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Community Impacts of Sudden Oak Death

Jessica Elgersma, Senior Thesis


Plant disease epidemics can radically impact the productivity of communities with high infection
and mortality rates, altering ecosystem balance. The disease Sudden Oak Death (SOD), caused
by the oocyte Phytophthora ramorum has spread rapidly and caused extensive mortality
throughout hardwood forests of California (Cobb et al., 2012). Notholithocarpus densiflorus is
particularly susceptible, and both understory seedlings and mature trees may be infected and

Infection and mortality of entire stands of N. densiflorus by P. ramorum has radically impacted
the physical structure and species composition of California forests, with the potential to cause a
cascade of long-term negative effects on coastal forests (Rizzo and Garbelotto, 2003). N.
densiflorus is a dominant hardwood species and a prolific acorn bearer in redwood and mixed
evergreen stands, with an average annual acorn production for a mature tree ranging from 35-
1000 lbs (Fryer, 2008). Over 99% of N. densiflorus acorns are consumed by insects, birds or
rodents (Fryer, 2008), so loss of N. densiflorus may represent a significant loss of food resources.

The Forest Ecology Research Plot (FERP) is in a coastal mixed evergreen forest on UCSC’s
Natural Reserves. N. densiflorus is the second most common tree species on the FERP, despite
many individuals being lost to SOD mortality in recent years. Two species of deer mice are also
found on the FERP, Peromyscus boylii and Peromyscus californicus. They are trapped and
tagged quarterly by the Small Mammal Undergraduate Research Foundation (SMURF) and
individuals encountered are recorded in a database. Approximately 16% of P. californicus’ diet,
and 51% of P. boylii’s diet is made up of N. densiflorus acorns (Reid et al., 2013). Since small
mammal distributions vary in response to seed abundance (Schnurr et al., 2004), I hypothesize
that P. boylii will be more strongly affected than P. californicus by the declines in seed
Materials and Methods

The study site was in the Coastal Mixed Evergreen Forest on the University of California at
Santa Cruz’s Forest Ecology Research Plot (FERP). Tan Oak is the dominant hardwood
understory tree on the UC Natural Reserves in the Santa Cruz mountains, at approximately 300m
elevation. This study sampled the original 6 hectares of the FERP, established in 2007 and
containing over 1700 tagged Tan Oak individuals over 1 cm in diameter at breast height.
To assess the spread of Sudden Oak Death (SOD), tagged individuals were located and rated on
presentation of Sudden Oak Death symptoms on a scale of 0-5, and 99, with ratings interpreted
as follows:
• 0 = No indication of SOD, plant healthy
• 1= An individual leaf or two dead (somewhat suspicious)
• 2=A complete branch with dead leaves (suspicious, but may be other causes)
• 3=Several branches with dead leaves (very suspicious)
• 4=Much of tree with classic SOD symptoms of dead leaves, often with cankers on
the stem.
• 5 =All leaves on tree dead, often with cankers on stem, classic SOD
• 99=Dead for some other reason

This qualitative classification system was identical to that used in 2007 and 2012 assessments of
Sudden Oak Death on the FERP, so that data could be compared between years.

Using R, we created a map of Tan Oaks on the FERP, highlighting individuals displaying classic
symptoms of SOD infection or mortality (Tan Oaks with rating of four or above). This map was
compared to the corresponding maps created for 2007 and 2012 to determine which regions on
the FERP are most impacted by Sudden Oak Death.

The Small Mammal Undergraduate Research Foundation (SMURF) database was also analyzed,
and locations of recorded captures of Peromyscus species during 2012 and 2017 were overlaid
on the map of Sudden Oak Death, to create a visual representation of the correlation between
small mammal relative abundance and distribution and Sudden Oak Death infection and
We graphed a scatter plot of encounter probability (defined as the number of Peromyscus caught
per trapping event, divided by the number of total traps set), using data points collected during
quarterly trapping events beginning in 2009. We also created maps of Peromyscus encounters in
traps, and compared these with encounter probability to analyze trends in Peromyscus relative
abundance and distribution on the FERP.


In 2007, SOD was present on the Northern edge of the FERP (Figure 1). Less than a hundred
trees were affected, few individuals had died and the infection was mostly clustered above 280
meters North on the FERP.

By 2008 more trees were infected than uninfected on the FERP’s Northern edge (Figure 2).
There were only 4 tree deaths between 2007 and 2008, but SOD had spread significantly and
extended southward. At present, SOD is ubiquitous throughout the FERP and has created entire
groves of standing dead or dying tan oak (Figure 4).

From 2009-2016, an average of 61 (S.D. = 23) unique individuals of Peromyscus californicus

and 95 (S.D. = 23) unique Peromyscus boylii individuals were encountered annually in SMURF
traps (Table 1). The difference in mean encounters between Peromyscus californicus and
Peromyscus boylii was statistically significant (95% confidence interval, p = 0.01082).

Apart from 2015, more unique Peromyscus boylii were encountered between 2009 and 2016 than
Peromyscus californicus. However, the number of encounters of Peromyscus californicus in
SMURF trapping events had a strong positive trend, more than doubling from 2009-2016. In
contrast, Peromyscus boylii has displayed a negative population trend, decreasing every year
from 2010-2015, although P. boylii encounters dramatically increased in 2016.
Figures and Tables

Figure 1 – SOD occurrence on the FERP, 2007 (Gilbert, 2017)

Figure 2 – SOD occurrence on the FERP, 2008 (Gilbert, 2017)
Figure 3 – Tan Oak Distribution and Sudden Oak Death incidence on the FERP
Unique Peromyscus Captures







2008 2009 2010 2011 2012 2013 2014 2015 2016 2017

Peromyscus boylii Peromyscus californicus

Figure 4 – Total unique Peromyscus individuals caught in SMURF trapping events, 2009-2016
(Cheng, 2017)
Figure 5 - Peromyscus distributions on the FERP for 2009 & 2016 mapped by coordinates (Cheng, 2017)
Year Peromyscus Peromyscus
boylii californicus
2009 94 38

2010 129 48

2011 103 40

2012 90 69

2013 85 60

2014 73 54

2015 63 73

2016 124 109

Table 1 – Unique Peromyscus encounters by year (Cheng, 2017)


Spatial mapping of Sudden Oak Death (SOD) occurrence on the FERP shows substantial
infection and mortality on the northern edge of the plot by 2008. Infections tended to occur in
clusters, but frequently did not correlate with centers of highest Tan Oak density.

There were significantly more unique P. boylii individuals caught in SMURF trapping events
from 2009-2016 than there were P. californicus. However, P. boylii showed a strong negative
trend in unique captures from 2009 to 2015, in contrast with the positive trend demonstrated by
P. californicus. The decline in P. boylii cannot be solely attributed to the drought, as P.
californicus was increasing during the same time. The results support the hypothesis that SOD
has a greater negative effect on P. boylii than P. californicus, but does not explain the P.
californicus population increase.

Additionally, P. boylii encounter frequency nearly doubled from 2015 to 2016. This may have
been the effect of abiotic factors such as increased primary production due to abundant winter
rains, but more long-term monitoring will be needed to determine if this represents a reversal of
the previous trend or a stochastic event.

Mapping coordinates where individuals were caught in SMURF trapping events showed that,
between 2009 and 2016, P. californicus captures on the Northern edge of the plot tripled. This is
the location shown by vegetation surveys to have been the initial epicenter of the SOD epidemic,
so it is possible that P. californicus is able to take advantage in the decline of the tan oak acorns
to expand into habitat where they had previously been excluded by competition with P. boylii. If
this is the case, SOD mortality may result in shifts in species composition of other mixed oak
woodlands in California.

Cheng, Tina. (2017) Personal Communication

Cobb, Richard C. et al. (2012) “Common Factors Drive Disease and Coarse Woody Debris
Dynamics in Forests Impacted by Sudden Oak Death.” Ecosystems 15(2): 242–255.

Davidson, J. M., et al. (2003) "Sudden oak death and associated diseases caused by
Phytophthora ramorum." Plant Health Progress 7.

Gilbert, Gregory. (2017) Personal communication

Leis, Sherry A., et al. (2008) "Small mammals as indicators of short-term and long-term
disturbance in mixed prairie." Environmental Monitoring and Assessment 137(1-3): 75-84.

Reid, Rachel EB, et al. (2013) "Dietary niche partitioning by sympatric Peromyscus boylii and P.
californicus in a mixed evergreen forest." Journal of Mammalogy 94(6): 1248-1257.

Sollmann, Rahel, et al. (2015) "Investigating the effects of forest structure on the small mammal
community in frequent-fire coniferous forests using capture-recapture models for stratified
populations." Mammalian Biology 80(4): 247-254.

Schnurr, Jaclyn L., et al. (2004) “Neighborhood analyses of small-mammal dynamics: impacts
on seed predation and seedling establishment." Ecology 85(3): 741-755.