A Testudo graeca graeca lays her eggs.
Failure to provide adequate nesting sites in captivity can have serious consequences for heath, including an increased danger of egg-retention leading to possible fatal peritonitis. The lack of acceptable nesting sites can also lead to increased stress, and negatively impact upon captive breeding success.
The consequences of eggs being retained too long due to lack of suitable nesting sites can be very serious, including extreme over-calcification as seen here.
Tortoise nesting behaviour can be broken down into several distinct phases:
Nest site selection
Preparation of the nest site
Excavating the nest
Laying
Filling in and covering the nest
Different species exhibit markedly different preferences and behaviour in each of these phases, so it is important to understand the particular requirements of the species in question. For example, Mediterranean Testudo species tend to exhibit a strong preference for nesting on gentle slopes, with sandy, well-drained soils. Damp clay soils or soil that is too stony is likely to be rejected as unsuitable, as are nest sites on flat surfaces.
A very elderly wild female Testudo ibera excavates a nest on a sloping hillside in Southwestern Turkey.
Some examples of attempts to replicate similar conditions in Northern Europe (former Tortoise Trust site in Wales, UK). Note the inclusion of 'mini greenhouses' made using wood and polycarbonate 'twinwall' to enhance basking options. These pens were seeded with edible wild flowers and were well-drained and with a steep slope.
Indications that a female might be preparing to nest include:
Not feeding
Hyperactivity
Digging test nests
Unusual movements in the back legs
'Sniffing' the ground frequently
Testudo hermanni nesting in warm, dry soil.
By contrast, many tropical species such as Redfoot tortoises (Chelonoidis carbonaria) will accept flat nest sites readily, but typically prefer the soil to be rich in organic content, moist, or even muddy. Mediterranean Testudo species usually prefer to lay in full sun, on a dry day, from mid-day to late afternoon; other species, such as Chelonoidis carbonaria or American Box turtles (Terrapene carolina) frequently lay at dusk, and especially during episodes of light rain when humidity is particularly high. It is important to be aware of these traits, as the prevailing weather conditions or time of day can give a good general indication as to when nesting may occur. Temperature, humidity and light levels then, are just as important to nesting tortoises as the substrate itself. Use an infra-red non- contact thermometer to check surface temperatures and at nest depth. Often, subsoil temperatures of between 25C to 29C are preferred.
Where it is suspected that a female has eggs, but may be experiencing problems, we highly recommend getting your vet to conduct an x-ray to determine the number, position and calcification status of the eggs.
A normal and healthy clutch of eggs from a Testudo graeca revealed by x-ray.
Those species that typically prefer to nest in sandy soil frequently prefer to excavate their nests close to shrubs, or in sand that is infiltrated by fine plant roots. It is possible that this provides extra cohesion during excavation, making collapses less likely.
Testudo kleinmanni (Egyptian tortoise) nesting
Species from rainforest type environments often utilise leaf litter in the construction of their nests; one species in particular, Manouria emys (the Burmese Brown tortoise) actually constructs a nest mound from leaf litter. The Yellowfoot tortoise (Chelonoidis denticulata) from South America is another species that is sometimes known to nest on the ground, under decaying leaf litter rather than excavate a nest in the usual manner. Though the ones we kept did excavate nests. Unlike Manouria emys, however, it appears to make use of pre-existing accumulations of leaf litter rather than actively constructing a mound specifically for nesting purposes.
A Three-toed American box turtle (Terrapene carolina triunguis) nests in warm, moist substrate at dusk.
The depth of substrate available is a very important factor for all species that excavate nests. If insufficient depth is available, nesting will usually be terminated. In captive situations, therefore, it is necessary to ensure that the laying area does provide enough depth of substrate to avoid this. It is only possible to offer general guidelines, as different species do vary in their nest depths. For most species the depth of substrate should be at least equal to the length of the hind limbs plus 70% of the length of the carapace.
We have found that many terrestrial species will nest in captivity in a substrate comprised of 60% soft (play) sand mixed with 40% loamy compost. If outdoor nesting sites are not possible, indoor artificial sites based upon such a mixture will often yield good results, especially if positioned under a basking lamp and of sufficient size and depth. We would recommend that for a medium-sized Mediterranean tortoise, and area of at least 1 square meter should be provided, and that this should be of adequate depth according to the formula described above. If the artificial nest site can slope gently, this may well increase the chances that it will be accepted by the female as a suitable and safe location.
For small tortoises, a large plastic or even cardboard box can be filled with suitable substrate and raised to an appropriate temperature. Large tortoises will require an artificial nesting mound constructed of a concrete block or strong wooden enclosure. The temperature of the substrate is certainly important - few tortoises will lay in a cold substrate. You may see the female touching the surface with her nose, as if inspecting it carefully. What she is actually doing is sensing the surface temperature to see if it is suitable for incubating her eggs. Some tortoises also appear to use the hind limbs to check that the subsurface temperature is also satisfactory. We allow artificial nest sites to stabilize to (warm) room temperature before use, and then, as mentioned previously, provide surface heat for a few hours by means of an overhead basking lamp.
One rather odd phenomenon we have noted is that gravid females, who may be reluctant during daytime to use an artificial nesting site, can often be persuaded to accept one if placed on it after dusk, under a basking lamp. We do not fully understand why this should be the case, but we have witnessed it often enough to be convinced that there is some merit in this technique! It may go right back to the nesting behaviours of sea turtles. Many species of terrestrial tortoises do seem to enter a similar 'trance-like' state while depositing their eggs.
Once the eggs have been laid, these should be removed to an incubator for safe, controlled incubation. We will have a newly updated article on that topic shortly.
To receive updates and notifications of new articles, videos and other important news please consider 'Subscribing' to the Tortoise Trust. This supports our work and allows us to make this information available. The Tortoise Trust also offers a comprehensive online course that covers all aspects of behaviour, ecology and captive care.
All text & images (c) Tortoise Trust/A. C. Highfield 2002-2024
I wonder if the ones that nest after dusk but with artificial heat/light do so because of some instinct to ensure the nest isn't seen by potential predators?