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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

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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. 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