The Relative Abundance of the Juvenile Phase of the
Eastern Red-Spotted Newt at Harvard Forest Prior to the
Arrival of the Hemlock Woolly Adelgid
Brooks G. Mathewson*
Abstract - The invasive insect pest Adelges tsugae (Hemlock Woolly Adelgid) threatens the
ecologically unique Tsuga canadensis (Eastern Hemlock)-dominated forest type throughout
its range. Relatively little is known about how the loss of this forest type will affect the relative
abundance of amphibians. This study assessed the relative abundance of the juvenile
phase of Notophthalmus viridescens viridescens (Eastern Red-spotted Newt, Red Eft) in
Eastern Hemlock-dominated stands (n = 5) and mixed deciduous stands (n = 5) at Harvard
Forest in Petersham, MA, using both transect surveys of the forest floor surface (n = 368
Red Eft observations over four seasons), and intensive searches of quadrats (n = 27 Red
Eft observations over two seasons). Using data from transect surveys, the average relative
abundance of Red Efts was more than two times greater in Eastern Hemlock-dominated
stands than in mixed deciduous stands, however the differences were not statistically significant
(P = 0.146). Quadrat surveys yielded relative abundance estimates for Red Efts
that were more than 5 times greater in Eastern Hemlock-dominated stands than in mixed
deciduous stands, but again the differences were not statistically significant (P = 0.213).
The long-lived, shade tolerant conifer species Tsuga canadensis Carrière (Eastern
Hemlock) has been described as a foundation species that creates unique habitat
and impacts core ecosystem processes (Ellison et al. 2005a). This ecologically
important species is threatened throughout its range by the invasive insect, Adelges
tsugae Annand (Hemlock Woolly Adelgid [HWA]) (Hemiptera: Adelgidae; Orwig
2002, Orwig and Foster 1998). Native to Japan, HWA was first discovered in Virginia
in the 1950s (Souto et al. 1996) and has spread throughout a great percentage
of Eastern Hemlock’s range via a number of dispersal agents including wind, birds,
deer, and humans (McClure 1990). As of 2004, when this study was conducted,
HWA was present in 50% of Eastern Hemlock-dominated stands in Massachusetts,
but was not yet present at Harvard Forest (Orwig and Povak 2004). Unfortunately,
no natural predators of the aphid-like insect occur in the United States (McClure
1995). HWA can cause mortality in all age classes of Eastern Hemlock within
4–10 years of infestation (McClure 1991). In central Massachusetts, cold winter
temperatures have slowed mortality of Eastern Hemlock in infested stands, though
anticipated warming trends threaten to accelerate rates of mortality and dispersal
(Orwig et al. 2012). At Harvard Forest, Eastern Hemlock will likely be replaced by
mixed deciduous species such as Betula lenta L. (Black Birch), Quercus rubra L.
*Harvard Forest, Harvard University, Petersham, MA 01366; firstname.lastname@example.org.
Manuscript Editor: Jeff Houlahan
Forest Impacts and Ecosystem Effects of the Hemlock Woolly Adelgid in the Eastern US
2014 Southeastern Naturalist 13(Special Issue 6):117–129
Vol. 13, Special Issue 6
(Red Oak), and Acer rubrum L. (Red Maple) (Orwig and Foster 1998, Sullivan and
Eastern Hemlock-dominated forests are structurally unique, providing important
habitat to assemblages of invertebrates, amphibians, birds, and mammals (Ellison et
al. 2005b, Ingwell et al. 2012, Mathewson 2009, Tingley et al. 2002, Yamasaki et al
2000). For example, the dense canopy of Eastern Hemlock-dominated forests provides
breeding habitat preferred by several songbird species including Dendroica
virens Gmelin (Black-throated Green Warbler), Vireo solitarius Wilson (Solitary
Vireo), and Dendroica fusca Müller (Blackburnian Warbler) (Benzinger 1994, Tingley
et al. 2002, Yamasaki et al. 2000). These dense canopies greatly reduce light
penetration resulting in forest floors that are cooler, darker, and with moister soil
than surrounding mixed deciduous stands (Benzinger 1994, Lustenhouwer et al.
2012, Rogers 1980). Many groups of invertebrates are more abundant in Eastern
Hemlock litter than mixed deciduous litter including collembolans, mites and ticks,
coleopterans, hymenopterans, and dipterans (8,5,4,2.5, and 2.5 times more abundant,
respectively; Hartman 1977).
Soils in Eastern Hemlock-dominated forests are more acidic than in mixed
deciduous forests due to the species ability to thrive in acidic conditions and the
acidity of hemlock needles themselves (Benzinger 1994). This association with
high soil acidity led to the perception that amphibians are less abundant in Eastern
Hemlock forests. Wyman and Jancola (1992) suggested the relative abundance of
Plethodon cinereus Green (Eastern Red-backed Salamander) was found to be lower
in Eastern Hemlock-dominated stands than in Fagus grandifolia Ehrh. (American
Beech) stands in Albany County, NY due to higher soil acidity in the former. However,
the relative abundance of Eastern Red-backed Salamanders was found to be
greater in Hemlock-dominated stands than in mixed deciduous stands at Harvard
Forest using surveys of artificial cover objects (ACOs; Mathewson 2009). No difference
was found in the relative abundance of Eastern Red-backed Salamanders
in the two forest types using intensive searches of quadrats (Mathewson 2009). At
Harvard Forest, soil pH in Eastern Hemlock-dominated stands, while lower than
in mixed deciduous stands, is above the level that negatively impacts the relative
abundance of Eastern Red-backed Salamanders (Mathewson 2009, Wyman and
Notophthalmus viridescens viridescens Rafinesque (Eastern Red-spotted Newt)
is the second most widely distributed salamander in North America (Petranka 1998).
It is also perhaps the most familiar salamander, especially as a terrestrial juvenile,
or Red Eft, due to its bright coloration and active diurnal behavior on the surface of
the forest floor (Petranka 1998). This bright coloration serves as a warning to potential
predators of the Red Eft’s highly toxic skin (Hurlbert 1970). Although there
are several variations, the most common life cycle involves 4 distinct stages—egg,
aquatic larva, terrestrial Red Eft, and aquatic adult (Petranka 1998). The Red Eft
stage usually lasts from 4–7 years (Petranka 1998). Despite their ubiquity, little
is known regarding differences in the relative abundance of Red Efts in different
forest types, and no study has ever assessed the relative abundance of Red Efts in
2014 Vol. 13, Special Issue 6
forests dominated by Eastern Hemlock. The only estimate of the relative abundance
of Red Efts comes from an oak-pine woodland located 800 m from a breeding pond
in central Massachusetts where the density of Red Efts was 0.03 individuals/m2
Red Efts do not appear to be affected by low soil pH likely because their skin
is coarser than the lungless salamanders, making them less sensitive to acidic soils
(Wyman and Jancola 1992). Therefore, low soil pH does not likely impact the
relative abundance of Red Efts in Eastern Hemlock-dominated forests at Harvard
Forest. However, food supply and moisture are important factors in habitat selection
by Red Efts, and both factors may be favorable in Eastern Hemlock-dominated
stands (Healy 1975). In addition, Eastern Hemlock-dominated stands have an
abundance of mushrooms, and Red Efts are often observed feeding on invertebrates
found around decaying mushrooms (MacNamara 1977). Finally, anecdotal observations
of Red Efts in Eastern Hemlock-dominated stands at Harvard Forest suggests
that the abundance of this species is greater in Eastern Hemlock stands than in
mixed deciduous stands (B.G. Mathewson, pers. observ.).
If the relative abundance of Red Efts is higher in Eastern Hemlock-dominated
stands than in mixed deciduous stands, a transition to mixed deciduous stands
due to HWA could lead to a reduction in the relative abundance of Red Efts at
these sites. Less desirable terrestrial habitat may also impact aquatic communities.
Only 1–2% of Eastern Red-spotted Newts survive the larval stage to become
Red Efts (Petranka 1998). Therefore, a change in survivorship of the juvenile
phase could have an important impact on population densities of the Eastern
Salamanders inhabiting the forest floor are ecologically important due to their
significant contribution to the overall biomass of vertebrates in the forest, and
their position in the middle of the food web (Burton and Likens 1975a, Welsh
and Droege 2001). At Hubbard Brook Experimental Forest in Coos Country, NH,
salamander biomass equaled small-mammal biomass, and was twice the biomass
of breeding birds (Burton and Likens 1975a). Red-backed Salamanders accounted
for 93.5% of salamander biomass with a density of 0.25 individuals/m2 at Hubbard
Brook, while Red Efts were rare due to the lack of suitable aquatic breeding
habitat within 1 km of study sites (Burton and Likens 1975a).
While no research has been conducted on the role of Red Efts in nutrient cycling
or decomposition rates, it has been hypothesized that predation by Red-backed
Salamanders on soil invertebrates that break down leaf litter reduces the rate of soil
decomposition by decreasing the amount of surface area available to bacteria and
fungi (Wyman 1998). Slowing down decomposition of organic matter on the forest
floor slows down the rates of CO2 emitted into the atmosphere (Wyman 1998).
Thus, reducing predation on invertebrate decomposers may have major implications
on the global carbon budget as it is estimated that six times as much CO2 is
emitted into the atmosphere by the breakdown of leaf litter as by any anthropogenic
source (Wyman 1998). As predators of soil invertebrates, it is also possible that Red
Efts have an impact on decomposition rates. Red Efts prey on a great diversity of
Vol. 13, Special Issue 6
invertebrates, including representatives from 25 orders and 58 families (MacNamara
1977). MacNamara (1977) and Burton (1976) reported that Red Efts’ preferred
prey (by percentage of overall diet by weight) are land snails (23.8% and 59.7%),
mites and ticks (13.8% and 3.4%), springtails (10.6% and 9.1%), dipteran adults
(9.7% and 8.8%), and lepidopteran larvae (7.9% and 2.3%).
Salamanders of the forest floor are also important as prey to larger vertebrates
such as snakes, birds, and small mammals (Welsh and Droege 2001). Due to low
metabolic rates, salamanders are extremely efficient at converting prey into protein,
which is then passed up the food chain (Burton and Likens 1975b). However,
Red Efts may not be as important prey to larger vertebrates as other salamanders
because of toxins in their skin (Brodie 1968, Hurlbert 1970, Uhler 1939). Many
potential diurnal predators including Charadrius vociferus L. (Killdeer), Buteo
jamaicensis Gmelin (Red-tailed Hawk), Falco sparverius L. (American Kestrel),
and Thamnophis sirtalis L. (Common Garter Snake) find Red Efts to be unpalatable
(Hurlbert 1970, Uhler 1939). Other predators such as Rana catesbeiana Shaw
(American Bullfrog), Procyon lotor L. (Raccoon), and Bufo americanus Holbrook
(American Toad) appear to be less sensitive to the toxins in Red Efts’ skin (Brodie
1968a, Hurlbert 1970).
I hypothesized that the relative abundance of Red Efts would be higher in Eastern
Hemlock-dominated stands than in mixed deciduous stands based on preliminary
field observations as well as the presence of biotic and abiotic habitat features preferred
by Red Efts (Benzinger 1994, Hartmann 1977, Healy 1975, Lustenhouwer
et al. 2012, Rogers 1980). In addition to testing this hypothesis, a secondary goal
of this study was to look for relationships between the relative abundance of Red
Efts and the average daily minimum and maximum temperatures in the spring and
fall, soil pH, and estimated distances to potential breeding habitat. The third goal
of this study was to establish baseline data on the relative abundance of Red Efts at
Field Site Description
This study was conducted in 10 second-growth stands at Harvard Forest in Petersham,
MA (42.533°N, 72.190°W; 338 m elev.). I chose 1 mixed deciduous stand
and 1 Eastern Hemlock-dominated stand at the Prospect Hill, Tom Swamp, and Slab
City tracts, and 2 mixed deciduous and 2 Eastern Hemlock-dominated stands on the
Simes tract. One of the stands, the mixed deciduous stand in the Tom Swamp tract
(hereafter referred to as TS-MD), was selectively logged in 1998. The average distance
from potential breeding habitats to the center of stands, estimated using maps
in the lab, was 545 m in Eastern Hemlock-dominated stands and 430 m in mixed
deciduous stands (Table 1). When this study was conducted, HWA was not known
to be present in any of the stands studied.
I used tree species composition data from the Harvard Forest Archives to select
stands and then qualitatively verified stand type in the field (Foster 1992). Eastern
Hemlock contributed 63% of the total basal area in the Eastern Hemlock-dominated
stand at Simes 1, and 60% in Simes 2 (Ellison et al. 2010). The dominant
2014 Vol. 13, Special Issue 6
overstory tree species in the mixed deciduous stand at Simes 1 were Red Oak
(36%), Black Birch (24%), Red Maple (13%), and Acer saccharum Marsh. (Sugar
Maple) (11%) (Ellison et al. 2010). In the mixed deciduous stand at Simes 2, the
dominant overstory tree species were Pinus strobus L. (Eastern White Pine) (35%),
Red Maple (17%), Black Birch (15%), and Red Oak (15%) (Ellison et al. 2010).
Quantitative data for overstory tree species composition data was not available in
the Prospect Hill, Slab City, or Tom Swamp sites. I qualitatively assessed Eastern
Hemlock-dominated stands at these sites to be greater than 50% Eastern Hemlock.
The primary species in mixed deciduous stands at these sites were Red Oak, Black
Birch, Eastern White Pine, and Red Maple (Table 1).
Table 1. Description and measurements of environmental variables in 10 forest stands at 5 sites at
Harvard Forest in Petersham, MA. Site codes are as follows: PH = Prospect Hill, S1 = Simes 1, S2 =
Simes 2, SC = Slab City, TS = Tom Swamp. FT indicates forest type (EH = Eastern Hemlock-dominated;
MD = mixed deciduous). Tree species codes are as follows: TSCA = Tsuga canadensis (Eastern
Hemlock), PIST = Pinus strobus (Eastern White Pine), QUVE = Quercus velutina Lam. (Black Oak),
QURU = Quercus rubra (Northern Red Oak), QUAL = Quercus alba (White Oak), BEPO = Betula
populifolia Marshall (Gray Birch), ACRU = Acer rubrum (Red Maple), BELE = Betula lenta (Black
Birch). Area = area covered by transects; dist = estimated distance to potential breeding habitat; low
temp = average daily low temperature; high temp = average daily high temperature. Spring = 22 April
2004–7 June 2004; Fall = 22 September 2004–12 November 2004. SD = standard deviation.
Stand Low High Low High
Tree Latittude size Area Dist Soil temp temp temp temp
Site FT species comp longitude (ha) (m2) (m) pH (ºC) (ºC) (ºC) (ºC)
PH MD QUVE-QURU 42°32.441' 1.0 180 700 4.2 7.4 22.9 7.6 13.1
PH EH TSCA-PIST 42°32.372' 1.0 180 500 4.1 7.2 18.9 5.4 11.4
S1 MD BELE-QURU 42°27.956' 1.0 180 500 4.4 7.6 25.6 5.2 13.6
S1 EH TSCA-QURU 42°28.313' 3.0 540 50 4.0 7.2 21.0 4.9 11.9
S2 MD PIST-BELE 42°28.758' 1.0 180 500 4.5 7.6 20.4 5.6 13.1
S2 EH TSCA-BELE 42°28.511' 3.0 540 500 4.2 7.6 18.4 5.6 11.7
SC MD QURU-ACRU 42°27.076' 0.4 185 1000 4.3 7.2 24.1 4.8 11.4
SC EH TSCA-QURU 42°27.192' 0.5 248 850 4.1 7.4 18.7 5.2 11.4
TS MD QUAL- QURU 42°30.232' 1.0 312 25 4.4 7.6 24.8 5.7 12.8
- ACRU 72°12.683'
TS EH TSCA-PIST 42°30.400' 1.0 248 250 4.0 7.6 18.5 5.1 11.2
MD Avg 0.9 207 545 4.4 7.5 23.6 5.8 12.8
SD (0.3) (59) (355) (0.1) (0.2) (2.0) (1.1) (0.8)
EH Avg 1.7 351 430 4.1 7.4 19.1 5.2 11.5
SD (1.2) (175) (301) (0.1) (0.2) (1.1) (0.3) (0.3)
Vol. 13, Special Issue 6
Red Eft sampling
My first method of assessing the relative abundance of Red Efts was daytime
visual surveys of the surface of the forest floor (hereafter referred to as transect
surveys). I conducted transect surveys 13–16 times in each stand on a total of 51
sampling days from 14 August 2003–29 October 2004 by walking transects and
counting all Red Efts that were visible on the surface of the forest floor entirely
within 0.5 m to the left and right of the transect. Natural cover objects such as
rocks and stones were not turned over during these searches. Transects varied in
length from 76–108 m because I established lengths so as to not extend the transects
beyond the edge of the stand, and the distance to the edge of the stand was
not constant. Each transect origin was chosen randomly. The area sampled in each
stand was between 180–540 m2, depending on the size of the stand (Table 1). I also
randomly chose the order in which I sampled the stands; not all sites were sampled
on the same day, but both forest types at a site were sampled on the same day.
The second method of sampling for Red Efts was 2-minute time-constrained
searches of natural cover objects (NCO) such as coarse woody debris, stones, and
leaves in 1-m2 quadrats on the surface of the forest floor (hereafter referred to as
quadrat surveys). In each stand, I searched 20 quadrats during fall 2003 and 20
quadrats during spring 2004. I placed quadrats at randomly selected points along
the same transects used in transect surveys, used a random number generator to determine
sampling points along transects, and flipped a coin to determine whether to
place the quadrat to the left or right of the transect. The order in which stands were
sampled was random, and the same quadrat was never searched twice. Following
searches, I returned all NCOs to their original position. I sampled both forest types
on the same day at all sites except at Simes 2 when I sampled the two stands on
consecutive days during the fall.
Measurements of habitat variables
I measured average daily high and low temperatures for each stand in the spring
(22 April 2004–7 June 2004) and fall (22 September 2004–12 November 2004)
using remote temperature sensors (1-Wire Thermochron iButtons ± 1 °C, Maxim
Integrated, San Jose, CA) that I placed on the soil surface in the center of each transect.
These sensors recorded temperature every half hour in spring 2004 and every
hour in fall 2004. To determine soil pH, I took 10–30 random samples from the
organic layer of the soil just below the leaf litter in each stand, and used a Thermo
Orion model 290 pH meter (± 0.005) to measure the pH of a slurry of 2.0 g of soil
from each sample in 20 ml deionized water (Hendershot et al. 1993).
Measurements of precipitation, relative humidity, and hourly temperature from
the Fisher Meteorological Station on the Prospect Hill Tract at Harvard Forest
were used to report the weather conditions on all sampling days for both transect
and quadrat methods. Weather conditions during transect surveys were reported
for sites as opposed to individual stands because transect surveys of both forest
types took only a few hours to complete and were completed in succession.
2014 Vol. 13, Special Issue 6
Weather conditions during quadrat surveys were reported for individual stands as
these surveys took more time and stands within a given site were often conducted
at different times of the day, or in one case on different days. On several occasions
the exact time of transect surveys was not recorded, and on one occasion exact
time of quadrat surveys was not recorded. In these instances, weather conditions
at noon on sampling day were used since this was the most frequent time searches
All observations of Red Efts during transect and quadrat surveys were
pooled to calculate the average relative abundance for each stand expressed as
salamanders/m2. In addition, I calculated the average relative abundance of Red
Efts for surveys that were conducted when a rain event had and had not occurred in
the prior 24 hours. I then used t-tests to test for differences in the average relative
abundance of Red Efts in the two forest types using the two methods of sampling
for all samples and for samples conducted within 24 hours of a rain event. All of
these tests were run with and without TS-MD. Analyses without TS-MD were run
because previous studies have found that selective harvesting can reduce the relative
abundance of Plethodontids, and the same may be true for Red Efts (Harpole
and Haas 1999). Further, piles of slash and dense stands of young vegetation may
have reduced the probability of detection in this stand. I also conducted t-tests to
evaluate differences in the estimated distance to potential breeding habitat, soil pH,
and average daily high and low temperatures for each stand in the spring (22 April
2004–7 June 2004) and fall (22 September 2004–12 November 2004) in the two
forest types. I used standard least squares regression analyses to test for individual
relationships between each of the above variables and the relative abundance of
Red Efts derived from the average of all transect surveys in a stand, both with and
without TS-MD. All statistical tests were run using the statistical software program
JMP IN version 5.1 (SAS Institute).
The average relative abundance of Red Efts derived from transect surveys (n =
368 observations) was higher in Eastern Hemlock-dominated stands than in mixed
deciduous stands, but the difference was not statistically significant (with TS-MD:
0.020 individuals/m2 vs. 0.009 individuals/m2, P = 0.146; without TS-MD: 0.020 individuals/
m2 vs. 0.011 individuals/m2, P = 0.230). The same was true for the average
relative abundance of Red Efts derived from quadrat surveys (n = 27 observations)
(with TS-MD: 0.115 individuals/m2 vs. 0.020 individuals/m2, P = 0.213; without
TS-MD: 0.115 individuals/m2 vs. 0.025 individuals/m2, P = 0.234). The average
relative abundance of Red Efts derived from transect surveys conducted within
24 hrs of a rain event (n = 307 observations) was also higher in Eastern Hemlockdominated
stands (with TS-MD: 0.036 individuals/m2 vs. 0.016 individuals/m2, P =
0.136; without TS-MD: 0.036 individuals/m2 vs. 0.019 individuals/m2, P = 0.209),
as was the average relative abundance of Red Efts derived from quadrat surveys
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within 24 hrs of a rain event (n = 16 observations) (with TS-MD: 0.146 individuals/
m2 vs. 0.025 individuals/m2, P = 0.153; without TS-MD: 0.146 individuals/m2 vs.
0.033 individuals/m2, P = 0.175) (Table 2), though, again, the differences were not
Soil pH was significantly lower in Eastern Hemlock-dominated stands than in
mixed deciduous stands (4.1 vs. 4.4; P < 0.01) as was the average high temperature
in the spring (19.1 °C vs. 23.6 °C; P < 0.01) and the average high temperature in
the fall (11.5 °C vs. 12.8 °C; P < 0.05) (Table 1). The difference in estimated distance
to potential breeding habitat in Eastern Hemlock-dominated stands versus
mixed deciduous stands was not significant (430 m vs. 545 m; P = 0.600) (Table 1).
Neither was the difference in average low temperature in spring (7.4 °C vs. 7.5 °C;
P = 0.524) (Table 1) and the average low temperature in fall (5.2 °C vs. 5.8 °C; P =
0.332) (Table 1).
Regression analyses did not reveal a statistically significant relationship between
any of the variables and the average relative abundance of Red Efts derived
from transect surveys. However, when removing TS-MD, a statistically significant
relationship was found between distance to potential breeding habitat and the average
relative abundance of Red Efts derived from transect surveys (n = 9, r2 adj =
0.76, P < 0.01) (Fig. 1). When TS-MD was included the results were not significant
(n = 10, r2 adj = 0.20, P < 0.19).
Table 2. Measurements of the average relative abundance (given in individuals/ m2) of Red Efts in10
forest stands at Harvard Forest. Transect surveys of the forest floor surface conducted from fall 2003
to fall 2004 (excluding winter). Quadrat surveys of 1-m2 quadrats conducted in fall 2003 and spring
2004. Site codes are as follows PH = Prospect Hill, S1 = Simes 1, S2 = Simes 2, SC = Slab City, TS =
Tom Swamp. FT indicates forest type (EH = Eastern Hemlock dominated; MD = mixed deciduous).
NA indicates that no sampling was conducted under these conditions.
Average relative abundance of Red Efts
Transect surveys Quadrat surveys
Within 24 hrs rain event Within 24 hrs rain event
Site FT All of rain event in prior 24hrs All of rain event in prior 24hrs
PH MD 0.012 0.017 0.002 0.050 0.050 NA
PH EH 0.020 0.029 0.000 0.050 0.050 NA
S1 MD 0.016 0.033 0.001 0.025 0.050 0.000
S1 EH 0.031 0.061 0.007 0.350 0.200 0.500
S2 MD 0.015 0.021 0.006 0.025 NA 0.025
S2 EH 0.028 0.041 0.000 0.150 0.300 0.000
SC MD 0.002 0.006 0.000 0.000 0.000 0.000
SC EH 0.000 0.000 0.000 0.000 NA 0.000
TS MD 0.001 0.001 0.001 0.000 0.000 0.000
TS EH 0.019 0.048 0.009 0.025 0.033 0.000
MD Avg. 0.009 0.016 0.002 0.020 0.025 0.006
SD (0.007) (0.013) (0.002) (0.021) (0.029) (0.013)
EH Avg. 0.020 0.036 0.003 0.115 0.146 0.125
SD (0.012) (0.023) (0.004) (0.143) (0.127) (0.250)
2014 Vol. 13, Special Issue 6
While these results do not confirm the hypothesis that the relative abundance
of Red Efts is significantly greater in Eastern Hemlock-dominated stands than in
mixed deciduous stands, they are suggestive of this hypothesis. Indeed, using data
from transect surveys, 4 of 5 stands with the highest relative abundance of Red Efts
were Eastern Hemlock-dominated stands. Further study with a larger sample size
may yield statistically significant results. The extent of the differences in the relative
abundance of Red Efts in the two forest types may be large given that transect
surveys yielded estimates in Hemlock-dominated stands that were more than two
times greater and quadrat surveys yielded estimates which were almost six times
greater than those for mixed deciduous stands. If the relative abundance of Red Efts
is greater in Eastern Hemlock-dominated forests, a shift from this forest type to the
mixed deciduous forest type due to HWA would likely negatively impact populations
of Eastern Red-spotted Newt at Harvard Forest.
The Red Eft, of course, is just one phase in the life cycle of the Eastern Redspotted
Newt. The loss of Eastern Hemlock along wetland borders may impact
Figure 1. Relationship between the average relative abundance of Red Efts as measured by
transect surveys and the estimated distance to potential breeding habitat at Harvard Forest
(42.533ºN, 72.190ºW; 338 m elev.)—including the Tom Swamp mixed deciduous stand (n =
10, r2 adj = 0.20, P < 0.19)—excluding the Tom Swamp mixed deciduous stand (n = 9, r2
adj = 0.76, P < 0.01).
Vol. 13, Special Issue 6
aquatic adult and larval phases as well. For example, a shift from Eastern Hemlock
to mixed deciduous species along wetland borders could cause increases in
solar radiation in the late winter and early spring before mixed deciduous trees
Table 3. Sampling effort and average weather conditions at the time of transect surveys. All weather
data were measured at the Fisher Meteorological Station on the Prospect Hill Tract at Harvard Forest
(42.533ºN, 72.190ºW; 338 m elev.). Site codes are as follows PH = Prospect Hill, S1 = Simes 1, S2 =
Simes 2, SC = Slab City, TS = Tom Swamp. RH = relative humidity. The first number in each cell
is the average for all sampling dates. The second number in each cell is the average for all sampling
dates conducted within 24 hours of a rain event. The third number in each cell is the average for all
sampling dates conducted when a rain event had not occurred within the previous 24 hours. SD =
Avg total days sampled
precipitation within 24 hrs
Site n Temp (°C) RH (%) prior 24 hrs (mm) of precipitation
PH 13 (9, 4) 15.0 (15.1, 14.7) 76 (89, 50) 8.3 (11.9, 0.0) 71 (100, 0)
S1 18 (11, 9) 17.5 (15.3, 20.2) 64 (72, 56) 2.8 (5.0, 0.0) 55 (100, 0)
S2 17 (10, 7) 16.5 (15.6, 20.6) 66 (77, 52) 6.0 (10.3, 0.0) 59 (100, 0)
SC 18 (8, 10) 17.4 (17.3, 18.4) 63 (84, 51) 2.3 (5.8, 0.0) 43 (100, 0)
TS 18 (7, 11) 17.6 (14.2, 19.8) 65 (77, 58) 4.9 (12.7, 0.0) 39 (100, 0)
Avg 17 (9, 8) 16.8 (15.5, 18.7) 67 (80, 53) 4.9 (9.1, 0.0) 53 (100, 0)
SD 2 (2, 3) 1.1 (1.1, 2.4) 5 (7, 3) 2.4 (3.5, 0.0) 13 (0, 0)
Table 4. Average weather conditions at the time of quadrat surveys in 10 forest stands at Harvard
Forest. All weather data were measured at the Fisher Meteorological Station on the Prospect Hill
Tract at Harvard Forest (42.533ºN, 72.190ºW; 338 m elev.). Site codes are as follows PH = Prospect
Hill, S1 = Simes 1, S2 = Simes 2, SC = Slab City, TS = Tom Swamp. FT indicates forest type (EH =
Eastern Hemlock-dominated; MD = mixed deciduous). The first number in each cell is the average
for all sampling dates. The second number in each cell is the average for all sampling dates conducted
within 24 hours of a rain event. The third number in each cell is the average for all sampling dates
conducted when a rain event had not occurred within the previous 24 hours. SD = standard deviation.
Number of Average total
quadrats Relative precipitation
Site FT surveyed Temperature (°C) humidity (%) prior 24 hrs (mm)
PH MD 40 (40, 0) 20.1 (20.1, na) 60 (60, na) 10.3 (10.3, na)
PH EH 40 (40, 0) 19.8 (19.8, na) 63 (63, na) 10.3 (10.3, na)
SI1 MD 40 (20, 20) 19.8 (14.9, 24.8) 60 (74, 47) 9.3 (18.5, 0.0)
SI1 EH 40 (20, 20) 19.2 (14.4, 24.0) 60 (71, 49) 5.3 (10.5, 0.0)
SI2 MD 40 (0, 40) 19.7 (na, 19.7) 60 (na, 60) 0.0 (na, 0.0)
SI2 EH 40 (20, 20) 18.9 (18.9, 18.8) 65 (96, 34) 7.2 (14.3, 0.0)
SC MD 40 (20, 20) 13.5 (16.0, 11.0) 47 (39, 55) 0.2 (0.3, 0.0)
SC EH 40 (0,40) 15.0 (na, 15.0) 43 (na, 43) 0.0 (na, 0.0)
TS MD 40 (30, 10) 10.4 (7.9, 16.7) 56 (61, 45) 1.9 (2.5, 0.0)
TS EH 40 (30, 10) 12.3 (10.9, 16.7) 45 (45, 45) 1.9 (2.5, 0.0)
MD avg (n = 5) MD 40 (22, 18) 16.7 (14.7, 18.1) 57 (59, 52) 4.3 (7.9, 0.0)
SD MD 0 (15, 15) 4.5 (5.1, 5.8) 6 (14, 7) 5.1 (8.3, 0.0)
EH avg (n = 5) EH 40 (22, 18) 17.0 (16.0, 18.6) 55 (69, 43) 4.9 (9.4, 0.0)
SD EH 0 (15, 15) 3.3 (4.1, 3.9) 10 (21, 6) 4.1 (5.0, 0.0)
2014 Vol. 13, Special Issue 6
leaf out. This increased radiation could result in higher water temperatures and a
reduction in size or even a complete disappearance of these wetlands. A comparison
of populations of Eastern Red-spotted Newts in aquatic habitat within Eastern
Hemlock-dominated and mixed deciduous forests would provide a more detailed
understanding of how the relative abundance of this species may change with the
loss of Eastern Hemlock.
Results from this study suggest that distance to breeding habitat may be the
most important factor in driving differences in the relative abundance of Red Efts.
Caution should be exercised when interpreting these findings, however, because the
distances to breeding habitat were estimations and confirmation of actual breeding
populations of Eastern Red-spotted Newts were not made. Additional research
investigating the relationship between the relative abundance of Red Efts and distance
to breeding habitat is warranted.
Data from this study, collected prior to the arrival of HWA at Harvard Forest, can
be used in before-after analyses to directly monitor potential changes in the relative
abundance of Red Efts in Eastern Hemlock-dominated stands throughout their decline
and transformation into mixed deciduous stands. HWA was first discovered at
Harvard Forest in 2006 near the Eastern Hemlock-dominated stand at Simes 1 (Ellison
et al. 2010). As of 2009, it was present in 44% of the Eastern Hemlock trees in the
two Eastern Hemlock-dominated stands at Simes 1 and Simes 2 (Ellison et al. 2010).
Therefore, it makes sense to repeat sampling of Red Efts as soon as possible.
A long-term study of the relative abundance of Red Efts would be an important
contribution to our understanding of populations of Eastern Red-spotted Newts
because no similar study has ever been conducted. As a long-term ecological research
(LTER) site, Harvard Forest is a perfect place for future studies to build on
the baseline data presented here. The most efficient use of sampling time would be
to conduct sampling within 24 hours of a rain event, if possible (Table 2). When
comparing future data with data from this study, it is important to take into account
average temperature and relative humidity at time of sampling along with total
precipitation prior to sampling (Tables 3, 4).
This study was conducted as part of my thesis research for the Master of Liberal Arts
Degree from Harvard University Extension School, as well as part of my thesis research
for the Masters in Forest Science at Harvard University. I would like to thank A. Benson,
J. Morris, B. Colburn, D. Foster, and S. Mathewson for their guidance and support
throughout this project. In addition, I would like to thank M. Bank and A. Ellison for
their statistical assistance, Jess Butler and S. Jefts for their help in the lab, and A. Barker-
Plotkin for her assistance with the selection of study sites at Harvard Forest. D. Foster,
G. Motzkin, D. Orwig, A. Ellison, A. Barker-Plotkin, M. Bank, J. O'Keefe, B. Colburn,
two anonymous reviewers, and manuscript editor, Jeff Houlahan, all provided extremely
valuable comments on earlier versions of this manuscript. Funds from the National Science
Foundation (DEB-0080592) and the Richard Thornton Fisher Fund for Research at
Harvard University supported this study. This work is a contribution of the Harvard Forest
Long Term Ecological Research Program.
Vol. 13, Special Issue 6
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