Triturus pygmaeus. Juvenile. Photo © Paolo Mazzei Amphibians & Reptiles of Europe

Triturus pygmaeus. Photo © Henrique Caetano, hcaetano@hotmail.com, courtesy of Marco Caetano, mcaetano2905@hotmail.com, Alentejo, Portugal.

Triturus pygmaeus. Photo © Henrique Caetano, hcaetano@hotmail.com, courtesy of Marco Caetano, mcaetano2905@hotmail.com, Alentejo, Portugal.

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Breeding (General)
European newts are famous for their remarkable sexual dimorphism during the breeding season, and sometimes elaborate mating rituals. Males of many species develop elaborate physical characteristics during this time, the most noted being an extended dorsal and tail crest, but may also include webbed toes, and bright coloration on the tail and flanks. The courtship behaviors of European newts are well analyzed and documented, as the following tables show (tables adapted from Arntzen & Sparreboom (1989), and Sparreboom and Arntzen (1987), respectively).

Table 1.1 - Display phase techniques performed by male newts of European newts

Fan	Tail is folded against the side of the body nearest the female and oscillated rapidly.
Wave	The male presents a full lateral view of his body to the female.
Whip	Tail is lashed against the side of the body.
Whiplash	An aggressive version of the Whip, with the hind legs elevated off the ground.
Flick	Tail is bent, and flicked repetitively toward the female.
Flamenco	Tail is raised to a 90° angle from the body and the tip is waved from side to side.
Wiggle	Tip of the tail is wiggled in a worm-like manner, also called distal luring.
Rocking	The male rocks on fore limbs while beating the tail in the direction of the female.
Head-Bob	Vertical up and down movements of the head.
Cat-Buckle	The body is raised on the hind legs and kinked in the middle to emphasize the size of the crest.
Lean-In	The male leans sideways toward the female from a cat-buckle position, displaying a lateral view of the body and crest.
Flurry	Rapid oscillations of the tail on the side of opposite of the female, while in Lean-In position.
Retreat	A female follows a male, who retreats in front of her while performing a particular type of tail-fanning.

Table 1.2 - Species specific display phase techniques of European newts

Fan

Wave

Whip

Whip-lash

Flick

Flam-enco

Wiggle

Rocking

Head-bob

Cat-Buckle

Lean-In

Flurry

Retreat

M. alpestris

o

x

x

x

x

x

x

x

x

o

o

x

x

"Cristatus" group

x

x

x

o

x

x

x

o

x

o

o

x

x

T. marmoratus

x

x

x

o

x

x

x

o

x

o

o

x

x

T. pygmaeus

x

x

x

o

x

x

x

o

x

o

o

x

x

T. vittatus

x

x

x

o

x

x

x

x

o

o

o

o

x

L. boscai

o

x

x

x

o

o

o

x

x

x

x

x

x

L. italicus

o

x

x

x

o

o

o

x

x

x

x

x

o

L. helveticus

o

o

o

x

x

x

o

x

x

x

x

x

o

L. vulgaris

o

o

o

x

x

x

o

x

x

x

x

x

o

L. montandoni

o

o

o

x

x

x

o

x

x

x

x

x

o

Table 1.3 - Outline of breeding habits of newts of European newts 

Some of the information provided in this table was compiled from information provided from experienced breeders of the listed species, and includes of a brief summary of procedures and results from such individuals. Natural breeding information contained in this table is composed of a summary of information collected from various reference materials (see the reference section at the bottom for more information). Please note that it is not always necessary to provide a dormant period with temperatures as low as would be encountered in the wild. Most species and individuals requiring a cooling period will enter breeding mode after a short period at temperatures from 45°F - 50°F.
	Natural Breeding Season	Clutch Size and other Characteristics
M. alpestris	Early or pre-spring through early autumn, sometimes starting later, after frozen pools have melted.	Usually around 150-180 eggs are produced, and attached individually to leaves, with the tips folded around the egg. Larvae are pond-type, with a frequency of neoteny.
T. carnifex	Mainly March through May, but adults can be found in water year-round.	Females may produce 200-250 eggs in a season. Females will wrap individual eggs in small-leaved plant varieties. Larvae are pond-type. Larvae metamorphose around September, and sexual maturity is reached after 2-6 years. Neoteny has been noted, but is rare in T. carnifex. This species possesses a chromosomal abnormality, described below.
T. cristatus	Late February to early June, after a dormant winter period spend on land.	Females produce up to 200-250 eggs in a season. Females will wrap individual eggs in small-leaved plant varieties. Larvae are pond-type. Larvae metamorphose around the end of summer or beginning of autumn, and sexual maturity is reached after 2-6 years. This species possesses a chromosomal abnormality, described below.
T. dobrogicus	Migration begins mid-Feb., and adults may remain aquatic until August. There may also be a second migration in autumn.	Females produce up to 200-250 eggs in a season. Females will wrap individual eggs in small-leaved plant varieties. Larvae are pond-type. Larvae metamorphose between the end of August and the end of September, and sexual maturity is reached after 2-6 years. This species possesses a chromosomal abnormality, described below.
T. karelinii	Assumed to be similar to T. cristatus	Females produce up to 200-250 eggs in a season. Females will wrap individual eggs in small-leaved plant varieties. Larvae are pond-type. Larvae metamorphose around the end of summer or beginning of autumn, and sexual maturity is reached after 2-6 years. This species possesses a chromosomal abnormality, described below.
T. marmoratus	March through May, after a cold winter period.	Females may produce up to 400 eggs in a season. Females will wrap individual eggs in small-leaved plant varieties. Larvae are pond-type. This species possesses a chromosomal abnormality, described below in the Breeding section.
T. pygmaeus	May begin as early as January, after a short, cooler winter period.	Females may produce up to 400 eggs in a season. Females will wrap individual eggs in small-leaved plant varieties. Larvae are pond-type. This species possesses a chromosomal abnormality, described below.
T. vittatus	Spring through early summer in the northern ranges. During the rainy season in the warmer southern ranges, sometimes lasting through winter.	Usually around 150-180 eggs are produced. In the warmer southern regions, metamorphosis may begin as early as 2 months after deposition, and later in the cooler, mountain regions. Larvae are typical pond-type.
L. boscai	December and March.	Females may produce 100-250 eggs in a season. Eggs are deposited singly, and folded between leaf tips of submerged vegetation. Larvae are pond-type. Larvae may spend their first winter aquatic, metamorphosing the following spring or summer.
L. italicus	Spring through autumn.	Females may produce 300-400 eggs in a season. Eggs are attached singly to aquatic vegetation, sometimes folded between leaf tips. Sexual maturity is reached late in the first to the second year.
L. helveticus	Northern populations may enter the water in February or March, and remain aquatic until June or July.	Females may produce 100-250 eggs in a season. Eggs are deposited singly, and folded between leaf tips of submerged vegetation. Larvae are pond-type.
L. vulgaris	Spring through autumn.	...
T. montandoni	Spring through autumn.	Females may produce 100-300 eggs in a season. Eggs are deposited singly, or in small masses to submerged vegetation. Individually deposited eggs are usually wrapped in leaf tips. Larvae are pond-type. Larvae may spend their first winter aquatic, metamorphosing the following spring or summer.

During the breeding season, captive newts can be moved into a tank with a deeper water area (10+ inches) and a few land areas. The newts should be placed on land, so they can make the transition into the water themselves. Males of the crested species will not develop crests until they enter the water, and if after a week or so one or more have not entered the water, they may not attempt to breed and should be moved back into their terrestrial home. For large 50%-50% water/land tanks, the newts will enter the water area on their own, provided the right conditions exist. Some species may require a cooling or dormant period prior to entering breeding mode, as summarized above in Table 1.3. 

Males of the crested species use their stored food reserves to produce the tail fin and crest, and so should be fed in abundance prior to the cooling period (if a cooling period is required). During cooling periods, the newts will be less active and should not be fed in abundance, as their metabolism slows considerably during this time. Crest size is an important criteria females use to judge their perspective mates' worth (Griffiths, 1996). The female wants to choose mates that display exceptional characteristics to insure the lowest mortality rate of her offspring. Because the crest size is related to the males food reserves, large crests may indicate a good forager and hunter, characteristics that would be passed on to their young. Larger crests may also indicate an older male, one that has survived for many years, as newts continue to grow throughout their adult life. These superior characteristics would be inherited by the offspring, which may account for the females' fastidious behavior when choosing the right mate.

The elaborate courtship behavior of European newts is similar, but modified, for all species, and, interestingly, does not consist of amplexus at any point. Males become increasingly active, and sometimes aggressive during the breeding season, incessantly trying to thwart competing males' attempts at courtship. Courtship begins with the male actively pursuing the female, sniffing her cloaca, and moving in front of her to block her path. If the female is interested, she will remain stationary, signaling the male to continue his intricate courting ritual. The male will display a complex array of movements and tail-fanning to prove himself worthy to the female. He will begin by fanning pheromones toward the female with the tail using various methods, as described above in Tables 1.1 and 1.2. Tail-fanning by the crested species may be interspersed with various gestures meant to display, and possibly exaggerate the the size of the crest.

The final stage of the courtship routine is spermatophore deposition and transfer, which varies only slightly for certain species. In all Triturus species, when the male feels that his display has been successful, he will lead the female away by creeping slowly in front of her, oscillating the tail. If the female is still interested, she will follow, touching the males cloaca with her snout, after which the male will deposit a spermatophore. Males of most species will manipulate the female to collect the spermatophore by turning perpendicular to her to block her from proceeding, just about when her cloaca is above the spermatophore. The female will continue to touch the males cloaca with her snout, and the male will then push her back by gently unfolding the tail. This pushes the female over the spermatophore, allowing it to adhere to her cloaca, and ensuring its collection. Only the "Cristatus" group, T. marmoratus, and T. pygmaeus do not perform this push-back technique, possibly because the manipulation is delicate, and not easily performed in highly vegetated waters (Griffiths, 1996). The spermatophore deposition phase may be repeated several times, enabling the female to collect several spermatophores from a single male, although typically only those dropped later on in the process are collected (Griffiths, 1996).

Egg deposition is similar for Triturus, Mesotriton, and Lissotriton, and consists of the female depositing single eggs or strings on small leafs, and folding the leaf tip or edge over the egg. Females may produce several hundred eggs per season, depositing them over the course of several months. In captivity, several small-leaved, aquatic plants should be including in the water area for egg deposition. Many species are especially fond of consuming their own eggs, so the eggs should be promptly removed and reared separately from the adults. The average clutch size and characteristics for each species are outlined above in Table 1.3.

T. cristatus, T. carnifex, T. dobrogicus, T. karelinii, T. marmoratus, and T. pygmaeus possess an abnormality in larval development that results in the termination of 50% of all eggs produced each season. In these species, half of the larvae produced stop developing around the time the tail begins to develop, or shortly after, and the embryo spontaneously aborts. This occurs as a result of the method in which chromosomes combine after fertilization. Of the twelve pairs of chromosomes in newt cells, the pair in Cell No. 1 differ from each other, and have been labeled 1A and 1B. These are the chromosomes that pass into the gametes (sex cells; eggs and sperm) of the parents. In the normal instance, one chromosome, either 1A or 1B, is contributed by each parent, so that the combination is always 1A + 1B. The chromosomal abnormality occurs when the chromosome contributed are of the same type, that is, when the combination is 1A + 1A, or 1B + 1B. The evolutionary aspects of this anomaly are discussed in detail in Malmgren, 2001.

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References

Arntzen, J.W., and M. Sparreboom. 1989. A phylogeny for the Old World newts, genus Triturus: biochemical behavioural data. Journal of Zoology 219: 645-664.

Arntzen, J.W., and G.P. Wallis. 1999. Geographic variation and taxonomy of crested newts (Triturus cristauts superspecies): morphological and mitochondrial DNA data. Contributions to Zoology 68(3): 181-203.

Garcia-Paris, M., Herrero, P., Martin, C., Dorda, J., Esteban, M., and Arano, B., 1993. Morphological characterization, cytogenetic analysis, and geographical distribution of the Pygmy Marbled Newt, Triturus marmoratus pygmaeus (Wolterstorff, 1905) (Caudata, Salamandridae). Bijdr. Dierkd., 63: 3-14.

García-París, M., A. Montori, and P. Herrero. 2004. Amphibia: Lissamphibia. Fauna Iberica Vol. 24. Madrid: Museo Nacional de Ciencias Naturales and Consejo Superior de Investigaciones Científicas.

Griffiths, Richard A. Newts and Salamanders of Europe. San Diego, CA: Academic Press Inc., 1996.

Litvinchuk, S.N., and L.J. Borkin. 2000. Intraspecific taxonomy and nomenclature of the Danube crested newt, Triturus dobrogicus. Amphibia-Reptilia 21(4) 419-430.

Malmgren, J.C. 2001. Evolutionary Ecology of Newts. PhD Thesis in Biology. Örebro, Sweden: Örebro University Press.

Obst, Fritz Jugen, Udo Jacob, and K. Richter. Completely Illustrated Atlas of Reptiles and Amphibians for the Terrarium. Neptune City, NY: T.F.H. Publications, Inc., 1989.

Sparreboom, M., and J.W. Arntzen, 1987. A survey of behavior in the Old World newts (genus Triturus). Proceedings 4th OGM of Societas Europea Herpetologica, pp.369-372, Nijmegen: Faculty of Science.

Zajc, I., and J.W. Arntzen. 1999. Phylogenetic relationships of the European newts (genus Triturus) tested with mitochondrial DNA sequence data. Contributions to Zoology 68(2): 73-81.

Other Resources
Amphibians and Reptiles of Europe
amphibiaweb.org
Great Crested Newt Conservation
amphibian.co.uk/alpine.html
amphibian.co.uk/triturus_vittatus/banded.html
perso.club-internet.fr/ccage/triturus_vittatus.htm
webfaculty.aub.edu.lb/~rsadek/Research/Triturusvittatus.htm
agurodela.de/archiv/tagungen_2000.php3
darkwave.org.uk/~caleb/trispe.html
wallonie.be/dgrne/sibw/especes/ecologie/amprep/triturus.alpestris.html
wallonie.be/dgrne/sibw/especes/ecologie/amprep/triturus.cristatus.html
amphibien-net.de/Berichte/Christopher%20Schwerdt/bergmolch.htm
kent.gov.uk/sp/biodiversity/pdf/KBAP706G.PDF
bbc.co.uk/nature/wildfacts/factfiles/478.shtml
serpenti.it/schedeanfibi/triturusmarmoratus.htm
amiciinsoliti.it/tritonieuropei.html