Logging Road Effects on Breeding-site Selection in
Notophthalmus viridescens (Red-spotted Newt) and Three
Ambystomatid Salamanders in South-central Pennsylvania
David L. Chambers
Northeastern Naturalist, Volume 15, Issue 1 (2008): 123–130
Full-text pdf (Accessible only to subscribers.To subscribe click here.)
Access Journal Content
Open access browsing of table of contents and abstract pages. Full text pdfs available for download for subscribers.
Current Issue: Vol. 30 (3)
Check out NENA's latest Monograph:
Monograph 22
2008 NORTHEASTERN NATURALIST 15(1):123–130
Logging Road Effects on Breeding-site Selection in
Notophthalmus viridescens (Red-spotted Newt) and Three
Ambystomatid Salamanders in South-central Pennsylvania
David L. Chambers*
Abstract - This study examined possible effects of several abiotic parameters on
breeding-pool selection of Ambystoma jeffersonianum (Jefferson Salamander),
A. maculatum (Spotted Salamander), A. opacum (Marbled Salamander), and Notophthalmus
viridescens (Red-spotted Newt). Twenty-four ephemeral pools and
permanent ponds, all adjacent to a logging road, were observed in south-central
Pennsylvania in 2006. There was a significant correlative effect of distance from
the logging road on breeding-site selection. Specifically, the distance from the road
significantly differed between the breeding-sites of Jefferson Salamanders and Redspotted
Newts and between those of Marbled Salamander and Red-spotted Newts
with both ambystomatid species breeding farther from the road than Red-spotted
Newts. This study supports the idea that ambystomatid salamander breeding-site
selection can be influenced by habitat disturbance, while generalist species are not
as easily influenced.
Introduction
Habitat modifications, primarily as the result of anthropogenic disturbances,
are arguably one of the most obvious threats to pool-breeding
amphibians (Semlitsch 2003), and contribute to at least some amphibian
population declines (Wyman 1990). While several studies have addressed the
impact of habitat change on anurans (Bradford et al. 2005, Peltzer et al. 2003),
others have demonstrated that salamanders may be even more sensitive to such
change than anurans (Petranka et al. 1993). These habitat alterations may influence
the distribution of breeding sites (Rubbo and Kiesecker 2005), and ultimately,
cause the reduction or extirpation of populations (Gibbons et al. 2006).
The primary objective of this study was to ascertain the effects of a logging
road on the selection of breeding sites among four salamander species:
Ambystoma jeffersonianum Green (Jefferson Salamander), A. maculatum
Shaw (Spotted Salamander), A. opacum Gravenhorst (Marbled Salamander),
and Notophthalmus viridescens Rafinesque (Red-spotted Newt) in southcentral
Pennsylvania. In addition, the correlation of several other abiotic
factors recorded from each pool to breeding-site selection was tested. Most
amphibian pool-breeders, including Jefferson Salamander, Spotted Salamander,
and Red-spotted Newt, typically breed during the spring and early
summer months (Petranka 1998). The Marbled Salamander, however, is one
of only two ambystomatid salamanders to deposit eggs in dried ephemeral
pools during the early fall, with larvae hatching shortly after pool filling in
the winter, and completing metamorphosis in the spring (Petranka 1998).
*Department of Biological Sciences, Virginia Polytechnic and State University, 2119
Derring Hall, Blacksburg, VA 24061; dlchamb@vt.edu.
124 Northeastern Naturalist Vol. 15, No. 1
Materials and Methods
Study site
The observational data presented was collected in the Mount Cydonia
Ponds Natural Area within Michaux State Forest, Franklin County, PA (Fig.1).
Michaux State Forest covers approximately 350 km2 in Adams, Cumberland,
and Franklin counties, PA. Mount Cydonia Ponds Natural Area is approximately
0.6 km2 in area. All ephemeral pools and permanent ponds observed were
in the same contiguous mixed-deciduous forest, and all observations at these
pools indicated an absence of fish. Pools occurred at similar elevations and were
located in close proximity to each other. For example, some pools were within
10 m of another, but all were within dispersal ranges of focal caudate species
(Petranka 1998). However, each was a different distance from a logging road
that follows the southwest border of the Mount Cydonia Ponds Natural Area.
The logging road is approximately 650 m long by 4 m wide, and currently has
somewhat limited commercial usage. Despite having spotty distribution within
the state, both Jefferson Salamander and Marbled Salamander adults are common
at this site in addition to the Spotted Salamander, which has a much wider
state distribution (Hulse et al. 2001). Red-spotted Newts can also be observed
frequently in the area and throughout the state, in both terrestrial, red-eft stage,
and aquatic (larva and adult) stages (Hulse et al. 2001).
Figure 1. Map of the Mount Cydonia Ponds Natural Area (shaded region) within
Michaux State Forest, Franklin County, PA. Logging road is labeled and indicated
by the dotted line.
2008 D.L. Chambers 125
Data acquisition
A total of 18 ephemeral pools and 6 permanent ponds along the logging road
were sampled from late-April to July 2006. Several of the observed pools were
visible from the road, but some were located deeper in the woodland area. All
sites were discovered by conducting intensive ground searches. Once a pool
was discovered, the distance (m) from the closest edge of the logging road to the
closest pool edge was measured and recorded. Thorough dip-netting occurred
to determine larval or aquatic adult presence of all four focal species within
each pool (Kolozsvary and Swihart 1999). In addition, the following abiotic
characteristics were recorded from each site: pool type (ephemeral or permanent),
area size at sampling time, percent canopy cover, dominant substrate
type (mud sediment or leaf litter), and depth. Approximately 130 man-hrs were
spent collecting data for this study.
Statistical analyses
All observational data were log transformed prior to any statistical analysis
to assist with normalization. A one-way analysis of variance (ANOVA) for
unequal variances was performed to determine if there was a general signifi-
cant difference amongst the focal species concerning their distance from the
logging road, the focal variable of this study. In addition, a pair-wise comparison
with a Tukey-Kramer adjustment was utilized for determining significant
differences between species regarding distance from the logging road.
Several other abiotic parameters were recorded for each pool sampled.
A logistic regression was conducted to determine if any of these parameters
could also be correlated to breeding-site selection for each species. All statistical
analyses were conducted using SAS (SAS Institute 2004).
Results
Table 1 shows the mean distance from the logging road (m ± standard error)
and number of occurrences among all (N = 24) pools for each sampled caudate
species. Table 2 lists the abiotic parameter data recorded from each pool.
Overall, there was a significant difference amongst all species in terms
of distance from the logging road and breeding-site selection (F3,8.08 = 13.17,
P = 0.0018). More specifically, there was a significant difference in distance
from the road between sites chosen by Jefferson Salamanders and Red-spotted
Newts (Tukey-Kramer Test, P = 0.0013), and between those of Marbled Salamanders
and Red-spotted Newts (Tukey-Kramer Test, P = 0.0208). However,
Table 1. Breeding-Pool mean distance from the logging road (± SE) (m) and number of pool
occurrences for each focal species.
Breeding-pool # of pool
mean distance occurrences
Focal species from road ± SE (m) (of 24)
Ambystoma jeffersonianum (Jefferson Salamander) 22.89 ± 2.43 3
A. maculatum (Spotted Salamander) 14.33 ± 4.37 4
A. opacum (Marbled Salamander) 14.30 ± 2.06 8
Notophthalmus viridescens (Red-spotted Newt) 7.37 ± 1.12 22
126 Northeastern Naturalist Vol. 15, No. 1
there were no significant differences in distance from the road between the following
species comparisons: Jefferson Salamanders and Spotted Salamanders
(Tukey-Kramer Test, P = 0.2880), Jefferson Salamanders and Marbeled Salamanders
(Tukey-Kramer Test, P = 0.0526), Spotted Salamanders and Marbled
Salamanders (Tukey-Kramer Test, P = 0.9965), and Spotted Salamanders
and Red-spotted Newts (Tukey-Kramer Test, P = 0.1353). In addition, no
significant differences were detected among any abiotic parameter relating to
breeding-site selection for all focal species (Table 3).
Discussion
The results indicate a correlative relationship between breeding-site
selection among focal salamander species and distance from the logging
road. However, no other abiotic factor significantly influenced breeding-site
selection in any focal species. This study adds supportive evidence to the
concept that ambystomatid salamanders can be influenced by habitat alterations
as they only bred in 3 (Jefferson Salamander), 4 (Spotted Salamander),
and 8 (Marbled Salamander) of 24 possible pools (see Table 1). In fact, ambystomatid
salamanders are suggested as being highly sensitive to various
habitat modifications, such as water quality alterations and urban development
(Lannoo 2005). Taylor et al. (2006) demonstrated poor reproductive
Table 2. Recorded abiotic parameter data from each breeding pool. Focal amphibian species
abbreviations are as follows: Ambystoma jeffersonianum (AJ), A. maculatum (AM), A. opacum
(AO), and Notophthalmus viridescens (NV). Pool numbers were assigned arbitarily.
Focal Pool size Estimated Dominant
Pool species at sampling Depth canopy substrate
# present Pool type (m2) (cm) cover (%) type
1 AJ, AO Ephemeral 55.66 7.62 0 Mud
2 AJ, NV Permanent 891.82 5.08 5 Mud
3 AJ, AM, AO Ephemeral 977.52 25.40 60 Leaf litter
4 AM, NV Permanent 386.91 39.67 25 Mud
5 AM, AO, NV Ephemeral 283.10 24.76 100 Leaf litter
6 AM, AO, NV Ephemeral 258.96 62.23 40 Leaf litter
7 AO, NV Ephemeral 201.54 27.31 30 Mud
8 AO, NV Ephemeral 3.33 13.34 70 Leaf litter
9 AO, NV Permanent 1275.52 73.66 85 Leaf litter
10 AO, NV Ephemeral 192.44 56.41 10 Mud
11 NV Ephemeral 54.78 41.48 15 Mud
12 NV Ephemeral 375.82 19.86 5 Mud
13 NV Ephemeral 35.28 8.34 20 Mud
14 NV Permanent 238.86 11.84 45 Leaf litter
15 NV Ephemeral 114.74 20.05 60 Leaf litter
16 NV Permanent 120.47 66.32 5 Mud
17 NV Ephemeral 109.5 25.61 50 Leaf litter
18 NV Ephemeral 19.89 11.12 55 Leaf litter
19 NV Ephemeral 94.41 17.50 40 Leaf litter
20 NV Ephemeral 30.01 28.64 0 Mud
21 NV Permanent 277.60 23.78 95 Leaf litter
22 NV Ephemeral 24.10 16.33 15 Mud
23 NV Ephemeral 371.96 20.17 25 Leaf litter
24 NV Ephemeral 95.40 30.10 60 Leaf litter
2008 D.L. Chambers 127
success and complete reproductive failure of Marbled Salamanders in over
50% and 27% of their observation years, respectively, with possible causes
being susceptibility to wetland and/or watershed disturbances.
Generalist species with broad habitat specificity can exist, and in some
instances, thrive in altered habitats. Based upon this study’s data, the Redspotted
Newt does not appear to be as sensitive to the disturbance caused by
logging roads: it occurred in 22 of 24 pools regardless of distance from the
road. Red-spotted Newts have a widespread distribution in Pennsylvania.
Range-wide, the Red-spotted Newt is the second-most widely distributed
North American caudate species (Petranka 1998). Therefore, its lack of breeding-
pool location preference in the Mount Cydonia Ponds Natural Area within
Michaux State Forest was expected. In addition to its widespread distribution,
the Red-spotted Newt has also been noted as a colonizing species (Hunsinger
and Lannoo 2005), even of degraded systems (Petranka 1998). Some populations
of Red-spotted Newt also have paedomorphic capability by retaining
gills into adulthood (Brandon and Bremer 1966). Harris (1987) suggested that
paedomorphosis can arise as an adaptation to altered environmental conditions.
Furthermore, Rubbo and Kiesecker (2005) claim that newts, along with
other amphibian species such as toads and various ranid frogs, can tolerate
sites subjected to urban development, while ambystomatid salamanders are
highly sensitive to those same sites. This study supports their claim, as Red-
Table 3. Logistic regression statistical results for abiotic parameters’ effect on breeding-site
selection.
Species Abiotic parameter Wald chi-square P-value*
Notophthalmus viridescens
Pool type 0.7781 0.3777
Pool size (m2) 0.9843 0.3211
Depth (cm) 0.9200 0.3375
Canopy cover (%) 0.3783 0.5385
Substrate type 0.1358 0.7125
Ambystoma jeffersonianum
Pool type 0.0216 0.8830
Pool size (m2) 0.7420 0.3890
Depth (cm) 0.7903 0.3740
Canopy cover (%) 0.2985 0.5848
Substrate type 0.0888 0.7658
A. maculatum
Pool type 0.6985 0.4033
Pool size (m2) 0.6378 0.4245
Depth (cm) 0.6891 0.4065
Canopy cover (%) 0.7705 0.3801
Substrate type 0.1181 0.7311
A. opacum
Pool type 2.0193 0.1553
Pool size (m2) 1.4835 0.2232
Depth (cm) 1.7722 0.1831
Canopy cover (%) 2.2168 0.1365
Substrate type 1.9916 0.1582
*Significance determined at alpha = 0.05.
128 Northeastern Naturalist Vol. 15, No. 1
spotted Newts displayed an unbiased distribution, while all 3 ambystomatid
salamander species did show breeding-site preference to varying degrees in
relation to the logging road. However, caution must be employed when comparing
two phylogenetically different groups (Notophthalmus and Ambystoma).
Several evolved life-history traits from each group make comparing their
breeding-site selection complicated. For example, Red-spotted Newts favor
permanent ponds for late life-history stages because they inhabit these systems
for their entire life (Petranka 1998). Thus, long-term population success
of Red-spotted Newts in ephemeral pools would not be expected. The focal
Ambystoma of this study, while occasionally breeding in permanent pools,
favor ephemeral pools due to the lack of predators (Petranka 1998). However,
from a management and conservation perspective, such comparisons
are needed to determine optimal practices. For instance, Rubbo and Kiesecker
(2005) emphasize the importance of quantifying multiple habitat parameters
for determining distributions of several phylogenetically distant amphibians
in anthropogenically disturbed habitats.
Aside from alterations in breeding-pool selection, there are other potential
problems associated with timber harvest activities, including subsequent
road construction through and between suitable habitats for amphibians.
Some of these impacts include: migration inhibition, decreased abundance
and species richness, complete population elimination, loss of genetic
diversity, and increased road mortality (Carr and Fahrig 2001, Dodd and
Smith 2003, Fahrig et al. 1995, Hels and Buchwald 2001, Petranka et al.
1994). Because amphibians are considered to be relatively poor dispersers
as compared to other taxa (Semlitsch 2000), any habitat alteration could be
extremely detrimental to them as they may not possess the ability to seek out
other suitable habitats within the vicinity of their mobility limitations.
Overall, anthropogenic habitat modifications can potentially alter breeding-
pool selection, and thus distribution, of pool-breeding salamanders.
Pool-breeding amphibians may often naturally experience frequent population
turnovers or even local extinctions, even in undisturbed habitats (Hecnar
and M’Closkey 1996, Skelly et al. 1999). If populations of pool-breeding
salamanders are naturally in a state of flux (Pechmann and Wilbur 1994), then
detrimental effects (such as local population declines and/or extinctions) at the
population level could arise as a result of anthropogenic disturbances (Gibbs
1993, Semlitsch and Bodie 1998). Unfortunately, freshwater systems have been
under increasing pressure recently, primarily because of habitat modifications,
and this trend may continue for years to come (Sala et al. 2000).
While distance from the logging road was the only significant correlate
to breeding-site selection found in this study, other factors potentially could
contribute to breeding-site selection. Future studies could attempt to analyze
water chemistry, specifically pH and nitrate levels, as several studies
have shown these factors to have negative effects on amphibians (Horne
and Dunson 1995, Marco et al. 1999). In addition, prey type and abundance
could be assessed at each pond. Ambystomatid salamanders and newt larvae
typically are voracious carnivores in their aquatic habitats, feeding upon
various zooplankton and invertebrate inhabitants (Petranka 1998). Thus, it
would be advantageous for these species to select pools with sufficient prey
2008 D.L. Chambers 129
items for their larvae to feed upon throughout development. While I did
not observe predatory fish in any pools, other possible predators in these
systems could also influence breeding-site selection. For instance, several
odonate and trichoptera larvae can consume amphibian larvae (Rubbo et al.
2006, Skelly 1994). Red-spotted Newts can also predate upon Ambystoma
larvae (Petranka 1998). This may be partially responsible for the limited
Ambystoma breeding-site selection observed here, as the Red-spotted Newt
was nearly uniform in distribution. Future studies could attempt to determine
the presence or interaction of these predators and assess their influence on
amphibian breeding-site selection.
Acknowledgments
I wish to thank Timothy J. Maret for his assistance and generosity in sharing his herpetofaunal
knowledge. Megan Saylor assisted in various fieldwork endeavors. Li Wang,
Wen Wang, and Hongzhang Zheng assisted with statistical applications. Lisa K. Belden,
David D. Chambers, Robert S. Covert, Doyle L. Crosswhite, and two anonymous reviewers
greatly improved earlier versions of this manuscript. All observational data was
acquired under a Pennsylvania Scientific Collectors Permit (no. 150, type 1). Funding
for this project was awarded by the Virginia Tech Department of Biological Sciences
and a Graduate Research and Development Grant from Virginia Tech.
Literature Cited
Bradford, D.F., J.R. Jaeger, and S.A. Shanahan. 2005. Distributional changes and
population status of amphibians in the Eastern Mojave Desert. Western North
American Naturalist 65:462–472.
Brandon, R.A., and D.J. Bremer. 1966. Neotenic newts, Notophthalmus viridescens
louisianensis, in southern Illinois. Herpetologica 22:213–217.
Carr, L.W., and L. Fahrig. 2001. Effect of road traffic on two amphibian species of
different vagility. Conservation Biology 15:1071–1078.
Dodd, C.K., Jr., and L.L. Smith. 2003. Habitat destruction and alteration: Historical
trends and future prospects for amphibians. Pp. 94–112, In R.D. Semlitsch (Ed.).
Amphibian Conservation. Smithsonian Institution, Washington, DC.
Fahrig, L., J.H. Pedlar, S.E. Pope, P.D. Taylor, and J.F. Wegner. 1995. Effect of road
traffic on amphibian density. Biological Conservation 73:177–182.
Gibbons, J.W., C.T. Winne, D.E. Scott, J.D. Willson, X. Glaudas, K.M. Andrews,
B.D. Todd, L.A. Fedewa, L. Wilkinson, R.N. Tsaliagos, S.J. Harper, J.L. Greene,
T.D. Tuberville, B.S. Metts, M.E. Dorcas, J.P. Nestor, C.A. Young, T. Akre, R.N.
Reed, K.A. Buhlmann, J. Norman, D.A. Croshaw, C. Hagen, and B.B. Rothermel.
2006. Remarkable amphibian biomass and abundance in an isolated wetland: Implications
for wetland conservation. Conservation Biology 20:1457–1465.
Gibbs, J.P. 1993. Importance of small wetlands for the persistence of local populations
of wetland-associated animals. Wetlands 13:25–31.
Harris, R.N. 1987. Density-dependent paedomorphosis in the salamander, Notophthalmus
viridescens dorsalis. Ecology 68:705–712.
Hecnar, S.J., and R.T. M’Closkey. 1996. Regional dynamics and the status of amphibians.
Ecology 77:2091–2097.
Hels, T., and E. Buchwald. 2001. The effect of road kills on amphibian populations.
Biological Conservation 99:331–340.
Horne, M.T., and W.A. Dunson. 1995. Toxicity of metals and low pH to embryos
and larvae of the Jefferson Salamander, Ambystoma jeffersonianum. Archives of
Environmental Contamination and Toxicology 29:110–114.
130 Northeastern Naturalist Vol. 15, No. 1
Hulse, A.C., C.J. McCoy, and E.J. Censky. 2001. Amphibians and Reptiles of Pennsylvania
and the Northeast. Cornell University Press, Ithaca, NY. 419 pp.
Hunsinger, T.W., and M.J. Lannoo. 2005. Notophthalmus viridescens (Rafinesque,
1820): Eastern Newt. Pp. 889–894, In M. Lannoo (Ed.). Amphibian Declines:
The Conservation Status of United States Species. University of California Press,
Berkeley, CA.
Kolozsvary, M.B., and R.K. Swihart. 1999. Habitat fragmentation and the distribution
of amphibians: Patch and landscape correlates in farmland. Canadian Journal
of Zoology 77:1288–1299.
Lannoo, M. (Ed.). 2005. Amphibian Declines: The Conservation Status of United
States Species. University of California Press, Berkeley, CA.
Marco, A., C. Quilchano, and A.R. Blaustein. 1999. Sensitivity to nitrate and nitrite
in pond-breeding amphibians of the Pacific northwest, USA. Environmental
Toxicology and Chemistry 18:2836–2839.
Pechmann, J.H.K., and H.M. Wilbur. 1994. Putting declining amphibian populations in
perspective: Natural fluctuations and human impacts. Herpetologica 50:65–84.
Peltzer, P.M., R.C. Lajmanovich, and A.D. Beltzer. 2003. The effects of habitat fragmentation
on amphibian species richness in the floodplain of the Middle Parana
River, Argentina. Herpetological Journal 13:95–98.
Petranka, J.W. 1998. Salamanders of the United States and Canada. Smithsonian
Institution Press, Washington, DC. 587 pp.
Petranka, J.W., M.E. Eldridge, and K.E. Haley. 1993. Effects of timber harvesting on
southern Appalachian salamanders. Conservation Biology 7:363–370.
Petranka, J.W., M.P. Brannon, M.E. Hopey, and C.K. Smith. 1994. Effects of timber
harvesting on low-elevation populations of southern Appalachian salamanders.
Forest Ecology and Management 67:135–147.
Rubbo, M.J., and J.M. Kiesecker. 2005. Amphibian breeding distribution in an urbanized
landscape. Conservation Biology 19:504–511.
Rubbo, M.J., K. Shea, and J.M. Kiesecker. 2006. The influence of multi-stage predation
on population growth and the distribution of the pond-breeding salamander,
Ambystoma jeffersonianum. Canadian Journal of Zoology 84:449–458.
SAS Institute. 2004. SAS System for Windows. Release 9.1.2 edition. SAS Institute,
Cary, NC.
Sala, O.E., F.S. Chapin III, J.J. Armesto, E. Berlow, J. Bloomfield, R. Dirzo, E. Huber-
Sanwald, L.F. Huenneke, R.B. Jackson, A. Kinzig, R. Leemans, D.M. Lodge, H.A.
Mooney, M. Oesterheld, N.L. Poff, M.T. Sykes, B.H. Walker, M. Walker, and D.H.
Wall. 2000. Global diversity scenarios for the year 2100. Science 287:1770–1774.
Semlitsch, R.D. 2000. Principles of management of aquatic breeding amphibians.
Journal of Wildlife Management 64:615–631.
Semlitsch, R.D. 2003. Conservation of pond-breeding amphibians. Pp. 8–23, In R.D.
Semlitsch (Ed.). Amphibian Conservation. Smithsonian Institution, Washington, DC.
Semlitsch, R.D., and J.R. Bodie. 1998. Are small, isolated wetlands expendable?
Conservation Biology 12:1129–1133.
Skelly, D.K. 1994. Activity level and the susceptibility of anuran larvae to predation.
Animal Behaviour 47:465–468.
Skelly, D.K., E.E. Werner, and S.A. Cortwright. 1999. Long-term distributional dynamics
of a Michigan amphibian assemblage. Ecology 80:2326–2337.
Taylor, B.E., D.E. Scott, and J.W. Gibbons. 2006. Catastrophic reproductive failure,
terrestrial survival, and persistence of the Marbled Salamander. Conservation
Biology 20:792–801.
Wyman, R.L. 1990. What’s happening to the amphibians? Conservation Biology 4:
350–352.