Fistulina hepatica’s anamorphic version: Confistulina

Having been informed earlier by a friend that they have an article coming out soon in the journal Field Mycology and having then read the article, I considered it important to build on the information and photos shared in the manner I am most familiar: ramblings. I do not want to spoil the article so please do source the article yourself, though as a form of executive summary, this fungus, Confistulina hepatica, is the anamorphic (asexual) stage of the fungus Fistulina hepatica (beefsteak), which is common on oak and sweet chestnut. I have found it once on beech, though it generally sticks to the first two hosts. This anamorphic stage produces asexual spore, which adorn the fruiting body’s outer portion in abundance. The reason for its emergence is not known, though it might be weather-related. More information is absolutely required from across its host range – not just England!

Disclaimer: The older finds, because I didn’t know what I was looking at, are quite poor photographs. A huge shame, but alas!

Disclaimer 2: I suspect many of these are the anamorphic stage Confistulina hepatica, though none were confirmed so please do not assume they all are. This blog post is simply to begin building a knowledge base that builds of the limited resource pool of present.

The first oak that I’ll share is perhaps one of the more interesting of the bunch because, two years in a row (2015 and 2016), the outwardly anamorphic fruiting body appeared in the same location; albeit, in 2015, the fruiting body was larger. Nonetheless, it infers that, in a crude manner, the same mycelial colony has done the same thing two years in succession. Unfortunately, during 2015, I didn’t get a shot of the whole tree so the location of the arrow points to the 2016 occurrence. However, from the positioning of the ivy, we can see the location is the same.

Confistulina hepatica Fistulina anamorphic oak Quercus 1
The arrow points to the location, which is on the southern side of the tree right at the base.
Confistulina hepatica Fistulina anamorphic oak Quercus 2
The 2015 version in its rotten glory!
Confistulina hepatica Fistulina anamorphic oak Quercus 3
The fruiting body sports what looks like burns, in addition to exuding a tar-coloured liquid alongside the more routinely observed reddish exudations common to young fruiting bodies.
Confistulina hepatica Fistulina anamorphic oak Quercus 4
A closer look at the darker liquid from the 2015 version.
Confistulina hepatica Fistulina anamorphic oak Quercus 5
And a puncture wound (so it appears) in the fruiting body.
Confistulina hepatica Fistulina anamorphic oak Quercus 6
And the smaller 2016 emergence!
Confistulina hepatica Fistulina anamorphic oak Quercus 7
Again, we can see the darker liquid exudations. The morphology is also somewhat similar.
Confistulina hepatica Fistulina anamorphic oak Quercus 8
Closer still on the 2016 find.

The next series of shots were taken on my mobile phone back in 2015 and again from oak. Once more, the adornment of the fruiting body with exudations, of which some are darker, can be observed. 2016 saw the fungus return, with a vengeance, and in a different position, again, an anamorphic fruiting body.

Confistulina hepatica Fistulina anamorphic oak Quercus 9
Yes, this oak has been hammered. Poor tree! (2015)
Confistulina hepatica Fistulina anamorphic oak Quercus 10
At the base we can see the fruiting body.
Confistulina hepatica Fistulina anamorphic oak Quercus 11
Oozing everywhere!
Confistulina hepatica Fistulina anamorphic oak Quercus 12
Looks like something out of Alien, quite honestly!
Confistulina hepatica Fistulina anamorphic oak Quercus 34
2016 checking in. What a load of rubbish!
Confistulina hepatica Fistulina anamorphic oak Quercus 35
Further round the tree (note that the 2015 location was to the right) sits an anamorphic fruiting body.
Confistulina hepatica Fistulina anamorphic oak Quercus 36
A close(ish) look.
Confistulina hepatica Fistulina anamorphic oak Quercus 37
Closer still.
Confistulina hepatica Fistulina anamorphic oak Quercus 38
And closer yet further.

From here, we go to another 2015 find. This time, the fruiting body was a little elevated on the oak and evidently emanating from an area of burring / accumulation of dormant buds beneath the bark surface. This one quickly became senescent after perhaps a week so whether it’s Confistulina or not is tough to say – I include photos of both times I visited it. It did not reappear in 2016.

Confistulina hepatica Fistulina anamorphic oak Quercus 13
The first visit to this fungus, which was very small – maybe 5cm across.
Confistulina hepatica Fistulina anamorphic oak Quercus 14
A closer look.
Confistulina hepatica Fistulina anamorphic oak Quercus 15
A week later it had senesced.

The remainder of the photos take us into 2016 and most (but not all!) are taken with a better camera, which is good! First, we venture down to the New Forest and look at a well-decayed oak log, upon which a cluster of fruiting bodies sit atop the log.

Confistulina hepatica Fistulina anamorphic oak Quercus 16
Yes, yes, this photo was taken with a potato.
Confistulina hepatica Fistulina anamorphic oak Quercus 17
A very odd form but once again we can see the tarry liquid.
Confistulina hepatica Fistulina anamorphic oak Quercus 18
A side profile. Looks like a toe!
Confistulina hepatica Fistulina anamorphic oak Quercus 19
Yep, definitely a toe – a wrangled one, at that.

The next series are taken a month apart (late August and late September). What’s curious with this one is that we can see two anamorphic fruiting bodies and, come September, Laetiporus sulphureus fruiting directly alongside. The host, as can be seen, is oak, which is in a hedgerow of oak and ash.

Confistulina hepatica Fistulina anamorphic oak Quercus 20
Fading light caused the blur!
Confistulina hepatica Fistulina anamorphic oak Quercus 21
At the base we can see a dueo of fruiting bodies.
Confistulina hepatica Fistulina anamorphic oak Quercus 22
The first…
Confistulina hepatica Fistulina anamorphic oak Quercus 23
…and the gruesome second.
Confistulina hepatica Fistulina anamorphic oak Quercus 25
A month later a wild chicken appeared!
Confistulina hepatica Fistulina anamorphic oak Quercus 26
Two, in fact. Here we see the beefsteak and chicken side-by-side.
Confistulina hepatica Fistulina anamorphic oak Quercus 27
And here it’s in the distant murk.

Now, we move to a curious case of two fruiting bodies next to one another, again on oak, where one became a teleomorph and the other an anamorph. Why? Who knows. Very interesting, however, as I am sure you can appreciate.

Confistulina hepatica Fistulina anamorphic oak Quercus 28
An oak pollard by a swing. Look at the old pollard head.
Confistulina hepatica Fistulina anamorphic oak Quercus 29
A duo forming.
Confistulina hepatica Fistulina anamorphic oak Quercus 30
Looking pretty similar. Now watch…
Confistulina hepatica Fistulina anamorphic oak Quercus 31
…they change! No longer are they twins.
Confistulina hepatica Fistulina anamorphic oak Quercus 32
Exactly why this happened is a mystery though I’d really like to know – anamorph left) and teleomorph (right).
Confistulina hepatica Fistulina anamorphic oak Quercus 33
A side profile for comparison with the earlier one taken from the same position.

In October of 2016, I came across this majestic oak in a field. On a buttress root was what appeared to be an anamorphic fruiting body of Fistulina hepatica, which we can observe below.

Confistulina hepatica Fistulina anamorphic oak Quercus 39
Yup – majestic.
Confistulina hepatica Fistulina anamorphic oak Quercus 40
Also majestic??? (not sure!!)
Confistulina hepatica Fistulina anamorphic oak Quercus 41
Eh, nope. A bit more ugly, to be fair. Has the outward character of an anamorph though please don’t assume it is (wasn’t confirmed via microscopy).

Lastly, here’s one I suspect on sweet chestnut. I didn’t return to check how it did, though this year I’ll certainly keep more of an eye out and be more thorough in my inspections of the samples! Please share any examples you might have found of this, too – it’s important we build up a knowledge base.

Confistulina hepatica Fistulina anamorphic sweet chestnut Castanea 1
A lovely old coppice stool.
Confistulina hepatica Fistulina anamorphic sweet chestnut Castanea 2
Here sits the sample.
Confistulina hepatica Fistulina anamorphic sweet chestnut Castanea 3
Whether this is or is not is tough to say, though it does resemble an anamorphic stage somewhat.
Fistulina hepatica’s anamorphic version: Confistulina

David Attenborough on Richmond Park, London

Nothing much need be added, in light of who is narrating. As a 20-minute long film, it’s something you can readily watch at any point where you have some time going spare. There’s a few good segments on trees, including on ancient trees, deer, deadwood and wood-decay fungi. Really a fascinating watch!

Support the Friends of Richmond Park here.

David Attenborough on Richmond Park, London

Trees, forests and warfare

As has been highlighted previously in this blog (the series on state forestry, for example), trees have been used to fund the gluttonous cogs of the war machine, across both time and space. Usually, this timber consumption has manifested from the progressive land acclamation and legislatory enforcement by the state, until large tracts of forest are state-owned; or private forests can be utilised by the state in times of political emergency. This post therefore focusses not on repeating what has previously been discussed, and instead investigates how the forests themselves have been used for the arts of war – as in, the forest as a site of battle, or for the preparation of one; not that the forest as a site of battle is to be desired, for any attacking force must expect the unexpected, and typical formations and approaches to warfare cannot be applied in the enclosed forest setting (Clayton, 2012). Of course, the prior blog posts I did on state forestry highlight how armed guerrillas in Indonesia and Zimbabwe used the forests for cover and ambush, though this aspect of forest use extends far beyond just these two examples.

Beginning somewhat close to home (for the author), it can be recognised how the New Forest, in the county of Hampshire, UK, was used by the British and American armies, during the Second World War (Leete, 2014). Because of its strategic location relative to the coast of continental Europe, residing along the south coast of England, and complete with nearby ports in Southampton and Poole, the New Forest was used as the first line of defence against any invading Germans coming over from France. For this reason, the forest was used by both the Intelligence Service, and also by thousands of troops who would constitute the defending force if enemy ground invasion did occur. Furthermore, the extensive forest cover provided camouflage for over 30,000 troops in the moths before D-Day (Operation Neptune) in 1944, and the surrounding heathlands acted as airfields and storage areas of military vehicles. In total, 20,000 acres of the New Forest were utilised by the resident forces, during the war, though much like how the forest suddenly filled with troops it also quickly emptied, and almost immediately after the D-Day landing at Normandy the New Forest once again became very sparsely populated.

eastern-warfare-school-brockenhurst-2
Troops training near to Brockenhurst, in the New Forest. Source: The New Forest Guide.

The Second World War, beyond its association with the New Forest, was the site of actual battle. One example is that of the Battle of Hürtgen Forest, which took place between the US and German forces through September 1944 to February 1945. Situated on the border of Germany and Belgium, the Germans occupied the forest because of its strategic importance to future offensives on the Rhine. Fearing that these German troops would eventually therefore support the front line, the US Army sought to take control of the forest to stall this pursuit. However, because the terrain was very uneven, the access routes through the forest to constituent villages were narrow and almost non-existent, the trees were very dense in many locations, and forest clearings sudden and sporadically occurring, support from tanks was not feasible, and navigating the forest was often challenging and certainly very risky. Subsequently, the US forces suffered losses of over 30,000 men (at times, entire units were lost), eclipsing those incurred by the Germans; in spite of their much larger size. Granted, the Germans also suffered huge losses (Rush, 2001). The forest was thus named ‘The Death Factory’, by the US troops (Whiting, 2000), and became the grave of many individuals from both sides of the conflict.

hurtgen
The 28th Infantry Division of the US Army journey through the intrepid forest on 2nd November 1944. Source: History Net.

Curiously, the close of the Second World War also saw forests treated almost as bounty or reparation; at least, in Germany. Following the defeat Germany suffered, the country was subsequently segmented into various zones: the south-west of Germany became the French Zone, whilst the southern and south-east segments were under control by the Americans, the northern and north-west overseen by the British, and the east and north-east by the Soviets. The purpose of this was to enable Germany to ‘repent’ its ‘sins’, and the occupiers – the Americans, British, French, and Soviets – could harvest the forests as they saw fit, as long as such harvests were not in excess of the reparation quotas detailed after the Potsdam Conference in the summer of 1945.

Unfortunately, as such quotas usually were far greater than the rate at which the remaining forests (many were in an alarming state of disrepair, commercially-speaking) of Germany could be replenished, the Soviet zone saw fourteen years’ worth of timber logged in just four years. Alongside the purging of these now Soviet-controlled forests, those foresters who were not drafted into the war effort by the German government at the time were forced to work as hard labourers in the forests, and the traditionally scientific method that was German forestry was quashed by the inexperienced Soviets. Similar unsustainable levels of forestry were undertaken in the other occupied areas of Germany, by the Allied governments (Nelson, 2005).

Beyond the Second World War, Clayton (2012) remarks that the forest has been the site of battle as early as 9 A.D. In this year, the forest of Teutoburg was to plague three Roman legions and their auxiliaries – who were ambushed by the allied local Germanic tribes after an uprising in the region – quite cataclysmically. In this case, the Roman legions were headed by the reportedly inexperienced commander Publius Quinctilius Varus, whilst the commander of the allied tribes was the Germanic nobleman known as Arminius, who had himself been trained by the Roman army and was in fact part of the Roman legions who were tasked to deal with the uprising of the local tribes, though quickly defected to lead the Germans into battle.

Under the order of Varus, who was persuaded by Arminius (who at this point in the saga was still in the Roman army and appointed as an officer), the Roman legions headed into the forest to attempt to quell the uprising; at which point Arminius defected, and gathered up to 50,000 Germans to fight against approximately 7,000 Roman troops and their horses (including the three legions of eighty men each). In this forest, the now-defected Arminius used the terrain (including steep slopes, fallen trees, and dense forest cover) to confuse and disorientate the armour-clad Roman legions and support troops, who at first became surrounded and then were torn apart by the nimble Germanic warriors equipped with lightweight weapons (such as darts) and, for close combat, broadswords and spears. Most Roman troops were killed within the forest, in the small units that fled in all directions after Varus (who committed suicide) declared a retreat, though some unfortunate individuals were enslaved and / or tortured by the Germans. Ultimately, this situation manifested because the Roman troops were geared for close combat in the open setting, and the clever use of the forest by Arminius and his warriors led to what can only be considered a Roman tragedy – a tragedy that would not have occurred, and in fact likely have been reversed, if the battle was undertaken in the open (Clayton, 2012; Murdoch, 2006).

zpage506
Varus is defeated within the forest of Teutoburg, as is depicted through this illustration. Source: Heritage History.

The use of trees during conflict has also given rise to their use for hanging and other forms of execution (Stone, 2008). Certainly a macabre aspect of how warfare – and on a broader scale acts of genocide – ties man to the arboreal world, it is nonetheless an important point to consider, as it highlights how the tree, as a tool, has uses that extend beyond those aforementioned. In the genocide that plagued Cambodia from 1975-1979, for instance, the Khmer Rouge, who were followers of the community party led by Pol Pot, are said to have thrown children against trees until they died – because trees were cheaper than bullets. In these cases, Tyner (2009) remarks, the children were executed because their parents were considered enemies of the state. Lynching in the US, between 1889 to 1930, constitutes another form of warfare; albeit more a form of societal warfare, which can occur even during peacetime. During this period, an estimated 3,724 individuals were lynched, and before usually being hung from a tree and displayed for all to see the pursued individual was tortured, humiliated, dragged, and sometimes burned in front of potentially many thousands of onlookers (Dutton, 2007). In the UK, trees have also been the site of hangings; for example, for the execution of ‘rebels’ – whatever this loose term was deemed to define at the time by the ruling powers (Barnes & Williamson, 2011).

Running concurrently to the very human dynamics of wars and forests, exist more ecologically-based aspects worthy of consideration in this section. Principally, and notably over the past decades, one can identify the desire to safeguard forest biodiversity during times of war, by incorporating forest conservation into military projects (Machlis & Hanson, 2008). As ascertained prior to this point, the demands placed upon the forest in such a period unrest is possibly incredibly great, and particularly when the forest is being harvested for its timber, is being cleared to flush out a hiding enemy or to remove a hiding place, or the war is taking place largely within the forest (Reuveny et al., 2010). In recent years, tropical forests over South America and Africa have been the site of armed conflicts between the state and drug cartels, rebels, or otherwise, and McNeely (2003) astutely observes that such forests and their ecosystems can therefore be considered victims of war. Where these forests are considered hotspots for biodiversity, the impact is certainly markedly more severe and concerning for the scientific community (Hanson et al., 2009).

However, war is not always bad for forests. Where armed conflicts drive the general populace away, if the forests are not being actively utilised for resource to fuel the conflict, then they can undoubtedly benefit from the sudden drop in human pressures. Of course, the displaced populace is not purged from existence, and therefore where refugee camps associated with the conflict are constructed within – or adjacent to – forests, there can be a huge spike in deforestation. A pertinent example of such a phenomenon is when the Rwandan civil war displaced large numbers of people, who settled in the Democratic Republic of Congo in refugee camps and caused over 300km² of deforestation to nearby forests (Machlis & Hanson, 2008).

References

Barnes, G. & Williamson, T. (2011) Ancient Trees in the Landscape: Norfolk’s arboreal heritage. UK: Windgather Press.

Clayton, A. (2012) Warfare in Woods and Forests. USA: Indiana University Press.

Hanson, T., Brooks, T., da Fonseca, G., Hoffmann, M., Lamoreux, J., Machlis, G., Mittermeier, C., Mittermeier, R., & Pilgrim, J. (2009) Warfare in biodiversity hotspots. Conservation Biology. 23 (3). p578-587.

Leete, J. (2014) The New Forest at War: Revised and Updated. UK: Sabrestorm.

Machlis, G. & Hanson, T. (2008) Warfare ecology. BioScience. 58 (8). p33-40.

Murdoch, A. (2006) Rome’s Greatest Defeat: Massacre in the Teutoburg Forest. UK: Sutton Publishing.

Nelson, A. (2005) Cold War Ecology: Forests, Farms, & People in the East German Landscape, 1945-1989. USA: Yale University Press.

Reuveny, R., Mihalache-O’Keef, A., & Li, Q. (2010) The effect of warfare on the environmentThe effect of warfare on the environment. Journal of Peace Research. 47 (6). p749-761.

Rush, R. (2001) Hell in Hürtgen Forest: The Ordeal and Triumph of an American Infantry Regiment. USA: University Press of Kansas.

Stone, D. (2008) The Historiography of Genocide. UK: Palgrave Macmillan.

Tyner, J. (2009) War, Violence, and Population: Making the Body Count. USA: The Guilford Press.

Whiting, C. (2000) Battle of Hürtgen Forest. UK: Spellmount.

Trees, forests and warfare

A trip to Aldenham Country Park – trees and fungi

With the weather remaining fair, in spite of the onerous musings spouted from the verbal orifices of the meteorological office, getting out at the weekend to explore new sites is still very much on the cards. Today, a group of us went over to Aldenham Country Park in north-west London, to search for interesting fungi on trees; as if a weekend would yield any other result!

We started the day by doing something socially reprehensible: bringing in fungi collections for display. As the below photos show, my collection is growing in extent, though is dwarfed in literal size by another collection, which essentially involves monster brackets that are, in some cases, still clinging to the very substrate that provided them with their life.

polypore collection fungi
My collection, consisting of fruiting bodies of fungi including the genus Ganoderma (top left), the genus Trametes (bottom left), Fomes fomentarius (top middle), the genus Phellinus (bottom right) and Coriolopsis gallica (bottom right).
polypore collection 2
Another collection, set up almost like a demonstration of the solar system (with the Perenniporia fraxinea on the poplar being the sun, of course!), including Fomes fomentarius (a monster one), Daedalea quercina and, as stated, the Perenniporia fraxinea on the poplar wood.

Before sharing some finds from today, it’s almost important to share some images of more cross-sectional decay as caused by Ganoderma pfeifferi. For those of you with a memory that stretches back beyond a mere seven days, you might recall a recent post I made showing a decay cross-section on a failed beech. Below, we see how the fungus’ activity within a branch stub of a beech has resulted in zonal decay, which is somewhat comparable to the other example shared recently – particularly, with regards to the rosing pattern.

Ganoderma pfeifferi internal decay 1
A tiny Ganoderma pfeifferi within the opening of a branch stub wound on beech.
Ganoderma pfeifferi internal decay 2
The cross-section of decay produced by the fungus.

And so, on with the walk we did, quite early on we wandered past an old poplar stump with some quite extensive Rigidoporus ulmarius decay. Indeed, as is quite routine with this fungus, the internal hollow was clad aplenty with small brackets, whilst the outside sported a much more sizeable fruiting body still in an active phase of its existence. Evidently, a new hymenium has recently been laid down, suggesting that this fungus is soon ready to begin producing spore for the coming season.

Populus Rigidoporus ulmarius stump decay 1
Rigidoporus ulmarius acting as a saprotroph on this senescent stump.
Populus Rigidoporus ulmarius stump decay 2
Quite a nice one, actually! Good morphology.
Populus Rigidoporus ulmarius stump decay 3
Looking inside the hollow, not only can we see that it is used as a bin, but also to house many small fruiting bodies of this fungus.

Very soon after this sighting, a fallen poplar log with Oxyporus populinus was discovered. I admit to only having seen this fungus twice, of which this find was one, so for me this was particularly exciting. In fact, the single fruiting body was rather massive and easily discernible by the quite brilliant tube layers separated by narrow bands of mycelium. Almost directly adjacent to this was a fruiting body of Ganoderma applanatum, as could be determined morphologically by the very thin cuticle atop the bracket (that is crushed easily and cuts very easily) and the extensive damage to the fruiting body, as caused by the yellow flat-footed fly Agathomyia wankowiczii.

Oxyporus populinus Populus log 1
A poplar log hides amongst ivy.
Oxyporus populinus Populus log 2
On one of the cut ends sits this large fruiting body of the fungus Oxyporus populinus.
Oxyporus populinus Populus log 3
The demarcations between each growth spurt are incredibly distinct, in this fungus.
Ganoderma applanatum Populus log 1
A fruiting body of Ganoderma applanatum also sat nearby, on the same log.
Agathomyia wankowiczii Ganoderma applanatum Populus log 3
Underneath, we can see the distinct gall structures caused by the yellow flat-footed fly.
Agathomyia wankowiczii Ganoderma applanatum Populus log 2
We can also see the internal damage caused by the fly, as it develops into its adult form and leaves to lay eggs elsewhere. The very thin upper cuticle can also be seen, which is thicker on Ganoderma australe.

Following the sighting of copious amounts of Daedaleopsis confragosa, our attention was then drawn to a rather sorry-looking beech tree over a well-used footpath. Upon close inspection, both Kretzschmaria deusta and the rhizomorphs of Armillaria mellea could be found, which certainly puts the longevity of this beech as is into doubt. To be honest, in all likelihood it’ll be monolithed, in order to still provide habitat but with the risk removed.

Kretzschmaria deusta beech Fagus Armillaria 1
It even leans over the footpath!
Kretzschmaria deusta beech Fagus Armillaria 2
Both the anamorphic stage of Kretzschmaria deusta and cambial necrosis caused by Armillaria mellea can be seen, in this image.
Kretzschmaria deusta beech Fagus Armillaria 3
Not looking good for this beech!

Around the proverbial corner (it was more like a ten minute trundle) from this beech stood a massive stump of an old poplar. In its prime, this would have been a tree operating on beast-mode, though is now far more modest in size. However, to make up for its literal demise, it now is host to the fungus Trametes gibbosa, which can be seen around one of the two stems.

Trametes gibbosa Populus stump 1
A fortress of nettles guards this poplar stump.
Trametes gibbosa Populus stump 2
Too bad they can’t defend against a zoom lens and / or walking boots and jeans!
Trametes gibbosa Populus stump 3
Some fresh brackets adorn the opposite side of the stump.
Trametes gibbosa Populus stump 4
Quite pretty, to be honest!

Delightfully, this stump also housed a bird nest, which I found only by pure chance when noticing what looked like chocolate mini-eggs! Tucked away impossibly well within a bark crevice was a small robin’s nest (I think), complete with four eggs. Hopefully, this stump will offer enough privacy to enable the chicks to develop well and not get picked-off by predators.

Erithacus rubecula eggs poplar stump tree 1
The arrow shows where the nest is, as it’d otherwise be impossible to see!
Erithacus rubecula eggs poplar stump tree 2
There were four eggs in this tiny nest. Such a great place for shelter and quite absurd that I came across it!

Once we had come across yet more Daedaleopsis confragosa, which I was busy photographing, a friend spotted a single Sarcoscypha coccinea (scarlet elf cup). Somehow, this is the first time I have seen this fungus and I can understand why it’s such a popular one! An absolute gem.

Sarcoscypha coccinea 1
Cheeky! Hiding away under nettles. Almost doesn’t want to be discovered…
Sarcoscypha coccinea 2
Nature’s very own satellite dish!

And then came something I found very interesting: my first ever sighting of the fungus of willow known as Phellinus igniarius. Upon what was either a crack willow or white willow, a few fruiting bodies had grown and the decay had since led to failure of an upper limb, which has since been cut up and left on the ground. The resulting abundance of fruiting bodies on both the tree and sawn logs is a testamenrt to the extensive colonisation of this fungus within the host. The largest bracket, which was a casulaty of the failure, in fact did not senesce and instead reiterated its growth so that the hymenium and tube layer re-grew at an angle perfectly parallel with the ground (known as geotropism / gravitropsim).

Phellinus igniarius Salix alba fragilis sp decay 1
A willow not unlike any other willow – battered by the elements.
Phellinus igniarius Salix alba fragilis sp decay 2
Oh but wait – a fungus! Surely it’s a Ganoderma…
Phellinus igniarius Salix alba fragilis sp decay 3
…nope!
Phellinus igniarius Salix alba fragilis sp decay 4
As we shall see by what is on the floor, upon these logs…
Phellinus igniarius Salix alba fragilis sp decay 5
…Phellinus igniarius! Surprise! (assuming you didn’t read the text and look only at the pictures)
Phellinus igniarius Salix alba fragilis sp decay 6
Quite a significant number of new sporophores are forming, following the fragmentation of this limb.
Phellinus igniarius Salix alba fragilis sp decay 7
Around an old branch tear sits a single fruiting body, however.
Phellinus igniarius Salix alba fragilis sp decay 8
Not unlike a young Fomes fomentarius, really!
Phellinus igniarius Salix alba fragilis sp decay 9
And the main bracket has not perished!
Phellinus igniarius Salix alba fragilis sp decay 10
Using flash photography (literally), we can see the white spore print beneath the reiterated growth, following the change in orientation of this bracket.

To round off, I share a diabolically grotesque example of Ganoderma resinaceum upon Turkey oak. Enough to challenge the gargoyle statues of various catacombs (in both video games and real life, if there exist any!) for the prize of what’s the most vile in appearance, and we’re not talking about the Turkey oak here, this fungus is clearly a shadow of its former self. Nonetheless, it is important we can still identify them in such aberrant form, if we are to appropriate diagnose issues and enact management regimes. Thus, as a sort of encore, I present to you…

Ganoderma resinaceum Quercus cerris weird 1
Nice enough tree, eh!
Ganoderma resinaceum Quercus cerris weird 2
But what is that at the base!?
Ganoderma resinaceum Quercus cerris weird 3
Uhh………??
Ganoderma resinaceum Quercus cerris weird 4
Yeah; uhhh…….?
Ganoderma resinaceum Quercus cerris weird 5
Ganoderma resinaceum!
A trip to Aldenham Country Park – trees and fungi

Trees in the ecosystem pt V: Trees & slime molds

Single-celled organisms that may create larger structures as groups in order to reproduce, slime molds, whilst not considered active wood decayers, can be found colonising deadwood (Heilmann-Clausen, 2001). Deadwood of 10-22 years of age, Heilmann-Clausen (2001) alleges, is most optimal for slime molds – at least, for the species observed on the decaying beech logs that featured within the study. This correlates with current understanding of slime molds, which suggests species strongly prefer moist, well-decayed wood.

Enteridium lycoperdon Pyrus
The false puffball (Enteridium lycoperdon) on the well-decayed remains of a pear (Pyrus sp.) stem.

The presence of wood-decay fungi sporophores, or even simply mycelium within the wood substrate, may also act as a source of energy for slime molds (Ing, 1994). As mycelial networks and their associated sporophores may take some time to develop within deadwood, this may perhaps be a further reason for why slime molds are found in greater abundance on older woody debris. The presence of bacteria, also greater in abundance on older and heavily-decayed wood, may also influence slime mold presence, as bacteria can be utilised as a further source of energy (Heilmann-Clausen, 2001). Lodge (1997) describes some slime molds as “predators of decomposers”. Slime molds may also utilise decaying leaves as a habitat (Ko et al., 2009; Raper, 1941; Raper, 1951; Stephenson, 1989). Therefore, the decaying leaf litter-soil ‘zone’ is another potential niche for slime mold species (Landolt & Stephenson, 1986). Moreover, slime molds may be found upon the bark of living trees (Olive & Stoianovitch, 1973; Stephenson, 1989).

Fuligo septica Betula
Fuligo septica, known commonly as ‘dog sick slime mold’ or ‘scrambled eggs’, growing on birch (Betula pendula).

Away from wood, decaying leaves, and soil exclusively, the composition of a forest ecosystem may also have an impact upon slime mold density. Landolt et al. (2006) found that, whilst species diversity did not differ between deciduous-broadleaved and coniferous stands, the broadleaved sites were host to slime mold populations over four times more abundant than coniferous sites. The same study also identified that different species of slime mold would be found at different altitude levels within forests, and suggested different micro-habitats perhaps act as refugia for different slime mold species that may have once colonised greater ranges of forest.

References

Heilmann-Clausen, J. (2001) A gradient analysis of communities of macrofungi and slime moulds on decaying beech logs. Mycological Research. 105 (5). p575-596.

Ing, B. (1994) Tansley Review No. 62: The phytosociology of myxomycetes. New Phytologist. 126 (2). p175-201.

Ko, T., Stephenson, S., Jeewon, R., Lumyong, S., & Hyde, K. (2009) Molecular diversity of myxomycetes associated with decaying wood and forest floor leaf litter. Mycologia. 101 (5). p592-598.

Landolt, J. & Stephenson, S. (1986) Cellular slime molds in forest soils of southwestern Virginia. Mycologia. 78 (3). p500-502.

Landolt, J., Stephenson, S., & Cavender, J. (2006) Distribution and ecology of dictyostelid cellular slime molds in Great Smoky Mountains National Park. Mycologia. 98 (4). p541-549.

Lodge, D. (1997) Factors related to diversity of decomposer fungi in tropical forests. Biodiversity & Conservation. 6 (5). p681-688.

Olive, L. & Stoianovitch, C. (1974) A cellular slime mold with flagellate cells. Mycologia. 66 (4). p685-690.

Raper, K. (1941) Dictyostelium minutum, a second new species of slime mold from decaying forest leaves. Mycologia. 33 (6). p633-649.

Raper, K. (1951) Isolation, cultivation, and conservation of simple slime molds. The Quarterly Review of Biology. 26 (2). p169-190.

Stephenson, S. (1989) Distribution and ecology of myxomycetes in temperate forests. II. Patterns of occurrence on bark surface of living trees, leaf litter, and dung. Mycologia. 81 (4). p608-621.

Trees in the ecosystem pt V: Trees & slime molds

Some fungal finds from the week

Some say it’s written in the stars, though the only experience I have had with braille is from select old Nintendo games from the 1990s and early 2000s (revealing my age a little here!). Others say it’s just annoying. I’d probably agree with the latter! Regardless, here we have it: more pictures of fungi on trees.

As always, I keep my eye out for some interesting finds. This week has been pretty decent on the fungal side of things, though given the time of year only the perennial polypores are really observable – asides from the odd Flammulina velutipes / elastica and some enterprising Pleurotus species. Nonetheless, for the sake of showcasing unique finds and for educational purposes, here are a few species of polypore and some common agarics.

Firstly, we have a rather cool deck of Ganoderma resinaceum brackets around a rather pronounced buttress on an oak (Quercus robur). The fruiting between the two buttress roots is likely indicative of good reaction growth that is well-compartmentalised, which in turn infers respectable and probably sound (i.e. free of appreciable decay) buttressing from which the oak is supporting itself. We then have some shots of a rather aberrant duo of Trametes gibbosa on what is probably an old sycamore (Acer pseudoplatanus) stump, some Kretzschmaria deusta on (again!) sycamore, Ganoderma australe on a fallen ash (Fraxinus excelsior) and finally some Flammulina sp. and Pleutorus ostreatus on a very decayed stump of an unknown deciduous broadleaved species.

ganoderma-resinaceum-quercus-robur-buttress-1ganoderma-resinaceum-quercus-robur-buttress-2ganoderma-resinaceum-quercus-robur-buttress-3

trametes-gibbosa-stump-acer-2trametes-gibbosa-stump-acer-3trametes-gibbosa-stump-acer-4

acer-pseudoplatanus-kretzschmaria-deusta-monolith-1acer-pseudoplatanus-kretzschmaria-deusta-monolith-2acer-pseudoplatanus-kretzschmaria-deusta-monolith-3

fraxinus-excelsior-ganoderma-australe-fallen-1fraxinus-excelsior-ganoderma-australe-fallen-2fraxinus-excelsior-ganoderma-australe-fallen-3fraxinus-excelsior-ganoderma-australe-fallen-4

flammulina-velutipes-pleutorus-ostreatus-stump-1flammulina-velutipes-pleutorus-ostreatus-stump-2flammulina-velutipes-pleutorus-ostreatus-stump-3

Some fungal finds from the week

Trees in the ecosystem pt IV: Trees & arthropods

The arthropods are vast in terms of species, and include ants, beetles, butterflies, mites, moths, spiders, and so on. Therefore, covering the entire spectrum of arthropods in this section is impractical, though the general provisioning by trees will be outlined and species will be used to illustrate given examples.

Many arthropods are considered to be saproxylic in nature – they principally utilise dead woody material (both standing and fallen, in both dead and living trees) as habitat, for at least part of their life cycle, though they may also rely upon fungal sporophores associated with the presence of deadwood, as is to be detailed below (Gibb et al., 2006; Harding & Rose, 1986; Komonen et al., 2000). Of all the saproxylic arthropods, beetles are perhaps the most significant in terms of the proportion occupied of total saproxylic species worldwide (Müller et al., 2010), though saproxylic flies also feature in great numerical abundance (Falk, 2014; Harding & Rose, 1986).

Beetles may be either generalist or specialist in nature (on either broadleaved or coniferous hosts), and they will normally require a host with an abundance of deadwood (or large sections of coarse woody debris) usually over 7.5cm in diameter that resides within an area typically not heavily shaded (Müller et al., 2010; Siitonen & Ranius, 2015). This may be, in part, due to many beetle species (in their adult stage) requiring nectar from herbaceous plants, which would be lacking in woodland with significant canopy closure (Falk, 2014; Siitonen & Ranius, 2015). This means that veteran trees amongst wood pasture and parklands (including in urban areas) may be particularly suitable (Bergmeier & Roellig, 2014; Harding & Rose, 1986; Ramírez-Hernández et al., 2014; Jonsell, 2012; Jørgensen & Quelch, 2014), though this is not at all a steadfast rule as species may also be found abundantly in (perhaps more open) woodland, and particularly where there are large amounts of veteran trees and deadwood – around 60 cubic metres per hectare, according to Müller et al. (2010). Granted, they are found particularly in older (mature to veteran) trees, including within cavities that possess wood mould, water-filled rot holes, dead bark, exposed wood, sap flows, fruiting bodies (of fungi and slime moulds), mycelia of fungi, dead branches, and dead roots (Carpaneto et al., 2010; Falk, 2014; Harding & Rose, 1986; Siitonen & Ranius, 2015; Stokland et al., 2012). Beetle species may also not necessarily associate preferentially with a species (or group of species), but with the conditions aforementioned that are present within a tree (Harding & Rose, 1986; Jonsell, 2012). At times, preferable conditions may be an infrequent as one veteran tree in every hundred (Harding & Rose, 1986).

veteran-oak-tree
A veteran oak tree that is of prime habitat for a variety of organisms.

Despite this, species preference is observed. For broadleaved obligates, heavier shade may be more necessary, and in such instances there is a closer affinity of the beetles with fungal mycelium. Because fungi tend to produce more mycelium in cooler and more humid conditions (though this does, of course, vary with the species), the broadleaved obligates may therefore be found normally in greater abundance where conditions are more suited to fungal growth, and their presence may thus be associated with a canopy openness of as little as 20% (Bässler et al., 2010; Müller et al., 2010). This is, of course, not a steadfast rule, and many open wood pastures may support a great abundance of saproxylic beetles (Harding & Rose, 1986).

It is also important to recognise that many species of saproxylic beetle are reliant upon particular stages of the wood decay process. For instance, species that require fresh phloem tissue will only be able to colonise briefly in the first few summers following on from the death of the phloem tissue (Falk, 2014). Other species require significantly-decayed wood in a particular micro-climate, and even of a particular tree species (Harding & Rose, 1986). There also exist intricate associations between species of fungi and saproxylic insects. Inonotus hispidus, which is usually found upon ash, is the habitat for Triplax russica and Orchesia micans, whilst the coal fungus (Daldinia concentrica), also oft found upon the deadwood of ash (Fraxinus excelsior), is the main provider of habitat for Platyrhinus resinosus (Falk, 2014). The birch polypore (Fomitopsis betulina) is also host to numerous species of Coleoptera (Harding & Rose, 1986); as is the polypore Fomitopsis pinicola (Jonsson & Nordlander, 2006; Komonen, 2003; Komonen et al., 2000). This means that these species may be found where there is a suitable population of the fungus’ host species, where sporophores are present and will likely fruit again in the future, across numerous trees, and for many years. Most beetle species rely on oak more so than other tree species however, as oak generally lives for much longer and thus provides a wider array of different micro-habitats, and possesses increased compositional complexity as a result (Harding & Rose, 1986; Siitonen & Ranius, 2015).

ancient_orchard_malus_inonotus_hispidus4
A fruiting body of Inonotus hispidus on apple (Malus sp.). This fungus not only creates habitat in the wood that it degrades but also is a direct habitat through its sporophore.

Therefore, the loss of suitable habitat through active management programmes (including logging, and felling trees for safety reasons in urban areas) will have a very adverse impact upon saproxylic beetles, though also certain species of moth, and even species associated with saproxylic insects, including parasitic wasps, solitary wasps (which use beetle bore holes for habitat), and predatory Coleoptera (Harding & Rose, 1986; Komonen et al., 2000). Curiously, research by Carpaneto et al. (2010) concluded that trees that were ranked as the most evidently ‘hazardous’ were host to the most saproxylic beetle species, and their removal would therefore have a drastic impact upon local populations. Similarly, fragmentation of woodland patches suitable for saproxylic populations has led to a decline in the meta-populations (Grove, 2002; Komonen et al., 2000), as has deadwood removal in a managed site itself (Gibb et al., 2006). Interestingly, though not surprisingly, ‘deadwood fragmentation’ also has an adverse impact upon saproxylic insect populations (Schiegg, 2000).

Both ants and termites also benefit from the presence of deadwood. With regards to both, nests will usually form at the base of a tree or at an area where there is at least moderate decay – enough to support a viable population (Jones et al., 2003; Shigo, 1986; Stokland et al., 2012). Ants and termites both follow CODIT (compartmentalisation of damage in trees) patterns in relation to how their nests progress, and thus their territory will increase as fungal decay propagates further into the host. Ants will not feed on the decaying wood of the host however, and will simply use the decaying site as a nesting area. Conversely, termites will feast upon decayed wood and essentially control (perhaps by slowing down) the spread of fungal decay in a manner that provides as much longevity of the host as possible for a viable nesting site (Shigo, 1986). In tropical rainforests, termites are in fact considered to be one of the principal means of wood decomposition (Mori et al., 2014), and thus the provisioning of deadwood habitat is absolutely critical. Without decaying wood within trees therefore, ants and particularly termites will lack a potential habitat, and thus where a stand is actively managed populations may be markedly reduced (Donovan et al., 2007; Eggleton et al., 1995). Of course, termites are not necessarily to be desired when they are invading the wood structure of a property, and therefore deadwood is not universally beneficial (Esenther & Beal, 1979; Morales-Ramos & Rojas, 2001) – at least, when human properties are involved.

termites_1_007
Ecologically beneficial? Yes. Economically beneficial? No. Termites can – and do – damage timber-frames buildings, as is the case here. Source: Pestec.

The presence of deadwood may also be beneficial for ground-nesting and leaf-litter dwelling spiders, which can utilise downed woody debris (particularly pieces with only slight decay) for both nesting and foraging (Varady-Szabo & Buddle, 2006). In fact, research by Buddle (2001) suggested that such spiders may more routinely utilise downed woody material when compared to elevated woody material (dead branches and telephone poles) because of the greater array of associated micro-habitats, and particularly at certain life stages – such as during egg-laying, for females (Koch et al., 2010). Furthermore, as fallen woody debris can help to retain leaf litter (or even facilitate in the build-up leaf litter), spider populations are more abundant and more diverse in sites where such woody debris is present (Castro & Wise, 2010). Therefore, where woodlands are managed and areas are clear-cut, spider populations may be markedly reduced in terms of the diversity of species. However, generalist species may benefit from the amount of cut stumps (Pearce et al., 2004). Curiously, Koch et al. (2010) suggest that spiders may perhaps benefit from woodland clearance, because the vigorous re-growth of trees and the higher light availability to the woodland floor (promoting herbaceous plant growth) increases the abundance of potential prey. Despite this, old-growth species will suffer (Buddle & Shorthouse, 2008), and thus the population structure of spider populations may dramatically change.

Soil mites are a further group that benefit from coarse woody debris, though also from hollows and holes throughout the basal region of a tree (including water-filled cavities), and from fungal sporophores and hyphae associated with wood decay (Fashing, 1998; Johnston & Crossley, 1993). Typically, termites will use fungi and insects found within the wood as a food source, and the wood structure itself will provide for an array of niche micro-habitats that are critical at different life stages of a mite. Certain mite species are obligates that associate with coarse woody debris exclusively, and may in fact only be associated with certain species’ woody debris. Additionally, mites may utilise woody debris and hollows within trees to parasitise upon other species using the ‘resource’, with both lizards and snakes being parasitised by mites following their frequenting of such resources. Beetles may also be parasitised, though the mite in such an instance may use the beetle as a means of entry into woody debris (Norton, 1980).

It is not just deadwood that arthropods will utilise, however. Foliage, both alive and abscised, is also of use (Falk, 2014). For example, the ermine moth (Yponomeutidae) will rely upon the living foliage of a host tree as a food source, and the bird cherry ermine moth (Yponomeuta evonymella) is one example of this. During late spring, larvae will fully defoliate their host Prunus padus, before pupating, emerging, and then laying eggs upon the shoots ready for the following year (Leather & Bland, 1999). Many other moth species will, during their larval stage, also behave in such a manner and thus defoliate their host – either entirely, or in part (Herrick & Gansner, 1987). Other species may alternatively have larvae mine into the leaf and feed upon the tissues within (Thalmann et al., 2003), such as horse chestnut leaf miner (Cameraria ohridella). Flies, including the holly leaf-miner (Phytomyza ilicis), will also mine leaves in a similar fashion (Owen, 1978). Ultimately however, the same purpose is served – the insect uses the living tissues of a leaf to complete its life cycle, and fuel further generations.

1280px-yponomeuta_evonymella_on_prunus_padus
Bird cherry ermine moth having defoliated an entire tree. Source: Wikimedia.

Fallen leaf litter, as briefly touched upon earlier when discussing spiders, may also be of marked benefit to many arthropods. Ants, beetles, and spiders are but three examples of groups that will utilise leaf litter as a means of habitat (Apigian et al., 2006). Beetles will, for instance, rely upon leaf litter to attract potential prey, though also to provide niche micro-climates that remain relatively stable in terms of humidity, light availability, and temperature (Haila & Niemelä, 1999). Their abundance may, according to Molnár et al., (2001) be greatest at forest edges, perhaps because prey is most abundant at these edge sites (Magura, 2002). Of course, this does not mean that edges created through artificial means will necessarily improve beetle populations, as research has shown that there are few ‘edge specialists’ and therefore populations usually will go into decline where there has been significant disturbance. Unless management mimics natural mortality events of forest trees, then constituent beetle populations may thus suffer adversely (Niemelä et al., 2007).

With regards to ants, Belshaw & Bolton (1993) suggest that management practices may not necessarily impact upon ant populations, though if there is a decline in leaf litter cover then ants associated with leaf litter presence may go into – perhaps only temporary (until leaf litter accumulations once again reach desirable levels) – decline (Woodcock et al., 2011). For example, logging within a stand may reduce leaf litter abundance for some years (Vasconcelos et al., 2000), as may (to a much lesser extent) controlled burning (Apigian et al., 2006; Vasconcelos et al., 2009), though in time (up to 10 years) leaf litter may once again reach a depth suitable to support a wide variety of ant species. However, the conversion of forest stands into plantations may be one driver behind more permanently falling ant populations (Fayle et al., 2010), as may habitat fragmentation (Carvalho & Vasconcelos, 1999) – particularly when forest patches are fragmented by vast monoculture plantations of tree or crop (Brühl et al., 2003). The conversion of Iberian wood pastures to eucalyptus plantations is one real world example of such a practice (Bergmeier & Roellig, 2014).

Also of benefit to many arthropods are nectar and pollen. Bees, beetles, butterflies, and hoverflies will, for instance, use nectar from flowers as a food source (Dick et al., 2003; Kay et al., 1984), and generally (but not always) a nectar source will lack significant specificity in terms of the insect species attracted (Karban, 2015). Despite this, different chemicals secreted by different flowers, and the toxicity of certain nectar sources to particular insects, means certain tree species may only be visited by certain insect species (Adler, 2000; Rasmont et al., 2005). Tree diversity may therefore be key to sustaining healthy insect populations (Holl, 1995), and where species may prefer to frequent herbaceous plant species the presence of a diverse woodland canopy above may still be very influential (Kitahara et al., 2008). This may be because a diverse array of woody plant species increases the diversity of herbaceous species. At times, pollen may also be a reward, as may (more rarely) a flower’s scent. Karban (2015) remarks that all are collectively dubbed as ‘floral rewards’.

References

Adler, L. (2000) The ecological significance of toxic nectar. Oikos. 91 (3). p409-420.

Apigian, K., Dahlsten, D., & Stephens, S. (2006) Fire and fire surrogate treatment effects on leaf litter arthropods in a western Sierra Nevada mixed-conifer forest. Forest Ecology and Management. 221 (1). p110-122.

Bässler, C., Müller, J., Dziock, F., & Brandl, R. (2010) Effects of resource availability and climate on the diversity of wood‐decaying fungi. Journal of Ecology. 98 (4). p822-832.

Belshaw, R. & Bolton, B. (1993) The effect of forest disturbance on the leaf litter ant fauna in Ghana. Biodiversity & Conservation. 2 (6). p656-666.

Bergmeier, E. & Roellig, M. (2014) Diversity, threats, and conservation of European wood-pastures. In Hartel, T. & Plieninger, T. (eds.) European wood-pastures in transition: A social-ecological approach. UK: Earthscan.

Brühl, C., Eltz, T., & Linsenmair, K. (2003) Size does matter–effects of tropical rainforest fragmentation on the leaf litter ant community in Sabah, Malaysia. Biodiversity & Conservation. 12 (7). p1371-1389.

Buddle, C. (2001) Spiders (Araneae) associated with downed woody material in a deciduous forest in central Alberta, Canada. Agricultural and Forest Entomology. 3 (4). p241-251.

Buddle, C. & Shorthouse, D. (2008) Effects of experimental harvesting on spider (Araneae) assemblages in boreal deciduous forests. The Canadian Entomologist. 140 (4). p437-452.

Carpaneto, G., Mazziotta, A., Coletti, G., Luiselli, L., & Audisio, P. (2010) Conflict between insect conservation and public safety: the case study of a saproxylic beetle (Osmoderma eremita) in urban parks. Journal of Insect Conservation. 14 (5). p555-565.

Carvalho, K. & Vasconcelos, H. (1999) Forest fragmentation in central Amazonia and its effects on litter-dwelling ants. Biological Conservation. 91 (2). p151-157.

Castro, A. & Wise, D. (2010) Influence of fallen coarse woody debris on the diversity and community structure of forest-floor spiders (Arachnida: Araneae). Forest Ecology and Management. 260 (12). p2088-2101.

Dick, C., Etchelecu, G., & Austerlitz, F. (2003) Pollen dispersal of tropical trees (Dinizia excelsa: Fabaceae) by native insects and African honeybees in pristine and fragmented Amazonian rainforest. Molecular Ecology. 12 (3). p753-764.

Donovan, S., Griffiths, G., Homathevi, R., & Winder, L. (2007) The spatial pattern of soil‐dwelling termites in primary and logged forest in Sabah, Malaysia. Ecological Entomology. 32 (1). p1-10.

Eggleton, P., Bignell, D., Sands, W., Waite, B., Wood, T., & Lawton, J. (1995) The species richness of termites (Isoptera) under differing levels of forest disturbance in the Mbalmayo Forest Reserve, southern Cameroon. Journal of Tropical Ecology. 11 (1). p85-98.

Esenther, G. & Beal, R. (1979) Termite control: decayed wood bait. Sociobiology. 4 (2). p215-222.

Falk, S. (2014) Wood-pastures as reservoirs for invertebrates. In Hartel, T. & Plieninger, T. (eds.) European wood-pastures in transition: A social-ecological approach. UK:     Earthscan.

Fashing, N. (1998) Functional morphology as an aid in determining trophic behaviour: the placement of astigmatic mites in food webs of water-filled tree-hole communities. Experimental & Applied Acarology. 22 (8). p435-453.

Fayle, T., Turner, E., Snaddon, J., Chey, V., Chung, A., Eggleton, P., & Foster, W. (2010) Oil palm expansion into rain forest greatly reduces ant biodiversity in canopy, epiphytes and leaf-litter. Basic and Applied Ecology. 11 (4). p337-345.

Gibb, H., Pettersson, R., Hjältén, J., Hilszczański, J., Ball, J., Johansson, T., Atlegrim, O., & Danell, K. (2006) Conservation-oriented forestry and early successional saproxylic beetles: responses of functional groups to manipulated dead wood substrates. Biological Conservation. 129 (4). p437-450.

Grove, S. (2002) Saproxylic insect ecology and the sustainable management of forests. Annual Review of Ecology and Systematics. 33 (1). p1-23.

Haila, Y. & Niemelä, J. (1999) Leaf litter and the small‐scale distribution of carabid beetles (Coleoptera, Carabidae) in the boreal forest. Ecography. 22 (4). p424-435.

Harding, P. & Rose, F. (1986) Pasture-Woodlands in Lowland Britain: A review of their importance for wildlife conservation. UK: NERC.

Herrick, O. & Gansner, D. (1987) Gypsy moth on a new frontier: forest tree defoliation and mortality. Northern Journal of Applied Forestry. 4 (3). p128-133.

Holl, K. (1995) Nectar resources and their influence on butterfly communities on reclaimed coal surface mines. Restoration Ecology. 3 (2). p76-85.

Jones, D., Susilo, F., Bignell, D., Hardiwinoto, S., Gillison, A., & Eggleton, P. (2003) Termite assemblage collapse along a land‐use intensification gradient in lowland central Sumatra, Indonesia. Journal of Applied Ecology. 40 (2). p380-391.

Jonsell, M. (2012) Old park trees as habitat for saproxylic beetle species. Biodiversity and Conservation. 21 (3). p619-642.

Jonsell, M. & Nordlander, G. (2004) Host selection patterns in insects breeding in bracket fungi. Ecological Entomology. 29 (6), p697-705.

Johnston, J. & Crossley, D. (1993) The significance of coarse woody debris for the diversity of soil mites. In McMinn, J. & Crossley, D. (eds.) Proceedings of the Workshop on Coarse Woody Debris in Southern Forests: Effects on Biodiversity. General Technical Report SE-94.

Jørgensen, D. & Quelch, P. (2014) The origins and history of medieval wood-pastures. In Hartel, T. & Plieninger, T. (eds.) European wood-pastures in transition: A social-ecological approach. UK: Earthscan.

Karban, R. (2015) Plant Sensing & Communication. USA: University of Chicago Press.

Kay, Q., Lack, A., Bamber, F., & Davies, C. (1984) Differences between sexes in floral morphology, nectar production and insect visits in a dioecious species, Silene dioica. New Phytologist. 98 (3). p515-529.

Kitahara, M., Yumoto, M., & Kobayashi, T. (2008) Relationship of butterfly diversity with nectar plant species richness in and around the Aokigahara primary woodland of Mount Fuji, central Japan. Biodiversity and Conservation. 17 (11). p2713-2734.

Koch, J., Grigg, A., Gordon, R., & Majer, J. (2010) Arthropods in coarse woody debris in jarrah forest and rehabilitated bauxite mines in Western Australia. Annals of Forest Science. 67 (1). p106-115.

Komonen, A. (2003) Distribution and abundance of insect fungivores in the fruiting bodies of Fomitopsis pinicola. Annales Zoologici Fennici. 40 (6). p495-504.

Komonen, A., Penttilä, R., Lindgren, M., & Hanski, I. (2000) Forest fragmentation truncates a food chain based on an old-growth forest bracket fungus. Oikos. 90 (1). p119-126.

Leather, S. & Bland, K. (1999) Naturalists’ Handbook 27: Insects on cherry trees. UK: The Richmond Publishing Co. Ltd.

Magura, T. (2002) Carabids and forest edge: spatial pattern and edge effect. Forest Ecology and Management. 157 (1). p23-37.

Molnár, T., Magura, T., Tóthmérész, B., & Elek, Z. (2001) Ground beetles (Carabidae) and edge effect in oak-hornbeam forest and grassland transects. European Journal of Soil Biology. 37 (4). p297-300.

Morales-Ramos, J. & Rojas, M. (2001) Nutritional Ecology of the Formosan Subterranean Termite (Isoptera: Rhinotermitidae) – Feeding Response to Commercial Wood Species. Journal of Economic Entomology. 94 (2). p516-523.

Mori, S., Itoh, A., Nanami, S., Tan, S., Chong, L., & Yamakura, T. (2014) Effect of wood density and water permeability on wood decomposition rates of 32 Bornean rainforest trees. Journal of Plant Ecology. 7 (4). p356-363.

Müller, J., Noss, R., Bussler, H., & Brandl, R. (2010) Learning from a “benign neglect strategy” in a national park: Response of saproxylic beetles to dead wood accumulation. Biological Conservation. 143 (11). p2559-2569.

Norton, R. (1980) Observations on phoresy by oribatid mites (Acari: Oribatei). International Journal of Acarology. 6 (2). p121-130.

Niemelä, J., Koivula, M., & Kotze, D. (2007) The effects of forestry on carabid beetles (Coleoptera: Carabidae) in boreal forests. Journal of Insect Conservation. 11 (1). p5-18.

Owen, D. (1978) The effect of a consumer, Phytomyza ilicis, on seasonal leaf-fall in the holly, Ilex aquifolium. Oikos. 31 (2). p268-271.

Pearce, J., Venier, L., Eccles, G., Pedlar, J., & McKenney, D. (2004) Influence of habitat and microhabitat on epigeal spider (Araneae) assemblages in four stand types. Biodiversity & Conservation. 13 (7). p1305-1334.

Ramírez-Hernández, A., Micó, E., de los Ángeles Marcos-García, M., Brustel, H., & Galante, E. (2014) The “dehesa”, a key ecosystem in maintaining the diversity of Mediterranean saproxylic insects (Coleoptera and Diptera: Syrphidae). Biodiversity and Conservation. 23 (8). p2069-2086.

Rasmont, P., Regali, A., Ings, T., Lognay, G., Baudart, E., Marlier, M., Delcarte, E., Viville, P., Marot, C., Falmagne, P., & Verhaeghe, J. (2005) Analysis of pollen and nectar of Arbutus unedo as a food source for Bombus terrestris (Hymenoptera: Apidae). Journal of Economic Entomology. 98 (3). p656-663.

Schiegg, K. (2000) Are there saproxylic beetle species characteristic of high dead wood connectivity?. Ecography. 23 (5). p579-587.

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Siitonen, J. & Ranius, T. (2015) The Importance of Veteran Trees for Saproxylic Insects. In Kirby, K. & Watkins, C. (eds.) Europe’s Changing Woods and Forests: From Wildwood to Managed Landscapes. UK: CABI.

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Trees in the ecosystem pt IV: Trees & arthropods