Life-history Aspects of Stereochilus marginatus,
with a Comparison of Larval Development in
Syntopic S. marginatus and Pseudotriton montanus (Amphibia: Plethodontidae)
Richard C. Bruce
Southeastern Naturalist, Volume 7, Number 4 (2008): 705–716
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2008 SOUTHEASTERN NATURALIST 7(4):705–716
Life-history Aspects of Stereochilus marginatus,
with a Comparison of Larval Development in
Syntopic S. marginatus and Pseudotriton montanus
Richard C. Bruce*
Abstract - The plethodontid salamanders Stereochilus marginatus (Many-lined
Salamander) and Pseudotriton montanus (Mud Salamander) have overlapping distributions
in the southeastern Atlantic Coastal Plain, where they often co-occur in
low, swampy habitats. The main objective of this study was to document life-history
traits of S. marginatus at Cool Springs in eastern North Carolina, and compare these
findings with life-history parameters of a population surveyed in the late 1960s at a
nearby locality (Croatan Forest). A second objective was to compare larval development
of S. marginatus and P. montanus at Cool Springs. I found that S. marginatus
has a larval period of 13–14 months, hatching in early spring and undergoing metamorphosis
late in the second spring. Males may breed initially as early as the autumn
following metamorphosis, at 19–21 months of age; females probably require an additional
year to attain maturity, ovipositing initially at 3 years. Clutch sizes, based
on counts of yolked ovarian follicles of dissected females, ranged from 42 to 60.
Compared to S. marginatus, the larval period of P. montanus is slightly longer (14–17
months), extending from hatching in winter to metamorphosis in the second spring.
Although larval body sizes of S. marginatus and P. montanus overlap considerably,
larvae of the latter species tend to grow larger and metamorphose at slightly larger
sizes. The phenologies of the life cycles of both species corroborate earlier studies,
both across years and across southeastern localities. However, growth and developmental
rates of S. marginatus at Cool Springs appear to be accelerated relative to
those reported previously for the Croatan Forest population.
Stereochilus marginatus Hallowell (Many-lined Salamander) is a small,
non-descript, lungless salamander endemic to the Atlantic Coastal Plain
of the United States. It is highly aquatic, living in and along the margins of
swamps, ponds, and sluggish streams. Means (2000) included S. marginatus
in an assemblage (assemblage 1) of Coastal Plain plethodontids that inhabit
low, swampy habitats, in contrast to a second assemblage (assemblage 2) of
Coastal Plain species that are associated with ravines. Various aspects of the
life history, feeding ecology, and reproductive biology of S. marginatus have
been documented by Bruce (1971), Foard and Auth (1990), Rabb (1956),
Schwartz and Etheridge (1954), and Wood and Rageot (1963).
In this paper, I report on the life history of S. marginatus in a population
inhabiting a swamp ecosystem in the Coastal Plain of eastern North
*Department of Biology, Western Carolina University, Cullowhee, NC 28723. Current
address - 200 Breeze Way, Aurora, NC 27806; firstname.lastname@example.org.
706 Southeastern Naturalist Vol. 7, No. 4
Carolina. Stereochilus marginatus was the most frequently encountered species
of salamander in the swamp. One objective was to compare life-history
traits in this population with those of a nearby population that had been
studied some years before (Bruce 1971). A second objective was to compare
the larval period of S. marginatus with that of Pseudotriton montanus Baird
(Mud Salamander), also an assemblage 1 species and the next most frequently
encountered species in the swamp. The two species are closely related,
and larvae are morphologically similar (Birchfield and Bruce 2000). The life
history of P. montanus is known mainly from studies conducted in western
North and South Carolina (Bruce 1974, 1975, 1978). Valuable summaries of
the natural history of both S. marginatus and P. montanus can be found in
Petranka (1998), and in the accounts by Ryan (2005) on S. marginatus and
Hunsinger (2005) on P. montanus in Lannoo (2005).
Study Area, Materials, and Methods
I conducted the study at the Cool Springs Environmental Education Center
administered by the Weyerhaeuser Corporation. The Cool Springs tract
encompasses about 680 ha adjacent to the confl uence of the Neuse River and
Swift Creek in Craven County, NC (35o11.4'N, 77o04.9'W). Elevations are
≤6 m above sea level. Natural habitats include pine-oak sand ridges, mixed
pine-hardwood forests, pocosins, and a cypress swamp (Hall 2006).
The sampling area in the present study was a small section of the cypress
swamp in which Chamaecyparis thyoides L. (Atlantic White Cedar),
rather than Taxodium distichum L. (Bald Cypress), was a co-dominant canopy
species, together with Acer rubrum L. (Red Maple), Nyssa aquatica
L. (Water Tupelo), Quercus michauxii Nuttall (Swamp Chestnut Oak), and
Fraxinus caroliniana Miller (Carolina Ash). This area was fed by a shallow,
sluggish stream that discharged into the swamp. The latter consisted
of a network of isolated and semi-isolated shallow, mucky pools along the
principal, but indistinct, drainage channel. Many of the pools supported
mats of Sphagnum spp.
I collected salamanders by pulling a dip net along the mucky bottoms of
the aquatic habitats. I also raked leaf litter and turned rotting logs along the
banks of the stream and pools and in the intervening fl oodplain. A total of
195 S. marginatus (109 larvae, 86 adults) and 75 P. montanus (65 larvae, 10
adults) were captured. Counts of other species of salamanders observed were
recorded. For convenience, throughout this paper, I use the term “adult” to
refer to fully-metamorphosed individuals, whether immature (juveniles) or
Captured S. marginatus and P. montanus were returned to the Cool
Springs laboratory, where they were anesthetized in a solution of MS-222
and then measured for standard length (SL), to the nearest 0.1 mm, from
the tip of the snout to the posterior end of the cloacal slit. All measurements
given herein refer to SL. Most of the salamanders were subsequently revived
in swamp water and returned to the site of capture, usually on the same day.
2008 R.C. Bruce 707
Given that the salamanders were not marked, it was unknown how many
were subsequently recaptured. However, I assumed that some captures were
recaptures, which required a cautious approach to statistical evaluation in
order to avoid violations of independence (Scheiner and Gurevitch 2001). In
statistical tests, significance was evaluated at α = 0.05.
The main set of samples was taken from 21 November 2003 through
29 November 2004. During this period, I attempted to sample twice each
month, but in August and September 2004, only single searches were
made. Four supplementary samples were taken between 30 March and 25
May 2005, mainly to obtain additional data on early larval development
of both species.
For the last three collections of 2004 and for those of the spring of 2005,
I preserved all adults of S. marginatus (n = 23), in order to dissect the reproductive
organs and determine reproductive condition. These specimens were
euthanized in MS-222, fixed in 10% formalin, and transferred to 70% ethyl
alcohol. Preserved specimens have been deposited in the herpetological collection
of the North Carolina State Museum of Natural Sciences (catalog
Although I did not attempt to quantify habitat features of the two species,
larvae of P. montanus were more abundant in the upper section of the swamp,
especially in the small feeder stream and adjacent pools. Most of the firstyear
larvae of P. montanus taken in the spring of both 2004 and 2005 were
found in a seep adjacent to the stream, at the base of a gentle slope. I assumed
that the nesting sites had been in subsurface channels leading into the seep.
In contrast, larvae of S. marginatus were more abundant downstream, in
lentic habitats, especially shallow pools that were often bordered by Sphagnum
mats. However, few larvae were found in Sphagnum; most were taken
from decaying vegetation on the bottom of the pools. Notwithstanding these
differences, larvae of S. marginatus and P. montanus broadly overlapped
throughout the area of the swamp that I searched, and were sometimes taken
in the same pool.
Most adult S. marginatus were captured in dip-net samples in the same
aquatic habitats as larvae. During a dry period in June and July 2004, when
pools had little or no water, 19 adults were found by excavating by hand the
saturated muck at the bottom of drying pools. In contrast, other than two
small, newly transformed juveniles taken in dip-net samples, the adult P.
montanus were found on land, under logs or boards on damp soil. I did not
make a concerted effort to excavate burrows, a favored habitat for this species
elsewhere (Bruce 1975).
Other species of salamanders encountered included 43 Desmognathus
auriculatus Holbrook (Southern Dusky Salamander), 6 Eurycea
chamberlaini Harrison and Guttman (Chamberlain’s Dwarf Salamander),
708 Southeastern Naturalist Vol. 7, No. 4
3 E. cirrigera Green (Southern Two-lined Salamander), 13 Plethodon chlorobryonis
Mittleman (Atlantic Coast Slimy Salamander), and 2 juvenile Amphiuma
means Garden (Two-toed Amphiuma). Although D. auriculatus may
be the second-most abundant species in the swamp, there were only 6 larvae
among the 43 individuals recorded. No larvae of either species of Eurycea
were seen, suggesting that neither breeds at the immediate study site. A full
account of the amphibians and reptiles of the Cool Springs tract is provided
by Hall (2006).
Larval development of Stereochilus marginatus
The size distribution of larval S. marginatus for the 2003–2004 samples
is shown in Figure 1, and that of small larvae taken in the spring of 2005
in Figure 2. The smallest larva (10.5 mm), captured on 15 April 2005, had
a belly distended with yolk and thus had likely hatched within the past
month. The course of early development in the spring of the successive
years was similar; the mean SL of the eight small larvae of 21 May 2004
(mean ± SD = 15.6 ± 1.10 mm) and that of the six first-year larvae of 25
May 2005 (mean ± SD = 16.4 ± 1.16 mm) did not differ significantly (t =
1.35, df = 12, P = 0.203).
Among older larvae, there was a trend toward size increase from November
to May. The largest larvae, 44–46 mm, were found between 30
January and 29 April. Larger larvae disappeared from the population in May,
presumably due to metamorphosis of all 2nd-year larvae. This conclusion is
supported by the observation that the smallest adults were captured between
Figure 1. Distribution of standard lengths of 100 larval Stereochilus marginatus
(Many-lined Salamander) captured between 21 November 2003 (day 1) and 29 November
2004 (day 375). Some symbols in this and other graphs represent more than
one individual of equivalent sizes.
2008 R.C. Bruce 709
31 May and 9 July (Fig. 3); five of these, ranging from 38 to 45 mm, showed
obvious gill resorption scars, but were otherwise fully metamorphosed.
Figure 2. Distribution of standard lengths of first-year larvae of Pseudotriton montanus
(Mud Salamander) (triangles, n = 19) and Stereochilus marginatus (Many-lined
Salamander) (inverted triangles, n = 8) captured between 30 March and 25 May 2005.
The x-axis is indexed in days from 20 November, as in the other graphs.
Figure 3. Distribution of standard lengths of 80 adult Stereochilus marginatus
(Many-lined Salamander) captured between 21 November 2003 (day 1) and 29 November
2004 (day 375).
710 Southeastern Naturalist Vol. 7, No. 4
Thus, if embryos hatched in early spring, and if most larvae metamorphosed
in mid-late spring of their second year, then the larval period was approximately
13–14 months at Cool Springs during the course of this study.
Few larvae were found in summer and autumn of 2004; nevertheless,
there is a pattern of rapid growth of first-year larvae in the interval from
spring to late autumn. Larvae in the final sample, that of 29 November 2004,
had attained the same sizes as those taken in the initial sample of 21 November
2003 (2003: n = 12, mean ± SD = 35.4 ± 2.85 mm; 2004: n = 4, mean ±
SD = 35.2 ± 3.38 mm; t = 0.121, df = 14, P = 0.905).
The distribution of body sizes of adult S. marginatus captured in the
2003–2004 samples is shown in Figure 3. The additional six individuals
captured in the spring samples of 2005 fell within the same size range. The
exceptionally large individual (67 mm) was a male; otherwise, among the
23 individuals that were dissected, maximum sizes of the sexes were similar
(male = 56 mm, female = 57 mm). Following Bruce (1971), I have calculated
ages from February, the probable median month of entry of individuals into
the population as eggs.
Females. Of the 17 individuals collected 29 October–29 November 2004
and subsequently dissected, 10 were females. The two smallest (45 and 46
mm) were immature, having narrow, straight oviducts and small, compact
ovaries containing small, translucent follicles. In the remaining eight (52–56
mm), vitellogenesis was in progress with yolked follicles ranging from 1.2 to
2.2 mm in diameter. Follicle counts varied from 42 to 60 (mean ± SD = 51.4 ±
5.85). Although the smallest female had the fewest yolked follicles, the slope
of the regression of follicle number on SL was non-significant (b = 1.05, t =
1.33, df = 6, P > 0.20; Fig. 4). It is likely that these gravid females would have
oviposited in the approaching egg-laying season in mid- to late winter.
Figure 4. Counts of yolked
ovarian follicles in 8 gravid
females of Stereochilus
Salamander) collected in
autumn 2004. The slope of
the regression line is nonsignificant (see text).
2008 R.C. Bruce 711
If the two small, immature females, which overlapped in size with the
three largest larvae, had metamorphosed in the previous spring, it seems
likely that they had been scheduled to yolk their initial clutch the next autumn
(i.e., 2005) and oviposit initially at an age of 3 years. Based on the
limited data available, this is the best estimate of age at first reproduction of
females in this population.
The four females dissected from the April–May 2005 samples were
51–57 mm SL. All four appeared mature, and three were in an obvious spent
condition, showing convoluted oviducts and loosely organized ovaries having
a scattering of small, translucent follicles. The ovaries of one individual
had three orange-pigmented, atretic follicles.
Although adults captured in the earlier samples of 2003 and 2004 were
not dissected, I did identify gravid females by the presence of yolky ovarian
follicles visible through the body wall of the distended abdomen. Eight
such females were found between 21 November and 29 February, but none
thereafter until the next autumn. These females were similar in size (n = 8,
mean ± SD = 51.9 ± 1.76 mm) to the gravid females dissected in the late
2004 samples (n = 8, mean ± SD = 53.2 ± 2.66 mm), and the difference was
non-significant (t = 1.22, df = 14, P = 0.243).
Males. Seven males (46–67 mm) were collected and dissected in the period
29 October–29 November 2004. Although the testes varied in size and
shape, all seven had dark, coiled vasa deferentia, and were thereby scored
as mature. In five of these males, including the smallest (46 mm), a cord
of spermatozoa was detected in the vasa. Thus, it is likely that many males
breed initially in the autumn following metamorphosis, at approximate ages
of 19–21 months. Only two males were dissected from the spring 2005
samples, both collected on 25 May. The smaller, 46 mm, possibly a recent
metamorph, was immature; the larger, 50 mm, was mature based on the presence
of black, coiled vasa, but was not in breeding condition, inasmuch as
the testes were thin and the vasa were empty.
Larval development of Pseudotriton montanus
During the principal sampling period in 2003–2004, 42 larvae and eight
adults of P. montanus were captured. Most larvae were taken during the
earlier phase of the sampling interval (Fig. 5). The smallest were collected
on 19 April and, from that date to mid-June, two distinct size classes of
larvae were present, presumably representing the cohorts of 2003 and 2004.
In the former, a slight trend of increasing size was evident in the samples
taken from November to mid-June. The loss of this cohort in late June was,
in part at least, an effect of metamorphosis in the spring months. A 46-mm
larva collected on 24 March metamorphosed in the laboratory on 19 April.
Two fully metamorphosed individuals (44 and 49 mm), which retained larval
pigmentation (brown), were found on 31 May and 15 June, respectively; on
the latter date, a metamorphosing larva (41 mm) was taken as well.
The supplementary samples of spring 2005 included two adults, four
large larvae (33–46 mm), and 19 1st-year larvae; the size distribution of the
712 Southeastern Naturalist Vol. 7, No. 4
latter (Fig. 2) was similar to that of 1st-year larvae taken the previous year
(Fig. 5). The mean sizes in two pairs of samples of 1st-year larvae captured
on nearly the same dates of the two years were not significantly different at
α = 0.05/2 (Bonferroni adjustment) (19 April 2004 vs. 15 April 2005: t =
0.116, df = 10, P = 0.910; 29 April 2004 vs. 30 April 2005: t = 1.394, df = 8,
P = 0.201).
Given that the smallest larvae of P. montanus, already ≥15 mm (n = 6,
mean ± SD = 15.8 ± 0.61 mm), were found on 30 March 2005, it is likely
they had hatched 1–2 months earlier in both 2004 and 2005, in mid-winter,
as at other localities (Bruce 1974). Most larvae at Cool Springs probably
metamorphose in the spring of their second year, after a larval period of
In an earlier study of the life history of S. marginatus in Croatan National
Forest, NC (a locality 32 km south of Cool Springs), I estimated
that the larval period was usually 25–28 months, but that some larvae
transformed at 13–16 months (Bruce 1971). Although based on smaller
samples, the Cool Springs data nonetheless suggest strongly that most
larvae metamorphose in their second year, 13–14 months after hatching,
at an approximate age of 15 months. In the Croatan study, I estimated that
Figure 5. Distribution of standard lengths of 42 larvae and 2 recent metamorphs (31
May: 43.9 mm; 15 June: 48.9 mm) of Pseudotriton montanus (Mud Salamander) captured
between 21 November 2003 (day 1) and 14 October 2004 (day 329).
2008 R.C. Bruce 713
oviposition occurred in winter, with hatching following in late March–
early April; this is supported by the newer data from Cool Springs. Thus,
the presence of gravid females in the population in autumn and winter—as
late as late February—indicates that the oviposition season may extend to
late winter. The two tiny larvae (10, 11 mm) taken in mid- and late April at
Cool Springs were similar in size to hatchlings found at Croatan in April
and May (taking into account the difference in methodology of measuring
body length). At Croatan, I found that metamorphosis occurred in late
spring and summer, similar to the findings at Cool Springs. Thus, whereas
larval developmental rates appear to differ between the two populations
(studied 35 years apart), the phenologies of the larval phases were very
similar. Perhaps climatological changes during the interval and/or habitat
differences between the sites contributed to the observed differences. I was
unable to conduct parallel studies of the Cool Springs and Croatan populations
in 2003–2005 because of changes in the habitat and severe reduction
of the S. marginatus population at the latter site.
The course of the larval period of P. montanus at Cool Springs was similar
to that of S. marginatus; however, hatching occurred earlier in the year
in the former species, probably indicating an earlier oviposition season, in
late autumn–early winter, as at other localities (Brimley 1944, Fowler 1946,
Goin 1947). Inasmuch as the smallest larvae in 2004 and 2005 were taken
in late March 2005, and had already grown to 15 mm and larger, I presumed
that they had hatched at least 1–2 months earlier and had undergone early
development in the subsurface nesting sites. Given that metamorphosis occurred
in the spring and presumably included all (or nearly all) 2nd-year
larvae, the estimated duration of the larval period is 14–17 months. This
estimate is similar to estimates for populations in the Piedmont of western
South Carolina (Bruce 1974, 1978). In a higher-elevation population in the
Blue Ridge of North Carolina, the usual larval period was found to be a year
longer (Bruce 1978).
At Cool Springs, the earlier oviposition season of P. montanus versus
S. marginatus gave the former an advantage in growth during the first year.
Although larger larvae of the two species overlapped broadly in size, those
of P. montanus tended to grow larger and metamorphose at slightly larger
sizes (≈41–50 mm) than those of S. marginatus (≈38–45 mm). Moreover, as
larvae of S. marginatus have a more slender habitus than those of P. montanus
(Birchfield and Bruce 2000), metamorphs of the latter probably have
even greater masses than those of the former species.
Data on the reproductive cycles of male and female S. marginatus at
Cool Springs are incomplete, given the necessity of dissection to determine
reproductive status of both sexes. However, the data obtained on specimens
taken in late autumn 2004 and spring 2005 agree well with the more comprehensive
picture of reproduction at Croatan Forest in 1968–69 (Bruce
1971). In the latter population, males were in breeding condition, with vasa
714 Southeastern Naturalist Vol. 7, No. 4
deferentia packed with sperm, by 1 November; similarly, for the October–
November 2004 samples from Cool Springs, all seven males were in breeding
condition, including the smallest (46 mm). The two males dissected from
the spring samples of 2005 were likewise similar to their spring counterparts
at Croatan Forest. The estimated age at first reproduction of 19–21 months
at Cool Springs represents the lower limit of the earlier estimate (21–45
months) for the Croatan Forest population. I contend that most males in both
populations were initiating spermatogenesis immediately following metamorphosis;
as such, the earlier age of reproduction at Cool Springs likely
refl ects the earlier age at metamorphosis.
The dissection data suggest that an additional year may be required for
development to sexual maturity in females versus males. Thus, age at first
reproduction (i.e., oviposition) is estimated as 3 years in female S. marginatus
at Cool Springs, a year earlier than in the Croatan Forest population
(Bruce 1971). As in males, the difference reflects the difference in age at
metamorphosis. At Croatan Forest, counts of follicle numbers in the ovaries
of gravid females were obtained for only three individuals; the values
were lower (n = 22–29) than for the eight gravid females at Cool Springs
(n = 42–60). No deposited eggs clutches were found at Croatan earlier or at
Cool Springs recently, but have been found by other investigators. Whereas
Schwartz and Etheridge (1954) and Rabb (1956) provided data on single
deposited egg clutches, the most comprehensive report on nesting of S.
marginatus was provided by Wood and Rageot (1963), who discovered 43
clutches between late winter and early spring of 1954 and 1955. Number of
eggs per clutch ranged widely (6–92), and attending females were largely
absent. Thus, it is possible that the smaller clutches had been reduced by
predation and that the larger were products of more than one parent. However,
Noble and Richards (1932) reported a mean clutch size of 57 in 19
females of S. marginatus following implantation of frog pituitaries; subsequent
dissection of the females showed that the total ovarian complements
of “mature” eggs had averaged 70, with a range of 16–121. From this series
of studies, including the present report, it appears that clutch size in S.
marginatus is highly variable; whether there is a tendency of clutch size
to increase with body size, as in its closest relatives in the genera Gyrinophilus
and Pseudotriton (Bruce 1969), is uncertain. High residual variance
obscures such a correlation in small samples in species where the range of
female body size is narrow, as in S. marginatus.
It would be of interest to examine interactions between S. marginatus
and P. montanus in habitats like that of the Cool Springs swamp, where both
species are relatively abundant. A working hypothesis is that interactions,
especially interspecific competition, are stronger between larvae of the two
species, which are morphologically similar (Birchfield and Bruce 2000) and
overlap in habitat use, than between adults, which show greater divergence
in size, morphology, and habitat associations. However, it is possible that
2008 R.C. Bruce 715
adult P. montanus prey on S. marginatus, given Brimley’s (1944) observations
of predation by the former species on Eurycea cirrigera. The swamp
habitat is a difficult one for conducting natural manipulative experiments,
but laboratory and mesocosm experiments could prove fruitful.
I thank Jeff Hall, former manager of the Cool Springs Environmental
Education Center, for providing access to the site; he also assisted in the field
work. Scientific collecting licenses were issued by the North Carolina Wildlife
Birchfield, G.L., and R.C. Bruce. 2000. Morphometric variation among larvae of
four species of lungless salamanders (Caudata: Plethodontidae). Herpetologica
Brimley, C.S. 1944. Amphibians and Reptiles of North Carolina. Carolina Biological
Supply Co., Elon College, NC. 63 pp.
Bruce, R.C. 1969. Fecundity in primitive plethodontid salamanders. Evolution
Bruce, R.C. 1971. Life cycle and population structure of the salamander Stereochilus
marginatus in North Carolina. Copeia 1971:234–246.
Bruce, R.C. 1974. Larval development of the salamanders Pseudotriton montanus
and P. ruber. American Midland Naturalist 92:173–190.
Bruce, R.C. 1975. Reproductive biology of the Mud Salamander, Pseudotriton montanus,
in western South Carolina. Copeia 1975:129–137.
Bruce, R.C. 1978. A comparison of the larval periods of Blue Ridge and Piedmont
Mud Salamanders (Pseudotriton montanus). Herpetologica 34:325–332.
Foard, T., and D.L. Auth. 1990. Food habits and gut parasites of the salamander
Stereochilus marginatus. Journal of Herpetology 24:428–431.
Fowler, J.A. 1946. The eggs of Pseudotriton montanus montanus. Copeia 1946:105.
Goin, C.J. 1947. Notes on the eggs and early larvae of three Florida salamanders.
Chicago Academy of Sciences, Natural History Miscellanea 10:1–4.
Hall, J.G. 2006. Herpetological sampling in the North Carolina Coastal Plain: A
comparison between techniques across habitats. M.Sc. Thesis. East Carolina
University, Greenville, NC. 173 pp.
Hunsinger, T.W. 2005. Pseudotriton montanus Baird, 1849. Mud Salamander. Pp.
858–860, In M. Lannoo (Ed.). Amphibian Declines: The Conservation Status of
United States Species. University of California Press, Berkeley, CA.
Lannoo, M. (Ed.). 2005. Amphibian Declines: The Conservation Status of United
States Species. University of California Press, Berkeley, CA. 1094 pp.
Means, D.B. 2000. Southeastern US Coastal Plain habitats of the Plethodontidae:
The importance of relief, ravines, and seepage. Pp. 287–302, In R.C. Bruce,
R.G. Jaeger, and L.D. Houck (Eds.). The Biology of Plethodontid Salamanders.
Kluwer Academic/Plenum Publishers, New York, NY.
Noble, G.K., and L.B. Richards. 1932. Experiments on the egg-laying of salamanders.
American Museum Novitates 513:1–25.
716 Southeastern Naturalist Vol. 7, No. 4
Petranka, J.W. 1998. Salamanders of the United States and Canada. Smithsonian
Institution Press, Washington, DC. 587 pp.
Rabb, G.B. 1956. Some observations on the salamander, Stereochilus marginatum.
Ryan, T.J. 2005. Stereochilus marginatus (Hallowell, 1856). Many-lined Salamander.
Pp. 862–863, In M. Lannoo (Ed.). Amphibian Declines: The Conservation
Status of United States Species. University of California Press, Berkeley, CA.
Scheiner, S.M., and J. Gurevitch (Eds.). 2001. Design and Analysis of Ecological
Experiments. Oxford University Press, Oxford, UK. 415 pp.
Schwartz, A., and R. Etheridge. 1954. New and additional herpetological records
from the North Carolina Coastal Plain. Herpetologica 10:167–171.
Wood, J.T., and R.H. Rageot. 1963. The nesting of the Many-lined Salamander in the
Dismal Swamp. Virginia Journal of Science 14:121–125.