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Susceptibility of
Persea
spp. and Other Lauraceae to Attack
by Redbay Ambrosia Beetle,
Xyleborus glabratus
(Coleoptera:
Curculionidae: Scolytinae)
Author(s): J. E. Peña, D. Carrillo R. E. Duncan, J. L. Capinera, G. Brar, S. McLean,
M. L. Arpaia, E. Focht, J. A. Smith, M. Hughes and P. E. Kendra
Source: Florida Entomologist, 95(3):783-787. 2012.
Published By: Florida Entomological Society
URL: http://www.bioone.org/doi/full/10.1653/024.095.0334
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Scientific Notez 783
SUSCEPTIBILITY OF PERSEA SPP. AND OTHER LAURACEAE TO
ATTACK BY REDBAY AMBROSIA BEETLE, XYLEBORUS GLABRATUS
(COLEOPTERA: CURCULIONIDAE: SCOLYTINAE)
J. E. PEÑA
1,*
, D. CARRILLO
1
, R. E. DUNCAN
1
, J. L. CAPINERA
2
, G. BRAR
2
, S. MCLEAN
2
, M. L. ARPAIA
3
, E. FOCHT
3
,
J. A. SMITH
4
, M. HUGHES
5
AND P. E. KENDRA
6
1
University of Florida, Tropical Research and Education Center, Homestead, FL 33031
2
University of Florida, Department of Entomology and Nematology, Gainesville, FL 32611
3
University of California, Department of Botany ad Plant Sciences, Riverside, CA 92521
4
University of Florida, School of Forest Resources and Conservation, Gainesville, FL 32611
5
University of Florida, Department of Plant Pathology, Gainesville, FL 32611
6
USDA-ARS, Subtropical Horticulture Research Station, Miami, FL 33158
*Corresponding author; E-mail: jepe@ifas.ufl.edu
Redbay ambrosia beetle (RAB), Xyleborus gla-
bratus Eichhoff (Coleoptera: Curculionidae: Sco-
lytinae), a native of Asia, was first discovered in
the USA near Savannah, Georgia in 2002 (Haack
2001; Rabaglia et al. 2006). RAB is an effective
vector of Raffaelea lauricola T.C. Harr., Fraedrich
& Aghayeva (Harrington et al. 2008) that causes
laurel wilt (LW), a lethal disease of several trees
in the Lauraceae in the southeastern USA (Crane
et al. 2008; Mayfield et al. 2008). Ambrosia beetle
adults bore through the bark and into the xy-
lem (wood) where they lay eggs, then adults and
larvae cultivate and feed on symbiotic ambrosia
fungi that grow in the galleries. Native Persea
(Laurales: Lauraceae) species appear to be pre-
ferred hosts. LW is responsible for high mortality
of redbay [P. borbonia (L.) Spreng.], swampbay
[P. palustris (Raf.) Sarg.], and sassafras [Sassa-
fras albidum (Nuttall) Nees] in Alabama, Flor-
ida, Georgia, Mississippi, North Carolina and
South Carolina (Fraedrich et al. 2008; Hanula
et al. 2008; Gramling 2010). As LW encroaches
upon the Lake Wales Ridge ecosystem in south-
central Florida, silkbay (P. humilis Nash) is also
showing susceptibility to LW and is dying. Addi-
tional species affected by LW include avocado (P.
americana Mill.), spicebush [Lindera benzoin (L.)
Blume], and other woody Lauraceae (Fraedrich et
al. 2008) (Table 1).
The susceptibility of 5 avocado cultivars of
Mexican, Guatemalan or West Indian origin to
RAB and LW was demonstrated by Mayfield et
al. (2008). However, with more than 23 West In-
dian cultivars grown in Florida, it is necessary
to determine their susceptibility. Moreover, as an
adventive species to the North American conti-
nent, RAB might affect other valuable New World
species. Most Persea species are of Mexican, Cen-
tral American, or South American origin. These
Persea may have significant value in germplasm
collections, some have been discovered recently,
and some have resistance to diseases that afflict
their commercial relative, the avocado (Skutch
et al. 1992; Scora & Bergh 1992; Zentmyer &
Schieber 1992). Thus, their susceptibility to RAB
and LW warrants evaluation. Another member of
the Lauraceae of much concern is the California
bay laurel [Umbellularia californica (Hook. &
Arn.) Nutt.], a dominant hardwood species of the
U.S. Pacific Coast. Through inoculation experi-
ments, Fraedrich (2008) demonstrated that U.
californica is susceptible to LW. With continued
westward spread of LW, the host status of U. cali-
fornica needs to be confirmed.
The 3 studies presented here evaluate sus-
ceptibility to RAB and LW in: 1) 13 West Indian
avocado cultivars; 2) 10 non-commercial Persea
spp., 1 Beilschmidia sp. (a genus related to Per-
sea), and 3) U. californica. First, no-choice ex-
periments were conducted to determine if RAB
would bore into avocado cultivars not screened
previously by Mayfield et al. (2008) and follow-
ing similar methodology. Avocado cultivars ‘Ber-
necker’, ‘Beta’, ‘Brooks late’, ‘Choquette’, ‘Don-
nie’, ’Dupuis’, ‘Hall’, ‘Loretta’, ‘Lula’, ‘Monroe’,
‘Simmonds’, ‘Tower 2’ and ‘Waldin’ (4 plants each)
were planted in 10-gallon pots in a screenhouse at
the Plant Sciences Research and Education Unit,
University of Florida (UF), Citra, Florida in VI-
2008. Two plants of each cultivar were infested
by enclosing 4 newly emerged RAB (UF colony-
reared) within a mesh sleeve on the lower trunk.
Two plants per cultivar were uninfested controls.
Entrance holes and perseitol (white exudate from
wounds) were monitored for 4 wk. Severity of wilt
symptoms was scored using the following LW in-
dex: 0 = no wilt; 1 = wilt, no leaf necrosis; 2 = wilt,
10% necrosis or defoliation; 3 = wilt, 30% necrosis/
defoliation; 4 = 50% necrosis/defoliation; 5 = 75%
necrosis/defoliation; 6 = 100% necrosis/defoliation
784 Florida Entomologist (95)3 September 2012
TABLE 1. LIST OF REPORTED AND POTENTIAL HOSTS OF XYLEBORUS GLABRATUS.
Species name Provenance
Evidence of
Reproduction
by RAB
Evidence of
infection by
R. lauricola
Evidence of
boring by
RAB Reference
Beilschmidia sp. * reported here
Cinnamomum camphora (L.) J. Presl. Taiwan, China, Japan * Smith et al .2009
Cinnamomum jensenianum Hand.-Mazz. China * Fraedrich (ppt)
Laurus noblis L. Southern Europe * Fraedrich (ppt)
Lindera benzoin (L.) Blume Eastern USA * * Fraedrich et al. 2008
Lindera latifolia Hk.f Wood and Bright 1992
Lindera melissifolia (Walter) Blume * Fraedrich et al. 2008
Lindera strychnifolia (Sims) Kosterm China, Japan * Fraedrich (ppt)
Litsea aestivalis
* * * -Hughes et al. 2011
Litsea elongata (Nees) Hk.f Wood and Bright 1992
Machilus thunbergii Siebold & Zucc. Taiwan, China, Japan * Fraedrich (ppt)
Ocotea coriacea (Sw.) Britton * Fraedrich (ppt)
Persea americanaMills. cv. ‘Hass’ Mexico, Central America * * Mayfield et al. 2008
P. americanaMills. cv. ‘Simmonds’ * * Mayfield et al. 2008
P. americanaMills. cv. ‘Monroe’ * * Mayfield et al. 2008
P. americanaMills. cv. ‘Winter Mexican’ * * Mayfield et al. 2008
P. americanaMills. cv. ‘Catalina’ * * Mayfield et al. 2008
P. americanaMills. cv. ‘Bernecker’ * reported here
P. americanaMills. cv. ‘Beta’ * reported here
P. americanaMills. cv. ‘Brookslate’ * reported here
P. americanaMills. cv. ‘Choquette’ * reported here
P. americanaMills. cv. ‘Donnie’ * reported here
P. americanaMills. cv. ‘Dupuis’ * reported here
P. americanaMills. cv. ‘Hall’ * reported here
P. americanaMills. cv. ‘Loretta’ * reported here
P. americanaMills. cv. ‘Lula’ * reported here
P. americanaMills. cv. ‘Tower 2’ * reported here
P. americanaMills. cv. ‘Waldin’ * reported here
Persea borbonia (L.) Spreng. Southern USA * * * Fraedrich et al. 2008, Hanula et al. 2008
Persea humilis Nash Southern USA, only Florida? * * Fraedrich et al. 2008, Hanula et al. 2009
Persea palustris (Raf.) Sarg. Eastern USA * * * Fraedrich et al. 2008
Persea caerula (Ruiz and Pav.) Mez Peru, Amazon * * reported here
Persea pachypoda Ehrenb Mexico, California, South America * * reported here
Persea floccosa Mez.,
* reported here
Scientific Notez 785
(Peña et al. 2011). After 2 wk, all cultivars had 1-2
entrance holes (df
8,36
; F = 2.18; Pr > F = 0.052) (Ta-
ble 2). This is an important indication of success-
ful beetle boring. The LW index assessed during
the last wk of the experiment fluctuated between
0 - 1.8 (df
8,38
; F = 1.85; Pr > F = 0.10), which cor-
responded to an average between wilt only and
10% leaf necrosis (Table 2). On 22-VIII-2008 all
plants were harvested, and wood chips were col-
lected, surface sterilized, and plated on medium
selective for Raffaelea lauricola (Mayfield et al.
2008). Isolation frequency ranged from 0-50% for
the different cultivars (Table 2). These results are
in in agreement with the results of Mayfield et al
(2008) who reported that when given no choice,
RAB can bore into and transmit the pathogen R.
lauricola into the xylem of avocado, which char-
acteristically presents as dark discoloration of the
outer sapwood
In the second study, RAB attraction to 10 Per-
sea spp. and 1 Beilschmidia sp.was tested in the
field and laboratory. Four replicate logs (33 cm
long × ~2.5 cm diam.) of P. caerulea Ruiz and
Pav., P. borbonia, P. pachypoda Ehrenb [syn: Cin-
namomum pachypodum (Nees) Kosterm., Phoe-
be pachypoda (Nees) Mez], P. floccosa Mez., P.
skutchii C. K. Allen, P. nubigiena L. O. Williams,
P. indica (L.) Spreng., P. tolimanensis Zentmyer
& Schrieber (also called “aguacate de mico”, a
Central American species), P. cinerascens S. F.
Blake, P. tilarensis and Beilschmidia were hung
in full sun ~1.2 m high near infested redbay trees
with an approx. distance of 10 m between adja-
cent treatments in Hastings, Florida from IX to
X-2009. After 30 d, the logs were collected and the
number of entry holes determined under a ste-
reomicroscope. Logs were placed individually in
cardboard containers for beetle emergence at 26
°C and 70-80% RH for 60 d. Bolts of P. skutchii,
P. cinerascens and P. indica appeared to be pre-
ferred by ambrosia beetles over other Persea spp.,
including P. borbonia (Table 3). Unfortunately, no
beetle emerged from these bolts, perhaps due to
desiccation of bolts under field or storage condi-
tions.
Another survey was set up at Hickory Ham-
mock, a 4,000-acre (1,619 ha) natural preserve in
Highlands County, Florida (27°25’35”N, 81°9’42”
W). This site was known since 2009 to have LW
and RAB. Bolts (same species as above) were
hung on 23-II-2010 on the sunny edge of a trail,
removed 30 d later, and stored as described above.
With the exception of P. floccosa, bolts of all spe-
cies had entrance holes (df
10,10
; F = 2.04; Pr > F =
0.13). RAB emerged from P. caerula and P. tilar-
ensi; Ambrosiodmus lecontei Hopkins (Curculion-
idae: Scolytinae) from P. nubigena, P. pachypoda
and P. tilarensis; and Xylosandrus crassiusculus
(Motschulsky) (Curculionidae: Scolytinae) (df
10,10
;
F = 1.61; Pr > F = 0.23) (Table 3). No beetle
emerged from other Persea spp. However, lack
TABLE 1. (CONTINUED) LIST OF REPORTED AND POTENTIAL HOSTS OF XYLEBORUS GLABRATUS.
Species name Provenance
Evidence of
Reproduction
by RAB
Evidence of
infection by
R. lauricola
Evidence of
boring by
RAB Reference
Persea skutchii L. O. Williams * reported here
Persea nubigiena L. O. Williams * * reported here
Persea indica Zentmyer & Schrieber * reported here
Persea tolimanensis Zentmyer & Schrieber * reported here
Persea cinerascens
* reported here
Persea tilarensis
* * reported here
Persea liebmanni (= P. podedemia)
Mexico, Guatemala * Fraedrich (ppt)
Sassafras albidum (Nuttall) Nees Central and Eastern USA *
* * Fraedrich et al. 2008
Umbellularia californica
* * Fraedrich 2008; ?reported here
786 Florida Entomologist (95)3 September 2012
of emergence from those species could be due to
desiccation and not the result of plant resistance.
On 24-II-2010, host boring bioassays were set
up in the laboratory, using methods similar to
Kendra et al. (2011). Bolts (10.9 × 2.7 cm diam) of
the same species above (2 replicates per species)
were cut and immediately placed individually
in glass jars (0.95 liter) with 200 mL of water to
prevent desiccation. Five newly emerged RAB
were placed on top of each bolt and kept for 24
h at 22 ± 2 °C and 12:12 h L:D. RAB boring was
recorded at 1, 2, 3, 4 and 24 h. RAB bored into all
species except P. floccosa, and infestation varied
from 1.5 to 4 beetles boring per bolt (Table 3) (
df,
10, 11
; F = 3.79, Pr > F = 0.02) (GLM procedure,
Tukey’s Studentized Range (HSD) Test (SAS,
2008). P. floccosa is a Guatemalan-type species,
which is believed to be the most ancient form of
Persea (Scora & Bergh 1992).
In the third experiment, bolts of redbay, avoca-
do and California bay laurel were hung at Ordway-
Swisher Biological Station, University of Florida,
Gainesville (N 29° 41.040, W 082° 22.109). Nine
logs of each species were hung in an area where
both diseased and healthy red bay were present,
and left in the field for 1 mo (18 IX-19-X-2009).
Logs were brought into the laboratory, bore holes
were measured, and those of appropriate diam-
TABLE 2. MEAN INFESTATION BY XYLEBORUS GLABRATUS AND SEVERITY OF LAUREL WILT OBSERVED AFTER 4 WK IN NO-CHOICE TESTS
WITH 13 WEST INDIAN AVOCADO CULTIVARS. YOUNG POTTED TREES (2 PER CULTIVAR) WERE EXPOSED TO 4 X. GLABRATUS
FEMALES ENCLOSED IN A SLEEVE AT THE BASE OF THE TRUNK.
Avocado Cultivar
Number of
holes/plant ± SE
Holes with
Perseitol/plant ± SE
LW Severity Index
R. lauricola
isolation frequency
Bernecker 1.50 ± 0.50 1.50 ± 0.00 1.50 ± 0.27 0.25
Beta 2.00 ± 0.00 2.00 ± 0.00 0.40 ± 0.16 0.50
Brookslate 1.50 ± 0.50 1.50 ± 0.50 0.00 ± 0.00 0.00
Choquette 2.00 ± 0.00 1.00 ± 0.00 0.00 ± 0.00 0.25
Donnie 2.50 ± 0.50 2.50 ± 0.50 1.00 ± 0.33 0.50
Dupuis 1.50 ± 0.50 1.50 ± 0.50 0.00 ± 0.00 0.00
Hall 2.00 ± 0.00 2.00 ± 0.00 1.30 ± 0.26 0.50
Loretta 1.00 ± 1.00 1.00 ± 1.00 0.00 ± 0.00 0.00
Lula 2.00 ± 0.00 2.00 ± 0.00 0.70 ± 0.00 0.25
Monroe 2.00 ± 0.00 2.00 ± 0.00 0.40 ± 0.16 0.25
Simmonds 1.50 ± 0.50 1.50 ± 0.50 1.70 ± 0.57 0.50
Tower 2 2.00 ± 1.00 2.00 ± 1.00 1.50 ± 0.17 0.00
Waldin 2.00 ± 0.00 2.00 ± 0.00 1.80 ± 0.13 0.50
Numbers followed by a different letter were significantly different at _ = 0.05; GLM procedure; Tukey’s Studentized Range
(HSD) test; SAS, 2008.
TABLE 3. SUSCEPTIBILITY OF PERSEA SPECIES TO ATTACK BY XYLEBORUS GLABRATUS AND OTHER SCOLYTINAE IN FLORIDA. BOLTS
WERE HUNG FOR 30 DAYS IN FIELD TESTS. BOLTS WERE EXPOSED TO 5 FEMALE X. GLABRATUS IN NO-CHOICE LABORATORY
BIOASSAYS.
Plant Species Tested
Field Test 1 Field Test 2 Laboratory Test
Entry
Holes/Bolt ± SE
Entry
Holes/Bolt ± SE
1
Emerging
Beetles/Bolt ± SE
Number of X. glabratus
entrances/bolt ± SE
1
P. caerulea
2.81 ± 2.57 0.50 ± 0.50 0.50 ± 0.50
3.00 ± 0.00 a
P. borbonia
0.37 ± 0.14 1.00 ± 1.00 0.00 ± 0.00 3.50 ± 0.50 a
P. pachypoda
1.73 ± 1.00 3.50 ± 1.50 1.00 ± 1.00 3.00 ± 1.00 a
P. floccosa
0.76 ± 0.28 0.00 ± 0.00 0.00 ± 0.00 0.00 ± 0.00 b
P. skutchii
5.61 ± 2.28 1.50 ± 1.50 0.00 ± 0.00 2.50 ± 0.50 a
P. nubigiena
2.14 ± 2.11 7.00 ± 3.00 2.50 ± 2.50
2.50 ± 0.50 a
P. indica
4.11 ± 2.71 0.50 ± 0.50 0.00 ± 0.00 1.50 ± 0.50 a
P. tolimanensis
1.57 ± 1.13 2.00 ± 2.00 0.00 ± 0.00 2.50 ± 0.50 a
Beilschmidia sp. 2.14 ± 1.95 0.50 ± 0.50 0.00 ± 0.00 3.00 ± 1.00 a
P. cinerascens
8.49 ± 6.71 2.00 ± 1.00 0.00 ± 0.00 4.00 ± 0.00 a
P. tilarensis
1.81 ± 1.57 4.00 ± 4.00 4.00 ± 4.00 3.50 ± 0.50 a
1
Means followed by a different letter are significantly different ( GLM Procedure; Tukey Studentized Range (HSD) Test; SAS
2008).
Scientific Notez 787
eter for RAB (0.8 mm, Hanula et al. 2008) were
counted and recorded. No dissection of the galler-
ies was made. Surprisingly, no RAB entry holes
were found on redbay logs, while entrance holes
were recorded on avocado (0.55 ± 0.29) and Cali-
fornia bay laurel (0.22 ± 0.14). However, no beetle
emerged from these bolts. Field tests and lab bio-
assays conducted during 2011 have shown that
female RAB are highly attracted to, and will bore
into freshly-cut bolts of U. californica (P. E. Ken-
dra, unpubl.; A. E. Mayfield, unpubl.).
SUMMARY
These preliminary results indicate that there
are numerous New World species of the Laura-
ceae potentially at risk of attack by X. glabratus.
More research is needed to fully determine the
susceptibility of Persea spp. and other genera
within the Lauraceae to both the pathogen and
vector. However, because of the difficulty in ob-
taining bolts of non-native Persea species and
other genera, efforts should be directed particu-
larly at those species where an indication of non-
susceptibility to X. glabratus has been observed.
A
CKNOWLEDGMENTS
We thank Drs. R. Giblin-Davis and R. E. Litz for
suggestions to improve the manuscript. We thank Jo-
se Alegria, Ana Vargas and the personnel of the Plant
Science Research Station of the University of Florida,
IFAS, Citra, Florida for their help. This research was
partially funded by the Florida Avocado Committee.
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