the coleopterists bulletin, 63(3):239–263. 2009. · [email protected] ... elevated /...

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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 Entomology Louisiana State University AgCenter 402 Life Sciences Building Baton Rouge, LA, 70803, U.S.A. [email protected] [email protected] KYLE E. HARMS Department of Biological Sciences Louisiana State University 202 Life Sciences Building Baton Rouge, LA, 70803, U.S.A. [email protected] AND CHRISTOPHER E. CARLTON Louisiana State Arthropod Museum, Department of Entomology Louisiana State University AgCenter 402 Life Sciences Building Baton Rouge, LA, 70803, U.S.A. [email protected] Abstract We conducted a study to explore which beetles utilize dead twigs in a Louisiana secondary forest and the effect of debris position on the beetle community. Twigs averaging 14 mm in diameter from one tree of Quercus falcata Michaux (southern red oak) were placed randomly 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 ground against the branch of a living woody plant. The bundles were collected 10 months later and each was placed into an emergence chamber. More than 400 adult Coleoptera specimens were 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 placed on the ground had the lowest richness, aboveground bundles had the highest, and propped bundles were intermediate. Twelve species (34%) were represented by singletons. To our knowledge, no general survey of the saproxylic beetles associated with fine woody debris has been undertaken in the United States. However, several studies have shown considerable beetle richness in dead branches and twigs. The exemplary research on saproxylic beetle succession conducted by Blackman and Stage (1918, 1924) showed that beetle assemblages in limbs less than 6.35 cm in diameter were distinct from those in larger portions of the dead tree and at times had greater species richness (up to 32 species). Mecke et al. (2001) reared 34 beetle The Coleopterists Bulletin, 63(3):239–263. 2009. 239

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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.

[email protected]

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

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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

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246 THE COLEOPTERISTS BULLETIN 63(3), 2009

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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.

Literature Cited

Beer, F. M. 1949. The rearing of Buprestidae and delayed emergence of their larvae. TheColeopterists’ Bulletin 3(6):81–84.

Blackman, M. W., and H. H. Stage. 1918. Notes on insects bred from the bark and wood ofthe American larch. The New York State College of Forestry, Technical PublicationNo. 10:11–115.

Blackman, M. W., and H. H. Stage. 1924. On the succession of insects living in the bark andwood of dying, dead, and decaying hickory. The New York State College ofForestry, Technical Publication No. 17:3–268.

Cramer, K. L. 1998. Effects of twig morphology on oviposition behavior and hatchingsuccess of the twig-girdling beetle Oncideres cingulata (Say) (Coleoptera: Ceramby-cidae). The Coleopterists Bulletin 52:186–193.

Darwin, F. (ed.). 1887. The life and letters of Charles Darwin, including an autobiographicalchapter. Volume 1. John Murray, London. 558 pp.

Elton, C. S. 1966. The Pattern of Animal Communities. Methuen & Co. Ltd., London.432 pp.

Fager, E. W. 1968. The community of invertebrates in decaying oak wood. Journal ofAnimal Ecology 37:121–142.

Forcella, F. 1981. Twig nitrogen content and larval survival of twig-girdling beetles,Oncideres cingulata (Say) (Coleoptera: Cerambycidae). The Coleopterists Bulletin35:211–212.

Forcella, F. 1984. Trees size and density affect twig-girdling intensity of Oncideres cingulata(Say) (Coleoptera: Cerambycidae). The Coleopterists Bulletin 38:37–42.

Hovore, F. T. 1980. A new genus and species of Cerambycidae from southern Texas(Coleoptera). The Coleopterists Bulletin 34:115–119.

Hovore, F. T., and R. L. Penrose. 1982. Notes on Cerambycidae co-inhabiting girdles ofOncideres pustulata LeConte (Coleoptera: Cerambycidae). The SouthwesternNaturalist 27:23–27.

Iwata, R., Y. Hirayama, H. Shimura, and M. Ueda. 2004. Twig foraging and soil-burrowingbehaviors in larvae of Dinoptera minuta (Gebler) (Coleoptera: Cerambycidae). TheColeopterists Bulletin 58:399–408.

Landau, D., and D. Prowell. 1999a. A partial checklist of moths from longleaf pine savannasin Louisiana (Insecta: Lepidoptera). Transactions of the American EntomologicalSociety 125:127–138.

Landau, D., and D. Prowell. 1999b. A partial checklist of moths collected from mixedmesophytic hardwood forests in Louisiana (Insecta: Lepidoptera). Transactions ofthe American Entomological Society 125:139–150.

Landau, D., D. Prowell, and C. E. Carlton. 1999. Intensive versus long-term sampling toassess lepidopteran diversity in a southern mixed mesophytic forest. ConservationBiology and Biodiversity 92:435–441.

Linsley, E. G. 1940. Notes on Oncideres twig girdlers. Journal of Economic Entomology33:561–563.

MacArthur, R. H. 1971. Patterns of terrestrial bird communities [pp. 189–221]. In: AvianBiology, volume 1 (D. S. Farner and J. R. King, editors). Academic Press, NewYork. 586 pp.

Manly, B. F. J. 2007. Randomization, Bootstrap and Monte Carlo Methods in Biology.Third Edition. Chapman and Hall/CRC, New York. 455 pp.

250 THE COLEOPTERISTS BULLETIN 63(3), 2009

Mecke, R., M. H. M. Galileo, and W. Engels. 2001. New records of insects associated withAraucaria trees: phytophagous Coleoptera and Hymenoptera and their naturalenemies. Studies on Neotropical Fauna and Environment 36:113–124.

Petrice, T. R., and R. A. Haack. 2006. Effects of cutting date, outdoor storage conditions,and splitting on survival of Agrilus planipennis (Coleoptera: Buprestidae) in firewoodlogs. Journal of Economic Entomology 99:790–796.

Polk, K. L., and D. N. Ueckert. 1973. Biology and ecology of a mesquite twig girdler,Oncideres rhodosticta, in west Texas. Annals of the Entomological Society ofAmerica 66:411–417.

Pollock, D. A. 1988. A technique for rearing subcortical Coleoptera larvae. TheColeopterists Bulletin 42:311–312.

Prado, P. I., and T. M. Lewinsohn. 2004. Compartments in insect–plant associations andtheir consequences for community structure. Journal of Animal Ecology73:1168–1178.

Prowell, D. 2001. Additions and corrections to Macrolepidoptera in Landau and Prowell1999 (a & b), partial checklists of moths from longleaf pine savannas and mesophytichardwood forests in Louisiana. Transactions of the American Entomological Society127:239–244.

Rice, M. E. 1989. Branch girdling and oviposition biology of Oncideres pustulatus(Coleoptera: Cerambycidae) on Acacia farnesiana. Annals of the EntomologicalSociety of America 82:181–186.

Rogers, C. E. 1977. Bionomics of Oncideres cingulata (Coleoptera: Cerambycidae) onMesquite. Journal of the Kansas Entomological Society 50:222–228.

Sanborn, C. E. 1911. The twig girdler. Bulletin of the Oklahoma Agricultural ExperimentStation 91:1–13.

Solomon, J. D., F. I. McCracken, R. L. Anderson, R. Lewis, Jr., F. L. Oliveria, T. H. Filer,and P. J. Barry. 1987. [slightly revised October 1999] Oak Pests: A Guide to MajorInsects, Diseases, Air Pollution, and Chemical Injury. Protection Report R8-PR7.USDA Forest Service, Southern Forest Experiment Station New Orleans. 69 pp.

Stehr, F. W. 1991. Immature Insects. Volume 2. Kendall/Hunt Publishing Company,Dubuque, IA. 975 pp.

Stevens, R. E., C. K. Lister, and J. L. Linnane. 1979. Outbreak of a twig beetle,Pityophthorus opaculus LeConte, in Colorado. The Coleopterists Bulletin 33:268.

Wang, Q., G. Shi, D. Song, D. J. Rogers, L. K. Davis, and X. Chen. 2002. Development,survival, body weight, longevity, and reproductive potential of Oemena [sic] hirta(Coleoptera: Cerambycidae) under different rearing conditions. Annals of theEntomological Society of America 95:563–569.

Wood, S. L. 1982. The bark and ambrosia beetles of North and Central America(Coleoptera: Scolytidae), a taxonomic monograph. Great Basin Naturalist Memoirs6:1–1359.

(Received 8 September 2008; accepted 21 April 2009. Publication date 28 September 2009.)

THE COLEOPTERISTS BULLETIN 63(3), 2009 251

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yS

coly

tus

qu

adri

spin

osu

sS

ay

(Sco

lyti

na

e)

NY

Bla

ckm

an

an

dS

tage

19

24

252 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

Bu

pre

stid

ae

An

tha

xia

qu

erca

ta(F

ab

rici

us,

18

01)

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

An

tha

xia

viri

dic

orn

is(S

ay,

18

23)

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

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