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Geocoris uliginosus (Say) (Hemiptera: Geocoridae): Associationwith Spanish Moss (Tillandsia usneoides) on Trees and ShrubsAuthor(s): A. G. Wheeler, Jr. and Juang-Horng ChongSource: Proceedings of the Entomological Society of Washington,114(3):424-429. 2012.Published By: Entomological Society of WashingtonDOI: http://dx.doi.org/10.4289/0013-8797.1143.424URL: http://www.bioone.org/doi/full/10.4289/0013-8797.1143.424

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PROC. ENTOMOL. SOC. WASH.

114(3), 2012, pp. 424–429

NOTE

Geocoris uliginosus (Say) (Hemiptera: Geocoridae): Association with SpanishMoss (Tillandsia usneoides) on Trees and Shrubs

Geocoris uliginosus (Say) is a com-mon North American bigeyed bug thatranges from southern Canada (New-foundland, Ontario, and Quebec) southto Florida and west to Colorado, NewMexico, and Texas (Ashlock and A. Slater1988, Maw et al. 2000). It also is knownfrom Cuba and North Caicos in the WestIndies (Baranowski and Slater 2005).

Adults are about 3.3 to 3.5 mm long,oval, and reddish-black to black. East oftheMississippi River, adults are uniformlypiceous to black, a color form that dif-fers from that of sympatric congeners(Hoffman 1996, Sweet 2000). Geocorisuliginosus is both a ground and plant-canopy inhabitant, although it is foundless frequently on vegetation than is thesometimes syntopic G. punctipes (Say)(Crocker and Whitcomb 1980, Readioand Sweet 1982, Sweet 2000). A spe-cies of temporary environments, it oc-curs mostly in disturbed habitats suchas roadsides and agroecosystems, withthe adults always fully winged or mac-ropterous (Readio and Sweet 1982, Sweet2000). This generalist predator feeds onchinch bugs, Blissus spp., in turfgrasses(Dunbar 1971, Reinert 1978, Carstenset al. 2008) and is found in crops such asalfalfa (Barney et al. 1984, Braman et al.1985), collards (Muckenfuss et al. 1990),corn (Roach 1980, Tillman 2010), cotton(Whitcomb and Bell 1964, Ruberson et al.1994), peanuts (Tillman 2010), soybean(Roach 1980, Braman et al. 1985), andtobacco (Roach 1980, Torres et al. 2004)

where it feeds on a wide range of pestinsects and sometimes engages in intra-guild predation (Crocker and Whitcomb1980, Sweet 2000). This geocorid alsooccurs on weedy forbs near marshesand streams (Blatchley 1926) and in (ornear) crop fields (Altieri and Whitcomb1979, 1980), as well as under matsof Chamaesyce maculata (L.) Small(Euphorbiaceae) (Wheeler 1981). In nat-ural communities, nymphs and adultsare found on a prostrate phlox, Phloxsubulata L. (Polemoniaceae), in mid-Appalachian shale barrens (Wheeler2011). Under conditions of low preydensities, G. uliginosus can switch toscavenging (necrophagy) and phytoph-agy, with some plant feeding apparentlyneeded for optimal performance (Crockerand Whitcomb 1980, Sweet 2000). Thebug will feed on extrafloral nectaries(Crocker and Whitcomb 1980) and can-nibalize in the laboratory (Sweet 1960,2000; Crocker and Whitcomb 1980;Readio and Sweet 1982).

Geocoris uliginosus is more geo-philous than plant inhabiting (Crockerand Whitcomb 1980, Sweet 2000). Be-cause adults rarely fly, even when dis-turbed (Readio and Sweet 1982), wewere surprised to find them in Spanishmoss, Tillandsia usneoides (L.) L. (Bro-meliaceae), festooning branches of oaks(Quercus spp.; Fagaceae) and other trees.Unaware of arboreal habits in this geo-corid, wewere interested in the frequencyof its occurrence in Spanish moss. Wereport our collections ofG. uliginosus fromSpanish moss in coastal South Carolina,suggest that the bug’s occupation of thisepiphytic bromeliad is more than incidental,

* Edited by Thomas J. Henry; accepted by Michael

W. Gates

and offer explanations for its occurrence onthe plant.

We independently began studies on in-sects associated with Spanish moss anddid not learn of each other’s work untilearly in 2010. From November 2007 toDecember 2009, JHC took biweeklysamples from pendant clumps of T.usneoides on branches of 16 crapemyrtle(Lagerstroemia indica L. and L. indica xL. faurei Koehne hybrids; Lythraceae)trees at the Clemson University CoastalResearch & Education Center (hereafterCharleston REC site 1), Charleston(West Ashley area), SC (32°47.6169N,80°03.9359W). Amat of moss was shakenvigorously for 1 min over a beating sheet,and dislodged arthropods were preservedin jars of ethanol for later identification.On 3 October 2008, AGW beat two adultsof G. uliginosus from Spanish moss onbranches of oak in another area of theClemson University REC (CharlestonREC site 2; 32°47.3259N, 80°04.1039W)and encountered the species on T. us-neoides at two other sites in the coastalplain of South Carolina. Because he hadnot observed this bigeyed bug on trees orshrubs during previous fieldwork insoutheastern states, AGW began to recordthe number and gender of allG. uliginosusfound during the periodic sampling ofSpanish moss on oaks and other woodyplants from late October 2008 to lateNovember 2010 in coastal South Carolina.An ax handle was used to beat clumps ofSpanish moss over an insect net witha shallow bag; geocorids were collectedinto plastic vials and point mounted in thelaboratory. At nearly all sites, other plants,both herbaceous and woody, also weresampled for possible individuals of G.uliginosus. Voucher material of the ge-ocorid has been deposited in the Clem-son University Arthropod Collection,Clemson, SC (JHC’s specimens) and theNational Museum of Natural History,

Smithsonian Institution, Washington,DC (AGW’s specimens).

We collected a total of 32 individualsof G. uliginosus—31 adults and one fifthinstar—at 12 sites in early to mid-April,late June, early August, mid-September,early and late October, and early Novem-ber. Three adults were collected fromSpanish moss at Charleston REC site 1during biweekly sampling, whereas theother specimens were taken during theperiodic sampling at Charleston REC site2 and 10 other sites. Geocoris uliginosuswas found not only on Spanish mossdraping branches of crapemyrtle, Darlingtonoak (Quercus hemisphaerica Bartr. exWilld.), live oak (Q. virginiana Mill), andwater oak (Q. nigraL.; Fagaceae), but also onSpanish moss on Callery pear (Pyrus call-eryana Dcne.; Rosaceae), coastal red cedar(Juniperus virginianaL. var. silicicola (Small)J. Silba; Cupressaceae), maple (Acer sp.;Aceraceae), sugarberry (Celtis laevigataWilld.; Ulmaceae), sweetgum (Liquid-ambar styraciflua L.; Hamamelidaceae),and yaupon (Ilex vomitoria Ait.; Aqui-foliaceae). The record from Spanish mosson the Asian Callery pear (Walterboro,SC), was from a landscape planting ina suburban shopping mall, whereas othercollections were from native plants (ex-cept for the Asian crapemyrtle) in morenatural settings.

Specimens examined.—South Caro-lina: Beaufort Co., Callawassie Island,32°19.8939N, 80°50.4969W, 17Apr. 2009,2 ♀. Berkeley Co., Clements Ferry Rd.,nr Cainhoy, 32°54.0519N, 79°55.1079W,8 Oct. 2008, 2 adults (sex unknown).Charleston Co., Charleston REC site 1,2 May 2008, 1 ♂ & 8 Aug. 2008, 1 ♂ &7 Nov. 2008, 1 ♂; Charleston REC site2, 3 Oct. 2008, 2 adults (sex unknown)& 25 June 2009, 1 fifth instar; Rt.17, Charleston (West Ashley area),32°47.7489N, 80°04.1009W, 2 Oct.2009, 1 ♀; Johns Island, 32°46.8239N,

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80°06.2459W, 31 Oct. 2008, 1♂, 1♀; Rt.165, Meggett, 32°43.0489N, 80°14.3819W,2 Oct. 2009, 1 ♂; Rt. 700, Rockville,32°37.1609N, 80°11.3439W, 17 Sept.2009, 1 ♂, 1 ♀; Cherry Point Landing, SEof Rockville, 32°36.0339N, 80°11.0319W,17 Sept. 2009, 5 ♂, 1 ♀ & 3 Apr. 2010,1 ♂, 2 ♀. Colleton Co., Bear IslandWildlife Management Area, 32°35.5719N,80°27.7509W, 4 Oct. 2008, 1 adult (sexunknown) & 3 Apr. 2010, 1 ♂, 4 ♀;Walterboro, 32°54.4289N, 80°40.0139W,3 Apr. 2010, 1 ♀. Dorchester Co., Rt. 165,Summerville, 32°59.4209N, 80°11.3799W,2 Oct. 2009, 1 ♂.

Records of North American speciesof Geocoris from trees and shrubs arescant. Tamaki (1972) studied predationby G. bullatus (Say) on aphids that fellfrom peach trees to the orchard floor; theonly adult geocorid observed on peachduring the six-year study in WashingtonState followed the mowing of grass inthe orchard. Geocoris punctipes has beenrecorded from crapemyrtle (Mizell andKnox 2006). In addition, an adult of therarely collected G. scudderi Stal wasbeaten from Spanish moss in Texas(Readio and Sweet 1982).

Geocoris uliginosus appears not tohave been reported previously fromstudies of the arthropod associates ofSpanish moss in the Southeast: Newelland Dougherty (1909), Rosenfeld (1911,1912), and Rainwater (1941) in Louisiana;and Young and Lockley (1989) in Mis-sissippi. Although numerous other insectspecies have been recorded from Spanishmoss, the nature of their relationship tothe bromeliad, whether incidental ormore intimate, mostly is unknown (Garth1964). The relationship of insects foundin Florida on four native species ofTillandsia other than Spanish moss (T.usneoides) was uncertain except for afew species that are obligate or poten-tially obligate inhabitants. For arthropods

of unknown relationship to FloridianTillandsia, the null hypothesis was thatthey “just happened to be there,” and thespecies also might be present in treecanopies or the leaf litter beneath, whichcould not be tested owing to lack of in-formation on the canopy and leaf-litterfauna at study sites (Frank et al. 2004).The same null hypothesis regarding therelationship of G. uliginosus to Spanishmoss in South Carolina cannot be testedfor similar reasons. We can, however,propose and evaluate several explanationsfor the bug’s occupation of Spanish moss.Our comments will pertain to adults; theonly nymph observed during our fieldworkwas a fifth instar beaten from a moss-draped branch about 2 m off the ground.

Species of Geocoris can attain largedensities (Tamaki 1972, Readio andSweet 1982), includingG. uliginosus (16bugs/0.09 m2) in chinch bug-infestedturfgrass in Florida (Reinert 1978). Wedid not try to determine the relativeabundance of G. uliginosus at any of thesites where we found the bug in Spanishmoss, but spillovers onto vegetationcould have occurred under conditions ofhigh densities. If such large populationsof the geocorid had prevailed at oursample sites, we probably would haveencountered adults on bare branches oftrees whose Spanishmoss-draped branchesyielded the bug, as well as on woodyplants devoid of T. usneoides. Instead,we found the bug only by beating fes-tooned branches of trees and shrubs.

Entomologists have inventoried Spanishmoss mainly as a potential overwinteringsite for economically important insects,and numerous crop pests have been foundin moss clumps during late fall and winter(Rosenfeld 1911, 1912; Rainwater 1941).Whitaker and Ruckdeschel (2010), how-ever, pointed out that Rosenfeld did notstate whether he examined Spanish mosscollected from or beneath trees.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON426

Adults of G. uliginosus overwinterin protected places such as under logs(Blatchley 1926, Froeschner 1944) andin plant material (“surface woods trash”)near the edges of cotton fields in easternSouth Carolina (Rainwater 1941). We didnot, however, collect adults of the geo-corid in winter during the biweekly sam-pling of Spanish moss at Charleston orin mid-December, early February, or mid-March on trees at other coastal sites inSouth Carolina where adults had beentaken in Spanish moss in spring, summer,or fall. Even though Spanish moss ontrees is used by other insects for over-wintering (Rosenfeld 1911; JHC, unpub-lished data), it might not provide suitableoverwintering quarters for G. uliginosus.

If the presence of G. uliginosus is notdue to spillover effects from large den-sities and the bugs cannot overwintersuccessfully in Spanish moss, then whyare they found in this epiphytic brome-liad? The answer might lie partly in theplant’s water-absorbing trichomes andability to retain water during periods ofdrought (Billings 1904, Penfound andDeiler 1947, Johnson 1975). Althoughthe moisture content of Spanish mossdecreases under drought stress, the plantconsistently maintains a high water con-tent (Penfound and Deiler 1947) andduring drought would provide a moistenvironment for adult Geocoris. Thedrought status in Charleston County,South Carolina, was severe in late 2007when sampling of Spanish moss beganat Charleston, and continued into early2008, but was normal, or, at most, in-cipient through the remainder of thesampling period (SCSCO 2010). Becausethe geocorid was found in Spanish mossin larger numbers when precipitation re-turned to normal levels, its use of thebromeliad during drought might be ofminimal importance. The bugs might alsoobtain supplemental nutrients by imbibing

from Spanish moss. Certain “host” trees,such as oaks, exhibit high rates of foliarleaching, allowing Spanish moss to obtainminerals (e.g., Ca, Mg, K, and P) fromatmospheric precipitation through thecanopy (Schlesinger and Marks 1977). YetSpanish moss might be most important toG. uliginosus by providing a consistentsource of various soft-bodied arthropods(JHC, unpublished data) as potential prey.

Crocker and Whitcomb (1980) notedthat species of Geocoris do not colonizeall plant species that harbor potentialprey and offer apparently suitable habi-tat. Spanish moss is regularly frequentedby adults of G. uliginosus in coastal SouthCarolina, but whether this plant associa-tion might benefit the bugs—and if so,how—remains unanswered.

We thank Merle Shepard and JamesRushing (Clemson University, CoastalREC, Charleston) for facilitating ourwork and Peter Adler (Clemson Univer-sity, Department of Entomology, Soils,and Plant Sciences, Clemson) and FrancisReay-Jones (Clemson University, Depart-ment of Entomology, Soils, and PlantSciences, Pee Dee REC, Florence) forcomments that improved an early draftof the manuscript. Fran Wimberly, LeslieStrickland, and Jessie Strickland assistedJHC in collecting specimens from crape-myrtle trees. This manuscript is basedupon studies supported by NIFA/USDAunder project number SC-1700351 (JHC).This manuscript is Technical Contribu-tion no. 5894 of the Clemson UniversityExperiment Station.

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A. G. Wheeler, Jr. and Juang-HorngChong, (AGW) Department of Entomology,Soils and Plant Sciences, Clemson University,Clemson, South Carolina 29634-0315,USA (e-mail: awhlr@clemson.edu); (JHC)Department of Entomology, Soils and PlantSciences, Clemson University, Pee DeeResearch & Education Center, 2200 PocketRoad, Florence, South Carolina 29506, USA(e-mail: juanghc@clemson.edu)

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