Canopy inverterbrate diversity, Literature Review, Duncan Allen

Duncan Allen Biol 3001

Canopy invertebrate diversity in tree collections

A Literature Review

D. W. Allen

December 2008

Bsc Wildlife Conservation

School of Biological Sciences

University of Plymouth

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Contents

1.0 Introduction 4

2.0 Urban and Agricultural trees history and uses 5

3.0 Trees as habitats 8

4.0 Canopy Invertebrates 13

5.0 Sampling canopies methods/difficulties 14

6.0 Implications 17

7.0 Study aims 18

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Abstract

It is often sited that trees hold vast arrays of invertebrates with some

trees such the English Oak (Quercus robur) holding as many as 300 plus

species. This literature review will be looking at the importance of urban

forested areas as habitats and niches for invertebrate fauna. Urban

forests can include stately home gardens and grounds as well as town

and city parks, these areas of urban forest may prove useful as habitats

for invertebrate communities. It may also be possible to judge the health

of an urban environment by the communities of invertebrate that are

found. Urban forests may also prove useful as wildlife corridors for

migrating wildlife and immigrating wildlife in a patchy and fragmented

habitat landscape. There is some evidence to support this idea that they

are useful to wildlife as corridors. Assessing the invertebrate

biodiversity of trees may also have benefits for planning and planting of

future forested areas or even have implications when replanting

hedgerows, certain parasatoid wasps used in bio control may live out

part of the life cycle in or on a tree. The species of tree can then be

planted as an aid to farmers. Understanding the biodiversity is also a

worthy pursuit for the potential new medicines, foods, and materials that

can be found within the worlds rain forests.

Key Words: Invertebrates, Biodiversity, Insect Sampling, Tree Canopy, Urban Forests,

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1.0 Introduction

The worlds forests play host to almost two thirds of the worlds species making

them very important for biodiversity (GFW 2008), woodland is declining all

over the world in the last three decades half the world’s forests have been

cleared (UN Earth-watch 2008, WWF 2007). In Europe alone forested areas

have gone from 4,690 square km to 1,521 square km in the last 5000 years

(GFW 2008). Roughly 90% of the UK’s forests have been lost in the last 5000

years; forest now only makes up 10% of the UK’s land cover (around 2.7

million hectares). The cover of forest in the UK’s countries is as follows

England 7%, Scotland 15%, Wales 12%, and Northern Ireland 6% (Forestry

commission 1999, Spencer et al 1992).

The world’s forests serve as crucial habitats for a variety of species; therefore

they play an important role safeguarding biodiversity (UN Earth-watch 2008).

Old trees in collections can also be important as ‘’habitat trees’’ generally tree

collections and parks are of an age now that the individual trees within these

collections are on average about 150 years old these trees will have bore

holes and hollows which provide nesting holes and important roosts for

nocturnal species. In the Australian rain forests near Queensland there have

been studies that show how important habitat trees are. It has been found that

42 % of mammals, 28 % of frogs, 28 % of reptiles and 17 % of birds use and

depend on tree hollows for nesting and roosting. Many of these trees may

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take up to 200 years or more to form hollows that are big enough to support

this array of fauna (ARCS 1999).

These habitat trees that can be found in urban forested areas might play an

important role as vital habitats to the UK’s invertebrate fauna, as well as

providing important corridors for other wildlife.

2.0 Urban and Agricultural Trees history and uses

The term urban forest can be defined as ‘’all areas comprising all tree

dominated green areas in and around urban areas’’ (Konijnendijk et al 2004).

This can encompass parks, cops, as well as certain stately home gardens and

arboretums. Gardens/arboretums are collections of trees and plants that have

been planted for recreation and enjoyment and not necessarily for any

scientific use. They received much popularity in the Victorian period among

the aristocracy and as a result the UK has a large collection of

gardens/arboretums housing in some cases up to 3000 different taxa of trees

(Forestry commission national arboreta 2008).

The 1800’s was an important time in the UK’s history botanically. It was about

this time that the plant hunters were out in the furthest reaches of the British

Empire collecting and acquiring plant samples that may have potential uses

and sending them home (Musgrave et al 1998). The UK’s tree collections

house a considerable amount of tree taxa not just from abroad but also from

England itself. Many stately homes have large oaks in there grounds as well

as other species of tree for example RHS Rosemore in Devon has 182

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species representing 36 genus (RHS Rosemore personal contact 2008, for

full list see appendix 1). It is these trees that are of interest and importance

when it comes to invertebrate diversity as most of these trees now are

approaching ages of 150 years or more, they start to become habitat trees

and play host to many invertebrates as well as the species that prey upon

them. Urban forests are used all over the world in a variety of ways in the

west they are mainly for aesthetic and recreation uses, although there is

growing evidence towards their uses in carbon capture and pollution filters in

cities (Nowak et al 2006, McHale et al 2007). In other parts of the world

especially areas of Southeast Asia and parts of Africa they are important

sources of fuel wood and timber, as well as non wood products such as fruit

and mushrooms. In rural Africa farmers regularly plant and maintain trees that

help to enhance food, fuel, and medicines this is especially true of low income

farmers in poorer rural areas (Thompson et al 2004, Boffa et al, 2008).

Other trees of importance although not strictly urban are stand alone trees in

agricultural systems. Stand alone trees are a common feature all over the

world in agricultural systems they provide important habitats for many

invertebrates. Most of the terrestrial insect orders can be found living in trees

as many rely on the trees foliage for food. These are known as phytophagus

insects these include species of the orders Lepidoptera (the butterflies and

moths), the main consumption of tree foliage is carried out in there larval

stage, as adults they then rely on trees for suitable hiding places from

predators as well as for egg laying, Coleoptera (the beetles) both adults and

larva feed on tree foliage as well as fruits seeds and even in some cases

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roots. Species such as the stag beetles (Lucanidae) rely on dead wood from

trees to continue out there life cycles, Hemiptera/Homoptera (the true bugs)

although they lack biting mouth parts they can still use their proboscis to feed

off the trees sap the most common example of this is with the aphids

(Aphidoidea). The Orthoptera (the grasshoppers and crickets) have evolved

strong mandibles which are used to eat a wide variety of foliage and leaf

cover.

As a result of this abundant insect food source many predatory insects and

spiders hunt for insects within trees. Parasatoid wasps (Hymenoptera) such

as the Ichneumonoidea and the Braconidae use the larval stages of

Lepidoptera species as hosts for their own larva. As well as playing host to

many insect orders they also provide ecological services and functions, they

have been stated as keystone structure because of there ecological

importance relative to there abundance (Gibbons et al 2008). Evidence shows

that the presence of stand alone trees especially oak can help to boost local

invertebrate biodiversity especially with the order coleoptera (Ohsawa 2007).

Urban and stand alone trees may also play an important role as wildlife

corridors. Wildlife corridors are strips or patches of habitat that permit

dispersal between reserves or larger habitats (Hambler 2004). Corridors can

be seen to have several important advantages they can be used by species to

‘’top up’’ populations that are too small to be viable on their own, they can also

help by bringing in new genetic stock to a population (Hilty et al 2006). Wildlife

corridors can also act to help boost biodiversity. Reviews have been made of

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the usefulness of wildlife corridors and have found that they are beneficial in

most cases and could be found to do no harm to ecological systems (Beier et

al 1998).

Urban forests can be seen as wildlife corridors as they can potentially house

many species of invertebrate these are in turn food for other species of birds

and animals. Different species can use these trees and parks as stepping

stones in order to disperse further afield and into other habitats.

3.0 Trees as habitats

Trees can be considered an ecosystem in themselves and can act as small

ecological islands even from crown to crown or species to species in a forest,

thus stand alone trees can be considered ecological islands (Müller et al

2007). This can be advantageous when it comes to sampling and collecting

data on tree dwelling arthropods as trees discrete habitats were the total

invertebrate fauna can be sampled although it is not with out its difficulties or

problems (Southwood et al 2004).

One such example of this is the English oak (Quercus robur), it has been

estimated to have 284 (see Table 1) insect species associated that totally or

partially depend on the oak as food, shelter or as overwintering sites as adults

or eggs/larvae (Feltwell 1989, Southward 1961, Boreham 1999). The oak can

also have well over 300 species of lichen associated. In one year an oak can

be host to 110 butterfly species as they live out their lives in and around the

oak. It is no wonder that the oak tree has the most associated insect species

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of any tree in Britain (Kennedy et al 1984) but why is this? This can be partly

due to the amount of time that oak species have been in Britain the English

oak (Q. robur) has been present in England since the Pleistocene period as a

result of its occurrence in Britain for this amount of time more species will

have adapted to use its available niches (see Table 2) (Feltwell 1989,

Southwood 1961,). Even dead wood standing or fallen is a valuable habitat to

many invertebrate species this is especially true of broad leaved tree species

(Wu et al 2008).

Table 1: Broad leaf and coniferous* tree species and numbers of associated insects species.

Tree spp Numbers of associated insectsOak (Quercus) 284Willow (Salix) 266Birch (Betula) 229Hawthorn (Crataegus) 149Poplars (Populus) 97Apple (Malus) 93*Pine (Pinus) 91Alder (Alnus) 90Elm (Ulmus) 82Hazel (Corylus) 73Beech (Fagus) 64Ash (Fraxinus) 41*Spruce (Picea) 37Lime (Tilia) 31Hornbeam (Carpinus) 28*Larch (Larix) 17*Fir (Abies) 16Holly (Ilex) 7

(Adapted from: The Number of Species of Insect Associated with Various Trees, Southwood 1961)

The oak (Q. Robur) can be split into 11 habitat zones which are used by animal’s plants or fungi they are as follows: (see Figure 1).

These habitats afford a range of opportunities for colonization and temporal use by numerous flora and fauna (see Table 3).

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Table 3: Niche and occupant and uses in the 11 habitat zones of a tree

Niche Occupant / uses

Canopy: This area of crown is used by many insects as an assembly area especially by purple emperor butterflies (Apatura iris) and long horn moths (Adela spp) as well as providing food for some phytophagus insects and larvae (Feltwell 1989).

Buds: The ‘’sticky buds’’ that are produced in the spring are use as food by small insects as well as providing vital egg laying sites for the purple hairstreak butterfly (Quercusia Quercus) (Feltwell 1989).

Catkins: Produced by both male and female flowers these provide homes for many of the tiny cynipid wasps as well as the weevil beetles (Curculionidae).

Acorns: The caterpillars of the tortricid moth (cydia splendana) are internal feeders of acorns many of the cynipid wasps also lay there eggs in the acorns.

Leaves: Many insect larvae and weevil species feed on oak leaves, as well as the oak tortrix moth (Tortrix virdiana).

Twigs: Sawfly (symphyta spp) larvae bore into oak twigs also the winter moth (Operophtera brumata) over winter there eggs on twigs.

Trunk: Peppered moths (Biston betularia) rely on there camouflage to blend into the rough fissured trunks as they rest during the day to avoid predation. Lichens and mosses grow upon oak trunks and form another micro habitat for other arthropod species such as the arachnids, isopods, and Collembola.

Rot holes: Rot hole form where limbs of the tree break away and the exposed area collects water that cannot drain away, while this is dangerous for the tree as a possible route for infection it does provide an important breeding pool for many fly diptera species especially the mosquitoes (culicidae). They also provide water holes for bird species.

Macro fungi: These fruiting bodies are signs of internal fungal infection within the tree, these show themselves in the form of bracket fungi which are used by diptera species as a mating

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platform.Deadwood: As the upper branches develop and prosper

some of the lower limb start to die off and form deadwood which then fall to the ground, this is then used by host of forest floor arthropods. Such as stag beetles (lucanidae spp).

Roots: These are out of site and can also be out of mind but they play host too many bacteria and fungi especially the mycorrhizal fungi associated with the oak.

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Figure 1: Available niches of a tree (Q ilex) to fauna

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Roots

Canopy

Buds

Catkins

Acorns

Twigs/branches

Trunk

Rot-holes

Deadwood

Macro fungi


4.0 Canopy Invertebrates:

Arthropods are some of the most numerous creatures on the planet there are

to best approximations 1065,000 described arthropod species in the world

(Gaston & Spicer 1998). Arthropods carry out many vital ecosystem functions

they are as E.O. Wilson put it "little things that run the world,’’ (Wilson, 1987).

Trees are home to a variety of arthropods especially insects with most

terrestrial orders represented from some 90 families (Southwood et al 1982),

on the whole the species of insects living on and in trees are confined to a

small period of seasonality either for mating or hatching/feeding in the

summer and spring months with a slackening period of less abundance in the

winter due to death migration or hibernation/reduced activity (Burley et al

2004, Recher et al 1996, Southwood et al 2004).

Other factors that can affect the abundance of arthropod species in trees is

the morphological features of the tree being sampled (Southwood et al 1982).

For example the long thin leaves of some willow species offer less leaf

surface area (potential resource) compared to the large broader leaf surface

of oak species.

At present we have a very limited knowledge about the abundance and

makeup of arthropod communities in the canopy environment. This can

possibly be put down to the way in which canopy invertebrates are sampled.

Arthropods that are found in epiphytes and also in the collected leaf litter on

branches and in hollows can be seriously under represented in collections as

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a result the abundance of several orders such as Acari, Collembola, and

Isoptera are grossly underestimated. A wide range of techniques is needed in

order to fully ascertain the numbers and complexity of canopy arthropod

communities. Although having stated this forest canopies show a high level of

insect diversity from the tropics through to temperate biomes although this

depends on the tree species sampled from (Lowman et al 1995, Southwood

1961, 1982, Basset et al 2001).

5.0 Sampling canopy inverts Methods and Difficulties:

Currently there are many techniques that can be employed in order to

establish and estimate arthropod abundance. But as many techniques as

there are, they all have there limitations such as time to set up, cost, and

difficulty in accessing the sample site. (Gibb et al 2006, Carrel 2002).

The most common and widely used method of sampling canopy invertebrates

is with insecticide fogging (Ellwood 2006), a knock down insecticide that is

released into a canopy this then kills all the invertebrates and they then fall to

the floor and can be sampled with the use of funnels and sheets that have

been laid out. While this is a versatile and effective technique it cannot

provide a complete inventory of species for example some species that are

living in plant tissue can go unrecorded due to the protection from the knock

down insecticide offered by the vegetation (Yanoviak 2003). Fogging was first

used in 1951 by Collyer (Colleyer, 1951 cited in Southwood et al 2004)

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subsequently most of the sampling studies have been based on this one

occasion.

Light traps or any bright light will attract adult moths as well as some other

nocturnal flying insects; this form of trapping usually takes the form of high

pressure mercury vapour bulb suspended in a box topped with a hollow cone

for insects to drop through (Sutherland 2006). While this is great for nocturnal

species it doesn’t give a representation of diurnal species.

Pheromone traps are also a very effective way of capturing insects from

canopies, although they tend to be specifically designed to target certain

species. Many insect pheromones have been identified and synthesized they

are proven useful for the major insect orders Lepidoptera, Coleoptera,

Homoptera, Diptera, Hymenoptera, Isoptera, Blattodea, and Hemiptera

(Pimentel et al 2007). Some work is currently being carried out into multi

species lures (Collins et al 2007) that can attract a variety of insects, but until

this lure system is better developed lure traps will remain a species specific

way of capturing insect species.

Arial pitfall traps are a novel way to catch canopy invertebrates. These traps

are especially good at sampling aerial insects. They can be cheap and easy

to use, and can be made from household products such as plastic bottles,

James E Carrel developed a simple yet effective aerial trap from four plastic

drink bottles for a cost of $1-2 (2002). The trap consisted of four bottles with a

17cm wide × 13cm high strip removed to allow insects to fly in that were then

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attached to a ply wood platform. The traps can then be suspended from a tree

or over scrub to access arthropod numbers. Collection of samples was

through the use of a meat baster to suck up the insects, which were then

filtered through a tea strainer and put into a cup with lid to be taken back to

the lab (for design see figure 2). Due to the simplicity of the design Carrel was

able to empty and set a dozen traps in less than an hour. With some

modification this trap system may also be able to give an insight into other

tree dwelling species without the need to use a knock down fog which will be

beneficial for the tree invertebrate communities as well as the surrounding

area.

Figure 2: Carrel’s aerial interception trap

(Adapted from: A novel aerial interception trap for arthropod sampling, Carrel 2002.)

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6.0 Implications:

From the break down of organic matter to forming the starting points in many

food webs the role of canopy invertebrates are numerous and varied. Yet

there is a risk of loosing invertebrate diversity before we even know they were

there due to the rate of logging and conversion to agriculture there is an

increased risk of loosing the very forests that hold these amazing creatures.

The world’s tropical forested areas are being lost at a current rate of 1–4%

annually, (Basset et al 2004).

In order to understand the complex web of biodiversity and ecological roles

both the canopy and associated micro fauna have on both ecological function

and future exploitation the scientific community needs to be able to

demonstrate detailed understanding of what is in the canopies in order to plan

the correct action to take with regards to management (Myers 2001)

There are many implications for elucidating the invertebrate assemblages in

trees, for example trees may house beneficial insects for crop pest

management, many parasitic and parasatoid wasps live out there lives in and

on trees and forage for potential hosts for there eggs. Through knowing the

species of tree that many of these parasatoids favour, planning and planting

hedgerows could use this knowledge and incorporate it into planting regimes

to aid agricultural ecosystems. One such known beneficial insect is the

weaver ant (genus Oecophylla). Large colonies of Oecophylla by necessity

have to consume significantly large amounts of food; this takes the form of a

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http://en.wikipedia.org/wiki/Genus


variety of arthropods more often than not insects. Insects not only provide

food for the workers but are also used as a protein source necessary for the

development of the larval brood. The weaver ant workers hunt and kill a verity

of insects some of which are potentially harmful plant pests. Tree species that

harbor weaver ant nests also benefit from having decreased levels of

herbivory. There has been a long history of the use of Oecophylla as a

biological control stemming from China and Southeast Asian in the citrus

orchards from 400 AD (VanMele et al 2001, 2008).

Through knowing the ecology of tree dwelling insects you may be able to

predict the possibilities of surges in tree crop pests with regards to possible

population surge factors, climate or precipitation etc. This knowledge could

help forest managers manage and control pests efficiently by targeting

specific known species rather than a broad sweep insecticide to kill off all

invertebrates.

7.0 Study Aims

The aim of this proposed study is to try and evaluate the biodiversity of tree

canopies in three tree species, (Pedunculate oak; Quercus robur, Holm oak;

Quercus ilex and the common English beech; Fagus sylvatica) which are

found in the garden and grounds of Saltram house on the out skirts of the city

of Plymouth. It is proposed to use aerial pitfall traps as to coincide with data

collected already using this method, so a comparison in biodiversity can be

made between urban forest trees and farmland forest trees.

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http://en.wikipedia.org/wiki/Herbivory

http://en.wikipedia.org/wiki/Protein

http://en.wikipedia.org/wiki/Arthropods


Appendix 1

SOUTH ARBORETUM TREES – Alphabetical list

Abies cephalonica Carpinus betulus AGM

Abies concolor var. lowiana Carpinus carolinianaAbies fargesii Carya cordiformis AGM

Abies magnifica Castanea sativa AGM

Abies nordmanniana AGM Catalpa fargesiiAbies pindrow Catalpa speciosaAbies procera AGM Cedrus deodara AGM

Abies spectabilis Cedrus libani AGM

Abies veitchii AGM Cercidiphyllum japonicum var. magnificumAcer campestre AGM Cercis siliquastrum AGM

Acer capillipes ‘Candy Stripe’ Chamaecyparis lawsonianaAcer cappadocicum Chamaecyparis nootkatensisAcer carpinifoilum Chamaecyparis obtusaAcer circinatum AGM Clethra delavayi AGM

Acer cissifolium Cornus ‘Eddie’s White Wonder’ AGM

Acer davidii ‘Madeleine Spitta’ Cornus kousa ‘Milky Way’Acer giraldii Cornus kousa var. chinensis ‘China Girl’Acer macrophyllum Cornus macrophyllaAcer maximowiczianum Cornus nuttallii ‘Colrigo Giant’Acer pensylvanicum AGM Crataegus laevigataAcer platanoides AGM Crataegus phaenopyrumAcer saccharum subsp. nigrum Crataegus pinnatifida var. majorAcer sempervirens Crataegus tanacetifoliaAcer spicatum Cryptomeria japonica AGM

Acer sterculiaceum Davidia involucrata var. vilmoriniana AGM

Acer trautvetteri Fagus crenataAcer triflorum Fagus orientalis Acer velutinum var. vanvolxemii Fraxinus americanaAesculus californica Fraxinus mandshuricaAesculus flava AGM Ginkgo biloba AGM

Aesculus glabra Halesia monticola var. vestita AGM

Aesculus hippocastanum AGM Juglans nigra AGM

Aesculus indica AGM Larix decidua AGM

Aesculus pavia AGM Larix kaempferi AGM

Aesculus turbinata Larix occidentalisAlnus japonicaAmelanchier lamarckii AGM

Liquidambar formosana Monticola Group

Betula alleghaniensis Liquidambar styracifluaBetula ermanii ‘Grayswood Hill’ AGM Liriodendron chinenseBetula lenta Liriodendron tulipifera AGM

Betula mandshurica var. japonica Magnolia acuminata ‘Whitespire Senior’ Magnolia asheiBetula medwediewii AGM Magnolia campbelliiBetula papyrifera Magnolia campbellii ‘Alba’Betula pendula ‘Tristis’ AGM Magnolia campbellii ‘Darjeeling’Betula pubescens Magnolia campbellii subsp. mollicomata

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Betula szechuanica Magnolia dawsonianaBetula utilis Magnolia kobusBetula utilis var. jacquemontii ‘Silver Shadow’ Magnolia obovataCalocedrus decurrens AGM

Magnolia officinalis var. biloba Pterostyrax hispida AGM

Magnolia salicifolia AGM Pyrus nivalisMagnolia sargentiana var. robusta Pyrus ussuriensisMagnolia sieboldii var. sinensis AGM Quercus alienaMagnolia tripetala Quercus bicolor Malus baccata var. mandshurica Quercus canariensis AGM

Malus coronaria Quercus coccineaMalus florentina Quercus ellipsoidalisMalus floribunda AGM Quercus frainettoMalus sikkimensis Quercus glanduliferaMalus trilobata Quercus libaniMalus yunnanensis var. veitchii Quercus macrantheraMetasequoia glyptostroboides AGM Quercus marilandicaNyssa sinensis AGM Quercus palustris AGM

Nyssa sylvatica AGM Quercus pontica Ostrya carpinifolia Quercus velutinaPicea abies Sequoia sempervirens AGM

Picea brachytyla Sorbus americanaPicea glauca Sorbus ariaPicea likiangensis Sorbus aucupariaPicea omorika AGM Sorbus commixta ‘Embley’ AGM

Picea smithiana AGM Sorbus cuspidataPinus armandii Sorbus devoniensisPinus bungeana Sorbus harrowianaPinus densiflora Sorbus insignisPinus gerardiana Sorbus sargentiana AGM

Pinus jeffreyi AGM Sorbus vilmorinii AGM

Pinus nigra AGM Staphylea colchica AGM

Pinus peuce Stewartia malacodendronPinus thunbergii Stewartia monadelphaPinus wallichiana AGM Stewartia ovataPlatanus orientalis AGM Stewartia pseudocamellia var. koreana AGM

Populus balsamifera Stewartia sinensis AGM

Populus szechuanica Styrax japonicus AGM

Prunus avium AGM Styrax obassia AGM

Prunus hirtipes Thuja plicataPrunus jamasakura Tilia cordata AGM

Prunus maackii Tilia henryanaPrunus mahaleb Tilia oliveriPrunus padus Tsuga canadensisPrunus rufa Tsuga heterophylla AGM

Prunus serotina Tsuga mertensianaPrunus serrulata var. hupehensis Zelkova carpinifoliaPseudotsuga menziesii AGM

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Canopy inverterbrate diversity, Literature Review, Duncan Allen