Table Of ContentPROC. ENTOMOL. SOC. WASH.
97(2), 1995, pp. 258-330
A RECLASSIFICATION AND PHYLOGENY OF THE TERMATOPHYLINI
(HETEROPTERA: MIRIDAE: DERAEOCORINAE), WITH A TAXONOMIC
REVISION OF THE AUSTRALIAN SPECIES, AND A
REVIEW OF THE TRIBAL CLASSIFICATION OF
THE DERAEOCORINAE
Gerasimos Cassis
Australian Museum, 6-8 College Street, P.O. Box A285, Sydney South, N.S.W. 2000,
Australia.
Abstract.—The genera of Termatophylini are described, including two new genera,
DemocorisandKundakimuka, fromAustralia. Akeyisgiventothetermatophylinegenera.
The type species of all the genera are described, and six new species ofAustralian ter-
matophylines, Kundakimukacarvalhoi, K. queenslandica, Democorisiugens, D. leptocytus,
Termatophylum melaleucae and T. weiri, are described. A key is given to the Australian
termatophylinespecies. Thetribeisre-definedonthebasisoftheshortfirstlabialsegment,
the presence ofantero-lateral pronotal setae, and the exposed metathoracic spiracle. The
genera Conocephalocoris, Hesperophylum and Termatomiris are removed from the Ter-
matophylini and placed in the Deraeocorini. A cladistic analysis ofthe Termatophylini
is provided on the basis of 29 morphological characters. The sister-group relationships
are discussed and the characters are reviewed. Characters of the pterothoracic pleura,
particularly the metathoracic spiracle and the scent efferent system ofthe metathoracic
glands are extensively utilized in this study. These characters are also discussed for other
supragenericgroups ofMiridae. The tribal classification ofthe Deraeocorinae is reviewed,
and it is concluded that none ofthe other tribes are adequately defined. It is also firmly
established that the termatophylines are mirids, and that any similarity with the Antho-
coridae is due to convergence.
Key Words: Heteroptera, Miridae, Deraeocorinae, Termatophylini, revision, phylogeny,
new genera, new species, Australia
TheTermatophylinihavehadalabilepo- the Deraeocorinae, there has been little dis-
sition within the classification ofthe Cim- pute as to their monophyly nor their fa-
icomorpha, varying from family status milial or subfamilial position. This view is
(Reuter 1910, Wagner 1970), asa subfamily supported in this work despite the present
within the Anthocoridae (Reuter 1884a, b) conclusion that the previous tribal defini-
and Miridae (China and Myers 1929), and tion,basedprimarilyonaporrecthead, nei-
more recently as a tribe of the mirid sub- ther holds for the tribe, nor is exclusive for
family Deraeocorinae (Carvalho 1952, other Deraeocorinae.
1955a, 1957). Mention has been made of This work was initiated by the receipt of
their salient similarity with some members aboxofmirid specimensfrom the Museum
of the Anthocoridae (Schuh 1976). Curi- and Art Galleries of Northern Territories,
ously, afterCarvalho (1952) placedthem in amongst which, one specimen ofa terma-
VOLUME 97, NUMBER 2 259
tophyline from Smith Point, near Darwin, termatophylines, and a cladistic analysis of
broughtforththequestion, "whyisthismir- the genera. Jose Carvalho's contributions
id so Hke an anthocorid?" This exaggerated on the Miridae, including the Australian
specimen has gigantic hind femora, some- fauna, remains unparalleled. Cassis and
what Hke those found in members of the Gross (in press) report that Carvalho de-
Halticini (Miridae: Orthotylinae). Aside scribed more than half of the Australian
from this apomorphy, the similarity of its mirid fauna. He had also commenced nu-
overall facies with anthocorids is striking, merous projects on the Australian Miridae,
and the absence ofocelli and presence ofa and much of the termatophyline material
hemelytral membrane cell, suggested an al- in this work has passed through his hands,
ternative taxonomic arrangement. Charac- including Democohs lugens n. gen. and n.
ters ofthe male genitalia and the presence sp. I have provided a dedication to him by
of femoral trichobothria strongly support naming the unique Smith Point species in
the placement ofthe termatophylines with- his honour, under the new generic name,
in the Miridae. The anthocorid-like fea- Kundakimuka.
tures, however, provided the impetus for In this work I have recognized eight gen-
making an analysis ofthe termatophylines, era and 30 species ofTermatophylini. The
mindful ofpresent definitions ofthe Miri- genus Democoris is restricted to south tem-
dae, Anthocoridae and other cimicomor- perate Australia. The genera Kundakimuka
phan families (Schuh and Stys 1991). and TermatophylumReuterarebroadlydis-
In this work I also support the placement tributed in the Eastern Hemisphere, includ-
of the termatophylines in the Deraeocori- ing Australia. Termatophylella Carvalho,
nae, but in redefining them as a monophy- Termatophylidea Reuter and Poppius, Ter-
letic tribe, I have discovered that the other matophyloidesCarvalho are restrictedto the
deraeocorine tribes are presently inade- Western Hemisphere. TermatophylinaCar-
quatelydefined. I havegiven acommentary valho is restricted to the Oriental region.
on the characters presently used in these Arygrotelaenus Reuter and Poppius is re-
classifications and suggest alternative char- stricted to the Middle East. I have provided
acters, particularly of the pterothoracic a key to the world genera, which is based
pleura. In particular I have introduced a entirely on external characters, and a key to
morphological comparison ofthe metatho- all known Australian species. In redefining
racic spiracular opening and the scent ef- theTermatophylini I havetransferred Con-
ferent system ofthe metathoracic glands in ocephalocorisKnight, Hesperophylum Reu-
the Miridae, to reappraise previous char- ter and Poppius and Termatomiris Ghauri
acters, such as the basally cleft claws, pri- to the Deraeocorini.
marily used to place the termatophylines
Materials and Methods
within the Deraeocorinae.
An invitation to provide a contribution About 1 10 specimens oftermatophylines
to a Festschrift for Dr. Jose Carvalho, ren- were examined in this study. This material
dered this idea into its present form. The was borrowed from the Natural History
Termatophylini have not been previously Museum, London and the National Muse-
recorded from Australia. In accumulating um ofNatural History, Smithsonian Insti-
all the Australian termatophyline material tute, Washington, andthe majorcollections
itwasclearthat notalltermatophylineswere found within Australia. The specimens of
anthocorid-like, which led me to study the the type specimens were examined. No
Australian fauna, redescribe the genera maleso{ArygrotelaenuselegansReuterand
worldwide, including a checklist ofspecies, Poppiuswereavailableforexamination.The
phylogenetic definition and position ofthe generic definitions incorporate observa-
260 PROCEEDINGSOFTHEENTOMOLOGICALSOCIETY OFWASHINGTON
tions of most termatophyline species. Not clature of the scent efferent system com-
all species of Termatophylum and Terma- ponentshasnotprovenasuseful, andI have
tophylidea were examined, but the descrip- preferred to use a translation ofsome ofthe
tions in the literaturewere sufficient to con- terminology of Carayon (1971). I refer to
fidently retain within them all previously the gutterlike channel which emerges from
assigned species. the ostiole as the peritreme. This structure
The male genitalic characters were ex- is usually tonguelike and extends to the lat-
amined on temporary mounts in glycerin, eral margins ofthe metepistemum. Under
usingbothaZeissSRStereomicroscopeand light microscopy it often appears shiny,
a Leitz Laborlux S Compound microscope. whereas under scanning electron micros-
The line drawings were prepared using a copy it is usually covered with microsetae.
camera lucida. Scanning electron micro- The peritreme is surrounded by highly dis-
graphsweremadefromdriedmuseum spec- sected cuticle, which has a granulate ap-
imens, using a Cambridge SI20 micro- pearance and is referred to as the evapo-
scope. rativeareas. Theevaporativeareasarecom-
posed ofnumerouscomponents, and in this
Taxonomic Characters
work I referchiefly to the evaporative caps,
The morphological terminology used in which are raised and flattened. The char-
this work is derived from various sources. acters of the scent efferent system are la-
Many of the terms used for somatic char- beled in Figs. 57, 119 and 123. I have in-
acters are now conventional, following the cluded in the Phylogenetic analysis section
modemworksofauthorssuchasSchuh(e.g. below an evaluation ofthese characters in
1984). Some original interpretation was in- other Miridae.
volved in my treatment ofthe forewing, the The forewing venation of the termato-
pterothoracic pleura and the malegenitalia. phylines has been used extensively in this
In this work I introduce the use ofcom- work, both as diagnostic and phylogenetic
parative morphology of the pterothoracic characters. The terminology and homology
pleura, particularly thecondition ofthe me- oftheheteropteran forewinghasbeenpartly
tathoracic spiracle and the scent efferent resolved by Tanaka (1926), Wootton and
system ofthemetathoracicscentglands. The Betts (1986), and Betts (1986). Carveret al.
homologies are partly derived from Cara- (1991) refer to the posterior margin ofthe
yon(1971) and Staddon (1979) forthe Het- embolium in the Miridae asthe R+M vein,
eroptera,andLarsen(1945)andSouthwood whereas Knight(1941)consideredthistobe
(1953) for the Miridae. Thejunction ofthe the costal vein. I accept the former inter-
mesepimeron and metepistemum contains pretation and concur with Betts (1986) that
in some Miridae, a visible opening of the the vein anteriad to the R+M vein is the
metathoracic spiracle (Figs. 57, 65). Its po- subcostal vein, which is evident in some
sition and associated structures are both Miridae. In this work I refer to the groove
R+M
phylogenetically and taxonomically signif- posteriad to vein as the median flex-
icant (Cassis 1984, Cassis in preparation). ion line. Knight (1941) referred to this
The external cuticular component of the grooveastheradialvein, butitsappearance
metathoracic scentglandsyields a wealth of suggests a flexion line, rather than a tra-
systematic characters. The terminology of cheate vein. Its position, posteriad to the
this system remains greatly confused, and posterior margin of the embolium, elimi-
R+M)
requires standardization across the Heter- natesitsinterpretationasaradial(or
optera. Staddon (1979) has referred to this vein. The median flexion line cannot be in-
system as the scent efferent system, which terpreted as the CuA vein, which according
I have adopted in this work. His nomen- to Betts (1986), is an unintermpted vein.
VOLUME NUMBER
97, 2 261
parallel and adjacent to the claval furrow. sclerites (Schwartz 1987) which originate at
The vein ofthe clavus is referred to as the or distally on the membranous lobes, and
first anal vein by Betts (1986), which I have as simple outgrowths ofthe membrane cu-
adopted in this work, and is synonymous ticle (equivalent to spicules or spiculae of
with theclaval veinofKnight (1941). Other other authors).
workers, such as Davis (1961), have pro- Stonedahl and Cassis (1991) in a revision
ducedvenationalterminologiesforthe Het- ofthe deraeocorine genus Fingulus Distant
eroptera, but it is beyond the scope ofthis described the male genitalia, and reported
paper to contrast in detail the various sys- homologiesandterminologiesthathaveap-
tems. The characters ofthe forewing are la- parent general value in the Deraeocorinae,
beled in Fig. 47. including the Termatophylini. They de-
The morphology ofthe male genitalia of scribed the male genitalia of Fingulus as
the Deraeocorinae has received little atten- possessing a basal tubular, sclerotized skirt
tion. Kelton (1959) gave a comparative surrounding the distal region ofthe ductus
morphological account, and reported con- seminis, a simple secondary gonopore, and
siderable diversity in form throughout the a distal multilobed membranous sac with
subfamily. He stated that the most distinc- lobal sclerites and/or fields ofspines. They
tive features included the flexible ductus found no evidence ofsclerotized processes
seminis, and the bulbous spiculate process- or basal processes.
es of the membranous vesica. Schwartz IntheTermatophylini theductusseminis
(1987) in a review ofMirinae and Deraeo- terminates at the base ofthe membranous
corinae male genitalia suggested that the componentofthe vesica (Figs. 1 1 1-1 18), as
secondary gonopore was ofcritical impor- reported for the Deraeocorini by Schwartz
tance, and that it differed in the two sub- (1987). The secondary gonopore is small,
families. He reported that the secondary simple and undifferentiated, as is found in
gonopore of the Deraeocorini terminated Fingulus. Aside from the genus Termato-
adjacent and usually basad of the mem- phylidea, the gonopore is associated with a
brane, and that the aperture is large and differentiated basal portion ofthe membra-
opens into a depressed trough, often with nous sac, which can be recognized as a V or
associatedserratesclerotizedplates. Healso U-shaped cavity marked by compact fields
stated that the secondary gonopores in the ofspines (Figs. 111-113, 1 15-1 18), and is
otherDeraeocorinaetribesweresmallerthan here referred to as the gonoporal cavity. It
those found in the Deraeocorini. Schwartz is difficult to ascertain in the Termatophy-
(1987) reviewed the terminology ofthe ves- lini whether the sclerites which are associ-
ical processes used in previous works (Kel- ated with the gonoporal cavity are lobal
ton 1959, Clayton 1982, Stonedahl 1983), sclerites (Figs. 111-113, 1 15-1 18), or rep-
suggesting that they were too simplified. resentauniquetypeofvesicalprocess. They
Stonedahl(1988)differentiatedtwotypes of are clearly outgrowths ofthe membranous
vesical processes in the mirine genus Phy- cuticle,butareneverfoundontheperiphery
tocoris Fallen, as follows: (1) sclerotized ofthe membranous lobe. This could not be
processes(equivalenttotheribbonlikestrap clarified because of the limited material
of Schwartz (1987)) which originate at or available for examination. In the Terma-
near the level of the gonopore, which is tophylini, the vesica is small and appears
sometimes continuous or associated with to be unilobed, and does not show any ob-
the basal sclerite (originating basad to the vious inflation as is evident in other Der-
secondary gonopore and not present in the aeocorinae and Mirinae. I have tentatively
Deraeocorinaeexamined), andvariouslyat- referred to these processes as lobal sclerites
tached to the membranous sac; (2) lobal pending further study. The distal regions of
262 PROCEEDINGSOFTHE ENTOMOLOGICALSOCIETY OFWASHINGTON
the membranous sac often have fields of achievedthroughoutgroupcomparison.The
spines (Figs. Ill, 116-118). selection ofoutgroupswas madedifficult by
The vesica of Termatophylidea is unlike current deficiencies in the tribal classifica-
any other termatophyline, in possessing a tion ofthe Deraeocorinae (Akingbohungbe
sclerotized process connected to the base of 1974, 1978, Stonedahland Cassis 1991). At
the ductus seminis and extending to near present six tribes are recognized; Clivine-
the apex of the membranous lobe. It also mini, Deraeocorini, Satumiomirini, Hy-
lacksanyobviousgonoporalcavityandthere aliodini, Surinamellini and Termatophyli-
arenolobal scleritesorfieldsofspines. This ni. A selection ofgenera from each of the
is reminiscent ofthe condition found in the tribes, aside from the ant-mimetic Surina-
dicyphine genus Singhalesia China and mellini, were initially included in the anal-
Carvalho and in some species ofHyaliodes ysis, to examine the distribution of char-
Reuter. The latter genus and Termatophy- acters throughout the Deraeocorinae. This
lideaaresuperficiallysimilar, butareclearly produced cladograms with low consistency
differentiated on characters ofthe head and indices, and the outgroups were then re-
the pterothoracic spiracle, and the presence stricted to two members ofthe Clivinemini
of a sclerotized process in both genera is {AmbraciusandBothynotus),andthegenera
considered to be independently derived. Conocephalocoris and Hesperophylum,
Stonedahl (personal communication) has which previously had been included in the
indicated that the sclerotized process isalso Termatophylini. The last two genera show
present in the clivinemines, Ambracius StM many similarities with the Termatophylini,
and Bothynotus Fieber. aspresentlydefined,particularlyinthehead
and hemelytral structure, and are consid-
ered as sister-groups. The Clivinemini gen-
Phylogenetic Methods erawere also included in adherence to mul-
The phylogenetic relationships ofthe ter- tiple outgroup principles (Maddison et al.
matophyline genera were examined using 1984), and were selected above other der-
cladistic methods (Wiley 1981). The clado- aeocorines because ofsome structural sim-
gram was produced, using in tandem, the ilarities with termatophylines, particularly
computer software HENNIG86 and CLA- in the form of the callosite region of the
DOS. These programsare designed forpro- pronotum. Genera of the Hyaliodini (An-
ducing cladograms of minimal length, and nona Distant and Hyaliodes Reuter), ini-
the manipulation ofthe characterdata. The tially included in the analysis, are similar
character data matrix was built in a text tosome Termatophylini, particularlythege-
editorandtransferredtoHENNIG86, where nus Termatophylidea, but were excluded
tree buildingalgorithmswere applied to the because they provided no additional clari-
input file. The final result was produced by fication to any ofthe character state trees.
using the Branch and Bound option, where The male genitalic characters were not ex-
the data were treated in a stepwise manner. amined in any ofthe outgroups because of
Choice between multiple trees of equal a lack of specimens. Stonedahl (personal
length was made by reweighting characters communication) provided information on
accordingto the rescaled consistency index. the male genitalia ofsome ofthe outgroup
The characters were coded as either bi- taxa.
nary or multistate characters. Most of the
multistate characters were ordered, except
Acknowledgments
for characters 4, 23 and 27 (see Table 2),
which were coded as non-additive, because I thank the following individuals and in-
there wasno evidence oftransformation se- stitutions for providing the specimens that
ries. Character state polarization was were usedin thisstudy. GaryM. Stonedahl,
VOLUME NUMBER
97, 2 263
International InstituteofEntomology, Lon- tarsal characters. Reuter (1884b) in his
don, Michael Webb, Natural History Mu- monograph ofthe anthocorids, recognized
seum, London (NHM), Thomas J. Henry, three subfamilies, Anthocorina, Microphy-
Systematic Entomology Laboratory, ARS, sina and Termatophylina, and provided a
USDA, % National Museum of Natural key to the subfamilies, differentiating the
History, Washington, D.C. (USNM), John twolattersubfamiliesfromtheAnthocorina
F. Donaldson, Queensland Department of bythe four-segmentedlabium. Subsequent-
Primary Industry, Brisbane (QDPI), Gra- ly, Reuter (1910) in his review of the sys-
ham Brown, Museum and Art Galleries of tematics ofthe Miridae, recognized the ter-
the Northern Territories (NTMAG), Jan matophylines as a family, separating them
Forrest, South Australian Museum, Ade- from the mirids by the presence ofan em-
laide (SAMA), and Tom A. Weir, Austra- bohum. Poppius (1910) supported the fam-
lian National Insect Collection, Canberra ily status, in describing a new Termatophy-
(ANIC). Carmen Zurl provided the habitus lum species from the Ethiopian Region.
illustrations ofKundakimuka queenslandi- Reuter and Poppius (1912) in the only
ca and Democohs lugens, and assisted in previous comprehensive paper on the ter-
the production ofthe photographic plates. matophylines retained the suprageneric
Sue Lindsay and GeoffAvem prepared the group as a family, givinga detailed descrip-
scanningelectron micrographs. Mary Carv- tion, and distinguishing them from the An-
er ofthe Australian National Insect Collec- thocoridae. They made mention ofthe por-
tion provided useful advice on forewingve- rect head and the short first labial segment,
nation. Gary Stonedahl assisted in many but erroneously concluded that the pretarsi
ways in this work and his prompt delivery lackedarolia (= parempodia). Schuh (1976)
of critical specimens, particularly of Con- inareviewofthemiridpretarsushas shown
ocephalocoris nasicus and Hesperophylum that setiform and fleshy parempodiaareho-
heidemanni, was vital. Both Gary Stone- mologous. Early European workers, such as
dahl and Toby Schuh provided useful re- Reuterand Poppius (1912), regarded fleshy
views ofthis paper. The Australian Biolog- parempodia as the only true "arolia," and
ical Resources Study provided a grant to their conclusion that termatophylines lack
producethiswork, andtheircontinued sup- "arolia" is a misinterpretation of setiform
port of mirid taxonomy is duly acknowl- parempodia. ReuterandPoppius(1912)de-
edged. Lastly, I would like to thank Tom scribed three new genera, Arygrotelaenus,
Henry and Al Wheeler for the opportunity Hesperophylum and Termatophylidea, and
to contribute to the Festschrift. seven new species. They also included a key
to the four known termatophyline genera.
Parshley (1923) described the family, large-
Historical Review
ly repeating Reuter and Poppius' (1912) di-
The first termatophyline, Termatophy- agnosis, but curiously suggested that ter-
lum insigne, was described by Reuter matophylinegenitalia were "much as in the
(1884a), who recognized this species as Anthocoridae."
unique, and erected a new subfamily, the China and Myers' (1929) seminal paper
Termatophylina, ofAnthocoridae forit. He on the classification of the cimicoid fami-
diagnosedthegrouponthebasisofthefour- lies, reappraisedtherelationshipsoftheter-
segmented labium, including the short first matophylines, and placed them within the
segment, the absence of ocelli, hemelytral Miridae. They concluded that apart from
structure, lack ofa hamus in the hindwing, the anthocorid-type head and the short first
and the three-segmented tarsi. He distin- labial segment, the termatophylines were
guished it from anotheranthocorid subfam- conclusively mirids, and treated them as a
ily, the Microphysina, on hemelytral and subfamily. They indicated that the porrect
264 PROCEEDINGSOFTHE ENTOMOLOGICALSOCIETY OF WASHINGTON
headconditionoccursintheCylapinae,and including prey and plant associations. This
noteditssimilaritytotermatophylines. Car- information was taken from the literature
valho (1951, 1952) analyzed the position of and label data. Myers (1935) was the first
the termatophylines, and concluded that to report termatophylines as predators, de-
they were a tribe of Deraeocorinae on the scribing Termatophylidea pilosa as an im-
basis ofthe basally cleft claws and setiform portant predator ofthe cacao thrips, Selen-
parempodia. He differentiated them from othrips rubrocinctus (Giard), a serious pest
other deraeocorines by the porrect head, of cacao in the tropical parts of the Neo-
large eyes, short antennae, and small body tropicalregion. Callan(1943, 1975)andvan
size. Carvalho (1955a) provided a key to Doesburg (1964) recorded other Termato-
thetermatophylinegenera, recognizingsev- phylidea speciesaspredatorsofthrips. Both
en genera, Hespewphylum, Termatophyli- T. maculataand T. opacaarealsopredators
dea, Arygrotelaenus, Conocephalocoris, of the cacao thrips, and Callan (1975) re-
Termatophylella, Tennatophyloides and ported that the former species also feeds on
Termatophylum. He distinguished the gen- the larvae of the grass thrips, Calliothrips
eramostly on headandpronotalcharacters, insularis(Hood). There are indications that
and the short antennae. othertermatophylines feed on moth larvae.
Aside from Wagner (1970), who retained Kundakimuka queenslandica feeds on the
the termatophylines as a family, no subse- xyloryctine moth, Xylorycta luteotactella
quent author has deviated from the tribal (Walker), which feeds on a paperbark spe-
placement ofthis group within the Deraeo- cies, Melaleuca integrifolia. I have exam-
corinae. Other works mostly include de- ined specimens of Termatophylina Indiana
scriptions of new genera and species. Car- from the Natural History Museum, which
valho (1955b) described new taxa from the has the label data: "associated with larval
Neotropical region, recognizing two new galleries of Lamida moncusalis,'' which
genera, Termatophyloidesand Termatophy- suggests that termatophylines may com-
lella, from Central America, and two new monly feed on moth larvae.
species of Termatophylidea. Miyamoto The termatophylines are found in a va-
(1965)gavedetailed descriptionsoftwo new riety of habitats, including flowers, foliage
termatophylines from Japan, providing ad- and moth larvalgalleries. Five termatophy-
equate genitalic and habitus illustrations. line species, Kundakimuka queenslandica,
Maldonado (1970) reviewed the distinctive Termatophyloides pallipes, T. pilosulus,
genus Termatophylidea, describing three Termatophylidea maculata and T. opaca,
newspecies, witha keytoallknown species. havebeen foundonleaves, with thelasttwo
Carvalho (1988) described a new genus, species restricted to the underside ofleaves
Termatophylina, from India, and gave il- (van Doesburg 1964, Callan 1975). Two
lustrationsofthemalegenitalia. Linnavuori species of Termatophylum, T hikosanum
(1974) described a new species of Terma- and T melaleucae, havebeencollected from
tophylum from Nigeria, and gave a key to flowers. Van Doesburg(1964) reported that
the African species. Poppius (1910, 1915), T opaca and its prey were found under fine
China (1929), Usinger (1935), Carvalho spider webs. The association of Termato-
(1983) and Ren (1983) described the other phylina Indiana with moth larval galleries,
species. suggests that termatophylines may be com-
monly encountered in sheltered microhab-
Biology itats. The prey ofKundakimuka queenslan-
The little that is known ofthe biology of dica, Xylorycta luteotactella, is also known
Termatophylini suggeststhatthespeciesare to live in small tunnels, which the moth
predators (Callan 1975). In Table 1 the bi- bores in the branches of their food plant
ological information that is known is listed. (Common 1990). Also ofinterest is the fre-
VOLUME 97, NUMBER 2 265
quency with which termatophylines are Termatophylinae: China and Myers 1929:
found at light. Five species of the genera 97 (subfamily ofMiridae).
Kundakimuka, Termatophylidea and Ter- Termatophylini: Carvalho 1951: 133 (tribe
matophylum have been taken at Hght or in ofDeraeocorini); Carvalho 1952: 32, 42,
Hght traps. This suggests that the termato- 43, 50 (checklist); Carvalho 1955a: 22
phyHnes are nocturnal predators. This hy- (generic key); Carvalho 1957: 34 (cata-
pothesisismorphologicallysupportedbythe logue).
presence of very large eyes, with enlarged
ommatidia. There is no evidence of Ter- Diagnosis.—TheTermatophyliniarebest
matophylina Indianabeingcollectedatlight, diagnosed by the short first labial segment
which isthe onlytermatophylinetaxon that (Figs. 19-20)whichdoesnotextendbeyond
does not have enlarged ommatidia, and the the posterior margin of the bucculae. The
eyes are ofmoderate size. head is moderately (Figs. 11, 14, 16, 17) to
Termatophylines are known from a wide strongly porrect (Figs. 12, 13, 15, 18), with
variety ofplants. They have been recorded the apex ofthe clypeus in lateral view ter-
from three monocot families, the Gramin- minatingaboveoratleveloftheantennifers
eae, Musaceae and Orchidaceae, and seven (Figs. 1 1-18). Unlike any other Deraeocor-
dicot families, the Anacardiaceae, Betula- inae the metathoracicspiracle isexternal (=
ceae, Bombacaceae, Compositae, Legumi- visible) as an oval or lanceolate opening,
nosae, Myrtaceae and Sterculiaceae (Table usually bounded by evaporative bodies on
1). The association oftermatophylines with the dorso-caudal arm ofthe dorsal margin
monocots is restricted to the Neotropical ofthe mesepimeron (Figs. 65, 67, 68). The
genera Termatophylella, Termatophylidea pronotum has antero-lateral projecting se-
and Termatophyloides. Thereisnoapparent tae on the anterior angles of the callosite
overall pattern of plant association in the region (Figs. 32-34).
termatophylines, with some species having Description.—Macropterous, either
multiple plant associations, and Termato- smooth or coriaceous; ovoid (Fig. 1), elon-
phylideaopacaknownfrombothamonocot gate-ovoid(Fig. 2), orparallel-sided; mostly
and dicot. Some sympatric termatophyline with rows ofpunctures on pronotum (Figs.
taxa are found on the same plant genera: 31, 33, 34, 38^2) and hemelytra (Figs. 47,
Arygrotelaneussimoniand Termatophylum 51-56); sparse to dense cover of setiform
insignearebothknownfromAcaciaspecies, vestiture, rarely intermixed with adpressed,
and Kundakimuka queenslandica and Ter- flattened scalelike setae. Head: moderately
matophylum melaleucae are known only (Figs. 11, 14, 16, 17) to strongly (Figs. 12,
from Melaleuca species. 13, 15, 18) porrect; elongate (Figs. 4, 5, 7,
10) or transverse (Figs. 3, 6, 8, 9); vertex
rounded, flattened, or narrowly bicom-
Systematics pressed, posterior margin sometimes weak-
ly carinate; frons moderately to strongly
Tribe Termatophylini
narrowed in front, barely to weakly decli-
Termatophylina: Reuter 1884b: 167; Reu- vent; clypeus short, weakly to moderately
ter 1884a: 218 (subfamily ofAnthocori- declivent, terminating above (Figs. 12, 13,
dae). 15) or at level ofantennifers (Figs. 11, 14,
Termatophylidae: Reuter 1910: 70 (family 16, 17, 18) in lateral view; bucculae usually
status); Poppius 1910: 56; Reuter and short and terminating caudally at or just
Poppius 1912:2(diagnosis); Reuter 1912: beyond antennifers, most often very nar-
52 (key; diagnosis); Parshley 1923: 665 row, with lateral margins either subparallel
(description); Wagner, 1970: *** (foot- (Figs. 20, 22, 23), weakly to moderately ar-
note). cuate (Figs. 19, 21, 26) sometimes strongly
PROCEEDINGSOFTHE ENTOMOLOGICALSOCIETY OF WASHINGTON
266
o o O I^
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VOLUME 97, NUMBER 2 267
Fig. 1. Democohs lugens, dorsal habitus offemale.
arcuate (Fig. 24), and in one case extremely dense matt ofwhitish, microsetae (Figs. 22,
broad, laterally almost contiguous with an- 23). Eyes: moderately sized (Figs. 12, 13,
tennifers, with margins also explanate (Fig. 15, \l\ 18) to very large (Figs. 11, 14, 16),
25); gula elongate, narrow (Figs. 22, 23) to most often contiguous, or nearly so, with
moderatelybroad(Figs. 19, 20, 21, 24),con- pronotal collar, rarely removed; ommatidia
cave (Figs. 19, 20, 22-26) to weakly convex usually very large, rarely small (Fig. 13),
(Fig. 21), often with transverse ridges, mar- with short to elongate setiform interocular
gins rarely carinate, sometimes with rows setae (Figs. 11-14). Antennae: inserted be-
of setae on lateral margins, rarely with a lowmidheightofeyes; usually short; mostly
