Bryophagy in the Auchenorrhyncha: Seasonal history and habits of a moss specialist, Javesella Opaca (Beamer) (Fulgoroidea: Delphacidae)

12 Pages·2003·3.8 MB·

by  A G Wheeler

PROC. ENTOMOL.SOC.WASH.
105(3). 200.3.pp. .599-610
BRYOPHAGY IN THK AUtHENORRHYNCHA: SEASONAL HISTORY AND
HABITS OF A MOSS SPECIALIST,JAVESELLA OPACA (BEAMER)
(FULGOROIDEA: DELPHACIDAE)
A. G. WntifiLRR. Jr.
Department ofEntomology. Clemson University, Clemson, SC 29634-0365, U.S.A. (e-
mail: [email protected])
—
Abstract. Specialization on mosses, previously unknown in the Fulgoroidea and ap-
parently also for any auchenorrhynchan, is reported for the delphacid planthopperJave-
sella opcicci (Beamer). New state records for this seldom-collected species are Alabama,
Georgia, Massachusetts, New York, North Carolina, South Carolina, and Tennessee.
Nymphs developed mainly on the gametophores ofcommon hair-cap moss. Polytrichum
commune Hedw. (Polytrichaceae),ongraniteflatrocksandinothercommunities,andonce
were found on anotherpolytrichaceous moss, Polytrichastniiualpinum (Hedw.)G.L. Sm.
At Boggs" Rock, a granite Hatrock in northwestern South Carolina, late instars overwin-
tered in mats ofPolytrichum commune, and adults developed by mid- to late March. The
only macropterous adults (n = 5) observed from June 1996 to July 1998 were found
between 10 April and 2 May 1997; all other adults were brachypters. First instars ofthe
season's first generation appearedfrom mid- to late May, with adultsappearingfrom mid-
June to early July. The first instars observed in August were those of an overwintering
generation. Fluctuations in the numbers of planthoppers are discussed in relation to the
condition of moss colonies (green vs. drought stressed) and weather (precipitation, tem-
perature, and evaporation). In the laboratory, nymphs fed on thicker stems ofthe moss.
The use of mosses is considered a derived host association in the evolution of the Del-
phacidae.
Key Words: Insecta, Delphacidae,mossfeeding,Polytricluinicoinnuine.Polytrichustrum
(ilpinimi. granite outcrops
The large gaps in our knowledge of the son 1969, Lawrey 1987), the insect fauna
bryosystem, coupled with the ubiquity of as.sociated with bryophytes evidently is not
mosses, leaves great scope for original re- ^^tably diverse when only those phytopha-
search by bry^otaunal eco—los,Ii..sMt..s.,K,m.ch,i.n 1„9„9„0 o"2ens tk,hneosewnplt^aontcsomaprrl-eeit.econt.ihseiid•rerdjeedv.elAormpmoenngt
Studies on insect-plant interactions have bryophy—tes, insects are found mai—nly on
emphasized vascular plants, the Tracheo- mosses Bryophyta sensu stricto rather
phyta, including seedless groups such as than on hornworts and liverworts. Moss-in-
fems, butespecially the moreeconomically habiting insects, which are recorded for
important gymnosperms and angiosperms most of the hemimetabolous and holome-
among the seed plants. Even though insects tabolous orders (Gerson 1969), can be
ofnonvascular plants (Bryophyta) general- found on both aquatic and terrestrial moss-
ly have been neglectedby researchers(Ger- es. In many cases, whether the species use
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600 PROCEEDINGSOFTHEENTOMOLOGICALSOCIETYOFWASHINGTON
mosses for food or only for shelter is not Materials and Methods
known.
Study area.—Boggs" Rock (34°48.4'N.
In the Homoptera. used in its traditional 82°41.6'W) in northwestern South Carolina
sDeonhsleen(tahnedgMroourpanis1p9a95r]a)p.hymloetsiscf[eee.dge..rsvionn- was the main study site. It is 2.1 km north
of Liberty. Pickens County, and lies just
clude several sternorrhynchan groups:
west of the intersection of U.S. highway
aphids. such as species ofMelaphis Walsh.
178 with state secondary road S-39-317
Mnscaphis Borner, and Myzodiiim Borner
(Quairy Road). This granite-gneiss flatrock
(e.g.. Patch 1938. Borner 1952. Muller ranges in elevation from 262 to 293 m
1973. Smith and Knowlton 1975, Richard- (Knox 1974). Severalplantspeciesendemic
son 1981. Moran 1989. Dolling 1991.
togranite flatrocksinthesoutheasternUnit-
ThomasandLombard 1991),aswellascer- ed States (e.g., McVaugh 1943, Shure
tain ensign scales(Ortheziidae)(e.g.,Kozar 1999) are found at Boggs' Rock, but be-
and Miller 1999. 2000) and mealybugs cause permanent depression pools are ab-
(Pseudococcidae) (Williams 1985). The use sent (Knox 1974), flatrock endemics re-
of mosses as true hosts by ortheziids is stricted to such pools also are lacking.
largely undocumented (Dolling 1991). Host plants. Polytrichiim commune
knBorwynopihnatghye aApupcahreenntolryrhhyanschraema(eivnieddenutnl-y d(aMrukscgir:eePno,lyrtorbiucshta,lepse:rePnonliyatlrimcohsacseaoef)wiisdea
also a paraphyletic taxon [e.g., Campbell et distribution in the New andOld Worldsex-
al. 1994, Sorensen et al. 1995, Hamilton cept in tropical areas. Its leafy stems, rigid
1996]). Dolhng (1991) stated that all au- and erect (acrocarpic), are connected by an
chenorrhynchans feed on vascular plants. extensive system ofsubterranean rhizomes.
Herein bryophagy in the auchenorrhynchan Stems of this tall moss can reach a height
superfamily Fulgoroidea is reported. I sum- of45 cm (Crum 1976, Crum and Anderson
marize dataon the seasonality andhabitsof 1981, Derda and Wyatt 1990). This dioe-
the delphacid planthopper Javesella opaca cious moss reproduces asexually by vege-
(Beamer) on common hair-cap moss. Po- tati\ebranchingandpropagationfromplant
lytrichitm commune Hedw.. and record fragments, with some populations being
nymphs and adults from the moss Polytri- maintainedentirely by vegetativereproduc-
cluistrifiii cilpiiuim (Hedw.) G.L. Sm.: dis- tion; it can also become established sexu-
cuss fluctuations in the numbers of plant- ally from spores (Leslie 1975, Derda and
hoppers relative to the condition of host Wyatt 1990,WyattandDerda 1997). Found
colonies (green vs. brown and drought on various acidic substrates in disturbed
stressed) and weather conditions: precipi- andnaturalcommunities,P. communeoften
tation, temperature, and evaporation; and is abundant on granite flatrocks. It isan im-
suggest that mosses representanevolution- portant soil builder ofthe annual-perennial
arily derived host association in the Del- herb community of granite outcrops and a
phacidae. Seven new state records also are characteristic plant of that community. Its
given for this seldom-collected species. coloniesormatsarepersistentandoftenex-
This contribution is dedicated to my tensive (Burbanckand Phillips 1983. Derda
friend and colleague. Craig A. Stoops, who and Wyatt 1990. Quarterman et al. 1993).
called my attention to a delphacid he col- The moss variety onwhichthe planthop-
lectedfrom mosswhen wewerein thefield perJ. opaca was collected at Boggs" Rock
in June of 1996. He helped make field col- andotherrockoutcrops isP. communevan
lections and laboratory observations before perigoniale (Michx.) Hampe. Plants ofthis
he left Clemson University in January variety are shorterand grow in densertufts
1997. than those of the nominate variety (Crum
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VOLUME 105. NUMBER 3
and Anderson 1981). Ofthe many varieties terial of both nymphs and adults was col-
of P. commune that have been described. lected and deposited in the Clemson Uni-
peri^oniale has been the one most consis- versity Arthropod Collection (CUAC);
tently recognized by authorities (Zouhairet adults also were deposited in the collection
al. 2000). Polytrichum commune van peri- ofS.W. Wilson.CentralMissouriStateUni-
goniale, however, is notgeneticallydistinct versity. Warrensburg.
from the nominate variety (Derda and Wy- Sampling at Boggs' Rock wasconducted
att 1990). Overlapping in range with P. similarly in 1997 and 1998 butat morereg-
commune van commune, P. commune van ular intervals than in 1996. I recorded the
perigoniale might represent only a "dry sta—ge of the first 10 individuals ofJ. opa-
land phase" ofa species that growsmainly ca as earl—y or late instars and adult males
in wet habitats (Crum and Anderson 1981). or females found on the moss in Knox's
Derda (1998) considered P. commune var. (1974) outcrop division C, a colony ofthe
perigoniale merely a phenotype ofdry. ex- host plant that became desiccated and
posed soils (cf. Zouhair et al. 2000). brown without frequent rain. I also often
In the southern Appalachians, J. opaca recordedthestageofthefirst 10individuals
was found on Polytrichum commune var. encountered in asupplemental samplefrom
commune and Polytrichastrum alpinum. a moss in a wetter area of the outcrop
polytrichaceous moss that also has been (Knox'sdivisionB). In 1997.afteraninitial
placed in Pogonatum and Polytrichum collectioninearlyJanuary.Isampledweek-
(Smith 1971. Crum and Anderson 1981. lyorbiweekly fromearlyMarchtolateAu-
Smith Merrill 1992). This coarse species, gust; I also sampled in mid-Septemberand
usually 4-16 cm high with erect stems, is earlyNovember.In 1998.samplesweretak-
Holarctic and particularly widespread at en weekly (occasionally biweekly orevery
northern latitudes (Crum and Anderson three weeks)from lateFebruary tolateJuly
1981). — when nomoreplanthopperswerebeingcol-
Field sampling. AfterJ. opaca wasdis- lected, presumably becauseofdroughtcon-
coveredatBoggs" RockinJune 1996. I sur- ditions. Afterthat, Isampledsixmoretimes
veyed other southeastern granite outcrops from late August to late November in an
(flatrocks and monadnocks) for the pres- attempt to collect J. opaca. Six additional
ence of the delphacid on Polytrichum spe- attempts to collect the planthopper at
cies. The seasonality ofJ. opaca was fol- Boggs' Rock were made from early April
lowed at Boggs' Rock, the study areas cor- 1999 to late May 2002.
respondingwithgraniteoutcropdivisionsB Fluctuations in planthopper numbers.
and C in Knox's (1974) floristic study. Data from the two nearest weather-record-
I (or C.A. Stoops) visited Boggs' Rock ing stationswereusedtobetterinterpretthe
seven more times at irregular intervals in numbers of planthoppers observed relative
1996 andcollected planthoppers by placing to the condition of moss colonies. The
a white enamel pan (28 X 40 cm) into a weather station at Pickens (34°53'N,
thickmatofthemossandusinganaxehan- 82°43'W) is about 10 km north of Boggs'
dle to tap the overlying stems overthe pan. Rock: the Clemson station (34°4rN.
Planthopper nymphs were field sorted by 82°49'W) is about 18 km southeast of the
size and recorded as either early (I, II) or sainple site. —
later (III-V) instars, and the sex of adults Laboratory observations. The work
andtheconditionofthemosscolonieswere with J. opaca in the laboratory involved
noted. Thenumberofindividualsof7. opa- placing nymphs ofseveral instars in plastic
ca observed was not standardized but gen- petri dishesorsnap-capvialsthatcontained
erally was 10. Nearly all individuals were the gametophyte (1-5 stems) of common
replacedonthemoss,althoughvoucherma- hair-cap moss. The feeding behavior of
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PROCEEDINGSOFTHEENTOMOLOGICALSOCIETYOFWASHINGTON
nymphs, presence of exuviae, and number McPherson 1981), was observed at Boggs"
of clays ofobservation were noted, but the Rock by early Novemberin 1996and 1997
duration ol nymphal stadia was not deter- (late instars collected in New York in mid-
mined. August and some early instars from Mas-
sachusetts in early September also were
Jcivi'sella opuca (Beamer) dark). Adults firstwere seen in 1997during
(Fig. 1) sampling on 16 March and were presenton
Jdvcscllii opiua was described in the ge- 29 March in 1998. In both years, malesap-
nus Delpluicodes Fieber from Connecticut peared first, but sex ratios usually became
by Beamer (1948), who selected as holo- female biased within three weeks after
lype a male collected at Willimantic t)n 8 adults began to appear. Overwintered fifth
August 1946. His type series also included instars were last seen on 10 April in 1997
adults taken from 3 August to 2 September (2 in sample of 10) and 5 April in 1998 (I
1946, at Storrs, Conn. (14 6. 8 9): Ding- of 20).
mans FeiTy, Pa. (6 cJ, 5 9); and Mountain First instars ofthe season's new genera-
Lake, Va. (1 6). The only additional pub- lion first wereobservedon 23 May in 1997
lished record is that of Maw et al. (2()()()) and 12 May in 1998. Adults ofthis gener-
Uom Quebec. Canada; this delphacid was ation, almost all males initially, first were
listed as a provincial record without a spe- seen in the 19June sample in 1997 andthe
cihc k)cality and wasrecognizedasbelong- 22June sample in 1998. Several lateinstars
ing to the genusJavesella (as ''Javasella") (7 of 20) of the new generation were still
Fennah (Maw et al. 2000). According to present in 1997 on 20July. Early instarsof
K.G.A. Hamilton(personalcommunication, a partial (overwintering) generation were
2002), the Quebec record of J. opaca is found on 4 August 1997. In 1998, appar-
based on a total of 13 males and 3 females ently because of drought conditions, no
from Covey Hill (5 June) and Laniel (5 nymphs or adults could be collected after
June, 20-21 July). Newstaterecords,based early July.
on the present study (see Material exam- Nearly all the adults observed from June
ined), are Alabama, Georgia, Massachu- 1996 to July 1998 were brachypters. Only
setts, New York, North Carolina, South five macropters were seen during the study,
Carolina, and Tennessee. all between 10 April and 2 May 1997; the
The transfer of opaca to Javesella by three that were collected were females.
Maw et al. (2000), made in a checklist Adults were not found after mid-August,
without discussion of this nomenclatural with only nymphs observed from then until
change, is the appriipriate generic place- mid- to late March ofthe following year.
ment (S.W. Wilson, personal communica- Fluctuations in planthopper numbers.
tion, 1996). Mainly on thebasisofgenitalic The numbers ofJ. opaca observed fluctu-
characters, the polyphyletic genus Delplui- ated widely during the period ofstudy, de-
codes should be restricted to about 10 Old pending mainly on the amount of precipi-
World species (Asche 1985, Bartlett and tation at Boggs" Rock. Available moisture,
Dietz 2()()()). — in turn, affected the condition of the moss
Seasonal history. Lateinstarsof7. opa- colonies inhabited by the planthopper.
ca overwintered in mats of common hair- Despite the wide variation in the num-
cap moss, hi contrast to the yellow or yel- bers of planthoppers that could be beaten
k>w-brown nymphs found during spring from Polytrichitnu at least 10 individuals
and summer, the overwintered nymphs usually could be found on each sample
were darker and sometimes fuscous. The date; few, however, were found in July of
darker coloration, similar to that observed 1998. Precipitation that yearwasabout 160
incertainotherdeiphacids(e.g.,Wilsonand cm at Pickens and about 156 cm at Clem-
VOLUME 105. NUMBER3
Fig. I. Jiivcsc'Uaopcuii. (A) Females(brachypters)on lealygametophytc immon hair-capmoss,Pol\-
rriiliiini< iiiiiiiiinc:(B)dorsaland(C)lateral viewsoffemale.
son, figures that actually exceeded annual with nine consecutive days of 36.1-36.7°C
averages of 142 cm tor Pickens (1952- (97-98°F). Evaporation at Clemson (data
2001) and 134 cm tor Clemson (1930- are unavailable for Pickens) exceeded rain-
2001). Looking only at total precipitation fall by 15.2 cm for the 30 days following
for 1998, fiowever, is misleading; aftersub- 20 June.
stantial rains on 1 May (nearly 6 cm) and Only four adults and no nymphs were
8 May (>3.6 cm), rainfall was scant in the observed during regularsamplingon4July
weeks preceding the planthopper's disap- 1998; seven adults were seen when a sup-
pearance. At Pickens, precipitation for the plemental sample was taken on 6 July. No
last half of May was only 1.4 cm; rainfall additional individuals were found during
for June totaled 13.6 cm, with nearly half samplingon seven dates from 24July to29
ofthe total fallingon the month's 20thday. November 1998. Two adult males were
After 20June, only 0.3 cm ofrain fell dur- found inearlyApril 1999, butJ. <>i?acawas
ing the next 30 days, a period when daily notobservedat Boggs' Rock in lateMarch,
maximum temperatures substantially ex- mid-May, and mid-August 2000; in late
ceeded normal highs. Temperatures were at March andearly November2001; orin late
least32.2°C(90°F)from 21 Juneto 14July. May 2002.
— —
604 PROCEEDINGSOFTHEENTOMOLOGICALSOCIETYOFWASHINGTON
Habits. Jcivesello opaca was found at were maintainedin apetri dish onthe same
Boggs" Rock inthictc matsofcommonhair- sprigs of P. commune for 16 days and
cap moss in marginal areas of rock out- nymphal exuviae were observed. A first in-
crops. Moss colonies iiarboring the plant- starplaced ina4-dram vial withtwomoist-
hopper grew beneath shrubs and small ened stems of moss molted iince and was
trees, particularly Chinese privet (Liiiiis- still alive after 27 days—.
rniiii sinciisc Lour.; Oleaceae) and eastern Material examined. Adults from a lo-
red-cedar(Jiiiiiperus virginianaL.;Cupres- cality are listed first, followed by a listing
saceae). The delphacid was not found in of any nymphs; when nymphs at Boggs'
more exposed colonies of the moss. Even Rock were recorded in the field as early (I,
when nymphs and adults were most nu- II) and later(III-V) instars. the numbers of
merous, they were difficult to detect on early and later instars are separated by a
mats of PolyUiclnim; only occasionally hyphen. An asterisk beside the word
wereafew individualsobservedonthe sur- nyniphis) indicates that voucher material is
faceofP. commune m—ats. The largestnum- housed in the CUAC. Collections were
bers of planthoppers early instars only madebytheauthor(nocollectorindicated),
were found on 8 June 1997. Javesellaopa- C.A. Stoops (C.A.S.). both ofus (C.A.S. &
ca had been difficult to find on 23 May A.G.W.). or W.K. Reeves (W.K.R.) from
1997 when moss colonies were brown (a Polytriclnim commune, except in Alleghany
total of9 individuals was observed in both County, NorthCarolina,wherethehostwas
the regular and supplemental samples), but Polytriclutstrum alpiniim and in Pennsyl-
on 8Juneafterrecentrainfall,morethan.30 vania where the host was identified as Po-
first and second instars were found in each lytriclnim sp.
ofthree I-pan samples from now green ar- ALABAMA: Cherokee Co.. Rt. 35. SE
easofmoss. When nymphswerenumerous, of Fort Payne. 31 Aug. 1996. 2 adults (sex
exuviae dislodged from moss alsowereob- undetermined). 10 Apr. 1997 (I d, 9 9);
served during sampling. Jackson Co., sandstone outcrop, Rt. 117,
In the southern Appalachians, J. opaca 0.3 km NW ofjet. Rt. 71. Flat Rock. 10
was found at an altitude ofabout 1.130 m Apr. 1997 (3 d.4 9; 1 fifth instar).GEOR-
on Polytrichaslrum alpimim (Alleghany GIA: Columbia Co.. Heggie's Rock, E of
Co., NC). The moss was mostly obscured Appling. 5 Apr. 1997 (4T, 11 2), 26 Mar.
byvariousgrasses. Populationsoftheplant- 2002 (1 d; 1 fifth instar);GwinnettCo..No
hopper also were found at about 1,685 m Business Creek granitic outcrop. 2.6 km
in mats ofPolytriclnim commune van com- SSE of Snellville, 7 July 1996 (1 c?). 26
nnnie (CarterCo., TN). Mar. 2002 (1 cJ,4 9; 2 fifth instars). MAS-
When nymphs were placed in petri dish- SACHUSETTS: MontagueSand Plains.3.5
ePsoloyrtrviicallnsiwmitchosmpmruingse,oftghaemyetqoupihcyktleyf(r<o5m k9;meNarloyf Minosnttarasg)u.e.NE5WSepYt.OR2K00:2S(u\lliiv,an\
min.) settled on the thicker stems and in- Co.. Rt.42, Forestburg,3Sept. 2002(8first
serted their stylets. While feeding, nymphs through third instars); Ulster Co., Sam's
(n = 10+) remained motionless, except for Point Dwarf Pine Ridge Preserve, NE of
anoccasionaltwitchingofthelegs,andtyp- Cragsmoor. 13 Aug. 2000 (12 S. 10 9;
ically stayed at one site for two hours or nyniphs). NORTH CAROLINA: Alleghany
more. Only once wasacleai"dropofexcre- Co., ca. 7 km ENE of Laurel Springs, 16
tion or honeydew seen at the end of the June 2002 (4 d, 1 9; late instars); Swain
anus. First instars were not reared to adult- Co., Great Smoky Mountains National
hood, but preliminary observations suggest Park, nr. Ravens Fork. 8 Jan. 2001. W.K.R.
thatJ. opacacouldbeeasilyrearedonmoss (late instars). PENNSYLVANIA: Potter
inthelaboratory. Nymphsofvariousinstars Co.. NE ofKeatina Summit. 18 May 1997.
VOLUME 10?.NUMBER3
C.A.S. (3 late instars). SOUTH CAROLI- DlSCU-SSION
oANf.AG:.LiWPb.ierctk(yea,ndusl1tC.6so-,.1.7sBeoxJgugnusen"de1Rt9oe9cr6k,m.inC2e..d1A;.kS.m25&N+ pmraetWssheenocnfePJo.onloymptoarscicsahuwwmaassincfioJrnsustniecdoelrolefedct1ie9nd9c6if,dreointm-s
nnyymmpphhss=)^;: 12-22);Ju6nJeuly1919969,6,CC..AA..SS..(&adAul.tGs.; gtaolr.oIidwsa,saungarowuapreinofwahniychmoasdsu-lftesedoifngmafnuly-
W. (7 (5, 10 9; nymphs*:0-ll); 20 July species, including delphacids, can be found
1996, C.A.S. (adults; nymphs^^O-4); 18 on plants that do not support nymphal de-
Aug. 1996 (5 adults, sex undetermined; velopment (e.g., DuBose 1960, Wilson et
nymphs:! 1-4); 29 Sept. 1996, C.A.S. & al. 1994). Because seeds ofvarious angio-
A.G.W. (nymphs:3-7); 13 Oct. 1996, sperms accumulate, germinate, and survive
C.A.S. & A.G.W. (nymphs:10-0); 9 Nov. in mats of mosses (McVaugh 1943), even
1996, C.A.S. & A.G.W. (nymphs:10-0); 4 the planthopper nymphs collected from
Jan. 1997 (nymphs*:0-19); 7 Mar. 1997 moss might have been associated with vas-
(nymphs*: 0-11); 16 Mar 1997, C.A.S. & cular plants, such as grasses orsedges, that
A.G.W. (2 6: nymphs: 0-10);21 Mar. 1997 were growing within the Polytrichiiiii col-
(3 adults, sex undetermined; nymphs: 0-7); onies.Delphacidsdevelopmainlyonmono-
4 Apr 1997 (4 cj, 4 9; nymphs*:0-2); 10 cots, especially grasses and sedges (Wilson
Apr 1997 (3 (5, 5 9; nymphs; 0-2); 18 et al. 1994). Javesella kilnuini (Van Duzee)
Apr 1997 (4 S.b 9); 25 Apr 1997 (3 6. andJ. stall(Metcalf)areassociatedwithan
1 9); 2May 1997(3 6,1 9); 10May 1997 aberrant host, the vascular cryptogam
(3 (5, 6 9; nymph:(3-l); 23 May 1997 (2 (sphenophyte) horsetails, Equlsetwn spp.
cJ, 4 9; nymphs:2-0); 8 June 1997 (Equisetaceae) (Strickland 1940, Ossian-
n(nyymmpphhss**::07--94));;2199JJuunnee 11999977 ((23 (<55,, 42 99 nilAssocnon1s9i7s8t)e.nt collection of nymphs and
nymphs*:l-3); 5 July 1997 (3 6: nymphs* adultsofthedelphacid from P. coinimineat
2-5); 13 July 1997 (11 d, 3 9; nymphs* Boggs" Rock and elsewhere, coupled with
2-4); 20 July 1997 (10 d. 3 9; nymphs* observations of its moss-feeding behavior
2-5);4 Aug. 1997 (6 d, 2 9; nymphs*:11- in the laboratory, soon indicated that J.
1); 17 Aug. 1997 (nymphs*:8-l); 14 Sept. opaca was a true bryophage. It can be re-
1997 (nymphs*:2-0); 2 Nov. 1997 garded as a bryophilous sp—ecies or a bry-
(nymphs: first through third instars): 28 oibmiaolnta{ssseoncsiitatGeedrseoxncl1u9s8i2v)elythwatitihs,abnryaon--
(Fenby.mph1s9:908-1(0n)y;mp2h2s*Ma:0r-6)1;99815(nMyamrphs1:9098- pkhnyotewsn. Abrsyosupchha,gJo.usopafuclagobreocidomaensdthaeppfairrs-t
10); 29 Mar 1998 (3 i: nymphs:0-7); 5 ently also the first auchenorrhynchan
Apr 1998 (7 i. 3 9); 12 Apr 1998 (5 S, known to develop on mosses. Somewhat
5 9); 24Apr 1998 (1 .5,9 9);5 May 1998 similarly, the firstknown bryophagous spe-
(1 c?, 2 9); 12 May 1998 (2 9; nyniphs:8- ciesofthe largestfamilyofHeteroptera,the
0); 18 May 1998 (nymphs:10-0); 27 May Miridae, only recently was discovered (Ya-
1998 (1 9; nymphs:9-0); 6 June 1998 sunaga 2000; see also Wheeler 2001).
(nymphs:8-12); 22 June 1998 (12 <5, 1 9; My collection ofJ. opaca in North Car-
nymphs:l-6); 4 July 1998 (3 d, 1 9); 6 olina on Polytrichastriim alpinum might
July 1998 (6 d, 1 9); 4 Apr 1999 (2 d); shed light on how Beamer (1948) was able
York Co., granite flatrock. Clover, 20 Apr to collect 22 adults ofthis planthopperata
1997 (3 9; 1 fifth instar). TENNESSEE: locality in Connecticut and 11 adults in
Carter Co., Carver's Gap, Roan Mountain, Pennsylvania without associating the spe-
27 October 2002 (late instars). cies with mosses. When 1 collectedthe first
PROCEEDINGSOFTHEENTOMOLOGICALSOCIETYOFWASHINGTON
specimens of J. opaca in North Carolina 2000), the long-distance conducting sys-
while sweeping grasses. I was unaware of tems ofcertain members ofthe Polytricha-
thepolytrichaceousmossgrowingunderthe ceae are similarin structure andfunctionto
grasses. Beamer probably would not have thoseofsimplevascularplants(Tansleyand
looked for delphacids in extensive mats of Chick 1901, Thomas et al. 1988). Water
Polytrichum commune because no auchen- moves in elements called hydroids; photo-
orrhynchanswerethenknownfrommosses. synthate, principally sucrose, moves in lep-
He more likelywouldhave swepttheplant- toids that are similar to sieve elements in
hopperfrom theerect stemsofPolytrichas- thephloemofhigherplants. Amongmosses
triim alpinum (or another moss of similar that have been studied, theconductingcells
growth habit) that grew beneath grasses, of Polytrichum commune, Polytrichastrum
plants that are common hosts of Delphaci- alpinum, and certain other polytrichaceous
dae. Delphacids should be more readily species are the most structurally complex
swept from amoss suchasPolylrichostnim (Collins and Oechel 1974, Behnke 1975,
alpinum that grows in loosetufts than from Scheirer 1980, Reinhart and Thomas 1981,
one such as Polytrichum commune that has Richardson 1981, Thomas et al. 1990).
a more densely tufted (cespitose) growth The moss-feeding mirid described as a
habit. Even if Beamer had noticed the new species by Yasunaga(2000)belongsto
mosses while he swept grasses, he likeh a group of heteropterans that are mainly
v\ould not have associated the planthopper cell-content feeders on mesophyll rather
uith a nonvascularplant. than vascular-tissue feeders (Wheeler
The Delphacidae appear not to have di- 2001). The mirid would not necessarily
versified on mosses. Except for J. opaca. need to feed on a moss with a conducting
all other delphacids for which nymphal system similar in structure to that of vas-
hostsareknowndeveloponvascularplants. cularplants.Javesellaopaca. however,asa
Fulgoroids andotherhomopteransthatfeed presumed phloem feeder, is likelyrestricted
on plants otherthan angiospermsoronfun- to polytrichaceous mosses with well-devel-
gi likely arederived fromangiospermfeed- oped conducting cells. It might be more
ers (e.g., Wilson et al. 1994; see also Ham- host restricted than are many other bry-
ilton 1990). Even though J. opaca belongs ophagous insects (Gerson 1982, Lawrey
to an "advanced" lineage (Stenocraninae- 1987). In fact, most delphacids (74%) are
Plesiodelphacinae-Delphacinae) ofthe Del- recorded from a single plant genus (Wilson
phacidae and belongs to the most morpho- et al. 1994).
logically advanced tribe of the most ad- Javesella opaca might feed on gameto-
\anced delphacid subfamily (see Asche phores ofcommon hair-cap moss in a man-
1985, 1990: Emeljanov 1995), moss feed- ner similar to that of the aphid Myzodium
ing by this planthopper might represent an modestum Hottes on the same moss species
ancient orrelictual host relationship forthe (Thomas and Lombard 1991). The plant-
family. More likely, however, is that in the hopper, however, does not feed in aggre-
Delphacidae the use of mosses reflects a gations as does the aphid and might not al-
one-time host shift and an evolutionarily ter the translocation of sugars in internal
derived host association. conducting tissues ofthe moss, as Thomas
Bryophagy in the Delphacidae, a phlo- and Lombard (1991) demonstrated for the
em-feeding group (e.g., O'Brien and Wil- aphid. Occasional production ofhoneydew
son 1985. Denno et al. 1987, Wilson et al. droplets by the sucrose-feeding M. modes-
1994, WatanabeandKitagawa2000),inight tum (Thomas and Lombard 1991) agrees
have been predicted for polytrichaceous with my observations on J. opaca, which
mosses. Although mosses are considered presumably also feeds on photosynthate of
nonvascular plants (Buck and Goffinet common hair-cap moss.
VOLUME 105. NUMBER3 607
Javcsellci Dpiicahasbeencollectedmain- of J. opaca in the two sample areas at
ly on thick mats of Polytrichiim commune Boggs" Rock. Adults and nymphs were no
in both natural and disturbed communities. longer seen, even in the usual wetter moss
This planthopper might also be found on colonies,whichhadbecomedesiccated.Al-
Polytrichastriim ohioense (Ren. & Card.) though temperatures within green mats of
G.L. Sm. Infact. Knox(1974)recordedPo- P. commune generally are lowerthan those
lytrichastriim ohioense as the only polytri- at the surface, desiccated mats are marked
chaceous moss at Boggs' Rock, but hi.s by internal temperatures higher than those
voucher material actually is Polytrichum of the ambient air (Leslie 1975). Other
commune (L.E. Anderson, personal com- groups ofarthropods, such as collembolans
munication. 2002). Both polytrichaceous and mites, were scarce in 1998 when moss
species can be found on granite outcrops colonies were brown anddry. In contrastto
(e.g.. Costing and Anderson 1939, Mc- the planthopper, collembolans and mitesre-
Vaugh 1943). but Polytrichum commune is appeared in numbers following abundant
a characteristic plant of granite flatrocks rainfall. That only two adults of J. opaca
and apparently is the dominant hair-cap have been collected at Boggs" Rock since
moss in granite outcrop communities in July 1998. even when moss was lush, sug-
Georgia and elsewhere in the southeastern gests that its populations were substantially
states (e.g.. Burbanck and Phillips 1983. reducedduringperiodsoflowrainfallinthe
Quarterman et al. 1993, Murdy and Carter summer of 1998 rather than being found
2000). deeper in moss and. therefore, perhaps
GiventhepersistenceofPolytrichumcol- more difficult to dislodge from desiccated
onies and the stability of granite flatrock colonies ofthe host.
communities, the observed low incidence In the southern Appalachians, mats of
{<3i%) of macroptery might have been ex- Polytrichum commune in high-elevation
pected forJ. opacabecausedispersal might mesic communities, such as grassy balds,
not be critical to the continuity ofa plant- would be constantly moist and should be
hopper population that lives in such a rel- considered an even more permanenthabitat
atively stable or permanent habitat (e.g., than the periodically dry mats ofthe moss
Wagner and Liebherr 1992). From June
1996 to July 1998, the delphacid typically in granite outcrops. The degree ofbrachyp-
was observed at Boggs" Rock in small tery in J. opaca might be studied in popu-
numbers when rainfall was scant and its lations occupying both permanent and less
bhoestcowlalsecmteedtabionligcreaaltleyrinnaucmtibveer,sbuftriotmcosutlildl smtaarbilleyhianbgietsattss.suTchraotsethtirsanpslloacnattheodppienrmoprsis-
green colonies of the moss in wetter areas leptoids, asapparently isthecase inseveral
of the flatrock. The more consistent avail- other moss-feeding hemipterans, requires
ability of moisture in these wettercolonies verification. Also needed are surveys for
apparently allowed egg hatch to continue, additional moss species that might serve as
which sometimes led to asynchrony in the hosts and studies of basic life-history pa-
development of hopper populations in des- rameters such as fecundity, longevity, and
iccated compared to greener host patches. duration ofthe immature stages (eggs were
In 1998, a yearofabove-average precip- not found in the present study), as well as
itation, the delphacid was not found after observations on mating and oviposition be-
early July. Apparently the low and uneven havior. In addition, the studieson nutrition-
distribution of rainfall in late spring and al ecology that Lawrey (1987)suggestedbe
early summer, combined with consecutive conducted on moss-feeding arthropods in
days of unusually high temperatures and general are appropriate to the research
evaporation, severely reduced the densities needs forJ. opaca.
—
608 PROCEEDINGSOFTHHENTOMOLOGICALSOCIETYOI- WASHINGTON
Acknowledgments pha). Marburger Enli)nu)li)j;iM.'he I'lihlikalioiien
baendTrhececuormri(ods«eisdt-ytb"eoecfeaduCisrnegaiohgfabStihtteosoiponsfi,tiJa.nloocpwoallcaeanctceianon-n pl2e(hr1aa).e::ildylFay-ieoyi.ilwOgi.oVtriho/i.rcdeaevyaii)sn—iaoen.aoasfingVneiiwzfciacsyauanbtfMaupmihiyrlly(oHgooetnnieUotcpil-c-
tomologist with the U.S. Navy. I also am link.BishopMuseumOccasionalPapers.^0: 154-
grateful toS. W. Wilson (DepartmentofBi- 187.
ology, Central Missouri State University, Uarlletl,C. K. and L. L. Dietz. 2000. Revisionofthe
New World delphacid planthoppergenusPisson-
Waanrdreinnsfbourrmgi)ngformeideonftifitysingcortrheectdelgpehnaecriidc (l>iHci.a\ti(oHnesmiipnleErnat:omEoullgoogryo:ideMao)n.ogTrhaopmhass..ESanytoPmuob--
placement, L. E. Anderson (Department of logicalSocietyofAmerica,Lanhani.Md.234pp.
Botany, Duke University, Durham, N.C.) Beamer. R. H. 1948. .Somenewspecieso\'Delphaco-
foridentifyingmosses,P. D. McMillan(De- dex(continued) (Homoptera. l-"uIgoridae. Delpha-
partment of Biological Sciences, Clemson cinae).PartIII.JournaloftheKansasEntomolog-
University) for preparing moss specimens icalSociety21: 1-10.
and sending them to L. E. Anderson, W. K. Behnkine,alHg.ae-,D.moIs97s.eSs.Pahnldofeemrnst,ispspu.ea1n8d7-s2i1e0v.eeIlneAmreonnt-s
Reeves (Department ofEntomology,Clem- oft,S.,J.Dainty,P R.Gorham.I..M.Srivastava.
son University) forci)llecting the planthop- and C. A. Swanson. eds. Phloem Transport. Ple-
per in Great Smoky Mountains National nuin Press,NewYork.
Park, K. G. A. Hamilton (Agriculture and Borner, C. 1952. Die Blattlau.sgattungen Myziis und
Agri-Food Canada, Eastern Cereal and Oil- Myzodes. BeilriigevurEntomologie2: 122-127.
seed Research Centre, Ottawa) for provid- Buck.claWs.sifRi.caatinodnBo.fmGoofsfsiense,t.pp2.00701.-12M.o3r.pIhnolSohgayw,anAd.
ing collection data forCanadian specimens .1.and B.Goffinet,eds. BryophyteBiology.Cam-
of J. opaca, D. E. Linvill (Department ol bridge UniversityPress,Cambridge.
Agricultural and Biological Engineering, Hurbanck.M.PandD.L.Phillips. 1983.Evidenceof
Clemson University) for providing meteo- plantsuccessionongraniteoutcropsoftheGeor-
rological data, J. F. Stimmel (Bureau of gia Piedmont. American Midland Naturalist 109:
APlgarnitcullituuirues,lryH,arPreinsnbsuyrlgv)anfiorapDheoptaorgtrmaepnhtinogf Campb9Gie4ll-ll.1.0419.B9.4.C.EvoJl.utDi.onSatreyffoerni-gCinamopfbewlhli.tclalnidesR(.HeJ-.
J. opaca. P. C. Huth (Mohonk Preserve, miptera: Sternorrhyncha: Aleyrodidae) inferred
New Paltz, N.Y.) for facilitating access to from I8S rDNA sequences. Insect MolecularBi-
Sam's Point DwarfPine Ridge Preserve, J. ology3: 73-88.
R. Allison (Georgia Department ofNatural Collins, N.J. andW.C.Oechcl. 1974. Ihcpatternof
Resources, Social Circle)foraccompanying growth and translocation of photosynlhaie in a
me in the field at granite outcrops in Geor- JtouunrdnraalomfosBso.taPnoyly5t2r:ic3h5i5i-m36c3d.piniini. Canadian
gia, R. W. Brooks (Department of Ento- Criim,II.|A.|. 1976.MossesoftheGreatLakesForest.
mology, University of Kansas, Lawrence) University Herbarium. University of Michigan,
for attempting to locate R. H. Beamer's Ann Arbor.404 pp.
field notes, and P. H. Adler(Departmentof Crum, H. A. and L. E. Anderson. 1981. Mosses of
Entomology, Clemson University) for pro- EasternNorthAmerica.Vol.2.ColumbiaUniver-
viding comments that improved the manu- sity Press,NewYork,pp.665-1328.
script. Two reviews secured by theeditor DennOol.msRt.eaEd..M.19E8.7.ScPhraaucftfi,caSl.dWi.agWniolssison,anadndnaKt.uraI.l.
one submitted anonymously, the other by historyoftwosiblingsaltmarsh-inhabitingplant-
C. R. Bartlett (Department of Entomology hoppersinthegenusProkelisia(Homoptera:Del-
& Applied Eco—logy, University of Dela- phacidae). ProceedingsoftheEntomological So-
ware, Newark) also improved the manu- cietyofWashington89: 687-700.
script. Deida,G.S. 1998.Geneticvariationandrclalionships
among the hair-cap mosses (Polytrichaceae).
LniiKArtiRE Cited Ph.D.Dissertation.UniversityofGeorgia,Athens.
Aschc, M, mS.S. /ill Phykigenie dcr Dclphacidae 197 pp.
Leach, ISl.'i (Honioptcia: Cicadiiia: Eulgoniinor- Derda.G.S.and R. Wvatl. 1990.Genetic variationin