Facultative paedomorphosis is an environmentally induced polymorphism that is well known for many caudate species including newts. Although facultative paedomorphosis has been documented in some smooth-newt populations, records of entirely paedomorphic populations outside the Balkans are limited. Here we present the first evidence of a paedomorphic population of the smooth newt in the Czech Republic with discussion of potential causes that need to be further tested

Oldřich Kopecký

Veronika Javůrková

Václav Gvoždík

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First evidence of a paedomorphic population of the smooth newt (<em>Lissotriton vulgaris</em>) in the Czech Republic

Gvoždík, Václav

Javůrková, Veronika

Kopecký, Oldřich

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Acta Herpetologica 8(1): 53-57, 2013
First evidence of a paedomorphic population of the smooth newt 
(Lissotriton vulgaris) in the Czech Republic
Václav Gvoždík1,2, Veronika Javůrková3,4, Oldřich Kopecký5,*
1 Department of Zoology, National Museum, Cirkusová 1740, 193 00 Prague, Czech Republic
2 Laboratory of Molecular Ecology, Institute of Animal Physiology and Genetics, Academy of Sciences of the Czech Republic, 277 21 
Liběchov, Czech Republic
3 Department of Zoology, Faculty of Science, Charles University in Prague, Viničná 7, 128 44, Prague, Czech Republic
4 Institute of Vertebrate Biology v. v. i., Academy of Sciences of the Czech Republic, Květná 8, 603 65 Brno, Czech Republic
5 Department of Zoology and Fisheries, Faculty of Agrobiology, Food and Natural Resources, Czech University of Life Sciences, Kamýcká 
957, 165 21 Prague, Czech Republic. * Corresponding author. E-mail: kopeckyo@af.czu.cz
Submitted on 2012, 20th December; revised on 2013, 13th March; accepted on 2013, 27th March. 
Abstract. Facultative paedomorphosis is an environmentally induced polymorphism that is well known for many 
caudate species including newts. Although facultative paedomorphosis has been documented in some smooth-newt 
populations, records of entirely paedomorphic populations outside the Balkans are limited. Here we present the first 
evidence of a paedomorphic population of the smooth newt in the Czech Republic with discussion of potential causes 
that need to be further tested.
Keywords. Facultative paedomorphosis, urodeles, “Triturus”, Central Europe.
Facultative paedomorphosis, the case of phenotypic 
heterochrony when some sexually mature individuals 
from a population (or some populations of a species) 
retain some states of larval morphology, has been widely 
documented for many caudates (review in Denoël et al., 
2005; Laudet, 2011). In Europe, facultative paedomor-
phosis regularly occurs in newts (former genus “Tritu-
rus”) with high frequency in the Alpine newt (Ichthyosau-
ra alpestris), especially in some areas of the Balkans and 
Alps (Denoël et al., 2001). Beside the Alpine newt pae-
domorphosis has also been recorded in other newt spe-
cies (summarized by Wells, 2007): Lissotriton helveticus, 
L. italicus, L.  vulgaris, Triturus cristatus, T. macedonicus, 
and Ommatotriton ophryticus (Başkale et al., 2011). In the 
smooth newt (L. vulgaris), this phenomenon is also rela-
tively common, although usually only a small part of a 
population or single individuals are paedomorphic. Cases 
of peadomorphosis in the smooth newt have been report-
ed from various places within almost the whole distri-
bution range (review in Schmidtler and Franzen, 2004), 
but entirely paedomorphic populations of L. vulgaris are 
rarely reported (Dely, 1967; Dolmen, 1981; Denoël et al., 
2009). One such population from northern Hungary was 
described like a different subspecies, Triturus vulgaris 
tataiensis (Dely, 1967), but validity of this subspecies was 
rejected (Raxworthy, 1990).
Here, we report a presumably almost completely 
paedomorphic population of L. vulgaris from the Czech 
Republic. This is the first formally reported recent record 
of urodelan paedomorphosis for the country. Paedomor-
phic smooth newts (Fig. 1) were discovered (by VG and 
VJ) on 28 April 2012 in a water reservoir in Stará Lysá 
(50°13’31.6”N, 14°47’57.1”E, ~190 m a.s.l.). The finding 
place is situated in “Polabí”, which is a lowland area along 
the Labe (Elbe) River in the Central Bohemian Region. 
This part of the Czech Republic is one of the warmest, 
with dry and mild winters and average annual tempera-
ture between 8-9 °C (Quitt, 1971). The local landscape 
54 Václav Gvoždík et al.
	  Fig. 1. Smooth newts observed in the Stará Lysá water reservoir, Czech Republic: A) Paedomorphic male in nuptial colouration; B) paedo-
morphic female in nuptial colouration; C) close-up view on another paedomorphic male showing the structure of gills; D) aquatic imma-
ture specimen with open gill slits (A-D photographed on 7 May 2012); E) paedomorphic male, and F) female later in the season with 
already reduced gills; G) different pair (adult male and female) demonstrating variation in the level of persistence of larval traits; H) imma-
ture paedomorphic specimen with large gills; I) metamorph-like aquatic immature specimen with open gill slits (E-I photographed on 23 
August 2012); J) Stará Lysá water reservoir.
55Paedomorphic smooth newts in the Czech Republic
has been historically used for agriculture and so the 
locality is from three quarters surrounded by crop fields 
and from one quarter by a relatively dry cultivated pine 
forest (cf. Covaciu-Marcov et al., 2011).
Newts inhabit a water reservoir (originally used for 
fire protection) sized 25 × 15 m with maximal water 
depth of 1.5 m made from concrete blocks (Fig. 1J). 
Shorter sides of the reservoir are formed by vertical 
walls while longer sides slope in approximate 45°. The 
reservoir is supplied from a deep borehole (~60 m), and 
water is changed approximately once per three years and 
occasionally refilled (last change in 2010; V. Vaněk, pers. 
comm.). No fish were observed. The reservoir is with-
out vegetation, only filamentous algae grow later in the 
season over submerged objects and newts use them for 
wrapping eggs. Apart from newts, several adult individu-
als of marsh frogs (Pelophylax ridibundus) were observed 
and their numerous offspring later in the season. The 
water is rich on planktonic micro-crustaceans. Generally, 
the reservoir meets conditions for occurrence of paedo-
morphic newts: does not freeze to the bottom in winter, 
do not dry in summer, few predators (no fish), abun-
dant food resources (Litvinchuk et al., 1996; Denoël and 
Poncin, 2001). 
The locality has been visited in several occasions 
(April, May, July, August, September, October 2012), and 
opportunistic visual surveys and dip-netting have been 
done. Paedomorphic individuals were observed during all 
visits, with a visually assessed abundance in the spring. 
During the assumed peak of reproductive season (7 May 
2012), we observed 70 adult smooth newt individuals 
when counting specimens visible along the reservoir walls 
during one walk around the tank. Except of two seem-
ingly metamorphic females (but without success to catch 
them and inspect for morphology) and one almost meta-
morphic juvenile (but with open gill slits; Fig. 1D), all 
individuals were clearly paedomorphic (Fig. 1A-C) and 
only three of them were males. Sex ratio bias toward the 
females is common in paedomorphic individuals of this 
species (e.g., Kalezić and Džukić, 1986; Litvinchuk et al., 
1996; Çiçek and Ayaz, 2011), however the observed ratio 
is unusual. Also situations when paedomorphic smooth 
newts outnumber metamorphic individuals have been 
only rarely reported (e.g., Kalezić and Džukić, 1986; Rot-
Nikcević et al., 2000) compared to the more common 
metamorph-biased population ratio in this species (e.g., 
Banks, 1985; Kalezić and Džukić, 1985; Çiçek and Ayaz, 
2011). However, it is important to mention that we can-
not make clear conclusion about the ratio of peadomorphs 
vs. metamorphs in our smooth-newt population as we did 
not make a quantitative survey. Potential bias toward pae-
domorphs might be also caused by differences in behav-
iour of the morphs with respect to the relatively high water 
depth. During the spring visits, many paedomorphic newts 
were resting in the water column along the walls to 50 cm 
below water surface and were easily counted.
To observe reproductive behaviour, three paedomor-
phic individuals (one male, two females) were taken into 
captivity (7 May 2012). Water temperature in the aquari-
um ranged 16.7-19. 4 °C and newts were kept under nat-
ural light conditions. The newts were housed together in 
an aquarium (50 × 25 × 25 cm) filled by settled tap water 
and with several waterweed plants (Egeria densa). Dur-
ing the following days we had observed the male’s court-
ship behaviour (fan display), which is in newts similar 
for both morphs (Denoël, 2002), and deposition of eggs 
by both females. As we expected (cf. Banks, 1985; Litvin-
chuk et al., 1996; Litvinchuk, 2001), the male metamor-
phosed after 19 days, followed by the females which met-
amorphosed after 42 and 45 days in captivity, respectively 
(including closed gill slits).
Later in the season (23 August 2012), all captured 
adults (Table 1) were already almost without crests and 
fin appendices, and with small, sometimes rather rudi-
mental gills, nonetheless, still in aquatic phase and with 
open gill slits (Fig. 1E-G). Immature individuals, accord-
ing to the body size and shape and level of development 
of the cloaca, were of two types: some had large gills and 
fins (Fig. 1H), while some others resembled the adults, 
but being substantially smaller, by having rudimentary 
gills or only open gill slits (Fig. 1I). Beside these paedo-
morphic specimens (open gill slits as a common charac-
ter), some typical larvae were also observed. It seems that 
adult individuals reduce their larval traits (gills) toward 
the winter (cf. Denoël, 2003). However, it is currently 
unknown if the same individuals grow the larval traits 
again in the next season. A study of the demographic 
structure is currently ongoing using the capture-mark-
recapture method and shall shed more light onto this and 
other questions in near future.  
Despite relatively strong evidence for the benefi-
cial role of facultative paedomorphosis as an alternative 
Table 1. Morphometric characteristics in cm (SVL - snout-vent 
length, Lt - length of tail, Dbl - distance between front and hind 
limbs) of adult peadomorphic smooth newts (females n = 7; males 
n = 3) from Stará Lysá, Czech Republic.
    mean min. max. SD
females SVL 3.45 3.13 3.82 0.27
Lt 3.10 2.80 3.47 0.22
Dbl 1.69 1.41 1.93 0.18
males SVL 3.68 3.34 4.01 0.34
Lt 3.50 3.19 3.82 0.32
  Dbl 1.67 1.57 1.77 0.10
56 Václav Gvoždík et al.
ontogenetic pathway in many tailed amphibians, impor-
tance and influence of particular factors lying behind its 
evolution is not fully resolved. Whilst paedomorphosis 
can be a highly heritable trait (e.g., Harris et al., 1990; 
Voss et al., 2003), the occurrence of facultative paedo-
morphosis in individuals and its maintenance in a popu-
lation is most probably a result of interplay between gen-
otype and heterogeneity of the environment (reviewed 
by Denoël et al., 2005). Among many factors, streambed 
or pond-bank structures (Bennett and Chippindale, 
2006), slow desiccation rate (Semlitsch et al., 1990), and 
high food availability in water (Ryan and Semlitsch, 
2003) have been observed to induce occurrence of pae-
domorphs and the maintenance of resulted polyphenism 
in a population. All the above-listed factors, particularly 
high food availability and resource partitioning between 
different phenotypes seem to be feasible as possible fac-
tors responsible for the maintenance of paedomorphosis 
in our smooth-newt population. Retention of larval feed-
ing apparatus in paedomorphs may allow more efficient 
foraging on plankton and aquatic prey (Whiteman et 
al., 1996; Denoël, 2004). As a result, both microhabitat 
segregation and utilization of different food has been 
observed in particular morphs, with paedomorphs main-
ly foraging on plankton and distributed in both shallow 
and deep parts of water column compared to foremost 
shallower distributed metamorphs (Denoël and Joly, 
2001; Denoël and Schabetsberger, 2003). As the water 
reservoir is rich in planktonic micro-crustaceans and 
areas of deep water column, we suppose the predomi-
nance of paedomorphs in our population to be inherent 
in these factors. It is however worth noting that predom-
inance of paedomorphs in our smooth newt population 
could also be a result of an environmental-hormonal 
interplay, as both stress and steroid hormones were 
observed to significantly affect metamorphosis of larvae 
and paedomorphs to the adult stage (reviewed by Lau-
det, 2011). Detailed endocrinological studies are needed 
to shed light on these proximate mechanisms.
ACKNOWLEDGEMENTS
The study was carried out in accordance to the permit 
SZ-092744/2012KUSK/3 issued by the Regional Office of the 
Central Bohemian Region of the Czech Republic and with a 
support by Vladimír Vaněk (the mayor of Stará Lysá). We also 
would like to thank Jiří Moravec (National Museum in Prague) 
and an anonymous reviewer for valuable discussion and com-
ments, respectively. The research was supported by the Ministry 
of Culture of the Czech Republic (DKRVO 2013/13, National 
Museum, 00023272). VJ is also grateful to the Institutional 
Research Support of the Charles University in Prague and pro-
ject SVV-2012-265-206. 
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First evidence of a paedomorphic population of the smooth newt (Lissotriton vulgaris) in the Czech Republic

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First evidence of a paedomorphic population of the smooth newt (<em>Lissotriton vulgaris</em>) in the Czech Republic

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