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Palatability and feeding preferences of Uresiphita maorialis (Lepidoptera: Crambidae) for three Sophora species

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In a three-hour bioassay, we tested the palatability and feeding preferences of Uresiphita maorialis (kōwhai moth) for Sophora tetraptera, Sophora microphylla and Sophora prostrata. Palatability tests showed no differences among the Sophora species. Feeding preferences, on the other hand, showed that S. tetraptera and S. microphylla leaves are preferred over S. prostrata leaves. Our results support our field observations in Wellington city parks and gardens showing that S. tetraptera and S. microphylla plants frequently have higher densities of larvae than S. prostrata
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Palatability and feeding preferences
of Uresiphita maorialis (Lepidoptera:
Crambidae) for three Sophora species
EA Mundacaa, EC Macayab & HA Vargasc
a Escuela de Agronomía, Facultad de Ciencias Agrarias y
Forestales, Universidad Católica del Maule, Curicó, Chile
b Laboratorio de Estudios Algales ALGALAB, Departamento de
Oceanografía, Facultad de Ciencias Naturales y Oceanográficas,
Universidad de Concepción, Concepción, Chile
c Departamento de Recursos Ambientales, Facultad de Ciencias
Agronómicas, Universidad de Tarapacá, Arica, Chile
Published online: 21 Apr 2015.
To cite this article: EA Mundaca, EC Macaya & HA Vargas (2015): Palatability and feeding
preferences of Uresiphita maorialis (Lepidoptera: Crambidae) for three Sophora species, New
Zealand Journal of Zoology, DOI: 10.1080/03014223.2015.1018281
To link to this article: http://dx.doi.org/10.1080/03014223.2015.1018281
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SHORT COMMUNICATION
Palatability and feeding preferences of Uresiphita maorialis (Lepidoptera:
Crambidae) for three Sophora species
EA Mundaca
a
*, EC Macaya
b
and HA Vargas
c
a
Escuela de Agronomía, Facultad de Ciencias Agrarias y Forestales, Universidad Católica del Maule, Curicó,
Chile;
b
Laboratorio de Estudios Algales ALGALAB, Departamento de Oceanografía, Facultad de Ciencias
Naturales y Oceanográficas, Universidad de Concepción, Concepción, Chile;
c
Departamento de Recursos
Ambientales, Facultad de Ciencias Agronómicas, Universidad de Tarapacá, Arica, Chile
(Received 19 May 2014; accepted 9 February 2015)
In a three-hour bioassay, we tested the palatability and feeding preferences of Uresiphita maorialis
(kōwhai moth) for Sophora tetraptera,Sophora microphylla and Sophora prostrata. Palatability tests
showed no differences among the Sophora species. Feeding preferences, on the other hand, showed that
S. tetraptera and S. microphylla leaves are preferred over S. prostrata leaves. Our results support our
field observations in Wellington city parks and gardens showing that S. tetraptera and S. microphylla
plants frequently have higher densities of larvae than S. prostrata.
Keywords: host plant; multi-choice; New Zealand; Sophora microphylla;Sophora prostrata;Sophora
tetraptera
Introduction
The relationship between moths of the genus
Uresiphita (Hübner) and Sophora (L.) plants has
been widely studied by Leen (1992,1995,1997)
and Conant (1975). Generally speaking, the asso-
ciation of the genus Uresiphita with its host plants
has mostly been attributed to the presence of
quinolizidine alkaloids (Banko et al. 2002) in the
plant tissues (e.g. Nihei et al. 2002). Species such as
Uresiphita reversalis, for example, are known to
sequester quinolizidine alkaloid from its host plants
and use it as a defence as they become aposematic
or toxic to predators (Bernays & Montllor 1989;
Montllor et al. 1990).
In New Zealand, Uresiphita maorialis (Felder
& Rogenhofer) is considered a garden and orna-
mental pest of Sophora (kōwhai) plants (Mulvay
1978; Spiller & Wise 1982;Leen1997). It also feeds
on Lupinus arboreus (lupin) (Molloy et al. 1991)
and it may also feed on Trifolium sp. (clover),
Cytisus scoparius (broom) and Ulex europaeus
(gorse) (Spiller & Wise 1982). Uresiphita maorialis
was previously known as Uresiphita polygonalis
maorialis (Felder), but it is now regarded as an
endemic New Zealand species (Nuss et al. 2003
2013). In many cases, the host plants of the Sophora
species become completely defoliated, though this
generally does not result in the death of the host
plant. Other species of legumes, such as Lupinus
arboreus (lupin), have been reported to be killed by
U. maorialis (Molloy et al. 1991), suggesting that
this moth species can potentially kill its host plants.
Among the Sophora species included in this
study, S. tetraptera has larger leaves than the other
two Sophora species. Due to their lower lignin
content and high water content, larger leaves may
be easier for herbivores to digest (Poorter et al.
2004) and have a longer lifespan (Wright &
Cannon 2001). Based on the high levels of infesta-
tion and defoliation observed on individuals of
*Corresponding author. Email: emundaca@ucm.cl
New Zealand Journal of Zoology, 2015
http://dx.doi.org/10.1080/03014223.2015.1018281
© 2015 The Royal Society of New Zealand
Downloaded by [Universidad Catolica De Maule], [Enrique Mundaca] at 09:06 24 April 2015
S. tetraptera in Wellington parks and gardens, in
addition to its anatomic leaf features, we hypothe-
sised that under laboratory-controlled conditions
S. tetraptera would be the most palatable Sophora
species for U. maorialis. We also hypothesised that
caterpillars would prefer S. tetraptera to S. micro-
phylla and S. prostrata in choice and multi-choice
experiments.
For practical purposes, we defined palatability
as the quality of a plant to be accepted as a feeding
substrate by a caterpillar, and feeding preference
as the caterpillars preference for feeding on one
species of host plant when compared to another.
Methods
We examined leaf palatability of leaves from three
species of Sophora spp: S. tetraptera,S. micro-
phylla and S. prostrata (Fig. 1). Second and third
instar larvae were collected in the field from trees
of S. molloyi. To remove traces of plant material
from their digestive systems, and to reduce the
potential interference from the previous host plant,
larvae were kept in tubes containing a special
multipurpose diet (Singh, 1983) until they reached
later instars (fourth-fifth). The palatability experi-
ment was a no-choice trial, where a single cater-
pillar was individually placed in a 100 × 15 mm
Petri dish with 0.50.8 g of leaves. Thirty-fourth or
fifth instar caterpillars of U. maorialis were used in
each treatment. We used late instar caterpillars as
they move more actively than early instars. In the
field, late instar caterpillars can drop from their
host plants and move to other plants located in the
vicinity (Mundaca E. pers. obs.). The petri dishes
were held out in temperature-controlled environ-
ments at 22 ± 2°C for 3 h with an unvented lid.
Simultaneously, 10 Petri dishes (replicates) of each
treatment were kept without larva as controls to
check for water loss of the plant tissue under those
conditions.
To test feeding preferences we carried out two
separate experiments, where caterpillars were
simultaneously exposed to different feeding sub-
strates. Both experiments utilised the same plants
used for the palatability experiment. We exposed
0.50.8 g of leaves to one caterpillar in an indi-
vidual Petri dish, in paired and multi-choice food
treatments. Thirty-fourth and fifth instar caterpil-
lars were used to carry out both experiments.
For both, palatability and feeding preference
trials, we measured the initial weight of the leaf
material of each species before exposing it to a
caterpillar. After 3 hours, we measured the final
weight of the sample. For each plant species in a
dish, we first recorded the occurrence of consump-
tion as yesor nobecause some larvae did not
consume any leaf tissue during the trials. Weight
measures were carried out using a Mettler Toledo
®
AG204 scale, with 0.00001 g of accuracy.
To correct for leaf weight reduction due to
water loss, we multiplied each final weight meas-
urement observed in the control dish by the
percentage of real consumption. Hence, we calcu-
lated the plant tissue consumed or food consump-
tion through the following formula:
FC ¼ðFw IwÞRC
Where:
FC = Food consumption
Fw = Final plant weight
Iw = Initial plant weight
RC = Real food consumption or frequency of
observed consumption in relation to the total
number of replicates per treatment.
Figure 1 Leaet morphology of the three species of
Sophora employed in this study. ST, S. tetraptera; SM,
S. microphylla;SP,S. prostrata.
2EA Mundaca et al.
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ShapiroWilk normality and homoscedasticity
tests were employed to analyse the datas distribu-
tion for each experiment. Normally distributed data
were analysed using a one-way analysis of vari-
ance for no-choice and multi-choice tests, while
t-tests were used in paired choice tests. Kruskall
Wallis and Wilcoxons independent paired tests
were used to analyse data that were not norm-
ally distributed. A Tukey HSD post hoc test was
carried out to check for variations within the
treatments for the one-way analysis of variance.
The data were analysed using the STATISTICA
7.1
©
statistical package.
Results
We tested palatability by placing larvae in a no-
choice situation where there was only feeding
substrate available. The results of the experiments
showed no differences (F= 0.57, NS, d.f. = 2)
among the consumption options of leaf material,
showing that all treatments were equally accepted
by the larvae. Although many caterpillars did not
feed during the trials, the measured levels of
effective consumption were above 50% in all
treatments (Fig. 2).
The first feeding preferences trial was carried
out through a paired choice experiment (Fig. 3).
The larvae could choose only between two feeding
substrates at the time. Sophora prostrata was the
least preferred feeding substrate when paired with
S. tetraptera (Z= 4.72, P< 0.01) and S. micro-
phylla (Z= 4.782, P< 0.01). The larvae did not
show specific preferences for either the S. tetra-
ptera or S. microphylla substrate (Z=0.4564,
NS). When testing the three species simultaneously
(Fig. 4), we found significant differences in the
consumption levels of leaf material, with S. tetra-
ptera showing higher levels of consumption than
S. microphylla and S. prostrata (H= 58.76, P<
0.01, d.f. = 2). In a number of cases, we observed
that some larvae refused to feed during the trial. We
were not able to explain this particular behaviour,
although, in some cases, larvae have been reported
not to feed for unknown reasons in laboratory trials
(Sagers 1992).
Discussion
In this study, we defined palatabilityas the quality
of the plant material to be accepted as a feeding
substrate by our herbivore. All three species of
Sophora offered in the trials were equally palatable
to U. maorialis. Interestingly, S. prostrata, which is
widely used as an ornamental plant in Wellington
gardens, had never been reported as a host plant.
Although U. maorialis fed on S. prostrata in labor-
atory bioassays, further experiments are necessary
to test the capacity of the larvae to develop into
adult stages and to produce viable progeny when
reared on S. prostrata.
The objective of the choice and multi-choice
experiments was to test a situation commonly found
in gardens and parks where two or more potential
host plant species occur in the same physical space.
The paired choice experiment confirmed our hypo-
thesis that S. tetraptera was the most preferred
feeding substrate. Although we did not analyse the
influence of chemical compounds and leaf anatomy
on the caterpillars feeding choices, our results are
consistent with those observations made in the field.
Hence, in terms of palatability and feeding prefer-
ences, we found that the larvae in captivity have
Figure 2 No-choice experiment. Palatability exhibited by
Uresiphita maorialis larvae for three species of Sophora
plants±SD(n= 30) during the 3-hour trials. The shaded
portion of the circles represents the proportion of dishes of
each plant species in which a caterpillar ate some part
of the leaf. ST, S. tetraptera;SM,S. microphylla;SP,
S. prostrata. Different letters indicate signicant differ-
ences between treatments (P< 0.01).
Palatability and feeding prefferences of the kōwhai moth 3
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the ability to move and choose different feeding
substrates, which seems to suggest that the larvae
may be able to discriminate among different
food substrates. Consequently, last instar larvae of
U. maorialis could, to some extent, shift their food
consumption towards alternative species of host
plants. This could be an important factor driving the
defoliation of the plants observed in parks and
gardens where, contrary to what happens in natural
conditions, planted individuals of different species
of Sophora occur at short distances from each other.
This allows the larvae to move from plants of one
species to another. The importance of this discrim-
inating capacity could be less relevant in natural
environments, however, where Sophora species
tend to occur in non-specific patches. Finally, the
extent to which this larval capacity explains the
defoliation patterns observed in parks and gardens
remains unclear, as, like most Lepidoptera species,
it is the female adult of U. maorialis the one that
chooses the oviposition substrate.
According to what we have observed in the
field, our findings advance our understanding of the
patterns of defoliation observed in the Wellington
city area. The dramatic fluctuations that some popu-
lations of U. maorialis show during the summer
months on individuals of, mainly, S. tetraptera, and,
secondarily, on S. microphylla could indicate, for
example, that U. maorialis has managed to adapt
to a host plant widely available in the Wellington
urban area.
Figure 3 Choice experiment. Feeding preferences exhibited by Uresiphita maorialis larvae when testing the three
species of Sophora ±SD(n= 30) during the 3-hour trials. The shaded portion of the circles represents the
proportion of dishes of each pair of plant species in which a caterpillar ate some part of the leaf. ST, S. tetraptera;
SM, S. microphylla; and SP, S. prostrata. Different letters indicate signicant differences between treatments
(P< 0.01).
Figure 4 Multi-choice experiment. Feeding preferences
exhibited by Uresiphita maorialis larvae when testing the
three species of Sophora ±SD(n= 30) during the 3-hour
trials. The shaded portion of the circles represents the
proportion of dishes of each plant species in which a
caterpillar ate some part of the leaf. ST, S. tetraptera; SM,
S. microphylla;SP,S. prostrata. Different letters indicate
signicant differences between treatments (P< 0.01).
4EA Mundaca et al.
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Acknowledgements
We would like to thank Victoria University of Wellington
for providing the physical space and materials to carry
out this research. We would also like to specially thank
Dr Phil Lester for his guidance and comments that
contributed to improve the quality of this paper. We
would also like to thank the government of Chile and
CONICY for funding the PhD programme of which this
research is part. Finally, thanks to all our enthusiastic
collaborators who lent us their gardens to carry out
observations and tag their plants, and to the field
assistants who helped us to collect larvae for our
experiments: Dr Mariana Lazzaro-Salazar, Jaime Pizarro,
Dr Claudio Sáez and Rodrigo Gomez.
Associate Editor: Dr Rob Cruickshank.
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Summary The spectrum between species with low leaf mass per area (LMA), short-lived leaves and high photosynthetic rate to those with high LMA, long-lived leaves and low photosynthetic rate is one of the major spectra of variation between plant species, and is of particular relevance to the ‘carbon-gain strategy’ of plants. In this study the relationship between physical properties of leaves and their lifespan was quantified for 17 sclerophyllous species from a nutrient-poor woodland in eastern Australia. Fracture properties of leaves (force of fracture, tissue toughness) and other leaf traits [LMA, thickness, dry-matter content (DMC), leaf area] were measured for each species and evaluated as predictors of leaf lifespan in cross-species and phylogenetic analyses, and for intercorrelation with one another. The LMA, mean force of fracture, leaf thickness and leaf area each explained approximately 30–40% of variation in leaf lifespan. Leaf toughness explained 25% of variation in leaf lifespan, and DMC 12%. Leaf toughness and DMC were correlated with each other, but not with leaf thickness. Leaf thickness and toughness were related closely to LMA, while DMC and LMA were only marginally correlated. Nutrients can be withdrawn prior to leaf death and redeployed elsewhere in the canopy when leaf death is initiated by a plant. However, when control is external to the plant these nutrients are lost. There may be advantages to increasing defence to give a high likelihood that the plant has control over the timing of leaf death.
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