I

Vol. 6, pp. 120-131

Asiatic Herpetological Research

June 1995::

On the Inheritance of the Mid-dorsal Stripe in the Iranian Wood Frog
(Rana macrocnemis)

DAVID N. TARKHN1SHVILI
Department of Biology, Tbilisi State University, University Str. 2, Tbilisi 380077, Georgia
Abstract. -The frequency of the phenotype Striata (presence of a light mid-dorsal stripe) is often used in
Simple hereditary nature of this character was establishedinvestigations of anurans.

population

Nevertheless, animals with a poorly developed stripe can be found in populations of somein the frog, Rana macrocnemis, the expression
species, together with non-striped specimens and clear Striata,

experimentally.

of the mid-dorsal stripe varies continually. Poorly striped frogs prevail in some populations. We examinedmorph distribution among froglets obtained from seven R. macrocnemis pairs whose eggs and larvae
developed at 18, 25, and 29°C. Non-striped froglets predominate in the offspring of non-striped parents andwith a clear
striped in the offspring of striped frogs. Non the less, all phenotype spectra, from specimens

The

distribution iscontinual

quite
morph
stripe to non-striped animals, are represented in almost all groups.
When the developmental temperature increases, the frequency ofand the character varies as a quantitative one.

the morph predominating in optimal conditions (18°C) declines with the increasing frequency of poorly
conditions shadows genetic
striped froglets. Modifications of the character under the influence of thermal
the morph distribution in froglets was foundspecifics of sibling groups. Similar influence of conditions on
distribution
in natural populations. Morphamong newly emerged froglets affects morph distribution of adults
in the populations two years later

Key words Amphibia, Anura, Ranidae, Rana macrocnemis, inheritance, Georgia

Introduction
Specimens with a high mid-dorsal stripe
can be found among a lot of anuran species.
This character (phenotype Striata) appears in
representatives of systematically distantanuran families. Sometimes a light stripe
appears as a specific character (e.g. in thefrog Rana amurensis or among some boreal
toads). In other species, Striata appears as amorph and within the same population nonstriped as well as striped animals can befound. Prevalence of this morph is species-

specific.

For

instance,

Discoglossus

among
some
or

if

pictusEleutherodactylus stripedanimalsandTrueb,
1986),(Duellman
predominate
in

Rana

temporaria they are sparse(Istchenko, 1978) and in Occidozyga laevis
only a few striped juveniles could be found(our unpublished data).

Anuran species among which a middorsal stripe appears can be divided into twogroups independently of their systematic
position. The first group includes species inwhich the mid-dorsal stripe appears as avery clear and discrete character, and striped© 1995 by Asiatic Herpetological Research

animals can be divided from non-striped
ones easily. There are, for example, some
representatives of the genus Rana:: Rana cf.
esculenta, R. nigromaculata, brown frogs
R. sylvatica, R. arvalis (Istchenko, 1978).
The second group includes species havingor
always softened stripe,characterized by the absence of a clear
border between the stripe and ground color
These include someof dorsal surface.

oftenbrown frogs (/?. temporaria, R. chensinesis,
R. macrocnemis) among others. Specimens
with a clear mid-dorsal stripe, as well aswith a softened stripe, can be found in R.
(Istchenko, 1978).
Moreover, in populations of these species,
the degree of development of the mid-dorsal
If numerous
stripe varies continually.

macrocnemis only

specimen samples are taken, a range fromquite unstriped to very clear Striata-s may beconstructed.

These extreme phenotypes

may be connected by animals with softened
stripe (Pseudostriata phenotype group).

Inheritance of the Striata phenotype was
examined experimentally. It was establishedthat, in different anuran families, thehereditary mechanism may be similar andApril 1995

Vol. 6, p. 121

Asiatic Herpetological Researchdependent on the effect of the fullydominant autosomal gene. It was shown,for instance, on Eleutherodactylus ricordii(Goin, 1947), Rana nigromaculata (Morya,1952), R. sylvatica (Browder et al., 1966),
R. arvalis (Stchupak, 1977; Stchupak andIstchenko, 1981), R. ridibunda (Berger andImietowski, 1982).
Simple hereditarynature was established for the coloration of
the mid-dorsal stripe of -4cm crepitans, aswell (Pyburn, 1961). That is the reason ofwide exploitation of the character in
population-genetic investigations, especiallyof frogs from the genus Rana (Stugren,
1966; Fishbeck and Underhill, 1971;
Istchenko, 1978; Pikulik, 1985; Gogoleva,
1985; Shibata, 1988; Kubantsev andNonetheless, thePeskova, 1989 etc.).
oftheStriatamorph has beenfrequency
used as the index of isolation degree(Stugren, 1966; Pikulik, 1985; Shibata,1988) or as the instrument of the natural
selection investigations (Fishbeck and1978;
1971; Istchenko,
Underhill,
Vershinin, 1987 etc.). Presence of a poorly
expressed stripe in some species made ususe mid-dorsal stripe in the phenetical
Inheritance ofinvestigations with care.
Pseudostriata is not clear. For R.

Materials and Methods

Experiment. Adults of R. macrocnemis(7 pairs) were obtained in 1989 from ahibernation site in Satovle Mountain ridge(eastern Georgia, surroundings of Tbilisi).
Mid-dorsal stripe expression differs inWedivided frogs
different specimens.
inones(Maculata;
conditionallynon-stripedM), specimens with poor enlightment in the
middle- or hind part of the dorsal surface(Pseudo-pseudostriata; PPS), with poorly
developed (Pseudostriata; PS) or clear

mid-dorsal stripe.
The
the
of
parents (maledescription
phenotype/female phenotype) is: pair 1M/M; 2-M/M; 3-M/PPS; 4-M/PPS; 5-PS/M;
6-M/PS; 7-PS/S. We selected 3 groups of
40 eggs from each of the 7 clutches
obtained.
Groups were placed in 40-1
withaeration and stable temperature
aquariaof 18, 25 and 29°C. Tadpoles were fed
boiled spinach. Half of the water volume
was renewed every second day. After(Striata;

S)

metamorphosis, the complete phenotype ofeach froglet (M, PS or S) was described.
Therefore, 20 experimental sites were

limnocharis, Moriwaki (1953) supposed
polyallelic inheritance; there are no data forother species. In any case, connections
between characters Pseudostriata and Striata
must be present. Continual connections

elaborated (tadpoles from the 7th clutch

between morphs, e.g. in R. macrocnemis

in a local

makes possible modifications of stripe
development under environmental

from Borjomi Canyon (central Georgia), theratio of phenotypes M, PPS, PS and S wereconditions, like most other quantitative
characters. Variations of mid-dorsal stripe
expression during the life span of individual

separately analyzed theinvestigated.
distribution of different morphs in adult

Rana temporaria specimens was noted(Heran, 1986). Earlier (Tarkhnishvili andMamradze, 1989; Mamradze, 1990) we
showed that Striata and Pseudostriata
phenotypes frequency in R. macrocnemis
sibling groups can be modified as a result of

temperature variability by which larval
development takes place. This present
article describes the results of the moredetailed experiments and field observations
on the Iranian wood frog populations.

developed

under

29°C died before

metamorphosis had begun).
Field Data.

During 1989, 1990 and 1991

population of R. macrocnemis

Weand in 16 groups offromdifferent ponds andfroglets emerged
included
20-44 froglets).
(eachsamplepools
There are more than 60 spawning sites in thefrogs, juveniles,

population investigated, but every year 2 or
3 water bodies ensure about 90% of thewhole generation. Comparative part of eachwater body was established as well as thenumber of" clutches deposited in each ofthem (Tarkhnishvili, in press).
For the comparison of morph distribution
webetweendifferent
samples,Vol. 6, p. 122

TABLE 1.

-test).

At 18°C

Asiatic Herpetological Research

January 1995

Significance of the differences in morph distribution between experimental groups (values of X

January 1995

TABLE 2.
Groups

Asiatic Herpetological ResearchInter-group differences in the frequency of each morph separately (F-test).

Vol. 6, p. 123

TABLE 3.

January 1995

Asiatic Herpetological ResearchVol. 6, p. 124

Significance of the phenotype frequency changes under the thermal conditions.

Difference of thermal conditions between
Sibling

Group

1

M

8°C to 25°C
(PS)

S

25°C to 29°C

M

2

4
5

6
7

1

S

M
++

1

3

(PS)

-

8°C to 29°C
(PS)

++

S

Asiatic Herpetolo^ical Research

January 1995

TABLE 4.

Vol. 6, p. 125

Significance of the differences in morph distribution between groups emerged
1 1 (1989) and 12 (1990), and that which emerged from another site (values of the X-test).

la

2a

3a

from the sites 9,small water bodies were described. Results
are shown on Fig. 3. Because of the small
volume of sample differences between them,appears non-significant in most cases.
Only froglets from the water bodies 9 andit

11,moreover

June 1995

Asiatic Herpetological ResearchVol. 6, p. 126

from

12

(1990)

aresignificantly different from most othergroups (Table 4). Spawning sites 9 and 12
are well warmed pools, filled withvegetation. Daily temperature during thedevelopmental period often exceeds 30°C.
Among the froglets that emerged from thesepools, the non-striped morph predominates.
The percent of the S-morph (in site 9, thismorph is absent), as well as PS and even

PPS, is low. Inversely, site 2 is selfVegetation is
flowing and shadowed.
absent and the water temperature do not
exceed 20°C, even during the warmest days.
PS and PPS morphs predominate here, andthe percent of the S-morph exceeds 20.
There is the question of whether thedifferences noted above can reflect non-random distribution of the parent's morphs
along the different spawning sites. On the
one hand, the displacement of warmed andshadowed sites are irregular in the habitat
examined, and striped frogs have equal
possibility of spawning in different water
bodies.
On the other hand, only a few
clutches (4 and 7 correspondingly) were
deposited in sites 9 and 11 during 1989.

Morph distribution in the froglets could be
the consequence of random processes as aresult. But, as many as 93 clutches werestructure between froglets that emerged fromdifferent water bodies as well as between theyoung generation and adult animals. The
stability of the morph composition of groups
emerging from the same site, but in different
waters, is additional evidence.

Above, we told about the increase of thenon-striped morph in 1991 to 65%. It may
be connected with the increasing of thecomparative part of warmed pools during
previous years. For instance, spawning site
5 produced at least 45% of the total numberof the 1989 generation (about 20,000). In
that time, the frequency of M-morph
approached 55% and not one froglet with aclear stripe was found. Most of the otherproducing metamorphosis in 1989
were warmed. Shadowed site 2 (17%and 23% S) produced not more than 25% of
sites

M

the generation.

In

1990, the situation

changed in Maculata's favor still more.
There were 68% of froglets of thisphenotype that emerged from site 12 (site 12
produced about 70% of the generation
numbering 44,000 froglets). Perhaps in the
future, we can suppose still more increase ofnon-striped morph's comparative part and a
decline of the striped animals' frequency inthe investigated population.

Discussion

Twodirections

of

the

S-morphfrequency investigations in amphibian
population can be noted. The first is thedeposited in site 12 (1990). It is quite arepresentative sample of the population,
numbering 1037 adult females in all (1990).
This time, the frequency of non-striped
froglets emerged from site 12 (68%) isinstantly higher than M-morph frequency
among adult frogs (45%). Moreover,during 1991 in site 2, already 37 clutches
were deposited, but the frequency of S-

species. The second one is to study the
dynamics of polymorphism of separate
populations including age-dependentThechanges of morph distribution.
summarizing bibliography following must

morph among

froglets even slightly
increased. Therefore, we can suppose non-Geographic variation. There is no goodevidence of clear clinal variations ofrandom reasons of the morph distribution in
the groups of froglets emerged fromdifferent spawning sites. The frequency of
one or another morph depends on thephysical conditions under which larval
development takes place. This is argued byfrequency of the morph Striata in a single
species. Stugren's conclusion (1966) about
increasing of frequency of striped frogs in
Rana arvalis toward the east was refuted by
Istchenko (1978). In some cases (i.e. R.
arvalis and R. temporaria in Belorussia), thethe significant differences in phenotypic

analysis of geographic variations of morph
frequency in populations of different

be noted.

June 1995

Asiatic Herpetological ResearchVol. 6, p. 127

percent of striped frogs increase toward thenorth (Istchenko, 1978; Pikulik, 1985), but
in the populations of R. arvalis from theUrals or of R. semiplicata from the Russian
Far East, the inverse situation was noted

elective elimination of larvae accordingly to(Toporkova, 1965; Istchenko, 1978). On
other hand, irregular but clear

rates (Schvarz and Istchenko, 1968). On
the other hand, the adult Striata-s are lessthetheir genotypes.
For instance, larvae ofStriata genotype are more vulnerable to thehigh density and shortage of the oxygen
because of the abnormally high metabolic

S-morphsusceptible than other frogs to drying andfrequency are common in different species'

are characterized by high migration ability.

populations (Masalikin, 1985; Panchenko,
1985; Shibata, 1988 etc.). The frequency of
the S-morph in R. arvalis may increase
under the hardening of antropogenous

Correspondingly, in some ecosystems withunfavorable conditions, striped specimens

interpopulational differences in

Therefore,pressing (Vershinin, 1987).
frequency distribution of phenotypes in theseparate populations of some species depend
highly on the climatic or microhabitat
conditions but not on historical reasons.
Interestingly, when the coexistence of R.
temporaria and R. arvalis populations takes
place (in the Middle Volga) the sameenvironmental conditions ensure parallelvariations in population morphology

predominate (Istchenko, 1978; Vershinin,1987).

Therefore, the influence of thedevelopmental conditions on the ratio of
striped and non-striped morphs in thegeneration may be considered wellgrounded. The role of the developmentalsites' type is obvious and our data are in
All theaccordance with this fact.
researchers

agree that the phenomenaontheelective elimination.
depends(Lebedinski et al., 1989)

Intrapopulation dynamics.
Comparative frequency of different morphs
may vary between years and separate
generations (Istchenko, 1978). Influence ofthe pond conditions on the morph
distribution in R. macrocnemis froglet
groups was described by this author.
Besides, morph distribution in thegeneration could be changed in relation toanimal age.
For some R. arvalis

populations, decreasing of S-morph
frequency in adult animals was establishedin comparison to juveniles (Pikulik, 1985;
Vershinin, 1987).

Inversely, in thesamespecies, increasing of the comparative partof striped animals was recorded for the older

age groups (Istchenko, 1978). Therefore,
the morphological features of the population
may be changed rapidly and instantly byenvironmental conditions. There is themean adaptation on the population level.
Istchenko (1978) supposes that rapid
reactions of morph distribution on theenvironmental conditions take place becauseof the elective elimination of animals. Rapid
modifications of the genotypes' distribution
is the result of this process. Selection may
be connected not only with preferences ofstriped or non-striped frogs but also with

It is not possible in practice to only
conduct field investigations. What is thepart of individual modifications in theprocess of morph distribution changing?
High natural mortality, especially during thelarval stages, does not allow us to exclude
the effect of natural selection. Thus, eachexplanation depends on a prior opinion of
the concrete researcher. For instance, infield conditions, information principally
similar to our experimental data was
obtained by Stchupak (1975). Increasing
the frequency of genotype S or
has
inverse reaction of larval grouping as theresult: the comparative part of the dominantphenotype decreases by the completion ofmetamorphosis. The author explains thisfact as the result of frequency-dependentM

selection.

Part of the phenotypic modification
during the early development can beestablished only in experiments where larvalmortality can be neglected. The experimentdescribed above illustrates the main part ofmodifications taking place during larval
development. We conclude that in somespecimens displacement of the characters in
the road of alternative variants could bechanges as a result of temperature changes.

Vol. 6, p. 128

Asiatic Herpetological Research

M

PS

January 1995

S^;-

lw ** 4rS fj£
v

9

1

-400

FIG. 4. Distribution of the pigmental cells in the mid-dorsal area of froglets that have different phenotypes.
Magnification is X400.

brings us to suppose that morphological
specifics of froglet groups is provided not
only by elective elimination but by
phenotypic modifications as well. What isIt

the comparative part of this latter could beshown by further investigations.
Experimental results described here andevidence in favor of the main part ofphenotypic modifications are obtained
macrocnemis, notnamely for R.
surprisingly. This is the only boreal brownwhich non-striped and stripedmorphs are connected by continual interfrog in

gradations (Istchenko, 1978) and practicallythe character "presence of the mid-dorsal
stripe" varies as the quantitative one. On the
one hand, it may be connected with thespecific nature of the character inheritance inthis species; but it is less possible. Morerealistic is taking in our opinion that every

character, both quantitative and qualitative,

determinedbyquantitativeThusmorphogenetic processes.
aredevelopment of coloration of the mid-dorsal
stripe in Acris crepitans is determined by thesize of pigmental cells and geneticalspecifics of morphs may be shaded (Gray,
1972). We observed the dorsal surface of

R. macrocnemis froglets with different
coloration patterns under a binocular.
Presence of the mid-dorsal stripe depends
on the rate of pigment concentration
declining toward the dorso-medial line (Fig.
4). The lower density of the pigmental cells
along the mid-dorsal line in all froglets canbe observed. But to distinguish the stripe
with the naked eye, the gradient ofpigmental cells concentration must be
expressed enough or exceed a certain
threshold. Morphogenetically, the presenceof the gradient is determined by the kinds ofmelanophorisation of the dorsal surface
before metamorphosis:
the mid-dorsal

surface is filled in by pigmental cells at theend. The phenotype of the froglet: S, PS,
PPS orwhich appears to correspond withM

the degree of filling in the mid-dorsal

surface by melanophores just beforeThe
metamorphic climax has begun.
hereditary mechanisms keep the main part of
this
but
of
process
regulation
morphogenesis by the environment isimportant, too.
Perhaps comparative
significance of the environment increases in

non-optimal conditions.

June 1995

Asiatic Herpetological Research

Perhaps in the base of phenotype
formation in species characterized by severe
alternative variation of the character (e.g. R.
arvalis or R. ridibunda) the samemorphological processes occur. But, inthese species, inter-gradations between Sand M- morphs are absent. It may depend
either on greater part of hereditary in stripe
determination among these species or on amore expressed threshold level. It must be
explained. Developmental processes which
form the ground of coloration patternformation may finish in some developmental
stage before metamorphosis has began.

Vol. 6, p. 129

stripe formation also affect intensification of
If intensity of theseenergetic exchange.

neithertheirchange,morphological nor physiological resultsappear. There is an interesting example ofprocesses

phenomena when modifications duringthethe separate animal's life results similarly toa short-term selection at the population level.

appears as one of the hard moments in
micro-evolutional investigations but would
not be neglected.
It

AcknowledgmentsOneor another phenotype could be formed
depending on if the process exceeds somethreshold level or not. Depending on theexpression of this threshold or prolongationof final stage of color formation (when basic
processes are completed) takes place either
in continual distribution of morphs (as in R.
macrocnemis) or in bi-modal distribution (asin R. arvalis).

The scheme described above, as we candoes not exclude identity of theinheritance nature of the Striata in R
see,I

greatly

appreciate

my

wife,

R.

Mamradze, for her help in the elaboration oftheI
also appreciate J.
experiment.
andS.
Tzabadze
for their help in
Manukjan
the experimental and field work, and to J.

Ilieva for reading the manuscript.

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species with continual distribution of the
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