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Habitat Associations Saproxylic Beetles SE USA_08-09-P[1]

Habitat Associations Saproxylic Beetles SE USA_08-09-P[1]

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A research paper from 2008 providing detailed evidence of the importance of retaining entire standing dead trees over stumps or logs when it comes to the deadwood habitat required by many saproxylic beetles and the higher fauna that rely on them as a direct and indirect food source.
A research paper from 2008 providing detailed evidence of the importance of retaining entire standing dead trees over stumps or logs when it comes to the deadwood habitat required by many saproxylic beetles and the higher fauna that rely on them as a direct and indirect food source.

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Categories:Types, Research, Science
Published by: Veteran Tree Group Australia on Nov 21, 2010
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Habitat associations of saproxylic beetles in the southeastern UnitedStates: A comparison of forest types, tree species and wood postures
Michael D. Ulyshen*, James L. Hanula
USDA Forest Service, 320 Green Street, Athens, GA 30602, United States
1. Introduction
Althoughitisnowwidelyrecognizedthatsaproxylicbeetlesarehighly sensitive to long-term losses of dead wood, virtuallynothingisknownaboutthestatusofthisdiversecommunityintheintensively managed forests of the southeastern United States.However, this region faces a number of continuing (e.g., timberharvesting), intensifying (e.g., urbanization and habitat fragmen-tation), and emerging (e.g. biofuel production) threats tosaproxylic organisms (Wear, 1996; Harding, 2007). The south-easternUnitedStatescontains40%ofthecountry’stimberlandand90% of the forests within the region are controlled by privatelandowners (Wear, 1996, and references therein). Around 22% of the land held by private landowners is managed intensively(typically on 30–50 yr rotations) for timber production by forestindustries (Wear, 1996). The remainder is managed variously by awide variety of landowners (Wear, 1996). Conserving saproxylicbeetlesandotherorganismsinthiscomplexlandscapewillrequiregreat care and understanding. Unfortunately, too little is knownabout the basic life histories and habitat requirements of mostspecies to prioritize actions or to make informed decisions. Hereweinvestigatethehabitatassociationsofsaproxylicbeetlesontheupper coastal plain of the southeastern United States. The mainfactors of interest are summarized below.
Forest type. The coastal plain of the southeastern United Statesis dominated by pines on relatively dry upland sites and bymixed hardwoods on mesic bottomland sites. The relativeimportance of these two main forest types to saproxylic beetlesremains unknown. Upland pine forests are more extensive thanbottomland hardwood forests throughout the region. However,bottomlandhardwoodforestssupportmorediversetreespecies
Forest Ecology and Management 257 (2009) 653–664
 Article history:
Received 29 April 2008
Received in revised form 4 August 2008
Accepted 28 September 2008
BiodiversityCoarse woody debrisDead woodForest canopyResource partitioningVertical stratification
Saproxylic beetles are highly sensitive to forest management practices that reduce the abundance andvarietyofdeadwood.However,thisdiversefaunacontinuestoreceivelittleattentioninthesoutheasternUnited States even though this region supports some of the most diverse, productive and intensivelymanagedforestsinNorthAmerica.Inthisreplicatedthree-wayfactorialexperiment,weinvestigatedthehabitat associations of saproxylic beetles on the coastal plain of South Carolina. The factors of interestwere forest type (upland pine-dominated vs. bottomland hardwood), tree species (
Quercus nigra
L.)andwoodposture(standinganddowneddeadwood,i.e.,snagsandlogs).Woodsamplesweretakenatfourpositionsalongeachlogandsnag(lowerbole,middlebole,upperbole and crown)
11 months after the trees were killed and placed in rearing bags to collect emergingbeetles. Overall, 33,457 specimens from 52 families and
250 species emerged. Based on an analysis of covariance, with surface area and bark coverage as covariates, saproxylic beetle species richness differedsignificantly between forest types as well as between wood postures. There were no significantinteractions. Species richness was significantly higher in the upland pine-dominated stand than thebottomland hardwood forest, possibly due to higher light exposure and temperature in upland forests.Although
L. styraciflua
yielded more beetle species (152) than either
Q. nigra
(122) or
P. taeda
(125),therewere no significant differences in species richness among tree species. There were also no relationshipsevident between relative tree abundance and observed or expected beetle species richness. Significantlymore beetle species emerged from logs than from snags. However snags had a distinct fauna includingseveral potential canopy specialists. Our results suggest that conservation practices that retain or createentire snags as opposed to high stumps or logs alone will most greatly benefit saproxylic beetles insoutheastern forests.Published by Elsevier B.V.
* Corresponding author.
E-mail address:
mulyshen@hotmail.com(M.D. Ulyshen).
Contents lists available atScienceDirect
Forest Ecology and Management
journal homepage: www.elsevier.com/locate/foreco
0378-1127/$ – see front matter. Published by Elsevier B.V.doi:10.1016/j.foreco.2008.09.047
DE- AI09-00SR22188 JOURNAL ARTICLE 2008 08-09-P
assemblages and may therefore be disproportionately impor-tant to the saproxylic beetle fauna. A number of factors likely todiffer between forest types, such as canopy coverage (Økland,2002), light exposure (Lindhe et al., 2005) and humidity (Warriner et al., 2004), may also have important consequencesfor the structure and species richness of saproxylic beetlecommunities. In this study we sampled saproxylic beetles inbothanuplandandbottomlandforest.Wepredictedthatoverallbeetlerichnesswouldbehigherinthebottomlandforestthaninthe upland forest due to the higher diversity of tree species inbottomland forests.
Tree species. We sampled wood from three tree species, each of which differed in abundance between upland and bottomlandforests, to evaluate the effects of relative tree abundance on thediversity and composition of saproxylic beetles. This questionhas particularly important implications for saproxylic beetleconservation, but remains largely unstudied. We predicted asignificant interaction between tree species and forest type dueto differences in relative tree species abundances between thetwo forest types.
Wood posture. A large volume and variety of resources areavailable to saproxylic insects above the ground in the form of standing dead trees (i.e., snags), dead branches and twigs, androttingheartwood(FonteandSchowalter,2004).Forexample,ina temperate broadleaved forest in Sweden,Norde´n et al. (2004)found snags made up about 22% of total dead wood volume andanother 6% was attributed to dead branches attached to livingtrees. Standing or suspended dead wood is generally drier anddecays more slowly than wood in contact with the ground( Jomura et al., 2008), possibly reducing the abundance anddiversity of insects present (Larkin and Elbourn, 1964). Severalstudies from Europe support this notion ( Jonsell and Weslien,2003;Gibbetal.,2006;McGeochetal.,2007;Hja¨lte´netal.,2007;Franc, 2007). However, many threatened species and otherinsects appear to favor snags ( Jonsell et al., 1998; Sverdrup-Thygeson and Ims, 2002; Kappes and Topp, 2004; Hedgren andSchroeder, 2004).Unfortunately,previouseffortstosample fromsnags have generally limited sampling to within a few meters of the ground. Until the upper reaches of snags are adequatelysampled, it will be impossible to reach definite conclusionsregarding the relative importance of snags and logs. Here wecompare the beetle communities inhabiting snags and logs frombase to crown to better understand the relative importance of these two habitats in southeastern forests. We predicted thatoverall species richness would be higher in logs than in snagsbased on previous research and on the idea that the upper bolesections and crowns of snags would be less accessible andtherefore less readily colonized than those of logs.
2. Methods
 2.1. Study site
This research took place on the 80,267-ha Savannah River Site(SRS) located in the upper Coastal Plain Physiographic Province of South Carolina. The SRS, a facility owned and operated by theUnited States Department of Energy, was established in 1951, andwas designated a National Environmental Research Park in 1972(Kilgo and Blake, 2005). Most of the land now owned by theSavannah River site was formerly used for agricultural purposesandmostforestscurrentlystandingwereplantedorregeneratedinthe early 1950s (Kilgo and Blake, 2005).TheSRSissomewhattypicalofthesoutheasterncoastalplaininthatitisdominated(68%)bypineforestsgrowingonrelativelydryupland sites and by mixed hardwoods (22%) occupying swampsand riparian bottomlands (Kilgo and Blake, 2005). However, theupland and bottomland sites do not consist purely of pines andhardwoods, respectively. At least three tree species are relativelycommon in both forest types. Sweetgum (
Liquidambar styraciflua
Quercus nigra
L.)growmostcommonlyonmesicsites dominated by mixed hardwoods but also appear sporadicallyamong pines on dry upland sites. Similarly, loblolly pine (
L.) is currently the dominant pine species growing in uplandpine forests but was historically restricted to moist bottomlandsites (Schultz, 1997) and continues to grow there at low densities.Kilgo and Blake (2005)provide percent basal areas for tree speciesindifferentforesttypesontheSavannahRiverSite.Forashortleaf-loblolly pine slope, comparable to the upland forest used in thisstudy,
madeup80%, 2% and 1% of the total basal area, respectively. In contrast, theaverage percent basal areas in bottomland forests bordering riversand large streams for
P. taeda
L. styraciflua
Q. nigra
were 2.2%,10.6% and 3.5%, respectively (Kilgo and Blake, 2005).The upland and bottomland forests used in this study wereapproximately 25 km apart. One Hobo Data Logger was placed ineachforesttypeforapproximatelyoneyear(2006–2007)torecordtemperature and humidity. On average, the upland forest waswarmerthanthebottomlandforest(18.8and17.8
C,respectively)whereas relative humidity was on average lower there than in thebottomlandforest(72.2and76.6%,respectively).Thesedifferenceswere most pronounced during the growing season (Fig. 1).
 2.2. Experimental design
Our sampling followed a 2
2 factorial design with therespective factors being forest type (upland pine forest vs. bottom-land hardwood forest), tree species (
L. styraciflua
P. taeda
), and posture (log vs. snag). There were three replicates.On June 5–6, 2006, we created 9 snags and 9 logs in the uplandsitesandthe samenumber inthe bottomland sites,equally dividedamong
(i.e.,threesnagsandlogsof eachspeciesateachsite).Snagswerecreatedbygirdlingthetreestoadepthof3 cmormoreusingachainsawandsprayingfullstrength(53.8%)glyphosate(Foresters’
,Riverdale ChemicalCompany,BurrRidge, IL, USA) into the wounds. To prevent the herbicide fromtraveling up the tree and possibly affecting insect colonization, asecond girdle was created about 15cm above the first beforeherbicidewasapplied.Onlythelowergirdlewastreated.Allgirdledtrees examined two weeks after treatment were dead.
Fig. 1.
Temperature and relative humidity over time for a bottomland hardwoodforest and an upland pine-dominated stand in South Carolina, USA.
M.D. Ulyshen, J.L. Hanula/Forest Ecology and Management 257 (2009) 653–664
Approximately 11 months later, in May 2007, we returned tocollect sections from the three logs and snags of each species ateachsite.Afterfellingthesnagswithchainsaws,weremoved0.5 msections from the lower bole, middle bole, and upper bole of eachsnag and log. The position of each section was measured from thetree base (Appendix A). We also collected three 0.5 m crownsections taken from major limbs or sometimes the upper-mostportion of the main bole. The tops of all but one of the sweetgumsnagshadbroken,sothosecrownsectionshadbeenincontactwiththegroundforanunknownlengthoftime.Theupperbolesectionsfrom these trees were taken directly below the point of breakage.Alltheothersnagswereintact.Allboleandcrownsectionscutonagiven day (May 3 and 8 for upland and bottomland forests,respectively) were labeled and transported to Athens, Georgia.We recorded the diameter (measured at the center) and barkcoverage (visual estimation) of each bole and crown section(AppendixA)inthelaboratory.Weusedthesedatatocalculatethetotal surface area (not including ends) and bark surface area(product of surface area and visual estimate of bark coverage)sampled from each snag and log.
 2.3. Insect rearing 
Emergingbeetleswerecollectedinthelaboratoryusingrearingbags. Rearingbags havebeenshown to beoneof the mostefficientmethods for collecting saproxylic beetles from dead wood ( Jonselland Hansson, 2007). Bole (108) and crown sections (36) (i.e., thethree branch sections from each tree were tied together) weresuspended from wooden beams with synthetic rope and enclosedwithin large (170 l) extra-strength black plastic trash bags. In onebottomcornerofeachbagwe attacheda clearplasticcollectingjarcontaining propylene glycol. To prevent mold problems, wecontinuously ventilated the bags using an electric blower(HADP9-1 Cast Aluminum Pressure Blower, Americraft Manufac-turing Co., Cincinnati, OH, USA). Air from the blower flowedthrough a plastic PVC pipe (
10 cm in diameter) that ran thelength of the rearing facility near the ceiling. Each side of the pipehad rows of holes into which were inserted sections of clear vinyltubing (0.95 cm o.d., 0.64 cm i.d.). Each section of tubing led fromthe pipe to one of the rearing bags. The bags became inflated withair, thus forming effective funnels. Excess air escaped through asingle small hole (
2 mm) drilled near the top of each collecting jar.Overheadfluorescentlightswereleftonatalltimes.Wedidnotattempt to control temperature or humidity in the rearing facility,but all samples experienced the same conditions. Screenedwindows were opened along both sides of the facility to allowfor air movement and to match ambient conditions as closely aspossible. However, it was typically warmer inside the facility thanoutside. Samples were collected about once a month for 20 weeks(4 May–21 September and 9 May–26 September for upland andbottomland samples, respectively) and transferred to 70% ethanol.Beetles were identified using the classification system of ArnettandThomas(2001,2002).Voucherspecimenshavebeendepositedin the Georgia Museum of Natural History, Athens, Georgia.
 2.4. Data analysis
To test whether there were any differences in the amount of surface area sampled, we conducted a three-way analysis of variance with total surface area sampled (summed for each log orsnag) as the response variable. The analysis was repeated for totalbark surface area sampled.Bole and crown samples from each snag or log were combinedbefore conducting an analysis of covariance on a three-wayfactorialdesign(SASInstitute,1990).Surfaceareaandbarksurfaceareawerethecovariatesandthemaineffectswereforesttype,treespeciesand woodposture. Alleffects werefixed and there werenomissing or incomplete samples.Species richness estimates, based on the Chao1 estimator, werecalculated using EstimateS (Colwell, 2006). The Chao1 estimator iscalculated as follows: Chao1 =
+ (
) where
is theobserved species richness,
is the number of singletons and
isthenumberofdoubletons(ColwellandCoddington,1994).Thisisanappropriate estimator for this study given that Chao1 is thought toperform well on large datasets with large numbers of rare species(Colwell and Coddington, 1994, and references therein). Speciesrichnessestimatesare useful because,byfactoringinspeciesrarity,theygiveanindicationofhowthoroughlyanassemblage ofspecieshasbeensampled.Becauseitispossibleforobservedrichnesstrendstodifferfromestimatedrichnesstrends,itisusefultoexamineboth.Indicator species analysis (Dufreˆne and Legendre, 1997) wasperformed four times using PC-ORD (McCune and Mefford, 2006)to determine which species were significantly associated with (1)upland or bottomland forests; (2) snags or logs; (3) oak, pine orsweetgum; (4) lower bole, middle bole, upper bole or crown.Indicator values ranging from 0 (no association) to 100 (perfectassociation) were tested for statistical significance using a MonteCarlo randomization with 2500 permutations (McCune and Grace,2002).
3. Results and discussion
 3.1. Data set 
Overall, 33,457 specimens from 52 families and 250 ‘‘species’’emergedoverthe20wksamplingperiod(AppendixB).Aneffortwasmade to identify all specimens to the lowest taxonomic unitspossible given available time and expertise. All specimens wereidentified to family, 79% were identified to genus and 59% wereidentified to species. Several species rich groups (e.g., Ciidae,Corylophidae and Ptiliidae) were not sorted below family level andwere treated as single taxonomic units even though they likelyconsisted of multiple species. The estimates of species richnesspresented in this paper are therefore conservative. At least oneundescribedspecies,ahisteridbelongingtothegenus
,wascollected in this study (A. Tishechkin, personal communication).
 3.2. Surface area and bark surface area
Surfaceareadidnotvarysignificantlyforanyofthefactors(datanot shown). However, bark surface area varied significantly amongtreespecies(
= 31.30,
0.0001), being lowerfor
P. taeda
Therewasalsoasignificantinteractionbetween tree species and posture (
= 9.6,
= 0.0009) due to thefactthat
P. taeda
snags had considerablyless barkthan
P. taeda
0.12 and 1.22
0.14 m
, respectively).
 3.3. Species richness and habitat associations
Overall, species richness differed significantly between foresttypes and wood postures but not among tree species (Table 1).Because there were no significant interaction terms (Table 1), theresults for each factor are discussed individually below.
Forest type. In total, 189 and 175 beetle species were collectedfrom the upland and bottomland forests, respectively. Meanspecies richness was significantly higher in the upland forestthan the bottomland forest (Fig. 2A). We attribute this todifferences in light intensity and temperature between the twoforest types. The upland pine-dominated forest was more open
M.D. Ulyshen, J.L. Hanula/Forest Ecology and Management 257 (2009) 653–664

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