SHRE Grant Final Report

1. Project Title: Evaluating Zauschneria (Epilobium) canum cultivars for floral traits
and attraction of beneficials

2. PI: Rachel Vannette, Assistant Professor, University of California Davis,

Department of Entomology and Nematology
Co-PI: David Fujino, Executive Director
California Center for Urban Horticulture

3. Introduction

There is a clear need for horticultural plant species and cultivar types that promote
wildlife health and related ecosystem services, including supporting populations of
pollinators and beneficial organisms. Horticultural plants represent a primary resource
for these organisms in urban and suburban landscapes, providing nutrition, shelter and
other resources for pollinators, predators, and parasitoids. Promoting the cultivation and
sale of plant species and cultivars that support these key goals benefits ecological
communities in these human-dominated landscapes. Additional ecosystem services
associated with these plantings include improving pest management in suburban
landscapes, reducing chemical inputs and conservation of pollinator populations.
Despite the potential for these multiple ecosystem services, current horticultural
offerings are typically bred for showy displays rather than ample floral resources.

Increasingly, it is recognized that native plants can be used in horticulture to provide

multiple positive ecosystem functions. In California, drought-resistant native plant
species can provide many benefits including low water use (Hilaire et al., 2008), the
provision of resources for native animal species, while also proving attractive and robust
to cultivation. However, the propagation and sale of native plants in California in the
horticultural industry remains relatively limited and is often restricted to small-scale
vendors and nurseries. Improving understanding of the benefits of native plant species
and generating scientific data on their attractiveness to wildlife in horticultural plantings
could provide the impetus for larger-scale demand for these plant species or cultivars
and generate greater sales and marketing potential.

Cultivars within in the genus Zauschneria (Epilobium) are a prime candidate for
increased cultivation and marketing. Zauschneria canum is a native California plant
species that is robust, attractive, and provides ample resources for pollinators and other
beneficial organisms during the late season, when other floral resources are often
dwindling. These showy plants frequently produce prolific floral displays and provide
ample nectar and pollen resources for pollinators and other beneficial insect populations
through the late summer and well into autumn. Current sales and distribution of cultivars
within this genus and species are relatively limited, and mainly focus on a few specific
cultivars. Previous trials on Zauschneria canum cultivars conducted by Bart O’Brien
indicate that many cultivars can be reliably propagated and out planted successfully
(O’Brien & Grant 2005, white paper). In this study, recommendations for best cultivars

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were made by expert assessment of attractiveness and form. However, which cultivar(s)
are optimal for pollinator benefit or for attracting parasitoids is currently unknown.
Indeed, in the O’Brien trial, those participating in the scoring agreed nearly unanimously
on foliage, but had widely varying opinion of which cultivar type had the most attractive
flowers.

Cultivars of Zauschneria vary widely in floral form, size and color (Fig 1). Rewards for
pollinators, including nectar and pollen, flowering phenology or floral display, may be
similarly varied, but have not been previously assessed quantitatively. Understanding
which floral traits are attractive to specific pollinator groups and other beneficial insects
is a key goal not only in horticulture, but breeding programs and ecology and evolution
of plant-pollinator systems. In other plant species, previous research documents that
bee pollinators forage based on floral display or number of flowers (Thomson, 1988); in
other cases, the amount of floral reward is correlated with flower size or color, which
increase pollinator foraging (Spaethe et al., 2001). For hummingbirds, artificial flower
studies indicate that larger floral displays and petal size increase visitation rates
(Fenster et al., 2006). However, as floral traits are often correlated and bird response to
artificial flowers may not be reflective of response to flowers in natural settings, further
work is needed. In addition, floral traits that attract non-pollinating organisms, including
predatory insects and parasitoids are poorly understood (Gómez & Zamora, 2006).

4. Materials and Methods

Part 1. Assessment of plant traits and floral visitation using established cultivars

We sampled established Zauschneria canum plants within the UC Davis arboretum

(Figure 1). In the late summer 2017 and 2018, approximately 3 representative plants of
each focal cultivar were tagged and GPS located (plants indicated on the map do not
always survive year to year or may move main location slightly). More or fewer plant
individuals per cultivar were selected based on availability and flowering (e.g. some
plants or even entire cultivars did not produce flowers during the focal years and are not
included in the focal analyses). Chosen plants were monitored weekly for the presence
and the number of flowers produced per plants. The number of open flowers on each
plant was counted. Each plant was monitored for 5 minutes between 10 am and noon,
and all visits to flowers were recorded. Most visits to flowers were performed by honey
bees, although visits by hummingbirds, carpenter bees, and flies were also observed.
To visually estimate indirect evidence of floral visitation, 10 open flowers per plant were
be chosen and features indicating visitation noted, including the presence of pollen on
the stigma and evidence of a slit in the corolla (made by carpenter bees).

Individual flowers were collected from each focal plant and floral traits collected on each
flower. Corolla length was measured from the base of the corolla to the longest petal,
corolla width was measured as the widest distance between petals (Appendix Fig 1).
The number of pollen tetrads (four pollen grains produced by Z. canum) was assessed
using a hand lens. For each flower, the presence of a slit in the corolla, made by
carpenter bees, was also assessed. Finally, floral nectar was collected using 10 µl

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microcapillary tubes and nectar volume assessed. For a subset of nectar samples, the
brix (sucrose equivalents) were measured using a handheld refractometer. In addition,
for a subset of floral samples, digital photographs were taken of individual flowers in a
standardized light environment against a black background. The amount of red, green
and blue in the corolla were assessed in 3 locations per flower using the Apple “Digital
Color Meter” with aperture size set to medium. Color values for R, G, and B were
recorded and averaged for each flower.

For insect collections, a subset of plants were chosen for intensive observation and
collections. Sweep net samples were taken three times during the fall for each plant,
and all insects identified to family, and other arthropods classified to order.

All statistical analysis was performed using R v. 3.5.0 and RStudio Version 1.1.453.
Floral traits were compared among cultivars using ANOVA. To assess the relationship
between floral traits and visitation by different visitors frequently observed
(hummingbirds, carpenter bee (slits), and honey bees), we used linear regression and
plant traits, including plant height, corolla length, corolla width, the presence of a
carpenter bee slit, and total flower number. Regression coefficients were extracted and
plotted with 95% confidence intervals around each coefficient.

Part 2. Installation of two common gardens

In collaboration with the Center for Urban Horticulture and the UC Davis Arboretum, we
established three additional plantings, each containing at least 8 cultivars of Z. canum.
were The additional plantings will allow for consistent evaluation of traits among cultivar
types in a common environment and better assessment of the relative attractiveness of
each cultivar type. Using three common gardens where all cultivars are present will
allow rigorous assessment of the relative attractiveness of cultivars to make
scientifically valid recommendations. In other words, this additional set of plants will
allow us to separate the effect of location (shade, near other attractive plants etc) from
that of the cultivar itself. Plant individuals of each cultivar type have been be acquired
from California flora nursery.

5. Results

Part 1. Assessment of plant traits and floral visitation using established cultivars

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 Existing plants were chosen to maximize
cultivar diversity and span environmental
variation in the UC Davis Arboretum (Fig 1).
We assessed flowering phenology (floral
display) every other week between June and
December, counting open flowers on each
focal plant. Between 2-6 representatives of
each cultivar were chosen, based on
availability, with 84 plants monitored over the
entire season representing 21 cultivar types.

Flowering phenology
Cultivars varied in flowering phenology.
Cultivars that flowered early in the season
included “Cloverdale”, “Marin Pink”, and
“Everett’s choice”, which all had the highest
floral displays relatively early in the season,
whereas “Bowmans’ best”, “Catalina”, “Sierra
Salmon” and “Uvas Canyon” instead flowered
later in the season, with many cultivar types
exhibiting the highest floral display in mid-
Figure 1. Map of marked Zauschneria fall (Fig 2).
canum plants in the UC Davis
Arboretum monitored 2017-2018. Floral traits
Cultivars also varied in all floral traits
measured (Fig 3), including corolla length (P<0.001), corolla width (P<0.001), floral
color (P<0.001), maximum floral density (P<0.001), nectar volume (P<0.001) and sugar
concentration and visitation rates (P<0.001). Visitation frequency also differed among
cultivars after floral display (total number of flowers) was controlled for (P=0.05). The
highest visitation rate overall, largely driven by honey bee visitation, was observed to
“Everett’s choice”, Zauschneria canum ‘canum’, “Brilliant Smith” and “UC Hybrid”.

Floral visitation: direct observations

The majority of observed visits were made by honey bees, with 1446 observations over
the 2035 minutes of observation. We observed 39 visits by hummingbirds and 104 visits
by carpenter bees. We observed the greatest number of visits by honey bees to the
cultivars or varieties: ‘canum’, “Carmen’s gray”, “Cloverdale”, “Mark West” and “Sierra
Salmon”. The most frequent cultivars observed being visited by hummingbirds included
“Catalina”, “Sierra Salmon” and “Carmen’s gray”, although we caution that bird
observations were relative rare. Carpenter bees were frequently observed visiting plants
from the variety “Mark West and also “Sierra Salmon”.

Floral visitation: Indirect measurements of visitation

Hummingbird visitation, quantified by the total number of pollen tetrads deposited on
floral stigmas, was highest on the cultivars “Carmen’s gray”, “el Tigre”, “Catalina”, and

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“UC Hybrid”, but was lowest on “Wayne’s silver”, “Schieffelin’s choice” and “Everett’s
choice” (Fig 3).

Bowman's best Bowman's Hybrid Brilliant smith Calistoga canum

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Carmen's Gray Catalina chaparral silver Cloverdale Dublin

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Marin Pink Mark West Schieffelin's choice Sierra Salmon Summer Snow
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Date (2018)

Figure 2. Flowering phenology of Zauschneria canum by cultivar type. Between 2-6

individual plants were monitored every other week for each cultivar type. Cultivars
“Summer snow” and “John Bixby” did not flower in year 1 and showed low survivorship
in the Arboretum conditions at both transplant sites. Colored lines indicate the line of
best fit and gray confidence intervals indicate the 95% CI around the estimated
smoothing line. Floral counts were assessed in 2018 starting in June 2018 and ending
in December.

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Insect collections
Insects from the families Apidae (including honey bees), Vespidae (wasps), Berytidae
(stilt bugs), Aphididae (aphids) and Formicidae (ants) were the most commonly
encountered insects. Notably, cultivars including “Cloverdale” and “Wayne’s silver”
hosted fewer insects (total number) and diversity (Shannon-Weiner index) than did
other cultivars, with an average of 5.5 insects/sweep +/-3.4 for “Cloverdale” and 6.5
insects/sweep +/- 5.3 for “Wayne’s silver” compared to 40.66 insects/sweep +/- 12.7 for
the most attractive cultivar. Differences in total insect numbers were driven mainly by
the lack of any insects on Wayne’s silver and Cloverdale save ants, stiltbugs and
aphids.

Relationship between plant traits and floral visitation patterns

Visitation to plants was significantly predicted by plant-level variables, however, the
plant traits associated with visitation differed depending on the identity of floral visitor.
Honey bee visitation was most strongly associated with the floral display, where plants
with large numbers of flowers attracted the most honey bee visitation (Fig 4a). In
addition, plants with wide flowers were also visited at a higher rate than plants with
narrow flower corollas, and robbing by carpenter bees, indicated by a high proportion of
carpenter bee activity, also increased visitation by honey bees. For carpenter bees,
plants with longer and wider corollas showed increased frequency of robbing, and
plants with a larger floral display also had a higher proportion of flowers robbed than
flowers with a small floral display. For hummingbirds, plant height and corolla length
were the two strongest predictors of pollen deposition (Fig 4b). Tall plants and those
with narrow flowers had the highest average pollen loads (Fig 5b).

Part 2. Installation of two common gardens

Garden installation was performed at in gateway garden in the UC Davis arboretum and
near the Environmental Horticulture building. Establishment success was initially good
after outplanting but survival to flowering was mixed and varied among cultivars
(Appendix Table 1). Floral sampling was conducted on plants that survived to flowering
but some varieties did not survive well or flower at all at any location, including “Summer
snow” and “John Bixby” ; “Liz’ choice” also showed poor survival so these could not be
included in most of the analyses below. However, establishment of these gardens will
aid future research efforts to compare plant phenotypes, floral traits, or relative
attractiveness among cultivars. In addition, our efforts here have given us experience
with establishing Z. canum and can help with future experiments and plant
establishment.

7
 Coefficient Plot Coefficient Plot

A) B)
scale(Proportion_CarpSlit) scale(Proportion_CarpSlit)

scale(Corolla_width_avg) scale(Corolla_width_avg)
Coefficient

Coefficient
Model
scale(Corolla_length_avg) Honeybee scale(Corolla_length_avg)
Carpenter bee

scale(log10(Total_flowers + 1)) scale(log10(Total_flowers + 1))

scale(Plant_height) scale(Plant_height)

-0.05 0.00 0.05 0.10 0 2 4

Value Value

Figure 4. Regression coefficients comparing visitation between plant-level variables of

Zauschneria canum and A) visitation by honey bees or carpenter bees, and B) visitation
by hummingbirds, measured by pollen deposition on the stigma surface. Plant-level
variables include the proportion of flowers with a carpenter bee slits, the average corolla
width, average corolla length, the total flowers (floral display), and plant height. Values
above 1 indicate that a greater value of that trait increases visitation. If 95% CI do not
include zero (dotted line), the relationship between the trait value and visitation is
statistically significant.

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Figure 5. Relationship between plant traits and floral visitation. In a), honey bee visits
increase with the total number of flowers per plant. In b) pollen on the stigma, a
measure of hummingbird visitation, is positively related to plant height. Points indicate
individual plants, and lines indicate the line of best fit (linear model).

8
Table 1. Plant and floral characteristics of Z. canum varieties assessed in Part 1 of the
current study and relative value for pollinators including hummingbirds, honey bees and
other beneficial insects.
Average plant Average flower corolla Average flower corolla Total floral display Flowering Humming Honey Non-pollinator
Variety height (m) length (mm) width (mm) (average) time birds bees beneficial insects
Bowman's
best 0.830946154 30.05238095 11.33571429 151.1764706 late +++ +++ +++
Brilliant
smith 0.645571429 28.06578947 13.72105263 23.42105263 mid +++ + +

canum 0.468857143 27.52916667 11.59166667 106.3333333 mid ++ ++ ++

Carmen's
gray 0.8675 41.75 16.175 475.5 mid-late +++ +++ +++

Catalina 0.742727273 33.54 13.23 49.83333333 late +++ + +

chaparral
silver 0.529666667 42.71666667 14.96666667 2.75 mid ++ + +

Cloverdale 0.343076923 38.73125 18.55625 50.55555556 early + ++ +

Dublin 0.646375 31.065 16.82 59.22222222 late +++ ++ ++

el Tigre 0.6193375 29.36333333 11.22333333 55.16666667 late +++ ++ +

Everett's
choice 0.406 24.3 8.45 66.14285714 early + ++ ++
Hurricane
point 0.6075 28.28 14.04 97.6 late +++ ++ ++

Marin Pink 0.515 NA NA 41.33333333 early ++ + +

Mark West 0.604428571 35.61666667 16.95545455 260.4444444 late ++ +++ +++

Schieffelin's
choice 0.478333333 19.80571429 8.994285714 148.75 mid ++ +++ +++
Sierra
Salmon 0.38 36 17.6 167 mid-late + +++ +++

UC Hybrid 0.479 32.11428571 13.54285714 76.28571429 mid-late ++ ++ ++

Uvas Canyon 0.501322222 39.63636364 19 43.25 mid-late ++ + +

Wayne's througho
silver 0.2775 28.03333333 14.46666667 22.16666667 ut + + +

9
 6. Discussion

The plant species Zauschneria canum (Onagraceae) is endemic to California, produces

abundant red tubular flowers. This plant species is a valuable food resource for many
organisms, including hummingbirds, bees, and other arthropod species that rely on
nectar and floral resources, particularly late in the season. Our work on Z. canum
cultivars in this project has revealed that horticultural varieties vary widely in floral traits,
including in floral morphology, standing nectar crop, and in attractiveness to insects. We
acknowledge that the data gathered in this survey may not represent an unbiased
comparison among cultivar types, as not all cultivars were represented in each
environment. Further work will be required to compare floral traits using plant species
growing in the common garden environments established in Part 2 above.
Nevertheless, we observed abundant visitation by hummingbirds and honey bees
throughout the arboretum, and cultivars were represented often in multiple locations
throughout the arboretum, so our results are unlikely to be explained wholly by spatial
autocorrelation.

Our results allow us to suggest specific plant traits in Zauschneria canum that are
particularly attractive to specific visitor taxa. For example, plants that had many flowers
(large floral display) were more attractive to honey bees. In contrast, tall plants with a
narrow corolla were more attractive to hummingbird pollinators. We suggest that these
traits could be targets for increasing hummingbird attraction and varieties with these
traits may be particularly useful to hummingbirds. In contrast, flowers with wide corollas
were found to be accessible and attractive to honey bees. We outline these differences
in Table 1 and provide recommendations on their relative attractiveness to
hummingbirds, honey bees and non-pollinator beneficial insects based on the data from
our regression analysis and other sampling data.

In addition, Z. canum was found to attract a number of insect families, including

pollinators (Apidae) and some pests (Aphidae) but also a number of natural enemies
including wasps (Vespidae), ants, and other predatory insects including Hemipterans in
the families Reduviidae and Berytidae (stilt bugs). Notably, insects from the Berytidae
were commonly observed feeding on flowers particularly as nymphs, but are known to
be omnivorous. In particular, stiltbugs are often found on “glandular-hairy” plants, where
they tend to feed on small insects including aphids and insect eggs (Wheeler and
Schaefer 1982). These insects were not detected on neighboring plant species (visual
inspection, pers. obs), suggesting that the hairy leaves of Z. canum can support this
omnivorous and potentially important predator. Future work should directly quantify
‘hairiness’ of Z. canum leaves and examine this relationship with stilt bug abundance
and effects on aphids.

Future work
The common gardens established will enhance future work comparing the relative
benefit of Z. canum cultivars on pollinators and non-pollinator arthropods. Future

10
classes including ABI50A and a pollination ecology class will likely make use of such
plantings in future years and research in the Vannette lab will continue on such topics.
We plan to submit this work for publication after we have finished our data analysis from
this past season

We also look forward to investigating other factors that may influence floral traits in Z.
canum, and benefits to beneficial insects and animals. This year, the fall rains, which
typically arrive in October, was very late, only beginning in late November. We propose
that investigating the effects of deficit irrigation and drought conditions on Z. canum, and
if cultivars are differentially resilient to such conditions, will be important for enhancing
resilience to changing climate conditions in California, particularly for pollinator and
insect populations.

Outreach and Impacts

The current grant resulted in a number of products and outreach activities. First, we
established three new common garden areas on campus. Although establishment
success was variable among cultivars, these efforts have increased the representation
of novel cultivar types in the UC Davis Arboretum in high-traffic, high-visibility locations.
Second, we have increased the visibility of Z. canum through the installation of signage
on campus summarizing our research efforts, working with arboretum staff to increase
planting, and outreach efforts, including giving talks to the Western Apiculture Society
and to the Master Gardeners of Solano and Yolo County highlighting our work on
Zauschneria canum. The research associated with this project also engaged over 140
students in the ABI 50A class on the UC Davis campus, who participated in research
related to this project, monitored plants and measured floral traits. Many of these
students have gone on to pursue further research or outreach opportunities as a result
of this class project. Details of each activity are below:

Outreach activities and other products:

• Vannette, RL, The role of plant traits and microorganisms on pollinator
attraction. Pollinator Workshop, “Bee-ing a better bee gardener”, Sept 23, 2017.
(~90 in attendance), Master Gardeners of Solano and Yolo County.
• Vannette, RL, Changes in nectar that affect bee foraging: a role for floral
microbes? Western Apiculture Society meeting; Davis, CA Sept 8, 2017. (~40 in
attendance).
• Provided input on additional pollinator plantings in the gateway garden in the UC
Davis arboretum.
• Produced signage on research activities related to Zauschneria canum in high-
traffic areas throughout the UC Davis arboretum.
• Engaged ~140 students in ABI 50A on projects related to horticultural plants and
plant-insect interactions.

11
 Appendix

A) B) C)

yes

Appendix Figure 1. Floral traits measured using Zauschneria canum flowers. In A) the
length of the corolla was measured; B) corolla width was measured and C) the
presence of a slit made by a carpenter bee (both flowers contain a slit) was assessed
on all collected flowers, with N=2134 flowers measured in total.

Appendix Table 1. Cultivars established in common gardens and establishment

success
Cultivar Establishment Notes
success
Liz’s choice 33% Poor survival
Bowman’s 100% Good survival,
hybrid good flowering

Catalina 66% Inconsistent

Calistoga 100% Good survival
but some
plants had
large sections
of dead
material
Marin Pink 100% Good survival
but poor
flowering early
on.
Cloverdale 67% Mixed survival
but good
flowering
Schieffelin’s 100% Good survival
choice and flowering

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John Bixby 16% High mortality
early after
planting
Summer snow 0% High mortality
early after
planting
Wayne’s silver 100% All plants
survived,
although some
appeared in
poor condition

13