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Group Title: Florida. University, Gainesville. State Museum. Bulletin. Biological sciences
Title: Pattern variation in the frog Eleutherodactylus nubicola Dunn
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Permanent Link: http://ufdc.ufl.edu/UF00001559/00001
 Material Information
Title: Pattern variation in the frog Eleutherodactylus nubicola Dunn
Series Title: Bulletin of the Florida State Museum ; volume 5, number 5
Physical Description: 244-258 p. : illus., tables. ; 23 cm.
Language: English
Creator: Goin, Coleman Jett, 1911-
Publisher: University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 1960
 Subjects
Subject: Eleutherodactylus nubicola   ( lcsh )
Zoology -- Variation   ( lcsh )
Genre: bibliography   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
non-fiction   ( marcgt )
 Notes
Bibliography: "Literature cited": p. 258.
 Record Information
Bibliographic ID: UF00001559
Volume ID: VID00001
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: ltqf - AAA0867
notis - ACK0932
alephbibnum - 000440466
oclc - 05069538
lccn - a 61009073

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        Page 243
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        Page 245
        Page 246
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        Page 250
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        Page 252
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    Back Cover
        Page 259
Full Text




BULLET


IN


OF THE


FLORIDA STATE MUSEUM


BIOLOGICAL


SCIENCES


Volume 5


Number 5


PATTERN VARIATION IN THE FROG
ELEUTHERODACTYLUS NUBICOLA DUNN

Coleman J. Goin


UNIVERSITY


OF FLORIDA


Gainesville







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S Publisi ed 17 August 19.60


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PATTERN VARIATION IN THE FROG
ELEUTHERODACTYLUS NUBICOLA DUNN

COLEMAN J. GOIN 1

SYNOPSIS: An analysis is made of the color pattern variation in the Jamaican
frog, Eleutherodaciylus nubicola Dunn. A basic pattern, Mottled, may be over-
laid by no less than six modifying patterns as follows: Dorsolateral stripes, Mid-
dorsal stripe, Broad middorsal stripe, Picket, Interocular bar, and Pelvic spots.
It is possible to have more than one of these modifying patterns present in a
single individual; one specimen had, at one and the same time, Dorsolateral
stripes, Middorsal stripe, and Picket. Evidence is presented that these modifying
patterns are inherited in a mendelian manner and Dorsolateral stripes and Mid-
dorsal stripe seem to be dominant. Evidence is not yet available that will permit
us to determine if the other pattern modifiers are dominant or recessive.
It is pointed out that certain Rhacophoridae from Ceylon and Microhylidac
in the Papuan region not only have developed terrestrial breeding habits like
those of Eleutherodactylus, but that they have several of the pattern modifications
described for E. nubicola. The significance of the parallelism is at present un-
known.

With the possible exception of the African genus Hyperolius, the
neotropical frog genus Eleutherodactylus is the most "difficult" tax-
onomically of any of the large anuran genera. The reason seems to
be threefold-the phenomenal amount of speciation, the morphologi-
cal similarity of the species, and the intraspecific diversity of color
pattern. In earlier papers (Goin, 1947, 1950) I presented evidence
that much of this variability in color pattern is due to a series of
genes that are inherited in a mendelian manner. So little is known
of amphibian genetics that the determination of the genetic basis of
color pattern inheritance in these species is of interest in itself, but
more importantly, an understanding of it may help to throw light
on the evolution and relationship of species within this complex
genus, and possibly on the even more fundamental problems of evo-
lution in natural populations.
Eleutherodactylus contains nearly 250 recognizable forms. Many
of the species are rather variable in color pattern, with the same basic
pattern modifications occurring in species after species. The evi-
dence, discussed below, suggests that these pattern variations reflect
the segregations of a number of different genes (and their alleles)
which are homologous in the different species.

'The author is Professor of Biological Sciences, University of Florida, and
Research Associate, Carnegie Museum. Manuscript submitted 16 February 1960.







244 BULLETIN FLORIDA STATE MUSEUM,. Vol. 5

My interest in this problem first centered on E. ricordi planirostris,
a species with two phenotypes, which was introduced into Florida
and has been abundant locally for a number of years. At the sugges-
tion of my friend, W. Gardner Lynn, I later turned my attention to
E. nubicola of Jamaica, because this species is known to have a large
number of phenotypes and because its eggs can be collected easily
along the mountain trails in the Blue Mountains of Jamaica. In the
summer of 1948 I was able to collect 25 clutches of nubicola eggs which
produced 1002 hatchlings. Analysis of the data (Coin, 1950) indi-
cated that, as in planirostris, variability in nubicola is produced by
genes that modify the color pattern and that are inherited in a simple
mendelian manner. Additional data gathered on two related species,
alticola and pantoni, indicate a similar mode of pattern inheritance in
these forms. I returned to Jamaica in 1957 to gather additional ma-
terial on nubicola; 43 clutches, comprising 1341 hatchlings, were col-
lected on this trip. These, together with the previous material, make
a total of 2,343 young nubicola from 68 clutches of eggs now available
for analysis.
ACKNOWLEDGMENTS
In the course of this work I have become indebted to a great many
institutions and people. My field work in 1948 was financed by the
American Academy of Arts and Sciences, that in 1957 by the American
Philosophical Society. I obtained data on the variability of other
species in Jamaica in 1952 by field work done with the aid of a grant
from George R. Cooley. As is true of anyone doing natural history
work in Jamaica, I am deeply indebted to C. Bernard Lewis and his
staff at the Science Museum, Institute of Jamaica. The Forest De-
partment of Jamaica, through its Conservators, E. W. March, E. M.
Brown, and J. A. N. Burra, made the Forest Hut at Clydesdale avail-
able to me as a base of operations. My neighbor in the Blue Moun-
tains, Bonner E. C. Shekell, formerly of Chester Vale, St. Andrew,
rendered me a number of courtesies that made my stay in the moun-
tains more pleasant and profitable. My field companions, Dennis
E. Miller, Byrum W. Cooper, and Robert H. Barth in 1948, Albert M.
Laessle in 1952, and my wife and two children in 1957, helped appre-
ciably in the collecting of the eggs. My research assistant, Ronald
Baxter, aided in tabulating the results. My colleague, Kenneth W.
Cooper, contributed materially by his help in analyzing the data and
Henry W. Wallbrunn assisted in some of the statistical calculations.
Curt Stern most kindly took time from his busy schedule to go over the
manuscript and has made several helpful suggestions. As usual, I







COIN: PATTERN IN ELEUTHERODACTYLUS


am deeply obligated to my wife, Olive Bown Goin, for her skillful
assistance in typing and editing the manuscript. This work was com-
pleted while I was under tenure of a Grant (G-5628) from the National
Science Foundation. To all of the above I am deeply grateful.

Eleutherodactylus nubicola Dunn
Eleutherodactylus nubicola is a montane species found in the
Blue Mountains of Jamaica between the altitudes of 4000 and 6000
feet. Lynn (1942) described in detail its life history and embryonic
development. The slopes of the Blue Mountains are so steep it is prac-
tical to collect only beside the foot trails. The female lays from 26
to 75 eggs which are fertilized by a single male at the time of deposi-
tion. The eggs are not scattered, but are gathered together in a single,
compact bunch under a rock. Thus it is certain that each clutch is
the product of a single set of parents. The female remains with the
clutch during intraovular development. It is thus possible for one
to determine the phenotype of the female parent of many clutches of
eggs collected in the field, although at times the female escapes be-
fore her pattern can be noted.
A study of live individuals in the field and preserved specimens
in the laboratory indicates that there is a basic pattern, Mottled, with
at least six distinctive modifications as follows: Dorsolateral stripes,
Middorsal stripe, Broad middorsal stripe, Picket, Interocular bar, and
Pelvic spots. Data compatible with a simple mendelian interpreta-
tion arc now available for all of these patterns except Broad mid-
dorsal stripe and Pelvic spots.
In the discussion that follows, the locus for genes affecting Dor-
solateral stripe is indicated by S, that for Picket by P, that for Mid-
dorsal by M. A dominant gene at the S locus is designated by S, its
recessive allele by s, and correspondingly P and p, and M and m,
for the other loci.
The following data are now available for pattern inheritance
analysis:
(1) For the phenotypic ratios in the wild population, 300 specimens collected
along the mountain trails in the same area where the eggs were taken showed the
following patterns or pattern combinations (females with eggs are included):
Mottled-222
Dorsolateral stripes--32
Picket-21
Middorsal stripe-21
Dorsolateral stripes plus Middorsal stripe-2
Picket plus Middorsal stripe-2


1960








BULLETIN FLORIDA STATE MUSEUM


(2) Front mothers of known pattern 1544 hatchlings from 45 clutches of eggs:


Mother-Mottled (34 clutches)
Collected in 1948
26 Mottled, 20 Dorsolateral
stripes
25 Mottled, 28 Middorsal stripe
25 Mottled, 26 Middorsal stripe
26 Mottled
43 Mottled
58 Mottled
47 Mottled
42 Mottled
53 Mottled
21 Mottled
51 Mottled
33 Mottled
41 Mottled








Mother-Dorsolateral stripes (5 clu
Collected in 1948
31 Dorsolateral stripes
9 Mottled, 8 Dorsolateral
stripes
5 Mottled


Mother-Middorsal stripe (4 clutches)
Collected in 1948
29 Mottled, 16 Middorsal stripe
34 Mottled, 39 Middorsal stripe


Mother-Picket (2 clutches)
Collected in 1948
7 Mottled, 4 Dorsolateral stripes,
3 Picket, 4 Middorsal stripe,
3 Dorsolateral stripes plus
Picket, 5 Picket plus Mid-
dorsal stripe, 5 Dorsolateral
stripes plus Picket plus Mid-
dorsal stripe


Collected in 1957


33 Mottled,
5 Mottled,
19 Mottled,
3 Mottled,
43 Mottled,
26 Mottled,
16 Mottled,
53 Mottled,
13 Mottled,
22 Mottled
21 Mottled
13 Mottled
14 Mottled
29 Mottled
26 Mottled
36 Mottled
29 Mottled
39 Mottled
41 Mottled
11 Mottled
37 Mottled


Dorsolateral stripes
Dorsolateral stripes
Dorsolateral stripes
Dorsolateral stripes
Dorsolateral stripes
Middorsal stripe
Middorsal stripe
Picket
Picket


tchcs)
Collected in 1957
6 Mottled, 16 Dorsolateral stripes
21 Mottled, 30 Dorsolateral stripes


Collected in 1957
5 Mottled, 5 Middorsal stripe,
10 Picket
16 Mottled, 19 Middorsal stripe


Collected in 1957
13 Mottled, 11 Picket


Vol. 5








GOIN: PATTERN IN ELEUTHERODACTYLUS


(3) From parents of unknown phenotype 799 hatchlings from 23 clutches of
eggs:
Collected in 1948 Collected in 1957
5 Mottled, 4 Dorsolateral 4 Mottled, 4 Dorsolateral stripes
stripes 11 Mottled, 15 Dorsolateral stripes
23 Mottled, 22 Middorsal stripe 15 Mottled, 29 Dorsolateral stripes
48 Mottled 31 Mottled, 3 Dorsolateral stripes
60 Mottled 20 Mottled, 14 Dorsolateral stripes,
34 Mottled 10 Interocular bar, 11 Dorso-
39 Mottled lateral stripes plus Interocular
bar
13 Mottled, 16 Midorsal stripe
26 Mottled, 17 Picket
25 Mottled, 25 Picket
21 Mottled
29 Mottled
35 Mottled
41 Mottled
27 Mottled
20 Mottled
29 Mottled
26 Mottled
47 Mottled

Mottled
The Mottled pattern consists of a brown background overlaid by
a dark brown or black mottling. The most consistent element is a
dark W-shaped mark in the suprascapular region, extending from
above the arm on one side to above the arm on the opposite side. The
apex of the middle portion of the W is directed anteriorly and located
on the midline of the dorsum. The area immediately caudad to the
two posteriorly directed apices of the W is generally somewhat lighter
in color than the surrounding dorsum. A faint, wavy line of dark pig-
ment is discernible caudad of this lighter area. A dark interocular
bar is present. Intensity of pigmentation shows a great deal of indi-
vidual variation and generally tends to increase with age. In hatch-
lings the pattern is immediately apparent, but in large, mature indi-
viduals it is often obscured by a general darkening of the dor-
sum.
The main features of this basic pattern may be discerned in the
majority of the species of this genus. In E. nubicola it seems to be
the basic wild pattern upon which the pattern modifications discussed
below are superimposed.


1960








BULLETIN FLORIDA STATE MUSEUM


Figure 1. Diagrammatic sketch of Figure 2. Diagrammatic sketch of
Mottled pattern. Dorsolateral stripes pattern.

Dorsolateral stripes
In this pattern two broad cream-colored bands, one on each side,
originate on the posterior margins of the upper eyelids, pass poster-
iorly above the tympani directly backward through the arms of the
W, and terminate above the insertions of the hind limbs. These
cream-colored bands obliterate the arms of the W leaving only the
A -shaped portion between the two stripes in the suprascapular region.
This is one of the most widely distributed pattern modifications
in the genus. I have demonstrated fairly conclusively (Goin, 1947)
that in E. ricordi planirostris this pattern is determined by a dominant
gene that is inherited in a simple mendelian fashion.
No clutches of eggs of E. nubicola were taken for which the color
patterns of both parents were known, but indirect evidence for domi-
nance or recessivity may be sought by use of the Hardy-Weinberg
equilibrium formula
q2 SS : 2q(1-q) Ss : (l-q)2 ss
to determine the genotypic ratio in the wild population.
If we assume Dorsolateral stripes to be dominant, then in a pop-
ulation in which 84/300 showed Dorsolateral stripes,


Vol. 5








COIN: PATTERN IN ELEUTHERODACTYLUS


2q(1-q) + q2 = 34/300 = 0.1133
therefore
(1-q)2 = 0.8867 and 1-q = 0.9416
hence
q = 0.0584
in which case
SS = 0.0034 Ss = 0.1100 ss = 0.8867
If the assumptions necessary for the application of the Hardy-
Weinburg law are made, we can calculate the phenotypic constitu-
tion of the clutches of the offspring. Thus of 68 clutches of eggs only
approximately 0.46 clutches would be expected to be made up of
offspring with Dorsolateral stripes only, 14.08 clutches should con-
tain offspring both with and without Dorsolateral stripes, and 53.46
clutches should be made up of nonstriped individuals only.
Going through the same steps, but assuming Dorsolateral stripes
to be recessive rather than dominant, it can be calculated that of 68
clutches approximately 0.87 should contain only offspring with Dor-
solateral stripes, 20.44 clutches should contain both types of offspring,
and 46.69 clutches should contain no offspring having Dorsolateral
stripes.
These two sets of estimates are compared with the actual results
in table 1.
TABLE 1

Theoretical clutches Theoretical clutches
Dorsolateral stripes Dorsolateral stripes Actual
dominant recessive clutches
Dorsolateral stripes only 0.46 0.87 1
Mixed 14.08 20.44 16
Without Dorsolateral
stripes only 53.46 46.69 51

Total 68.00 68.00 68
x2 = 0.53* x' = 2.140
P = <0.5>0.3 P = <0.2>0.1
SLines 1 and 2 are summed for calculation of xt.

While these results give no certain statistical basis for choosing
between the two assumptions, I believe that it logically may be as-
sumed that the gene for Dorsolateral stripes is dominant in E. nubicola
because it has been shown to be dominant in both E. alticola (Coin,








250 BULLETIN FLORIDA STATE MUSEUM Vol. 5

1950) and E. r. planirostris (Goin, 1947). As Sturtevant (1948: 230)
has concluded, "there can be no doubt that, in general, related species
have essentially the same complements of genes."
The pattern of Dorsolateral stripes has been recorded in at least
the following species of Eleutherodactylus and is probably present
in many others as well.
alhipes intermedins polytychus
alticola fimori portoricensis
armstrongi lentus rhodopis
atkinsi ockendeni richmondi
audanti orcutti ricordi
bogotensis orientalis rufifemoralls
cuneatTu pantoni turquinensis
gossei pictissimus wienlandi
gundlachi planirostris

Picket
This pattern consists of a light
area on the dorsum, bounded an-
teriorly by the posterior margin
of the middle section of the W
mark. The lateral margins of this
light area apparently coincide with
the median margins of the cream-
colored bands in individuals with
Dorsolateral stripes. When both
Dorsolateral stripes and Picket are
present in the same individual, the
entire area between the stripes pos-
terior to the A-mark is occupied
by the light picket.
On the basis that 23 out of the
sample of 300 individuals of the
wild population had the Picket pat-
tern and 277 lacked it, an analysis
similar to that made for the inheri- Figure 3. Diagrmmatic sketch of
Picket pattern.
tance of Dorsolateral stripes may
be made for Picket (see table 2).
Here the evidence is consistent with the assumption that the gene
for Picket pattern is dominant.
I have found this pattern in only three other species: gossei, pan-
toni, and polytychus.








GOIN: PATTERN IN ELEUTHERODACTYLUS


TABLE 2

Theoretical clutches Theoretical clutches Actual
Picket dominant Picket recessive clutches
Picket only 0.22 0.40 0
Mixed 9.81 15.07 7
Without Picket only 57.97 52.52 61
Total 68.00 67.99 68
x2 = 1.074 x' = 6.003
P = 0.3 P = <0.02>0.01
Middorsal stripe
In adults showing this pattern, the dorsum is marked by a narrow,
cream-colored median line which extends from the snout to the vent
and there branches to continue along the posterior margin of each leg
to the sole of the foot, where it terminates at the juncture of the fourth
and fifth toes. Ventrally a narrow, cream-colored, median line extends
from the tip of the chin to the
crotch. This is the only one of the
patterns that is manifest on the ven-
tral side of the body. The midven-
tral portion of this line tends to be-
come obliterated with increasing '
size (hence age) but even in the
largest specimens a median line
on the chin and throat is still evi-
dent. In hatchlings the middorsal
portion of the stripe extends only
as far forward as the anteriorly-
directed apex of the W but in ma-
ture individuals it continues on to
the snout.
Using the same method of
analysis, on the basis that 25 in a
sample of 3(X) individuals of the
wild population had the Middorsal
stripe pattern and 275 lacked it,
similar calculations were made. Figure 4. Diagrammatic sketch of
The results are shown in table 3. Middorsal stripe pattern.
Some additional evidence may be obtained from the clutches of
females having Middorsal stripe. Of five such clutches, each con-
tained offspring showing Middorsal stripe. This is to be expected if


1960








BULLETIN FLORIDA STATE MUSEUM


the gene for Middorsal stripe is dominant; if Middorsal stripe is re-
cessive, each of these females must have mated with a male carrying
the recessive gene. Since only 25 out of 300 adults in the population
showed the pattern, then
25/300 = 0.0833 = (1-q)2
and
(1-q) = 0.2886 and q = 0.7114.
TABLE 3
Theoretical clutches Theoretical clutches
Middorsal stripe Middorsal stripe Actual
dominant recessive clutches
Middorsal stripe only 0.24 0.47 0
Mixed 10.62 16.12 11
Without Middorsal
stripe only 57.14 51.40 57
Total 68.00 67.99 68
x2 = 0.0021 x2 = 2.4915
P = >0.95 P = >0.1<0.2

The distribution of the genotypes in the population may accord-
ingly be estimated:
MM = 0.506
Mm = 0.411
mm = 0.083
1.000
Still assuming the gene for Middorsal stripe to be recessive, each
of the five females must have been mm in genotype. Since each
female produced both Middorsal striped and non-Middorsal striped
progeny, all five females must have mated with heterozygous males
(Mm). But genotypically Mm frogs make up only 0.411 of the popu-
lation, and the likelihood that, on chance alone, a given set of five
females would each be mated with an Mm male is only (0.411)5, or a
P of 0.012-a poor likelihood. It seems most probable, then, that
Middorsal stripe is also dominant.
This Middorsal stripe pattern occurs in at least the following
additional species:
abbotti baker lanciformis
alticola cundalli matudai
antillensis dimidiatus pantoni
auriculaloides gossei portoricensis
avocalis funori


Vol. 5








GOIN: PATTERN IN ELEUTHERODACTYLUS


Independent Assortment
For considering whether the three modifications of the basic Mot-
tled phenotype that have been discussed so far are determined by
genes that are multiple alleles, or whether these genes occur in differ-
ent loci, evidence is provided by the presence of three modifications
of the pattern in a single clutch. The most complex series of offspring
from a single clutch is the set of 31 young from a Picket mother which
contains 7 of the 8 combinations of patterns possible with these three
modifiers. The presence in one clutch of all three modifications plus
the basic pattern precludes the possibility that the three pattern
modifiers are all allelomorphic inter se or to the basic pattern gene.
Let us assume then that three different pairs of genes are involved,
and that the gene modifying the color pattern is dominant.
Since there are unmodified Mottled offspring in this clutch, it
would not have been possible for either parent to have been homozy-
gous for Dorsolateral stripes, or Picket, or Middorsal stripe. As both
Dorsolateral stripes and Middorsal stripe occur among the offspring
in approximately a : 1 ratio, and as the mother showed neither, the
dominant genes for both of these patterns should have been present
in the father. Since Picket is present in the mother and in approxi-
mately one-half of the young, the dominant gene for this character
should not have been present in the father. Thus, on the basis of
the argument given above, the mother must have been heterozygous
for Picket and homozygous recessive for Dorsolateral stripes and Mid-
dorsal stripe, and the father must have been heterozygous for both
Dorsolateral stripes and Middorsal stripe, and homozygous recessive
for Picket. There should be, according to the laws of chance, a nearly
equal distribution of the eight possible phenotypes in the offspring.
The theoretical phenotypical composition of a clutch from this cross
is compared below with the actual ratio of the 31 offspring of the
female with Picket pattern.
ss Pp mm X Ss pp Mm
Offspring
Theoretical Actual
Mottled 3.875 7
Dorsolateral stripes 3.875 4
Picket 3.875 3
Middorsal stripe 3.875 4
Dorsolateral stripes plus Picket 3.875 3
Dorsolateral stripes plus Middorsal stripe 3.875 0
Dorsolateral stripes plus Picket plus
Middorsal stripe 3.875 5
Picket plus Middorsal stripe 3.875 5


1960








254 BULLETIN FLORIDA STATE MUSEUM Vol. 5

Moreover, if the parents were ss Pp mm X Ss pp Mm, each of
the three modifiers (Dorsolateral stripes, Picket, and Middorsal stripe),
when considered individually, should be equally distributed among
the offspring. Below the theoretical number is compared with the
actual number of each of the three traits considered independently:
Theoretical Actual P"
Dorsolateral stripes present 15.5 121


Dorsolateral stripes absent

Picket present

Picket absent

Middorsal stripe present

Middorsal stripe absent


Figure 5. Diagrammatic sketch of
Interocular bar pattern.

Binomial estimate of likelihood,
worse fit to : 1 ratio.


0.14
15.5 19J


15.5 161

15.5 15J

15.5 141

15.5 17 J


0.50


0.36


The actual ratios obtained for
Middorsal stripe and Picket pat-
terns are obviously very close to
the expected ratios, and a ratio as
divergent or worse than 12 to 19, as
represented in the Dorsolateral
stripes series, would be expected
in a random sample about 14 per-
cent of the time. The data are thus
consistent with the hypothesis that
there are three pairs of independ-
ent genes.

Interocular bar
A sharply defined, cream-col-
ored, slightly convex bar passes
across the top of the head from
the upper eyelid of one side to the
upper eyelid of the other side.
Both anteriorly and posteriorly it
is margined by buff-brown or black.
I had no data on this character

on sampling error, of getting as bad or








COIN: PATTERN IN ELEUTIIERODACTYLUS


when I first published on inheritance in nubicola (Goin, 1950). When
I discussed the evolution of the gossci group (Goin, 1954) I reported
its occurrence in gossei and junori, and later (Goin, 1957) I recorded
it in nubicola. I am now able to report that Interocular bar in nu-
bicola, like other pattern modifications, is apparently mendelian in
nature.
In a clutch of 56 eggs from an unknown parent, 55 of the eggs
hatched. The hatchlings showed the following patterns: 20 Mottled,
10 Interocular bar, 14 Dorsolateral stripes, and 11 Dorsolateral stripes
plus Interocular bar. Thus it would seem that Interocular bar well
might be the phenotypic effect of a simple dominant, or recessive,
color-pattern gene-21 with the bar, 34 without-and also that it
assorts independently of Dorsolateral stripes. Such a thesis is cer-
tainly consistent with what we know concerning the inheritance of
the other color pattern modifiers. The presence of the character in
two other species in the group is also suggestive. The fact that Inter-
ocular bar has not yet been found with color patterns other than Dor-
solateral stripes is, I suspect, due simply to the rarity of the gene
in the population.

Broad middorsal stripe
A sharply defined, broad,
cream-colored stripe runs from
back of the head to the vent along
the median dorsal line. For most \\
of its length it is about as wide as
the greatest diameter of the eye.
Its margins are sharply defined by
black or very dark brown.
This is the only pattern modi-
fication so far known that might
completely mask another. It is
quite possible that the presence of
Broad middorsal stripe would pre-
vent the phenotypic expression of
Middorsal stripe.
There are at present no data
on the inheritance of this charac-
ter. The possibility that it may be
an allele to the gene for Middorsal Figure 6. Diagrammatic sketch of
stripe should not be overlooked. Broad middorsal stripe pattern.








BULLETIN FLORIDA STATE MUSEUM


I know of this character in the following species:


alticola
gossei


intermedius
nubicola


rhodopis


Pelvic spots
A rather small but conspicuous, black, more or less rounded patch
is present on each side of the back above the groin.


This pattern differs in two
cussed. It varies greatly in its c


Figure 7. Diagrammatic sketch of
Pelvic spots pattern.


major respects from those so far dis-
onspicuousness and it is expressed by
the addition of pigment rather than
by deletion. (All of the other pat-
tern modifications in nubicola are
brought about by diminution or
absence of pigment, resulting in
the presence of pale, cream-col-
ored areas.) In some mature in-
dividuals the pigmented pelvic
spots are fairly conspicuous, in
others they are rather dim. This
variability seems to be due in part
to variation in intensity of the spots
themselves and in part to variation
in the dorsal ground color of the
individual.
Very dark, conspicuous Pelvic
spots are invariably present in E.
andrewsi, and I have seen paler,
less conspicuous Pelvic spots in
pantoni and gossei as well as nubi-
cola. Judging from published de-
scriptions, this character seems to
be fairly widespread in the genus.


Corollary Discussion
The genus Elcatherodactylus is remarkable not only for the extent
of its pattern variability but also for its life history and mode of em-
bryonic development. Mating takes place and the eggs are laid on
land. The young hatch, not as tadpoles, but as miniature replicas
of the adult. It is not simply a case of metamorphosis before hatch-
ing, for the embryo fails to develop many of the typical tadpole char-


Vol. 5







COIN: PATTERN IN ELEUTIIERODACTYLUS


acters (e.g., functional gills, mouth parts, sucking disk) and does de-
velop a flattened, respiratory tail.
A most remarkable instance of parallelism is shown by certain
members of two other unrelated families of frogs (Rhacophoridae and
Microhylidae) on the opposite side of the world. They have inde-
pendently developed similar terrestrial breeding habits and similar
patterns of embryonic development (tadpole structures lacking, res-
piratory tail present) and show, moreover, series of color pattern mod-
ifications essentially similar to those found in Eleutherodactylus.
Rhacophorus microtympanum (Giinther), a member of the family
Rhacophoridae, lives in the mountainous regions of Ceylon. It lays
about 20 eggs on land in situations similar to those occupied by
E. nubicola, and its embryo is of the eleutherodactyloid type. Kirti-
singhe (1957: 11, 66-67) has pointed out the striking parallelism be-
tween this species and E. nubicola of Jamaica. The following pat-
terns, as described for nuhicola, also occur in R. microtympanum:
(1) mottled pattern with dark pigment arranged in the shape of a
W-mark on the back in the suprascapular region,
(2) a pair of pale, dorsolateral stripes, one on each side from the
eye to the groin,
(3) a narrow middorsal stripe,
(4) a pale interocular bar bordered posteriorly and anteriorly by
darker pigment. (See Kirtisinghe, 1957, fig. 50)
In the Papuan region live a number of species belonging to sev-
eral genera (Sphwnophryne, Oreophryne, Asterophrys, etc.) of the
family Microhylidac. Many of these species are highly variable in
color pattern. Indeed, Parker's (1934: 163) description of color pat-
tern variation in Oreophryne variahilis (Boulenger) from Mt. Bon-
thain, southern Celebes, reads like the description of some species of
Eleutherodactylus. The following patterns are described:
(1) a mottled pattern with a distinct, dark W-shaped mark on the
back in the suprascapular region,
(2) a pair of broad, yellow, dorsolateral light stripes from eye to
groin,
(3) a narrow middorsal stripe from snout to groin.
Parker also describes interocular light bars and ocellar dark spots
in the groin in other species of this genus.
These Papuan microhylid genera are all probably rather closely
related and are the only genera of Microhylidae anywhere in the


1960








258 BULLETIN FLORIDA STATE MUSEUM Vol. 5

world known to have not only terrestrial breeding habits but also an
eleutherodactyloid pattern of embryonic development with the young
hatching as tiny frogs.
Thus members of three different families of frogs, from three dis-
tinct geographic regions, have independently developed very similar
reproductive habits and embryos and at the same time show the same
kinds of color patterns and pattern variability. It is perhaps signifi-
cant that all three types are found in mountainous regions, and while
not at present confined to mountains, they may well have evolved as
montane forms. But the evolutionary significance of this striking
parallelism is still not clear.

LITERATURE CITED

Coin, Coleman Jett
1947. Studies on the life history of Elenuherodactylus ricordii planirostris (Cope)
in Florida with special reference to the local distribution of an allelo-
morphic color pattern. Univ. Florida Studies, Biol. Sci. Series, vol. 4,
no. 2, xi + 66 pp., 6 pls., 7 text figs.
1950. Color pattern inheritance in some frogs of the genus Eleutherodactylus.
Bull. Chicago Acad. Sci., vol. 9, no. 1, pp. 1-15, 1 pl.
1954. Remarks on evolution of color pattern in the gossei group of the frog
genus Eleutherodactylus. Ann. Carnegie Mus., art. 10, pp. 185-195,
2 figs.
1958. Further studies on color pattern inheritance in the frog, Eleutherodacty-
lus nubicola. Year Book Amer. Philo. Soc. for 1957, pp. 248-250.
Kirtisinghc, P.
1957. The amphibia of Ceylon. Privately published, Ceylon, xiii + 112 pp.,
74 figs.
Parker, H. W.
1934. A monograph of the frogs of the family Microhylidae. British Mus.
(Nat. list.), London, viii + 208 pp., 67 figs.
Sturtevant, Alfred Henry
1948. The evolution and function of genes. Amer. Sci., vol. 36, no. 2, pp.
225-236.








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