the coleopterists bulletin, 63(3):239–263. 2009. · [email protected] ... elevated /...
TRANSCRIPT
THE BEETLE COMMUNITY OF SMALL OAK TWIGS IN LOUISIANA, WITH A
LITERATURE REVIEW OF COLEOPTERA FROM FINE WOODY DEBRIS
MICHAEL L. FERRO, MATTHEW L. GIMMEL
Louisiana State Arthropod Museum, Department of EntomologyLouisiana State University AgCenter
402 Life Sciences BuildingBaton Rouge, LA, 70803, U.S.A.
[email protected]@gmail.com
KYLE E. HARMS
Department of Biological SciencesLouisiana State University202 Life Sciences Building
Baton Rouge, LA, 70803, [email protected]
AND
CHRISTOPHER E. CARLTON
Louisiana State Arthropod Museum, Department of EntomologyLouisiana State University AgCenter
402 Life Sciences BuildingBaton Rouge, LA, 70803, U.S.A.
Abstract
We conducted a study to explore which beetles utilize dead twigs in a Louisiana secondaryforest and the effect of debris position on the beetle community. Twigs averaging 14 mm indiameter from one tree of Quercus falcata Michaux (southern red oak) were placedrandomly into bundles of ten. At each of three sites, three bundles were laid on the ground,three were propped at the base of a living tree, and three were tied tightly above the groundagainst the branch of a living woody plant. The bundles were collected 10 months later andeach was placed into an emergence chamber. More than 400 adult Coleoptera specimenswere collected, representing 35 species within 16 families. Cerambycidae (longhorn beetles)and Curculionidae (weevils) exhibited the highest species richness, with nine and five species,respectively. Species richness was significantly different among treatments. Bundles placedon the ground had the lowest richness, aboveground bundles had the highest, and proppedbundles were intermediate. Twelve species (34%) were represented by singletons.
To our knowledge, no general survey of the saproxylic beetles associated withfine woody debris has been undertaken in the United States. However, severalstudies have shown considerable beetle richness in dead branches and twigs. Theexemplary research on saproxylic beetle succession conducted by Blackman andStage (1918, 1924) showed that beetle assemblages in limbs less than 6.35 cm indiameter were distinct from those in larger portions of the dead tree and at timeshad greater species richness (up to 32 species). Mecke et al. (2001) reared 34 beetle
The Coleopterists Bulletin, 63(3):239–263. 2009.
239
species from dead 3–12 cm diameter Araucaria limbs in Brazil. Numerous beetlespecies have been associated with twigs girdled by adults of various species in thecerambycid genus Oncideres Lepeletier and Audinet-Serville in Lacordaire(Linsley 1940; Polk and Ueckert 1973).
Twigs are important reservoirs of beetle richness and students of Coleopteraknow that twigs are an important habitat. Certain taxa are popularlycharacterized by their affinity to twigs. Many Scolytinae are reported from‘‘unthrifty twigs and branches’’ (Wood 1982). Some Bostrichidae are commonlycalled ‘‘Twig and Wood Borers’’ (Stehr 1991). Species of Oncideres are commonlycalled ‘‘Twig Girdlers’’ (Linsley 1940). Species of the genus ElaphidionoidesLinsley (5Anelaphus Linsley) are commonly called ‘‘Twig Pruners’’ (Solomon etal. 1999), and the curculionid Pityophthorus opaculus LeConte is commonly calledthe ‘‘Twig Beetle’’ (Stevens et al. 1979).
A review of the relevant literature (Table 1) shows that, aside from theeconomically important taxa mentioned above, little attention has been paid to thegeneral insect community that inhabits dead twigs. Here the term community ismeant to refer to the beetles inhabiting twigs sensu MacArthur (1971) (‘‘… any set oforganisms currently living near each other and about which it is interesting to talk’’).Based on literature records and limited rearing, Hovore and Penrose (1982) listed 19species of Cerambycidae and an additional 13 species within seven families of beetlesassociated with twigs girdled by Oncideres pustulata LeConte. Polk and Ueckert(1973) reported several families of Coleoptera, Hymenoptera, and Diptera (those ofthe latter not enumerated) reared from twigs girdled by Oncideres rhodosticta Bates.Rogers (1977) reported a bostrichid, a clerid, five species of cerambycids, and severalparasitoids that were associated with twigs girdled by Oncideres cingulata (Say). Beer(1949) reported rearing three species of Buprestidae from limbs of various trees.
The dead twig habitat is a difficult medium from which to collect beetles. Themost invasive method is direct dissection of the limb with removal of the (oftenimmature) insects (Sanborn 1911; Polk and Ueckert 1973; Rogers 1977). Whilethis may be the most expedient method of surveying twigs, the collector is biasedtoward species with large and easy-to-see adults and larvae, and against specieswith small adults and larvae (e.g., Scolytinae with mature larvae 2–10 mm long[Stehr 1991]). Additionally, larvae are often difficult or impossible to identify tothe species level using existing literature.
A second, and arguably more thorough, approach is to allow the emergence ofspecimens from the twig (Sanborn 1911; Linsley 1940; Beer 1949; Rogers 1977;Hovore 1980; Hovore and Penrose 1982). This has been accomplished using avariety of methods. Pollock (1988) reared subcortical larvae individually in 3-dramvials. Polk and Ueckert (1973) placed limbs into rearing chambers (the authors wereunclear about whether single or multiple limbs were in each chamber) and collectedpositively phototactic specimens that accumulated in translucent jars beneath thechamber. We have tabulated all published U.S. records of Coleoptera emergingfrom twigs in community level surveys (Appendix A).
The purposes of our experiment were to: 1) record which beetles (if any) utilizedead twigs in a secondary forest in Louisiana; and 2) determine if Coleopteraspecies composition is affected by twig position. Previous studies used a wide sizerange of branches or twigs, did not standardize for time since death, failed todifferentiate branch conditions (e.g., hanging, on ground, etc.), and may haveoverlooked species with small body sizes. In this study, tree species, size of limb, dateof death, treatment of limbs, inoculation time, and rearing time were known andheld constant, and a full census of the emergent beetle community was undertaken.
240 THE COLEOPTERISTS BULLETIN 63(3), 2009
Tab
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ty
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THE COLEOPTERISTS BULLETIN 63(3), 2009 241
Au
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242 THE COLEOPTERISTS BULLETIN 63(3), 2009
Au
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tes
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erg
ence
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sp
lan
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ma
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(Bu
pre
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(av
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ge
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)cm
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shly
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led
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nm
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ra
gen
to
fm
ort
ali
ty
Ta
ble
1.
Co
nti
nu
ed.
THE COLEOPTERISTS BULLETIN 63(3), 2009 243
Material and Methods
Study Area. The study was performed at Feliciana Preserve, a 61 ha tractlocated about 16 km east of St. Francisville, West Feliciana Parish, Louisiana(30u47.69N, 91u15.29W, WGS84). Feliciana Preserve encompasses a portion of theTunica Hills at the southern extreme of the Blufflands, a belt of thick loessoriginally blown from the Mississippi River floodplain (Delcourt and Delcourt1975). The resulting hills of easily erodible substrate have since formed a series ofdeep forested ravines. This area was a major refugium for mixed mesophyticforest species during the Wisconsin glaciation. Many taxa are holdovers from thisPleistocene event and occur nowhere else in Louisiana, or meet the southern limitof their range in the Tunica Hills (Delcourt and Delcourt 1975). This area alsocontains many subtropical elements not found further north, making this aunique collection of taxa in Louisiana.
The preserve consists of a secondary mixed mesophytic forest dominated bymagnolia (Magnolia grandiflora L.), holly (Ilex opaca Aiton), beech (Fagusgrandifolia Ehrhart), pine (Pinus spp.), and several species of oak (Quercus spp.).The area was clear-cut during the late 1950s (,55 years ago) and has remainedlargely undisturbed since. Feliciana Preserve is also the location of several studiesfocused on the Lepidoptera fauna of the area (Landau and Prowell 1999a, b;Landau et al. 1999; Prowell 2001).
Study Design. A single, healthy, ,30-year-old Quercus falcata Michaux(southern red oak) was felled 19 March 2006 in Feliciana Preserve. Followingthe cutting, 270 twigs averaging 36 cm (61 cm) in length and 14 mm (65 mm) indiameter were removed from the tree. Twigs were cut using saws and pruners sothat ends were square, not splintered. These were placed in a single pile and laterrandomly assigned to one of 27 bundles of ten twigs each. Each bundle was tiedwith two lengths of twine. The 27 bundles were then randomly sorted into threegroups of nine bundles each.
Three study sites, each situated ,300 m from each other, were used. Each sitewas within closed canopy forest and qualitatively similar with respect to standsize, age, substrate, amount of dead wood, litter, undergrowth, and distance fromforest openings. At each site, three random twig bundles were placed directly onthe ground (resting on top of the leaf litter), three twig bundles were propped atan approximately 45u angle, with the upper end against the trunk of a living treeand the lower end contacting the ground, and the remaining three bundles weretied horizontally against the limb of a small tree or woody shrub approximately1.5 m above the ground. Bundles placed directly on top of the leaf litter, bundlespropped against trees, and bundles tied 1.5 m above the ground are referred to asground, propped, and aboveground, respectively. Bundles were 2–4 m from theirnearest neighbor. A flag with an identity code was placed next to each bundle.The study was set up as quickly as possible (within about five hours) to ensurethat insects did not oviposit on the twigs prior to being positioned at the studysites, and to ensure that any volatiles or chemical attractants would still be presentin quantities strong enough to simulate a recent natural breakage event.
The bundles were left undisturbed until 28 January 2007 (ca. ten months later)when they were collected for emergence. Care was taken to ‘‘pounce’’ on eachbundle and transfer it to a container quickly so as to minimize loss of fast-movingindividuals fleeing the bundle. On site, each bundle was initially placed by itselfinto a clean white plastic ,19-L bucket (inside diameter 28 cm, height 36 cm),sealed, and removed to a central emergence area. Here, bundles were transferredto an emergence chamber that consisted of a similar bucket with an inverted foam
244 THE COLEOPTERISTS BULLETIN 63(3), 2009
bowl in the bottom. A small amount of full-strength propylene glycol antifreeze(PrestoneH Low Tox2 brand) was poured in the bottom of the chamber to serveas a killing and preservative agent. Twig bundles were transferred from theircollecting bucket to the emergence chamber and positioned more or less verticallywith the bottom end resting on the bowl so that the bundles were not in contactwith the propylene glycol. Any material left in the transfer bucket was dumpedinto the emergence chamber. Emergence chambers were sealed, labeled, andrandomly arranged in a covered, open air building.
On 12 July 2007 (ca. six months later), each chamber was opened and the twigbundleswereshakenandvisually inspectedforadults.Thepropyleneglycol/specimen/frass slurry was washed into a labeled Whirl-PakH and removed to the laboratory forsorting. Adult Coleoptera were pinned or pointed as needed, and labeled.Identification to the finest level possible (typically species) was performed with theappropriate taxonomic literature and/or comparison with authoritatively identifiedreference specimens. All other macroinvertebrates were sorted from the debris,labeled, and preserved in 90% ethanol. Specimens are deposited in the Louisiana StateArthropod Museum (LSAM), LSU AgCenter, Baton Rouge, Louisiana.
Because of the exploratory nature of this study, several design aspects weresomewhat arbitrary. Twigs of southern red oak were used because there is aconcurrent study within Feliciana Preserve that is exploring aspects of Coleopteracommunities within other portions of the same tree species. The twig diameterwas chosen because it falls below the diameter of the smallest wood used in adifferent concurrent study, but was judged robust enough to harbor multipleinsects. Twigs were placed in bundles of ten to reduce twig loss over time andbecause of convenient mathematical aspects. The length of the twigs was chosenso that the rearing chambers could comfortably accommodate them. Twigs werecut early in the spring to allow for ample colonization time, gathered before aputative spring pulse of emergence, and allowed to rear for an extended period oftime to accommodate the emergence of as many adults as possible.
Statistical Analysis. Statistical significance (a 5 0.05) was tested using MonteCarlo simulations to compare observed data to null expectations generated byappropriately randomizing those observed data (Manly 2007). Average speciesrichness per bundle and total number of specimens for each site and treatmentcombination (n 5 9 bundles for each site and treatment combination) wereindividually compared to null distributions, i.e., distributions of expected valuesgenerated from 1,000 randomizations. Each randomization reassigned eachbundle’s observed species richness and number of specimens to a randomlychosen site and treatment combination (without replacement), whereupon ninerandomized bundles’ values were chosen at random to calculate one randomizedvalue of average species richness and one randomized value of total number ofspecimens. The randomizations provided 1,000 randomized values for expectedspecies richness per bundle and 1,000 randomized values for expected totalnumber of specimens. A significant deviation from randomized expectationsoccurred when an observed value fell in the upper or lower 2.5% tail of thecorresponding frequency distribution of expected (randomized) values; observedvalues in the upper 2.5% tail were significantly higher than expected, and viceversa (Prado and Lewinsohn 2004; Manly 2007).
Results
Richness. A total of 414 adult Coleoptera specimens were collected,representing 35 species in 33 genera and 16 families (Table 2). Twig bundles
THE COLEOPTERISTS BULLETIN 63(3), 2009 245
Ta
ble
2.
Co
leo
pte
raem
erg
ent
fro
mo
ak
twig
bu
nd
les
inL
ou
isia
na
.T
ax
aa
reg
rou
ped
acc
ord
ing
top
rese
nce
intr
eatm
ents
.T
ho
sep
rese
nt
ina
lltr
eatm
ents
ap
pea
ra
tth
eto
p,
tho
sein
on
lytw
otr
eatm
ents
foll
ow
,a
nd
tho
sep
rese
nt
ino
nly
on
etr
eatm
ent
ap
pea
rla
st.
Fam
ily
Sp
ecie
s
Ind
ivid
ua
lsem
erg
ed:
To
tals
Gro
un
dP
rop
ped
Ab
ov
egro
un
d
Bip
hy
llid
ae
Dip
loco
elu
sru
dis
(LeC
on
te,
18
63
)2
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up
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sole
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)6
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pin
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alp
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y,
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27
)1
47
22
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246 THE COLEOPTERISTS BULLETIN 63(3), 2009
Fa
mil
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ity
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sey
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41
4
To
tal#
spec
ies
13
15
24
35
Tab
le2
.C
on
tin
ued
.
THE COLEOPTERISTS BULLETIN 63(3), 2009 247
placed directly on the ground yielded 130 specimens, representing 13 species, 13genera, and 10 families. Propped twig bundles yielded 91 specimens, representing15 species, 14 genera, and 8 families. Aboveground bundles yielded 193specimens, representing 24 species, 22 genera, and 10 families.
Within the entire collection, 13 species (37%) were represented by five or moreindividuals, whereas 22 species (63%) were represented by fewer than fiveindividuals. Twelve species (34%) were represented by singletons. The number ofspecies represented by fewer than five individuals was 10 (77%), 8 (53%), and 13(54%) for twig bundles placed on the ground, propped, and aboveground,respectively.
The buprestid Agrilus obsoletoguttatus Gory was the most abundant species(131 specimens), accounting for 32% of all individuals, and having emerged fromall three treatments. The scolytine curculionid Pseudothysanoes dislocatus(Blackman) had the second highest abundance with 56 individuals (13.5%), butonly emerged from twig bundles that were aboveground. Five species, A.obsoletoguttatus, Liopinus alpha (Say), Anelaphus villosus (Fabricius), Diplocoelusrudis (LeConte), and Melanophthalma distinguenda (Comolli), emerged from allthree treatments. A single species, Neoclytus acuminatus (Fabricius), was sharedbetween ground and propped treatments, but was not found in the abovegroundtreatment. Six species, Hypothenemus californicus Hopkins, Cyrtinus pygmaeus(Haldeman), Ecyrus dasycerus (Say), Laemosaccus nephele (Herbst), Anelaphuspumilus (Newman), and Attalus scincetus (Say), were shared among propped andaboveground treatments, but did not emerge from bundles placed on the ground.Seven species that emerged from bundles placed on the ground were unique tothat treatment, three species were unique to propped bundles, and 13 species wereunique to aboveground bundles.
Species represented by five or more individuals within a particular treatmentoccurred in only four families: Buprestidae, Cerambycidae, Curculionidae, andCleridae. However, Curculionidae were absent from ground bundles, andCleridae were unique to aboveground bundles. Cerambycidae were representedby ten species, followed by Curculionidae with five species. Six families wererepresented by two species and the remaining eight families were each representedby a single species.
Sites Comparison. A total of 146 adult Coleoptera emerged from all bundles atSite I, 183 adult Coleoptera emerged from bundles at Site II, and 85 emerged frombundles at Site III. Numbers of specimens emerged per bundle were notsignificantly different among sites (P . 0.05). The average number of species perbundle was 4.0, 4.3, and 3.4 for Sites I, II, and III, respectively. Number of speciesemerged per bundle were not significantly different among sites (P . 0.05).
Treatments Comparison. A total of 130 adult Coleoptera emerged from allbundles placed on the ground, 91 adult Coleoptera emerged from proppedbundles, and 193 emerged from aboveground bundles. Numbers of specimenswere not significantly different among treatments (P . 0.05). The average numberof species per bundle was 2.3, 3.8, 5.7, and 3.9 for ground, propped, aboveground,and all treatments combined, respectively. The average species richness perbundle placed on the ground was significantly less than expected (P , 0.05) fromthe null distribution. The average species richness per propped bundle was notsignificantly different (P . 0.05) from the null distribution. The average speciesrichness per aboveground bundle was significantly more than expected (P , 0.05)from the null distribution. Thus, species richness per bundle ranked as follows:ground , propped , aboveground.
248 THE COLEOPTERISTS BULLETIN 63(3), 2009
Discussion
While this may be likened to one of Charles Darwin’s ‘‘fool’s experiments’’(Darwin 1887), the results of this research were astounding. From a smattering offinger-sized twigs we collected over 400 specimens and 35 species of beetles. Halfthe species collected were wholly absent (seven species) or were represented by fiveor fewer specimens (ten species) in the Louisiana State Arthropod Museum(which houses approximately 600,000 Coleoptera specimens from the region).This should be of great interest to researchers conducting comprehensive faunalinventories, and we highly recommend addition of this or a similar method to thebiodiversity surveyor’s and ecologist’s toolkit.
Species richness results among treatments were also unexpected. Cramer (1998)reported complete mortality of the cerambycid O. cingulata caused by desiccationof twigs brought to the laboratory. However, Cerambycidae, while reared fromall three treatments, showed the highest number of individuals and diversity inpropped and aboveground bundles, presumably the driest treatments.
Aboveground twig bundles had the highest species richness, but presumably: 1)were more susceptible to desiccation; 2) experienced greater changes in dailytemperature; and 3) were generally less accessible to potential colonists. Incontrast, bundles placed on the ground had the lowest richness, but presumablynone of the above limitations.
Interestingly, propped bundles, which were presumably more accessible tocolonists than aboveground bundles but more prone to desiccation than groundbundles, showed a greater similarity to aboveground bundles. This initiallysuggests that desiccation of limbs may be more important at promoting speciesrichness than accessibility to colonists. Petrice and Haack (2006) reported thatdesiccation was a major cause of mortality in Agrilus planipennis Fairmaire (theemerald ash borer), a congener of A. obsoletoguttatus. Perhaps A. obsoletoguttatusloses its dominance in drier wood, allowing other species to utilize the freedresources.
Increased exposure to predation, especially by ants (Formicidae), probably didnot contribute to lower species richness of bundles placed in full contact with theground or propped. When considering this question, predation and ability tocolonize must be considered simultaneously. The number of specimens did notdiffer among treatments. This could occur if predation and ability to colonizewere equal for all treatments, or it could occur if predation and ability to colonizediffered among treatments, with predation increasing at the same rate as ability tocolonize. The relative proportion of the two factors would have had to be equalacross all treatments. This is unlikely due to great differences in ability to colonizeamong the bundles placed on the ground and those that were aboveground.Additionally, predation would probably not have resulted in the loss of allindividuals of a given taxon within a treatment if sufficient numbers of that taxonhad been present.
This research indicates that a rich, perhaps unique beetle fauna inhabits finewoody debris. Beetle communities among twigs that are in full contact with theground appear to be distinct from those that remain in the tree after death. Thesebasic observations may now be used to foster hypothesis-driven research.
Acknowledgments
We thank Dorothy Prowell (Louisiana State University) for allowing the studyto be conducted on her property and access to the outbuilding-turned-emergence
THE COLEOPTERISTS BULLETIN 63(3), 2009 249
facility. We thank Robert Rabaglia (Maryland Department of Agriculture) forthe scolytine identifications, Stephanie Gil (Louisiana State University) for helpwith the corylophid identification, and Sarah Samson (University of Missouri,Columbia) for assistance in the field. This publication was approved by theDirector, Louisiana Agricultural Experiment Station as manuscript number 2009-234-2782.
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THE COLEOPTERISTS BULLETIN 63(3), 2009 251
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252 THE COLEOPTERISTS BULLETIN 63(3), 2009
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stid
ae
An
tha
xia
viri
dif
ron
sG
ory
,1
84
1C
ary
ag
lab
ra(M
ill.
)S
wee
t[a
sH
ico
ria
gla
bra
Mil
l.]
,6
.4cm
Tre
ek
ille
db
yS
coly
tus
qu
adri
spin
osu
sS
ay
(Sco
lyti
na
e)
NY
Bla
ckm
an
an
dS
tage
19
24
Bu
pre
stid
ae
Ch
ryso
bo
thri
sa
na
lis
LeC
on
te,
18
60
Leu
caen
ap
ulv
eru
len
ta(S
chlt
dl.
)B
enth
.A
sso
cia
ted
wit
htw
igs
gir
dle
db
yO
nci
der
esp
ust
ula
taL
eCo
nte
(Cer
am
by
cid
ae)
TX
Ho
vo
rea
nd
Pen
rose
19
82
Bu
pre
stid
ae
Ch
ryso
bo
thri
sd
enti
pes
(Ger
ma
r,1
82
4)
La
rix
lari
cin
a(D
uR
oi)
K.
Ko
ch.
2.5
–5
cmD
ead
tree
,u
nk
no
wn
cau
seN
YB
lack
man
an
dS
tage
19
18
Bu
pre
stid
ae
Ch
ryso
bo
thri
sfe
mora
ta(O
liv
ier,
17
90)
Ca
rya
gla
bra
(Mil
l.)
Sw
eet
[as
Hic
ori
ag
lab
raM
ill.
]
,6
.4cm
Tre
ek
ille
db
yS
coly
tus
qu
adri
spin
osu
sS
ay
(Sco
lyti
na
e)
NY
Bla
ckm
an
an
dS
tage
19
24
Bu
pre
stid
ae
Ch
ryso
bo
thri
sro
tun
dic
oll
isG
ory
&L
ap
ort
e,1
83
7[a
sC
hry
sob
oth
ris
bla
nch
ard
iH
orn
]
La
rix
lari
cin
a(D
uR
oi)
K.
Ko
ch.
2.5
–5
cmD
ead
tree
,u
nk
no
wn
cau
seN
YB
lack
man
an
dS
tage
19
18
Bu
pre
stid
ae
Ch
ryso
bo
thri
sse
xsi
gn
ata
Sa
y,
18
39
Ca
rya
gla
bra
(Mil
l.)
Sw
eet
[as
Hic
ori
ag
lab
raM
ill.
]
,6
.4cm
Tre
ek
ille
db
yS
coly
tus
qu
adri
spin
osu
sS
ay
(Sco
lyti
na
e)
NY
Bla
ckm
an
an
dS
tage
19
24
Bu
pre
stid
ae
Ch
ryso
bo
thri
sse
xsi
gn
ata
Sa
y,
18
39
La
rix
lari
cin
a(D
uR
oi)
K.
Ko
ch.
2.5
–5
cmD
ead
tree
,u
nk
no
wn
cau
seN
YB
lack
man
an
dS
tage
19
18
Bu
pre
stid
ae
Dic
erca
luri
da
(Fa
bri
ciu
s,1
77
5)
Ca
rya
gla
bra
(Mil
l.)
Sw
eet
[as
Hic
ori
ag
lab
raM
ill.
]
,6
.4cm
Tre
ek
ille
db
yS
coly
tus
qu
adri
spin
osu
sS
ay
(Sco
lyti
na
e)
NY
Bla
ckm
an
an
dS
tage
19
24
THE COLEOPTERISTS BULLETIN 63(3), 2009 253
Ap
pen
dix
A.
Co
nti
nu
ed.
Co
leo
pte
rafa
mil
yC
ole
op
tera
spec
ies
Pla
nt
spec
ies
Su
bst
rate
dia
met
erS
ub
stra
teo
rig
inS
tate
Ref
eren
ce
Bu
pre
stid
ae
Ph
aen
op
sfu
lvo
gu
tta
ta(H
arr
is,
18
29)
[as
Mel
ano
ph
ila
fulv
og
utt
ata
(Ha
rris
)]
Lari
xla
rici
na
(Du
Ro
i)K
.K
och
.2
.5–
5cm
Dea
dtr
ee,
un
kn
ow
nca
use
NY
Bla
ckm
an
an
dS
tag
e1
91
8
Cer
am
by
cid
ae
Ach
ryso
nsu
rina
mu
m(L
inn
aeu
s,1
76
7)
Ass
oci
ate
dw
ith
twig
sg
ird
led
by
On
cid
eres
pu
stu
lata
LeC
on
te(C
era
mb
yci
da
e)
Lin
sley
19
40
Cer
am
by
cid
ae
Ach
ryso
nsu
rina
mu
m(L
inn
aeu
s,1
76
7)
Leu
caen
ap
ulv
erule
nta
(Sch
ltd
l.)
Ben
th.
Ass
oci
ate
dw
ith
twig
sg
ird
led
by
On
cid
eres
pu
stu
lata
LeC
on
te(C
era
mb
yci
da
e)
TX
Ho
vo
rea
nd
Pen
rose
19
82
Cer
am
by
cid
ae
Aeg
om
orp
hu
sm
od
estu
s(G
yll
enh
al,
18
17)
[as
A.
dic
ipie
ns
Ha
ldem
an
]
Ca
rya
gla
bra
(Mil
l.)
Sw
eet
[as
Hic
ori
ag
lab
raM
ill.
]
,6
.4cm
Tre
ek
ille
db
yS
coly
tus
qu
ad
risp
ino
sus
Sa
y(S
coly
tin
ae)
NY
Bla
ckm
an
an
dS
tag
e1
92
4
Cer
am
by
cid
ae
An
ela
ph
us
deb
ilis
(LeC
on
te,
18
54)
Leu
caen
ap
ulv
erule
nta
(Sch
ltd
l.)
Ben
th.
Ass
oci
ate
dw
ith
twig
sg
ird
led
by
On
cid
eres
pu
stu
lata
LeC
on
te(C
era
mb
yci
da
e)
TX
Ho
vo
rea
nd
Pen
rose
19
82
Cer
am
by
cid
ae
An
ela
ph
us
iner
mis
(New
ma
n,
18
40)
[as
An
op
liu
mtr
un
catu
mL
eCo
nte
]
Ass
oci
ate
dw
ith
twig
sg
ird
led
by
On
cid
eres
pu
stu
lata
LeC
on
te(C
era
mb
yci
da
e)
Lin
sley
19
40
Cer
am
by
cid
ae
Ast
ylo
psi
sse
xg
utt
ata
(Sa
y,
18
26
)[a
sL
epto
sty
lus
sex
-g
utt
atu
s(S
ay
)]
Lari
xla
rici
na
(Du
Ro
i)K
.K
och
.2
.5–
5cm
Dea
dtr
ee,
un
kn
ow
nca
use
NY
Bla
ckm
an
an
dS
tag
e1
91
8
Cer
am
by
cid
ae
Ata
xia
cry
pta
(Sa
y,
18
31)
Ass
oci
ate
dw
ith
twig
sg
ird
led
by
On
cid
eres
cin
gu
lata
tex
an
aH
orn
(Cer
am
by
cid
ae)
Lin
sley
19
40
254 THE COLEOPTERISTS BULLETIN 63(3), 2009
Ap
pen
dix
A.
Co
nti
nu
ed.
Co
leo
pte
rafa
mil
yC
ole
op
tera
spec
ies
Pla
nt
spec
ies
Su
bst
rate
dia
met
erS
ub
stra
teo
rigin
Sta
teR
efer
ence
Cer
am
by
cid
ae
Cy
rto
ph
oru
sve
rru
cosu
s(O
liv
ier,
17
95
)C
ary
ag
lab
ra(M
ill.
)S
wee
t[a
sH
ico
ria
gla
bra
Mil
l.]
,6
.4cm
Tre
ek
ille
db
yS
coly
tus
qu
adri
spin
osu
sS
ay
(Sco
lyti
na
e)
NY
Bla
ckm
an
an
dS
tage
19
24
Cer
am
by
cid
ae
Do
rca
sch
ema
cin
ereu
m(O
liv
ier,
17
95
)[a
sH
eto
emis
cin
erea
(Oli
vie
r)]
Ca
rya
gla
bra
(Mil
l.)
Sw
eet
[as
Hic
ori
ag
lab
raM
ill.
]
,6
.4cm
Tre
ek
ille
db
yS
coly
tus
qu
adri
spin
osu
sS
ay
(Sco
lyti
na
e)
NY
Bla
ckm
an
an
dS
tage
19
24
Cer
am
by
cid
ae
Do
rca
sch
ema
nig
rum
(Sa
y,
18
26
)C
ary
ag
lab
ra(M
ill.
)S
wee
t[a
sH
ico
ria
gla
bra
Mil
l.]
,6
.4cm
Tre
ek
ille
db
yS
coly
tus
qu
adri
spin
osu
sS
ay
(Sco
lyti
na
e)
NY
Bla
ckm
an
an
dS
tage
19
24
Cer
am
by
cid
ae
Ebu
ria
mu
tica
LeC
on
te,
18
53
Leu
caen
ap
ulv
eru
len
ta(S
chlt
dl.
)B
enth
.A
sso
ciate
dw
ith
twig
sg
ird
led
by
On
cid
eres
pu
stu
lata
LeC
on
te(C
era
mb
yci
da
e)
TX
Ho
vo
rea
nd
Pen
rose
19
82
Cer
am
by
cid
ae
Ecy
rus
arc
ua
tus
Ga
ha
n,
18
92
[as
Ecy
rus
tex
an
us
Sch
aef
fer]
Ass
oci
ate
dw
ith
twig
sg
ird
led
by
On
cid
eres
pu
stu
lata
LeC
on
te(C
era
mb
yci
da
e)
Lin
sley
19
40
Cer
am
by
cid
ae
Eud
erce
sre
ich
eiL
eCo
nte
,1
87
3[a
sE
ud
erce
sex
ilis
Ca
sey
]
Ass
oci
ate
dw
ith
twig
sg
ird
led
by
On
cid
eres
pu
stu
lata
LeC
on
te(C
era
mb
yci
da
e)
Lin
sley
19
40
Cer
am
by
cid
ae
Eud
erce
sp
icip
es(F
ab
rici
us,
17
87)
Ca
rya
gla
bra
(Mil
l.)
Sw
eet
[as
Hic
ori
ag
lab
raM
ill.
]
,6
.4cm
Tre
ek
ille
db
yS
coly
tus
qu
adri
spin
osu
sS
ay
(Sco
lyti
na
e)
NY
Bla
ckm
an
an
dS
tage
19
24
Cer
am
by
cid
ae
Ger
op
aco
nco
lor
(LeC
on
te,
18
73
)[a
sA
chry
son
conco
lor
LeC
on
te]
Ass
oci
ate
dw
ith
twig
sg
ird
led
by
On
cid
eres
pu
stu
lata
LeC
on
te(C
era
mb
yci
da
e)
Lin
sley
19
40
Cer
am
by
cid
ae
Ger
op
aco
nco
lor
(LeC
on
te,
18
73
)L
euca
ena
pu
lver
ule
nta
(Sch
ltd
l.)
Ben
th.
Ass
oci
ate
dw
ith
twig
sg
ird
led
by
On
cid
eres
pu
stu
lata
LeC
on
te(C
era
mb
yci
da
e)
TX
Ho
vo
rea
nd
Pen
rose
19
82
THE COLEOPTERISTS BULLETIN 63(3), 2009 255
Ap
pen
dix
A.
Co
nti
nu
ed.
Co
leo
pte
rafa
mil
yC
ole
op
tera
spec
ies
Pla
nt
spec
ies
Su
bst
rate
dia
met
erS
ub
stra
teo
rig
inS
tate
Ref
eren
ce
Cer
am
by
cid
ae
Gn
ap
ha
lodes
tra
chy
der
oid
esT
ho
mso
n,
18
60
Leu
caen
ap
ulv
erule
nta
(Sch
ltd
l.)
Ben
th.
Ass
oci
ate
dw
ith
twig
sg
ird
led
by
On
cid
eres
pu
stu
lata
LeC
on
te(C
era
mb
yci
da
e)
TX
Ho
vo
rea
nd
Pen
rose
19
82
Cer
am
by
cid
ae
Het
era
chth
esq
ua
dri
macu
latu
sH
ald
ema
n,
18
47
Ca
rya
gla
bra
(Mil
l.)
Sw
eet
[as
Hic
ori
ag
lab
raM
ill.
]
,6
.4cm
Tre
ek
ille
db
yS
coly
tus
qu
adri
spin
osu
sS
ay
(Sco
lyti
na
e)
NY
Bla
ckm
an
an
dS
tag
e1
92
4
Cer
am
by
cid
ae
An
ela
ph
us
vill
osu
s(F
ab
rici
us,
17
92
)[a
sH
yp
erm
all
us
vill
osu
s(F
ab
rici
us)
]
Ca
rya
gla
bra
(Mil
l.)
Sw
eet
[as
Hic
ori
ag
lab
raM
ill.
]
,6
.4cm
Tre
ek
ille
db
yS
coly
tus
qu
adri
spin
osu
sS
ay
(Sco
lyti
na
e)
NY
Bla
ckm
an
an
dS
tag
e1
92
4
Cer
am
by
cid
ae
Kn
ull
ian
aci
nct
a(D
rury
,1
77
3)
[as
Ch
ion
cin
ctu
sD
rury
]
Ca
rya
gla
bra
(Mil
l.)
Sw
eet
[as
Hic
ori
ag
lab
raM
ill.
]
,6
.4cm
Tre
ek
ille
db
yS
coly
tus
qu
adri
spin
osu
sS
ay
(Sco
lyti
na
e)
NY
Bla
ckm
an
an
dS
tag
e1
92
4
Cer
am
by
cid
ae
Lep
turg
esa
ng
ula
tus
(LeC
on
te,
18
52)
[as
Lep
turg
esa
ng
ula
tus
can
us
Ca
sey
]
Leu
caen
ap
ulv
erule
nta
(Sch
ltd
l.)
Ben
th.
Ass
oci
ate
dw
ith
twig
sg
ird
led
by
On
cid
eres
pu
stu
lata
LeC
on
te(C
era
mb
yci
da
e)
TX
Ho
vo
rea
nd
Pen
rose
19
82
Cer
am
by
cid
ae
Lep
turg
esin
fila
tus
Ba
tes,
18
72
Leu
caen
ap
ulv
erule
nta
(Sch
ltd
l.)
Ben
th.
Ass
oci
ate
dw
ith
twig
sg
ird
led
by
On
cid
eres
pu
stu
lata
LeC
on
te(C
era
mb
yci
da
e)
TX
Ho
vo
rea
nd
Pen
rose
19
82
Cer
am
by
cid
ae
Lio
pin
us
nr.
alp
ha
(Sa
y,
18
27
)[a
sL
eio
pus
nr.
alp
ha
(Sa
y)]
Ca
rya
gla
bra
(Mil
l.)
Sw
eet
[as
Hic
ori
ag
lab
raM
ill.
]
,6
.4cm
Tre
ek
ille
db
yS
coly
tus
qu
adri
spin
osu
sS
ay
(Sco
lyti
na
e)
NY
Bla
ckm
an
an
dS
tag
e1
92
4
Cer
am
by
cid
ae
Lio
pin
us
mim
etic
us
(Ca
sey,
18
91)
[as
Lei
op
us
ho
ust
on
iC
ase
ya
nd
Lei
op
us
tex
an
aC
ase
y]
Ass
oci
ate
dw
ith
twig
sg
ird
led
by
On
cid
eres
pu
stu
lata
LeC
on
tea
nd
O.
cin
gu
lata
tex
an
aH
orn
(Cer
am
byci
da
e)
Lin
sley
19
40
256 THE COLEOPTERISTS BULLETIN 63(3), 2009
Ap
pen
dix
A.
Co
nti
nu
ed.
Co
leo
pte
rafa
mil
yC
ole
op
tera
spec
ies
Pla
nt
spec
ies
Su
bst
rate
dia
met
erS
ub
stra
teo
rigin
Sta
teR
efer
ence
Cer
am
by
cid
ae
Lio
pin
us
mim
etic
us
(Case
y,
18
91
)[a
sS
tern
idiu
sm
imet
icu
s(C
ase
y)
an
dS
tern
idiu
ste
xa
nu
s(C
ase
y)]
Leu
caen
ap
ulv
eru
len
ta(S
chlt
dl.
)B
enth
.A
sso
ciate
dw
ith
twig
sg
ird
led
by
On
cid
eres
pu
stu
lata
LeC
on
te(C
era
mb
yci
da
e)
TX
Ho
vo
rea
nd
Pen
rose
19
82
Cer
am
by
cid
ae
Lio
pin
us
wil
tii
(Ho
rn,
18
80)
[as
Lei
op
us
wil
tii
Ho
rn]
Ass
oci
ate
dw
ith
twig
sg
ird
led
by
On
cid
eres
pu
stu
lata
LeC
on
tea
nd
O.
cin
gu
lata
tex
an
aH
orn
(Cer
am
by
cid
ae)
Lin
sley
19
40
Cer
am
by
cid
ae
Loch
ma
eocl
esco
rnu
tice
ps
corn
uti
cep
sS
cha
effe
r,1
90
6
Leu
caen
ap
ulv
eru
len
ta(S
chlt
dl.
)B
enth
.A
sso
ciate
dw
ith
twig
sg
ird
led
by
On
cid
eres
pu
stu
lata
LeC
on
te(C
era
mb
yci
da
e)
TX
Ho
vo
rea
nd
Pen
rose
19
82
Cer
am
by
cid
ae
Mo
lorc
hu
sb
imacu
latu
sS
ay,
18
24
Ca
rya
gla
bra
(Mil
l.)
Sw
eet
[as
Hic
ori
ag
lab
raM
ill.
]
,6
.4cm
Tre
ek
ille
db
yS
coly
tus
qu
adri
spin
osu
sS
ay
(Sco
lyti
na
e)
NY
Bla
ckm
an
an
dS
tage
19
24
Cer
am
by
cid
ae
Neo
cly
tus
acu
min
atu
s(F
ab
rici
us,
17
75)
Ca
rya
gla
bra
(Mil
l.)
Sw
eet
[as
Hic
ori
ag
lab
raM
ill.
]
,6
.4cm
Tre
ek
ille
db
yS
coly
tus
qu
adri
spin
osu
sS
ay
(Sco
lyti
na
e)
NY
Bla
ckm
an
an
dS
tage
19
24
Cer
am
by
cid
ae
Neo
cly
tus
leu
cozo
nus
Lap
ort
e&
Go
ry,
18
35
[as
Neo
cly
tus
lon
gip
es(K
irb
y)]
La
rix
lari
cin
a(D
uR
oi)
K.
Ko
ch.
2.5
–5
cmD
ead
tree
,u
nk
no
wn
cau
seN
YB
lack
ma
na
nd
Sta
ge
19
18
Cer
am
by
cid
ae
Neo
com
psa
excl
am
ati
on
is(T
ho
mso
n,
18
60
)[a
sIb
idio
nex
cla
ma
tio
nis
Th
om
son
]
Ass
oci
ate
dw
ith
twig
sg
ird
led
by
On
cid
eres
pu
stu
lata
LeC
on
te(C
era
mb
yci
da
e)
Lin
sley
19
40
THE COLEOPTERISTS BULLETIN 63(3), 2009 257
Ap
pen
dix
A.
Co
nti
nu
ed.
Co
leo
pte
rafa
mil
yC
ole
op
tera
spec
ies
Pla
nt
spec
ies
Su
bst
rate
dia
met
erS
ub
stra
teo
rigin
Sta
teR
efer
ence
Cer
am
by
cid
ae
Neo
com
psa
excl
am
ati
onis
(Th
om
son
,1
86
0)
Leu
caen
ap
ulv
eru
len
ta(S
chlt
dl.
)B
enth
.5
6m
mA
sso
cia
ted
wit
htw
igs
gir
dle
db
yO
nci
der
esp
ust
ula
taL
eCo
nte
(Cer
am
by
cid
ae)
TX
Ho
vo
rea
nd
Pen
rose
19
82
Cer
am
by
cid
ae
Neo
com
psa
mex
ica
na
(Th
om
son
,1
86
5)
[as
Ibid
ion
tow
nse
nd
iL
inel
l]
Ass
oci
ate
dw
ith
twig
sg
ird
led
by
On
cid
eres
pu
stu
lata
LeC
on
te(C
era
mb
yci
da
e)
Lin
sley
19
40
Cer
am
by
cid
ae
Neo
com
psa
mex
ica
na
(Th
om
son
,1
86
5)
Leu
caen
ap
ulv
eru
len
ta(S
chlt
dl.
)B
enth
.A
sso
cia
ted
wit
htw
igs
gir
dle
db
yO
nci
der
esp
ust
ula
taL
eCo
nte
(Cer
am
by
cid
ae)
TX
Ho
vo
rea
nd
Pen
rose
19
82
Cer
am
by
cid
ae
Ob
rium
ma
cula
tum
(Oli
vie
r,1
79
5)
Ass
oci
ate
dw
ith
twig
sg
ird
led
by
On
cid
eres
pu
stu
lata
LeC
on
tea
nd
O.
cin
gu
lata
tex
an
aH
orn
(Cer
am
by
cid
ae)
Lin
sley
19
40
Cer
am
by
cid
ae
Ob
rium
ma
cula
tum
(Oli
vie
r,1
79
5)
Leu
caen
ap
ulv
eru
len
ta(S
chlt
dl.
)B
enth
.A
sso
cia
ted
wit
htw
igs
gir
dle
db
yO
nci
der
esp
ust
ula
taL
eCo
nte
(Cer
am
by
cid
ae)
TX
Ho
vo
rea
nd
Pen
rose
19
82
Cer
am
by
cid
ae
Ob
rium
mo
zin
nae
Lin
ell,
18
97
Leu
caen
ap
ulv
eru
len
ta(S
chlt
dl.
)B
enth
.A
sso
cia
ted
wit
htw
igs
gir
dle
db
yO
nci
der
esp
ust
ula
taL
eCo
nte
(Cer
am
by
cid
ae)
TX
Ho
vo
rea
nd
Pen
rose
19
82
Cer
am
by
cid
ae
On
cid
eres
cin
gu
lata
tex
an
aH
orn
,1
88
5[a
sO
nci
der
este
xan
aH
orn
]
Aca
cia,
pro
ba
bly
als
oP
roso
pis
an
dE
ben
op
sis
[as
Pit
hec
olo
biu
m]
Liv
etw
igg
ird
led
by
pa
ren
tL
insl
ey1
94
0
258 THE COLEOPTERISTS BULLETIN 63(3), 2009
Ap
pen
dix
A.
Co
nti
nu
ed.
Co
leo
pte
rafa
mil
yC
ole
op
tera
spec
ies
Pla
nt
spec
ies
Su
bst
rate
dia
met
erS
ub
stra
teo
rig
inS
tate
Ref
eren
ce
Cer
am
by
cid
ae
On
cid
eres
pu
stu
lata
LeC
on
te,
18
54
Aca
cia
farn
esia
na
(L.)
Wil
ld.,
Eb
enop
sis
eba
no
(Ber
l.)
Barn
eby
&G
rim
es[a
sP
ith
ecolo
biu
mfl
exic
auli
s(B
enth
.)J.
M.
Co
ult
.],
Pro
sop
isg
lan
du
losa
To
rr.,
Aca
cia
ber
lan
die
riB
enth
.,P
ark
inso
nia
acu
lea
taL
.,M
imo
saa
cule
ati
carp
aO
rteg
a[a
sM
imo
sali
nd
hei
mer
iA
.G
ray
]
20
–4
0m
mL
ive
twig
gir
dle
db
yp
are
nt
Lin
sley
19
40
Cer
am
by
cid
ae
On
cid
eres
pu
stu
lata
LeC
on
te,
18
54
Leu
caen
ap
ulv
erule
nta
(Sch
ltd
l.)
Ben
th.
Liv
etw
igg
ird
led
by
pa
ren
tT
XH
ov
ore
an
dP
enro
se1
98
2C
eram
by
cid
ae
On
cid
eres
rho
do
stic
taB
ate
s,1
88
5P
roso
pis
gla
nd
ulo
saT
orr
.v
ar.
gla
ndu
losa
Av
era
ge
9.1
mm
Liv
etw
igg
ird
led
by
pa
ren
tT
XP
olk
an
dU
eck
ert
19
73
Cer
am
by
cid
ae
Pla
cost
ern
us
dif
fici
lis
(Ch
evro
lat,
18
62)
Ass
oci
ate
dw
ith
twig
sg
ird
led
by
On
cid
eres
pu
stu
lata
LeC
on
te(C
era
mb
yci
da
e)
Lin
sley
19
40
Cer
am
by
cid
ae
Pog
on
och
eru
sm
ixtu
sH
ald
ema
n,
18
47
Lari
xla
rici
na
(Du
Ro
i)K
.K
och
.2
.5–
5cm
Dea
dtr
ee,
un
kn
ow
nca
use
NY
Bla
ckm
an
an
dS
tag
e1
91
8C
eram
by
cid
ae
Psy
rass
au
nic
olo
r(R
an
da
ll,
18
38)
[as
Pse
ud
ibid
ion
un
ico
lor
(Ra
nd
all
)]
Ca
rya
gla
bra
(Mil
l.)
Sw
eet
[as
Hic
ori
ag
lab
raM
ill.
]
,6
.4cm
Tre
ek
ille
db
yS
coly
tus
qu
adri
spin
osu
sS
ay
(Sco
lyti
na
e)
NY
Bla
ckm
an
an
dS
tag
e1
92
4
Cer
am
by
cid
ae
Purp
uri
cen
us
ax
illa
ris
Ha
ldem
an
,1
84
7C
ary
ag
lab
ra(M
ill.
)S
wee
t[a
sH
ico
ria
gla
bra
Mil
l.]
,6
.4cm
Tre
ek
ille
db
yS
coly
tus
qu
adri
spin
osu
sS
ay
(Sco
lyti
na
e)
NY
Bla
ckm
an
an
dS
tag
e1
92
4
THE COLEOPTERISTS BULLETIN 63(3), 2009 259
Ap
pen
dix
A.
Co
nti
nu
ed.
Co
leo
pte
rafa
mil
yC
ole
op
tera
spec
ies
Pla
nt
spec
ies
Su
bst
rate
dia
met
erS
ub
stra
teo
rigin
Sta
teR
efer
ence
Cer
am
by
cid
ae
Sa
per
da
dis
coid
eaF
ab
rici
us,
17
98
Ca
rya
gla
bra
(Mil
l.)
Sw
eet
[as
Hic
ori
ag
lab
raM
ill.
]
,6
.4cm
Tre
ek
ille
db
yS
coly
tus
qu
adri
spin
osu
sS
ay
(Sco
lyti
na
e)
NY
Bla
ckm
an
an
dS
tage
19
24
Cer
am
by
cid
ae
Sp
ha
eno
thec
us
biv
itta
taD
up
on
t,1
83
8
Ass
oci
ate
dw
ith
twig
sg
ird
led
by
On
cid
eres
cin
gu
lata
tex
an
aH
orn
(Cer
am
by
cid
ae)
Lin
sley
19
40
Cer
am
by
cid
ae
Sp
ha
eno
thec
us
biv
itta
taD
up
on
t,1
83
8[a
sT
ara
no
mis
biv
itta
tab
ivit
tata
(Du
po
nt)
]
Leu
caen
ap
ulv
eru
len
ta(S
chlt
dl.
)B
enth
.A
sso
cia
ted
wit
htw
igs
gir
dle
db
yO
nci
der
esp
ust
ula
taL
eCo
nte
(Cer
am
by
cid
ae)
TX
Ho
vo
rea
nd
Pen
rose
19
82
Cer
am
by
cid
ae
Ste
no
sph
enu
slu
gen
sL
eCo
nte
,1
86
2L
euca
ena
pu
lver
ule
nta
(Sch
ltd
l.)
Ben
th.
Ass
oci
ate
dw
ith
twig
sg
ird
led
by
On
cid
eres
pu
stu
lata
LeC
on
te(C
era
mb
yci
da
e)
TX
Ho
vo
rea
nd
Pen
rose
19
82
Cer
am
by
cid
ae
Ste
no
sph
enu
sn
ota
tus
(Oli
vie
r,1
79
5)
Ca
rya
gla
bra
(Mil
l.)
Sw
eet
[as
Hic
ori
ag
lab
raM
ill.
]
,6
.4cm
Tre
ek
ille
db
yS
coly
tus
qu
adri
spin
osu
sS
ay
(Sco
lyti
na
e)
NY
Bla
ckm
an
an
dS
tage
19
24
Cer
am
by
cid
ae
Th
rya
llis
un
datu
s(C
hev
rola
t,1
83
4)
Leu
caen
ap
ulv
eru
len
ta(S
chlt
dl.
)B
enth
.A
sso
cia
ted
wit
htw
igs
gir
dle
db
yO
nci
der
esp
ust
ula
taL
eCo
nte
(Cer
am
by
cid
ae)
TX
Ho
vo
rea
nd
Pen
rose
19
82
Cer
am
by
cid
ae
Tra
chyd
eres
ma
ndib
ula
ris
(Du
po
nt
inA
ud
inet
-Ser
vil
le,
18
34)
[as
Den
dro
bia
sm
an
dib
ula
ris
(Au
din
et-S
erv
ille
)]
Leu
caen
ap
ulv
eru
len
ta(S
chlt
dl.
)B
enth
.A
sso
cia
ted
wit
htw
igs
gir
dle
db
yO
nci
der
esp
ust
ula
taL
eCo
nte
(Cer
am
by
cid
ae)
TX
Ho
vo
rea
nd
Pen
rose
19
82
Cer
am
by
cid
ae
Urg
lep
tes
celt
is(S
cha
effe
r,1
90
5)
Leu
caen
ap
ulv
eru
len
ta(S
chlt
dl.
)B
enth
.A
sso
cia
ted
wit
htw
igs
gir
dle
db
yO
nci
der
esp
ust
ula
taL
eCo
nte
(Cer
am
by
cid
ae)
TX
Ho
vo
rea
nd
Pen
rose
19
82
260 THE COLEOPTERISTS BULLETIN 63(3), 2009
Ap
pen
dix
A.
Co
nti
nu
ed.
Co
leo
pte
rafa
mil
yC
ole
op
tera
spec
ies
Pla
nt
spec
ies
Su
bst
rate
dia
met
erS
ub
stra
teo
rigin
Sta
teR
efer
ence
Cer
am
by
cid
ae
Urg
lep
tes
qu
erci
(Fit
ch,
18
58)
[as
Lep
turg
esq
uer
ci(F
itch
)]
Ca
rya
gla
bra
(Mil
l.)
Sw
eet
[as
Hic
ori
ag
lab
raM
ill.
]
,6
.4cm
Tre
ek
ille
db
yS
coly
tus
qu
ad
risp
ino
sus
Sa
y(S
coly
tin
ae)
NY
Bla
ckm
an
an
dS
tage
19
24
Cer
am
by
cid
ae
Xy
lotr
ech
us
colo
nu
s(F
ab
rici
us,
17
75)
Ca
rya
gla
bra
(Mil
l.)
Sw
eet
[as
Hic
ori
ag
lab
raM
ill.
]
,6
.4cm
Tre
ek
ille
db
yS
coly
tus
qu
ad
risp
ino
sus
Sa
y(S
coly
tin
ae)
NY
Bla
ckm
an
an
dS
tage
19
24
Cle
rid
ae
Ch
ari
essa
pil
osa
(Fo
rste
r,1
77
1)
Ca
rya
gla
bra
(Mil
l.)
Sw
eet
[as
Hic
ori
ag
lab
raM
ill.
]
,6
.4cm
Tre
ek
ille
db
yS
coly
tus
qu
ad
risp
ino
sus
Sa
y(S
coly
tin
ae)
NY
Bla
ckm
an
an
dS
tage
19
24
Cle
rid
ae
Cy
ma
tod
era
ino
rna
ta(S
ay,
18
35)
Ca
rya
gla
bra
(Mil
l.)
Sw
eet
[as
Hic
ori
ag
lab
raM
ill.
]
,6
.4cm
Tre
ek
ille
db
yS
coly
tus
qu
ad
risp
ino
sus
Sa
y(S
coly
tin
ae)
NY
Bla
ckm
an
an
dS
tage
19
24
Cle
rid
ae
En
ocl
eru
sq
ua
dri
sig
na
tus
(Sa
y,
18
35)
Leu
caen
ap
ulv
eru
len
ta(S
chlt
dl.
)B
enth
.A
sso
cia
ted
wit
htw
igs
gir
dle
db
yO
nci
der
esp
ust
ula
taL
eCo
nte
(Cer
am
by
cid
ae)
TX
Ho
vo
rea
nd
Pen
rose
19
82
Cle
rid
ae
En
ocl
eru
ssp
.P
roso
pis
gla
ndu
losa
To
rr.
va
r.g
lan
du
losa
Av
era
ge
9.1
mm
Ass
oci
ate
dw
ith
twig
sg
ird
led
by
On
cid
eres
rho
do
stic
taB
ate
s(C
era
mb
yci
da
e)
TX
Po
lka
nd
Uec
ker
t1
97
3
Cle
rid
ae
Ma
do
nie
lla
dis
loca
tus
(Sa
y,
18
25)
[as
Phy
llo
ba
enu
sd
islo
catu
s(S
ay)]
La
rix
lari
cin
a(D
uR
oi)
K.
Ko
ch.
2.5
–5
cmD
ead
tree
,u
nk
no
wn
cau
seN
YB
lack
man
an
dS
tage
19
18
Cle
rid
ae
Ma
do
nie
lla
dis
loca
tus
(Sa
y,
18
25)
[as
Phy
llo
ba
enu
sd
islo
catu
s(S
ay)]
Ca
rya
gla
bra
(Mil
l.)
Sw
eet
[as
Hic
ori
ag
lab
raM
ill.
]
,6
.4cm
Tre
ek
ille
db
yS
coly
tus
qu
ad
risp
ino
sus
Sa
y(S
coly
tin
ae)
NY
Bla
ckm
an
an
dS
tage
19
24
Cle
rid
ae
Mo
no
ph
yll
ap
all
ipes
Sch
aef
fer,
19
08
Leu
caen
ap
ulv
eru
len
ta(S
chlt
dl.
)B
enth
.A
sso
cia
ted
wit
htw
igs
gir
dle
db
yO
nci
der
esp
ust
ula
taL
eCo
nte
(Cer
am
by
cid
ae)
TX
Ho
vo
rea
nd
Pen
rose
19
82
THE COLEOPTERISTS BULLETIN 63(3), 2009 261
Ap
pen
dix
A.
Co
nti
nu
ed.
Co
leo
pte
rafa
mil
yC
ole
op
tera
spec
ies
Pla
nt
spec
ies
Su
bst
rate
dia
met
erS
ub
stra
teo
rigin
Sta
teR
efer
ence
Cle
rid
ae
Mo
no
ph
yll
ate
rmin
ata
(Sa
y,
18
35)
Ca
rya
gla
bra
(Mil
l.)
Sw
eet
[as
Hic
ori
ag
lab
raM
ill.
]
,6
.4cm
Tre
ek
ille
db
yS
coly
tus
qu
adri
spin
osu
sS
ay
(Sco
lyti
na
e)
NY
Bla
ckm
an
an
dS
tage
19
24
Cle
rid
ae
Py
tice
roid
esla
tico
rnis
(Sa
y,
18
35)
[as
Nei
chn
eala
tico
rnis
(Sa
y)]
Ca
rya
gla
bra
(Mil
l.)
Sw
eet
[as
Hic
ori
ag
lab
raM
ill.
]
,6
.4cm
Tre
ek
ille
db
yS
coly
tus
qu
adri
spin
osu
sS
ay
(Sco
lyti
na
e)
NY
Bla
ckm
an
an
dS
tage
19
24
Cu
rcu
lio
nid
ae
Ch
ram
esu
sh
ico
ria
eL
eCo
nte
,1
86
8C
ary
ag
lab
ra(M
ill.
)S
wee
t[a
sH
ico
ria
gla
bra
Mil
l.]
,6
.4cm
Tre
ek
ille
db
yS
coly
tus
qu
adri
spin
osu
sS
ay
(Sco
lyti
na
e)
NY
Bla
ckm
an
an
dS
tage
19
24
Cu
rcu
lio
nid
ae
Co
ph
esfa
lla
x(L
eCo
nte
,1
87
6)
[as
Cry
pto
rhy
nch
us
fall
ax
LeC
on
te]
Ca
rya
gla
bra
(Mil
l.)
Sw
eet
[as
Hic
ori
ag
lab
raM
ill.
]
,6
.4cm
Tre
ek
ille
db
yS
coly
tus
qu
adri
spin
osu
sS
ay
(Sco
lyti
na
e)
NY
Bla
ckm
an
an
dS
tage
19
24
Cu
rcu
lio
nid
ae
Hy
locu
rus
rud
is(L
eCo
nte
,1
87
6)
[as
Hy
locu
rus
bio
rbis
Bla
ckm
an
]
Ca
rya
gla
bra
(Mil
l.)
Sw
eet
[as
Hic
ori
ag
lab
raM
ill.
]
,6
.4cm
Tre
ek
ille
db
yS
coly
tus
qu
adri
spin
osu
sS
ay
(Sco
lyti
na
e)
NY
Bla
ckm
an
an
dS
tage
19
24
Cu
rcu
lio
nid
ae
Ma
gd
ali
so
lyra
(Her
bst
,1
79
7)
Ca
rya
gla
bra
(Mil
l.)
Sw
eet
[as
Hic
ori
ag
lab
raM
ill.
]
,6
.4cm
Tre
ek
ille
db
yS
coly
tus
qu
adri
spin
osu
sS
ay
(Sco
lyti
na
e)
NY
Bla
ckm
an
an
dS
tage
19
24
Cu
rcu
lio
nid
ae
Ma
gd
ali
sp
an
du
ra(S
ay,
18
31)
Ca
rya
gla
bra
(Mil
l.)
Sw
eet
[as
Hic
ori
ag
lab
raM
ill.
]
,6
.4cm
Tre
ek
ille
db
yS
coly
tus
qu
adri
spin
osu
sS
ay
(Sco
lyti
na
e)
NY
Bla
ckm
an
an
dS
tage
19
24
Cu
rcu
lio
nid
ae
Po
lyg
rap
hu
sru
fip
enn
is(K
irb
y,
18
37)
La
rix
lari
cin
a(D
uR
oi)
K.
Ko
ch.
2.5
–5
cmD
ead
tree
,u
nk
no
wn
cau
seN
YB
lack
ma
na
nd
Sta
ge
19
18
Cu
rcu
lio
nid
ae
Sci
aph
ilu
sa
sper
atu
s(B
on
sdo
rff,
17
85
)[a
sS
cia
ph
ilu
sm
uri
catu
s(F
ab
rici
us)
]Ca
rya
gla
bra
(Mil
l.)
Sw
eet
[as
Hic
ori
ag
lab
raM
ill.
]
,6
.4cm
Tre
ek
ille
db
yS
coly
tus
qu
adri
spin
osu
sS
ay
(Sco
lyti
na
e)
NY
Bla
ckm
an
an
dS
tag
e1
92
4
262 THE COLEOPTERISTS BULLETIN 63(3), 2009
Co
leo
pte
rafa
mil
yC
ole
op
tera
spec
ies
Pla
nt
spec
ies
Su
bst
rate
dia
met
erS
ub
stra
teo
rig
inS
tate
Ref
eren
ce
Cu
rcu
lio
nid
ae
Sco
lytu
sp
icea
e(S
wa
ine,
19
10
)[a
sE
ccop
tog
ast
erp
icea
eS
wa
ine]
La
rix
lari
cin
a(D
uR
oi)
K.
Ko
ch.
2.5
–5
cmD
ead
tree
,u
nk
no
wn
cau
seN
YB
lack
man
an
dS
tage
19
18
Cu
rcu
lio
nid
ae
Sco
lytu
sq
ua
dri
spin
osu
sS
ay,
18
24
[as
Ecc
op
tog
ast
erq
ua
dri
spin
osu
s(S
ay
)]
Ca
rya
gla
bra
(Mil
l.)
Sw
eet
[as
Hic
ori
ag
lab
raM
ill.
]
,6
.4cm
NY
Bla
ckm
an
an
dS
tage
19
24
Mel
an
dry
ida
eO
rch
esia
cast
an
eaM
elsh
eim
er,
18
46
Ca
rya
gla
bra
(Mil
l.)
Sw
eet
[as
Hic
ori
ag
lab
raM
ill.
]
,6
.4cm
Tre
ek
ille
db
yS
coly
tus
qu
ad
risp
ino
sus
Sa
y(S
coly
tin
ae)
NY
Bla
ckm
an
an
dS
tage
19
24
Mo
rdel
lid
ae
1sp
.L
euca
ena
pu
lver
ule
nta
(Sch
ltd
l.)
Ben
th.
Ass
oci
ate
dw
ith
twig
sg
ird
led
by
On
cid
eres
pu
stu
lata
LeC
on
te(C
era
mb
yci
da
e)
TX
Ho
vo
rea
nd
Pen
rose
19
82
Ten
ebri
on
ida
e1
sp.
Leu
caen
ap
ulv
eru
len
ta(S
chlt
dl.
)B
enth
.A
sso
cia
ted
wit
htw
igs
gir
dle
db
yO
nci
der
esp
ust
ula
taL
eCo
nte
(Cer
am
byci
da
e)
TX
Ho
vo
rea
nd
Pen
rose
19
82
Tro
go
ssit
idae
Tem
nosc
hei
lasp
.L
euca
ena
pu
lver
ule
nta
(Sch
ltd
l.)
Ben
th.
Ass
oci
ate
dw
ith
twig
sg
ird
led
by
On
cid
eres
pu
stu
lata
LeC
on
te(C
era
mb
yci
da
e)
TX
Ho
vo
rea
nd
Pen
rose
19
82
Ap
pen
dix
A.
Co
nti
nu
ed.
THE COLEOPTERISTS BULLETIN 63(3), 2009 263