Table Of ContentPROC. ENTOMOL. SOC. WASH.
96(4), 1994, pp. 701-712
REVISION OF THE MYRMECOMORPHIC PLANT BUG GENUS
SCHAFFNERIA KNIGHT (HETEROPTERA: MIRIDAE: ORTHOTYLINAE)
Thomas Henry
J.
Systematic Entomology Laboratory, Plant Sciences Institute, ARS, USDA, % National
Museum ofNatural History, MRC-168, Washington, D.C. 20560.
Abstract.—Tht myrmecomorphic plant bug genus Schaffneha is reviewed, the species
5. hungerfordi Knight is synonymized under S. davisi Knight, and S. bweni Knight and
5. schaffneri Knight are synonymized under 5. pilophoroides (Knight). The genus and the
two included species are redescribed, new host and distribution records and comments
on ant mimicry are provided, and the relationship of Schaffneria to other genera is
discussed.
Key Words: Heteroptera, Miridae, Schaffneria, synonymy, distribution, hosts, habits,
ant mimicrv. North America
Miridae of the ceratocapsine genus Kelton (1980) later reported S. davisi from
Schaffneria Knight are remarkable in their Manitoba and 5. schaffneri from Alberta
resemblancetocertain speciesofants. Until and Saskatchewan, considerable range ex-
recently, little was known of their habits. tensions for both species.
Knight (1966) proposed the genus Schaff- It was not until Wheeler's(1991)workon
neria to accommodate Pamillia davisi the mirids of scrub oak, Quercus ilicifolia
Knight, known only from New Jersey, Cer- Wangenh., that more life-history informa-
alocapsiispilophoroides Knight, known only tion on these interesting mimetic bugs be-
from Kansas; and his new species Schaff- cameavailable. Duringhis research on pitch
neria burenifrom Louisiana, S. hungerfordi pine-scrub oak barrens in the Eastern Unit-
from Michigan, and S. schaffnerifrom Tex- ed States, he discovered that both S. davisi
as. The last-named species was discovered and S. pilophoroides (as S. schaffneri) are
bythe lygaeid specialist M. H. Sweet (Texas closely associated with Dolichoderiis ants
A&M University, College Station), who tending certain species ofaphids.
found specimens running on the ground in Primarily because ofspecimens collected
a dry juniper habitat and speculated that by Wheelerand those accumulated through
these bugs might be feeding on fallen cedar our combined fieldwork, I have been able
fruits, a habit that would be highly unusual to reevaluate the species of Schaffneria.
for mirids. As a consequence ofSweet and Study of the types has allowed me to de-
A&M
J. C. Schaffner's (also Texas Univer- termine that three ofthe five described spe-
sity) observations. Knight (1966) entitled cies are junior synonyms. In this paper I
his paper ""Schaffneria. a new genus of synonymize 5. bureni and S. schaffneri un-
ground dwelling plant bugs ." In that der S. pilophoroides and 5'. hungerfordi un-
. . .
paper. Knightalsospeculatedthatthesebugs der S. davisi, diagnose and redescribe S.
were ". perhaps associating with ants." davisiandS.pilophoroides, providenewhost
. .
702 PROCEEDINGSOFTHE ENTOMOLOGICALSOCIETY OFWASHINGTON
anddistribution records, and discuss habits corium inside and basal area outside radial
and the relationship ofSchaffheria to other vein appearing dull or matlike at stereo-
genera ofCeratocapsini. scopicmicroscopemagnificationsof63x or
less (Figs. 4, 12), actual cuticular surface
Schaffneria Knight thickly covered by numerous tiny spicules
(Fig. 13); narrow band through middle of
Schaffheria Knight, 1966: 1. Type species: corium and clavus(dulled area) with a glau-
Schaffheria schaffneri Knight, 1966, aju-
coussheen, cuneusand most ofcorium out-
nior synonym of Ceratocapsus pilopho-
side ofradial vein polished; surface with a
roides Knight, 1930. Original designa-
few scattered, short, erect, bristlelike setae.
tion.
Ventral surface polished, except for the
Diagnosis.—This genus is unique in hav- glaucous metapleural area above the osti-
ing the combination ofa posteriorly flared olar peritreme and coxae. Male genital cap-
and convex pronotum, apically broadened sulewitharelativelysmall,shallow,circular
hemelytra, the irregulardulled band through aperture, dorsal edge irregularly tubercu-
the middle ofthe corium and clavus, a pol- late. Left paramere slender, sickleshaped;
ished cuneusand area ofthecorium outside right paramere short, sickleshaped to api-
the radial vein, the reduced male genital cally flared; phallotheca thickened, apically
aperture, and the sickle-shaped left and re- truncate.
duced right paramere. Remarks.—The relationships of genera
Description.—Orthotylinae: Ceratocap- within the Orthotylinae are poorly under-
sini. Myrmecomorphic. Length 2.80-3.80 stood, making it diflicult to place Schaff-
mm. Head subtriangular in dorsal aspect, neria in a phylogenetic context. Knight
elongate in lateral aspect, impunctate, basal (1966) placed Schaffheria in the tribe Cer-
margin straight, sharply but narrowly cari- atocapsini with the genera Ceratocapsus
nate, basal angles ofeyes weakly rounded, Renter, Pamiltia Uhler, and Pilophoropsis
eye elongate oval in lateral aspect, occu- Poppius. Although the tribe has been rec-
pyinghalfor more ofhead, buccula narrow, ognized in a number of recent papers (e.g.
extending nearly to base of head, rostral Henry 1979, 1985, Carvalho et al. 1983,
openingbetween bucculae broadly rounded Henry and Wheeler 1988), it also has been
to contain all but apex ofthickened rostral considered a synonym ofOrthotylini (Car-
segment I. Antenna evenly thickened, ex- valho 1958, Schuh 1974). Schuh (1974)
cept for gradually narrowing basal half of characterized what he called the Serico-
segment II. Pronotum trapeziform, im- phanes group for genera with a well-devel-
punctate, shiny, broadly flared posteriorly, oped collar, some degree of ant resem-
posteriorwidth much broaderthan anterior blance, usually lacking spiculi, hemelytra
width, lateral margins broadh concave from often with maculae or faciae, and restricted
humeral to anterior angle, disc moderately to the Western Hemisphere (with the ex-
to strongly convex, each anteriorangle with ception ofXenofiilvius Bergroth known from
an erect, bristlelike seta. Scutellum trian- the Philippines). Schuh's group includes es-
gular, shiny, impunctate, arched upward sentially the same taxa that are now placed
from base to meet posterior margin ofpro- in Ceratocapsini, plus the genus Serico-
notum, apical halfflattened to weakly swol- phaiies. Until a comprehensive cladistic
len or bulbous. Hemelytra subparallel on analysis of the Orthotylinae (now under-
basal half, broadly flared from apical half way) can be performed, I am maintaining
ofcorium to baseofcuneusgivingbasal half Ceratocapsini to include the above genera,
a constricted appearance; clavus and all of as well as a number of other New World
VOLUME NUMBER
96. 4 703
taxa suchasEwotusDistant. Marinonicoris that S. davisi could be separated from S.
Carvalho, Renodaeus Distant, Zanchisme hungerfordiby the wider vertex and slightly
Kiri^aldy, and the genera discussed below. thicker antennal segment III, my study of
Knight (1966) commented that Schaff- thetypesandotherspecimensindicatesthat
neriawasalliedtoPamilliaUhlerandcould Knight apparently mismeasured the head
bedistinguished by thepolishedareasofthe characters. The diameters of the antennal
corium and cuneus (versus frosted or pru- segments are variable and do not support
inose in Pamillia) and the strongly convex the recognition of two species. The male
pronotum (versus slightly convex in Pam- genitalia ofthe holotypes also are indistin-
illia). Both genera have a similar myrme- guishable and indicate that S. hungerfordi
comorphic habitus, a characteristic shared should be placed as ajunior synonym ofS.
by many Ceratocapsini as noted above and davisi.
appearing convergently in numerous other Description.—.Va/t' (n = 10): Length
subfamilies and tribes of the Miridae (e.g. 3.52-3.80 mm, basal width 1.08-1.18 mm,
Wheelerand Henry 1980. Schuh 1984. Pol- width across widest part of corium 1.36-
hemus and Polhemus 1985, Mclver and .46 mm. Head:Width0.84-0.86 mm, ver-
1
Stonedahl 1993). Schajfneria and Pamillia tex 0.38-0.42 mm. Rostrum: Length, 1.30-
alsoshareareducedgenitalaperture, sickle- 1.36 mm. extending to middle coxae. .4;?-
shaped left paramere, and stout, apically tenna: Segment L length 0.30-0.32 mm; IL
truncatephallotheca, placingthemnearsev- 1.04-1.10mm; IH,0.64-0.70mm; IV,0.60-
eral other ceratocapsine genera and species 0.66 mm. Pronotum:Length 0.80-0.86 mm,
groupssuchas CeratocapsellaCarvalhoand anterior width 0.72-0.74 mm, posterior
Schaffner. Tiicuruiella Carvalho. and the width 1.16-1.18 mm.
Ceratocapsuspunctulatus group. Female (n = 10): Length 3.52-3.84 mm,
basal width 1.12-1.16 mm, width across
Schaffneria davisi (Knight) widest partofcorium 1.50-1.58 mm. Head:
Figs. 1-9; map 1 Width0.88-0.90 mm.vertex0.42-0.44mm.
Rostrum: Length 1.30-1.34 mm. Antenna:
Pamillia davisi Knight. 1923: 535.
Pam1i2l1.ia [sic] davisisi [sic]: Carvalho. 1958: S1e.0g6memnmt;I,IIlI,en0g.t6h4-00..2686-0m.m30;mIVm,;0.II6,2-10..0626-
mm. Pronotum: Length 0.80-0.88 mm, an-
Schaffneria dayisi: Knight. 1966: 3; Kelton. teriorwidth 0.76-0.80 mm, posteriorwidth
1980: 213; HenryandWheeler. 1988: 400: 1.14-1.18 mm.
Wheeler, 1991: 424.
General coloration black to reddish
Schaffneria hungerfordi Knight. 1966: 3;
NEW brown. Head black to reddish brown, sur-
Henry and Wheeler, 1988: 400.
SYNONYMY. face finely grained oralutaceous. with a few
scattered short, erect, bristlelike setae. An-
—
Diagnosis. 5. davisi is distinguished tennal segments almost uniformly thick-
from S. pilophoroides by the greater length, ened, segment IIgraduallynarrowedtoward
broader body form, shorter 2nd antennal base; segment I yellowish brown, even on
segment that is shorterthan the basal width dark color forms, segments I, III. and IV
ofthe pronotum. the shorter 3rd antennal fuscous. Pronotum black to reddish brown,
segment that is shorter than the width of impunctate, surface shiny, weakly aluta-
thehead, theflattenedscutellumthatislevel ceous, nearly glabrous, except for a long,
with the hemelytra, and quite different erect, bristlelike seta at each anterior angle
broad, apically truncate right paramere. and a few scattered short setae around basal
Although Knight (1966) said in his key and lateral margins,discstronglyswollen or
PROCEEDINGSOFTHE ENTOMOLOGICALSOCIETY OFWASHINGTON
704
VOLUME 96, NUMBER 4 705
Figs. 2-5. Scanning electron microscope micrographs ofSchafjneria davisi. 2. lateral aspect ofhead, pro-
notum,and scutellum(54.2x). 3. peritremeandostiolarevaporativearea(278»). 4, dorsalaspect ofhemelytra
(46.3X). 5, caudal aspect ofmale genital capsule (212x).
convex, raised well above anterior half or cous, narrowly pale along edges bordering
level ofhemelytra. Scutellum black to red- cuneus and around apical margins. Ventral
dish brown, basal half strongly arched up- surface shiny to polished; abdomen always
ward to meet basal margin ofconvex pro- polished black, with a large, quadrate, glau-
notum, posterior halflevel with hemelytra: couspatchacrosssegment III, endingbefore
apexdulled, texturesimilartodulledhemel- lateral margins; thoracic sterna fuscous to
ytral surface. Hemelytra uniformly black or reddish brown; ostiolar peritreme whitish.
fuscous to bicolored (i.e. reddish brown on Legs fuscous to reddish brown; tarsomere
basal half fuscous to black apically): lateral I, basal halfof tarsomere II, and apices of
marginsofbasal halfsubparallel, apical half coxae pale. Male genitalia; left paramere
beginningat middleofcoriumbroadly flared (Fig. 6); right paramere (Fig. 7); aedeagus
around to cuneus; clavus and basal halfof and phallotheca (Fig. 8); dorsal edge ofap-
corium appearing dull or matlike (Fig. 4), erture (Fig. 9).
but see Fig. 13; middle ofclavus and basal Remarks.—The color polymorphism
area of corium with a glaucous sheen, set found in S. davisi is remarkable. Popula-
with short recumbent, pale setae; area out- tions of this species in Pennsylvania that
side ofradial vein polished, including em- were found running with the entirely black
bolium, apical half of corium, and all of ant Dolichoderus tascheubergi Mayrare en-
cuneus; membrane and veins fumateto fus- tirely black, whereas populations discov-
Fig. 1. Dorsal habitus ofSchaffneria davisi.
706 PROCEEDINGS OFTHE ENTOMOLOGICALSOCIETY OFWASHINGTON
Figs.6-9. MalegenitaliaofSchaffnenadavisi.6,leftparamere.7,nghtparamere.8,aedeagusandphallotheca.
9. dorsal edge ofmale genital aperture.
ered in Georgia that were found with the Wangenh., black oak, Q. velutina Lamb.,
red and black D. mariae Forel are similarly and upland willow, Salix humilis Marsh.
bicolored. SeeMclverandStonedahl(1993) The plant host of this bug appears sec-
for an excellent review of color and trans- ondary to its association with certain spe-
formational polymorphism in myrmeco- ciesofantsthattendaphids. Wheeler(1991)
morphic spiders and insects, including nu- observed S. davisi running with Dolichod-
merous examples in the Miridae. erustascheubergithatwastendingthe shiny
Distribution (Map 1).—New Jersey black aphid Lachints allegheuiensis Mc-
(Knight 1923); Michigan (Knight 1966, as Cook and the yellowaphid Myzocallisbella
5". himgerfordi); Manitoba (Kelton 1980); (Walsh). M. D. Schwartz (see "Specimens
and Maine, Massachusetts, New Hamp- examined") collected this species with D.
shire, and Pennsylvania (Wheeler 1991). laschenbergias well, but on Salix humilis it
New records are Georgia for the United tended another aphid Chailophonis nigri-
States and Ontario for Canada. ceiitris Richards. In Georgia, A. G. Wheeler
Hostsand habits.—This species is known and I collected this species with D. mariae
from bur oak, Q. macrocarpa Michx. (Kel- Forel tending the aphid Neosymdobius sp.,
ton 1980) and scrub oak, Qiiercus ilicifolia near quercihabitus Miller. At this locality,
(Wheeler 1991). On the basis of specimen we observed one individual preying on an
data, it also occurs on post oak, Q. stellata unidentified collembolan. Wheeler (1991)
-
3
VOLUME NUMBER
96, 4 707
%J^r^
'
\ ^
Map 1. Dislnbution ofSchaffnenadavisi. O = new records. old records.
noted that 5. davisi fed on crushed cater- tris tended by DoUchoderus taschenhergi
pillarsand on aphids unprotected from ants (CNC). UNITED STATES: GEORGIA-
in the laborator\', and speculated that in na- (5<?, 12 99, Columbia Co., Georgia. Burk's
ture aphids were its primary prey, perhaps Mt., nr. Pollard's Corner, 24 Apr. 1988, T.
supplementingthis diet with honeydewand J. Henry(TJH)&A. G. Wheeler,Jr. (AGW),
scavenging. Almost certainly these mimetic taken on Quercusstellatawith DoUchoderus
mirids gain a protective advantage from manac tending Neosymdobius sp., near
predatorsbycloselyresemblingants. Whee- qiiercihabitus (USNM). MAINE-2 SS, Ox-
ler (1991) "hypothesized that 5. davisi de- ford Co., Fryeburg Barrens, 10 Aug. 1990,
rives protection from visual predators that AGW, on Quercus ilicifolia (USNM); 1 <5, 3
have learned toavoid noxiousantsand rep- 99, York Co., Waterboro Barrens. 7 Aug.
resents a classic Batesian mimic." 1990, AGW, on Quercus ilicifolia: 1 $, 6 99,
Specimens examined.—CANADA: York Co., Shapleigh Barrens, 9 Aug. 1990,
MANITOBA- 1 9, Horton, 25 July 1953, AGW, on Quercusilicifolia(USNM). MAS-
Brooks & Kelton (CNC); 1 9, Russell, 21 SACHUSETTS-2 <5<?, 1 9, Franklin Co.,
July 1954. Brooks & Wallis (CNC). ON- Montague Barrens, 4 Aug. 1990, AGW, on
TARlO-4 6$, 18 99. Ottawa, Carlton Co., Quercus ilicifolia (USNM). MICHIGAN
Constance Bay, 16-18 July 1991, M. D. holotype c5, allotype 9 (of S. hungerfordi),
Schwartz, taken on Salix hiiinilis var. hu- Cheboygan Co., 14 July 1931, H. B. Hun-
milis. collected with Chailophonis nigricen- gerford (USNM); 2 SS, 2 99, Presque Isle,
708 PROCEEDINGSOFTHE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Map 2. DistributionofSchaffneriapilophoroidcs. O = new records. • = old records.
Rt. 33. at Montmorency Co. line, 26 July Frackville. 15 Aug. 1986. TJH & AGW.
1991. TJH & AGW, on Quercus velutina taken on Quercus ilicifolia associated with
(USNM). NEW HAMPSHIRE- 1 6. 1 9, Dolichoderus taschenbergi (USNM).
Carroll Co., Ossipee Barrens, 10 Aug. 1990,
AGW, taken on Quercus ilicifolia (USNM).
NEW JERSEY-2 29, Jamesburg. 4 July Schaffneria pilophoroidcs (Knight)
1902. H. G. Barber colln. (USNM); holo- Fig. 10-19; map 2
type 9 (ofPamillia davisi). Lakehurst, "IX- Ceralocapsus pilophoroidcs Knight. 1930:
4," H. H. Knight colln. (USNM); 1 S, 3 99, 197;Carvalho. 1958: 48.
Lakehurst. 20June 1903, H. G. Barbercolln. Schaffneria pilophoroidcs: Knight. 1966: 1;
(USNM); paratype 9 (of P. davisi), Manu- Henr> and Wheeler. 1988: 400.
muskin. 15 Sept. '02 (CNC). PENNSYL- Schaffneria bureni Knight. 1966: 5; Henry
VANIA-2 SS. 2 99. Centre Co.. Scotia Bar- and Wheeler. 1988: 400. NEW SYN-
rens. 19 July 1990. AGW. takenon Quercus ONYMY.
ilicifolia (USNM); 15 SS. 24 99. Schuylkill SchaffneriaschaffheriKn'ighx. 1966: 2; Kel-
Co., Rt. 81, 5 mi. S. Frackville, 22-23 July ton. 1980: 213; Henry and Wheeler. 1988:
1986. 1 Aug. 1986. 6 July 1988. 15 Sept. 400; Wheeler. 1991: 427. NEW SYN-
1990. AGW, taken on Quercus ilicifolia as- ONYMY.
sociated with Dolichoderus taschenbergi
tendingLachnus allegheniensis(USNM); 13 Diagnosis.—5. pilophoroidcs is distin-
SS, 9 99, Schuylkill Co., Rt. 81, 5 mi. S. guished from 5. davisibytheshorterlength.
VOLUME 96. NUMBER 4 709
Figs. 10-15. Scanning electron microscope micrographs o(Schaffneria pilophoroides. 10, lateral aspect of
head, pronotum, and scutellum (70.5x). 11. peritreme and ostiolarevaporativearea (285x). 12, dorsal aspect
ofhemelytra (63.0x). 13. split-image dorsal aspect ofhemelytra (438x; 1752x). 14, claw (925x). 15, caudal
aspect ofmale genital capsule (247x).
more slender body form, the longer 2nd an- Knight (1966) separated S. pilophoroides,
tennal segment that is much longer than the S. schaffneri, and S. bureni entirely on the
basal width ofthe pronotum, the longer 3rd color of the wing membrane and infusca-
antennal segment that is subequal to orlon- tions on the pronotum and hemelytra. This
ger than the width ofthe head, the apically variation occurs in longer series of speci-
swollen scutellum that is distinctly raised mens, except perhaps the mostly "white"
above the level of the hemelytra. and the membrane of S. pilophoroides (hoiotype),
short, slender, apically acute right para- which appears abnormally bleached. As
mere. noted above, specimens vary from reddish
710 PROCEEDINGS OFTHE ENTOMOLOGICALSOCIETY OFWASHINGTON
Figs. 16-19. Malegenitaliao{Schaffneriapilophoroides. 16, leftparamere. 17, rightparamere. 18,aedeagus
and phallotheca. 19. dorsal edge ofmalegenital aperture.
brown to almost completely fuscous or Width0.80-0.82mm,vertex0.38-0.44mm.
black. The greatest difference among types Rostrum: Length 1.32-1.34 mm. Antenna:
is the submacropterous condition of the Segment L length 0.30-0.32 mm; IL 1.12-
hemelytra on the holotype (S. schajfneii) 1.20 mm; III, 0.74-0.76 mm; IV, 0.62-0.64
from Texas. However, both fully macrop- mm. Pronotum: Length 0.66-0.76 mm, an-
terousandsubmacropterousindividualsare terior width 0.64-0.74, posterior width
found together in series collected in Penn- 0.96-1.04 mm.
sylvania. Male genitalia are indistinguish- General coloration fuscous to reddish
able among the types and other specimens brown. Head fuscous to reddish brown,
I have examined. finely alutaceous, weakly shiny, with a few
Description.—Ma/£'(n = 7): Length 2.88- scattered erect setae. Antenna uniformly
3.16 mm, width across base 0.92-0.96 mm. brown, but segments III. IV. and apical half
width across widest part of corium 1.00- of II often fuscous. Pronotum impunctate,
1.08 mm. Head:Width 0.76-0.78 mm. ver- black to reddish brown, shiny, finely alu-
tex 0.36-0.40 mm. Rostrum: Length 1.28- taceous, moderately convex, disc and an-
1.32 mm. extending to bases ofmetacoxae. terior half evenly rounded, with a single
Antenna: Segment L length 0.30-0.32 mm; bristlelike seta at each anterior angle and a
IL 1.20-1.22 mm: IIL 0.74-0.80 mm; IV. few scattered, short, recliningand semierect
0.60-0.62 mm. Pwnotum:Length0.66-0.70 setae. Scutellum fuscous to reddish brown,
mm, anterior width 0.66-0.70 mm, poste- uniformly shiny, basal third arched upward
rior width 0.94-0.98 mm. to meet base ofpronotum, distal two thirds
Female (n = 7): Length 2.92-3.20 mm, bulbous, swollen above level ofhemelytra.
basal width 0.98-1.04 mm, width across Hemelytrabicolored (i.e. reddish brown ba-
widest part ofcorium 1.22-1.38 mm. Head: sally; fuscous apically) to uniformly fuscous
