The University of Bristol

School of Life Sciences

MSci Practical Project Report

Veteran Trees in PAWS: A Comparison of the Use of

Different Ages and Species of Tree by Woodland
Fauna

Author:
Supervisor:

Thom Erritt

Dr. Jane Memmott

te0047@my.bris.ac.uk
1019047

Word Count: 5,998

Veteran Trees in PAWS: A Comparison of the Use of Different Ages and Species
of Tree by Woodland Fauna
Thom Erritt
School of Life Sciences
The University of Bristol

Abstract
Two distinct yet complementary investigations were carried out to examine the ecological roles of veteran trees in
Plantations on Ancient Woodland Sites (PAWS). The aim of the first investigation was to compare the abundances and
diversities of birds and invertebrates present on the two main categories of tree found in a PAWS; veteran broadleaf
trees and non-native conifers. It was found that the veteran trees were host to significantly higher abundances and
diversities of our survey species, providing evidence that they play an important part in maintaining the biodiversity
of these PAWS. The aim of the second investigation was to examine the relationship between the age of a veteran tree
and the number of invertebrates and microhabitats on its trunk. There was found to be a positive correlation between
the age of a veteran tree and both of these variables. This indicates that the biodiversity of veteran trees may increase
with age and therefore so does their ecological value.
Keywords: veteran tree, PAWS, ecology, birds, invertebrates, microhabitats

1. Introduction

The diverse range of ecological roles performed by veteran trees make them of huge value to the biodiversity

of woodland ecosystems across Great Britain (1) . Other than their superb ecological value, veteran trees are also

genetically, aesthetically, culturally and historically valuable. In combination, these values make veteran trees an

important part of the natural heritage of Great Britain. This project aims to quantify the ecological value of veteran

trees specifically in Plantations on Ancient Woodland Sites (PAWS). This should provide evidence that they support

a wide range of woodland species and help to increase the biodiversity of the ecosystems they belong to.

The term veteran tree can be fairly ambiguous. There are a number of slightly different definitions of the term;

however one of the most concise and accurate ones comes from Croft (2) , who defines a veteran tree as one that

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possesses all or a number of the following characteristics:

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1. Biological, aesthetic or cultural interest because of its great age.

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2. A growth stage that is defined as ancient or post-mature.

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3. A chronological age that is old relative to others of the same species.

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One of the reasons that veteran trees have captured numerous imaginations through the ages is due to their aesthetic

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appeal. Their unique visual charm stems from the morphological characteristics that a tree begins to develop as it ages.

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Some of the most important and recognisable of these veteran characteristics are as follows:

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1. Large trunk circumference (compared to other trees of same species) after many years of accumulation of annual
growth rings.

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2. Progressive diminution in width of successive annual growth rings.

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3. Decay, leading to branch failure and trunk hollowing.

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4. A progressive or episodic reduction in post mature crown size, often known as retrenchment.

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These characteristics, amongst many others, make veteran trees excellent habitats for a wide range of other wood-

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land species; which is why these trees are so ecologically valuable. These species include a multitude of fungi, lichens

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and bryophytes, with up to 2,500 species of fungi and 50 species of moss known to associate with veteran trees in the

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forests of Sweden (3) . Other species that are reliant on veteran trees include a wide array of arthropods; these species

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inhabit niches all over the tree, from pollinators in the foliage, to predatory spiders on the bark and fungivorous

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collembola around the base (4) . Finally, birds and mammals rely on veteran trees for foraging and nesting, with the

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great spotted woodpecker (Dendrocopus major) relying on insects found in fallen wood for 97% of its diet throughout

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winter (5) while Barbastelle (Barbastella barbastellus) and Daubentons (Myotis daubentonii) bats are known to roost

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and give birth in tree holes during summer months (6) .

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The majority of this project will conducted by surveying the trees found in Plantations on Ancient Woodland Sites.

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Most of the PAWS in the United Kingdom were created between the years of 1941 and 1970 when as much as 44% of

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the UKs surviving ancient woodland was converted to coniferous plantation in order to avoid timber shortages during

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future instances of war, resulting in the formation of around 220,000 hectares of PAWS (7) (8) . One interesting feature

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of these PAWS is that they often contain extant features of the ancient woodland that preceded them, which are now

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some of the most rare, yet biodiverse and ecologically valuable ecosystems in the UK (9) . These extant features are

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more often than not the veteran trees which were excluded from the initial clear felling and are now surrounded by

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conifer plantation. These are the two categories of tree that this project aims to compare, the dominant non-native

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conifers and the broadleaf veterans that occasionally intersperse them.

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The restoration of PAWS has become a top priority for many woodland managers after the Habitat Action Plans

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for Native Woodland set in motion ideas about enhancing the remaining features of ancient woodland through the

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conversion of PAWS to Restored Native Woodland on Ancient Sites or RNWAS (10) . This restoration involves the

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gradual replacement of non-native conifer species with broadleaf species in order to encourage growth of a forest that

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is more valuable for both recreation and biodiversity (11) (12) .

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We aim to study the trees within two separate PAWS located in Leigh Woods, North Somerset and Savernake

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Forest, Wiltshire. We will do this by surveying the abundances and diversities of the birds, arthropods and micro-

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habitats that are present on different species and ages of tree. We decided to survey these criteria as they are all
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known to be good indicators of the general ecological health of an area. For example, birds are known to be excellent

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ecological indicators because of their widespread presence, high detectability and sensitivity to natural and human

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alteration of environments (13) (14) . In addition to this, we chose to include collembola and arachnids in our arthropod

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surveys as collembola are known to be excellent indicators of soil fauna diversity whilst arachnids have been reliably

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used numerous times to determine the ecological value of many different habitats including woodlands, marshlands

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and grasslands (15) (16) . The final criteria that we decided to survey was the abundance of microhabitats on the trees,

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this is because microhabitats are known to be an excellent proxy for representing the biodiversity of species that are

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generally difficult to observe such as fungi, lichens, bats and invertebrates (17) (18) (19) (20) .

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The knowledge of whether these ecological indicator species are present in higher abundances on different trees

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in PAWS will provide numerous valuable insights with wide-ranging benefits. For example, the ascertainment of the

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fact that the veteran trees in PAWS are host to higher abundances and diversities of woodland fauna than the conifers

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that surround them will provide valuable evidence that veteran trees are helping to increase the levels of biodiversity

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found in PAWS. It should also provide evidence for the importance of future conversion of PAWS to RNWAS by

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replacements of the incumbent conifers with the potentially more ecologically valuable broadleaf trees. Furthermore,

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the ascertainment of whether different species and ages of broadleaf veteran are host to different assemblages of

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species will also be invaluable for the management of currently mature trees, ensuring the most ecologically valuable

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populations of veteran trees in the future.

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2. Methods

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This project was conducted through two distinct yet complementary investigations. The first investigation was an

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intensive study on a small sample of trees found around Bristol, with the aim of directly comparing the differences

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in abundances of birds and arthropods on veteran and conifer trees; this part of the project will be described under

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Objectives 1 and 2. The second investigation was a more extensive study on a much larger sample of veteran trees

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found in Savernake Forest, with the aim of observing how the abundances of invertebrates and microhabitats on

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a veteran tree vary with age and species; this part of the project will be described under Objectives 3 and 4. All

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statistical analyses throughout this project were completed on IBM SPSS 21.

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2.1. Objective 1: Comparing the abundance and diversity of birds found on veterans and conifers in PAWS

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Bird counts were recorded from nine veteran and fourteen conifer trees. Seven of the veteran trees were Sweet

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Chestnuts (Castanea sativa) and two were Oaks (Quercus robur), four of the conifers were Norway spruce (Picea

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abies) and ten of them were Scots Pine (Pinus sylvestris). The veterans were selected as they were the only ones

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located either in PAWS, or in close association with conifer trees in the area surrounding the city of Bristol, UK.

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Three of the nine veteran trees were located in a PAWS at Paradise Bottom, Leigh Woods (51.458269, -2.635398).

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The other six were located in Ashton Court, an area of parkland to the west of Bristol (51.445481, -2.642838). The
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conifer trees in our survey were randomly selected from all of the conifer trees that surrounded our chosen veteran

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trees. Four conifers were chosen at the Paradise Bottom site as they were estimated to occupy a similar amount of

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space as the veterans there whilst nine conifers were chosen at the Ashton Court site for the same reason.

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Count censuses were used to deduce the abundance and diversity of birds visiting our sample of veteran and

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conifer trees. Birds were counted if they landed in one of our survey trees, birds flying above or landing in trees

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other than our survey trees were ignored. A total of 8 hours of bird watching was carried out at Paradise Bottom as

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it was possible to watch the veterans and conifers simultaneously, while a total of 16 hours was spent bird watching

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at Ashton Court as the veteran trees and the conifers had to be surveyed separately due to the short distance between

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them. Bird species were identified using binoculars and a bird identification guide before being classified as either

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small, medium or large.

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A One-Way ANOVA was used to test for differences between the conifers and the veterans after checking visually

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for assumptions of independence, normality and equal variance. Shannon-Wiener Biodiversity Indices were calculated

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to compare the biodiversity of each group of trees.

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2.2. Objective 2: Comparing the abundance and biodiversity of invertebrates found in veterans and conifers in PAWS

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Bark brushing was used to sample the invertebrates present on the same population of veteran and conifer trees

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that were used in Objective 1. Our methodology was to place string transects around the veteran trees at heights of

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20, 120 and 220cms from the trees base. A large paintbrush was then used to gently brush the bark 10cm above and

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below the full length of the transect whilst catching any invertebrates that fell off in a plastic container. Each transect

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was sampled for five minutes. As the diameter of the conifer trees in our sample were much smaller, the whole basal

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two metres of their trunks was sampled for five minutes. The invertebrates were then euthanized and stored at -20 C

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until processing.

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The abundances and diversities of invertebrate morphotypes (collembola, arachnids, mites, flies, grubs, beetles,

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woodlice, other) that were present in each of our samples was determined in the laboratory using a Leica LED3000

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SLI on a Stereo Microscope. The number of invertebrates found per m2 of bark was then calculated using the circum-

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ference of the survey trees; this allowed us to compare the conifers and the veterans accurately. A One-Way ANOVA

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was used to compare the two data sets in the same way as in Objective 1.

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2.3. Objective 3: Comparing the abundance and biodiversity of invertebrates across different ages and species of

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veteran tree in PAWS

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Data was obtained by sampling every veteran tree located in a PAWS in the northern part of Savernake Forest,

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Wiltshire (51.406147, -1.667596). Samples of invertebrates were obtained using the same methodology of bark

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sampling used in Objective 2, the only difference being that only one transect was sampled per tree at a height of

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120cm in order to save time. Our sample initially included a total of 200 veterans comprised of Oaks, Beeches, Sweet

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Chestnuts, Horse Chestnuts, Silver Birches and Sycamores. However statistical analyses were only conducted on a
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total of 179 trees comprised of Oaks, Beeches and Sweet Chestnuts as these were the only species of tree whose ages

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could be reliably estimated using the exact methodology suggested by White (21) .

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The relationship between tree age and invertebrate population was tested using a linear regression in the same way

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as in Objective 3. A One-Way ANOVA was used to test for differences between invertebrate abundances of different

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species of veteran tree.

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2.4. Objective 4: Comparing the abundance and diversity of microhabitats in different ages and species of veteran

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trees found in PAWS

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Microhabitat abundances were obtained using a point count census of the entire basal two metres of the same sam-

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ple of veteran trees used in Objective 3. Microhabitats were classified into six easily identifiable and distinguishable

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categories: basal hole, trunk hole, flaking bark, crevice in trunk, gall/canker, sap run.

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A linear regression was used to test the relationship between tree age and number and diversity of microhabitats in

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the same way as in Objective 3. A One-Way ANOVA was used to test for differences between microhabitat abundances

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of different species of veteran tree.

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3. Results

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3.1. Objective 1: Comparing the abundance and diversity of birds found on veterans and conifers in PAWS

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A significantly higher number of birds were observed on the veteran trees than on the conifers (P<0.001, F46,1

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=7.218). The mean number of birds recorded per survey on veteran trees was 6.75 whilst the mean number of birds

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recorded per survey on conifer trees was only 1. There was also found to be significantly higher mean numbers of

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sightings of all sizes of birds (small, medium and large) in veteran trees than in conifers (Figure 1.).

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Calculation of Shannon-Wiener Biodiversity Indices shows that the species of bird sighted in the veteran trees were

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more diverse than those sighted in the conifer trees (veteran biodiversity index=1.75, conifer biodiversity index=1.66).

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3.2. Objective 2: Comparing the abundance and biodiversity of invertebrates found in veterans and conifers in PAWS

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A significantly higher number of invertebrates was observed on the bark of veteran trees than on the conifers

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(P<0.001, F182,1 =17.198). The mean number of invertebrates per m2 of veteran tree bark was found to be 5.86 while

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the mean for conifer trees was only 1.3. There was also a found to be significantly higher numbers of all morphotypes

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of invertebrate on the veteran trees than on the conifers (Figure 2.).

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Comparison of the Shannon-Wiener Biodiversity Indices of collected invertebrates show that veteran trees are host

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to more diverse assemblages of invertebrates than conifer trees (veteran biodiversity index=1.17, conifer biodiversity

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index=0.63).

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Figure 1: Mean abundances (+/-SE) of small, medium and large

Figure 2: Mean abundances (+/-SE) of each morphotype of inver-

birds observed on veteran and conifer trees.

tebrate per m2 of bark of veteran and conifer trees.

3.3. Objective 3: Comparing the abundance and biodiversity of invertebrates across different ages and species of
veteran tree in PAWS

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A weak yet positive correlation was found between the age of a veteran tree and the number of invertebrates found

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on its bark (Ra =0.292, R2 =0.085, F177,1 =16.459, P<0.001). Significant differences were also found between the mean

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number of insects found on each species of tree (P<0.001, F176,2 =17.708). Sweet Chestnuts were found to have the

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highest mean number of insects present on their bark (31.25), followed by Oak trees (14.78) and finally Beech trees

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(8.75).

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The trees in our sample did not appear to become more biodiverse as they aged as there was no significant corre-

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lation between the Shannon-Wiener Biodiversity index of a tree and its age. There was also no significant difference

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between the mean Shannon-Wiener indices of each species of tree in our sample.

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3.4. Objective 4: Comparing the abundance and diversity of microhabitats in different ages and species of veteran
trees found in PAWS

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Strong positive correlations were found between the age of veteran trees and the number of microhabitats they

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possessed across all species of tree in our sample. Oak trees tended to have the strongest correlation between age

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and number of microhabitats (Ra =0.879, R2 =0.772, F104,1 =378.976, P<0.001), followed by Beech trees (Ra =0.761,

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R2 =0.579, F49,1 =67.258, P<0.001), and finally Sweet Chestnuts (Ra =0.725, R2 =0.525, F20,1 =9.702, P<0.001) (Figure

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4.).

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The mean number of microhabitats found on each species of veteran tree was also found to vary significantly

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(P<0.001, F176,2 =67.982). Beech trees were found to have the highest mean number of microhabitats (31), followed

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by Sweet Chestnuts (14) and finally Oak trees (8).

Figure 3: Relationship between the age of all species of tree and

the number of invertebrates per

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m2

Figure 4: Relationship between the age of each species of tree and

of bark.

the number of microhabitats on its trunk.

Finally, It was also found that the veteran trees at the Bristol survey sites had significantly more microhabitats than

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the conifers present there (veteran mean=19.9, conifer mean=2.0, P<0.001, F23,1 =21.39)

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4. Discussion

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There are a number of conclusions that can be inferred from our results. The first of these is that in the PAWS

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which we surveyed, veteran trees tended to host higher abundances, and higher diversities of birds and invertebrates

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than the conifers that surrounded them. The second major finding was that, in the veterans that we sampled, there

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was a positive correlation between the age of the tree and the number of arthropods that were present on its bark.

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The final major finding of our survey was that there were positive correlations between all species of veteran tree that

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we sampled and the number of microhabitats on their trunks, however the strength of correlations differed between

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species, as did the mean number of microhabitats.

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Overall, these findings lead us to conclude that there are significant differences in the ecological roles of different

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species of tree in PAWS, specifically in terms of the abundances of organisms that they are supporting. The most

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prolific of these differences being that the veteran trees in PAWS are playing a more significant role in supporting the

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biodiversity of the area than the conifers that surround them. I will now consider the limitations of this project before

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assessing the findings of each objective in more depth and in association with other literature on the subject.

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4.1. Limitations

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Our surveys were subject to a number of minor limitations. The first of these was the size of the sample of trees

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that were surveyed in Objectives 1 and 2. In total, only nine veteran trees were sampled as these were the only ones

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located in PAWS in the area around the University of Bristol. Ideally, a sample size of at least double this number

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would have been desirable. However, because of the more in-depth and descriptive nature of this section of the study it
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was still possible to obtain significant and conclusive results. I do not believe a larger sample size would have altered

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our major findings however it may have allowed us to gain more insightful conclusions in terms of comparing conifers

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with different species of veteran tree rather than just the population as a whole.

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A second limitation to our study was the time of year that our data was collected. All experimental work was

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carried out over a ten week period from the middle of January to the end of March in order to comply with the term

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schedule of our university. The consequence of this is that many arthropod species would have still been overwintering

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and would not have been active on the bark (22) . Ideally, our surveys would have been completed during late spring and

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summer when arthropod activity would be at its peak. However, I do not believe this limitation will have significantly

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affected our findings as it was still possible to collect large numbers of arthropods from both the veterans and the

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conifers that should be proportional to the number of insects that would be active at the height of summer, and thus

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the differences between them would still be present.

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4.2. Objective 1 - Birds

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We can obtain a better understanding of why larger abundances and diversities of birds were sighted in veteran

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trees rather than conifers by considering why avifauna are using these trees in the first place. Birds are known to use

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trees for a diverse range of ecological functions including roosting, breeding, overwintering, perching and foraging (23) .

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I would hypothesise that the reason we saw more birds in the veteran trees was because the higher abundances of prey,

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microhabitats and epiphytes on the veterans allowed the birds to perform these functions more efficiently and safely

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than they could in the conifers.

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For example, it is highly likely that birds would have better foraging opportunities on veterans because, as our

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results have shown, they are host to higher abundances of invertebrates than conifers. It has also been proven that

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birds regularly feed on the morphotypes of invertebrate that we found on our survey trees; experiments by Gunnarson

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have shown that collembola and arachnid abundances were significantly higher in sites where bird predation was

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artificially removed (24) . Therefore, we can hypothesize that one of the reasons that we encountered more birds in the

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veteran trees than the conifers was that foraging opportunities were better on the veteran trees due to their significantly

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higher abundances of arthropod prey.

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The second reason that birds may find veteran trees more appealing than the surrounding conifers is because of the

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superior nesting and roosting sites on offer. We know that finding a suitable nest site is essential to the life histories

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of many species of bird, who generally rely on trees to provide them. For example, ten of the eleven species of

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European woodpecker and at least ten species of European owl are known to make their nests in tree holes (25) (26) .

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Furthermore, Laiolo states that the availability of tree holes is fundamental for woodland birds, and can result in

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severe competition between hole-nesters (23) . When considering these facts in combination with our findings from

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Objective 4, that veteran trees have more microhabitats compared to conifers, we can see that birds may have been

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choosing to land in veteran trees as they offer a higher probability of having a microhabitat that would be suitable for

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nesting.
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It is also possible that we may have seen more birds in the veterans compared to the conifers because of the

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higher number of epiphytes present on veteran trees. Nadkarni found that in forest ecosystems, a higher abundance

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of epiphytes led to a higher abundance of birds (27) . Possible reasons for this include that epiphytes increase the

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total amount of resources available, provide opportunities for resource specialisation and temporally spread resources

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throughout the year (28) . Although our surveys did not directly compare the number of epiphytes on each category of

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tree, I would consider it likely that there would be higher abundances of epiphytes on the veterans. This is because the

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presence of epiphytes is included as a specifically veteran characteristic in Crofts unifying description of veteran

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trees (2) .

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4.3. Objectives 2 and 3 - Invertebrates

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The second major finding of our surveys was that there were higher abundances and diversities of all morphotypes

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of invertebrates in older veteran trees than there were in conifers and younger veteran trees. In this part of the

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discussion I will consider numerous reasons for why we may have seen these differences. However, I feel that it will

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be easiest to explain these differences whilst focusing on only two of the morphotypes that appeared in our surveys:

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arachnids and collembola. This is because these two taxa are known to be excellent ecological indicators as they

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make significant contributions to arboreal food webs. Collembola contribute to food-webs as they are preyed upon

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by a variety of other species, notably arachnids (29) ; arachnids contribute to food-webs as they are intermediate level

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predators and also act as a food source for birds (30) .

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I hypothesise that one of the most probable reasons for why we recorded the highest abundances of arthropods

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on the bark of older veteran trees was because of their increased structural complexity and therefore increased habitat

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heterogeneity.

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The effect of arboreal structural complexity on arthropod populations has been thoroughly studied by Halaj (31) .

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This work involved devising a method for comparing the structural complexities of different species of tree in order to

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ascertain which physical characteristics were the best predictors of arachnid abundance and diversity. Halaj deemed

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that a tree was more structurally complex if it had a larger diameter at breast height, larger maximum vertical and

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horizontal branch spread, larger range of branching angles and higher biomass. He then carried out a pair of ex-

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periments whilst using this definition. The first experiment surveyed the natural arachnid abundances of trees with

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differing structural complexities and the second experiment manipulated the complexity of tree branches in a lab and

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viewed its effect on populations of arachnids and collembola.

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The results of the first experiment showed that higher abundances of arachnids were present on more complex

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trees (32) . The results of the second experiment showed that reducing the complexity of a habitat reduced the abun-

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dances of arachnids and collembola whilst increasing the complexity of a habitat significantly increased arachnid

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abundances and caused an eight-fold increase in abundances of collembola (31) .

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These conclusions allow us to put the results of our own experiments in perspective. We saw almost exactly

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the same pattern as Halaj, our results showed that in more complex environments, i.e. veteran trees with increased
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diameter, branch spread, branch angle variability and biomass there were higher abundances of arthropods than in

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less complex conifers. As we can see, the structural complexity of a habitat has a significant influence on the resident

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arthropod community.

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Arthropods use trees for a number of different ecological functions. They are used for shelter, foraging, oviposi-

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tion, sun-basking, sexual display, and for herbivores such as collembola, they provide nutritional value (31) . I would

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hypothesise that in a structurally more complex tree, arthropods are able to perform these essential parts of their

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life-histories more efficiently and safely. For example, increased branch biomass correlates positively with increased

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surface area, meaning larger populations and higher densities of arthropods can be supported with less intense com-

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petition. Furthermore, increased branching angles give spiders more options in terms of sites for web attachment as

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well as for finding suitable retreat sites that can be used for egg-laying and predator avoidance. Overall, what we are

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seeing is that increased complexity of veteran trees leads to increased habitat heterogeneity and therefore increased

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capacity to support larger and more diverse populations of invertebrates.

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These ideas can be taken one step further when we consider the complexity of the arboreal environment at an

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even finer scale. Many studies have shown that certain biological surfaces have a fractal, or at least semi-fractal

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structure (33) (34) (35) . A surface is fractal if its area increases when a progressively finer unit of measurement is used;

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this is because the finer units of measurement will detect certain irregularities that larger units of measurement would

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not, thus increasing the surface area (36) . Experiments by Florin have shown that bark has a fractal structure, meaning

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the surface area of bark becomes greater when the unit of measurement becomes smaller (35) .

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The degree of fractality of a certain surface can be worked out by measuring the surface and using a specific

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equation which produces a number between 1 and 2, this number is called the fractal dimension of the surface, or D.

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A surface with a D value nearer to 2 will be more fractal than a surface with a D value nearer to 1 (37) . Florin used

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this equation to work out the fractal dimension of the barks of varying ages of plum tree and showed that older, more

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complex, fissured and exfoliated bark tended to have a higher fractal dimension than younger, less complex bark.

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They also found that the age of a plum tree could be accurately predicted from the D value of its bark (35) .

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As we now know that different complexities of bark have different fractal dimensions, we can now consider

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what this means for the arthropods living upon it. Gunnarson has hypothesised that if the way an animal perceives

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its environment is related to its body size then an animal with a body length of 3mm will perceive the area of a

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surface with a D value of 1.5 as up to one order of magnitude larger than an animal with a body length of 30mm;

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this increased perceived surface area should be able to support higher population densities of animals with a body

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length of 3mm than 30mm. Gunnarson tested this hypothesis by allowing spiders to invade artificial plant habitats

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with different D values before comparing their population densities and abundances. He found higher densities of

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spiders in environments with high fractal dimensions and lower densities of spiders on environments with lower fractal

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dimensions (38) .

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If we now consider the findings of the Florin and Gunnarson experiments together we can obtain an interesting hy-

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pothesis for why we recorded more arthropods on veteran trees than on conifers as well as on older veteran trees than
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on younger ones. The Florin paper concluded that the more complex bark from older trees had a higher fractal dimen-

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sion. The Gunnarson paper concluded that structures with higher fractal dimensions supported higher abundances

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of arachnids than structures with lower fractal dimensions. Therefore we could hypothesise that the differences in

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fractal dimensions of bark between our comparison groups, which was deep and fissured in our veteran Oaks and

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Sweet chestnuts (high D value) while smooth and flaky in our conifers (low D value), could have contributed to

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differences in invertebrate abundances in our samples. This is because the high D value bark of older veterans had a

291

larger perceived surface area that could support larger populations of arthropods.

292

Overall, I believe our findings that invertebrate abundances and diversities are higher in veteran trees than conifers

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as well as in older veteran trees than younger veteran trees are well founded. There is strong evidence that higher

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structural complexity at both a habitat and fractal level can lead to higher abundances of both spiders and collembola,

295

reflecting exactly what we discovered in our experiments. These explanations also allows us to hypothesise why we

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saw differences between the invertebrate communities of different species of tree. In theory, the smooth, almost sterile

297

looking bark of Beech trees would have a had much lower structural and fractal complexity than the deep, fissured

298

bark of Sweet Chestnuts; leading to much lower abundances of invertebrates. This is exactly what we saw in our

299

experiments, Sweet Chestnuts were host to the highest abundances of invertebrates whilst Beeches were host to the

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lowest.

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4.4. Objective 4 - Microhabitats

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Our third and final major finding was that as the age of veteran trees increased, so did the number of microhabitats

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on their trunks. These findings support the hypothesis that veteran trees become more ecologically valuable as they

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get older. This is because the abundance of microhabitats in an ecosystem is known to correlate with the abundance

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of many arboreal species such as many passerines, bats and invertebrates (17) (18) .

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I believe this correlation exists because the number of microhabitats that a tree has will be likely to represent of

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the number of niches that the tree may have available. Having higher abundances of niches means that a tree has a

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higher capacity to support a larger abundances of certain species with minimal amounts of competition.

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4.5. Tree Size

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One final possible reason for why we observed different abundances and diversities of species in different cate-

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gories of tree is simply to do with their size. The veteran trees in our sample tended to be much larger than the conifer

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trees (mean veteran diameter=99.7cm, mean conifer diameter=46.2cm) as did the older veteran trees compared to the

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younger ones because tree diameter is directly related to age. When considering these differences in size in combina-

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tion with island biogeography theory we can gain some insight into why abundances of certain species differ between

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veterans and conifers.

316

It has been proposed that island theory could be useful in explaining the differences in arthropod abundances

317

between different species of plant (39) . This is because larger islands, in this case larger veteran trees, are more likely
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to support higher abundances and diversities of species simply because they offer more available space and are more

319

likely to be discovered in the first place (40) . Therefore, one possible reason for why we recorded more species in the

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veterans than the conifers was simply because the veterans were larger and were therefore more likely to be discovered

321

and populated by arthropods and birds.

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4.6. Conclusions - What Does This Mean for PAWS and Veteran Trees?

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This project has two major implications for PAWS and veteran trees. The first of these is that the current population

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of veteran trees located in PAWS are extremely beneficial to the ecological health and biodiversity of the ecosystems

325

that they belong to. Our surveys show that the veterans are acting as islands of high species abundance and richness

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in a sea of less diverse conifers; this has the effect of bringing up the average biodiversity of the PAWS as a whole.

327

This is clear evidence that the current population of veteran trees should be conserved and managed to maximise the

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biodiversity they support. For example, we saw that birds were choosing to land in veteran trees rather than conifers

329

and that this was most likely due to the better nesting and foraging sites provided by the veteran trees. As competition

330

between birds for nest sites is fierce, any loss of veteran trees and their abundance of nest sites from a PAWS would

331

almost definitely lead to a drop in bird populations of the area. In contrast, an increase in the number of nest sites

332

produced by correct management of veteran trees would likely lead to an increase in the bird population of the area,

333

leading to an increase in the overall biodiversity of the PAWS.

334

The second implication that can be gleaned from our findings is that gradual replacement of the conifers with

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broadleaf species would have a positive effect on the biodiversity and ecological health of the PAWS ecosystem. This

336

is because larger populations of broadleaf species would have the capacity to support higher abundances and more

337

diverse assemblages of species than the current population of conifers. In addition to this, the broadleaf trees would

338

eventually mature into veteran trees and would hopefully begin to develop characteristics and species assemblages

339

similar to those of the ancient woodland that once covered large quantities of the Great Britain. This idea of the

340

conversion of PAWS to RNWAS is currently being explored by woodland managers around Great Britain and the

341

results of our surveys provide evidence that it would be beneficial to the ecology of these woodlands and the country

342

as a whole.

343

Another interesting observation was that different species of veteran tree may have been carrying out slightly

344

different ecological functions in the PAWS. It was clear that the deep, complex bark of Sweet Chestnuts was an

345

excellent habitat for a wide range of invertebrate species. Whilst the complex branching and exposed roots of the

346

beech trees we surveyed were home to the greatest number of microhabitats, creating excellent perching, foraging and

347

nesting sites for birds. So, we can see that the conservation of a diverse range of veteran trees in PAWS would allow

348

the conservation of the greatest range and diversity of ecological roles.

349

Overall, we have observed clear differences in the use of different ages and species of trees in PAWS by woodland

350

fauna. This has allowed us to conclude that the presence of veteran broadleaf trees and the superior assemblages

351

of species that they support are clearly ecologically beneficial to the PAWS that they belong to. Therefore, the
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conservation of the veterans and the potential conversion of PAWS to RNWAS will ensure the enhancement and even

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restoration of the remnant assemblages of species and interactions that would have filled the ancient woodland of

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pre-1900s Great Britain.

355

5. Competing Interests

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357

This project has no competing interests.

6. Acknowledgements

358

Firstly, I would like to thank Ellie Lewis for being such an excellent and enthusiastic project partner. I would also

359

like to thank Professor Jane Memmott at the University of Bristol for her continued guidance and advice. Finally,

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I would also like to thank Paul Rutter and Tom Blythe at PlantLife and the Forestry Commission for their ongoing

361

support and recommendations throughout this project and Millie and Nick Carmichael for their hospitality during our

362

stay in Savernake Forest.

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7. Funding

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365

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This project was funded by the University of Bristol.

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