How do stag beetle larvae cope with the cold?

The photo below shows a stag beetle Lucanus cervus larva together with the larvae of other species found in the same habitat. These larvae were collected during the mild winter of 2014. On that day the soil temperature was 7.5° C, at a depth of 18 cm. Even though these overwintering larva were lethargic in the field, they responded to being handled; you can see by their body posture that they were rather lively by the time that this photo was taken.

Stag beetle, lesser stag beetle and rose chafer larvae. Photo by Maria Fremlin, 5 February 2014

Overwintering third instar larvae. From the left: stag beetle, lesser stag beetle and rose chafer.
Photo by Maria Fremlin. Colchester, Essex, UK. 5 February 2014.

Even though these three detritivore/saproxylic species live close together, a close examination of their gut revealed three different overwintering strategies.

1 - Stag beetle Lucanus cervus larva. Note how dark the body of this larva is; its gut is full of food. The gut is divided in three regions: foregut (hidden under the fat); the midgut is rather large; the hindgut also has three regions: the ileum which is rather large and has a ring of caeca (pouches) in the junction with the midgut; the colon forms an inverted S shape with the rectum which is hidden under the fat.

Dissected stag beetle larva. Photo by Maria Fremlin

Dissected overwintering stag beetle larva. Photo Maria Fremlin.

It seems that in a region with relatively mild winters, like Colchester where I live, stag beetle Lucanus cervus larvae carry on feeding at this temperature. And perhaps at even lower temperatures. I have observed an active larva in captivity feeding at 4 °C. These larvae are quite mobile and if the weather cooled down for a very long period they could always move down away from the frost line. There are stag beetles Lucanus cervus as far north as Southern Sweden, not to mention Central Europe - all regions with very cold winters. However, in those regions I do not know what their overwintering strategy would be.

2 - Lesser stag beetle Dorcus parallelipipedus larva. By contrast with the large stag beetle larva on the left in the top picture, the body of this one is almost translucent. This larva has stopped eating. And it has prepared itself for the winter by emptying its whole gut and filling it with a special winter anti-freeze fluid. Even though it is filled with antifreeze its shape is very similar to the stag beetle larva above.

Dissected lesser stag beetle larva. Photo by Maria Fremlin

Dissected overwintering lesser stag beetle larva. Photo Maria Fremlin.

Lesser stag beetle larvae spend their life in the wood above the ground; they are not as mobile as stag beetle larvae; therefore, are much more exposed to the cold. This strategy will protect them from freezing up. Again, this stag beetle's range in the continent covers areas, which have very severe winters. Indeed the wood where they are might even freeze, sometimes.

3 - Rose chafer Cetonia aurata larva. This larva has quite a different shape; note that its lower abdomen is rather dark. It has also stopped eating, but only its midgut is full of anti-freeze fluid; remarkably, the hindgut remains full; the hindgut is underneath the distended darkened area. See below.

Dissected rose chafer larva. Photo by Maria Fremlin

Dissected overwintering rose chafer larva. Photo Maria Fremlin.

Rose chafer larvae feed mostly below the soil, often together with stag beetle larvae, but they seem to be much more sensitive to the cold. I have observed that in my area when the soil cools down to around 10 °C they stop feeding, this is their hibernation strategy. In the Spring, when the soil warms up to around 9-10 °C, they resume their feeding. This beetle was thought of as a Southern beetle in the UK, but it seems to be expanding its range, recently. See some recent records here.

Further reading about overwintering strategies and gut morphology:
Edwards EE (1930) On the Morphology of the Larva of Dorcus parallelopipedus, L. (Coleoptera), Zoological Journal of the Linnean Society, 37(251):93–108, https://doi.org/10.1111/j.1096-3642.1930.tb02062.x. [PDF]
Insect winter ecology - in Wikipedia.
Neven LG, Duman JG, Beals JM, Castellino FJ (1986) Overwintering adaptations of the stag beetle, Ceruchus piceus: removal of ice nucleators in the winter to promote supercooling. Journal of Comparative Physiology, 156:707-716.
Xu, L, Neven, LG and Duman, JG (1990) Hormonal control of hemolymph lipoprotein ice nucleators in overwintering freeze susceptible larvae of the stag beetle Ceruchus piceus: Adipokinetic hormone and juvenile hormone. Journal of Comparative Physiology B, 160:51-59.
Walters KR Jr1, Serianni AS, Voituron Y, Sformo T, Barnes BM, Duman JG. (2011) A thermal hysteresis-producing xylomannan glycolipid antifreeze associated with cold tolerance is found in diverse taxa. Journal of Comparative Physiology B, 181(5):631-40.

Last modified: Sat April 13 2024