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228 HERPETOLOGICA Vol. 21, No. 3
LITERATURE CITED
REPRODUCTIONIN A NEOTIROPICALSALAMANDER,
BOLITOGLOSSAADSPERSA (PETERS)
AND J. R. TAMSITT
DARIOVALDIVIESO
INTRODUCTION
Salamanders of the genus Bolitoglossa which inhabit the high
Andean regions of Colombia have been considered to be ovovivip-
arous by some herpetologists (Peters, 1863; Posada, 1909; Noble,
1931). In reference to Bolitoglossa adspersa (Peters), the common
salamander of the eastern Colombian Andes, Ruthven (1922)
simply stated that, "this species is a viviparous species," basing his
conclusion only on the presence of eggs in the ovary and, as such,
having no direct evidence. Dunn (1944) referred to Neotropical
plethodontid salamanders and stated that, "some are oviparous,
whereas others, especially in the mountainous regions of Colombia,
are thought to be viviparous. However, it is not definitely known
if this is the case, and any information concerning the mode of
reproduction of these salamanders would be interesting." This
question remained unanswered until Niceforo Marla (19,60) pre-
sented evidence to show that B. aclspiersa was oviparous and not
viviparous as believed by earlier workers. He removed six eggs of
this species from beneath a rock at Dintel (2,700 m.) in the Eastern
1965 REPRODUCTION IN A SALAMANDER 229
Andes and later obtained newly hatched salamanders from the eggs.
During 1961-1962, the authors obtained additional data on the
life history and reproductive organs of this oviparous salamander
which supplement the original note of Niceforo Marla. Collections
were made in January, July, August, September, October, and
November, 1961, and in March, April, and May, 1962, near Bogota
(40 35' N, 740 04' W, 2,599 m.), Dept. of Cundinamarca, Colombia.
The salamanders, which are active throughout the year, were taken
from beneath rocks or rusted sheets of tin in wooded areas on
mountain slopes above this city.
The Bogot'a region is found in the lower montane dry forest
formation according to the classification of Holdridge (1947).
Rain, however, falls in every month, and at Bogot'a in 1961 most rain
fell in April, June, July, October, and November, the yearly average
being 925 mm. Temperatures may reach extremes, and the 1961
minimum was -1.1? C. and the maximum 23.0? C.; the annual
temperature mean was 13.40 C. The annual mean relative humidity
for 1961 at Bogot'a was 79 per cent (Empresa de Acueducto y
Alcantarillado de Bogot'a, 1961), and the total annual variation in
day length at latitude 40 N in Colombia is approximately 24 minutes
(Miller, 1959).
IDENTIFICATION OF SEX
Sexual dimorphism in body size occurs in this species, and the
adult female is considerably larger (snout-vent length 45-66 mm.)
than the adult male (snout-vent length no greater than 50-55 mm.).
230 HERPETOLOGICA Vol. 21, No. 3
Adult females with heavily yolked eggs and ready to breed have
been found in every month of the year except February, June, and
December, for which no data are available. Egg clutches have been
found in March, September, and October, and the clutch described
by Niceforo Marla (op. cit.) was encountered in November. Al-
though more observations would be desirable, even from these data
it can be seen that reproduction in B. adsp.ersa shows no seasonal
pattern and is apparently acyclic. There is no single egg-laying
season, and fecund females can be found in most months of the
year. Climatic conditions in the collecting area are relatively uni-
form, and a constant food supply is available to sustain continuous
reproduction. Under such conditions Neotropical salamander re-
productive patterns should be expected to be altered from the
seasonal cycles of North American temperate zone plethodontids
(see Organ, 1961; Highton, 1962; Martof, 1962).
Males.-Sixty-nine males of all ages, collected during 6 months
in 1961 and during 3 months in 1962, were examined to obtain the
following data. The testes of the smallest individuals possessed only
one lobe, the largest single-lobed testis measuring 8 mm. in in-
dividuals less than 40 mm. in snout-vent length. As the salamander
grows, the testes become elongate and clearly divided into two or
three lobes of unequal length. In some adult males, however, only
a single large lobe persisted, the largest single-lobed testis in the
sample measuring 13 mm. in an individual 42 mm. in snout-vent
length. In their sample Stebbins and Hendrickson (1959) found
that the smallest breeding male was 41.0 mm. in snout-vent length,
and we have found that males smaller than 39 mm. in snout-vent
length have small mental glands, slender sperm ducts and small,
single-lobed testes less than 8 mm. in length. In these males
spermatogenesis is only beginning, and they are not capable of
reproducing. Males less than 20 mm. in snout-vent length are
difficult to distinguish from females. In these the testes are
membranous and spermatogenesis has not begun. In all males
greater than 39 mm. in snout-vent length, the mental gland was
clearly seen, the central longitudinal duct was filled with sperm, and
the testes were single or bilobed. Only in one sexually mature male
(snout-vent length 47 mm.) was a small, posterior third lobe of the
testes present.
Histological examination of the testes of sexually mature males
indicates that structure and the spermatic cycle are very much as
those described by Humphrey (1922) for other urodeles. The
lobules of the elongated testes empty directly by very short ducts
into a central longitudinal duct. This duct receives mature sperm,
which are then transported through efferent tubules to the Wolffian
duct. There is no second longitudinal duct near the mesonephros as
has been found in other amphibia (Noble, 1931).
1965 REPRODUCTION IN A SALAMANDER 233
3-
JflI] 3 LOBES
5
:D
0 3
2 LOBES
Z 7-
5- LOBE
o1
20 25 30 35 40 45 50 55
SUMMARY
Field and laboratory studies of the plethodontid salamander
Bolitoglossa adspersa (Peters) at Bogota, Colombia, during 1961-
1962 present evidence that this species is oviparous and that re-
production is apparently acyclic. Fecund females and mature males
were found in 9 nonconsecutive months of the year. There is no
single egg-laying season, and females oviposited from 10-11 eggs
in March, September, October, and November, months for which
data were available.
All females larger than 45 mm. in snout-vent length were sexually
mature, irrespective of the month of capture, and there was no
correlation between the size of the female and the number of yolked
eggs, which averaged 10 in number in 44 adults of snout-vent length
45-66 mm.
Males larger than 39 mm. in snout-vent length were sexually
mature, and the number of lobes per testis varied from one to three.
A sample of 58 males collected in April and May, 1962, consisted of
six age groups as determined by the size of the male and the
number of lobes per testis, the oldest males being at least in their
fourth breeding season.
The acyclic reproductive pattern of B. adspersa is discussed in
relation to the cyclic behavior of temperate zone species.
LITERATURE CITED
BRAME, A. H., JR., AND D. B. WAKE. 1963. The salamanders of South
America. Los Angeles Co. Mus. Contr. in Sci., 69:5-72.
DUNN, E. R. 1944. Herpetology of the Bogota area. Rev. de la Acad.
Colomb. de Cien. Exact., Fisic. y Nat., 6:68-81.
EMPRESA DE ACUEDUCTO Y ALCANTARILLADO DE BOGOTA. 1961. Boletin
informativo, 1961:1-114.
HIGHTON, R. 1962. Geographic variation in the life history of the slimy
salamander. Copeia, 1962:597-613.
HOLDRIDGE, L. R. 1947. Determination of world plant formations from simple
climatic data. Science, 105:367-368.
HUMPHREY, R. R. 1922. The multiple testis in urodels. Biol. Bull., 43:45-67.
MARTOF, B. S. 1962. Some aspects of the life history and ecology of the
salamander Leurognathus. Amer. Mid. Nat., 67:1-35.
MILLER, A. H. 1959. Reproductive cycles in an equatorial sparrow. Proc.
Nat. Acad. Sci., 45:1095-1100.
NIc'FORO MARIA, HNO. 1960. Salamandras de la Familia Plethodontidae:
Hallazgo de los huevos de Bolitoglossa adspersa (Peters). Caldasia,
8:337-339.
236 HERPETOLOGICA Vol. 21, No. 3