xy=16,0,20 xy=16,0,26 xy=15,0,35 xy=16,0,39 place cone radius xy=16,0,4 .05 place text fvi place cone radius xy=16,0,35 xy=16,0,41 .05 xy=16,0,6 .05 20 .05 place text fvi place cone radius .05 place cone radius xy=15,0,15 xy=16,0,19 .05 35 .05 xy=15,0,0 .05 xy=16,0,15 xy=16,0,21 xy=15,0,30 xy=16,0,34 .05 17) Sample Arborograph Script xy=15,0,45 .05 .05 .05 .05 .05 place cone radius xy=16,0,45 xy=15,0,50 xy=15,0,55 xy=15,0,60 .05 .05 .05 place text fvi xy=16,0,50 xy=16,0,55 xy=16,0,60 xy=15,0,65 xy=15,0,70 .05 45 place text fvi place text fvi place text fvi xy=16,0,65 xy=16,0,70 xy=15,0,75 xy=16,0,44 50 55 60 place text fvi place text fvi xy=16,0,75 xy=16,0,46 xy=16,0,49 xy=16,0,54 xy=16,0,59 65 70 place text fvi xy=16,0,51 xy=16,0,56 xy=16,0,61 xy=16,0,64 xy=16,0,69 75 place cone radius place cone radius place cone radius xy=16,0,66 xy=16,0,71 xy=16,0,74 .05 .05 .05 place cone radius place cone radius xy=16,0,76 Lighting settings used for rendering light place xy=150,0,150 xy=0,0,0 light place xy=-150,0,150 xy=0,0,0 light place xy=0,150,150 xy=0,0,0 light place xy=0,-150,150 xy=0,0,0 light place xy=0,0,0 xy=0,0,150 18) Blank Data Sheet - Structure

18) Appendix 4. Blank Data Sheet

19) Sample Data Sheet

19) Appendix 5. Sample Data Sheet

Grand Total CD sticky Funnel Crawl Hangtraps All Traps 100yr Old Total 100yr Old Total 100yr Old Total Grand Total 6490 644 596 1240 1596 1128 2724 1245 1281 2526

Order Family Species binomial Acarina Acari-Grouped Acari-Grouped 100 4 1 5 69 9 78 12 5 17 Araneae Araneae- Grouped Araneae- Grouped 318 2 3 5 94 83 177 59 77 136 Salticidae YDB sp. 1 3 3 3 Blattodea Blatellidae Blatellidae YDB sp.1 18 11 6 17 1 1 Blatellidae YDB sp.2 41 9 26 35 6 6 Blatellidae YDB sp.3 37 26 4 30 2 5 7 Blatellidae YDB sp.4 75 1 1 34 34 68 4 2 6 Blatellidae YDB sp.5 54 23 25 48 3 3 6 Coleoptera Anobiidae Anobiidae TFIC sp.10 5 1 1 2 2 2 2 Anobiidae YDB sp.01 55 5 10 15 2 7 9 7 24 31 Dryophilodes TFIC sp.4 13 1 1 3 2 5 2 5 7 Hadrobregmus areolicollis 8 1 1 2 3 5 1 1 2 Ptinus exulans 1 1 1 Boganiidae Boganium armstrongi 1 1 1 Brentidae Apion tasmanicum 17 1 1 7 9 16 Auletobius TFIC sp.1 1 1 1 Buprestidae Cisseis oblonga 2 1 1 1 1 Carabidae Agonocheila ?fenestrata 2 1 1 1 1 Demetrida YDB sp.1 2 1 1 1 1 Mecyclothorax ambiguus 1 1 1 Cerambycidae Ancita crocogaster 1 1 1 Cerambycidae YDB sp.02 1 1 1 Cerambycidae YDB sp.03 2 2 2 Lygesis ?mendica 1 1 1 Zygocera canosa 2 2 2 Cerylonidae Philothermus tasmanicus 1 1 1 Chrysomelidae Arsipoda erichsoni 7 1 1 2 2 2 3 3 Chrysophtharta bimaculata 22 1 1 9 4 13 4 4 8 Chrysophtharta lignea 1 1 1 Monolepta TFIC sp. 02 3 1 1 1 1 2 Monolepta TFIC sp.02 3 2 1 3 Paropsis rupidipes 1 1 1 Sterromela subcostata 7 7 7 Trachymela YDB sp.01 2 1 1 1 1 Chrysomelidae larvae Chrysomelidae larvae YDB sp. 01 1 1 1 Clambidae Sphaerothorax tasmani 2 2 2 Cleridae Cleridae YDB sp. 01 4 1 1 1 2 3 Lemidia pallida 4 1 1 1 2 3 Neoscrobiger patricius 6 3 3 1 2 3 Parapylus bicinctus 2 2 2 Coccinelidae Cleobora mellyi 3 1 1 2 2 Rodatus TFIC sp. 01 27 25 2 27 Coccinellidae Rhyzobius TFIC sp.05 6 3 1 4 2 2 Rhyzobius TFIC sp.14 6 4 2 6 Rhyzobius TFIC sp.20 2 1 1 1 1 Cryptophagidae Cryptophagus tasmanicus 1 1 1 Curculionidae Aades cultratus 1 1 1 Acacicis abundans 1 1 1 Cryptorhyncinae TFIC sp.11 1 1 1 Curculionidae YDB sp.03 1 1 1 Curculionidae YDB sp.05 1 1 1 Curculionidae YDB sp.06 2 2 2 Curculionidae YDB sp.07 2 2 2 Curculionidae YDB sp.11 1 1 1 Curculionidae YDB sp.12 1 1 1 Elleschus wellingtoniensis 1 1 1 Emplesis TFIC sp.01 1 1 1 Exithius capucinus 1 1 1 Orthorhinus TFIC sp.1 1 1 1 Orthorhinus TFIC sp.2 3 1 2 3 Pentamimus australis 6 2 2 2 2 4 Pentarthrum? TFIC sp.03 1 1 1 Platypus YDB sp. 01 1 1 1 Tychiinae TFIC sp.8 1 1 1 Dermestidae Trogoderma YDB sp.1 3 2 1 3 Trogoderma YDB sp.2 3 1 1 2 1 1 Elateridae Augenotus quadriguttatus 2 2 2 Conoderus australasiae 1 1 1 Denticollinae YDB sp.4 13 2 1 3 4 6 10 Elateridae YDB sp.1 1 1 1 Elateridae YDB sp.2 1 1 1 Elateridae YDB sp.UNID 1 1 1 Erotylidae Erotylidae KH 1 6 1 1 2 1 1 1 2 3 Thallis vinula 5 1 4 5 Laemophloeidae Laemophloeidae TFIC sp.01 1 1 1 Larvae-Coleoptera Coleoptera YDB sp. 01- larvae 1 1 1 Coleoptera YDB sp. 02- larvae 1 1 1 Latridiidae Aridus costatus 4 3 3 1 1 Cortinicara TFIC sp.02 7 2 5 7 Latridiidae YDB sp. 01 2 1 1 1 1 Lymexylidae Atractocerus YDB sp. 01 1 1 1 Atractocerus YDB sp.1 1 1 1 Melandryidae Melandryidae TFIC sp.03 2 1 1 1 1 Melandryidae TFIC sp.04 5 2 2 4 1 1 Orchesia ?eucalyptii 2 1 1 1 1 Orchesia alphabetica 2 1 1 1 1 Melyridae Dasytes TFIC sp.02 2 2 2 Mordellidae Mordella promiscua 2 1 1 2 Mordellidae TFIC sp.05 4 3 1 4 Mycetophagidae Litargus intricatus 2 1 1 1 1 Mycteridae Trichosalpingus TFIC sp.01 12 2 2 4 3 7 2 1 3 Nitidulidae Soronia superba Reitter 1 1 1 Thalycrodes australe 2 2 2 Oedemeridae Dohrnia miranda 18 3 14 17 1 1 Oedomeridae Dorhnia simplex 43 13 23 36 2 5 7 Phalacridae Phalacridae TFIC sp.01 20 1 1 2 2 10 7 17 Prostomidae Dryocora cephalotes 1 1 1 Pselaphidae Pselaphidae rename 5 5 5 Pyrochroidae Binburrum ?ruficollis 1 1 1 Salpingidae Orphanotrophium frigidum 24 13 4 17 2 1 3 3 1 4 Scarabaeidae Heteronyx tasmanicus 2 1 1 1 1 Phyllochlaenia villosus 1 1 1 Grand Total CD sticky Funnel Crawl Hangtraps All Traps 100yr Old Total 100yr Old Total 100yr Old Total Grand Total 6490 644 596 1240 1596 1128 2724 1245 1281 2526

Order Family Species binomial Scirtidae Pryonocyphon TFIC sp.01 10 1 2 3 1 1 2 2 3 5 Scirtidae TFIC sp.04 1 1 1 Coleoptera Scraptiidae Scraptia laticollis 4 1 1 2 1 3 Scydmaenidae Euconnus sp. 1 1 1 1 Silvanidae Cryptamorpha victoriae 1 1 1 ?Silvanidae Silvanidae YDB sp. 01 2 1 1 1 1 Staphylinidae Pselaphinae YDB sp.01 1 1 1 Sepedophilus TFIC sp.01 1 1 1 Staphylinidae YDB sp.03 2 1 1 2 Staphylinidae YDB sp.04 1 1 1 Staphylinidae YDB sp.05 1 1 1 Staphylinidae YDB sp.06 1 1 1 Staphylinidae YDB sp.07 1 1 1 Tenebrionidae Appelatus tasmanicus 3 1 2 3 Throscidae Aulonothroscus elongatus 259 11 20 31 110 56 166 34 28 62 Trogossitidae Egolia variegata 2 2 2 Trogossitidae TFIC sp.01 41 2 12 14 4 4 8 4 15 19 Trogossitidae YDB sp. 01 1 1 1 Zopheridae Penthelispa fuliginosa 1 1 1 Collembola Paronellidae Paronellidae YDB sp.1 15 7 2 9 3 3 6 Dermaptera Spongophoridae Spongophoridae YDB sp.1 3 3 3 Diptera Diptera YDB sp. 01 2 2 2 Diptera YDB sp. 02 1 1 1 Diptera YDB sp. 04 1 1 1 Diptera YDB sp. 05 16 5 7 12 1 1 3 3 Diptera YDB sp. 06 668 103 135 238 120 53 173 128 129 257 Diptera YDB sp. 07 2 1 1 2 Diptera YDB sp. 08 10 7 3 10 Diptera YDB sp. 09 3 2 2 1 1 Diptera YDB sp. 10 4 2 2 4 Diptera YDB sp. 12 2 1 1 1 1 Diptera YDB sp. 13 1 1 1 Diptera YDB sp. 14 1 1 1 Diptera YDB sp. 15 11 11 11 Diptera YDB sp. 16 80 57 23 80 Diptera YDB sp. 17 1 1 1 Asilidae Asilidae YDB sp.1 3 1 1 1 1 1 1 Calliphoridae Calliphora YDB sp.1 12 2 4 6 3 3 6 Chloropidae Chloropidae YDB sp.1 8 1 3 4 2 2 4 Empididiae Empididiae YDB sp.2 6 1 5 6 Ironomyidae Ironomyidae YDB sp.1 1 1 1 Lauxiniidae Lauxiniidae YDB sp.1 7 6 6 1 1 Muscidae Muscidae YDB sp.1 13 4 2 6 5 2 7 Muscidae YDB sp.2 5 3 2 5 Mycetophilidae Mycetophilidae YDB sp.1 2 2 2 Mycetophilidae YDB sp.2 35 3 28 31 1 1 3 3 Mycetophilidae YDB sp.3 1 1 1 Mycetophilidae YDB sp.4 1 1 1 Mycetophilidae YDB sp.5 4 4 4 Mycetophilidae YDB sp.6 3 3 3 Mycetophilidae YDB sp.7 28 16 16 11 1 12 Phoridae Phoridae YDB sp.1 29 15 9 24 4 1 5 Phoridae YDB sp.2 3 3 3 Phoridae YDB sp.3 194 9 23 32 15 7 22 64 76 140 Phoridae YDB sp.4 9 1 1 2 5 2 7 Phoridae YDB sp.5 8 7 7 1 1 Psychodidae Psychodidae YDB sp.1 6 4 2 6 Psychodidae YDB sp.2 1 1 1 ?Scenopinidae Diptera YDB sp. 11 1 1 1 Sciaridae Sciaridae YDB sp.01 1 1 1 Sciaridae YDB sp.02 242 46 21 67 43 49 92 43 40 83 Sciaridae YDB sp.03 1082 20 14 34 229 96 325 381 342 723 Sciaridae YDB sp.04 1 1 1 Sciaridae YDB sp.05 17 12 12 2 3 5 Sciaridae YDB sp.06 1 1 1 Sciaridae YDB sp.07 2 1 1 1 1 Sciaridae YDB sp.08 2 2 2 Sciaridae YDB sp.09 78 60 11 71 1 1 6 6 Sciaridae YDB sp.10 153 72 55 127 8 7 15 7 4 11 Sciaridae YDB sp.11 80 45 2 47 21 7 28 4 1 5 Tabanidae Tabanidae YDB sp.1 4 3 3 1 1 Tachinidae Tachinidae YDB sp.1 16 3 3 4 1 5 3 5 8 Tipulidae Tipulidae YDB sp. 1 94 31 24 55 25 14 39 Tipulidae YDB sp. 2 35 1 1 1 1 2 18 14 32 Tipulidae YDB sp. 3 8 2 6 8 Tipulidae YDB sp. 4 72 5 5 28 11 39 7 21 28 Hemiptera Hemiptera YDB sp. 01 1 1 1 Hemiptera YDB sp. 02 1 1 1 Hemiptera YDB sp. 03 1 1 1 Acanthosomatidae Acanthosomatidae YDB sp.1 2 2 2 Cicadellidae Cicadellidae YDB sp. 01 8 1 1 2 1 3 1 3 4 Cicadellidae YDB sp. 02 21 2 2 15 3 18 1 1 Cicadellidae YDB sp. 03 6 1 2 3 1 2 3 Cicadellidae YDB sp. 04 1 1 1 Cicadellidae YDB sp. 05 1 1 1 Cicadellidae YDB sp. 06 40 1 1 17 19 36 1 2 3 Cicadellidae YDB sp. 07 9 1 8 9 Cicadellidae YDB sp. 08 110 53 51 104 3 3 6 Cicadellidae YDB sp. 09 11 1 1 3 7 10 Cixiidae Cixiidae YDB sp.01 13 1 1 2 3 5 4 3 7 Coccidae Coccidae YDB sp.1 1 1 1 Enicocephalidae Enicocephalidae YDB sp.1 3 1 1 2 2 Eurymelidae Eurymela YDB sp. 01 3 1 1 2 2 Eurymelidae YDB sp.01 5 1 1 2 2 2 1 1 Eurymelidae YDB sp.02 36 3 26 29 5 2 7 Flatidae Flatidae YDB sp.1 58 34 22 56 1 1 2 Flatidae YDB sp.2 1 1 1 Fulgoridae Fulgoridae YDB sp.1 51 13 15 28 11 3 14 7 2 9 Fulgoridae YDB sp.2 13 11 2 13 Fulgoridae YDB sp.3 18 6 6 12 1 5 6 Fulgoridae YDB sp.4 1 1 1 Lygaeidae Lygaeidae YDB sp.1 2 2 2 Grand Total CD sticky Funnel Crawl Hangtraps All Traps 100yr Old Total 100yr Old Total 100yr Old Total Grand Total 6490 644 596 1240 1596 1128 2724 1245 1281 2526

Order Family Species binomial Lygaeidae YDB sp.2 6 1 1 4 4 1 1 Lygaeidae YDB sp.3 1 1 1 Membracidae Membracidae YDB sp.1 1 1 1 Miridae Miridae YDB sp. 01 18 8 10 18 Hemiptera Miridae Miridae YDB sp. 02 5 1 3 4 1 1 Miridae YDB sp. 03 4 1 1 3 3 Miridae YDB sp. 04 1 1 1 Miridae YDB sp. 05 2 2 2 Miridae YDB sp. 06 1 1 1 Sejanus YDB sp. 01 2 1 1 1 1 Pentatomidae Pentatomidae YDB sp.1 1 1 1 Pentatomidae YDB sp.2 16 8 8 16 Pentatomidae YDB sp.3 106 56 48 104 1 1 2 Pentatomidae YDB sp.4 47 1 1 17 25 42 4 4 Psyllidae Psyllidae YDB sp. 2 1 1 1 Psyllidae YDB sp. 3 10 8 2 10 Psyllidae YDB sp. 4 1 1 1 Psyllidae YDB sp. 5 1 1 1 Psyllidae YDB sp. 6 1 1 1 Psyllidae YDB sp. 7 1 1 1 Psyllidae juvenile Psyllidae YDB sp. 1 2 1 1 2 Hymenoptera Hymenoptera YDB sp. 01 1 1 1 Hymenoptera YDB sp. 02 8 1 1 3 3 2 2 4 Hymenoptera YDB sp. 03 1 1 1 Hymenoptera YDB sp. 04 9 2 2 3 3 1 3 4 Hymenoptera YDB sp. 06 17 1 1 7 6 13 3 3 Hymenoptera YDB sp. 10 3 3 3 Hymenoptera YDB sp. 12 1 1 1 Hymenoptera YDB sp. 15 1 1 1 Hymenoptera YDB sp. 24 1 1 1 Braconidae Braconidae YDB sp.1 1 1 1 Braconidae YDB sp.2 1 1 1 Braconidae YDB sp.3 24 16 2 18 4 2 6 Chalcidoid? Chalcidoid? YDB sp.1 60 12 16 28 7 18 25 3 4 7 Evaniidae Evaniidae YDB sp.2 17 2 10 12 1 4 5 Firmicidae Firmicidae YDB sp.1 1 1 1 Formicidae Camponotus YDB sp. 01 7 3 2 5 1 1 2 Camponotus YDB sp. 02 13 2 4 6 7 7 (* Ants- grouped) Formicidae- Grouped 234 21 21 64 111 175 24 14 38 Hymenoptera YDB sp. 22 5 1 3 4 1 1 Gasteruptionidae Gasteruptionidae YDB sp.01 14 2 2 12 12 Ichneumonidae Ichneumonidae YDB sp.1 9 3 1 4 3 2 5 Ichneumonidae YDB sp.2 1 1 1 Ichneumonidae YDB sp.3 3 2 2 1 1 Ichneumonidae YDB sp.4 1 1 1 Ichneumonidae YDB sp.5 6 1 3 4 2 2 Microwasp Hymenoptera YDB sp. 08 3 1 1 2 2 Hymenoptera YDB sp. 09 1 1 1 Hymenoptera YDB sp. 11 2 2 2 Hymenoptera YDB sp. 13 1 1 1 Hymenoptera YDB sp. 14 6 1 5 6 Hymenoptera YDB sp. 21 1 1 1 Hymenoptera YDB sp. 25 3 1 1 2 1 1 Hymenoptera YDB sp. 26 1 1 1 Hymenoptera YDB sp. 27 7 1 3 4 1 1 2 1 1 Hymenoptera YDB sp. 30 10 4 4 1 1 1 4 5 MicroWasp? Hymenoptera YDB sp. 07 2 2 2 Hymenoptera YDB sp. 16 18 5 5 5 8 13 Hymenoptera YDB sp. 17 26 11 7 18 2 2 6 6 Hymenoptera YDB sp. 18 5 2 2 1 1 2 2 Hymenoptera YDB sp. 20 16 4 4 2 8 10 1 1 2 Hymenoptera YDB sp. 23 1 1 1 Hymenoptera YDB sp. 28 1 1 1 Hymenoptera YDB sp. 29 3 2 1 3 Pompilidae Pompilidae YDB sp.1 40 5 5 18 10 28 4 3 7 Proctotrupoidae Proctotrupoidae YDB sp.1 17 1 8 9 2 5 7 1 1 Stinging Wasp? Hymenoptera YDB sp. 05 46 28 11 39 1 3 4 1 2 3 Vespidae Vespula germanica 3 1 1 2 2 Wingless MicroWasp? Hymenoptera YDB sp. 19 3 3 3 Isoptera Isoptera YDB sp.1 29 2 1 3 7 6 13 4 9 13 Isoptera YDB sp.2 19 5 14 19 Larvae UNID Hexapod larvae- Grouped 13 12 1 13 Lepidoptera Lepidoptera YDB sp.1 1 1 1 Lepidoptera YDB sp.2 3 3 3 Lepidoptera YDB sp.3 4 1 1 2 1 3 Cosmopterygidae Cosmopterygidae YDB sp.1 41 26 15 41 Cosmopterygidae YDB sp.2 15 9 1 10 3 2 5 Geometidae Geometridae YDB sp.1 80 3 3 40 27 67 3 7 10 Oecophoridae Oecophoridae YDB sp. 1 50 8 8 25 17 42 Oecophoridae YDB sp. 10 6 4 1 5 1 1 Oecophoridae YDB sp. 2 57 1 1 2 30 16 46 6 3 9 Oecophoridae YDB sp. 3 2 1 1 1 1 Oecophoridae YDB sp. 4 10 2 8 10 Oecophoridae YDB sp. 5 13 9 9 1 3 4 Oecophoridae YDB sp. 6 3 3 3 Oecophoridae YDB sp. 7 32 1 1 12 12 14 5 19 Oecophoridae YDB sp. 8 7 1 1 3 3 3 3 Oecophoridae YDB sp. 9 4 3 1 4 Oecophoridae larvae Oecophoridae YDB sp. 11- larvae 1 1 1 Torticidae Tortricidae YDB sp. 3 4 3 3 1 1 Tortricidae Tortricidae YDB sp. 1 40 17 23 40 Tortricidae YDB sp. 2 15 1 1 10 4 14 Lepidoptera larvae UnID Lepidoptera larvae- Grouped 11 8 3 11 Neuroptera Hemerobiidae Micromus tasmaniae 2 1 1 1 1 Orthoptera Gryllacrididae Adult Kinemania YDB sp. 1 12 10 2 12 Juvenile Kinemania YDB sp. 1 5 5 5 Plecoptera Plecoptera YDB sp. 1 38 36 2 38 Eustheniidae Eusthenia spectabilis 10 9 1 10 Pseudoscorpionidae Pseudoscorpionidae YDB sp. 1 2 2 2 Psocoptera Psocoptera YDB sp. 1 2 2 2 Psocoptera YDB sp. 2 1 1 1 Psocoptera YDB sp. 3 1 1 1 Grand Total CD sticky Funnel Crawl Hangtraps All Traps 100yr Old Total 100yr Old Total 100yr Old Total Grand Total 6490 644 596 1240 1596 1128 2724 1245 1281 2526

Order Family Species binomial Psocoptera YDB sp. 4 2 2 2 Psocoptera YDB sp. 5 1 1 1 Trichoptera Trichoptera YDB sp. 1 1 1 1 Leptoceridae Leptoceridae YDB sp. 1 2 2 2 Unidentified Unidentified-Mangled Unidentified-Mangled 122 6 6 12 12 9 21 35 54 89 21) Appendix 7. Eucalyptus Canopy Arthropod Research Review

DRAFT

A Review of Canopy Arthropod Biodiversity Research in Australian Eucalyptus by Yoav Bar-Ness Written in October 2003.

Introduction

The Eucalyptus tree is the quintessential Australian tree, covering a vast range of habitats on the continent and occupying a critical position in the ecology of the continent. The genus Eucalyptus is a member of the biological subfamily Leptospermidae in the family Myrtaceae, and can be found over 35 degrees of latitude from the tropics of Indonesia to the temperate forests of Tasmania [Williams, 1997 #118]. In this article, the scientific literature reporting on arthropod biodiversity in the canopies of native Australian Eucalyptus trees is reviewed. Three criteria were used to select articles to include: 1. Adult Eucalyptus trees studied: Within the study, collected fauna were intended to be collected from specific Eucalyptus tree canopies, as opposed to ground level studies of other habitats within Eucalyptus forest, e.g. [Andersen, 2000 #44;Neumann, 1978 #114;Neumann, 1978 #113;Catterall, 2001 #86;Davies, 2000 #35;Michaels, 1995 #6;Oliver, 2000 #1;Peeters, 2002 #77;Peeters, 2001 #22;Richardson, 1999 #17;Yee, 2001 #9], or immature trees (Evelegh et al 2001). 2. Native forests: Research on native Eucalyptus forests in Australia, as opposed to urban plantings of native trees [e.g. Bhuller & Majer 2000] or forestry plantations in Australia or on other continents, [e.g. Withers, 2001 #10]. 3. Biodiversity focus: Fauna are examined at a wide range of taxa, as opposed to studies of a specific, narrower taxa, e.g.. [Neumann, 1978 #114;Bashford, 2001 #80;Davies, 2000 #35;Halaj, 1997 #96]. A synthesis of invertebrate research conducted in Eucalyptus forests has been written by [Majer, 1997 #139]. Their review article summarizes a wide range of research projects, such as those investigating native and plantation forests, above and below ground habitats, Australian forests, and foreign plantings. The research papers reviewed here are divided into two major categories: those focusing on arthropod or invertebrate biodiversity of the Eucalyptus canopy, and those focusing on the biodiversity phytophagous invertebrates. They are then arranged alphabetically and chronologically by the last name of the primary investigator and the year fieldwork began. Thus, scientific papers written about the same data are grouped together, in the order of their publication. Each review is presented of the goals, location, methods, and results of each investigation. When appropriate, quantitative figures or trends are given in relation to the stated goals of the research. The figure or table where these numbers are to be found or extrapolated is presented in the text of the review. When numbers are estimated from a graphical display, this is noted in the text. Basset (1997) has written a review of field methods used in studying canopy arthropods. Each method has biases and limitations, and the several different techniques used preclude an easy comparison of different investigations. [Basset, 1997 #83;Stork, 1997 #13] For comparison of the different investigations, a table is presented listing various information about the study: the location, time, Eucalyptus species, field methods, taxonomic resolution, primary investigator’s most recent contact organization, and potential comparisons possible from the study design. This last category represents the broad spectrum of hypotheses testable from the data collected by the study methods as reported, rather than the published results. For those seeking to begin a study on canopy arthropod biodiversity, papers on collecting and sorting techniques [Basset, 1997 #40;Richardson, 1999 #17], research directions [Stork, 1997 #13;Grove, 2000 #94], and study design and data processing [Marshall, 1994 #72] offer useful perspectives and experiences from other researchers. The “functional feeding groups,” or “guilds”, refer to the ecological role of invertebrates. There are seven guilds: 1) Phytophage chewers and Phytophage sapsuckers, 2) Epiphyte grazers, 3) Scavengers, dead wood, and fungus feeders, 4) Predators, 5) Parasitoids, 6) Ants, 7) Tourists. [Moran, 1982 #46;Southwood, 1982 #45]. Research in other forest regions of Australia and the world have created a context to compare canopy arthropod compositions in Eucalyptus forests [Basset, 1991 #39;Basset, 1991 #41;Basset, 2001 #42;Erwin, 1995 #33;Guilbert, 1994 #95;Hijii, 1984 #98;Floren, 2001 #90;Nadkarni, 1990 #112;Clement, 2001 #87;Longino, 1994 #24;Moran, 1982 #46;Southwood, 1982 #45;Parmenter, 1984 #76;Paoletti, 1991 #3;Peeters, 2001 #22;Peeters, 2002 #77;Stork, 2001 #12].

Studies not included: Articles which reiterate previously reviewed data in the context of studying another subject are not included. [Majer, 1992 #133;Recher, 1996 #146;Ohmart, 1983 #143] A significant body of work has been conducted on the specific ecological roles arthropods play in Eucalyptus ecosystems, especially herbivory [Lowman, 1995 #119;Fox, 1983 #61;Fox, 1986 #92;Haukioja, 1994 #70;Lamb, 1985 #56;Landsberg, 1989 #63;Landsberg, 1990 #66;Landsberg, 1995 #69;Landsberg, 1997 #34;Lowman, 1982 #26;Lowman, 1985 #55;Lowman, 1987 #60;Lowman, 1987 #58;Lowman, 1987 #64;Lowman, 1992 #74;Morrow, 1977 #71;Ohmart, 1984 #75;Ohmart, 1983 #115;Springett, 1978 #62;Stone, 1995 #68] and the phenomenon of Eucalyptus dieback [Palzer, 1983 #116;Lowman, 1987 #58;Landsberg, 1990 #66;Fox, 1983 #91].

Article Reviews:

Biodiversity studies not concentrating on phytophagous insects:

1982 Abbott Abbott, I. and Van Heurck, P. (1985). Tree species preference of foraging birds in Jarrah forest of Western Australia. Australian Wildlife Research. 12: 461-466. Abbott and Van Heurck’s 1982 field work and 1985 paper examined the arthropods in the crowns of 5 plant species (2 of which were eucalypts) of juvenile forest trees as a foraging resource for birds. The goal of this paper is to quantitatively analyse the foraging niches of forest birds. Sweep netting of arthropods on 10 trees within a 125 hectare forest watershed were compared to observations of birds foraging on trees. The number of individuals and proportion at the ordinal level within each tree are presented, as well as the arthropod biomass found within each tree. E. marginata had more individuals (1128) and average biomass (.222g) than C. calophylla (570 and .416g) (Table 3) Of 10 birds observed for foraging preferences, 7 were observed more often on E. marginata than C. calophylla (Table 1). Arthropod abundance and biomass are concluded to be important in predicting birds’ usage of different tree species.

1985 Abbott Abbott, I., Burbidge, T., Williams, M. and Van Heurck, P. (1992). Arthropod fauna of jarrah (Eucalyptus marginata) foliage in Meditteranean forest of Western Australia: Spatial and temporal variation in abundance, biomass, guild structure and species composition. Australian Journal of Ecology. 17: 263-274. Abbott, Burbidge, Williams, and Van Heurck’s 1985 field work and 1992 paper examine the canopy biodiversity of E. marginata. The three stated goals of the study are to: 1) Compare foliage invertebrates of coppice trees <2m high and pole trees ~14m high. 2) Examine diversity at a regional scale (7500 km2) rather than a more detailed smaller scale. 3) Examine patterns of fauna presence between trees. 4) Study the effect of the outbreak caterpillar Uraba lugens on the foliage invertebrate community. Branch clipping of foliage was used to collect invertebrates, which were sorted to morphospecies and functional feeding groups. The abundance and species richness of invertebrates per kg of dried foliage is presented to compare the coppice and pole foliage. This segment of the study was conducted on 3 pole and 3 coppice trees in each of 15 stands within a 50 km radius. Significant differences were detected with more ants in coppice than pole; and more U. lugens, and thus Lepidoptera in pole. Regional diversity and dyamics of foliage arthropod diversity is examined. Two samples were taken from 45 trees within a 50km radius, repeated 9 times over two years. Taxonomic composition, species accumulation curves for different taxa, and seasonality of major taxa are presented. Throughout the region, 396 species were collected, with 98% of all individuals being in one of 6 taxa: Araneae (5.4%), Lepidoptera (68.6%- mostly U. lugens), Diptera (3.8%), Hymenoptera (3.2%), Hemiptera (12.3%), and Coleoptera (5.0%) (Table 2). Arthropods were found to be mostly very common or very rare. Rate of species accumulation for these 6 taxa, plotting the discovery of new species against sampling effort, has leveled out for Araneae, Hymenoptera, and Diptera (Figure 3). This implies that the diversity of the other taxa- Hymenoptera, Hemiptera, and Lepidoptera was not sufficiently examined by the sample effort. January (summer) samples were the most species rich (Table 3). Tables comparing seasonal species richness, abundance and biomass are presented for the above 6 taxa, functional feeding groups, and taxa of note such as ants or U. lugens (Tables 3, 4, and 5). The spatial patterns of jarrah foliage expressed little difference between pole and coppice foliage. Arthropod community structure was not strongly correlated with structural features of the study trees. Comparison with other jarrah foliage studies shows different taxonomic compositions. The authors conclude “that there is no single fixed pattern of organization or predictable assemblage of invertebrates on jarrah foliage.” This conclusion of lack of spatial patterning is revisited in Burgman and Williams (1995). The U. lugens outbreak is examined and the biomass of this outbreak insect is concluded to be not significantly correlated with the biomass and abundances of other taxa. (Table 7). The hypothesis is presented that abundance of other invertebrates is limited by other factors than competition for foliage to feed upon.

1985 Abbott Burgman, M. and Williams, M. (1995). Analysis of the spatial pattern of arthropod fauna of jarrah (Eucalyptus marginata) foliage using a Mantel correlogram. Australian Journal of Ecology. 20: 455-457. Burgman and Williams re-analyse the data from the 1985 Abbot study and reveal a significant relationship between dissimilarities of tree arthropod diversity and distance from each other. A patch size of approximately 20 km is determined, within which tree are determined to have similar taxonomic composition. Abbott (2002) comments that this patch size may fluctuate during different seasons.

1990 Abbott. Abbott, I., Wills, A., Burbidge, T. and Van Heurck, P. (2002). Arthropod faunas of crowns of jarrah (Eucalyptus marginata) and marri (Corymbia calophylla) in mediterranean-climate forest: a preliminary regional-scale comparison. Australian Forestry. 63: 21-26. The method of branch clipping utilized in Abbott’s 1985 collection effort was repeated over in E. marginata and Corymbia calophylla. This research tests the hypothesis that adjacent pole sized trees of these two species have similar species richness, abundance, and biomass. Emphasis was placed on regional coverage over a wide area rather than temporal or local-scale comparisons. The investigators clipped one branch from each tree at approximately 15m using a cherry-picker vehicle. 45 trees were selected divided equally in 3 rainfall zones within a 35km radius. Insects were sorted to morphospecies and functional feeding group. For each tree species results are presented comparing species richness, abundance, and biomass of major taxa per kg of dried foliage. Of 80 species, 8 were found on both tree species, 29 were found only on E. marginata, and 43 found only on C. calophylla (Table 2). On average, E. marginata showed statistically insignificant higher totals of species recorded, individual abundance, and biomass. (Table 3). Significant differences were recorded with higher species richness, abundance and biomass of Lepidoptera and leaf chewers in E. marginata than C. calophylla, and a higher biomass of Araneae, and predators in C. calophylla. The authors report that the data does disprove the “hypothesis of no consistent diffeerences between the arthropod faunas of the crowns of jarrah and marri,” and recommend the construction of a canopy crane facility to properly investigate the matter. 1978 Bell Bell, H. (1985). Seasonal variation and the effects of drought on the abundance of arthropods in savanna woodland on the Northern Tablelands of New South Wales. Australian Journal of Ecology. 10: 207-221. Bell’s 1978 fieldwork investigated the seasonal changes of arthropod biomass and size one year before and during the two driest seasons yet recorded in Wollomombi, New South Wales. The arthropod data was a component of a bird foraging study. Bagging, shaking, and clipping of tree and shrub foliage was repeated monthly over three years. Eucalyptus species, sampled in the ratio of 6 stringybarks(E. calignosa, E. laevopinea): 4 boxes (E. conica, E. melliodora, E. bridgesiana): 1 gum (E. viminalis, E. amplifolia, E. blakelyi), were counted as one without differentiation. Data is presented showing results for Eucalyptus separately from the other plants. Graphs are presented showing monthly biomass recorded from Eucalyptus trees of arthropods, larvae, coleopterans, ants, scale insects, hemipterans, and spiders (Figures 4 through 11). The arthropod biomass on Eucalyptus trees was substantially lower during the drought years than the first year of study. Wintertime arthropods from Eucalyptus trees were significantly smaller than those collected in summer (Figure 13b). The summer to winter ratio of biomass on all plants was approximately 10:1 during the first year and approximately 2:1 during the later two drought years.

2002 Grove Grove, S., Bashford, R., Lillywhite, P. and Muirhead, A. (2002). Sampling the high life: Canopy fogging in Tasmanian wet Eucalyptus obliqua forest. 3rd International Canopy Conference poster, June 2002. Cairns, Queensland. A 2002 exploratory pilot project of canopy fogging at Warra Long Term Ecological Research site in the southern Tasmanian E. obliqua forest yielded 1813 individuals from 4 E. obliqua trees. The taxonomic composition of beetles on E. obliqua is compared between upper and lower crown samples, and to fogging of Nothofagus cuninghamii and Acacia melanoxylon. On E. obliqua, Hemiptera were most abundant (29%) followed by Coleoptera (26%), Diptera (10%), Collembola (9%), Hymenoptera- ants (9%), other Hymenoptera (5%), Acari (4%), Araneae (2%), Blattodea (2%), other insects (2%), Thysanura (1%) and Lepidoptera (1%) (Figure 1). 182 beetle morphospecies were distinguished and are listed on the poster. Beetle species within each tree (upper and lower samples) were more similar than between trees. (Figure 3.)

1986 Majer Majer, J. and Recher, H. (1988). Invertebrate communities of Western Australian eucalypts: a comparison of branch clipping and chemical knockdown procedures. Australian Journal of Ecology. 17: 383-393. The two methods of branch clipping and chemical knockdown are examined and compared in Majer and Recher’s 1986 study in the wandoo and jarrah forests of Western Australia. The Dryandra region (containing E. wandoo and E. marginata) and the Karragullen region (C. calophylla and E. marginata) were sampled by both methods, and another region of Dryandra (E. wandoo and E. accedens) was sampled by chemical knockdown only. The goals of this study were to compare the two methods for their detection of spatial patterns and abundance of arthropods, and to report of the animals detected on the three species sampled by both methods. For the chemical knockdown, 3-5 patches greater than 25m2 and containing 1- 10 trees were sampled. Single branches of at least 40 leaves from multiple trees were bagged and clipped. Animals collected were sorted, generally to order. Chemical knockdown collected an average per tree of 196 animals in E. marginata and 382 in E. wandoo at Dryandra, 3574 in C. calophylla and 311 in E. marginata at Karragullen, and 368 in E. accedens and 478 in E. wandoo at the second Dryandra site (Tables 1, 2, 5- note that C. calophylla is mislabelled as E. wandoo in Table 5.) The exceptionally high abundances reported for C. calophylla was due to a concentration of Hemiptera-psyllids. Branch clipping results are presented calibrated to leaf weight and to leaf area measured from a subset of 40 leaves. (Tables 4 and 6). Due to the presence of the psyllids, Karragullen results had much higher abundances of insect individuals per gram (0.5 for C. calophylla, 0.03 for E. marginata) than Dryandra (0.03, E. marginata, and 0.14 for E. wandoo). The authors report that with psyllids removed from data analysis, the difference between abundance in tree species at Karragullen is not statistically significant due to high variabilty. At Dryandra, E. wandoo had significantly higher abundance and biomass of invertebrates than E. marginata, as well as significantly higher abundances of Araneae, Blattodea, Thysanoptera, adult Coleoptera and adult Diptera. For all trees sampled, E. marginata showed the lowest abundance and biomass, a trend which is discussed in relationship to leaf levels of polyphenols. The two types of sampling method are concluded to report on relative abundance of arthropods. Knockdown samples animals living on the bark and mobile animals in the canopy, whereas clipping collects more immobile animals. The authors conclude that the effort involved in combining the two methods is minor enough to justify using them simultaneously.

1987 Majer Majer, J., Recher, H., Perriman, W. and Achutan, N. (1989). Spatial variation of invertebrate abundance within the canopies of Australian eucalypt forests. Studies in Avian Biology. 13: 65-72. The 1987 field work by Majer and Recher investigated the canopy arthropods of E. crebra and E. molucanna in New South Wales and C. calophylla and E. marginata in Western Australia using chemical knockdown (10 nets placed) and branch clipping (10 clips of 40 leaves removed). In both study regions, 10 trees were selected anew during each of four seasons. Non- flowering trees were sampled above 7 metres (canopy) and below (subcanopy). The relationship of these communities to birds (as a foraging resource) and to ground and foliage nutrients was investigated concurrently. In this paper, the stated goals are to study invertebrate abundances within a tree species and between tree species, and to compare the chemical knockdown and branch clipping results. Data is presented on mean arthropod abundances per tree at an ordinal and maturity level of resolution, with ants excluded from the knockdown totals. Within tree species, fauna collected in canopy chemical knockdown samples were generally more abundant than subcanopy samples. Only Araneae from E. crebra and Psocoptera on E. mollucana were significantly more abundant in subcanopy samples than upper canopy. The variability within trees was reported to be greater than the variability between trees Between the two sampled tree species, chemical knockdown resulted in higher levels of mean invertebrate abundances in C. calophylla (455) than E. marginata (351) in Western Australia, and higher levels in E. crebra (1797) than E. mollucana (1017) in New South Wales. (Tables 1 and 2) Branch clipping resulted in fewer taxa detected than chemical knockdown. (12 vs. 27 in WA and 17 vs. 27 in NSW). The ratio of relative abundances of insects between tree species were exaggerated in comparison to chemical knockdown. For each method, arthropod abundance was higher in the NSW samples than the WA samples.

1987 Majer Majer, J., Recher, H. and Ganeshanandam, S. (1992). Variation in foliar nutrients in Eucalyptus trees in eastern and Western Australia. Australian Journal of Ecology. 17: 383-393. In this paper the nutrient levels of the trees studied in Majer and Recher’s 1987 field work are analysed in relation to the arthropod fauna and bird fauna (which are both show higher values in eastern Australia than Western.) The goal of this paper is to explore the hypothesis that leaf nutritrion is responsible for the differences in invertebrate abundances in coexisting Eucalyptus species. Eastern Australian foliage was found to be more nutrient rich than Western Australian foliage. (Table 1) More nutrients were present in the leaves of E. crebra than E. mollucana and in C. calophylla over E. marginata. Nutrients levels were generally higher in the upper canopy than subcanopy. Invertebrate abundances from sampling in eastern Australia are higher than those from Western Australia. Abundances were generally higher on E. crebra than E. mollucana and in C. calophylla over E. marginata, and are generally higher in upper canopy than subcanopy. The possible impact of leaf polyphenols and leaf toughness are mentioned in the conclusion. The authors conclude that within the scope of their study, foliage nutrients influence invertebrate abundance.

1987 Majer Majer, J., Recher, H. and Postle, A. (1994). Comparison of arthropod species richness in eastern and western Australian canopies: a contribution to the species number debate. Memoirs of the Queensland Museum. 36:121-131. The results of the 1987 field work by Majer and Recher is again analysed in this paper, in which the data on species richness detected in the upper canopies by chemical knockdown (once in each season in both regions) are compared and contrasted Eastern forest sampling resulted in more individuals (50,900 vs. 16,500) than Western Australian forest sampling, as well as more species (977 vs. 691) The number of species in each arthropod family is reported in Table 2. The abundances of arthropods in each order collected on eastern trees is presented in Figure 1, and on Western trees in Figure 2. Species richness is plotted against arthropod order in Figure 3. The seasonal abundance of mean total individuals per tree collected is compared in Table 1. Totalling all seasons, E. crebra exhibited the highest total mean levels of abundance per tree (2917) followed by E. moluccana (2174), C. calophylla (893) and E. marginata (753). Both Western Australian species had exhibited the highest mean levels of abundance in the autumn season. In the east, the highest levels were detected in spring (E. crebra) and summer (E. moluccana). The authors comment on the absence of data in this study regarding other habitats (other plant species, soil, or litter), or host specificity between different plants or within a species. The concept of sigma diversity, or “seasonal turnover,” is indicated as an important factor to be studied. After addressing these ideas, the authors present some estimates of Australian arthropod diversity. They postulate that the high diversity levels found in their study’s limited scope suggest that these may be underestimates.

1987 Majer Majer, J. D., Recher, H. F. and Ganesh, S. (2000). Diversity patterns of eucalypt canopy arthropods in eastern and western Australia. Ecological Entomology. 25: 295-306. The fauna collected in the upper canopy nets used in Majer and Recher’s 1987 east-west canopy fogging efforts are reanalysed in this article. Species richness by family and estimated species accumulation curves are used to assess the adequacy of the sampling method in describing the theoretical total arthropod biodiversity attainable by this method. An updated listing of species richness within different taxa (generally families) is presented as Table 1, sorted by tree species. 976 species in 173 families were sorted from samples in eastern forests, and 687 species in 176 families in Western Australian forests. The total species richness per tree is presented as Table 2, (as distinct from the mean abundance per tree by season presented as Table 1 in Majer, Recher, and Postle (1994). E. crebra samples showed the highest level of species diversity (726), followed by E. moluccana (641), E. marginata (448) and C. calophylla (444). Bootstrap statistical techniques (random resampling of the original data) were used to create species accumulation curves for the total study, each tree species, and specific guilds by tree species. Tables 3 and 4 present the number of arthropod species sampled each season for each tree species, and the number of new species discovered with each sampling season for each tree species. These are shown in Figures 1, 2 and 3. In Table 5, an estimated asymptote predicting the theoretical total number of species to be found (as indicated by the species accumulation curve) is presented. The authors compare the total number of species actually found to this asymptote figure, and comment that their sampling was “reasonably complete.” These curves also suggest that sampling of carnivores and tourists was less complete than that of herbivores. A high correlation was detected between number of individuals collected in an order and number of species detected within that order. The authors comment that abundance data presented at the ordinal data may indicate the species richness within that order. The authors comment that sigma diversity, or temporal variation, should be included in for research on biodiversity. They conclude with a comparison of their results to estimates of global biodiversity.

1987 Majer Recher, H. F., Majer, J. D. and Ganesh, S. (1996). Seasonality of canopy invertebrate communities in eucalypt forests of eastern and western Australia. Australian Journal of Ecology. 21: 81-95. The seasonality of the arthropod fauna collected in Majer and Recher’s 1987 field is analysed in this 1996 paper with Ganesh (Ganeshenandam). The paper begins by outlining other research investigating the sigma (seasonal temporal) diversity of canopy arthropod fauna. This paper analyses their collection at an ordinal level for seasonal trends. The goal is to discover “seasonal trends of taxa in each forest,” and to determine if the “patterns of seasonality are the same in both areas.” Seasonal investigations for upper and lower (understorey) canopy sampling was performed on taxa sufficiently abundant for statistical analysis. Upper and lower canopy samples were considered separately, as lower canopy samples represemt groups of trees, and upper canopy samples represent single trees. Excluding ants, statistically significant differences were detected for total arthropod abundances 1) between eastern trees for the upper canopy samples (but not understorey), 2) between eastern seasons for understorey samples (but not upper canopy), 3) and between seasons for both upper and lower canopy Western samples (but not between tree species for either upper or lower samples)(Table 2). Figure 2 illustrates seasonal variations in man arthropod abundance per tree by taxa for all four tree species. The graphs for total arthropods (Figure 2q) show a larger range of values for eastern forests throughout the year than Western forests. This is the same data as used for Table 1 in Majer, Recher, and Postle (1994). Referring to that table in the earlier article for the precise figures on the graph, the highest absolute range of mean arthropod abundance per tree throughout the year was found in E. crebra (Spring high of ~844 individuals minus Winter low of ~490 equals ~354). This tree species was also found to be the most species diverse. (See Majer, Recher, and Ganesh (2000)). The next highest range was found in E. moluccana (Summer 742 minus Autumn 411 equals 331), followed by the Western Australian trees, E. marginata (Autumn 242 minus summer 125 equals 117) and C. calophylla (Autumn 252 minus spring 184 equals 68). Converted to proportional differences (Difference in abundances divided by high season abundances), the division between East and West is not so distinct: the eastern Australian trees do not yield uniformly higher values (E. crebra, .42 and E. moluccana, .45), but were intermediate between the Western Australian species (E. marginata, .48, and C. calophylla, .26). However, these simplifications of complex data sets are of limited value in determining seasonal variation, lacking values of variance between replications. To properly integrate the data with variance levels, the authors use a coefficient of variation (standard deviation divided by mean) as a measure of variation within for east and West samples. The mean and standard error of the mean abundance per tree was calculated for each season, each taxa, and each tree species. Comparing eastern and Western forests, the mean coefficient of variations was insignificantly different, with the in Western Australia was 79%, and in eastern, 61%. The authors comment on the rainfall pattern: Western Australia receives precipitation mostly during winter, while in New South Wales is spread more evenly across the year (Figure 1), and that this may be indicated by the slightly higher mean coefficient of variation for Western Australia. In conclusion, most taxa experience seasonal variation in abundance, and the patterns of seasonality in the different regions of the continent are distinct from each other. The abundance of eastern Australian arthropods tends to peak in the samples from the growing seasons of spring and summer, while in the Western samples, arthropods abundance tend to peak in autumn samples.

1987 Majer (west) Heterick, B., Majer, J., Recher, H. and Postle, A. (2001). A checklist of canopy, bark, soil and litter fauna of the Darling Plateau and adjacent woodland near Perth, Western Australia, with reference to the conservation of forest and woodland fauna. Curtin University of Technology School of Environmental Biology Bulletin. (21). This is a list of identified species and morphospecies from three forest entomology studies in Western Australia. The canopy arthropods presented in Table 1 are from the 1987 Majer investigation. Data analysis and discussion are minimized in this publication and the presented data is the taxonomic results of these studies. The study design and collection techniques are analyzed in detail in Majer et al (2002).

1987 Majer (west) Majer, J., Recher, H., Heterick, B. and Postle, A. (2002). The canopy, bark, soil, and litter invertebrate fauna of the Darling Plateau and adjacent woodland near Perth, Western Australia, with reference to the diversity of forest and woodland invertebrates. Pacific Conservation Biology. 7: 229-239. Data on four different habitats in Western Australian woodlands are presented from three research projects, and the taxonomic and feeding guild composition is analyzed. The canopy fauna obtained by chemical knockdown is the western portion of Majer and Recher’s 1987 collections. The bark fauna was obtained from eight study sites in a line southeast from Perth. C. calophylla was sampled at all eight sites, E. marginata at the start and end sites, and E. accedens and E. wandoo at the last site. Invertebrates were collected using bark drift fence traps and intercept traps between October 1998- October 1999. The soil and litter sampling was performed near Dwellingup, Western Australia during June 1980-September 1981 using litter removal and soil coring techniques to collect material. Tentative comparisons between the four habitats are discussed with regard to noteworthy differences in study design, taxonomic resolution, and study sites. For all tree species pooled, 561 species in 161 families were collected in the canopy habitat. 1234 species in 269 families in the bark habitat, 180 species in 25 families in the soil habitat, and 222 species in 38 families in the litter habitat (Table 1). In general, greater similarity of families was found within the aerial habitats (canopy and bark) and within the ground habitats (soil and litter) than between aerial and ground habitats (Figure 4). 126 families were found in both bark and canopy samples, 24 families were found in both ground habitats and bark samples, and 17 were found in both ground habitats and canopy samples. The relative composition of functional feeding guilds is presented as a proportion of each habitat’s total taxonomic diversity (Figure 5). Ground habitats were found to have a diverse array of fungivorous, scavenging, and saproxylic organisms, with approximately half of the identified taxa being assigned to this decomposer guild. Relative aerial habitat decomposer diversity was substantially lower (21.6% in the canopy and 16.9% in the bark habitat). Sapsuckers were more prevalent in ground habitats than in aerial habitats. Predators and parasitoids were more prevalent in aerial habitats than ground habitats. Epiphyte and leaf eaters were fond to be less diverse than other guilds. The authors conclude by comparing their results to estimates of global biodiversity and Australian insect diversity and comment that the results of their work suggest that these projections may be underestimates. They also comment that the recorded diversity of this Western Australian eucalypt ecosystem is less than that encountered in research in the tropical forests and greater than the temperate deciduous and coniferous forests.

1993 Majer Majer, J., Recher, H. and Keals, N. (1996). Branchlet shaking: A method for sampling tree canopy arthropods under windy conditions. Australian Journal of Ecology. 21: 229-234. Majer, Recher, and Keals compare the chemical knockdown techniques used in other studies with physical shaking of branches in their 1993 field work. E. wandoo in Western Australia and E. melliodora in New South Wales were sampled with both field techniques in the framework of a study examining habitat fragmentation and productivity. The Western Australian results of shaking branchlets in high winds are compared to chemical knockdown. 20 woodland (interior and edge) trees and 20 paddock trees were sampled in the two study areas during September and November 1993 and divided evenly between trees sampled by chemical knockdown and branchlet shaking. Arthropods collected were sorted to order and the 19 most abundant taxa (generally order and maturity) were analysed. Chemical knockdown collected more animals (mean 225 per tree) than branchlet shaking (mean 91 per tree). The authors remark that shaking results showed less of the smallest and largest animals than chemical knockdown. In the Western Australia woodland interior, when compared to chemical knockdown, branchlet shaking collected significantly less Acarina, significantly more Aranease, and significantly more Homoptera. In the woodland edge, significantly more Homoptera and Hymenoptera-others and significantly less adult Diptera and Hymenoptera-ants were collected by branchlet shaking. In the paddock sites, branchlet shaking collected signicantly less Acarina, Psocoptera, and Thysanoptera. For each combination of tree position and taxa (i.e. out of 3 positions x 19 taxa), 30 were more abundant in shaking, 20 less abundant and 7 equally abundant (Table 1). The authors remark that the extra step in sorting animals from branch litter in shaking may result in less smaller animals sampled, and that it does not record the presence of some foliage parasites that cling tightly to leaf surfaces. The authors conclude that branchlet shaking is a useful alternative if windy circumstances make chemical knockdown impossible.

1993 Majer Majer, J., Recher, H. and Keals, N. (1999). Canopy arthropod faunas in fragmented agricultural landscapes. In Temperate Eucalypt Woodlands in Australia: Biology, Conservation, Management, and Restoration (Eds, Hobbs, R. J. and Yates, C. J.) Surrey Beatty & Sons, Chipping Norton, New South Wales, pp. 235-247.

The branchlet shaking of Majer, Recher, and Keals’ 1993 field work is utilized in several sites and presented in this book chapter. Data from both New South Welsh E. melliodora and Western Australian E. wandoo is analysed in a study examining the canopy invertebrates and nutrient levels in isolated patches (remnant) of woodland reserves and corridors to a matrix of agricultural land (paddock). Three eucalypt woodland fragments (and adjacent wildlife corridors) in Western Australia and two in New South Wales were sampled in spring 1993. Sixty branchlets in the upper canopies of the study trees were shaken and invertebrates falling into a net were collected. Forty study trees were selected in each site, evenly divided into “remnant centre, remnant edge, paddock, and corridor” classes. Invertebrate data is presented as ordinal abundances per tree. In general, paddock and edge sites resulted in the most abundant collections of invertebrates (Table 3). The authors list detected significant results by taxa for study site location and tree location in Table 4, and quantify the general trend for tree locations by remarking that 23 of 31 statistically significant “most abundant” instances were in edge and paddock trees, and that 17 of 26 statistically significant “least abundant” instances were in centre and corridor trees. The postulated effect of habitat fragmentation is discussed in the conclusion. Possible reasons for the difference in abundances detected in different patch types put forth include more vigorous trees in paddock and edge sites, an aggregation effect on flying animals (analogous to a net), or lower predator levels. The authors state that these explanations do not account for the less abundant fauna in corridor sites.

1999 Radho-Toly Radho-Toly, S., Majer, J. D. and Yates, C. (2001). Impact of fire on leaf nutrients, arthropod fauna and herbivory of native and exotic eucalypts in Kings Park, Perth, Western Australia. Austral Ecology. 26(5):500-506. An urban park in Perth, Western Australia, was sampled for arthropods, nutrients, and herbivory damage by chemical knockdown in May 1999 by Radho- Toly, Majer, and Yates. Woodland unburnt since 1989 was compared to woodland burnt in 1996. Native E. marginata and E. gomphocephala were sampled alongside planted exotic E. cladocalyx and E. botyroides. The goal of the work was to document the arthropod fauna of native and exotic trees, examine the significance of fire on these animals, and to test the hypothesis that herbivory on native trees was higher than exotic trees. For the arthropod sampling, in each burn history, five trees of each species were fogged with insecticide and shaken onto plastic sheeting and the results identified to an ordinal level. Of 11 orders investigated, the most abundant were Araneae, Hemiptera, Coleoptera, and Formicidae. A remarkably high number of ants were found on E. gomphocephala (Table 4). On the burnt site, mean abundance of total arthropods excluding ants was highest on E. gomphocephala (34.6), followed by E. botyroides (22.0), E. cladocalyx (18.8), and E. marginata (16.0). For the unburnt site, mean abundance of total arthropods excluding ants was highest on lignotuberous E. marginata (11.2), followed by E. gomphocephala (9.8), E. botyroides (5.8) and E. caldocalyx (3.2). For all species except E. marginata, a significant difference in total arthropod abundance (without ants) was detected (Table 4). E. botyroides had the most nutritious foliage, as measured by nitrogen, potassium, and phosphorous, followed by E. cladocalyx, E. gomphocephala, and E. marginata (Table 1). The native plants had the foliage with the lowest nutrient levels. For all trees, total leaf damage was higher in the burnt area than the unburnt area (Table 2). E. gomphocephala had the lowest levels of leaf damage in both burnt and unburnt areas. The exotic plants showed lower levels of herbivory in the unburnt trees than E. marginata, but similar levels in the burnt trees (Table 2). The authors conclude that the their results “indicate there was no consistent difference in the levels of the various types of leaf damage or of arthropods on the exotic and native eucalypts.” This study suggests that the exotic plants are not eaten less, and that the higher nutrient levels in their leaves may attract local herbivores.

1979 Yen Yen, A. (1989). Overstorey invertebrates in the Big Desert, Victoria. In Mediterranean Landscapes in Australia: Mallee Ecosystems and their management (Eds, Noble, J. and Bradstock, R.) CSIRO, Melbourne, pp. 285-299. Yen’s 1979 study in desert Victoria studies the arthropod communities of foliage on mallee eucalypts. Yen’s 1989 paper investigates feeding guild composition among coppice and mature aged specimens of three species of Eucalyptus over different seasons. The goal of this study is to study the biodiversity of arthropod communities separately from herbivory, pest outbreak, or bird foraging. Arthropods were collected by beating the foliage of E. dumosa, E. foecunda, and E. incrassata in the Big Desert, and sorted to family level and functional feeding group. The collected animals are stored at the Museum of Victoria. 10 sampling efforts of 10 trees of each species, of each age, were conducted over 15 months. Coppice plants (regenerating after a 1977 fire) and mature plants (dating from 1959) are compared by order and functional feeding groups. Total number of individual animals collected was higher on coppice trees than mature trees (2937 vs. 2719)(Table 18.1). Overall, the most abundant fauna was collected on E. foecunda (2092), followed by E. dumosa (1914), and E. incrassata(1650). E. dumosa and E. incrassata were found to have more individuals on mature trees (983 and 925, respectively) than coppice trees ( 931 and 725) . E. foecunda had a more abundant fauna on coppice trees (1281 vs. 811) (Table 18.1) For all tree species, phytophagous invertebrates were more abundant on coppice trees than predators; with the reverse being true for mature trees. Coppice trees were marked by high abundance in summer, and mature trees by lowest abundance in winter. (Table 18.2) Seasonal variation in feeding group proportion was higher in mature than coppice trees. Seasonal changes in abundance of feeding group were remarked to be significant for scavengers and suckers and chewers. The hypothesis that coppice trees will suffer more from herbivory because of more abundant, and more nutritious young foliage is discussed. A survey of leaf area lost revealed more difference between species of tree than between age classes.

Studies concentrating on phytophagous insects:

1971 Burdon Burdon, J. and Chilvers, G. (1974). Fungal and insect parasites contributing to niche differentiation in mixed species stands of eucalypt saplings. Australian Journal of Botany. 22: 103- 114 The host specificity of phytophagous insects and leaf decay is examined by Burdon and Chilvers’ 1971 study. Observing the similar morphologies of co- dominant Eucalyptus trees, the investigators tested the hypothesis that the co- existence of these similar trees was due to fungal or insect parasites keeping one species from dominating over the other. In the Brindabella mountain range of the Australian Capital Territories, four sites at different elevations were chosen. Each site had codominant Eucalyptus species in the subgenera Symphomyrtus (730 metres altitude- E. viminalis; 1130m- E. dalrympleana; 1280m- E. dalrympleana; 1220m- E. dalrympleana) and in the subgenus Monocalyptus (730m- E. radiata; 1130m- E. dives; 1280m- E. delegatensis; 1220m- E. pauciflora): Adult beetles and leafhoppers were collected by leaf and stem search of 30-50 trees of each species at each site during the summer and autumn of 1971-1972. Animals were identified to morphospecies, or genus. Leaves with fungal decay were collected and the fungus analysed, and a sampling of leaves were analysed to generate statistics on leaf loss. 99 species of insects were identified. Percent of total and proportion on each subgenus is presented for the 17 most abundant species. Within these 17 taxa, more individuals were collected on Symphomyrtus (711) than Monocalyptus (354). 4 of these 17 were host specific- found on only one of the subgenera (Table 2). The host specificity of fungal and insect parasites is concluded to lend evidence to the hypothesis that at a subgeneric level, the ecological niche filled of these codominant eucalyptus species is in part defined by parasites rather than morphological differences.

1971 Burdon Burdon, J. and Chilvers, G. (1974). Leaf parasites on Altitudinal populations of Eucalyptus pauciflora Sieb. ex Spreng. Australian Journal of Botany. 22: 265-269 In the same issue of Australian Journal of Botany, Burdon and Chilvers present results of a preliminary survey of the impact of leaf parasites on regrowth saplings of the Monocalyptus species E. pauciflora. Individuals of E. dalrympeana were included in the lowest study site. The study was conducted in the Brindabella mountain range of the Australian Capital Territory at sites at elevation 1220, 1480, 1650, and 1750 metres. Individuals of E. dalrympeana were included in the 1220m study site. Leaf area loss and fauna collection methods were performed as described as in the earlier article, “Fungal and insect parasites contributing to niche differentiation in mixed species stands of eucalypt saplings”, reviewed above. In general, mean absolute leaf area on bud shoots, and percentage leaf area lost, decreased with altitude (Fig 1a and 1b). Concurrently, number of leaf parasite species decreased with altitude (Fig 1c). A listing of species or taxa is not included. Reading from the histogram of Figure 1c, an approximate total of 115 species of leaf parasite were found at 1220m, 85 species at 1480m, 80 species at 1650m, and 60 species at 1750m. Within the text, the authors remark that 53 species of beetles were found at 1220m altitude, and 23 species at 1750m. The authors discuss the relative impacts of high altitude environmental pressures, interspecific competition, and insect predation on E. pauciflora. They remark that at higher altitudes, physical environment may be more important in regulating growth than lower altitudes, where the biotic pressures are greater (i.e. competing Eucalyptus species and higher diversity of leaf parasites.

1987 Fensham Fensham, R. (1994). Phytophagous insect-woody sprout interactions in tropical eucalypt forest II. Insect community structure. Australian Journal of Ecology. 19: 189-196. The monsoon forests of Melville Island, Northern Territory, was studied by Fensham in 1987 in a study on herbivory on the regeneration of multiple stemmed trees. This 1994 paper on community structure reports on the seasonal change in taxonomic composition during 3 occasions- late dry season 1987, wet season 1988, and early dry 1988. 20 woody sprouts from each present Eucalyptus species were sweep sampled, leaf searched, and foliage beaten during each sampling effort in low forest (E. confertiflora, Buchanania obovota, Planchonia careya, Terminalia ferdinandiana, Erythrophleum chlorostachys), and tall forest (E. confertiflora, E. miniata, E. tetrodonta, Buchanania obovota, Planchonia careya, Terminalia ferdinandiana, Erythrophleum chlorostachys). A separate sampling effort in a second tall forest site was performed in the early dry 1988. Phytophagous animals were identified to morphospecies and are stored with CSIRO Tropical Ecosystem Research in Darwin. 3348 individuals, and approximately 271 identified species, were collected. More than half of the individuals were Psylloidea (36.9%) or other Hemiptera (39.8%). Data is presented showing mean abundance of insects per kg dry foliage in each season for Psylloidea, other Hemiptera, Lepidoptera, and all chewing insects on each plant species (Table 2). Insects were consistently more abundant in low forest than tall forest, with other Hemiptera being an exception, having about equal numbers in the late dry sample. Differences in Psylloidea abundance were dramatic: they were most abundant in the late dry season and mostly absent in the wet and early dry samples. They were approximately 7 times more abundant in low forest than wet forest. The authors conclude with a discussion of seasonal trends, host specificity, and fire regimes.

1973 Morrow Morrow, P. (1977). Host specificity of insects in a community of three co-dominant Eucalyptus species. Australian Journal of Ecology. 2: 89-106. The host specificity of insects collected in Mt. Kosciusko, New South Wales was analysed in Morrow’s 1977 paper describing her field work in 1973. The hypothesis that insect species have different herbivory influence on taxonomically close and distant Eucalyptus species is tested. Two Monocalyptus species, E. pauciflora and E. stellulata, and one Symphomyrtus species, E. perriniana, were sampled 13 times at least fortnightly in spring and summer 1973-1974. Branch shaking and sweep netting were used on lignotuberous regrowth dating from a 1960 power line clearance. Collected Coleoptera, Hemiptera and Orthoptera known to be phytophagous were sorted to species and are stored at the Australian National Insect Collection in Canberra. A total of 3901 individual insects were collected representing 141 species (Table 2). E. stellulata had the most abundant fauna (1741 individuals) followed by E. pauciflora (1093) and E. pauciflora (1741). The most species rich tree was E. pauciflora (88), followed by E. stellulata (83) and E. perriniana (62)(Table 2). A graph of seasonal arthropod abundance is presented for each tree species (Figure 1). December and January samples tended to be the most abundant for all three species. Host specificity data was processed using only those phytophagous species of which at least four individuals were collected at least four times. 63% of these animal species were recorded on only one of the Eucalyptus, and 32% were recorded on two. 27 insects were found solely on E. stellulata, 12 on E. perriniana, and 11 on E. pauciflora. More species were host specific on both E. stellulata and E. pauciflora (11) than E. pauciflora and E. perriniana (8) or E. stellulata and E. perriniana (2). Insect herbivores were concluded to discriminate between the single Symphomyrtus species and the two Monocalyptus species. This supports the hypothesis that the two more closely related species had different herbivory patterns than the other tree species. The author remarks that insect discrimination between the two Monocalyptus species was higher than she expected.

1981 Ohmart Ohmart, C., Stewart, L. and Thomas, J. (1983). Phytophagous insect communities in the canopies of three Eucalyptus forest types in south-eastern Australia. Australian Journal of Ecology. 8: 395-403. 1981 field work by Ohmart, Stewart, and Thomas explored the foliage insects on foliage of E. delegatensis, E. pauciflora, and E. dives in the Brindabella range of the Australian Capital Territory. The stated goal of this research is to see how insect herbivores contribute to nutrient flow and to learn about community structure of phytophagous insects in mature eucalyptus forests of south-eastern Australia. The ecological function of these insects is reported in Ohmart (1983a). Branch bagging and clipping was performed monthly over 5 months from the start of summer 1981 to spring 1982. Three sites, each dominated by one Eucalyptus species, were studied. Three sets of three trees in each site were selected on each occasion and four lower and six upper branches from each were gathered. All collected insects were identified to species and assigned as chewers or sapsuckers, and spiders counted. Data is presented with insect abundance calibrated to kg of dry foliage. 122 species of phytophagous species of insect were encountered: 39 on E. delegatensis, 35 on E. pauciflora, and 48 on E. dives (Table 2). Major taxa included Microlepidoptera, Chrysomelid beetles, Curculionid beetles, and leaf-hoppers. According to Appendix 1, in which “most of the species encountered in the study” are recorded, 73 species of insect are listed in 3 orders: 35 species of Lepidoptera, 31 of Coleoptera, and 7 of Hemiptera. Psyllidae and gallmakers were almost entirely absent, in contrast to Morrrow (1977) and Burdon and Chilvers (1974). By foliage weight, E. dives had the most phytophagous insects, followed by E. pauciflora and E. delegatensis (Table 3). A comparison of insects found in upper and lower foliage samples is presented as Table 4. For all 3 tree species , Chrysomelid beetles. Curculionid beetles, Lepidopterans, leaf hoppers and spiders were more abundant in the lower crown than upper crown samples, with 3 exceptions: Curculionid beetles and leaf-hoppers in E. delegatensis, and Curculionid beetles in E. dives. The authors conclude that the results suggest defoliation levels and defoliator abundance in Eucalyptus forests is approximately equal to reported values from northern temperate forest ecosystems.

1979 Woinarski Woinarski, J. and Cullen, J. (1984). Distribution of invertebrates on foliage in forests of southeastern Australia. Australian Journal of Ecology. 9:207-232. The 1979 field work by Woinarski and Cullen looked at leaf insects found on a range of plant species across the state of Victoria. More study sites and plant species were considered in this work than any of the other investigations. The goal of this project was to learn about the role of foliage invertebrates over a wide range without focusing on economic entomology, single taxa, specific questions of ecological function, or their availability as prey. The invertebrate communities detected are compared across plant species, seasons, and flower incidence. 16 study sites representing different ecosystems across the state of Victoria were selected and sampled at least seasonally between May 1979 and June 1982. In total, 22 Symphomyrtus and 9 Monocalyptus species of Eucalyptus were included, in addition to 125 other vascular plant species (Appendix A). Branch clipping of one branch from one plant of each species present at each collection yielded invertebrates larger than 3mm that were classified generally to order and larval or adult stage. (Arachnida; Psocoptera; Hemiptera- psyllids with lerps; Hemiptera psyllids without lerps; Hemiptera- other; Thysanoptera; adult Coleoptera; adult Diptera; adult Lepidoptera and Neuroptera; Hymenoptera-ants; adult Hymenoptera-other; larvae; other invertebrates). Shannon-Weiner diversity indexes were generated using these 13 taxonomic categories. Data is presented as mean density per m2 of foliage. Within the Eucalyptus tree species, subgeneric placement in Monocalyptus or Symphomyrtus was found to be an important factor in determining invertebrate communities. For all taxa except other Hymenoptera, and for all animals combined, Symphomyrtus showed higher density of invertebrates and higher mean diversity index than Monocalyptus (Figure 2c). The authors remark that the ranking of taxa by density was alike for both subgenera and for all Eucalyptus species studied, if not the overall density. When compared to the other plants studied, Eucalyptus showed a significantly lower density of animals collected. More specifically, Eucalyptus showed significantly higher densities of larvae, lerp-forming psyllids, and non-lerp- forming psyllids, and significantly lower densities of Arachnida, Psocoptera, other Hemiptera, Thysanoptera, Coleoptera, and Diptera. The seasonality of Eucalyptus invertebrates is presented by taxonomic group. (Figure 3) Total abundance was highest in spring and lowest in winter. Diversity was highest in summer and autumn and lowest in winter. The presence of flowers on all plants studied shows a significantly higher mean density and diversity index of invertebrates than flowers absent. Results confined to Eucalyptus are not presented. (Figure 2a) The authors conclude that these three factors- Eucalyptus subgenus, season, and flower presence- were the most important in differentiating invertebrate assemblages in their results.

1995 Wotherspoon Wotherspoon, K. P. (1998). Patterns of phytophagous insect herbiv ory and abundance on juvenile regrowth of Eucalyptus subgenera coexisting in southeastern Tasmania. Australian Journal of Ecology. 23: 430-442. The 1995 field work by Wotherspoon in late spring- early autumn 1995-1996 examined phytophagous insects on branches near Hobart, Tasmania in a coppice regrowth site beneath a power line clearance cut two years earlier. This research tested the hypothesis that juvenile trees in Symphomyrtus had more leaf damage and more insect abundance and diversity than Monocalyptus. The phyophagous insects found on two Monocalyptus species, E. obliqua and E. pulchella, and two Symphomyrtus trees, E. viminalis and E. globulus, were studied by bagging and clipping branches of juvenile foliage. 320 branches from each species were collected over 5 collection efforts, and the animals sorted into functional group – suckers or chewers-, and generally family within Coleoptera and Hemiptera. When occurring as 5 or more individuals, insects were sorted to morphospecies (Table 7). Vouchers are stored in the Centre for Environmental Studies at the University of Tasmania, Hobart. The most species of phytophagous insects was encountered on E. obliqua (10 suckers and 16 chewers) followed by E. viminalis (14 suckers and 8 chewers), E. pulchella (11 suckers and 9 chewers), and E. globulus (8 suckers and 6 chewers) (Table 6). The most abundant group encountered was Psyllidae, encountered at very high levels on foliage of E. globulus. The highest abundance of individuals per kg of dry foliage was found on E. globulus (3299.48), followed distantly by E. pulchella (146.46), E. obliqua (122.37), and E. viminalis (87.16) (Table 5). Sampling of leaves showed the mean percent leaf area lost to total chewing damage was significantly higher in Monocalyptus (4.76%)than Symphomyrtus (3.26%), but the difference for total insect damage was insignificantly higher (10.62% vs. 9.20%). The author concludes that the hypothesis of higher insect damage on Symphomyrtus was not upheld by the results, and that the hypothesis that Symphomyrtus has more individuals and more diversity than Monocalyptus was also not upheld by the results.

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Palzer, C. (1983). “Crown Symptoms of Regrowth Dieback.” Pacific Science 37(4): 465-470.

Paoletti, M., Taylor, R., Stinner, B., Stinner, D. and Benzing, D. (1991). “Diversity of soil fauna in the canopy and forest floor of a Venezualan cloud forest.” Journal of Tropical Ecology 7: 373-383.

Parmenter, R.R. and MacMahon, J.A. (1984). “Factors influencing the distribution and abundance of ground dwelling beetles (Coleoptera) ina shrub steppe ecosystem: the role of shrub architecture.” Pedobiologia 27: 21-34.

Paviour-Smith, K. and Elbourn, C. (1993). “A quantitative study of the fauna of small dead and dying wood in living trees in Wytham Woods, near Oxford.” English Nature Science 7: 373-383.

Peeters, P.J. (2002). “Correlations between leaf structural traits and the densities of herbivorous insect guilds.” Biological Journal of the Linnean Society 77: 43-65.

Peeters, P.J., Read, J. and Sanson, G.D. (2001). “Variation in the guild composition of herbivorous insect assemblages among co-occurring plant species.” Austral Ecology 26(4): 385-399.

Pik, A. and Beattie, A. (1999). “Taxonomic sufficiency in ecological studies of terrestrial invertebrates.” Australian Jounral of Ecology 24: 555-562.

Pinheiro, F., Diniz, I.R., Coelho, D. and Bandeira, M.P.S. (2002). “Seasonal pattern of insect abundance in the Brazilian cerrado.” Austral Ecology 27(2): 132-136.

Radho-Toly, S., Majer, J.D. and Yates, C. (2001). “Impact of fire on leaf nutrients, arthropod fauna and herbivory of native and exotic eucalypts in Kings Park, Perth, Western Australia.” Austral Ecology 26(5): 500-506.

Ranius, T. and Hedin, J. (2000). “The dispersal rate of a beetle, Osmoderma eremita, living in tree hollows.” Oecologia 126(3): 363-370.

Recher, H. and Majer, J. (1996). “One Humble Gum Tree: Home to 1000 Species.” GEO Australasia 18(6): 20-29.

Recher, H.F., Majer, J.D. and Ganesh, S. (1996). “Seasonality of canopy invertebrate communities in eucalypt forests of eastern and western Australia.” Australian Journal of Ecology 21: 81-95.

Recher, H.F., Majer, J.D. and Ganesh, S. (1996). “Eucalypts, arthropods and birds: On the relation between foliar nutrients and species richness.” Forest Ecology and Management 85(1-3): 177-195.

Richardson, B.J., Azarbayjani, F.F., Burgin, S. and Richardson, S. (1999). “Arboreal arthropod biodiversity in woodlands: Effect of collection procedures and geographic distance on estimates of diversity found on two species of Melaleuca.” Australian Journal of Ecology 24(5): 544-554.

Robson, B. and Chester, E. (1999). “Spatial patterns of invertebrate species richness in a river: the relationship between riffles and microhabitats.” Australian Journal of Ecology 24: 599-607.

Southwood, T.E., Moran, V. and Kennedy, C. (1982). “The richness, abundance, and biomass of the arthropod communities on trees.” Journal of Animal Ecology 51: 635- 649.

Springett, B. (1978). “On the ecological role of insects in Australian eucalypt forests.” Australian Journal of Ecology 3: 129-139.

Stone, C. and Bacon, P. (1995). “Leaf dynamics and insect herbivory in a Eucalyptus camaldulensis forest under moisture stress.” Australian Journal of Ecology 20: 473- 481.

Stork, N.E., Didham, R.K. and Adis, J. (1997). Canopy arthropod studies for the future.In: Canopy Arthropods. N. E. Stork, J. Adis and R. K. Didham. London, Chapman & Hall: 551-562. .

Stork, N.E., Hammond, P.M., Russell, B.L. and Hadwen, W.L. (2001). “The spatial distribution of beetles within the canopies of oak trees in Richmond Park, UK.” Ecological Entomology 26(3): 302-311.

Swengel, A. (2001). “A literature review of insect responses to fire, compared to other conservation managements of open habitat.” Biodiversity and conservation 10: 1141- 1169.

Taylor, R.J. and Doran, N. (2001). “Use of terrestrial invertebrates as indicatoirs of the ecological sustainability of forest management under the Montreal Process.” Journal of Insect Conservation 5: 221-231.

Williams, J. (1990). “The importance of herbivory in the population dynamics of three sub-alpine eucalypts in the Brindabella range, south-east Australia.” Australian Journal of Ecology 15: 51-55.

Williams, J. and Brooker, I. (1997). Eucalypts: an introduction.In: Eucalypt Ecology. J. Williams and J. Woinarski. Cambridge, Cambridge University Press. .

Withers, T.M. (2001). “Colonization of eucalypts in New Zealand by Australian insects.” Austral Ecology 26(5): 467-476.

Woinarski, J. and Cullen, J. (1984). “Distribution of invertebrates on foliage in forests of southeastern Australia.” Australian Journal of Ecology 9: 207-232.

Wotherspoon, K.P. (1998). “Patterns of phytophagous insect herbivory and abundance on juvenile regrowth of Eucalyptus subgenera coexisting in southeastern Tasmania.” Australian Journal of Ecology 23(5): 430-442.

Yee, M., Yuan, Z., and Mohammed, C. (2001). “Not just waste wood: decaying logs as key habitats in Tasmania's wet sclerophyll Eucalyptus obliqua production forests: the ecology of large and small logs compared.” Tasforests 13(1): 119-128.

Yen, A. (1989). Overstorey invertebrates in the Big Desert, Victoria.In: Mediterranean Landscapes in Australia: Mallee Ecosystems and their management. J. Noble and R. Bradstock. Melbourne, CSIRO: 285-299. . Table 1. A comparison of published research projects of Australian eucalyptus canopy arthropod biodiversity. Investigation Location Time Eucalyptus Method General Primary Comparisons Species Taxonomic Investigators potentially sampled Resolution Affiliation or investigated: Contact A. Spatial continental B. Spatial /fire local C. Spatial within tree D. Temporal/Seasonal E. Collecting methods F. Between Euc. species G. Within Euc. species 1982 Abbott 32o 51'S Oct marginata, Sweep net Order; F. Between Euc. 116o 25' E 1982 calophylla on foliage "Broad species Yarragil 4L up to 4.4 taxonomic G. Within Euc. Western meters categories" species Australia 1985 Abbott 34o 15' S Sep marginata Branch Morphospecies; B. Spatial /fire 116o 10' E 1985- clipping Functional local within 50 km Jan feeding group: C. Spatial within of Manjimup, 1988 tree Western G. Within Euc. Australia species

1990 Abbott. ~33o 21’ S Sep marginata, Branch Morphospecies CALM: Dept. B. Spatial /fire ~116o 09’ E 1990 callophylla clipping ; of local Collie, Functional Conservation F. Between Euc. Southwestern groups and Land species Western Management, G. Within Euc. Australia Western species Australia 1978 Bell 30o32’S Sep Studied as one: Branch Larvae, beetles, Dept of. B. Spatial /fire 152o 02’ E 1978- caliginosa, clipping ants, scale Zoology, Uni local Northern Aug laevopinea, insects, of New D. Tablelands, 1981 conica, hemipterans, England, Temporal/Seasonal New South melliodora, isopterans, Armidale, Wales bridgesiana, spiders, and New South viminilis, orthopterans; Wales amplifolia, Species blakelyi 1971 Burdon ~35o S Dec radiata, dives, Stem and Insect leaf Botany Dept. B. Spatial /fire ~141o E 1971- delegatensis, leaf parasites only; Australian local Brindabella Oct pauciflora, searching Species National F. Between Euc. Range 1972 viminalis University, species Australian dalrympleana, Canberra G. Within Euc. Capitol species Territory 1987 11o 44’S Sep confertiflora, Leaf Phytophagous Dept. of B. Spatial /fire Fensham 130o 41’E 1987- miniata, searching , insects only; Geography local Melville Jun tetrodonta Foliage morphospecies and D. Island, 1988 beating, Environmental Temporal/Seasonal Northern Sweep Studies, Uni. F. Between Euc. Territory sampling of Tasmania, species Hobart, G. Within Euc. Tasmania species 2002 Grove 43o 3’ S 2002 obliqua Chemical Order; beetles Forestry G. Within Euc. 146o 39’ E knockdown to Tasmania, species Warra LTER, morphospecies Hobart, Tasmania Tasmania 1986 Majer ~32o S Mar wandoo, Chemical Order B. Spatial /fire ~116o E 1986 marginata, knockdown; local Dryandra State wandoo, branch E. Collecting Forest accedens, clipping methods Western calophylla, F. Between Euc. Australia and marginata species Karragullen, G. Within Euc. Western species Australia 1987 Majer 33o 53' S Feb East: crebra, Chemical Morphospecies A. Spatial 150o 51' E 1987- moluccana knockdown; continental Scheyville, Jan West: branch C. Spatial within New South 1988 calophylla, clipping tree Wales (East) marginata D. Apr Temporal/Seasonal 32o 04' S 1987- E. Collecting 116o 07' E Nov methods Karragullen, 1989 F. Between Euc. Western species Australia G. Within Euc. (West) species 1993 Majer 31o 09' S Sep wandoo, Chemical Order Dept. of A.Spatial 117o 05' E 1993- melliodora knockdown; Environmental continental Amery Nov branchlet Biology, B. Spatial /fire Reserve, 1993 shaking Curtin Uni of local Western Technology, E. Collecting Australia Bentley, methods 31o 10' S Western G. Within Euc. 117o 14' E Australia species Namalcatching Reserve, Western Australia 31o 11' S 117o 23' E Wyalkatchem Reserve, Western Australia 30o 26' S 151o 36' E Newholme, New South Wales 30o 26' S 152o 00' E Thalgarrah, New South Wales

1973 Morrow 31o 12' S Oct perriniana, Foliage Phytophagous Dept. of D. 148o 43' E 1973- stellulata, shaking and Coleoptera, Ecology, Temporal/Seasonal Dainer's Gap, Mar pauciflora sweep Hemiptera and University of F. Between Euc. Mt. Kusciusko 1974 netting Orthoptera Minnesota, St. species New South only; Paul, G. Within Euc. Wales morphospecies Minnesota, species USA 1981 Ohmart 35o 20' S Dec delegatensis, Branch Spiders CSIRO Forest B. Spatial /fire 141o 50' E 1981- pauciflora, clipping counted; Research, local Brindabella Apr dives Phytophagous Canberra, C. Spatial within Range, 1982 insects only; tree Australian morphospecies F. Between Euc. Capital species Territory G. Within Euc. species 1999 Radho- 31o 56' S May marginata, Chemical Order Dept. of B. Spatial /fire Toly 115o 50' E 1999 gomphocephala, knockdown Environmental local Kings Park, cladocalyx, Biology, F. Between Euc. Perth, Western botyroides Curtin Uni of species Australia Technology, G. Within Euc. Bentley, species Western Australia 1979 16 sites across May behriana, Branch Invertebrates Dept. of A. Spatial Woinarski state of 1979- blakelyi, clipping larger than Zoology, continental Victoria June botryoides, 3mm: Divided Monash B. Spatial /fire 1982 amaldulensis, into 13 University, local cephalocarpa, categories, Melbourne, D. cypellocarpa, mostly order + Victoria Temporal/Seasonal dalrympleana, maturity F. Between Euc. dumosa, species foecunda, goniocalyx, incrassata, largiflorens, leucoxylon, meiliodora, microcarpa, ovata, polyanthemos, plybractea, sideroxylon, st.johnii, viminalis, viridis, baxteri, delegatensis, dives , macrorhynca, obliqua, pauciflora, radiata, regnans, sieberi 1995 42o 55' 30" S Nov globulus, Branch Phytophagous Dept. of F. Between Euc. Wotherspoon 147o 16' 48" E 1995- viminalis, clipping insects only: Geography species SW of Hobart, Mar obliqua, morphospecies; and G. Within Euc. Tasmania 1996 pulchella Feeding Group Environmental species Studies, Uni. of Tasmania, Hobart, Tasmania 1979 Yen 35o 42' S Sep dumosa, Foliage Family; Melbourne B. Spatial /fire 141o 52' E 1979- foecunda, beating Feeding Group Museum, local Lake Dec incrassata over tray Melbourne, D. Albacutya, 198 Victoria Temporal/Seasonal The Big F. Between Euc. Desert, species Victoria G. Within Euc. species

Table 2. Literature citations reporting on Australian eucalyptus canopy arthopod biodiversity research.

Investigation: Year and Selected citations reporting reults: First Author 1982 Abbott Abbott, I. and Heurck, P. V. (1985). Tree species preference of foraging birds in Jarrah forest of Western Australia. Australian Wildlife Research. 12: 461-466. 1985 Abbott Abbott, I., Burbudge, T., Williams, M. and Heurck, P. V. (1992). Arthropod fauna of jarrah (Eucalyptus marginata) foliage in Meditteranean forest of Western Australia: Spatial and temporal variation in abundance, biomass, guild structure and species composition. Australian Journal of Ecology. 17: 263-274. Burgman, M. and Williams, M. (1995). Analysis of the spatial pattern of arthropod fauna of jarrah (Eucalyptus marginata) foliage using a Mantel correlogram. Australian Journal of Ecology. 20: 455-457. Abbott, I. and Wills, A. (2001). Biodiversity of canopy arthropods in Jarrah forest of south-west Western Australia: review of ecological theory and conservation management. Pacific Conservation Biology. 7: 101-117. 1990 Abbott. Abbott, I., Wills, A., Burbidge, T. and Van Heurck, P. (2002). Arthropod faunas of crowns of jarrah (Eucalyptus marginata) and marri (Corymbia calophylla) in mediterranean-climate forest: a preliminary regional-scale comparison. Australian Forestry. 63: 21-26. 1978 Bell Bell, H. (1985). Seasonal variation and the effects of drought on the abundance of arthropods in savanna woodland on the Northern Tablelands of New South Wales. Australian Journal of Ecology. 10: 207-221. 1971 Burdon Burdon, J. and Chilvers, G. (1974). Fungal and insect parasites contributing to niche differentiation in mixed species stands of eucalypt saplings. Australian Journal of Botany. 22: 103- 114 Burdon, J. and Chilvers, G. (1974). Leaf parasites on altitudinal populations of Eucalyptus pauciflora Sieb ex. Spreng. Australian Journal of Botany. 22: 265-269. 1987 Fensham Fensham, R. (1994). Phytophagous insect-woody sprout interactions in tropical eucalypt forest II. Insect community structure. Australian Journal of Ecology. 19: 189-196. 2002 Grove Grove, S., Bashford, R., Lillywhite, P. and Muirhead, A. (2002). Sampling the high life: Canopy fogging in Tasmanian wet Eucalyptus obliqua forest. 3rd International Canopy Conference poster, June 2002. Cairns, Queensland. 1986 Majer Majer, J. and Recher, H. (1988). Invertebrate communities of Western Australian eucalypts: a comparison of branch clipping and chemical knockdown procedures. Australian Journal of Ecology. 17: 383-393. 1987 Majer East Majer, J., Recher, H., Perriman, W. and Achutan, N. (1990). Spatial variation of invertebrate & abundance within the canopies of Australian eucalypt forests. Studies in Avian Biology. 13: 65-72. 1987 Majer West Majer, J., Recher, H. and Ganeshanandam, S. (1992). Variation in foliar nutrients in Eucalyptus trees in eastern and Western Australia. Australian Jourral of Ecology. 17: 383-393. Majer, J., Recher, H. and Postle, A. (1994). Comparison of arthropod species richness in eastern and western Australian canopies: a contribution to the species number debate. Memoirs of the Queensland Museum. 36:121-131. Majer, J. D., Recher, H. F. and Ganesh, S. (2000). Diversity patterns of eucalypt canopy arthropods in eastern and western Australia. Ecological Entomology. 25: 295-306. Recher, H. F., Majer, J. D. and Ganesh, S. (1996). Eucalypts, arthropods and birds: On the relation between foliar nutrients and species richness. Forest Ecology and Management. 85: 177- 195. Recher, H. F., Majer, J. D. and Ganesh, S. (1996). Seasonality of canopy invertebrate communities in eucalypt forests of eastern and western Australia. Australian Journal of Ecology. 21: 81-95. 1987 Majer West Heterick, B., Majer, J., Recher, H. and Postle, A. (2001). A checklist of canopy, bark, soil and litter and fauna of the Darling Plateau and adjacent woodland near Perth, Western Australia, with reference to the conservation of forest and woodland fauna. Curtin University of Technology School of Environmental Biology Bulletin. (21). Majer, J., Recher, H., Heterick, B. and Postle, A. (2002). The canopy, bark, soil, and litter invertebrate fauna of the Darling Plateau and adjacent woodland near Perth, Western Australia, with reference to the diversity of forest and woodland invertebrates. Pacific Conservation Biology. 7: 229-239. 1993 Majer Majer, J., Recher, H. and Keals, N. (1996). Branchlet shaking: A method for sampling tree canopy arthropods under windy conditions. Australian Journal of Ecology. 21: 229-234 Majer, J., Recher, H. and Keals, N. (1999). Canopy arthropod faunas in fragmented agricultural landscapes. In Temperate Eucalypt Woodlands in Australia: Biology, Conservation, Management, and Restoration (Eds, Hobbs, R. J. and Yates, C. J.) Surrey Beatty & Sons, Chipping Norton, New South Wales, pp. 235-247.

1973 Morrow Morrow, P. (1977). Host specificity of insects in a community of three co-dominant Eucalyptus species. Australian Journal of Ecology. 2: 89-106. 1981 Ohmart Ohmart, C., Stewart, L. and Thomas, J. (1983). Phytophagous insect communities in the canopies of three Eucalyptus forest types in south-eastern Australia. Australian Journal of Ecology. 8: 395-403. Ohmart, C., Stewart, L. and Thomas, J. (1983). Leaf consumption by insects in three Eucalyptus forest types in Southeastern Australia and their role in short-term nutrient cycling. Oecologia. 59: 322-330. 1999 Radho-Toly Radho-Toly, S., Majer, J. D. and Yates, C. (2001). Impact of fire on leaf nutrients, arthropod fauna and herbivory of native and exotic eucalypts in Kings Park, Perth, Western Australia. Austral Ecology. 26(5):500-506. 1979 Woinarski Woinarski, J. and JM, C. (1984). Distribution of invertebrates on foliage in forests of southeastern Australia. Australian Journal of Ecology. 9:207-232. 1995 Wotherspoon Wotherspoon, K. P. (1998). Patterns of phytophagous insect herbivory and abundance on juvenile regrowth of Eucalyptus subgenera coexisting in southeastern Tasmania. Australian Journal of Ecology. 23: 430-442. 1979 Yen Yen, A. (1989). Overstorey invertebrates in the Big Desert, Victoria. In Mediterranean Landscapes in Australia: Mallee Ecosystems and their management (Eds, Noble, J. and Bradstock, R.) CSIRO, Melbourne, pp. 285-299.

22) Appendix 8. Spider Species Composition

The following work was performed by Lisa Boutin, of the Queen Victoria Museum and Art Gallery, Launceston Tasmania. She selected spiders from the archived specimens and compiled the following information between May and October 2005. These represent a subset of the collected animals, based on her taxonomic interests, and therefore is not a representative sample. "Static" traps are from ground-level flight-intercept traps placed by Dick Bashford (Forestry Tasmania) at Warra

DATE FAMILY(SUPER) GENUS (SUBFAM) SPECIES M F tree # old?/100 COLLECTING TYPE TRAP # notes 30.iii.2005 Theridiidae Achaearanea 1 7 100 Funnel crawl funnel 11 i-iii-2004 Gnaphosidae Acroceridae parasite immature 10 old Funnel crawl funnel 9 Acrocerid in spider i-iii-2004 Desidae Badumna 1 8 old Funnel crawl funnel 5 i-iii-2004 Clubionidae Clubiona A 2 1 100 Funnel crawl funnel 3 i-iii-2004 Clubionidae Clubiona A 3 1 100 Funnel crawl funnel 1 i-iii-2004 Clubionidae Clubiona A 1 1 100 Funnel crawl funnel 5 30.iii.2005 Clubionidae Clubiona A 1 1 100 Funnel crawl 8 funnel i-iii-2004 Clubionidae Clubiona A 1 1 100 01Hang 6 i-iii-2004 Clubionidae Clubiona cycladata 1 1 100 Hanging Trap 01Hang 4 i-iii-2004 Clubionidae Clubiona X 1 1 100 Flight intercept Tree 1 ant mimic i-iii-2004 Clubionidae Clubiona cycladata 1 2 100 Funnel crawl Funnel 5 i-iii-2004 Clubionidae Clubiona A 1 4 old Flight intercept 04Hang 6 30.iii.2005 Clubionidae Clubiona A 1 4 old Funnel crawl 11 funnel i-iii-2004 Clubionidae Clubiona A 1 5 100 Flight intercept 05 Hang 7 i-iii-2004 Clubionidae Clubiona A 1 7 100 Flight intercept 07Hang 3 i-iii-2004 Clubionidae Clubiona A 2 7 100 Funnel crawl funnel 6 i-iii-2004 Clubionidae Clubiona A 11 7 100 Funnel crawl funnel 1 i-iii-2004 Clubionidae Clubiona A 31 7 100 Funnel crawl 11 funnel i-iii-2004 Clubionidae Clubiona cycladata 1 7 100 Funnel crawl funnel 11 i-iii-2004 Clubionidae Clubiona A 1 8 old Funnel crawl funnel 7 i-iii-2004 Clubionidae Clubiona A 1 9 Funnel crawl funnel 5 i-iii-2004 Clubionidae Clubiona A 1 9 100 Funnel crawl funnel 11 i-iii-2004 Clubionidae Clubiona cycladata 1 9 100 Funnel crawl funnel 11 Clubionidae Clubiona I 1 10 old Flight intercept Tree 10 i-iii-2004 Clubionidae Clubiona A 1 12 Funnel crawl funnel 5 i-iii-2004 Clubionidae Clubiona cycladata 1 12 old Funnel crawl funnel 3 i-iii-2004 Clubionidae Clubiona A 1213 100 Funnel crawl funnel 5 i-iii-2004 Clubionidae Clubiona A 1114 old Funnel crawl Tree14 i-iii-2004 Clubionidae Clubiona A 1 14 old 14Hang 6 i-iii-2004 Clubionidae Clubiona A 1 14 old 14Hang 5 i-iii-2004 Clubionidae Clubiona cycladata 1 14 old Hanging Trap 14Hang 6 i-iii-2004 Clubionidae Clubiona A 1 15 Funnel crawl funnel 9 i-iii-2004 Clubionidae Clubiona A 1 15 100 Flight intercept 15Hang 05 30.iii.2005 Clubionidae Clubiona A 1 15 100 Funnel crawl 11 funnel i-iii-2004 Clubionidae Clubiona A 2 16 old Flight intercept funnel 7 i-iii-2004 Clubionidae Clubiona A 1116 old Funnel crawl funnel 5 30.iii.2005 Clubionidae Clubiona A 1116 old Static trap 11 funnel i-iii-2004 Clubionidae Clubiona cycladata 1 16 old Flight intercept hang trap i-iii-2004 Clubionidae Clubiona cycladata 1116 old Funnel crawl funnel 5 i-iii-2004 Clubionidae Clubiona cycladata 1 16 old Funnel crawl funnel 7 i-iii-2004 Clubionidae Clubiona cycladata 1 16 old Flight intercept Hang trap i-iii-2004 UNKNOWN*** cya ana micro? teeny 1 14 Flight intercept 14Hang 4 loose palp 30.iii.2005 Desidae D3 1 15 100 Funnel crawl funnel 11 30.iii.2005 Thomisidae Diaea 1 7 100 Funnel crawl funnel 11 i-iii-2004 Thomisidae Diaea immature W ACROCER 12 old Flight intercept 12 Hang i-iii-2004 UNKNOWN Dict Hahn Amau? quite squished1 1 15 Flight intercept 15 Hang 4 30.iii.2005 Desidae Dictinoid? 1 7 100 Funnel crawl funnel 11 30.iii.2005 Theridiidae Dipoena 1 14 old Funnel crawl funnel 1 30.iii.2005 Theridiidae Dipoena 1 16 old Funnel crawl funnel 11 i-iii-2004 Sparassidae Eodelena? nr. lapidicola? 1 N/A N/A check list i-iii-2004 Araneidae Eriophora 1 7 100 Funnel crawl funnel 5 30.iii.2005 Theridiidae Euryopis sp A 2 7 100 Funnel crawl funnel 11 i-iii-2004 Theridiidae Euryopis 112 Funnel crawl funnel 7 i-iii.2004 Theridiidae Hadrotarsinae 1 12 Funnel crawl funnel 5 30.iii.2005 Gnaphosoidea Hemicloea? tasmani? 1 Funnel crawl funnel 1 check list 30.iii.2005 Lamponidae Lampona ewens 1 16 old Funnel crawl funnel 11 NEW to LIST 30.iii.2005 Lamponidae Lampona immature sp 2 16 old Funnel crawl funnel 11 i-iii-2004 UNKNOWN Liny? banded legs 1 5 Flight intercept 05hang4 i-iii.2004 Theridiidae? Micropholcomat? flat head scped 1 13 Funnel crawl funnel 10 30.iii.2005 Desidae Namandia 1 10 old Funnel crawl funnel 1 i-iii-2004 Hahniidae Scotopsilis bicolor 1 3 na 03Hang 4 i-iii-2004 Thomisidae Stephanopis lata 1 13 100 Funnel crawl funnel 5 2.iii.2005 Corinnidae Supunna 1 Static trap 72 vane i-iii-2004 Thomisidae Tharraleah? new? 1 13 100 Funnel crawl funnel 5 new to list? i-iii-2004 Hahniidae 1 1 old Funnel crawl funnel 3 i-iii.2004 Theridiidae 11 3 Flight intercept 03Hang4 i-iii-2004 Gnaphosidae 1 4 old Funnel crawl funnel 7 i-iii-2004 Salticidae 1 4 old Funnel crawl funnel 7 i-iii-2004 Salticidae 1 4 old Flight intercept 04Hang4 i-iii-2004 Salticidae 1 6 old Funnel crawl funnel 7 i-iii-2004 Segestriidae 1 6 old Funnel crawl funnel 7 i-iii-2004 Theridiidae 1 6 Flight intercept 06Hang6 i-iii-2004 Salticidae 1 7 100 Funnel crawl funnel 1 30.iii.2005 Hahniidae 2 8 old Funnel crawl funnel 1 i-iii-2004 Salticidae 1 9 100 Flight intercept 09Hang 3 i-iii-2004 Hahniidae 1 10 old Funnel crawl funnel 7 30.iii.2005 Hahniidae 1 10 old Funnel crawl funnel 1 i-iii-2004 Segestriidae 1 10 old Funnel crawl funnel 3 i-iii-2004 Theridiidae 1 10 old Funnel crawl funnel 3 i-iii.2004 Theridiidae small dark 1 11 Funnel crawl funnel 5 i-iii-2004 Linyphiidae 1 13 Sticky tray funnel 11 i-iii-2004 Salticidae 1 13 Funnel crawl funnel10 i-iii-2004 Salticidae 1 13 100 Funnel crawl funnel 5 i-iii-2004 Corinnidae 1 15 100 Funnel crawl funnel 5 i-iii-2004 Desidae 1 15 100 Flight intercept 15Hang 06 30.iii.2005 Linyphiidae 1 15 100 Funnel crawl funnel 11 30.iii.2005 Salticidae 1 15 100 Funnel crawl funnel 11 i-iii-2004 Salticidae 1 15 100 Flight intercept 15Hang 6 i-iii.2004 Theridiidae 1 15 100 Funnel crawl funnel 11 i-iii-2004 Theridiidae 1 15 100 Flight intercept 15Hang 6 30.iii.2005 Linyphiidae 1 16 old Funnel crawl funnel 11 i-iii-2004 Salticidae spp? 1116 Funnel crawl funnel 9 i-iii-2004 Cycloctenidae 1 Funnel crawl funnel 7 2.iii.2005 Stiphidiidae 1 Static trap 68 vane http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :184 Order:Acarina (Mites) Family:Oribatida Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

Morphospecies # :085 Order:Acarina (Mites) Family: Species binomial: Total Collected:12 # in CD StickyTraps:0 # in Funnel Crawl Traps:12 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (1 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :201 Order:Acarina, Oribatida, Mesostigmata (Mites) Family: Species binomial: Total Collected:35 # in CD StickyTraps:3 # in Funnel Crawl Traps:22 # in HangTraps:10

Morphospecies # :005 Order:Acarina? Family: Species binomial: Total Collected:42 # in CD StickyTraps:1 # in Funnel Crawl Traps:36 # in HangTraps:5

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (2 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :038 Order:Arachnida- Araneae Family:Clubionidae Species binomial: Total Collected:107 # in CD StickyTraps:0 # in Funnel Crawl Traps:69 # in HangTraps:38

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (3 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :037 Order:Arachnida- Araneae Family:Salticidae Species binomial: Total Collected:3 # in CD StickyTraps:0 # in Funnel Crawl Traps:3 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (4 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :065 Order:Arachnida- Araneae Family:Sparassidae Species binomial:Delena cancerides Total Collected:3 # in CD StickyTraps:0 # in Funnel Crawl Traps:3 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (5 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :039 Order:Arachnida- Araneae Family:Thomisidae Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (6 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :036 Order:Arachnida- Araneae Family: Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (7 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :100 Order:Arachnida- Araneae Family: Species binomial: Total Collected:206 # in CD StickyTraps:5 # in Funnel Crawl Traps:103 # in HangTraps:98

Morphospecies # :012 Order:Blattodea Family:Blatellidae Species binomial: Total Collected:75 # in CD StickyTraps:1 # in Funnel Crawl Traps:68 # in HangTraps:6

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (8 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :022 Order:Blattodea Family:Blatellidae Species binomial: Total Collected:54 # in CD StickyTraps:0 # in Funnel Crawl Traps:48 # in HangTraps:6

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (9 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :024 Order:Blattodea Family:Blatellidae Species binomial: Total Collected:37 # in CD StickyTraps:0 # in Funnel Crawl Traps:30 # in HangTraps:7

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (10 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :046 Order:Blattodea Family:Blattelidae Species binomial: Total Collected:41 # in CD StickyTraps:0 # in Funnel Crawl Traps:35 # in HangTraps:6

Morphospecies # :057 Order:Blattodea Family:Blattelidae Species binomial: Total Collected:18 # in CD StickyTraps:0 # in Funnel Crawl Traps:17 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (11 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :102 Order:Coleoptera Family:?Silvanidae Species binomial: Total Collected:2 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (12 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :351 Order:Coleoptera Family:Anobiidae Species binomial:Anobiidae TFIC sp 10 Total Collected:5 # in CD StickyTraps:1 # in Funnel Crawl Traps:2 # in HangTraps:2

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (13 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :098 Order:Coleoptera Family:Anobiidae Species binomial:Anobiidae YDB sp 01 Total Collected:49 # in CD StickyTraps:9 # in Funnel Crawl Traps:9 # in HangTraps:31

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (14 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :123 Order:Coleoptera Family:Anobiidae Species binomial:Anobiidae YDB sp 01 Total Collected:6 # in CD StickyTraps:6 # in Funnel Crawl Traps:0 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (15 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :035 Order:Coleoptera Family:Anobiidae Species binomial:Dryophilodes TFIC sp 4 Total Collected:13 # in CD StickyTraps:1 # in Funnel Crawl Traps:5 # in HangTraps:7

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (16 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :352 Order:Coleoptera Family:Anobiidae Species binomial:Hadrobregmus areolicollis Total Collected:8 # in CD StickyTraps:1 # in Funnel Crawl Traps:5 # in HangTraps:2

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (17 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :353 Order:Coleoptera Family:Anobiidae Species binomial:Ptinus exulans Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:0

Morphospecies # :354 Order:Coleoptera Family:Boganiidae Species binomial:Boganium armstrongi Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (18 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :068 Order:Coleoptera Family:Brentidae Species binomial:Apion tasmanicum Total Collected:17 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:16

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (19 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :223 Order:Coleoptera Family:Brentidae Species binomial:Auletobius TFIC sp 1 Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (20 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :117 Order:Coleoptera Family:Buprestidae Species binomial:Cisseis oblonga Total Collected:2 # in CD StickyTraps:1 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (21 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :355 Order:Coleoptera Family:Carabidae Species binomial:Agonocheila ?fenestrata Total Collected:2 # in CD StickyTraps:1 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (22 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :279 Order:Coleoptera Family:Carabidae Species binomial:Demetrida YDB sp 1 Total Collected:2 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (23 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :356 Order:Coleoptera Family:Carabidae Species binomial:Mecyclothorax ambiguus Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (24 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :151 Order:Coleoptera Family:Cerambycidae Species binomial:Ancita crocogaster Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (25 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :078 Order:Coleoptera Family:Cerambycidae Species binomial:Cerambycidae YDB sp 02 Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (26 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :204 Order:Coleoptera Family:Cerambycidae Species binomial:Lygesis ?mendica Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (27 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :357 Order:Coleoptera Family:Cerambycidae Species binomial:Zygocera canosa Total Collected:2 # in CD StickyTraps:0 # in Funnel Crawl Traps:2 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (28 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :139 Order:Coleoptera Family:Cerambycidae Species binomial: Total Collected:2 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:2

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (29 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :359 Order:Coleoptera Family:Cerylonidae Species binomial:Philothermus tasmanicus Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (30 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :033 Order:Coleoptera Family:Chrysomelidae Species binomial:Arsipoda erichsoni Total Collected:7 # in CD StickyTraps:2 # in Funnel Crawl Traps:2 # in HangTraps:3

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (31 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :027 Order:Coleoptera Family:Chrysomelidae Species binomial:Chrysophtharta bimaculata Total Collected:22 # in CD StickyTraps:1 # in Funnel Crawl Traps:13 # in HangTraps:8

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (32 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :014 Order:Coleoptera Family:Chrysomelidae Species binomial:Chrysophtharta lignea Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:0

Morphospecies # :207 Order:Coleoptera Family:Chrysomelidae Species binomial:Monolepta TFIC sp 02 Total Collected:3 # in CD StickyTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (33 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm # in Funnel Crawl Traps:0 # in HangTraps:3

Morphospecies # :248 Order:Coleoptera Family:Chrysomelidae Species binomial:Monolepta TFIC sp. 02 Total Collected:3 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:2

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (34 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :115 Order:Coleoptera Family:Chrysomelidae Species binomial:Paropsis rupidipes Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (35 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :015 Order:Coleoptera Family:Chrysomelidae Species binomial:Sterromela subcostata Total Collected:7 # in CD StickyTraps:0 # in Funnel Crawl Traps:7 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (36 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :043 Order:Coleoptera Family:Chrysomelidae Species binomial:Trachymela YDB sp 01 Total Collected:2 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (37 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :397 Order:Coleoptera Family:Chrysomelidae larvae Species binomial: Total Collected:1 # in CD StickyTraps:1 # in Funnel Crawl Traps:0 # in HangTraps:0

Morphospecies # :209 Order:Coleoptera Family:Clambidae Species binomial:Sphaerothorax tasmani Total Collected:2 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:2

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (38 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :298 Order:Coleoptera Family:Cleridae Species binomial:Cleridae YDB 01 Total Collected:4 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:3

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (39 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :103 Order:Coleoptera Family:Cleridae Species binomial:Lemidia pallida Total Collected:4 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:3

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (40 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :034 Order:Coleoptera Family:Cleridae Species binomial:Neoscrobiger patricius Total Collected:6 # in CD StickyTraps:0 # in Funnel Crawl Traps:3 # in HangTraps:3

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (41 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :152 Order:Coleoptera Family:Cleridae Species binomial:Parapylus bicinctus Total Collected:2 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:2

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (42 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :028 Order:Coleoptera Family:Coccinelidae Species binomial:Cleobora mellyi Total Collected:3 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:2

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (43 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :017 Order:Coleoptera Family:Coccinelidae Species binomial:Rodatus TFIC sp. 01 Total Collected:27 # in CD StickyTraps:0 # in Funnel Crawl Traps:27 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (44 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :089 Order:Coleoptera Family:Coccinellidae Species binomial:Rhyzobius TFIC sp 05 Total Collected:6 # in CD StickyTraps:0 # in Funnel Crawl Traps:4 # in HangTraps:2

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (45 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :031 Order:Coleoptera Family:Coccinellidae Species binomial:Rhyzobius TFIC sp 14 Total Collected:6 # in CD StickyTraps:0 # in Funnel Crawl Traps:6 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (46 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :360 Order:Coleoptera Family:Coccinellidae Species binomial:Rhyzobius TFIC sp 20 Total Collected:2 # in CD StickyTraps:1 # in Funnel Crawl Traps:1 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (47 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :361 Order:Coleoptera Family:Cryptophagidae Species binomial:Cryptophagus tasmanicus Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (48 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :362 Order:Coleoptera Family:Curculionidae Species binomial:Aades cultratus Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (49 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :227 Order:Coleoptera Family:Curculionidae Species binomial:Acacicis abundans Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (50 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :380 Order:Coleoptera Family:Curculionidae Species binomial:Cryptorhyncinae TFIC sp 11 Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (51 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :363 Order:Coleoptera Family:Curculionidae Species binomial:Curculionidae YDB sp 03 Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (52 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :364 Order:Coleoptera Family:Curculionidae Species binomial:Curculionidae YDB sp 05 Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (53 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :294 Order:Coleoptera Family:Curculionidae Species binomial:Curculionidae YDB sp 06 Total Collected:2 # in CD StickyTraps:0 # in Funnel Crawl Traps:2 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (54 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :365 Order:Coleoptera Family:Curculionidae Species binomial:Curculionidae YDB sp 07 Total Collected:2 # in CD StickyTraps:0 # in Funnel Crawl Traps:2 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (55 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :366 Order:Coleoptera Family:Curculionidae Species binomial:Curculionidae YDB sp 11 Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (56 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :367 Order:Coleoptera Family:Curculionidae Species binomial:Curculionidae YDB sp UNID Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:0

Morphospecies # :112 Order:Coleoptera Family:Curculionidae Species binomial:Elleschus wellingtoniensis Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (57 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :249 Order:Coleoptera Family:Curculionidae Species binomial:Emplesis TFIC sp 01 Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (58 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :083 Order:Coleoptera Family:Curculionidae Species binomial:Exithius capucinus Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (59 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :165 Order:Coleoptera Family:Curculionidae Species binomial:Orthorhinus TFIC sp 1 Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (60 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :293 Order:Coleoptera Family:Curculionidae Species binomial:Orthorhinus TFIC sp 2 Total Collected:3 # in CD StickyTraps:0 # in Funnel Crawl Traps:3 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (61 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :162 Order:Coleoptera Family:Curculionidae Species binomial:Pentamimus australis Total Collected:6 # in CD StickyTraps:2 # in Funnel Crawl Traps:0 # in HangTraps:4

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (62 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :305 Order:Coleoptera Family:Curculionidae Species binomial:Pentarthrum? TFIC sp 03 Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (63 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :358 Order:Coleoptera Family:Curculionidae Species binomial:Tychiinae TFIC sp 8 Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (64 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :289 Order:Coleoptera Family:Dermestidae Species binomial:Trogoderma YDB sp 1 Total Collected:3 # in CD StickyTraps:0 # in Funnel Crawl Traps:3 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (65 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :307 Order:Coleoptera Family:Dermestidae Species binomial:Trogoderma YDB sp 2 Total Collected:3 # in CD StickyTraps:2 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (66 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :255 Order:Coleoptera Family:Elateridae Species binomial:Augenotus quadriguttatus Total Collected:2 # in CD StickyTraps:0 # in Funnel Crawl Traps:2 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (67 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :291 Order:Coleoptera Family:Elateridae Species binomial:Conoderus australasiae Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (68 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :163 Order:Coleoptera Family:Elateridae Species binomial:Denticollinae YDB sp 4 Total Collected:13 # in CD StickyTraps:0 # in Funnel Crawl Traps:3 # in HangTraps:10

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (69 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :368 Order:Coleoptera Family:Elateridae Species binomial:Elateridae YDB sp 1 Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (70 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :299 Order:Coleoptera Family:Elateridae Species binomial:Elateridae YDB sp 2 Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (71 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :369 Order:Coleoptera Family:Elateridae Species binomial:Elateridae YDB sp UNID Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:0

Morphospecies # :230 Order:Coleoptera Family:Erotylidae Species binomial:Erotylidae KH 1 Total Collected:6 # in CD StickyTraps:2 # in Funnel Crawl Traps:1 # in HangTraps:3

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (72 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :130 Order:Coleoptera Family:Erotylidae Species binomial:Thallis vinula Total Collected:5 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:5

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (73 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :191 Order:Coleoptera Family:Laemophloeidae Species binomial:Laemophloeidae TFIC sp 01 Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (74 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :258 Order:Coleoptera Family:larvae Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (75 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :280 Order:Coleoptera Family:larvae Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (76 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :381 Order:Coleoptera Family:Latridiidae Species binomial:Cortinicara TFIC sp 02 Total Collected:7 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:7

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (77 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :382 Order:Coleoptera Family:Latridiidae Species binomial:Latridiidae YDB sp. 01 Total Collected:2 # in CD StickyTraps:1 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (78 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :228 Order:Coleoptera Family:Latriidae Species binomial:Aridus costatus Total Collected:4 # in CD StickyTraps:3 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (79 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :260 Order:Coleoptera Family:Lymexylidae Species binomial:Atractocerus YDB sp 1 Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (80 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :262 Order:Coleoptera Family:Lymexylidae Species binomial:Atractocerus YDB sp. 01 Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (81 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :210 Order:Coleoptera Family:Melandryidae Species binomial:Melandryidae TFIC sp 03 Total Collected:2 # in CD StickyTraps:1 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (82 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :241 Order:Coleoptera Family:Melandryidae Species binomial:Melandryidae TFIC sp 04 Total Collected:5 # in CD StickyTraps:4 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (83 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :180 Order:Coleoptera Family:Melandryidae Species binomial:Orchesia ?eucalyptii Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (84 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :296 Order:Coleoptera Family:Melandryidae Species binomial:Orchesia ?eucalyptii Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (85 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :370 Order:Coleoptera Family:Melandryidae Species binomial:Orchesia alphabetica Total Collected:2 # in CD StickyTraps:1 # in Funnel Crawl Traps:1 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (86 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :371 Order:Coleoptera Family:Melyridae Species binomial:Dasytes TFIC sp 02 Total Collected:2 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:2

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (87 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :372 Order:Coleoptera Family:Mordellidae Species binomial:Mordella promiscua Total Collected:2 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:2

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (88 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :159 Order:Coleoptera Family:Mordellidae Species binomial:Mordellidae TFIC sp 05 Total Collected:4 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:4

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (89 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :142 Order:Coleoptera Family:Mycetophagidae Species binomial:Litargus intricatus Total Collected:2 # in CD StickyTraps:1 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (90 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :077 Order:Coleoptera Family:Mycteridae Species binomial:Trichosalpingus TFIC sp 01 Total Collected:12 # in CD StickyTraps:2 # in Funnel Crawl Traps:7 # in HangTraps:3

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (91 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :131 Order:Coleoptera Family:Nitidulidae Species binomial:Thalycrodes australe Total Collected:2 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:2

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (92 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :373 Order:Coleoptera Family:Oedemeridae Species binomial:Dohrnia miranda Total Collected:18 # in CD StickyTraps:0 # in Funnel Crawl Traps:17 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (93 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :013 Order:Coleoptera Family:Oedomeridae Species binomial:Dorhnia simplex Total Collected:43 # in CD StickyTraps:0 # in Funnel Crawl Traps:36 # in HangTraps:7

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (94 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :079 Order:Coleoptera Family:Phalacridae Species binomial:Phalacridae TFIC sp 01 Total Collected:20 # in CD StickyTraps:1 # in Funnel Crawl Traps:2 # in HangTraps:17

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (95 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :202 Order:Coleoptera Family:Prostomidae Species binomial:Dryocora cephalotes Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (96 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :398 Order:Coleoptera Family:Pselaphidae Species binomial: Total Collected:5 # in CD StickyTraps:5 # in Funnel Crawl Traps:0 # in HangTraps:0

Morphospecies # :174 Order:Coleoptera Family:Pyrochroidae Species binomial:Binburrum ?ruficollis Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (97 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :229 Order:Coleoptera Family:Salpingidae Species binomial:Orphanotrophium frigidum Total Collected:24 # in CD StickyTraps:17 # in Funnel Crawl Traps:3 # in HangTraps:4

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (98 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :173 Order:Coleoptera Family:Scarabaeidae Species binomial:Heteronyx tasmanicus Total Collected:2 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:1

Morphospecies # :277 Order:Coleoptera Family:Scarabaeidae Species binomial:Phyllochlaenia villosus Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (99 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :125 Order:Coleoptera Family:Scirtidae Species binomial:Pryonocyphon TFIC sp 01 Total Collected:10 # in CD StickyTraps:3 # in Funnel Crawl Traps:2 # in HangTraps:5

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (100 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :108 Order:Coleoptera Family:Scirtidae Species binomial:Scirtidae TFIC sp 04 Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (101 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :061 Order:Coleoptera Family:Scraptiidae Species binomial:Scraptia laticollis Total Collected:4 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:3

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (102 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :127 Order:Coleoptera Family:Scydmaenidae Species binomial:Euconnus sp. 1 Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

Morphospecies # :374 Order:Coleoptera Family:Silvanidae Species binomial:Cryptamorpha victoriae Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (103 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :375 Order:Coleoptera Family:Staphylinidae Species binomial:Pselaphinae YDB sp 01 Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (104 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :379 Order:Coleoptera Family:Staphylinidae Species binomial:Sepedophilus TFIC sp 01 Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (105 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :376 Order:Coleoptera Family:Staphylinidae Species binomial:Staphylinidae YDB sp 03 Total Collected:2 # in CD StickyTraps:0 # in Funnel Crawl Traps:2 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (106 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :377 Order:Coleoptera Family:Staphylinidae Species binomial:Staphylinidae YDB sp 04 Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (107 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :378 Order:Coleoptera Family:Staphylinidae Species binomial:Staphylinidae YDB sp 05 Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (108 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :290 Order:Coleoptera Family:Staphylinidae Species binomial:Staphylinidae YDB sp 06 Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (109 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :236 Order:Coleoptera Family:Staphylinidae Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (110 of 290) [12/01/2006 5:02:01 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :126 Order:Coleoptera Family:Tenebrionidae Species binomial:Appelatus tasmanicus Total Collected:3 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:3

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (111 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :016 Order:Coleoptera Family:Throscidae Species binomial:Aulonothroscus elongatus Total Collected:259 # in CD StickyTraps:31 # in Funnel Crawl Traps:166 # in HangTraps:62

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (112 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :124 Order:Coleoptera Family:Trogossitidae Species binomial:Egolia variegata Total Collected:2 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:2

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (113 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :129 Order:Coleoptera Family:Trogossitidae Species binomial:Trogossitidae TFIC sp 01 Total Collected:15 # in CD StickyTraps:14 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (114 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :282 Order:Coleoptera Family:Trogossitidae Species binomial:Trogossitidae TFIC sp 01 Total Collected:26 # in CD StickyTraps:0 # in Funnel Crawl Traps:8 # in HangTraps:18

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (115 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :128 Order:Coleoptera Family:Trogossitidae Species binomial:Trogossitidae? Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (116 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :405 Order:Coleoptera Family:Trogossitidae? Species binomial: Total Collected:1 # in CD StickyTraps:1 # in Funnel Crawl Traps:0 # in HangTraps:0

Morphospecies # :270 Order:Coleoptera Family:Zopheridae Species binomial:Penthelispa fuliginosa Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (117 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :086 Order:Collembola Family:Paronellidae Species binomial: Total Collected:15 # in CD StickyTraps:0 # in Funnel Crawl Traps:9 # in HangTraps:6

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (118 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :188 Order:Dermaptera Family:Spongophoridae Species binomial: Total Collected:3 # in CD StickyTraps:0 # in Funnel Crawl Traps:3 # in HangTraps:0

Morphospecies # :256 Order:Diptera

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (119 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm Family:Asilidae Species binomial: Total Collected:3 # in CD StickyTraps:1 # in Funnel Crawl Traps:1 # in HangTraps:1

Morphospecies # :189 Order:Diptera Family:Calliphoridae Species binomial:Calliphora Total Collected:12 # in CD StickyTraps:0 # in Funnel Crawl Traps:6 # in HangTraps:6

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (120 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :059 Order:Diptera Family:Chloropidae Species binomial: Total Collected:8 # in CD StickyTraps:0 # in Funnel Crawl Traps:4 # in HangTraps:4

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (121 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :395 Order:Diptera Family:Empididiae Species binomial: Total Collected:6 # in CD StickyTraps:6 # in Funnel Crawl Traps:0 # in HangTraps:0

Morphospecies # :252 Order:Diptera Family:Ironomyidae Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (122 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :181 Order:Diptera Family:Lauxiniidae Species binomial: Total Collected:7 # in CD StickyTraps:6 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (123 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :111 Order:Diptera Family:Muscidae Species binomial: Total Collected:13 # in CD StickyTraps:6 # in Funnel Crawl Traps:0 # in HangTraps:7

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (124 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :148 Order:Diptera Family:Mycetophilidae Species binomial: Total Collected:3 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:3

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (125 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :157 Order:Diptera Family:Mycetophilidae Species binomial: Total Collected:35 # in CD StickyTraps:31 # in Funnel Crawl Traps:1 # in HangTraps:3

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (126 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :169 Order:Diptera Family:Mycetophilidae Species binomial: Total Collected:28 # in CD StickyTraps:0 # in Funnel Crawl Traps:16 # in HangTraps:12

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (127 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :219 Order:Diptera Family:Mycetophilidae Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (128 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :266 Order:Diptera Family:Mycetophilidae Species binomial: Total Collected:4 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:4

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (129 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :143 Order:Diptera Family:Mycetophilidae? Species binomial: Total Collected:2 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:2

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (130 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :158 Order:Diptera Family:Phoridae Species binomial: Total Collected:29 # in CD StickyTraps:24 # in Funnel Crawl Traps:0 # in HangTraps:5

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (131 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :160 Order:Diptera Family:Phoridae Species binomial: Total Collected:3 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:3

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (132 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :176 Order:Diptera Family:Phoridae Species binomial: Total Collected:194 # in CD StickyTraps:32 # in Funnel Crawl Traps:22 # in HangTraps:140

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (133 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :193 Order:Diptera Family:Phoridae Species binomial: Total Collected:9 # in CD StickyTraps:2 # in Funnel Crawl Traps:0 # in HangTraps:7

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (134 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :194 Order:Diptera Family:Phoridae Species binomial: Total Collected:8 # in CD StickyTraps:7 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (135 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :197 Order:Diptera Family:Psychodidae Species binomial: Total Collected:6 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:6

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (136 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :208 Order:Diptera Family:Psychodidae Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (137 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :026 Order:Diptera Family:Sciaridae Species binomial: Total Collected:1082 # in CD StickyTraps:34 # in Funnel Crawl Traps:325 # in HangTraps:723

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (138 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :051 Order:Diptera Family:Sciaridae Species binomial: Total Collected:80 # in CD StickyTraps:47 # in Funnel Crawl Traps:28 # in HangTraps:5

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (139 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :054 Order:Diptera Family:Sciaridae Species binomial: Total Collected:153 # in CD StickyTraps:127 # in Funnel Crawl Traps:15 # in HangTraps:11

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (140 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :137 Order:Diptera Family:Sciaridae Species binomial: Total Collected:78 # in CD StickyTraps:71 # in Funnel Crawl Traps:1 # in HangTraps:6

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (141 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :138 Order:Diptera Family:Sciaridae Species binomial: Total Collected:242 # in CD StickyTraps:67 # in Funnel Crawl Traps:92 # in HangTraps:83

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (142 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :244 Order:Diptera Family:Sciaridae Species binomial: Total Collected:17 # in CD StickyTraps:12 # in Funnel Crawl Traps:0 # in HangTraps:5

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (143 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :254 Order:Diptera Family:Sciaridae Species binomial: Total Collected:2 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (144 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :281 Order:Diptera Family:Sciaridae Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (145 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :386 Order:Diptera Family:Sciaridae Species binomial: Total Collected:1 # in CD StickyTraps:1 # in Funnel Crawl Traps:0 # in HangTraps:0

Morphospecies # :394 Order:Diptera Family:Sciaridae Species binomial: Total Collected:1 # in CD StickyTraps:1 # in Funnel Crawl Traps:0 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (146 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :402 Order:Diptera Family:Sciaridae Species binomial: Total Collected:2 # in CD StickyTraps:2 # in Funnel Crawl Traps:0 # in HangTraps:0

Morphospecies # :167 Order:Diptera Family:Tabanidae Species binomial: Total Collected:4 # in CD StickyTraps:0 # in Funnel Crawl Traps:3 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (147 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm Morphospecies # :080 Order:Diptera Family:Tachinidae? Species binomial: Total Collected:16 # in CD StickyTraps:3 # in Funnel Crawl Traps:5 # in HangTraps:8

Morphospecies # :044 Order:Diptera Family:Tipulidae Species binomial: Total Collected:94 # in CD StickyTraps:0 # in Funnel Crawl Traps:55 # in HangTraps:39

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (148 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :063 Order:Diptera Family:Tipulidae Species binomial: Total Collected:72 # in CD StickyTraps:5 # in Funnel Crawl Traps:39 # in HangTraps:28

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (149 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :088 Order:Diptera Family:Tipulidae Species binomial: Total Collected:35 # in CD StickyTraps:1 # in Funnel Crawl Traps:2 # in HangTraps:32

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (150 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :178 Order:Diptera Family:Tipulidae Species binomial: Total Collected:8 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:8

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (151 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :234 Order:Diptera Family:Window Fly? Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (152 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :070 Order:Diptera Family: Species binomial: Total Collected:668 # in CD StickyTraps:238 # in Funnel Crawl Traps:173 # in HangTraps:257

Morphospecies # :105 Order:Diptera Family: Species binomial: Total Collected:2 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:2

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (153 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :134 Order:Diptera Family: Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

Morphospecies # :196 Order:Diptera Family: Species binomial: Total Collected:16 # in CD StickyTraps:12 # in Funnel Crawl Traps:1 # in HangTraps:3

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (154 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :198 Order:Diptera Family: Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

Morphospecies # :211 Order:Diptera Family: Species binomial: Total Collected:2 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:2

Morphospecies # :212

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (155 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm Order:Diptera Family: Species binomial: Total Collected:10 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:10

Morphospecies # :221 Order:Diptera Family: Species binomial: Total Collected:4 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:4

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (156 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :231 Order:Diptera Family: Species binomial: Total Collected:3 # in CD StickyTraps:2 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (157 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :253 Order:Diptera Family: Species binomial: Total Collected:2 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (158 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :268 Order:Diptera Family: Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (159 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :384 Order:Diptera Family: Species binomial: Total Collected:1 # in CD StickyTraps:1 # in Funnel Crawl Traps:0 # in HangTraps:0

Morphospecies # :391 Order:Diptera Family: Species binomial: Total Collected:1 # in CD StickyTraps:1 # in Funnel Crawl Traps:0 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (160 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :393 Order:Diptera Family: Species binomial: Total Collected:80 # in CD StickyTraps:80 # in Funnel Crawl Traps:0 # in HangTraps:0

Morphospecies # :396 Order:Diptera Family: Species binomial: Total Collected:11 # in CD StickyTraps:11 # in Funnel Crawl Traps:0 # in HangTraps:0

Morphospecies # :238 Order:Diptera? Family:Muscidae Species binomial: Total Collected:5 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:5

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (161 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :232 Order:Diptera? Family:Mycetophilidae Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (162 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :200 Order:Fragments, UNID Family: Species binomial: Total Collected:110 # in CD StickyTraps:12 # in Funnel Crawl Traps:19 # in HangTraps:79

Morphospecies # :297 Order:Hemiptera Family:Acanthosomatidae Species binomial: Total Collected:2 # in CD StickyTraps:0 # in Funnel Crawl Traps:2 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (163 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :019 Order:Hemiptera Family:Cicadellidae Species binomial: Total Collected:110 # in CD StickyTraps:0 # in Funnel Crawl Traps:104 # in HangTraps:6

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (164 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :021 Order:Hemiptera Family:Cicadellidae Species binomial: Total Collected:21 # in CD StickyTraps:2 # in Funnel Crawl Traps:18 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (165 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :049 Order:Hemiptera Family:Cicadellidae Species binomial: Total Collected:40 # in CD StickyTraps:1 # in Funnel Crawl Traps:36 # in HangTraps:3

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (166 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :058 Order:Hemiptera Family:Cicadellidae Species binomial: Total Collected:9 # in CD StickyTraps:0 # in Funnel Crawl Traps:9 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (167 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :113 Order:Hemiptera Family:Cicadellidae Species binomial: Total Collected:11 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:10

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (168 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :149 Order:Hemiptera Family:Cicadellidae Species binomial: Total Collected:6 # in CD StickyTraps:0 # in Funnel Crawl Traps:3 # in HangTraps:3

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (169 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :217 Order:Hemiptera Family:Cicadellidae Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (170 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :389 Order:Hemiptera Family:Cicadellidae Species binomial: Total Collected:1 # in CD StickyTraps:1 # in Funnel Crawl Traps:0 # in HangTraps:0

Morphospecies # :023 Order:Hemiptera Family:Cicadellidae? Species binomial: Total Collected:8 # in CD StickyTraps:1 # in Funnel Crawl Traps:3 # in HangTraps:4

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (171 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :106 Order:Hemiptera Family:Cixiidae Species binomial: Total Collected:13 # in CD StickyTraps:1 # in Funnel Crawl Traps:5 # in HangTraps:7

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (172 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm Morphospecies # :302 Order:Hemiptera Family:Coccidae? Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:0

Morphospecies # :121 Order:Hemiptera Family:Enicocephalidae Species binomial: Total Collected:3 # in CD StickyTraps:1 # in Funnel Crawl Traps:0 # in HangTraps:2

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (173 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :018 Order:Hemiptera Family:Eurymelidae Species binomial:EurymelaYDB sp. 01 Total Collected:3 # in CD StickyTraps:1 # in Funnel Crawl Traps:2 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (174 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :099 Order:Hemiptera Family:Eurymelidae Species binomial: Total Collected:36 # in CD StickyTraps:0 # in Funnel Crawl Traps:29 # in HangTraps:7

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (175 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :192 Order:Hemiptera Family:Eurymelidae Species binomial: Total Collected:5 # in CD StickyTraps:2 # in Funnel Crawl Traps:2 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (176 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :006 Order:Hemiptera Family:Flatidae Species binomial: Total Collected:58 # in CD StickyTraps:0 # in Funnel Crawl Traps:56 # in HangTraps:2

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (177 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :010 Order:Hemiptera Family:Flatidae Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (178 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :066 Order:Hemiptera Family:Fulgoridae Species binomial: Total Collected:51 # in CD StickyTraps:28 # in Funnel Crawl Traps:14 # in HangTraps:9

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (179 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :082 Order:Hemiptera Family:Fulgoridae Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (180 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :120 Order:Hemiptera Family:Fulgoridae Species binomial: Total Collected:18 # in CD StickyTraps:12 # in Funnel Crawl Traps:0 # in HangTraps:6

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (181 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :144 Order:Hemiptera Family:Fulgoridae Species binomial: Total Collected:13 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:13

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (182 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :045 Order:Hemiptera Family:Lygaeidae Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (183 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :029 Order:Hemiptera Family:Lygaeidae? Species binomial: Total Collected:6 # in CD StickyTraps:1 # in Funnel Crawl Traps:4 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (184 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :147 Order:Hemiptera Family:Lygaeidae? Species binomial: Total Collected:2 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:2

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (185 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :216 Order:Hemiptera Family:Membracidae Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (186 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :092 Order:Hemiptera Family:Miridae Species binomial:Sejanus YDB sp. 01 Total Collected:2 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (187 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :047 Order:Hemiptera Family:Miridae Species binomial: Total Collected:5 # in CD StickyTraps:0 # in Funnel Crawl Traps:4 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (188 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :091 Order:Hemiptera Family:Miridae Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (189 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :166 Order:Hemiptera Family:Miridae Species binomial: Total Collected:2 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:2

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (190 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :185 Order:Hemiptera Family:Miridae Species binomial: Total Collected:4 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:3

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (191 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :276 Order:Hemiptera Family:Miridae Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (192 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :183 Order:Hemiptera Family:Miridae? Species binomial: Total Collected:18 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:18

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (193 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :003 Order:Hemiptera Family:Pentatomidae Species binomial: Total Collected:16 # in CD StickyTraps:0 # in Funnel Crawl Traps:16 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (194 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :011 Order:Hemiptera Family:Pentatomidae Species binomial: Total Collected:106 # in CD StickyTraps:0 # in Funnel Crawl Traps:104 # in HangTraps:2

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (195 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :030 Order:Hemiptera Family:Pentatomidae Species binomial: Total Collected:47 # in CD StickyTraps:1 # in Funnel Crawl Traps:42 # in HangTraps:4

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (196 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :168 Order:Hemiptera Family:Pentatomidae Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (197 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :094 Order:Hemiptera Family:Psyllidae Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (198 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :140 Order:Hemiptera Family:Psyllidae Species binomial: Total Collected:10 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:10

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (199 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :171 Order:Hemiptera Family:Psyllidae Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (200 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :235 Order:Hemiptera Family:Psyllidae Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (201 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :263 Order:Hemiptera Family:Psyllidae Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (202 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :400 Order:Hemiptera Family:Psyllidae Species binomial: Total Collected:1 # in CD StickyTraps:1 # in Funnel Crawl Traps:0 # in HangTraps:0

Morphospecies # :154 Order:Hemiptera Family:Psyllidae juvenile Species binomial: Total Collected:2 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:2

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (203 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :237 Order:Hemiptera Family: Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (204 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :286 Order:Hemiptera Family: Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (205 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :292 Order:Hemiptera Family: Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (206 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :155 Order:Hymenoptera Family:Braconidae Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (207 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :177 Order:Hymenoptera Family:Braconidae Species binomial: Total Collected:24 # in CD StickyTraps:18 # in Funnel Crawl Traps:0 # in HangTraps:6

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (208 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :401 Order:Hymenoptera Family:Braconidae Species binomial: Total Collected:1 # in CD StickyTraps:1 # in Funnel Crawl Traps:0 # in HangTraps:0

Morphospecies # :052 Order:Hymenoptera Family:Chalcidoid? Species binomial: Total Collected:60 # in CD StickyTraps:28 # in Funnel Crawl Traps:25 # in HangTraps:7

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (209 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :069 Order:Hymenoptera Family:Evaniidae Species binomial: Total Collected:17 # in CD StickyTraps:0 # in Funnel Crawl Traps:12 # in HangTraps:5

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (210 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :172 Order:Hymenoptera Family:Firmicidae Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (211 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :074 Order:Hymenoptera Family:Formicidae Species binomial:Camponotus YDB sp. 01 Total Collected:7 # in CD StickyTraps:0 # in Funnel Crawl Traps:5 # in HangTraps:2

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (212 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :075 Order:Hymenoptera Family:Formicidae Species binomial:Camponotus YDB sp. 02 Total Collected:13 # in CD StickyTraps:0 # in Funnel Crawl Traps:6 # in HangTraps:7

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (213 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :071 Order:Hymenoptera Family:Formicidae Species binomial: Total Collected:26 # in CD StickyTraps:0 # in Funnel Crawl Traps:7 # in HangTraps:19

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (214 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :073 Order:Hymenoptera Family:Formicidae Species binomial: Total Collected:5 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:4

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (215 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :205 Order:Hymenoptera Family:Formicidae Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (216 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :224 Order:Hymenoptera Family:Formicidae Species binomial: Total Collected:3 # in CD StickyTraps:1 # in Funnel Crawl Traps:1 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (217 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :269 Order:Hymenoptera Family:Formicidae Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (218 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :273 Order:Hymenoptera Family:Formicidae Species binomial: Total Collected:17 # in CD StickyTraps:16 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (219 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :156 Order:Hymenoptera Family:Gasteruptionidae Species binomial: Total Collected:14 # in CD StickyTraps:2 # in Funnel Crawl Traps:0 # in HangTraps:12

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (220 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :002 Order:Hymenoptera Family:Ichneumonidae Species binomial: Total Collected:9 # in CD StickyTraps:4 # in Funnel Crawl Traps:5 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (221 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :008 Order:Hymenoptera Family:Ichneumonidae Species binomial: Total Collected:3 # in CD StickyTraps:0 # in Funnel Crawl Traps:2 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (222 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :050 Order:Hymenoptera Family:Ichneumonidae Species binomial: Total Collected:6 # in CD StickyTraps:4 # in Funnel Crawl Traps:2 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (223 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :214 Order:Hymenoptera Family:Ichneumonidae Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (224 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :285 Order:Hymenoptera Family:Ichneumonidae Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (225 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :072 Order:Hymenoptera Family:Microwasp Species binomial: Total Collected:7 # in CD StickyTraps:4 # in Funnel Crawl Traps:2 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (226 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :182 Order:Hymenoptera Family:Microwasp Species binomial: Total Collected:10 # in CD StickyTraps:4 # in Funnel Crawl Traps:1 # in HangTraps:5

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (227 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :222 Order:Hymenoptera Family:Microwasp Species binomial: Total Collected:3 # in CD StickyTraps:1 # in Funnel Crawl Traps:0 # in HangTraps:2

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (228 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :250 Order:Hymenoptera Family:Microwasp Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (229 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :257 Order:Hymenoptera Family:Microwasp Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (230 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :271 Order:Hymenoptera Family:Microwasp Species binomial: Total Collected:3 # in CD StickyTraps:0 # in Funnel Crawl Traps:2 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (231 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :383 Order:Hymenoptera Family:Microwasp Species binomial: Total Collected:1 # in CD StickyTraps:1 # in Funnel Crawl Traps:0 # in HangTraps:0

Morphospecies # :403 Order:Hymenoptera Family:Microwasp Species binomial: Total Collected:2 # in CD StickyTraps:2 # in Funnel Crawl Traps:0 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (232 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :404 Order:Hymenoptera Family:Microwasp Species binomial: Total Collected:6 # in CD StickyTraps:6 # in Funnel Crawl Traps:0 # in HangTraps:0

Morphospecies # :406 Order:Hymenoptera Family:Microwasp Species binomial: Total Collected:1 # in CD StickyTraps:1 # in Funnel Crawl Traps:0 # in HangTraps:0

Morphospecies # :056 Order:Hymenoptera Family:MicroWasp? Species binomial: Total Collected:18 # in CD StickyTraps:5 # in Funnel Crawl Traps:13 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (233 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :133 Order:Hymenoptera Family:MicroWasp? Species binomial: Total Collected:2 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:2

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (234 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :136 Order:Hymenoptera Family:MicroWasp? Species binomial: Total Collected:26 # in CD StickyTraps:18 # in Funnel Crawl Traps:2 # in HangTraps:6

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (235 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :164 Order:Hymenoptera Family:MicroWasp? Species binomial: Total Collected:3 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:3

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (236 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :179 Order:Hymenoptera Family:MicroWasp? Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (237 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :187 Order:Hymenoptera Family:MicroWasp? Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (238 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :225 Order:Hymenoptera Family:MicroWasp? Species binomial: Total Collected:16 # in CD StickyTraps:4 # in Funnel Crawl Traps:10 # in HangTraps:2

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (239 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :226 Order:Hymenoptera Family:MicroWasp? Species binomial: Total Collected:5 # in CD StickyTraps:2 # in Funnel Crawl Traps:1 # in HangTraps:2

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (240 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :009 Order:Hymenoptera Family:Pompilidae Species binomial: Total Collected:40 # in CD StickyTraps:5 # in Funnel Crawl Traps:28 # in HangTraps:7

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (241 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :055 Order:Hymenoptera Family:Proctotrupoidae Species binomial: Total Collected:17 # in CD StickyTraps:9 # in Funnel Crawl Traps:7 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (242 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :119 Order:Hymenoptera Family:Stinging Wasp? Species binomial: Total Collected:46 # in CD StickyTraps:39 # in Funnel Crawl Traps:4 # in HangTraps:3

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (243 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :007 Order:Hymenoptera Family:Vespidae Species binomial:Vespula germanica Total Collected:3 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:2

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (244 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :060 Order:Hymenoptera Family:Wingless MicroWasp? Species binomial: Total Collected:3 # in CD StickyTraps:0 # in Funnel Crawl Traps:3 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (245 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :053 Order:Hymenoptera Family: Species binomial: Total Collected:17 # in CD StickyTraps:1 # in Funnel Crawl Traps:13 # in HangTraps:3

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (246 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :090 Order:Hymenoptera Family: Species binomial: Total Collected:9 # in CD StickyTraps:2 # in Funnel Crawl Traps:3 # in HangTraps:4

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (247 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :093 Order:Hymenoptera Family: Species binomial: Total Collected:8 # in CD StickyTraps:1 # in Funnel Crawl Traps:3 # in HangTraps:4

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (248 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :199 Order:Hymenoptera Family: Species binomial: Total Collected:3 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:3

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (249 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :251 Order:Hymenoptera Family: Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (250 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :275 Order:Hymenoptera Family: Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (251 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :287 Order:Hymenoptera Family: Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (252 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :385 Order:Hymenoptera Family: Species binomial: Total Collected:1 # in CD StickyTraps:1 # in Funnel Crawl Traps:0 # in HangTraps:0

Morphospecies # :392 Order:Hymenoptera Family: Species binomial: Total Collected:1 # in CD StickyTraps:1 # in Funnel Crawl Traps:0 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (253 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :190 Order:Hymenoptera? Family:Formicidae Species binomial: Total Collected:5 # in CD StickyTraps:0 # in Funnel Crawl Traps:4 # in HangTraps:1

Morphospecies # :101 Order:Hymenoptera- Ant Family: Species binomial: Total Collected:181 # in CD StickyTraps:4 # in Funnel Crawl Traps:166 # in HangTraps:11

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (254 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm Morphospecies # :064 Order:Isoptera Family: Species binomial: Total Collected:27 # in CD StickyTraps:3 # in Funnel Crawl Traps:11 # in HangTraps:13

Morphospecies # :390 Order:Isoptera Family: Species binomial: Total Collected:19 # in CD StickyTraps:19 # in Funnel Crawl Traps:0 # in HangTraps:0

Morphospecies # :301

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (255 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm Order:Larvae UNID Family: Species binomial: Total Collected:13 # in CD StickyTraps:0 # in Funnel Crawl Traps:13 # in HangTraps:0

Morphospecies # :041 Order:Lepidoptera Family:Cosmopterygidae Species binomial: Total Collected:15 # in CD StickyTraps:0 # in Funnel Crawl Traps:10 # in HangTraps:5

Morphospecies # :150 Order:Lepidoptera Family:Cosmopterygidae

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (256 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm Species binomial: Total Collected:41 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:41

Morphospecies # :040 Order:Lepidoptera Family:Geometidae Species binomial: Total Collected:80 # in CD StickyTraps:3 # in Funnel Crawl Traps:67 # in HangTraps:10

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (257 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :020 Order:Lepidoptera Family:Oecophoridae Species binomial: Total Collected:32 # in CD StickyTraps:1 # in Funnel Crawl Traps:12 # in HangTraps:19

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (258 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :025 Order:Lepidoptera Family:Oecophoridae Species binomial: Total Collected:57 # in CD StickyTraps:2 # in Funnel Crawl Traps:46 # in HangTraps:9

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (259 of 290) [12/01/2006 5:02:02 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :042 Order:Lepidoptera Family:Oecophoridae Species binomial: Total Collected:7 # in CD StickyTraps:1 # in Funnel Crawl Traps:3 # in HangTraps:3

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (260 of 290) [12/01/2006 5:02:03 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :048 Order:Lepidoptera Family:Oecophoridae Species binomial: Total Collected:2 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (261 of 290) [12/01/2006 5:02:03 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :067 Order:Lepidoptera Family:Oecophoridae Species binomial: Total Collected:6 # in CD StickyTraps:0 # in Funnel Crawl Traps:5 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (262 of 290) [12/01/2006 5:02:03 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :116 Order:Lepidoptera Family:Oecophoridae Species binomial: Total Collected:50 # in CD StickyTraps:0 # in Funnel Crawl Traps:8 # in HangTraps:42

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (263 of 290) [12/01/2006 5:02:03 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :118 Order:Lepidoptera Family:Oecophoridae Species binomial: Total Collected:10 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:10

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (264 of 290) [12/01/2006 5:02:03 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :146 Order:Lepidoptera Family:Oecophoridae Species binomial: Total Collected:3 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:3

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (265 of 290) [12/01/2006 5:02:03 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :153 Order:Lepidoptera Family:Oecophoridae Species binomial: Total Collected:13 # in CD StickyTraps:0 # in Funnel Crawl Traps:9 # in HangTraps:4

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (266 of 290) [12/01/2006 5:02:03 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :243 Order:Lepidoptera Family:Oecophoridae Species binomial: Total Collected:4 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:4

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (267 of 290) [12/01/2006 5:02:03 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :247 Order:Lepidoptera Family:Oecophoridae larvae Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (268 of 290) [12/01/2006 5:02:03 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :264 Order:Lepidoptera Family:Torticidae Species binomial: Total Collected:4 # in CD StickyTraps:0 # in Funnel Crawl Traps:3 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (269 of 290) [12/01/2006 5:02:03 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :114 Order:Lepidoptera Family:Tortricidae Species binomial: Total Collected:40 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:40

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (270 of 290) [12/01/2006 5:02:03 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :145 Order:Lepidoptera Family:Tortricidae Species binomial: Total Collected:15 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:14

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (271 of 290) [12/01/2006 5:02:03 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :242 Order:Lepidoptera Family: Species binomial: Total Collected:4 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:3

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (272 of 290) [12/01/2006 5:02:03 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :265 Order:Lepidoptera Family: Species binomial: Total Collected:3 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:3

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (273 of 290) [12/01/2006 5:02:03 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :274 Order:Lepidoptera Family: Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (274 of 290) [12/01/2006 5:02:03 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :062 Order:Lepidoptera larvae UnID Family: Species binomial: Total Collected:11 # in CD StickyTraps:0 # in Funnel Crawl Traps:11 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (275 of 290) [12/01/2006 5:02:03 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :175 Order:Mangled UNID? Family: Species binomial: Total Collected:11 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:11

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (276 of 290) [12/01/2006 5:02:03 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :084 Order:Mesostigmata Family: Species binomial: Total Collected:10 # in CD StickyTraps:1 # in Funnel Crawl Traps:8 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (277 of 290) [12/01/2006 5:02:03 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :087 Order:Neuroptera Family:Hemerobiidae Species binomial:Micromus tasmaniae Total Collected:2 # in CD StickyTraps:0 # in Funnel Crawl Traps:1 # in HangTraps:1

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (278 of 290) [12/01/2006 5:02:03 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :097 Order:Neuroptera? Family: Species binomial: Total Collected:2 # in CD StickyTraps:0 # in Funnel Crawl Traps:2 # in HangTraps:0

Morphospecies # :095 Order:Orthoptera Family:Gryllacrididae Species binomial:Adult Kinemania sp. YDB 1 Total Collected:12 # in CD StickyTraps:0 # in Funnel Crawl Traps:12 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (279 of 290) [12/01/2006 5:02:03 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :081 Order:Orthoptera Family:Gryllacrididae Species binomial:Juvenile Kinemania sp. YDB 1 Total Collected:5 # in CD StickyTraps:0 # in Funnel Crawl Traps:5 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (280 of 290) [12/01/2006 5:02:03 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :001 Order:Plecoptera Family:Eustheniidae Species binomial:Eusthenia spectabilis? Total Collected:10 # in CD StickyTraps:0 # in Funnel Crawl Traps:10 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (281 of 290) [12/01/2006 5:02:03 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :004 Order:Plecoptera Family: Species binomial: Total Collected:38 # in CD StickyTraps:0 # in Funnel Crawl Traps:38 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (282 of 290) [12/01/2006 5:02:03 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :304 Order:Pseudoscorpionidae Family: Species binomial: Total Collected:2 # in CD StickyTraps:0 # in Funnel Crawl Traps:2 # in HangTraps:0

http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm (283 of 290) [12/01/2006 5:02:03 PM] http://www.geog.utas.edu.au/yoav/pho/voucher/vouchergallery.htm

Morphospecies # :203 Order:Psocoptera Family: Species binomial: Total Collected:2 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:2

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Morphospecies # :233 Order:Psocoptera Family: Species binomial: Total Collected:2 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:2

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Morphospecies # :240 Order:Psocoptera Family: Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

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Morphospecies # :272 Order:Psocoptera Family: Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

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Morphospecies # :399 Order:Psocoptera Family: Species binomial: Total Collected:1 # in CD StickyTraps:1 # in Funnel Crawl Traps:0 # in HangTraps:0

Morphospecies # :300 Order:Skink Family: Species binomial: Total Collected:2 # in CD StickyTraps:0 # in Funnel Crawl Traps:2 # in HangTraps:0

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Morphospecies # :213 Order:Trichoptera Family:Leptoceridae Species binomial: Total Collected:2 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:2

Morphospecies # :161 Order:Trichoptera Family: Species binomial: Total Collected:1 # in CD StickyTraps:0 # in Funnel Crawl Traps:0 # in HangTraps:1

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