Hemiptera, Lygaeidae)

AMERICAN MUSEUM

Novta tes PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2754, pp. 1-23, figs. 1-24 November 12, 1982

Lilliputocorini, a New Tribe with Six New Species of Lilliputocoris, and a Cladistic Analysis of the Rhyparochrominae (Hemiptera, Lygaeidae)

JAMES A. SLATER' AND T. E. WOODWARD2

ABSTRACT The Lilliputocorini, a new tribe based on the ferred to Lilliputocoris, redescribed and reported genus Lilliputocoris Slater and Woodward, is from Queensland and Northern Territory (Aus- described and placed in the subfamily Rhyparo- tralia) and Dauan Island (Torres Straits). Dorsal chrominae. The relationships of the new tribe are view illustrations ofLilliputocoris neotropicalis, L. discussed and it is assigned to a clade also con- terraereginae, L. seychellensis, L. exiguus, and L. taining the Antillocorini and Lethaeini. A discus- punctatus are included as are illustrations of the sion of wing polymorphism and of zoogeography nymph of terraereginae, ofthe genitalia and abdo- is included. A key is given to the species of Lil- men, and scanning electron photographs of the liputocoris. Lilliputocoris ghanaensis (Ghana), L. abdominal trichobothria, scent gland scars and grossocerata (Borneo), L. neotropicalis (Brazil), L. spiracles. seychellensis (Seychelles Islands), L. taylori (Bor- A discussion ofthe cladistic relationships ofthe neo), and L. terraereginae (Australia) are described tribes of the Rhyparochrominae is included as new. Tomocoris punctatus Woodward is trans- together with a cladogram and a character analysis.

INTRODUCTION The genus Lilliputocoris was described by present paper. At the time of the original Slater and Woodward (1979) based on Lil- description of L. exiguus, we tentatively liputocoris exiguus from Sri Lanka (Ceylon). placed Lilliputocoris in the tribe Antillocorini At that time we noted that before us were because of its concave apical corial margin, additional species, from a number of widely elongate buccular carinae, lack of basal iri- separated localities. One of these had been descent areas on the dorsal head surface, ven- described as Tomocoris punctatus by Wood- tral spiracles, and minute size. None of these ward (1959), the others are described in the reasons are very compelling. The nature of

' Research Associate, Department of Entomology, American Museum of Natural History; Biological Sciences Group, University of Connecticut, Storrs, Connecticut 06268. 2 Professor of Entomology, Department of Entomology, University of Queensland, Brisbane, Australia.

Copyright ©3 American Museum of Natural History 1982 ISSN 0003-0082 / Price $1.75 2 AMERICAN MUSEUM NOVITATES NO. 2754 the concave apical corial margin in Lillipu- We appreciate the financial assistance from tocoris is quite different from that of the the following: a University of Queensland Antillocorini in that it encompasses most of Research Grant and an Australian Biological the margin (figs. 1 1, 12, 17), whereas in most Resources Study Grant to the junior author. antillocorine species the concavity occurs as A United States National Science Founda- a more or less abrupt sinuation near the inner tion Grant to the senior author. angle. Most other Rhyparochrominae, except the Lethaeini, lack iridescent basal head areas LILLIPUTOCORINI, NEW TRIBE (although some have this surface area mod- Polished, shining or subshining, minute ified) and a great many have elongate buc- insects less than 2 mm. in length. Body slightly cular carinae. flattened. Clothed above and on abdominal The availability ofconsiderable additional venter with elongate erect hairs. Pronotum, material and especially the opportunity to scutellum, and hemelytra coarsely punctate. study a fifth instar nymph of Lilliputocoris Head porrect, lacking basal iridescent areas, terraereginae, new species, now makes it pos- sometimes with scattered punctures but fre- sible to interpret the systematic position of quently smooth across vertex, latter strongly the genus in a more definitive manner. convex. Bucculae joined mesally well behind We wish to thank all the collectors named base of labium. Eyes separated from prono- in the text; Mr. E. L. Dahms, Curator of tum by less than one-half eye length. Inter- Insects and the authorities ofthe Queensland ocellar distance approximately three times Museum for permission to examine the holo- ocellar width. First antennal segment exceed- type of Tomocoris punctatus; Mr. T. A. Weir ing apex of tylus, often greatly so; segments (Australian National Collection) for search- two and three robust and clavate; segment ing "berlesates" and extracting specimens; Dr. four broadly fusiform; all segments with long C. W. Schaefer (University of Connecticut) erect and suberect hairs. Pronotum of for aid in the interpretation ofthe ovipositor; macropterous form broadened posteriorly Dr. G. Schmitz (Musee Royal de l'Afrique with rounded humeral angles; lateral margins Central, Tervuren, Belgium) for the use of sinuate, frequently with crenulate edges; anatomical sketches; Ms. Jane O'Donnell meson often depressed; calli shining, large, (University ofConnecticut) for dissection and impunctate or nearly so. Scutellum broader illustration of the male genitalia; Mr. G. V. than long, lacking a median elevation. Meso- Hardy (Electron Microscope Centre, Queens- sternum anteriorly usually with a short, low, land University) for the scanning electron median carina. Hemelytra with clavus broad; micrographs; Mrs. G. Yeo, Mr. G. Thomp- a distinct commissure present; lateral corial son, and Ms. Mary Jane Spring for the dorsal margins sinuate, often crenulate on proximal view illustrations; Mr. G. Schmidt, Mr. M. third; apical corial margins evenly and shal- S. Upton, and Curators (Australian National lowly concave for most oflength. Membrane Insect Collection); the late Dr. Dennis Leston ofhemelytron without cross veins. Hind wing (formerly University of Ghana); Dr. R. T. (when present) lacking veins in cubital sector Schuh (American Museum of Natural His- and anal lobe. Abdomen with well-developed tory) for the loan of material; Mrs. R. Crom- dorsal abdominal scent gland scars between bie, Mrs. P. Haas, and Mrs. Elizabeth Slater terga three and four and four and five; lacking for aid in the preparation of the final manu- a scent gland scar between terga five and six script. (fig. 6). Inner laterotergites absent.

FIGS. 1-8. Anatomical details of species of Lilliputocoris. 1. L. taylori, new species, genital capsule dorsal view. 2. L. taylori, new species, genital capsule lateral view. 3. L. taylori, new species, right clasper. 4. L. seychellensis, new species, head and thorax lateral view. 5. L. seychellensis, new species, spermatheca. 6. L. seychellensis, new species, abdomen dorsal view. 7. L. seychellensis, new species, abdomen lateral view. 8. L. seychellensis, new species, metatarsus. 1982 SLATER AND WOODWARD: LILLIPUTOCORIS 3

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All abdominal spiracles ventral (fig. 7). the males are macropterous and the females, Abdominal suture between sterna four and in part or always, micropterous or brachyp- five usually incomplete, occasionally com- terous. In the Lygaeidae when wing dimor- plete. Trichobothria of sternum five either phism occurs both sexes are equally involved. with one trichobothrium anterior to spiracle Second, as noted by Slater (1977-(1978)), it and two placed together behind it (fig. 23) is very unusual to find flightless Lygaeidae in (plesiomorphic condition) or lacking two lowland rainforests. posterior trichobothria on sternum five. The disjunct distribution, degree of mor- Suture between sterna three and four some- phological isolation and the nature ofthe wing times incomplete dorsolaterally and that reduction all seem congruent with the idea between five and six often nearly or com- that this is a very old group. pletely obliterated. Metathoracic scent gland auricle somewhat scimitar-shaped, curving strongly anteriorly (fig. 20); often elevated SYSTEMATIC POSITION along outer margin to form rim of evapora- Stys (1964) has suggested that the Lygae- tive area. Fore femora mutic. Tarsi two seg- idae probably constitute a paraphyletic group mented (fig. 8). Ovipositor reduced, scarcely and that the general lygaeoid-like habitus of laciniate, second valvifers fused across mid- members of the pentatomoid family Thau- line; no suture present between ninth tergite mastellidae is the retention of a plesio- and ninth paratergite; first valvifers lightly morphic body shape. sclerotized distally; first and second valvulae It is true that species of Lilliputocoris do membranous; tenth segment solidly sclero- resemble species of Thaumastella and also tized, tenth tergum closely applied to ninth species of the lygaeid genus Camptocera tergite, the two separated by a suture but not (Lethaeini). These three taxa are all com- by a membrane. Spermatheca without flanges; posed of small, yellowish brown insects with bulb ovoid; pump short; duct relatively elon- convex heads, subquadrate pronota and sub- gate (fig. 5). Sperm reservoir bulb apparently flattened bodies. The overall resemblance of ofuniform thickness; wings straplike curving Lilliputocoris to Thaumastella is enhanced ventrad around bulb; no vesical coils; large by both taxa containing species with a similar membranous lobes present laterally just staphylinoid morph and species with crenu- proximad of sperm reservoir (figs. 14, 15). late pronotal and hemelytral margins. Frequently flightless, in which case hem- Therefore it is important to consider the elytra staphylinoid or micropterous,3 com- possible relationships between Thaumastella pound eyes much reduced, placed laterally and Lilliputocoris. Females ofboth taxa have on head, ocelli lacking or vestigial and prono- a reduced ovipositor. In Thaumastella the tum quadrate with posterior lobe much ovipositor is platelike, whereas in Lilliputo- reduced. coris, despite the reduction, it is distinctly laciniate. TYPE GENUS: Lilliputocoris Slater and Wood- Apomorphic features found in Lilliputo- ward coris and not in Thaumastella are the incomplete abdominal sutures (fig. 7), two-seg- Although nothing is known of the biology mented tarsi (fig. 8), lack of flanges on the of these insects other than that some live in spermathecal bulb (fig. 5), and the uniquely rainforest ground litter, the wing dimorphism shaped, forwardly directed metapleural scent is distinctive in two ways. First, this is the gland auricle (fig. 20). The Thaumastellidae only case known to us in the Lygaeidae where have derived features in the five-segmented there is a sexual difference in wing develop- antennae, a unique hemelytral corium that ment. In some other families of Hemiptera, distally is separated into two parts with the such as the Miridae and Anthocoridae, it is membrane extending a short distance into not uncommon to observe species in which this distal corial area, fusion of the eighth ventral laterotergites, trichobothria arranged 3 Staphylinoid and micropterous used in the sense of in a two-plus-two pattern on sterna two Slater (1975, p. 53). through six and a one-plus-one pattern on 1 982 SLATER AND WOODWARD: LILLIPUTOCORIS 5 sternum seven. Stys (1964) and others have concluded that despite its lygaeid-like habitus Thaumastella should be placed as a distinct family in the superfamily Pentatomo- idea. The ovipositor does not relate Lilliputo- coris to any pentatomoid taxa. In the Cyd- nidae which is certainly a pentatomoid group (and one that has many "primitive" features) the fusion of the second valvifers is incomplete, the two structures being separated by a membrane (McDonald, 1966; Schaefer, 1968). In Lilliputocoris the second valvifers are fused. This is true ofsome Pentatomoidea but there the second valvifers are more reduced and lack a close association with the ninth paratergites. The fusion of the second valvifers in Lilliputocoris also rules out a direct relationship with either the Coreoidea or Pyrrhocoridae as in these groups no midline fusion occurs in any ofthe paired genital structures (Schaefer, 1964, 1965). The structure of the ovipositor thus appears to be an FIG. 9. Lilliputocoris terraereginae, new independent reduction phenomenon and does species, abdomen of fifth instar nymph. Dorsal not indicate unambiguously cladistic rela- view. tionship with the pentatomoid or coreoid complexes. dorsal abdominal scent gland between terga The incomplete, fused suture between five and six. We recognize that this is a reduc- abdominal sterna four and five is a strong tion phenomenon and we have not been able synapomorphy which places Lilliputocoris in to establish other synapomorphies. Never- the lygaeid subfamily Rhyparochrominae. theless the reduction ofthe posterior abdom- Sweet (in Slater, Sweet and Baranowski, inal scent gland is a constant feature that 1977) discusses an important synapomorphy occurs throughout the many and varied taxa that is possessed by members of the rhypa- of the Lethaeini. The Lethaeini is certainly a rochromine tribes Antillocorini and monophyletic assemblage as evidenced by the Lethaeini. This is the presence ofa double or unique sperm reservoir (Slater and O'Don- "troughed" suture in the nymphs between nell, 1978), lack of a Y-chromosome, linear abdominal terga three and four and four and trichobothria on abdominal sternum five and five. These troughs lead from the lateral edges iridescent areas at the base of the dorsal sur- ofthe dorsal abdominal scent gland openings face ofthe head. Unfortunately, none ofthese to the lateral margins of the abdomen where derived character states occurs in any species there is usually a field ofspines and tubercles of Lilliputocoris. present. The nymph of Lilliputocoris terra- Our original decision to place Lilliputo- ereginaepossessesjust such a pair oftroughed coris in the tribe Antillocorini, as noted above, sutures and a lateral field of spines (fig. 9). was a tentative one. Ashlock (1964) originally We believe that this establishes Lilliputocoris defined the Antillocorini as having as syn- as part of a monophyletic group with the apomorphies linear trichobothria on abdom- Antillocorini and Lethaeini. inal sternum five and a concave apical corial The position of Lilliputocoris within the margin. As more and more Antillocorini are above clade is a more difficult problem. We being discovered with the trichobothria of have, in the cladogram (fig. 10), placed Lil- abdominal segment five in the plesiomorphic liputocoris as the sister group ofthe Lethaeini position (and some species with straight api- on the basis of the reduction or loss of the cal corial margins) (Slater, 1980, and in prep.), 6 AMERICAN MUSEUM NOVITATES NO. 2754 it is becoming evident that the Antillocorini PARAPHYLY: Unlike many cladists we are may well be a paraphyletic group. While rec- not necessarily opposed to the use of para- ognition of such a taxon may or may not be phyletic taxa in a classification. We see con- warranted, depending on one's systematic siderable merit in the views of Gauld and philosophy, it is evident that Lilliputocoris is Mound (1982) on this subject. We do believe not closely related to any of the known taxa that if such groups are included that they be in the Antillocorini. The various species of clearly recognized as such so that they are not Lilliputocoris in fact form a closely related used in biogeographic analyses in the same monophyletic group held together by several way that monophyletic groups are. An exam- synapomorphies such as the absence ofa dor- ple of the value of an analysis of this kind sal abdominal scent gland between terga five and of the functional limitations of probable and six, the clavate second and third antennal paraphyletic groups is the status of the tribe segments, the reduced and modified ovipos- Stygnocorini. This taxon as currently under- itor, the two-segmented tarsi, the uniquely stood is an assemblage whose limits Slater formed metapleural scent gland auricle and and Sweet (1970) considerably restricted from adjacent evaporative area, and by the loss of a former polyphyletic assemblage recognized abdominal inner laterotergites. by Scudder (1957). As the cladogram (fig. 10) Inner laterotergites are present in the illustrates the Stygnocorini belong to a clade Lethaeini. Thus Lilliputocoris cannot be con- that contains most of the tribes of the Rhy- sidered a member ofthe Lethaeini. We there- parochrominae. However, genera at present fore conclude that the most appropriate taxo- placed in the Stygnocorini do not have a sin- nomic procedure is to treat the genus as a gle synapomorphy that segregates them from distinct tribe and tentatively as the sister group the other tribes. of the Lethaeini. We recognize that such Slater and Sweet (1970) discuss the Styg- action necessitates the evolution ofthe linear nocorini as a group of "primitive" rhyparo- arrangement of the trichobothria on abdom- chromines within the "Y-suture" assem- inal sternum five independently in the Antil- blage. The present cladistic analysis therefore locorini and in the Lethaeini. This does not is important in emphasizing that at present seem unreasonable as it certainly has arisen the distribution of the Stygnocorini cannot independently in the Targaremini (see dis- be considered of zoogeographic significance cussion below). as the taxon may be paraphyletic. That this may indeed be so is at least suggested by the SYSTEMATIC RELATIONSHIPS similarity in habitus of some Stygnocorini to WITHIN THE RHYPAROCHROMINAE some Ozophorini. Indeed the loss ofthe inner When this manuscript was originally sub- laterotergites alone would place some taxa in mitted Dr. Randall Schuh suggested that it the Ozophorini rather than the Stygnocorini. might not be inappropriate to include a pre- (Note that the cladogram suggests three inde- liminary cladogram and a discussion of the pendent losses of the inner laterotergites.) cladistic relationships of the tribes currently Other suggested cases of paraphyly are recognized in the subfamily Rhyparochro- revealed by the cladogram. The Antillocorini minae. To do so might further clarify the which were erected by Ashlock (1964) to reasons for our decision to recognize Lilli- include taxa without a nymphal Y-suture, but putocoris as a distinct tribe as well as place with linear trichobothria on abdominal ster- current views on the relationships ofthe tribes num five and a deep concavity on the inner into the form of a testable hypothesis. The portion of the apical corial margin, are now following discussion is a working hypothesis. known to contain a number of taxa with We wish it to be viewed as exactly that. Much straight apical corial margins and non-linear more evidence is needed to firmly establish trichobothria. There thus does not appear to (or refute) several ofthe suggested sister group be a synapomorphy for this tribe. relationships. The analysis does reveal where Sweet (1967) erected the Udeocorini for a major cladistic character strengths and weak- number ofchiefly Australian genera and sep- nesses lie and suggests which tribes may be arated it from the Myodochini by the pres- paraphyletic. ence ofinner laterotergites in the Udeocorini 1 982 SLATER AND WOODWARD: LILLIPUTOCORIS 7

z z C) 0 0 0 IaI z z I z Z 0 0It 0 z 0 0 U) 0 C-) z I H Z H w 0 z 0 a: a- 0 0cl a: z z 0 z 0I -i 0 0 I 2: L CD 7- ar- 0 z w z wD 0 I -J z -J w N Hr 0 w a<. 0 a- < Ji -J H 0- 0 C]

FIG. 10. Cladogram of tribes of Rhyparochrominae. and their absence in the Myodochini. Pres- to include a short discussion ofwhat we con- ence ofinner laterotergites is a plesiomorphic sider to be "strong" and "weak" characters condition in the Rhyparochrominae and thus in the cladogram. Once again we recognize the Udeocorini, like the Antillocorini and the that this is a matter of philosophical contro- Stygnocorini, do not at present possess a sin- versy, some cladists believing that the dis- gle definitive synapomorphic feature and tribution ofcharacters without consideration again may well represent a paraphyletic group. of their relative importance is necessary. We For a cladistic analysis of this type to be do not believe this is the place to enter such valuable for future workers it seems desirable a debate but we do believe that some char- 8 AMERICAN MUSEUM NOVITATES NO. 2754 acters are much more important than others near the base of the cladogram (elimination as the following discussion indicates. of character 2). The presence of a fused suture between Recognition of the troughed suture (char- abdominal terga four and five is the single acter 2) as a synapomorphy requires the rec- character that appears to be a strong synap- ognition ofthe secondary loss ofthe Y-suture omorphy holding the Rhyparochrominae in the ancestor ofthe Gonianotini and Mega- together as a monophyletic group. Two points lonotini while at the same time rejecting the should be made. The Plinthisini, which are secondary loss ofthe suture in the Cleradini. usually included in the Rhyparochrominae, That secondary loss of the Y-suture in the do not have this suture fused. Sweet (1967) Gonianotini and Megalonotini is probably retained the Plinthisini in the subfamily on correct is supported by the dorsalization of the basis of overall habitus, similar feeding some ofthe anterior abdominal spiracles and habits, and the possession of head tricho- the development of a strongly sclerotized bothria. The last feature is surely plesio- spermathecal pump, both conditions occur morphic. The general habitus and feeding in tribes with a Y-suture present (characters habits are suggestive but not useful cladisti- 5, 7, 19). cally. Putshkov (1958) has already suggested The position ofthe Cleradini on the clado- that the Plinthisini (which possess a number gram is arbitrary. We have placed them near of autapomorphies) might better be recog- the base ofthe cladogram to emphasize their nized as a subfamily. anomalous position and the large number of The fused and incomplete suture between autapomorphies present in the tribe. The abdominal sterna four and five is also pos- overall habitus of many Cleradini suggests a sessed by several species ofLargidae. Whether possible relationship with the Drymini. The this is evidence that they form part of a fact that the species of Cleradini whose feed- monophyletic assemblage with the Rhyparo- ing habits are known are blood suckers living chrominae or that this apomorphic condition in the nests of vertebrates, and that at least has developed independently is beyond the one drymine also lives in nests and is at least scope ofthis paper and must await the results in part predaceous (Slater and Carayon, 1963), ofa phylogenetic analysis ofthe Largidae and and that several drymines are arboreal (Gas- Pyrrhocoridae currently underway by Stehlik trodes, Apollonius) suggests the possibility that and Stys. the cleradines may have once had a common We have placed the taxa with "troughed ancestor with the Drymini. However, we have sutures" on the nymphal tergum together by not found a synapomorphy for the two groups. considering this a true synapomorphy. (See The cleradine complex is currently under character 2.) This is a parsimonious hypoth- study by both Malapatil and Harrington and esis but there are reasonable alternative pos- we should have a more sophisticated analysis sibilities. The hypothesis assumes that the available within a few years. troughed suture between terga three and four We have previously discussed the clade developed first and subsequently two mono- containing the Antillocorini, Lilliputocorini, phyletic assemblages "diverged." The first and Lethaeini and here only wish to reem- retained the troughed suture between terga phasize that the double-troughed sutures three and four and developed a similar (character 4) appear to be a very strong syn- troughed suture between terga four and five. apomorphic feature that establishes a mono- This is represented by the present clade which phyletic group. The presumed synapomor- contains the Antillocorini, Lilliputocorini, and phy (character 10) that unites the Lethaeini. The hypothesis also assumes that Lilliputocorini and Lethaeini is a loss feature another ancestral type elaborated the troughed and therefore the evidence for monophyly is suture between terga three and four into the much less compelling. Y-sutured condition. It is of course possible The polychotomy that includes the Styg- that these two events occurred independently nocorini, Phasmosomini, Ozophorini, Tar- in which case the monophyly of the two garemini, and Drymini will, we believe, groups would not be destroyed but it would require a great deal of work before it can be necessitate the recognition of a trichotomy resolved. We have noted above the possible 1 982 SLATER AND WOODWARD: LILLIPUTOCORIS 9 paraphyletic nature of the Stygnocorini. The acter 16) can at least equally well be consid- presumed synapomorphies used in the clado- ered to be two independent developments. gram to group the Phasmosomini with the Our decision to accept it as a working hypoth- Ozophorini and the Targaremini with the esis is partly because many species ofthe two Drymini are not compelling. In particular the tribes have a similar habitus and the bio- forward movement ofthe trichobothria used geography is intriguing although beyond the to group the Targaremini and Drymini (char- scope of this paper to discuss.

LIST OF APOMORPHIC CONDITIONS USED IN THE ANALYSIS OF CLADISTIC RELATIONSHIPS IN THE RHYPAROCHROMINAE 1. Fusion of sutures between abdominal in the Rhyparochrominae. We have accepted sterna four and five. this conclusion as it seems to be consistent 2. Development ofa scent conducting trough with other evidence. between abdominal terga three and four in the nymphs. 6. Secondary loss of the "Y-suture." 3. Elaboration and bifurcation of the above 7. Movement ofspiracle three from a ventral trough laterally between abdominal terga to a dorsal position. three and four into a "Y-suture." 8. Movement of spiracle two from a ventral 4. Development of a trough in the nymphs to a dorsal position. between abdominal terga four and five. 9. Loss ofthe abdominal inner laterotergites. 5. Movement of abdominal spiracle four Sweet (1967) has used the loss of inner from a ventral to a dorsal position on the laterotergites as an important feature to abdomen. establish the tribes Ozophorini and Udeo- Dorsally placed spiracles on anterior corini. We have used it but have considerable abdominal segments is considered to be a reservations about its value in phylogenetic derived condition. The polarity appears to analysis. It is after all a loss character. Also have been from a dorsal spiracle only on seg- in taxa where it is present its degree ofdevel- ment four, to dorsal spiracles on segments opment is extremely diverse ranging from three and four and finally to dorsal spiracles wide very heavily sclerotized plates to slender on segments two, three, and four. Both the lightly sclerotized strips that in some cases Megalonotini and Rhyparochromini have can scarcely be seen unless the abdomen is dorsal spiracles on segments three and four. cleared and stained. Therefore, the Ozo- Either the dorsalization of spiracle three has phorini and Myodochini which are separated developed independently twice or the Y-su- from the Stygnocorini and Udeocorini chiefly ture has been lost twice. The cladogram (fig. by this feature will certainly benefit from a 10) indicates an independent development of further analysis of relationships. a dorsal spiracle on segment three. The heavy 10. Reduction and loss ofthe dorsal abdom- sclerotization of most nymphs of Goniano- inal scent gland between terga five and tini and Megalonotini (as well as a similar six. distribution) suggests a close relationship 11. Loss ofthe dorsal abdominal scent gland between these two tribes. between abdominal terga three and four. Ventral spiracles are the plesiomorphic 12. Loss of secondary veins in the cubital condition in the Heteroptera. However, this sector of the metathoracic wing. is not necessarily true ofthe Lygaeidae. Sweet 13. Loss of the cubital furrow in the meta- (1981) has introduced the interesting possi- thoracic wing. bility that the abdominal sclerotization ofthe 14. Fusion of veins R and M in the meta- Lygaeidae is close to the plesiomorphic con- thoracic wing for a considerable distance dition but with the connexivum shifted dor- distad of their juncture. sally. This would result in primitive dorsal 15. Linear placement of trichobothria on spiracles in the family. Sweet himself, how- abdominal sternum five. ever, believes that dorsalization is secondary 16. Forward movement ofthe posterior pair to AMERICAN MUSEUM NOVITATES NO. 2754

of trichobothria on abdominal sternum 30. Modification of the scent gland auricle five. and evaporative area. This character state is tentatively consid- 31. Reduction of the ovipositor. ered to establish a synapomorphy for the 32. Concavity of the apical corial margin. Drymini and Targaremini. Since the tricho- 33. Loss of a tarsal segment. bothria are linear in the latter and the two 34. Development of a secondary longitudi- posterior trichobothria are more or less dor- nal abdominal suture. soventrally situated in the former this may not be a true relationship. The habitus is often LILLIPUTOCORIS SLATER AND WOODWARD very similar in the two taxa. This is a rela- Lilliputocoris Slater and Woodward, 1979, p. 16. tionship worthy of a detailed analysis. TYPE SPECIES: Lilliputocoris exiguus Slater 17. Development of "pores" anteriorly on and Woodward. Monobasic. the abdominal sternum. DIAGNOSIS: Same as for tribe. 18. Anterior movement of both posterior trichobothria of sternum five. KEY TO SPECIES OF LILLIPUTOCORIS 19. Development ofa heavily sclerotized and 1. Length offirst antennal segment considerably transversely ridged spermathecal pump. greater than width ofhead across eyes (Bor- Scudder (1957) used this character state as neo) (macropters only known) ...... the most important synapomorphy for an ...... grossocerata,new species inclusive tribe which included all of the la. Length of first antennal segment less than rhyparochromine taxa with dorsal spiracles. (rarely nearly subequal to) width of head across eyes ...... 2 Sweet (1967) notes that transverse ridging can 2. Macropterous ...... 3 be seen to be weakly developed in other taxa 2a. Micropterous or staphylinoid ...... 7 when viewed under a high power of a light 3. First antennal segment relatively elongate, microscope. Nevertheless, the condition in much more than three-fourths as long as the taxa with dorsal spiracles is in our opinion width of head across eyes ...... 4 a strong indication of a close cladistic rela- 3a. First antennal segment relatively short, at most tionship. only slightly more than two-thirds and always much less than three-fourths as long 20. Development of a hardened, heavily as width of head across eyes ...... 5 sclerotized nymphal abdomen. 4. Basal one-halfoflateral corial margins bearing 21. Development of "corrugated" head a series of conspicuous crenulate teeth (fig. grooves. 17); antennae nearly unicolorous, chiefly yellowish brown (Australia) ......

Sweet (1967) uses the presence of "corru- ...... terraereginae,new species gated head grooves" as a synapomorphy use- 4a. Basal one-halfoflateral corial margins smooth ful for recognition of the tribe Ozophorini. or nearly so, lacking a series ofconspicuous However, very similar grooves are present in crenulate teeth; antennae chiefly dark choc- some Stygnocorini (Slater and Sweet, 1970). olate brown; distal ends of antennal seg- What appear to be at least analogous head ments two and three contrastingly pale (Ghana) ...... ghanaensis, new species surfaces are present in some Myodochini as 5. Punctures of pronotal lobe fine and incon- well. spicuous, not remotely approaching diam- 22. Development of a pseudoperculum on eter of claval punctures (fig. 12) (Brazil) . . the egg...... neotropicalis,new species a 5a. Punctures ofposterior pronotal lobe relatively 23. Development of bifurcate paramere. coarse and conspicuous, nearly of same 24. Utilization of vertebrate blood. diameter as those of clavus ...... 6 25. Lateral placement of ocelli. 6. Head and pronotal calli smooth and polished, 26. An expanded connexival membrane. former with only a few scattered punctures 27. Loss of the Y-chromosome. present (fig. 11) (Sri Lanka) ......

28. Extreme modification of the sperm res- ...... exiguusSlater and Woodward ervoir. 6a. Head finely but thickly punctate over entire 29. Development of"iridescent" head areas. surface; calli granulose or minutely punctate 1 982 SLATER AND WOODWARD: LILLIPUTOCORIS I1I

(Papua New Guinea; Australia) ...... by the lack of crenulate teeth on the basal ...... punctatus (Woodward) portions of the lateral corial margins. 7. Micropterous, hemelytral wing pads not DESCRIPTION: General color testaceous. extending over abdominal tergum three, Scutellum and a macula at apex of corium narrowly in contact at midline (Borneo) dark red-brown to chocolate brown. Pleural ...... taylori, new species 7a. Staphylinoid, hemelytra covering third and ventral surfaces red-brown. Abdomen abdominal tergum, broadly in contact at becoming pale yellow laterally. Antennal seg- midline (fig. 18) ...... 8 ment one ochraceous; proximal two-thirds of 8. First antennal segment short not exceeding antennal segments two and three dark choc- tylus by as much as one-half its length (fig. olate brown contrasting strikingly with 13) (Papua New Guinea; Australia) ...... ochraceous coloration of distal one-third of ...... punctatus (Woodward) each segment; fourth antennal segment tes- 8a. First antennal segment relatively elongate taceous, infuscated with dark brown proxi- exceeding tylus by much more than one- mally. Head, pronotum, and scutellum thickly half its length ...... 9 9. Suture between abdominal terga four-five and and coarsely punctate. Dorsal surface clothed five-six smooth, lacking a series of ridges with numerous, conspicuous, elongate, sil- and furrows; lateral margins of hemelytra very, upstanding hairs. nearly smooth, lacking well-developed Head only slightly declivent anteriorly; crenulate teeth (Seychelles) ...... vertex strongly convex. Tylus extending only ...... seychellensis, new species to proximal one-third of first antennal seg- 9a. Suture between abdominal terga four-five ment. Ocelli very prominent, located as close bearing a series of ridges and furrows (fig. to midline as to compound eyes. Length head 24); lateral margins of hemelytra with 0.26,4 width 0.34, interocular space 0.27. prominent outwardly directed teeth (fig. 18) Pronotum with lateral margins deeply sin- (Australia) .... terraereginae, new species uate, weakly crenulate; meson broadly and deeply impressed, this impressed area nar- Lilliputocoris exiguus Slater and Woodward rowing posteriorly; transverse impression Figure 11 incomplete; anterior collar area well delim- Lilliputocoris exiguus Slater and Woodward, 1979, ited; calli widely separated, nearly impunc- pp. 17-18. tate. Length pronotum 0.28, width 0.56. Scu- DIAGNOSIS: Recognized by the smooth pol- tellum somewhat elevated mesally but lacking ished head and pronotal calli and the rela- a distinct median carina. Length scutellum tively short first antennal segment which is 0.14, width 0.36. Hemelytra with clavus very only two-thirds as long as the width of the broad; three more or less anastomosing rows head across the eyes. of coarse punctures present, claval commis- DIscussIoN: This is a very small (1.62 mm.) sure noticeably longer than scutellum. Length nearly uniformly bright brown species. The claval commissure 0.20. Corium with lateral hemelytra and legs are pale yellow. The lat- margins deeply concave or sinuate lacking eral margins of the pronotum and hemelytra conspicuous crenulations; midline distance lack conspicuous crenulations. apex clavus-apex corium 0.34, midline dis- Lilliputocoris exiguus was described from tance apex corium-apex membrane 0.36. a single female from Sri Lanka and remains Labium reaching metacoxae. Length labial known only from the holotype. segments I 0.20, II 0.20,111, and IV obscured. ETYMOLOGY: The name refers to the small Antennae stout, segments two and three size of the insect. moderately clavate, segment four broadly fusiform. Length antennal segments I 0.29, Lilliputocoris ghanaensis, new species II 0.30, III 0.20, IV 0.30. Total body length 1.62. DIAGNOSIS: Recognized by its elongate, dark ETYMOLOGY: Named for Ghana, West chocolate brown first antennal segments, the bicolored second and third antennal segments which have the proximal two-thirds 4In the following descriptions all measurements are dark and the distal one-third ochraceous and in millimeters. 12 AMERICAN MUSEUM NOVITATES NO. 2754

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Africa, the country from which the holotype distance apex corium-apex membrane 0.38. was collected. Abdominal tergum with scent gland openings HOLOTYPE: Female, GHANA: Tafo present on sutures between terga three and November 6, 1965, Dennis Leston collector; four and four and five, a groove extending deposited in the American Museum of Nat- along suture part way to lateral margin from ural History. edge of each gland opening. Labium extending almost to metacoxae. Length labial seg- Lilliputocoris grossocerata, new species ments I 0.16, II 0.18, III 0.14, IV 0.19. DIAGNOSIS: Recognized by the extremely Antennae very conspicuous, all segments elongate, uniformly dark chocolate brown clothed with elongate upstanding hairs, seg- antennae that contrast strongly with the honey ments two, three, and four fusiform. Length yellow coloration ofthe remainder ofthe body antennal segments I 0.36, II 0.31, III 0.20, and by the crenulate lateral pronotal margins. IV 0.28. Total body length 1.50. DESCRIPTION: General coloration of body ETYMOLOGY: Named for the very large including legs bright yellow. Membrane of elongate conspicuous "horns" or antennae. forewing bicolored, a dark fumose irregular HOLOTYPE: Female, BORNEO: Sarawak, band running transversely across membrane Baco National Park near Kuching, SL (=soil from distal ends ofapical corial margins, this litter) rainforest July 1, 1968, berlesate, dark band very much broadened mesally; a accession number 68.767 (Robert W. Taylor second fumose area present on distal end of collector); deposited in The Australian membrane and extending anteriorly to mid- National Insect Collection number 9446. dle of membrane, remainder of membrane PARATYPE: One female, BORNEO: Sara- translucent. Distal portion of apical corial wak, East Malaysia, Semongok Forest margin dark brown. All femora with dark Reserve January 3, 1978, from litter in pri- brown spots. Entire antennae a very strongly mary rainforest (Thomas E. Woodward col- contrasting dark chocolate brown. Head lector); deposited in The University of nearly smooth. Pronotum, scutellum, clavus, Queensland Insect Collection. and corium coarsely punctate. Dorsal surface appearing nearly glabrous but with a few scat- Lilliputocoris neotropicalis, new species tered semideclivent hairs. Figure 12 Head only slightly declivent anteriorly. DIAGNOSIS: Recognized by its minute Tylus extending to basal one-fourth of first inconspicuous pronotal punctures and short antennal segment. Eyes well removed from first antennal segment. anterior margin ofpronotum by a short thick DESCRIPTION: General coloration pale tes- straight sided "neck." Ocelli present slightly taceous. Distal one-half of apical corial mar- behind eyes, the latter small and extending gin narrowly dark brown. Membrane marked only slightly above bases of antenniferous with pale brown very much as in terraere- tubercles. Length head 0.32, width 0.32, ginae. First antennal segment pale basally, interocular space 0.23. Pronotum with a dis- becoming fuscous on distal two-thirds. tinct anterior collar; lateral margins ofprono- Antennal segments two and three chiefly dark tum deeply sinuate with posterior lobe much chocolate brown with strongly contrasting expanded and conspicuously crenulate; pale distal ends. Head shining and polished, transverse impression complete; posterior almost impunctate across convex vertex. margin slightly convex between swollen Pronotal punctures large and coarse, ante- humeral angles. Length pronotum 0.27, width riorly delimiting a "collar-like" area; coarse pronotum 0.50. Scutellum lacking a median punctures also present transversely along the elevation. Length scutellum 0. 16, width 0.20. incomplete transverse impression and on Lateral corial margins sinuate, narrowest posterior pronotal lobe immediately behind at level of middle of claval commissure. it, remainder of posterior lobe nearly Length claval commissure 0. 16. Midline dis- impunctate, at most with small, shallow tance apex clavus-apex corium 0.30, midline widely separate inconspicuous punctures. 14 AMERICAN MUSEUM NOVITATES NO. 2754

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Scutellum with large punctures but these antennal segment. Length head 0.26, width widely separated from one another, never 0.32, interocular space 0.19. Lateral margins giving surface an anastomosing appearance. of anterior pronotal lobe very feebly crenu- Dorsal surface clothed with numerous elon- late. Length pronotum 0.26, width 0.53. Scu- gate upstanding pale hairs. tellum nearly smooth, not noticeably ele- General body shape including head con- vated mesally. Length scutellum 0.18, width vexity, medially depressed and laterally sin- 0.32. Clavus with punctures very much uate pronotum and noncarinate scutellum reduced, these conspicuous only along mar- very similar to ghanaensis. Tylus elongate gins of scutellum and along claval suture, extending well beyond basal one-third offirst greater part of claval surface essentially 1 982 SLATER AND WOODWARD: LILLIPUTOCORIS 15 impunctate. Length claval commissure 0.18. Central area of corium hyaline, lateral margins only shallowly sinuate, lacking crenulations; midline distance apex clavus-apex corium 0.34, midline distance apex corium-apex membrane 0.30. Labium partially obscured, probably reaching metacoxae. Length labial segments I 0.25,11 0.14, III and IV obscured. Antennae relatively slender. Length antennal segments I 0.20, II 0.20, III 0.14, IV-missing. Total body length 1.38. ETYMOLOGY: Named for the Neotropical faunal region, from which the holotype was collected. HOLOTYPE: Female, BRAZIL: Para, Jaca- reacanga June 1970, F. R. Barbosa collector; deposited in the American Museum of Nat- ural History. (Note: this locality is at 6°09'S, 5801 5'W.) Lilliputocoris punctatus (Woodward) Figures 13-15 Tomocoris punctatus Woodward, 1959, pp. 51- 53. Tomocoris punctatus Woodward, 1963, p. 220. FIG. 13. Lilliputocoris punctatus (Woodward), DIAGNOSIS: Recognized by the thickly dorsal view. punctate dorsal surface and by the short first antennal segment which is never more than three-fourths the width ofthe head across the "male" should read "female"; (2.) p. 52, line eyes. 5 "0.33" should read "0.20." DISCUSSION: This species was originally MEASUREMENTS: Macropterous form- described from a single staphylinoid speci- Length head male 0.25, female 0.22; width men from near Port Moresby, Papua New head male 0.30, female 0.31; interocular space Guinea. Macropterous forms now available male 0.18, female 0.20. Length pronotum for study differ from the sympatric terraere- male and female 0.24, width male 0.48, ginae by the much shorter first antennal seg- female 0.50. Length scutellum male 0.14, ment, the almost completely dark color of female 0.16; width male 0.23, female 0.26. the antennae, the lack ofa median brown area Length claval commissure male 0.16, female on the hemelytral membrane, a less promi- 0.19. Midline distance apex clavus-apex co- nently mesally depressed pronotum, and the rium male 0.35, female 0.30. Midline dis- finer pronotal, scutellar and hemelytral punc- tance apex corium-apex membrane male and tures. Staphylinoids differ as noted in the pre- female 0.30. Length labial segments I male ceding key and by the finer dorsal punctures. and female 0.18; II male 0.17, female 0.15; The crenulations along the lateral margins of III male 0.10, female 0.08; IV male 0.22, the hemelytra are usually less prominently female 0.20. Length antennal segments I male developed than they are in terraereginae. 0.23, female 0.20; II male 0.23, female 0.20; The 34 specimens available for study show III male 0.17, female 0.15; IV male and female striking sexual differences in wing develop- 0.25. Width across hemelytra male 0.51, ment. All 15 males are macropterous, 17 of female 0.50. Total body length male and 19 females are staphylinoid. female 1.43. The following corrections should be made Staphylinoid form (measurements indi- to the original description: (1.) pp. 51-52 cate range): Length head 0.26-0.28, width 16 AMERICAN MUSEUM NOVITATES NO. 2754

(I. Yeo). Torres Strait: one female Dauan [Cornwallis] Island 6.V. 1953 (leaf mold near creek) (E. N. Marks). Northern Territory: one male, four females West Alligator Mouth 22.VII. 1979 (WAI) (Queensland Museum berlesate number 113. 12°12'S, 132°13'E, rainforest, sieved litter) (G. B. Monteith); one male, one female as above except 21.VII. 1979 l (WA2) (berlesate number 112). One female East Alligator Crossing 18.VII. 1979 (Queensland Museum berlesate number 107.

,- 1 2025'S, 1 32058'E, rainforest, sieved litter) (G. B. Monteith). Three males, one female Gorge northeast of Mount Gibruth 12.VII. 1979 (Queensland Museum berlesate number 101. 13°02'S, 133°05'E, rainforest, sieved litter) (G. B. Monteith); six males, six females (macrop-

1% terous) same except 10.VII.1979 (berlesate number 100). Two males Radon Creek, I Mount Brockman 14.VII.1979 (Queensland I Museum berlesate number 104. 12045'S, I 132053'E, rainforest, dry site sieved litter) (G. 14 15 B. Monteith); one male same data except 15.VII. 1979 (Queensland Museum berlesate FIGS. 14, 15. Lilliputocoris punctatus (Wood- number 105, stick brushing) deposited in ward). 14. Sperm reservoir dorsal view. 15. Sperm reservoir lateral view. Queensland Museum, University ofQueens- land Insect Collection, Australian National Insect Collection and J. A. Slater Collections.

0.29-0.30, interocular space 0.22-0.23. Lilliputocoris new Length pronotum 0.20, width 0.37-0.39. seychellensis, species Length scutellum 0.13-0.16, width 0.22-0.26. Figures 4-8, 16 Length forewing (to mesal margin) 0.13-0.16. DIAGNOSIS: Recognized by its strongly Length labial segments I 0.17-0.19, II punctate staphylinoid hemelytra, lack of 0.17-0.19, III 0.10, IV 0.22. Length antennal ridges and furrows on sutures between segments I 0.18-0.21, II 0.19-0.20, III abdominal terga four and five and five and 0.13-0.14, IV 0.17-0.20. Width across abdo- six and lack of the posterior pair of tricho- men 0.49-0.57. Total body length 1.30-1.34. bothria on sternum five. MATERIAL EXAMINED: Holotype female DESCRIPTION: General coloration reddish from PAPUA, NEW GUINEA. Brown River brown. Legs and distal one-halfto two-thirds near Port Moresby August 4, 1956, from leaf of antennal segments two and three testa- litter in rainforest near a river bank, Thomas ceous. E. Woodward collector. AUSTRALIA: Shape, punctation, pubescence, and Queensland: one female Iron Range, Cape structure very similar to staphylinoids of L. York Peninsula 1 1-17.V.1968 (from leaf lit- terraereginae. Lateral margins of pronotum ter) (G. Monteith): one male as above except at most very feebly crenulate. Lateral margins 26-31 .V. 1971. Three females (one macrop- of hemelytra lacking spinelike crenulations, terous) Pirate Beach, 30 miles (about 48 km.) appearing smooth. Tylus extending to prox- north of Cairns 23.VII. 1968 (Australian imal one-third of first antennal segment. National Insect Collection berlesate number Sutures between abdominal terga three and 98, Melaleuca litter in grass) (L. A. Mound). four and four and five fine and narrow, lack- One female Lotus Creek, Mackay district ing a series of ridges and grooves. Length 2.VI. 1956 (from leaf litter, brigalow scrub) head 0.29, width 0.32, interocular space 0.26. 1982 SLATER AND WOODWARD: LILLIPUTOCORIS 17 Z.

FIG. 16. Lilliputocoris seychellensis, new species, dorsal view.

Length pronotum 0.24, width 0.43. Length wing pad 0.38, width wing pad 0.52. Maxi- scutellum 0. 17, width 0.27. Maximum length mum width abdomen 0.56. Length labial seg- 18 AMERICAN MUSEUM NOVITATES NO. 2754 ments I 0.20, II 0.20, III 0.10, IV obscured. tum at level of calli. Length pronotum 0.22, Length antennal segments I 0.28, II 0.22, III maximum width 0.37. Scutellum depressed 0.16, IV missing (Paratype 0.24). Total body posteriorly lacking any indication ofa median length 1.46. elevation. Length scutellum 0.14, width 0.31. ETYMOLOGY: Named for the Seychelle Micropterous hemelytra reduced to short Islands from which the species was collected. broad posteriorly truncate pads that barely HOLOTYPE: Female, SEYCHELLES: Pras- reach base ofabdomen and narrowly meet at lin: Vallee de Mai. Foret de Palmacees July midline; posterior margin slightly angled 22-23, 1972, Mission Zoologique Belge Aux anteriorly from caudolateral angles; lateral Sechelles Benoit and J. J. van Moi collectors; margins strongly convex and strongly cren- deposited in the Musee Royal de l'Afrique ulate. Width across wing pads 0.50, length Central, Tervuren, Belgium. wing pad 0.20. Abdomen elliptical, dorsal PARATYPE: One female: same data as holo- surface convex, anterior margin of tergum type deposited in James A. Slater Collection. three thickened and tilted upward as a sharp rim. Maximum width across abdomen 0.62. Dorsal abdominal scent gland scars present Lilliputocoris taylori, new species between terga three and four and four and Figures 1-3 five, but intersegmental sutures curving cau- DIAGNOSIS: Recognized by the minute wing domesad so that gland scars are placed behind pads that are only narrowly in contact mes- suture lines. Labium exceeding metacoxae. ally and do not extend posteriorly over ter- Length labial segments I 0.22, II 0.22, III gum three, by the scent scars on the abdom- 0.12, IV 0.24. Length antennal segments I inal dorsum appearing to be placed 0.30, II 0.22, III 0.16, IV 0.22. Total body intersegmentally and the lack ofthe posterior length 1.54. pair of trichobothria on sternum five. ETYMOLOGY: Named for Dr. R. W. Taylor DESCRIPTION: General coloration bright of the Australian National Insect Collection reddish brown becoming almost orange-yel- who collected a large part of the type series. low on head and femora. Antennae chocolate HOLOTYPE: Female, BORNEO: Sarawak, brown on proximal two-thirds ofsegment two Semengok Forest Reserve. 11 miles south- and proximal one-half of segment three; west of Kuching, June 30, 1963, rainforest, fourth segment pale yellow, slightly darkened berlesate accession number 68.778, Robert at proximal end. Conspicuous upstanding W. Taylor collector; deposited in the Austra- hairs present on dorsal surface. Head lian National Insect Collection number 9445. impunctate across vertex, weakly punctate PARATYPES: One female same data as holo- anteriorly, a single row of coarse punctures type; one female same except 2-3.VII. 1968 present at anterior end where "neck" inserts (berlesate number 68.784); one female same into prothorax. Pronotum coarsely punctate as above except (berlesate number 68.781); anteriorly and behind calli, latter and pos- one female same except 28-31 .V. 1968 (leaf terior portion ofpronotum impunctate. Scu- mold, berlesate number 68.197 rainforest); tellum and wing pads coarsely punctate. two females same as above except (berlesate Head moderately declivent anteriorly, tylus number 68.201); one female same as above reaching only basal one-third offirst antennal except (berlesate number 68.196); one female segment. Compound eyes very much reduced, same except 1-4.VI. 1968 (berlesate number only a few ommatidia present. Ocelli absent. 68.259); three females same except (berlesate Vertex strongly convex. Length head 0.26, number 68.261). Two females Baco National width 0.32, interocular space 0.28. Pronotum Park near Kuching SL [=soil litter] rainfor- quadrate, anterolateral margins evenly est 1.VII.1968 (berlesate number 68.757). One rounded, lateral margins parallel-sided, female Borneo: Sabah, mile 45 Labok Road coarsely crenulate. Calli very large, convex, from Sandakan (Lungmanis) 12-13.Vl. 1968 and impunctate. Pronotal surface slightly (rainforest, berlesate number 68.502); depos- impressed mesally behind calli, posterior lobe ited in the Australian National Insect Col- greatly reduced, maximum width of prono- lection, Queensland Museum, American 1982 SLATER AND WOODWARD: LILLIPUTOCORIS 19

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FIG. 17. Lilliputocoris terraereginae, new species, dorsal view, macropter. 20 AMERICAN MUSEUM NOVITATES NO. 2754

one across base, one median and broadening to apex. Vertex finely punctate with a row of coarser punctures behind ocelli and a transverse impunctate area between each eye and ocellus. Dorsal surface with elongate upstanding hairs. Shape ofhead, pronotum and hemelytra as in ghanaensis. Lateral corial margins with distinct prominent crenulate-like teeth. Labium reaching metacoxae. Length head 0.34, width 0.36, interocular space 0.24. Length pronotum 0.30, width 0.61. Length scutellum 0.17, width 0.25. Length claval commissure 0.23. Midline distance apex clavus-apex corium 0.43; midline distance apex corium-apex membrane 0.40. Length labial segments I 0.23, II 0.22, III 0.14, IV 0.28. Length antennal segments I 0.35, II 0.34, III 0.23, IV 0.27. Total body length 1.80. Staphylinoid females differ from macropterous males as follows: yellowish brown; antennal segment four and tylus usually paler brownish yellow; abdomen shining brown or yellowish brown; ventral surface ofhead, thorax, and base of abdomen yellowish brown to dark brown. Length head 0.30-0.35; width across eyes 0.35-0.37; interocular space 0.28-0.32 (ca. 8-lOX width of eye); ocelli vestigial. Length antennal segments I 0.28-0.31, II 0.24-0.28, III 0.17-0.19, IV FIG. 18. Lilliputocoris terraereginae, new 0.24-0.25. Length labial segments I species, dorsal view, staphylinoid. 0.23-0.26, II 0.23-0.25, III 0.11-0.14, IV 0.25-0.28. Length pronotum 0.25-0.28; posterior width 0.45-0.51. Length scutellum Museum of Natural History, University of 0.15-0.18; width 0.28-0.34. Forewings leav- Queensland, Agricultural Research Institute ing abdomen exposed from tergum IV to apex; Semongok Sarawak, and James A. Slater Col- entire surface of hemelytra with large, deep, lection. mostly contiguous punctures, interspaces raised, giving surface a coarsely granulate Lilliputocoris terraereginae, new species appearance; apical margins broadly and shal- Figures 9, 17-24 lowly concave, posterolateral angles acutely DIAGNOSIS: Recognized by the relatively or subacutely produced posteriorly; length long first antennal segment, the nearly pale mesal margin 0.18-0.20. Total body length yellowish brown color of the antennae and 1.61-1.7 1; width across abdomen 0.67-0.74. the conspicuous crenulate teeth on the lateral ETYMOLOGY: Named for Queensland, the margins of the pronotum and basal half of Australian State from which the species was the corium. collected. DESCRIPTION: Coloration nearly uniformly HOLOTYPE: Male (macropterous), AUS- pale yellowish-testaceous. Base of scutellum, TRALIA: Queensland 730 m. Mount Tip- apical corial macula and pleural and ventral tree, June 29, 1971, Taylor and Feehan col- areas of thorax brown. Membrane of fore- lectors, berlesate Australian National Insect wing with extensive brown to fuscous areas, Collection 345, rainforest, 17°03'S, 145°37'E; 1 982 SLATER AND WOODWARD: LILLIPUTOCORIS 21

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FIGS. 19-24. Lilliputocoris terraereginae, new species, scanning electron micrographs. 19. Abdominal terga three through six. 20. Metathoracic scent gland auricle and evaporative area. 21. Ventral view of head with labium and antennae removed. 22. Detail ofanterior trichobothrial area ofabdominal sternum five showing surrounding pectinate processes. 23. Spiracles and trichobothria of abdominal sterna five and six. 24. Mesal portion of suture between abdominal terga four and five. 22 AMERICAN MUSEUM NOVITATES NO. 2754 deposited in the Australian National Insect shallowly concave and posterolateral angles Collection number 9444. acutely or subacutely produced backward. PARATYPES: Queensland, two macropter- Whereas all specimens of L. terraereginae ous males, nine staphylinoid females same were collected at altitudes of over 600 m. in data as for holotype; two macropterous males northeast Queensland, punctatus was col- (one dissected), seven staphylinoid females, lected only at or near sea level in coastal or data as above except berlesate Australian subcoastal areas in Papua New Guinea and National Insect Collection (=ANIC Below) northeast Queensland, in some instances geo- number 346; one staphylinoid female, data graphically very close to the sites of occur- as above except 840 m. berlesate ANIC num- rence of terraereginae. ber 347, 17004'S, 145037'E; one macropter- In most morphological features whereby ous male, two staphylinoid females (one with terraereginae differs from punctatus, the for- abdomen separated and aluminium coated), mer appears to be the more apomorphic. The data as above except 900 m. berlesate ANIC longer first antennal segment, proportion- 348, 17004'S, 145037'E; one staphylinoid ately more reduced eyes in the staphylinoids, female, 1140 m. near Mt. Haig, 30.VI.1971, the coarser and deeper punctation, the prom- Taylor and Feehan, berlesate ANIC 349, inent setiferous tubercles along the lateral rainforest, 17006'S, 145037'E; one staphyli- pronotal and corial margins, and the back- noid female, data as above except 750 m., wardly produced caudolateral angles of the berlesate ANIC 350, 17010'S, 145036'E; one forewings of staphylinoids all appear to be macropterous male, 20 km. south of Rav- derived features. This indicates the proba- enshoe, 800 m., 3.VII. 1971, Taylor and Fee- bility of terraereginae and related species han, berlesate ANIC 358, rainforest, 1 7°49'S, being derived from a punctatus-like ancestor 145°32'E; two staphylinoid females (alumi- and at least in this case becoming secondarily num coated), Beatrice River, Palmerston adapted to higher altitude conditions. Highway, 30.VII. 1956, from leaf litter, rainforest, about 700 m., T. E. Woodward; one staphylinoid female, Black Mountain Road, LITERATURE CITED 30 km. north of Kuranda, 4.XI. 1969, ANIC Ashlock, P. D. berlesate number 165, leaf mold, rainforest, 1964. Two new tribes of Rhyparochrominae: J. G. Brooks, deposited in Australian National a reevaluation of the Lethaeini (Hem.- Insect Collection, Queensland Museum, Het.: Lygaeidae). Ann. Ent. Soc. Amer- American Museum ofNatural History, Uni- ica, vol. 57, pp. 414-433, figs. 1-7. versity of Queensland and J. A. Slater Col- Gauld, I. D., and L. A. Mound lections. 1982. Homoplasy and the delineation of ho- DISCUSSION: Lilliputocoris terraereginae lophyletic genera in some insect groups. Syst. Ent., vol. 7, no. 1, pp. 73-86, figs. differs from L. punctatus the other Queens- 1-4. land species as follows: both macropters and McDonald, F. J. D. staphylinoids larger; antennae proportion- 1966. The genitalia of North American Pen- ately longer, segment I extending farther tatomoidea (Hem.-Het.). Quaestiones beyond apex of head and with length sub- Ent., vol. 2, pp. 7-150, figs. 1-520. equal to head width in macropters and sub- Putshkov, V. G. equal to interocular space in staphylinoids 1958. Larvae of Hemiptera-Heteroptera. I. (less in each instance in punctatus); antennal Lygaeidae. Rev. Entomol. URSS., vol. segment four ofstaphylinoids without a large 37, no. 2, pp. 392-413, figs. 1-39. dorsal concavity; pronotum, scutellum, and Schaefer, C. W. more 1964. The morphology and higher classifica- forewings coarsely and deeply punctate; tion ofthe Coreoidea (Hem.-Het.): Parts hairs along lateral margins of pronotum and I and II. Ann. Ent. Soc. America, vol. corium arising from conspicuous acute tuber- 57, no. 6, pp. 670-684, figs. 1-17. cles; macropters with an extensive dark 1965. The morphology and higher classifica- median area on hemelytral membrane; fore- tion of the Coreoidea (Hemiptera-Het- wings of staphylinoids with apical margins eroptera). Part III. The families Rho- 1982 SLATER AND WOODWARD: LILLIPUTOCORIS 23

palidae, Alydidae and Coreidae. Misc. Slater, J. A., and M. H. Sweet Publ. Ent. Soc. America, vol. 5, no. 1, 1970. The systematics and ecology ofnew gen- pp. 1-76, figs. 1-176. era and species ofprimitive Stygnocorini 1968. The morphology and higher classifica- from South Africa, Madagascar and Tas- tion of the Coreoidea (Hemiptera-Het- mania (Hemiptera: Lygaeidae). Annals eroptera). Part IV. The Acanthocephala- Natal Museum, vol. 20, no. 2, pp. group and the position of Stenoscelidea 257-292. Westwood (Coreidae). Univ. Connecti- Slater, J. A., M. H. Sweet, and R. M. Baranowski cut Occas. Papers Biol. Sci., vol. 1, no. 1977. The systematics and biology ofthe genus 3, pp. 153-199, figs. 1-46, tables 1-7. Bathydema Uhler (Hemiptera: Lygae- Scudder, G. G. E. idae). Ann. Ent. Soc. America, vol. 70, 1957. The higher classification of the Rhypar- no. 3, pp. 343-358, figs. 1-23. ochrominae (Hem., Lygaeidae). Ento- Slater, J. A., and T. E. Woodward mologists Monthly Mag., vol. 93, pp. 1979. In Slater, J. A., Hemiptera, Heteroptera: 152-156, figs. 1-6. Lygaeidae from Sri Lanka (Ceylon). Ent. Slater, J. A. Scandinavica. Report No.45 Lund Univ. 1975. On the biology and zoogeography of Ceylon Expedition of 1962, pp. 1-27, Australian Lygaeidae (Hemiptera; Het- figs. 1-6, tables 1-3 (preprint). eroptera), with special reference to the Stys, P. southwest fauna. Jour. Australian Ent. 1964. Thaumastellidae-a new family of pen- Soc., vol. 14, pp. 47-64, figs. 1-26, tables tatomoid Heteroptera. Acta Soc. Ent. 1-9. Czechoslovakia, vol. 61, no. 3, pp. 197741978). The incidence and evolutionary 238-253, figs. 1-14. significance of wing polymorphism in Sweet, M. H. lygaeid bugs with particular reference to 1967. The tribal classification of the Rhypar- those of South Africa. Biotropica, vol. ochrominae (Heteroptera: Lygae- 9, no. 4, pp. 217-229, figs. 1-6, tables idae). Ann. Ent. Soc. America, vol. 60, 1-7. no. 1, pp. 208-226. 1980. Systematic relationships of the Antillo- 1981. The external morphology ofthe pre-gen- corini of the Western Hemisphere ital abdomen and its evolutionary sig- (Hemiptera:Lygaeidae). Syst. Ent., vol. nificance in the order Hemiptera 5, no. 2, pp. 199-226, figs. 1-53. (Insecta). Rostria (Trans. Hemiptero- Slater, J. A., and J. Carayon logical Soc. Japan), no. 33, supplement 1963. Ethiopian Lygaeidae IV: a new preda- pp. 41-5 1, figs. 1-3. tory lygaeid from Africa with a discus- Woodward, T. E. sion of its biology and morphology 1959. Two new species ofMegalonotinae from (Hemiptera: Heteroptera). Proc. Royal New Guinea (Hemiptera: Lygaeidae). ent. Soc. London. Series A, vol. 38, parts Proc. Roy. Soc. Queensland, vol. 70, pp. 1-3, pp. 1-11, figs. 1-10. 51-56, figs. 1-2. Slater, J. A., and J. O'Donnell 1963. A new species of Tomocoris Woodward 1978. A new species of Cistalia from Brazil and a new related genus (Hemiptera: Ly- and comments on systematic characters gaeidae: Rhyparochrominae). Univ. in the Lethaeini (Hemiptera: Lygaeidae). Queensland Papers, Dept. Entomology, Florida Ent., vol. 61, no. 2, pp. 49-55, vol. 14, pp. 217-223, figs. 1-8. figs. 1-12.