From the Literature: Oxygen, Temperature, and Giant Insects

I hope everyone liked the giant insects post last week!  It was one of my favorites to write.  The topic is just so fun!  I continue with the subject this week by describing a scientific paper that was released in July.  It combines several things I love (giant insects, aquatic insects, and respiration)  into one manuscript of pure science fabulousness!  Let’s get to it, shall we?

You probably learned as a kid that insects are ectothermic (aka, cold-blooded).  Ectothermic organisms are largely unable to regulate their body temperatures, so their bodies remain close to the temperature of their environments.  As the temperature increases, processes like metabolic rates speed up.  The opposite happens at cold temperatures and everything slows way down.  Ectotherms survive best under a range of temperatures where their body processes work efficiently, but the animal is still able to get everything it needs (food, water, oxygen, etc) from the environment.  They’re like Goldilocks: they don’t like things too hot or too cold and prefer for things to be just right.

Oxygen plays a big role in the interaction of ectotherms with their environments, especially at extreme temperatures.  Let’s consider a hypothetical insect, say a grasshopper.  As the grasshopper gets warmer, its metabolic rate increases and its body processes become more efficient.  However, as the grasshopper’s metabolic rate increases, so does its oxygen consumption.  At some point, the oxygen demand of the grasshopper may become greater than its oxygen availability and all sorts of bad things start to happen as its body processes start to break down.  Oxygen plays a role at very cold temperatures as well, leading scientists to propose that oxygen can set thermal limits (the maximum and minimum temperatures our grasshopper can survive) in ectotherms.

One problem though: terrestrial insects don’t fit the pattern observed in many other ectothermic animals.  This may be because their respiratory systems do not depend on lungs and blood to deliver oxygen to their cells and instead deliver oxygen directly to their cells via a series of tubes that connect to the outer environment.  This creates a terribly efficient system that provides enough oxygen even at high temperatures for many terrestrial insects.  Quite simply, their respiratory system provides enough oxygen even under the worst conditions.  But what if the insects live in oxygen-limited environments, such as water?  Might oxygen play a role in setting those upper thermal limits then?


Image from bilder/030416_dinocras_cephalotes.htm.

Researchers Wilco Verberk and David Bilton considered this question and determined that if the thermal limits of any insects were to be limited by oxygen levels, aquatic insects were the most likely suspects.  So, they chose an insect that requires a low temperature and flowing water as their subject, the stonefly Dinocras cephalotes.  If the maximum temperature the stonefly could tolerate was limited by oxygen consumption, the maximum tolerable temperature would decrease in low oxygen water while it would increase in high oxygen water.  They then developed a simple experiment to determine whether this was the case.

The team placed stoneflies in flow-through chambers in a water tank and ran 10°C water containing various mixtures of oxygen and nitrogen (20% O2/80% N2 = normal, 5% O2 /95% N2 = low oxygen treatment, and 60% O2/40% N2 = high oxygen treatment) through them.  After letting the stoneflies acclimate for an hour, they ramped the temperature of the water up 0.25 degrees per minute until the critical temperature was reached, i.e., the stoneflies started showing signs of thermal stress such as lack of movement and leg twitching.  Then they compared the critical temperatures for each treatment to determine if their hypothesis was correct.

And it was!  They discovered that the upper thermal limit increased almost 3°C in the high oxygen water compared to water containing normal levels of oxygen.  Conversely, the upper thermal limit decreased in low oxygen water by about 1.5°C compared to that at normal oxygen levels.  The conclusion: oxygen levels can set upper thermal limits in larval aquatic insects!

Now you might be wondering why this is exciting or what any of this has to do with giant insects.  The results are interesting for several reasons, but largely because they show that some insects do experience oxygen-induced changes in their upper thermal limits.  This means that, while terrestrial insects might be able to obtain enough oxygen at any temperature to meet their needs, aquatic insects and other insects that live in oxygen limited environments can reach a temperature at which their oxygen demand outstrips the oxygen available to them.  Consider how an insect such as a stonefly gets the oxygen it uses.  They don’t have any spiracles (the pores through which terrestrial insects “breathe”), so oxygen is simply absorbed through the exoskeleton.  Many stoneflies have gills to make this process more efficient (the bigger your body surface, the more oxygen you can absorb from the water), but it’s still a very slow process.  The size of these insects may be limited as a result.  Aquatic insects that rely on absorbing oxygen from the water rather than going to the surface to breathe are also unable to regulate their oxygen uptake very well.  They can do various behaviors to increase the flow of oxygen into their bodies when they become oxygen stressed, but oxygen becomes toxic at very high concentrations.  Aquatic insects can’t do much to prevent oxygen from flowing into their bodies, so this can be a problem.

And this brings us to the giant insect part of the paper.  Verberk and Bilton propose that oxygen limitation at temperature extremes may have contributed to the rise of insect gigantism in the late Carboniferous and early Permian.  This makes sense considering how many of the giant insects were insects that probably had aquatic nymphs (proto-dragonflies, mayflies, and stoneflies, among other aquatic organisms).  The high levels of oxygen at the end of the Palaeozoic meant that oxygen could be absorbed more efficiently by aquatic insects and allowed them to become larger.  I covered this hypothesis last week, so check that post for more details.

Alternatively, Verberk and Bilton suggest that oxygen toxicity may have played a significant role in promoting insect gigantism.  How can an aquatic insect cope with increasing levels of oxygen in water and prevent oxygen poisoning?  They can get bigger!  If insects increased in size as oxygen levels in water rose, then they could counteract the negative effects of high oxygen levels on their bodies.  Oxygen levels at the end of the Palaeozoic were so high that aquatic insects likely had to become very large to prevent oxygen poisoning.  Giant immatures then led to giant adults.  Hence, giant insects that resulted from the limits of their respiratory systems in very high oxygen environments!  It’s a very interesting, new idea.  I suspect many people will do further tests in the future to determine whether this might really have been possible, so we’ll see if it holds up to further study.

I love this hypothesis!  Still, I have to point out that there is one major assumption that the entire hypothesis is built upon, that the giant proto-dragonflies, mayflies, stoneflies, etc had aquatic nymphs.  Modern dragonflies, mayflies, and stoneflies have aquatic immatures, so it’s likely that their predecessors did too.  However, there is no fossil evidence of aquatic nymphs for these groups at the time of the giant insects.  For all we know, the griffenflies and giant mayflies may have had terrestrial nymphs, which would make Verberk and Bilton’s hypothesis fall apart completely.  While the authors did acknowledge this assumption, I think their position would be strengthened if a fossil of even one aquatic immature could be found from that time period.  Without that piece of evidence, I fear this hypothesis is built upon a shaky foundation, one that might not hold up to scrutiny.

But wow!  A new explanation for how giant insects may have evolved!  And focused on the aquatic stages of insects!  You can see why I’m excited by it.  I can’t wait to see the research generated by this paper in the future.  It’s going to make for some very interesting reading!


Literature Cited:

Verberk WC, & Bilton DT (2011). Can oxygen set thermal limits in an insect and drive gigantism? PloS one, 6 (7) PMID: 21818347

This paper is open access!  Full text available online here:


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Mayflies, Damselflies, and Stoneflies: What’s the Difference?

I haven’t done an identification post for a while, so its high time that I write another one!  I find that a lot of people have a hard time distinguishing the aquatic insect nymphs with tails sticking off the back, the mayflies, the damselflies, and the stoneflies.  They’re easy to tell apart once you learn a few basics!  A lot of people have read my post on how to tell the damselflies and dragonflies apart as nymphs, so let’s start with them.

Behold, the mighty damselfly:

damselfly nymph

Damselfly nymph

There are several things to look for that will let you know this is a damselfly nymph and not a stonefly or mayfly.  However, the mouthpart is a dead giveaway!  If you don’t know about the awesome odonate mouthpart, allow me to enlighten you.  Odonates have highly adapted mouthparts that form a long, hinged structure that they can thrust out toward prey to capture it and draw it back to the chewing mouthparts to be eaten.  There are pictures of this structure available on the post linked above and you can see a little part of it sticking out past the head of the damselfly in the image above.  Odonates are the only insects that have this style of mouthpart, so if you have a nymph with tails sticking off the back-end and you can see a long, folded mouthpart under the head, you’re looking at a damselfly for sure.

But perhaps you’re looking at an insect in the water and you aren’t able (or willing) to pull it out to look at the mouthpart – what then?  Well, take a look at the location and structure of the gills:

Damselfly gills

Damselfly gills

The three damselfly “tails” are really gills that they use to help them breathe and swim!  They are always located at the back-end of the insect and they tend to be broad and leaf-shaped with varying levels of pointy-ness.  As you’ll see in a moment, the stoneflies and the mayflies have gills in other locations and do not have broad, leaf-like tails.  If you see gills that look like the image above, you’re looking at a damselfly nymph!

Let’s move along to the mayflies:



You should notice some differences between the mayfly and the damselfly right away.  First, look at the tails:

mayfly tails

Mayfly tails

Nothing broad and leaf-like about these tails!  Mayflies have long, filamentous tails, often longer than their bodies.  They also usually have three tails like the damselflies, but some groups only have 2.  Clearly, the flat-headed mayfly in the photo falls into the latter category.  This causes some confusion when distinguishing the mayflies from the stoneflies, as you’ll see in a moment.  However, if you see 3 filamentous tails, you’ve got a mayfly on your hands!

Now let’s take a look at the location of mayfly gills:

mayfly gills

Mayfly gills

The gills  are always attached along the sides or the bottom of the abdomen in the mayflies, never on the thorax or sticking off the back. If you see gills in another location, you’re not looking at a mayfly.  Mayfly gills tend to be broad and leaf-like as in the damselflies, though they may be fringed or sharply pointed in some groups.  They usually have a pair of gills on nearly every abdominal segment, though the exact placement on the abdomen varies by group.

Now we’re left with the stoneflies:



Stoneflies and mayflies look a lot alike in most cases.  The mayfly in my photos above is a specialized species adapted for living in fast flowing water, but a lot of mayflies are shaped more like the stonefly depicted here.  How do you tell them apart when the body shapes are similar?  Let’s look at the tails first:

Stonefly "tails"

Stonefly "tails"

Stoneflies always have two tails.  Like the mayflies, they’re long and filamentous.  In some species, these tails are very long.  In others, they’re shorter than the length of the abdomen.  They’re never leaf-like.

Let’s check out the location of the gills too.

Stonefly armpit gills

Stonefly gill location

Unlike the damselflies and mayflies, stonefly gill placement is quite variable.  Many species don’t have gills.  Some species that do have gills don’t get them until they’ve matured to some specific point.  Some species have gills on the abdomen, but if they do they’re located only on the first few abdominal segments and never further down.  (This helps distinguish them from the mayflies, which almost always have gills on the 3rd-6th abdominal segments.)  But in most stoneflies with gills, you’ll find them in their armpits, as indicated in the photo.  Stonefly gills are very different from the broad, flattened gills of damselflies and mayflies.  They typically have a round main stalk with multiple branches.  These are called “finger-like” gills for some reason, but I think the structure is rather similar to the boojum tree, just on a smaller scale:

Boojum Tree

Boojum Tree. Photo by Bernard Gagnon, from wiki/File:Boojum_Tree.jpg.

I find that people have the most trouble telling the mayflies and stoneflies apart.  If the mayfly has three tails, no problem!  It’s a mayfly for sure.  However, you have to remember those pesky two-tailed mayflies that throw a wrench in the whole system.  Plus, mayflies are notorious for losing their gills.  If you’re working with preserved specimens, sometimes it’s hard to figure out where the gills did or did not attach.  How then do you tell a two-tailed mayfly with no gills apart from a similarly shaped stonefly with no gills?  It’s easy!  Look at the claws on the legs.  Mayflies have one claw on every foot.  Stoneflies have two.  It couldn’t be simpler.

As with any identification, the more animals you see, the easier this gets.  For those of you who have little experience collecting and identifying insects, getting a specimen IDed to order can be a challenge at times!  Remembering the characteristics of tons of insects can be hard too.  I thus present this handy-dandy chart that summarizes the information I covered above:

Mayfly Damselfly Stonefly
Location of Gills abdomen end of abdomen when present, thorax, base of abdomen
Shape of Gills leaf-like, plate-like, or fringed leaf-like finger-like
Style of Mouthparts chewing chewing + hinged segment folded under head chewing
Number of Tails 2-3 3 2
Shape of Tails filamentous leaf-like filamentous
Number of Claws 1 2 2

If you forget the characteristics of the mayflies, damselflies, and stoneflies, use this chart as a quick reminder of what to look for!

Next up: another thrilling edition of Friday 5!  This week’s will feature 5 places I’ve found a particular type of tiny insect in my home.  Check it out to discover where these little beasts may be lurking!


Unless otherwise stated, all text, images, and video are copyright © 2011