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Ecology of Root-Feeding Beetles and Their Associated Fungi on Longleaf Pine in Georgia


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Root-feeding beetles, particularly of the curculionid subfamilies Scolytinae and Molytinae, are known to be effective vectors of Ophiostomatoid fungi. Infestation by these insects and subsequent infection by the Ophiostomatoid fungi may play an important role in accelerating symptom progression in pine declines. To examine the relationship between beetles and fungi in longleaf pine stands, root-feeding curculionids were collected in pitfall traps baited with ethanol and turpentine for 62 wk, and Ophiostomatoid fungi were isolated from their body surfaces. The most abundant root-feeding beetles captured were Hylastes tenuis, H. salebrosus, Pachylobius picivorus, Hylobius pales, and Dendroctonus terebrans. The number of insects captured peaked in spring and fall, although peaks for different insect taxa did not coincide. The most frequently isolated fungi were Grosmannia huntii, Leptographium procerum, L. terebrantis, and L. serpens. Other Ophiostomatoid fungi recovered included Ophiostoma spp. and Pesotum spp. Insect infestation data suggest that Hylastes spp. share an ecological niche, as do Hb. pales and P. picivorus, because the ratios of their fungal symbionts were similar. The fungi associated with D. terebrans suggest that it did not share habitat with the other principle vectors.
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Ecology of Root-feeding Beetles and Their Associated Fungi on
Longleaf Pine in Georgia
School of Forestry and Wildlife Sciences, Auburn University, 602 Duncan Drive, Auburn University, AL 36849
Environ. Entomol. 39(2): 415Ð423 (2010); DOI: 10.1603/EN09261
ABSTRACT Root-feeding beetles, particularly of the curculionid subfamilies Scolytinae and Mo-
lytinae, are known to be effective vectors of Ophiostomatoid fungi. Infestation by these insects and
subsequent infection by the Ophiostomatoid fungi may play an important role in accelerating symptom
progression in pine declines. To examine the relationship between beetles and fungi in longleaf pine
stands, root-feeding curculionids were collected in pitfall traps baited with ethanol and turpentine for
62 wk, and Ophiostomatoid fungi were isolated from their body surfaces. The most abundant root-
feeding beetles captured were Hylastes tenuis,H. salebrosus,Pachylobius picivorus,Hylobius pales, and
Dendroctonus terebrans. The number of insects captured peaked in spring and fall, although peaks for
different insect taxa did not coincide. The most frequently isolated fungi were Grosmannia huntii,
Leptographium procerum,L. terebrantis, and L. serpens. Other Ophiostomatoid fungi recovered in-
cluded Ophiostoma spp. and Pesotum spp. Insect infestation data suggest that Hylastes spp. share an
ecological niche, as do Hb. pales and P. picivorus, because the ratios of their fungal symbionts were
similar. The fungi associated with D. terebrans suggest that it did not share habitat with the other
principle vectors.
KEY WORDS bark beetles, regeneration weevils, Ophiostomatoid fungi, Pinus palustris,Lep-
tographium spp
Bark beetles and regeneration weevils (Coleoptera:
Curculionidae, subfamilies Scolytinae and Molyti-
nae, respectively) live in a close association with
Ophiostomatoid fungi (Ascomycota: Ophiostoma-
tales) (WingÞeld et al. 1993). These fungi are com-
monly referred to as blue stain fungi because of the
cosmetic damage some species cause in logs and lum-
ber (Seifert 1993). Most members bear their meioti-
cally or mitotically derived spores in slimy matrices
suitable for transfer by insects (Malloch and Blackwell
1993), and they include the sexual genera Ophiostoma
Sydow and P. Sydow, Ceratocystiopsis Upadhyay and
Kendrick, and Grosmannia Goidanich, as well as the
asexual form-genera Pesotum Crane and Schoknecht
emend Okada and Seifert, Sporothrix Hektoen and
Perkins, Hyalorhinocladiella Upadhyay and Kendrick,
and Leptographium Lagerberg and Melin (Zipfel et al.
Some members of the genus Grosmannia (ana-
morph: Leptographium, sensu Zipfel et al. 2006) are
known for their role in forest diseases and declines.
Black stain root disease affects several conifer species
in the PaciÞc Northwest and is caused by G. wageneri
(Kendrick) Zipfel, de Beer and WingÞeld (Wagener
and Mielke 1961, Cobb 1988). The causal fungus is
vectored by bark beetles including Hylastes nigrinus
(Mannerheim) (Witcosky et al. 1986) and H. macer
LeConte (Goheen and Cobb 1978), and it is the only
Leptographium sp. known to be a primary pathogen of
conifers. In South Africa, L. serpens (Goidanich) Si-
emaszko, L. lundbergii Lagerberg and Melin and other
Ophiostomatoid fungi have been isolated from the
roots of dying Pinus spp. after early infestation by the
European bark beetles H. angustatus (Herbst) and
Hylurgus ligniperda (Fabricius) (WingÞeld et al. 1988;
Zhou et al. 2001, 2002). Pinus spp. have been intro-
duced in the southern hemisphere, in contrast with
northern hemisphere plantations where native pine
species are typically planted. Likewise, exotic pi-
nophagous bark beetles and their associated fungi
have been introduced into other southern hemisphere
countries; notably, H. ater (Paykull) has been found in
New Zealand accompanied by G. huntii (Robinson-
Jeffrey and Grinchenko) Zipfel, de Beer and Wing-
Þeld, L. procerum (Jacobs and WingÞeld 2001), and O.
ips (Reay et al. 2002). In Chile, H. ater and Hg. lig-
niperda have been found carrying G. huntii, O. ips, and
other Ophiostomatoid fungi (Zhou et al. 2004).
In the northeastern United States, white pine (Pinus
strobus L.) root decline has been linked to the pres-
ence of L. procerum (Kendrick) WingÞeld, L. tere-
brantis Barras and Perry (WingÞeld 1986, Alexander et
al. 1988) and their associated curculionid vectors in-
cluding the weevils Hylobius pales (Herbst) and Pis-
Corresponding author, e-mail:
0046-225X/10/0415Ð0423$04.00/0 2010 Entomological Society of America
sodes nemorensis Germar, and a suite of scolytine bee-
tles (Nevill and Alexander 1992). The same fungi have
been implicated in red pine (P. resinosa Aiton) decline
(Klepzig et al. 1991). The associated vectors were
more typically bark beetles, D. valens LeConte and H.
porculus Erichson (Klepzig et al. 1995), although the
weevils Pachylobius picivorus (Germar), Hb. pales, and
Hb. radicis Buchanan were also found to be infested
with L. procerum, L. terebrantis, and other Ophiosto-
matoid fungi including O. ips and G. huntii (Klepzig et
al. 1991).
In the southeastern United States, Ophiostomatoid
fungi and their vectors have been found to be impor-
tant contributing factors in loblolly pine (P. taeda L.)
decline (Eckhardt et al. 2007). In loblolly pine stands
experiencing decline, Hb. pales, P. picivorus, and Ps.
nemorensis were captured, as well as southern scoly-
tines of the same guild: H. salebrosus Eichhoff, H. tenuis
Eichhoff, and D. terebrans (Olivier). These insects
were found to be infested with L. procerum and L.
terebrantis, as well as L. serpens (Eckhardt et al. 2007).
In pathogenicity tests, L. serpens and L. terebrantis
had greater effects on seedling growth than L. pro-
cerum, with L. serpens producing larger lesions than
L. procerum or L. terebrantis in seedlings and mature
trees (Eckhardt et al. 2004b). Infestation with Lep-
tographium spp. was shown to increase brood size in
Hylastes spp., which in turn was effective in inoculat-
ing the fungi into uninfected root sections (Eckhardt
et al. 2004a).
Longleaf pine (P. palustris Miller) has been re-
ported to experience decline-like symptoms in the
southeastern United States (Otrosina et al. 1999). Al-
though a pyrophytic species (Frost 2006), mature lon-
gleaf pine were dying after prescription burning. A
survey of the roots showed L. procerum and L. tere-
brantis, but no connection was made to insect vectors
at that time (Otrosina et al. 1999). The effects of Þre
on insect populations in longleaf pine stands have
been examined after wildÞre (Hanula et al. 2002) and
prescription Þre (Sullivan et al. 2003). Hylastes spp.
and other phloem-feeders were more abundant in
unburned areas than in burned sites after a wildÞre in
Florida, whereas the trend with xylem-feeding scoly-
tines was an increase in populations with burn severity
(Hanula et al. 2002). In a similar study, applying pre-
scription burns of differing intensity in southern Al-
abama, the opposite trend was observed in phloeopha-
gous scolytines (Sullivan et al. 2003). Some of the
regeneration weevils were examined for the presence
of Ophiostomatoid fungi in the Florida study and
showed the presence of L. procerum and L. terebrantis
(Hanula et al. 2002).
Relatively little is known regarding the occurrence
or biology of the insects that infest pine roots or the
fungi that they may carry. For this reason, and asso-
ciated with a concern regarding the decline, the dis-
tribution and abundance of root-feeding bark beetles
and regeneration weevils, and their phoretic Ophios-
tomatoid fungi, were studied at Fort Benning Military
Reservation, GA. The study site is located in the san-
dhills ecoregion near the Fall Line separating the
coastal plain from the piedmont region in west central
Georgia. Fort Benning (FB) maintains an active pre-
scription burning program to enhance longleaf pine
habitat. In addition to investigating seasonal differ-
ences in beetle populations, the patterns of association
of the fungi that they carry were considered.
Materials and Methods
Plot Setup and Collection of Pinophagous Beetles.
Thirty-two 0.07-ha plots were installed at FB, in Mus-
kogee and Chattahoochee Counties in Georgia, and
Russell County, AL. Plot design followed the protocol
of the USDA Forest ServiceÕs Forest Inventory and
Analysis program (Dunn 1999). Each total plot was an
array of a central plot and three subplots: all circles of
7.62 m. Plots were blocked into four age classes (10,
10 Ð19, 20Ð 40, and 40 yr at the beginning of the study,
February 2006), with eight replicates of each age class.
Plot locations were selected by querying for appro-
priate stand type and age class in ArcGIS 3.2a (ESRI
1996), and 15 random points were generated per age
class. Final plot locations were selected by ground
truthing of the random points for appropriate stand
conditions (longleaf pine dominant, accurate age clas-
siÞcation). Although plots were occasionally situated
within a single stand, no two plots were closer than
150 m.
Pitfall traps were placed in each outlying subplot
(3/plot) to collect vector beetles. The traps (adapted
from Klepzig et al. 1991) were constructed of sections
of PVC tubing 10 cm diameter and 20 cm long with a
Þxed cap at the bottom end and a loose cap at the top.
Eight radial holes (6 mm diameter) were drilled to
allow insects to enter and fall into a cup placed inside
the trap. A plastic skirt was Þtted on the trap to reduce
the risk of ßooding (Menard 2007), such that the entry
holes on the trap were within 5 cm of ground level.
Steam-distilled southern pine turpentine (Hercules,
Wilmington, DE) and 95% ethanol (in two 8-ml vials)
were used as baits. Three sections of longleaf pine
twig, 5 cm long by 2 cm diameter, were also placed
in the cup as substrate for the captured insects and as
additional bait.
Pitfall trap collection was performed weekly from 4
March 2006 to 5 May 2006 (10 collections) and from
24 August 2006 to 26 August 2007 (52 collections).
Thus, data were collected for two spring seasons (2006
and 2007) and one continuous year (2006 Ð2007). Dur-
ing each trap collection, the contents of the pitfall
traps were transferred to a clean specimen cup and
stored at 4C for laboratory analysis. The trap cups
were sprayed with 70% ethanol to disinfect them and
swabbed with liquid Teßon (Northern Products,
Woonsocket, RI) to deter insects from crawling out of
the traps. Fresh twig sections were replaced in the trap
cups. The turpentine and ethanol baits were evacu-
ated and reÞlled. In the laboratory, insects were sorted
from each specimen cup and identiÞed to species and
Insect numbers were pooled at the plot and species
levels throughout the course of the sampling period
and effects of tree age class and sampling week tested
using repeated-measures analysis with the Þrst-order
autoregressive model (AR1) for the covariance struc-
ture (PROC MIXED; SAS Institute 2003).
Effects of Weather Variables. Effects of tempera-
ture and precipitation were assessed using weather
data from the National Climatic Data Center (http:// Data from the Colum-
bus Airport weather station were used, because data
from the FB weather station during the study period
are incomplete. Average weekly maxima and minima
were calculated from daily maxima and minima.
Weekly precipitation was summed from the daily pre-
cipitation data during the period between collection
dates. Weekly totals of H. tenuis, H. salebrosus, Hb.
pales, P. picivorus, and D. terebrans were pooled by
plot for correlation analysis with temperature and
precipitation (PROC CORR; SAS Institute 2003). To
assess seasonal effects, weekly capture numbers for
each plot and week were compared by coding 13-wk
periods with dummy variables and analysis (PROC
CORR and PROC GLM; SAS Institute 2003). For anal-
ysis of variance (ANOVA), means were separated
using the Tukey-Kramer procedure.
Infestation With Ophiostomatoid Fungi. Each cap-
tured beetle of interest was rolled across the surface
of one petri plate of malt extract agar amended with
cycloheximide and streptomycin (CSMA) (Hicks et
al. 1980) and one plate of unamended 2% malt extract
agar (wt:vol, MEA) to isolate fungal propagules. Cy-
cloheximide tolerance is a character shared by most
Ophiostomatoid fungi; unamended MEA was included
to allow for growth of rare cycloheximide-intolerant
species. Plates were inspected after 5 d and periodi-
cally thereafter for 4 wk for the presence of Ophios-
tomatoid fungi. These fungi where serially transferred
via spore droplets or hyphal tips to CSMA until pure
cultures had been obtained. The fungi were trans-
ferred to MEA for storage before species determina-
tion. IdentiÞcations were made based on microscopic
examination of morphological and cultural characters
as described in Jacobs and WingÞeld (2001) for Lep-
tographium spp. Other Ophiostomatoid fungi (Peso-
tum spp., Ophiostoma spp.) were identiÞed to genus
and pooled for subsequent study.
The distribution of fungi on insects was compared
tests (PROC FREQ; SAS Institute 2003). The
correlation between fungal isolation from beetles and
weather variables was tested in a similar manner.
Incidence of Pinophagous Beetle Activity and Sea-
sonal Effects. A total of 3,351 root-feeding beetles
were captured (Table 1). Four species of scolytine
bark beetles (H. tenuis, H. salebrosus, H. porculus, and
D. terebrans) and two species of molytine weevils (Hb.
pales and P. picivorus) were captured most frequently.
Voucher specimens of these beetles have been depos-
ited in the Louisiana State Arthropod Museum, Baton
Rouge, LA, as part of the dissertation work of the
primary author. Other scolytines captured included
Ips avulsus (Eichhoff) (n67), I. grandicollis (Eich-
hoff) (n34), Xyleborinus saxesenii (Ratzeburg)
(n162), and Xylosandrus crassiusculus (Motschul-
sky) (n84).
Hylastes tenuis was the most frequently captured
insect, with 1,726 individuals collected. A post hoc
examination of 250 randomly selected Hylastes indi-
viduals did not show any specimens of H. opacus Erich-
son, a morphologically similar but exotic insect. Ex-
cept where indicated below, data for H. salebrosus and
H. porculus were pooled, because both were tallied as
H. salebrosus during the collection period. A post hoc
examination of intact captured specimens showed 25%
(97/385) of beetles originally identiÞed as H. salebro-
sus were H. porculus. The ratio of these species varied
by season.
Week of capture was a signiÞcant factor in all spe-
cies tested by repeated-measures analysis (Table 2).
Stands in the 40-yr age class yielded more insects
than the 10-yr age class in all species tested, and
were higher than all other ages for H. tenuis, H. sale-
brosus, and D. terebrans (Fig. 1).
Table 1. Most frequently captured root-feeding beetles,
summed by stand age class at Fort Benning, GA
Age class (yr) Total
10 10Ð19 20 Ð40 40
H. tenuis 409 332 369 616 1,726
H. salebrosus
93 113 168 222 596
P. picivorus 82 184 122 179 567
Hb. pales 58 67 54 68 247
D. terebrans 16 55 47 97 215
Includes H. porculus.
Table 2. Summary statistics for repeated-measures analysis for effects of age class and week of capture at Fort Benning, GA, Aug.
2006 to Aug. 2007
Species Overall model Age class Week
F(df) PF(df) P
H. tenuis 130.84 0.0001 6148 7.39 (3,28) 0.001 8.60 (50,1550) 0.0001
H. salebrosus
53.2 0.0001 3598 3.67 (3,28) 0.02 4.87 (50,1550) 0.0001
P. picivorus 81.47 0.0001 4056 5.11 (3,28) 0.006 4.19 (50,1550) 0.0001
Hb. pales 1.85 0.17 1349 1.97 (3,28) 0.14 1.74 (50,1550) 0.001
D. terebrans 77.52 0.0001 2635 5.10 (3,28) 0.006 2.43 (50,1550) 0.0001
Null model likelihood ratio test for overall model,
Akaike information criterion.
Includes H. porculus.
Weekly temperature maxima ranged from 9 to 37C,
with an average of 25C, and the weekly temperature
minima ranged from 4.4 to 25C, with an average of
13C during the sampling period. Weekly precipita-
tion peaked at 13.2 cm and averaged 2.3 cm. Insects
were captured in all weeks except during 3 wk (Feb-
ruary 2007) when temperature minima were below
freezing. Weekly precipitation did not correlate with
numbers of insects captured, but annual precipitation
was below average overall.
Numbers of captured beetles peaked in spring and
fall, with a maximum of 100 beetles captured in spring
2006 and 149 in fall 2006. All species were present
during the spring trapping season in both years (Figs. 2
and 3). The number of insects captured, dropped to
zero for 3 wk in February 2007, corresponding to a
period of low temperature. D. terebrans was negatively
correlated with weekly temperature maxima and min-
ima (
⫽⫺0.368, P0.003 and
⫽⫺0.373, P0.003,
respectively), with a population peak during winter
trapping (
0.438, P0.0004). Although Hylastes
spp. peaked in spring and fall, the numbers of H. tenuis
were higher in the fall (
0.552, P0.0001), and the
number of H. salebrosus peaked in the spring (
0.633, P0.0001; Fig. 2A and B). The numbers of P.
picivorus captured were positively correlated with
temperature maxima and minima (
0.667, P
0.0001 and
0.643, P0.0001, respectively) and
were higher during the summer (
0.745, P0.0001;
Fig. 2C) and lower in fall and winter (
⫽⫺0.285, P
0.0249 and
⫽⫺0.420, P0.0007, respectively). The
number of Hb. pales captured was numerically higher,
but not signiÞcantly so, in spring (
0.214, P
Comparisons between numbers of insects captured
in spring 2006 and 2007 (Fig. 3) were similar for D.
terebrans (F7.37; df 1,8; P0.03), and H. sale-
brosus (including H. porculus, F 5.53; df 1.8; P
0.04). Comparisons of insects captured in the two
spring seasons were dissimilar for H. tenuis (F2.17;
df 1,8; P0.14), Hb. pales (F0.03; df 1,8; P
0.85), and P. picivorus (F0.01; df 1,8; P0.92).
Fig. 1. Mean number of captures per plot visit by age class at Fort Benning, GA, August 2006 to August 2007. Columns
within panes bearing the same letter are not signiÞcantly different at P0.05. Mean separation by Tukey Kramer method.
(A) H. tenuis, (B) larger Hylastes spp. (H. salebrosus and H. porculus), (C) P. picivorus, (D) Hb. pales, and (E) D. terebrans.
The subset of H. salebrosus that were examined in
the post hoc diagnoses showed the ratio of H. sale-
brosus:H. porculus to vary throughout the years. Spring
and summer ratios ranged from 90% H. salebrosus
(166:20 Hs:Hp) to 73% (60:13 Hs: Hp), but in winter,
the ratio shifted to 65% H. porculus (16:31 Hs:Hp).
Differences between seasonal ratios were signiÞcant
80.1; df 4; P0.0001).
Infestation With Ophiostomatoid Fungi. The pre-
dominant Ophiostomatoid fungi recovered from the
beetle species considered in this study were L. pro-
cerum, L. terebrantis, G. huntii, and Pesotum spp. (Ta-
ble 3). Overall, 27% of the major scolytine and moly-
tine insects yielded isolates of Ophiostomatoid fungi,
suggesting that these fungi are facultatively associated
with their insect vectors. L. procerum (24%), G. huntii
(23%), L. terebrantis (20%), and pooled Pesotum spp.
(16%) were the most frequently observed fungi-in-
festing insects (Table 3). The distribution of fungi was
similar between the pine regeneration weevils, H.
pales, and P. picivorus (
8.14; df 6; P0.23), and
between Hylastes spp. (
5.58; df 5; P0.35). The
mycota of D. terebrans differed from the other most
commonly captured beetles (
623.33; df 14; P
0.0001). D. terebrans rarely yielded isolates of L. tere-
brantis and had a higher frequency of isolates resem-
bling G. aureum and O. ips but probably undescribed
fungal species (M. J. WingÞeld, personal communica-
Leptographium procerum isolates from insects were
positively correlated with high weekly temperature min-
ima (
0.197, P0.0297; Table 4) and were most
frequently collected during fall 2006 (
0.307, P
0.0151). Fewer L. procerum isolates were collected dur-
ing winter (
⫽⫺0.279, P0.0278), and L. terebrantis
isolates increased in spring (
0.375, P0.0027) and
decreased in summer and winter (
⫽⫺0.403, P0.0012
⫽⫺0.258, P0.043, respectively). Ophiostoma
spp. isolates (excluding O. ips-like) were collected more
frequently in spring (
0.337, P0.0075) and less
frequently in winter (
⫽⫺0.301, P0.013). Ophios-
toma ips-like isolates were less abundant during periods
Fig. 2. Weekly captures of root-feeding beetles in longleaf pine stands at Fort Benning, GA, from 24 August 2006 to 26
August 2007. (A) H. tenuis, (B) larger Hylastes spp. (H. salebrosus and H. porculus), (C) P. picivorus, (D) Hb. pales, and (E)
D. terebrans.
of temperature minima (
⫽⫺0.323, P0.0104). The G.
aureumÐlike fungus was more frequently isolated in fall
0.259, P0.0417).
The results presented here showed that root-feed-
ing beetles are active throughout most of the year at
FB and that they carry a suite of Ophiostomatoid fungi,
including species previously unreported in the region.
Several studies have examined the bark beetle (ex-
cluding southern pine beetle) and regeneration wee-
vil fauna of southeastern forests (Fatzinger, 1985; Sul-
livan et al. 2003; Campbell et al. 2008a, b). However,
relatively few studies (Barnard et al. 1991, Otrosina et
al. 1999) have considered the Ophiostomatoid fungi
Fig. 3. Weekly captures (SEM) of root-feeding beetles in longleaf pine stands at Fort Benning, GA, spring (early March
to early May) 2006 and 2007. (A) H. tenuis, (B) larger Hylastes spp. (H. salebrosus and H. porculus), (C) P. picivorus, (D)
Hb. pales, and (E) D. terebrans.
Table 3. Incidence of fungi isolated from exoskeletons of root-feeding curculionids at Fort Benning, GA
Insect spp.(spring 2006,
spring 2007, total
Lt Ls Ga Gh Oi O.
spp. P. spp. Total by
insect spp.
D. terebrans (11,15,215) 4 5 0 26 2 33 6 10 86 40
H. tenuis (242,286,1726) 119 95 31 2 192 19 16 129 603 35
H. salebrosus (166,205,596)
11 15 1 0 26 17 3 13 86 14
P. picivorus (91,76,567) 68 47 0 0 4 3 4 1 127 22
Hb. pales (98,26,247) 32 31 2 0 4 3 2 3 77 31
Total by fungal spp. 234 193 34 28 228 75 31 156 979
Total includes both springs (early Mar. to early May), and fall 2006 to fall 2007 (late Aug. 2006 to late Aug. 2007) data.
Lp, L. procerum; Lt, L. terebrantis; Ga, G. aureum-like; Ls, L. serpens; Gh, G. huntii; Oi, O. ips-like; O. spp., other pooled Ophiostoma spp.;
P. spp., pooled Pesotum spp.
Includes H. porculus.
associated with these insects. In addition, very few
studies (Hanula et al. 2002, Eckhardt et al. 2007, Me-
nard 2007) have directly considered the relationship
between phloeophagous root-feeding curculionids
and their mycota in the southeastern United States.
Beetles were present during 59 of 62 wk of sampling,
indicating that root-feeding beetles and their fungi are
active at FB almost year-round. In previous studies of
the fungi associated with root-feeding beetles, collec-
tions were restricted to the spring, from early March
to May (Eckhardt et al. 2007, Menard 2007). The
results presented here indicate that year round sam-
pling is necessary to assess population peaks in the
curculionid fauna.
Numbers of insect captured in this study were lower
than those in studies using other trap designs such as
stovepipe (Fatzinger 1985) or multiple funnel traps
(Hanula et al. 2002, Sullivan et al. 2003). However, this
study required living insects to maximize fungal iso-
lation success, and this precluded the use of lethal
traps. Therefore, direct comparisons should be made
with those in loblolly pine with similar plot layouts,
identical traps, and performed in the same area (Eck-
hardt et al. 2007, Menard 2007).
In general, the numbers of vector captures were
similar to predicted asymptomatic plots in studies of
loblolly pine decline (Eckhardt et al. 2007, Menard
2007). The number of H. tenuis captured was greater
than the larger Hylastes spp., in contrast with other
studies where the larger insects were predominant
(Bauman 2003, Eckhardt et al. 2007) or similar in
number (Menard 2007). P. picivorus outnumbered Hb.
pales by 5:2. The natural range of these weevils
largely overlap, with Hb. pales being predominant
throughout much of that area, but the ratios are known
to shift in favor of P. picivorus in the southeast (Nord
et al. 1984). Fluctuations in the ratio of these weevils
have been found in other studies in the southeastern
United States, with P. picivorus more abundant in some
cases (Sullivan et al. 2003, Menard 2007), Hb. pales in
others (Hanula et al. 2002), or ßuctuating between the
two species (Eckhardt et al. 2007). Although strong
correlations with weather were observed for D. tere-
brans and P. picivorus, the single yearÕs data are not
deÞnitive, as has been shown in other year-round
studies (Sullivan et al. 2003). However, the occur-
rence of a winter peak in D. terebrans and H. porculus
in the southeast re-emphasizes the need for year-
round surveys for these species. Also, although little is
known of the biologies of H. porculus and H. salebrosus,
data presented here suggest that their population
peaks are not synchronous in western Georgia.
The overall occurrence of Ophiostomatoid fungi in
this study was lower than in previous studies (Eck-
hardt et al. 2007). The proportion of fungal isolations
from insects suggests niche overlap between the re-
generation weevil species and between Hylastes spp.
The rarity of L. terebrantis on its eponymous host was
unexpected, as a previous study has shown 90% infes-
tation of this fungus on D. terebrans (Eckhardt et al.
2007) compared with 2% in this study. G. huntii and
L. serpens were isolated more frequently from Hylastes
spp. L. serpens, previously reported from loblolly pine
and associated insects in Alabama (Eckhardt et al.
2007), was isolated infrequently from insects (1% of
all major vectors and 3.5% of all insect isolates).
Pesotum spp. (like Ophiostoma and Leptographium
spp.) are cycloheximide tolerant fungi formerly as-
signed to the form-genus Graphium. Their biology is
similar to Ophiostoma and Leptographium (Okada et
al. 1998, Harrington 2005). Few Pesotum spp. are
known to be pathogenic, although members of the
Ophiostoma ulmi complex produce Pesotum ana-
morphs and represent an important exception (Up-
adhyay 1981). Their role has been ignored in previous
studies with this pathosystems (Otrosina et al. 1999,
Eckhardt et al. 2007), and a rigorous attempt to iden-
tify isolates to species level was not pursued in this
study. Future studies may reinforce the observed
niche differences between insect taxa in these fungi as
well, and they might play an important role in longleaf
pine decline.
Grosmannia huntii was the fungus most frequently
found on Hylastes spp., and this is the Þrst time that it
has been recorded from the southeastern United
States. This fungus is morphologically similar to L.
serpens, and the two fungi might have been confused
in the past (Eckhardt et al. 2007.). Both fungi have
only recently been found in the southeastern United
States, and the dominance of these fungi on root feed-
ing insects suggests that they may play some role in the
ecology of their vectors. Furthermore, the fact that L.
serpens has been implicated in tree diseases in the past
(WingÞeld et al. 1988, Eckhardt et al. 2004a) suggests
that they deserve further study.
Table 4. Correlation table between weather variables and root-feeding-beetle-derived fungal isolates at Fort Benning, GA
Lt Ls Gh Ga Oi O. spp. P. spp. Insects
Weekly max temp 0.19736 0.00071 0.04624 0.00605 0.06092 0.23445 0.11978 0.09182 0.10169
0.1242 0.9956 0.7212 0.9628 0.6381 0.0666 0.3538 0.4779 0.4316
Weekly min temp 0.27638*0.02521 0.01137 0.03184 0.03488 0.32334*0.06045 0.06965 0.07036
0.0297 0.8458 0.9301 0.8059 0.7878 0.0104 0.6407 0.5906 0.5869
Weekly precipitation 0.12075 0.12445 0.00524 0.12050 0.02175 0.13169 0.13810 0.15595 0.16807
0.3498 0.3352 0.9678 0.3509 0.8668 0.3076 0.2844 0.2261 0.1916
For each cell, the upper value is the Pearson correlation coefÞcient (
) and the lower value is the probability, P, of the absolute value of
0. SigniÞcant values (P0.05) of
are indicated by asterisks.
Lp, L. procerum; Lt, L. terebrantis; Ga, G. aureum-like; Ls, L. serpens; Gh, G. huntii; Oi, O. ips-like; O. spp., other pooled Ophiostoma spp.;
P. spp., pooled Pesotum spp.
Some of the fungi isolated from insects in this study
most likely represent novel taxa that bear only a re-
semblance to known species. These include isolates
that have been treated as O. ips and G. aureum. There
is also some indication (unpublished data) that iso-
lates of G. huntii recovered in this study are differ-
ent to this fungus typiÞed by isolates of the fungus
originally described from lodgepole pine (P. con-
torta Douglas) in Canada (Robinson-Jeffrey and
Grinchenko 1964). Although fungal identiÞcations
presented in this study are sufÞcient to understand
patterns of association with root feeding insects, more
accurate identiÞcations in the future will reÞne our
understanding of the relationships of the Ophiosto-
matoid fungi with insects for which the ecology has
been minimally studied in the past.
Fungal recovery from insects varied by season, and
in some cases, these trends followed host afÞnities. For
example, the O. ips-like isolates were most commonly
isolated from insects captured in winter, which is con-
sistent with the fact that this fungus is closely associ-
ated with D. terebrans that was also captured most
frequently in winter months. In other cases, fungal
isolations seemed to be less dependent on the insect
vectors. For example, L. terebrantis isolations were
most common in spring, although the insects that carry
this fungus were most commonly captured in fall.
Whether temperature effects on seasonal changes are
caused by temperature directly or may result from
seasonal changes in host phenology or vector behavior
cannot be tested from the results of this study. How-
ever, the results suggest that a further examination of
seasonal effects on the occurrence of Ophiostomatoid
fungi with their insect vectors deserves further study.
We thank the Department of Defense for funding this
research and the staff of the Land Management Branch, Fort
Benning Military Reservation, in particular J. Parker, for
support. M. WingÞeld (University of Pretoria, Pretoria, South
Africa) and S. Enebak (Auburn University, Auburn, AL)
reviewed the manuscript and provided invaluable comments.
Prior studies by R. Menard provided a foundation for this
study. We further acknowledge the technical support of
several undergraduate research assistants and T. Garren-
Grubbs for archiving insect-derived fungal isolates.
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... It is thus assumed that the fungus was introduced into South Africa with these insects. In the USA, G. alacris has been consistently isolated from Hylastes salebrosus and Hylastes tenuis as well as root tissue of Pinus taeda and Pinus palustris infested with these insects (Eckhardt et al., 2007;Zanzot et al., 2010). Isolates of G. alacris have also been collected from France, Portugal and Spain; however, thorough studies have neither been conducted on the insect association nor the host range of this species in those countries (Jacobs & Wingfield, 2001;Duong et al., 2012a). ...
... Grosmannia alacris has the widest distribution of all species in the recently described G. serpens complex (Duong et al., 2012a). It has been associated with pine decline in various studies as well as in fungal-insect association studies in the USA and South Africa (Zhou et al., 2001;Eckhardt et al., 2004aEckhardt et al., , 2007Matusick & Eckhardt, 2010a,b;Zanzot et al., 2010;Duong et al., 2012a). However, nothing is known regarding the centre of origin nor the genetic diversity of the fungus in these countries. ...
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... Rhizophagous beetles and the fungi that they carry have been suggested to be associated with SPD (Eckhardt et al. 2007, Zanzot et al. 2010. Several rhizophagous beetle species occur in the pine ecosystems of the southern United States: native species such as Dendroctonus terebrans (Olivier), Hylastes porculus Erichson, Hylastes salebrosus Eichhoff, Hylastes tenuis Eichhoff, Hylobius pales (Herbst), Pachylobius picivorus (Germar), and Pissodes nemorensis Germar, as well as the nonnative species, Hylastes opacus Erichson. ...
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... but has also been documented to attack red spruce (Picea rubens Sargent) when these are >10 cm diameter (Wood 1982a, Staeben et al. 2010). Also, they show a preference for older trees and those weakened by fire, mechanical damage from logging or construction operations, disease, lightning strike, or colonization by other forest pests (Hopkins 1909, Kucera et al. 1970, Merkel 1981, Mayfield and Foltz 2005, Zanzot et al. 2010). ...
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... Hylastes salebrosus Eichhoff (Coleoptera: Curculionidae) and Hylastes tenuis Eichhoff (Coleoptera: Curculionidae) are the most common species associated with stressed, diseased, or dying pines in the southeastern United States. (Sullivan et al. 2003;Eckhardt et al. 2004;Zanzot et al. 2010). Adult H. salebrosus are about 4.5mm long and H. tenuis about 2-to 3.5-mm long (Drooz 1985). ...
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Root-feeding beetles, including several species of Hylastes Erichson (Coleoptera: Curculionidae), Hylurgops LeConte (Coleoptera: Curculionidae), Hylobius Gemar (Coleoptera: Curculionidae), Pachylobius LeConte (Coleoptera: Curculionidae), Pissodes Germar (Coleoptera: Curculionidae), and Steremnius Schönherr (Coleoptera: Curculionidae), have emerged as serious problems in conifer plantations and forests in the United States and Canada in recent decades. Root-feeding beetles are particularly associated with stressed, diseased, or injured trees. Emerging adults kill seedlings by girdling them at the root collar and kill older trees, at least in the North and West, by transmitting fungal root pathogens in the genus Leptographium Lagerberg & Melin (Ophiostomatales: Ophiostomataceae). In the South seedling, mortality can be as high as 60% for seedlings planted in winter following fall harvest. For stands harvested after 1 July, planting should be delayed a full year. Broadcast insecticides can be used, but dipping seedlings in 0.75% permethrin prior to planting and physical barriers to feeding have proven effective. However, an integrated pest management approach that emphasizes a combination of measures to minimize attraction of beetles and to maintain health of host trees is recommended. Shelterwood harvest and soil scarification can create site conditions that minimize attraction of root beetles. Precommercial thinning and prescribed fire are often used to reduce tree competition and reduce vulnerability to stem-colonizing bark beetles. However, root beetles are attracted to thinned or burned stands, for at least 6-7 mo. Therefore, thinning should be avoided in areas of high risk for root disease transmission or, when necessary, thinning should be implemented during June or July following beetle dispersal in May. Semiochemicals can be used to monitor abundances of root beetles.
... One of the best-known examples is Ophiostoma novo-ulmi Brasier, causing the destructive Dutch elm disease of elm (Ulmus) and being dispersed by bark beetles (Coleoptera, Curculionidae, Scolytinae) (Moser et al., 2010). Also interactions between Ophiostomatoid fungi with bark beetles have been well studied in North America, Europe and Japan (Wingfield 1983;Nevill and Alexander, 1992;Eckhardt et al. 2007;Zanzot et al. 2010;Jankowiak and Bilański, 2013;Robert 2013;Yamaoka 2017). Another interesting case is the chestnut blight fungus Cryphonectria parasitica (Murr.) ...
In Europe, the most widely used Christmas tree species, Abies nordmanniana, has a long history of infestation with the adelgid Dreyfusia nordmannianae. Since 2011, the species has furthermore been increasingly damaged by the fungus Neonectria neomacrospora. The objective was to study whether infestations by N. neomacrospora were facilitated by the presence of adelgids in Nordmann fir and to explore the genetic components of this potential interplay. The damage caused by D. nordmannianae and N. neomacrospora in two Nordmann fir clonal seed orchards in Denmark was evaluated in 2014, and an experimental infestation was conducted to test the performance of Nordmann fir somatic embryogenesis-derived trees exposed to N. neomacrospora and D. nordmannianae individually or in combination by setting up an artificial infestation chamber. The genetic correlation between the two diseases was 0.54 at two clonal seed orchards, indicating that the trees susceptible to N. neomacrospora were also susceptible to D. nordmannianae or vice versa. In the experimental test, only trees in the treatment with both D. nordmannianae and N. neomacrospora developed needle loss and dieback symptoms. Thus, both the field evaluations and the experimental test revealed an interplay between D. nordmannianae and N. neomacrospora damage on Nordmann fir trees. Future studies should explore in more detail how D. nordmannianae interacts with N. neomacrospora. Our findings and future studies on this topic could have important implications for suitable breeding programs for Nordmann fir Christmas trees and forest pest management.
... Scolytinae and Platypodinae are components of what are increasingly being termed " saproxylic " beetle communities— species associated with dead wood and associated structures (such as woody fungi) (Ausmus, 1977; Swift, 1977; Ahnlund, 1996; Hammond et al., 2001; Ulyshen et al., 2004; Ødegaard, 2004; Tykarski, 2006; Lachat et al., 2006 Lachat et al., , 2012 Zanzot et al., 2010). Host trees are usually dead or severely weakened, and their colonization by these beetles, which often carry with them a complex community of fungi, bacteria, yeasts, and mites, initiates the breakdown of plant tissues and recycling of nutrients. ...
The construction of hydrogeological cities in China has attracted attention from researchers worldwide. This study surveyed the major urban hydroecological river channels and selected 59 stations for hydroecological field sampling. The Ecopath model was used to screen for key river species, and niche analysis was performed to study the niche breadth and overlap between the key species and water quality factors. Sixteen key fish species were screened out; Pseudorasbora parva had the highest niche breadth (2.952). Of the 16 screened key species, Carassius auratus had the largest niche overlap (11.977). Seven key zooplankton species were screened out. Phryganella nidulus had the highest niche breadth (2.070). Of the seven screened key species, P. hemisphaerica had the largest niche overlap (9.065). Of the nine screened key species, Bithynia fuchsiana had the largest niche overlap. These findings provide a reliable theoretical basis for ecological restoration, and serve as a methodological reference for sustainable water resources management.
Frequent prescribed burns are essential to pine forest restoration and management. Research studies have assessed effects of prescribed fire and burn frequency on plants and vertebrates, but impacts of fire on terrestrial invertebrate communities are still poorly understood. This case study investigated effects of burning frequency on species richness and community composition of social insects (ants, Hymenoptera: Formicidae and termites, Blattodea: Isoptera) in fire-managed Southern longleaf pine flatwoods in central Florida. Community response to different fire frequencies was assessed: burned annually, every 2 yr, or every 3 yr, 30 yr unburned and 75 yr unburned. Richness was similar across all treatments, but ant community composition and species density significantly differed between frequently burned (1, 2, and 3 yr) and long-unburned (30 and 75 yr) treatments. Long-unburned treatments had higher ant abundance, but the species present were less characteristic of open canopy longleaf pine habitat than ants in frequently burned treatments. The annual burn treatment differed from 2-yr burn in species density, but to a lesser degree. Exotic species abundance was highest in frequently burned sites; only native species were detected in the 75-yr unburned plot. The red imported fire ant, Solenopsis invicta Buren (Hymenoptera: Formicidae), was detected in all regularly burned plots but not in long-unburned sites. Frequent burning at this site increased habitat suitability for ant species adapted to the sunny, open canopy, and diverse niches characteristic of longleaf pine forest; however, regular fire disturbance also increased the likelihood of exotic ant species establishment.
Heterobasidion annosum was confirmed to be present in 17 of 30 thinned slash pine (Pinus elliottii) plantations in north and north central Florida. Symptoms of possible root disease were detected on only 8% of 1,840 live trees, 14% of 2,204 stumps, and 13% of 811 dead trees examined. Although H. annosum was isolated from 47% of root and wood samples with white stringy rot, this symptom was observed in only 0.6, 5, and 4% of the live trees, stumps, and dead trees, respectively. Resin-soaking and/or staining was observed in 6, 2, and 7%, respectively, of the live trees, stumps, and dead trees, and H. annosum was isolated from only 10% of root and wood samples exhibiting this symptom. Inonotus circinatus was isolated from 9% of root and wood samples displaying resin-soaking and/or staining. Other root and stump infecting fungi detected were Armillariella tabescens, Phaeolus schweinitzii, Leptographium procerum, Fomitopsis palustris, Monascus floridanus, and a Ganoderma sp. A. tabescens was a predominant root and stump colonizer in four of the 30 plantations and sometimes occurred in the same roots as H. annosum. H. annosum was confirmed present in only two of 11 plantations surveyed with the “annosus sampling procedure,” whereas the presence of the pathogen was confirmed in six of the same 11 plantations via a 20-unit plantation-row plot method. In one plantation, H. annosum was undetected using the annosus sampling procedure, despite the fact that the fungus was isolated, respectively, from 33, 83, and 60% of the live trees, stumps, and dead trees sampled via the plantation-row plot method.
Wildfires burned over 200,000 ha of forest lands in Florida from April to July 1998. This unique disturbance event provided a valuable opportunity to study the interactions of summer wildfires with the activity of pine feeding insects and their associates in the southeastern United States. We compared tree mortality with abundance of bark and ambrosia beetles, reproduction weevils and wood borers relative to fire severity. Over 27% of residual live trees in stands that experienced high fire severity died between October 1998 and May 1999. An additional 2–3% of trees that initially survived the fire died during the second year compared to <1% mortality in unburned stands. One year after the fire, more than 75% of the trees surviving in high fire severity stands had roots infected with one or more species of Leptographium and/or Graphium spp. and nearly 60% of the sampled roots were infected. No such fungi were recovered from roots of trees in unburned stands. Significantly, more root weevils, Hylobius pales and Pachylobius picovorus, were captured in unbaited pitfalls in the moderate and high fire severity stands than in the controls. Mean trap catches of Ips grandicollis, Dendroctonus terebrans and Hylastes salebrosus, three common bark beetles that feed on phloem tissue of pines, were lower in Lindgren traps in the fire-damaged areas than in the control stands. In contrast, catches of the ambrosia beetles, Xyleborus spp. and Monarthrum mali, were higher in burned stands than in control stands. The generalist predator, Temnochila virescens (Coleoptera: Trogositidae), showed a strong positive relationship between abundance and fire severity, while the flat bark beetle, Silvanus sp. (Coleoptera: Sylvanidae), exhibited the reverse trend. Our results show that most tree mortality occurred within 1 year of the fire. Ips or Dendroctonus bark beetle populations did not build up in dead and weakened trees and attack healthy trees in nearby areas. The prevalence of Leptographium spp. in roots may be a symptom of, or result in, weakened trees that may affect the trees’ susceptibility to bark beetles in the future.