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Writer's picturePhilippos Katsiyiannis

The pleasures of external fertilisation


The above male Fire-bellied Toad, is floating on the surface of the swamp in which it lives, and calling to attract a female. Calling day and night, the small toad will finally succeed to grasp a female in amplexus and pass its genes to the next generation. Amplexus is the special grip with which amphibian males hold the females during copulation. Although the animals will get tightly grasped together, there is no penis, no insertion, and the fertilisation takes place in the water, at least for the Anura, the group of frogs and toads (Anura in Greek means 'without tail). When the female release the eggs, the male is releasing sperm to fertilise them.


Tadpole of Yellow-Bellied Toad, Bombina variegata, photographed in situ in a mountain stream


Bufotes viridis pair, entering the water

Like these Green Toads in amplexus, that just enter the water. They spend most of their lives on dry land, but come spring, and they migrate to ponds where they breed. Males will grasp walking females in amplexus, far from the water, and the larger females will carry them on their backs to the water bodies where they spawn. There, in the water, a slow process starts, where most of the time the two animals are motionless, with bouts of activity every now and then. This is very true especially when temperatures are low. But in February, Green Toads will mate with air temperatures 11 degrees Celsius or lower, and water temperatures 13 degrees or lower.


Green Toads, Bufotes viridis, in amplexus, with eggs in ribbons all around

The ribbons of eggs seen in the above photo, quickly transform into tiny embryos and then tadpoles. Thousands of them.


Just hatched tadpoles of the Green Toad. The area where the head will take shape, can be seen


And this is after some days, thousands of tiny tadpoles feeding on algae.

A Common Toad, Bufo bufo, in its pool with eggs. All spawning happens by night, for a number of reasons

Sometimes the process can be more complicated. When males find a female, they try to grasp her, and sometimes more than one manage to get a grip.


Two male Common Toads, Bufo bufo, trying to mate with a female (lower left)

Two male Agile Frogs, Rana dalmatina, one in amplexus and the second trying to get a grip. Egg mass can be seen on the left

Green and Common Toads, are of the earlier breeders, starting in January, especially in Peloponnisos and southern Greece. But the same species can breed later too, and on the mountains breeding can be seen in May as well. Another early breeder is the Agile Frog, Rana dalmatina, with eggs found as early as Christmas.


Egg mass of Rana dalmatina in shallow, temporary water

Freshly laid eggs of Rana dalmatina

Rana dalmatina in amplexus. The female is much bigger than male, its belly very distended with eggs. They also become bright red

Another member of the Rana genus is the Greek Stream Frog, Rana graeca. It was a species that until recently the voice of it was not recorded. Despite the fact the they live in mountsin streams and there is a lot of noise form the water, Rana graeca have a quite, soft croaking. For the video, you can see here: https://herpinggreece.wixsite.com/herpinggreece/post/greek-stream-frog-call-and-amplexus


Rana graeca in amplexus. The males turn greenish and the females reddish in breeding season

In lowland wetlands of western Greece, a cacophony of croakings can be heard at night. Many species vie for attention, all together making the soundscape almost unbearable.


Pelophylax epeiroticus in situ, on the beginning of spring

Pelophylax ridibundus croaking and calling for a mate

Hyla arborea, one little guy with a very big voice

When it is mating season, male Tree Frogs Hyla arborea, become dark, golden or brown. In all other seasons they are bright green

And this is the product of Tree Frog love: their eggs

But at Western Peloponnisos, we still have to hear and record the voice of a largely secretive and silent toad: the Eastern Spadefoot. Although we have been at the right time at the right place, no one has heard this little guy up to this day. Its secrets keep calling us to their breeding grounds for years, but still nothing.


Phantom of the swamp. The secretive Eastern Spadefoot, Pelobates balcanicus chloeae

Pelobates balcanicus in amplexus. Note that the male grasps the female in front of the hind legs, not behind the front legs

Pelobates balcanicus tadpoles are huge

All Anura communicate with sound and they can be easily detected from their croaking. But there is another order of Amphibians, quite different from them, they are called Urodela. In this order of amphibians Salamanders and Newts are members. These are silent animals and they use body language and pheromones to mate.


Male Lissotriton graecus. They are really small animals, but if you bend and pay some attention, they are just gorgeous

Salamandra salamandra tadpoles in a brook. Most of the time, it is easier to find the species in an area from the tadpoles

Triturus macedonicus tadpole from western Greece. Note the external gills

Newts have a special way to mate. Males try to intercept the trajectory of a female, they swim in front of her waving their tail towards her, sending pheromones to her. If she agrees, the male deposits a spermatophore, a sac with sperm, and the female takes this inside her body with her cloaca.


Lissotriton graecus display. The male on the right waves the tail towards the female on the left, which is full of eggs. It is not apparent from the photo, but this is in the water

Ichthyosaura alpestris, the Alpine Newt, in display. The male is on the left trying to impress the uninterested female

Ichthyosaura alpestris male (below) touching and trying to intercept the female

Last but not least, in Greece we have two Lyciasalamanders living on Kastellorizo and Karpathos group of islands. Very restricted and beautiful animals, they have no need of water for their reproduction. They have evolved on very dry islands, and they managed to survive because they give birth to live young. Their amplexus is also special. Contrary to any other Greek amphibian, the male goes under the female, stretches the arms backwards to grasp the arms of the female in a strange amplexus, and uses the soft spike on top of its tail, to make its way to her cloaca


Male (bottom) and female on top. Lyciasalamandra luschani making out


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