HARVARD UNIVERSITY LIBRARY OF THE Museum of Comparative Zoology l^nisHto s $ i ^ mi THE UNIVERSITY OF KANSAS SCIENCE BULLETIN JUL •:•:•:•>:•:•:• JOL. ■ "* ''-^ R V , , J Host-relations of the Batfly Megistopoda aranea (Diptera: Streblidae) in Panama By William L. Overal S il s 11 Vol.52, No. l,pp. 1-20 June 27, 1980 ANNOUNCEMENT The University of Kansas Science Bulletin (continuation of the Kansas Uni- versity Quarterly) is an outlet for scholarly scientific investigations carried out at the University of Kansas or by University faculty and students. Since its incep- tion, volumes of the Bulletin have been variously issued as single bound volumes, as two or three multi-paper parts or as series of individual papers. Issuance is at irregular intervals, with each volume prior to volume 50 approximately 1000 pages in length. Effective with volume 50, page size has been enlarged, reducing the length of each volume to about 750 pages. 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Reprints of individual papers for personal use by investigators are available gratis for most recent and many older issues of the Bulletin. Such requests should be directed to the author. The International Standard Serial Number of this publication is US ISSN 0022-8850. Editor William L. Bloom Editorial Board Philip W. Hedrick Rudolf Jander Harvey Lillywhite Charles D. Michener Norman A. Slade Henry D. Stone George W. Byers, Chairman THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Vol. 52, No. l,pp. 1-20 June 27, 1980 Host-relations of the Batfly Megistopoda aranea (Diptera: Streblidae) in Panama^ ' William L. Overal^ CONTENTS Abstract 1 Introduction 1 Materials and Methods 2 Survey Results 5 Life Cycle of Megistopoda aranea 8 Behavior of Megistopoda aranea 12 Ecological Considerations 16 Summary 18 Resumen 18 Acknowledgments 19 Literature Cited 19 Abstract Megistopoda aranea (Coquillett, 1899) is the most common insect ectoparasite of the frugivorous bat Ar- biteus jamaicensis (Chiroptera; Phyllostomatidae) in Panama. This flightless batfly infests about 60% of indi- viduals of this host species, whereas its occurence on other hosts is sporadic and probably accidental. When these flies are separated from hosts, they survive only 5-20 h. Normally, they never leave the host except when females pupiposit. Puparia are placed in the bat roost in the vicinity of the host, usually on a rough surface above the host. The pupal stadium lasts about 23 days under conditions resembling those in a bat roost. Newly emerged adults can curvive about 36 h before feeding. Copulation was observed to occur two days after emergence. The larval stages which are passed within the female's body total 10 days. Adults of both sexes lived for about two months on caged A. jamaicensis. The main source of mortality was host grooming. M. aranea occupies the host's body fur and avoids the wing membranes. In choice arenas these batflies show a marked preference for rougher surfaces, behavior which is probably important for the maintenance of contact between batflies and their hosts. Increased locomotory activity was observed in response to sudden movements of air. In an "alert" posture the fly raises three legs while standing on its alternate legs. This often precedes a "procession" movement in which the animal moves sideways in an outward spiral on a hori- zontal surface. Host odor was not found to have attractiveness for M. aranea, although temperatures in the range of the host's skin temperature may be attractive. Megistopoda aranea is probably restricted to Artibetis jamaicensis by ecological factors, such as roosting be- havior of this host and colony size and stability, in addition to intrinsic factors, such as host odor or physiology. Introduction both the host and parasite. The present Tu ^ L ^ i ,. -u]- . study seeks to determuie how the associa- inat a host organism and its obligate ■> n ■ i ^^^^ ■ -^ ^ r \ , r 1 J lionhtiwttnrhthdiX.ny Megistopoda aranea parasite constitute a unihed system, hnked i.» u. •- .. u ; s f by physiological, ecological and behavioral 'Contribution no. 1705 from the Department of COmpatabilities, requires that answers to Entomology, University of Kansas, Lawrence, Kan- 11 r • r 1 ■ • sas 66045. problems of maintenance of the parasitic .p^^^^^^ ^^^^^^^. ^^^^^ p^^^^^^^ E^jlio Goeldi, relationship be sought in the biology of Caixa Postal 399, 66000 Belcm, Para, Brasil. The University of Kansas Science Bulletin (Coquillett, 1899) and its principal host, the Jamaican Fruit-eating Bat, Artibeits jamatcensis jamaicensis Leach, is main- tained. Aspects of this association which were investigated concern the hfe history, orientation behavior and occurrence of the batfly, the activities and habits of the host, and the co-existence of the host and para- site. Like other species of the dipterous fam- ily Streblidae, Megistopoda aranea is an obligate, blood-sucking parasite of bats. It is reported in the Neotropical Region from Mexico to Brazil, Paraguay and Peru, and throughout the West Indies (Wenzel, 1970). Determination of this species from the United States (Stiles and Nolan, 1931: 658) refers to an accidental intro- duction or is in error because none of the hosts of M. aranea are resident in that country. The batfly has been frequently collected in Panama and the Canal Zone (Wenzel et al., 1966), where the present study was conducted. Other specimens used in this investigation were collected in Costa Rica and Mexico. Contributions to our knowledge of the life cycles and natural history of streblids have been made by Muir (1911), Jobling (1949), Ross (1961), and Wenzel et al. (1966). Immature stages have been dis- cussed and figured by several authors (cf. Maa, 1971), and such findings are well summarized by Hennig (1952: 405). The biology of nycteribiid batflies is better known than that of streblids, primarily through the works of Rodhain and Be- quaert (1916), Hase (1931), Schulz (1939), Ryberg (1947), Hurka (1964), Leong and Marshall (1968), and Marshall (1970fl, 1970^). Aspects of hippoboscid biology relevant to this study have been presented by Coatney (1931), Bequaert (1953), and Hill (1963). Important questions pertain- ing to the adaptations and host-specificities of pupiparous Diptera have been raised and discussed by Theodor (1957), and Wenzel and Tipton (1966). Taxonomic placement of aranea in the genus Megistopoda Macquart (1852: 332) was by Maa (1965), when the genus to which it had originally been assigned, Pterellipsis Coquillett (1899), was placed in synonymy. Both Maa (1965) and Wen- zel et al. (1966) indicated that aranea may be a junior synonym of the type species of the genus, Megistopoda pilatei Mac- quart (1852). Megistopoda desiderata Speiser (1900) is a synonym of M. aranea (Coq.) (Aldrich, 1907). The morphology of M. aranea has been investigated by Job- ling (1949: 316; 1952: 134) and Machado- AUison (1966: 70). Megistopoda aranea was chosen for this study because of its relative abundance in Panama and because of the high degree of host specificity indicated by collections (Wenzel et al., 1966). Host bats can be kept in the laboratory (Novick, 1960), and their dietary and roosting habits are known (Goodwin and Greenhall, 1961). The fly is flightless and can be handled without damage. It is easily located on a host due to its size, light tan color, and long hind legs. Megistopoda aranea is dis- tinctive and not easily confused with other streblids. Even congeneric species may be distinguished under low magnification. Sexing is easily accomplished with ether- ized flies. The availability of natural colo- nies of infested Artibeus jamaicensis which could be observed directly permitted the formulation of hypotheses which could be tested with laboratory colonies of the host and parasite. Materials and Methods Field work was done in the Republic of Panama and the Canal Zone during December, 1970, and January, June, July and August, 1971. Laboratory work was done at the field station of the Smith- sonian Tropical Research Institute on Host-relations of the Batfly Megistopoda aranea Barro Colorado Island in the Canal Zone and at the Department of Entomology of the University of Kansas. Bats and batflies were caught in Pana- ma or the Canal Zone with the use of nets. Mist nets were employed in the man- ner illustrated by Greenhall and Paradiso (196(S), usually being placed over streams, along forest trails, or under fruiting fig trees known to be attractive to Artibeus bats. Within caves or buildings containing bat roosts, hand nets were used. In either case bats were placed individually in paper bags for transport to the laboratory or be- fore being etherized, to avoid the misasso- ciation of their parasites. Artibeus jamai- censis has been reported as a carrier of rabies virus in Panama (Constantine, 1970: 355; Keenan, pers. comm.). Because in- fected bats can transmit rabies to man, all bats were handled with gloves by person- nel vaccinated with duck embryo rabies vaccine. Batflies were removed with forceps from the bat or from the paper bag in which a bat was kept. Catching the flies by their hind legs did not appear to crip- ple them or afTect their behavior. Live flies were kept on a host until use. Dead flies were preserved in 70% ethanol, and hosts were preserved in formalin for iden- tification. Bats used in experimental infestations were housed in shaded areas outside the laboratory on Barro Colorado Island, in cages of 0.6 cm wire mesh. Although the batflies could easily pass through this screening, this was not found to be a source of batfly losses since the flies re- mained in contact with a host and did not wander off. Cages were cylindrical, 60 cm high and 45 cm diameter, with a detach- able bottom for convenience in cleaning. Generally, no more than 12 bats were kept in a cage of this size. A flight cage (10 m X 5 m X 3 m) with a concrete floor was used to house a colony of 155 bats. ■ The experimental bats, A. jamaicensis and other stenodermines, are frugivorous and can be maintained on diets of fruits and vitamin supplements. Fruits accepted by A. jamaicensis include bananas, figs, mangoes, melons and apples. These were peeled, sliced, and mixed with dried milk powder and vitamin syrup. Water was continuously available to the bats, but none was observed to drink. The environ- mental conditions under which the bats were kept varied more than did the condi- tions in bat roosts from which some bats were collected, but this did not appear to have a significant effect on the bats. The temperature varied from 16°C to 30°C and the relative humidity from 60% to saturation in a daily pattern (Fig. 1). In- dividual bats in the laboratory colony were marked by application of quick-drying paints to their hind claws which permitted identification from outside the cage. In addition to laboratory colonies, nat- ural colonies of A. jatnaicensis were ob- served in their roosts for comparative pur- poses. These colonies were located in three abandoned bunkers in Ft. Kobbe, C.Z., in Chilibrillo Caves, Chilibre, Panama, and in a culvert under Madden Forest Road, C.Z. Care was taken not to disrupt these colonies, and since individuals were not repeatedly observed, these bats were not banded or marked. Collections were made of bats and their parasites at several localities in order to survey the geographic distribution of M. aranea in Panama and its incidence and frequency of ocurrence on various hosts. Where possible, a sample of at least 25 Artibeus jamaicensis was taken at each locality in order to assess the effect of edaphic conditions on the host-parasite relationship. Those localities where this was possible are listed in Table 1 and mapped in Figure 2. Bats for experimental work were netted in June and July, 1971, at Balboa Heights, C.Z., under a fig tree The University of Kansas Science Bulletin 3 > O -T3 V 60 > > -a V c 'n C E '5 - n u a X J5 c • -5 ^ « OS o , S 3 O M U < 2 c pa B o u 6C o o "S 3 O U 3 - > u a. o Host-relations of the Batfly Megistopoda aranea near the railroad station, and at Summit Gardens (Botanical Park), C.Z. Voucher specimens have been deposited with the Museum at Kearney State College, Ne- braska (Alcoholic Mammal Collection, Catalog Nos. 1246-1271) in the care of Dr. J. Farney, who verified identifications made in the field. Specimens of M. aranea used in this study have been given to the Snow Entomological Museum of the Uni- versity of Kansas. Specific techniques relating to the care and use of batflies in behavioral and phys- iological experiments are presented in the pertinent sections below. Statistical tests are as given by Sokal and Rohlf (1969). Survey Results Megistopoda aranea has been reported from a variety of hosts. Reports include Phyllostomus sp. (Stiles and Nolan, 1931: 658), Artibeus yiwatanicus in Mexico (Hoffmann, 1953), Artibeus planirostris polax in British Guiana, Artibeus planiro- stris trinitatis in Trinidad and Tobago (Machado-Allison, 1966; Goodwin and o o UJ q: h- < UJ Q. UJ 35 30 25 20 15 1 _L JL 01:00 06 00 TIME 12:00 OF 18:00 2400 D AY 3 X UJ > Ul q: 100 90 80 70 60 I L _L X 0100 06:00 12:00 18:00 24:00 M E 0 F DAY Fig. 2. Map of Panama showing localities where bats and their parasites were collected. Place names and coordinates for these localities are given in Table 1. The University of Kansas Science Bulletin Greenhall, 1961), Artibeus jamaicensis par- vipes in Cuba (Matheson, 1928), Sturnira ludovici in Costa Rica (Tonn and Arnold, 1963), Artibeus j. jamaicensis, A. literatits palmaritm, Carollia perspicillata azteca, Desmodus rotiindus miirinus, and Phyllo- stomus discolor in Panama (Wenzel et al., 1966). The published host records are difficult to interpret because of the unset- tled and specialized nature of bat taxon- omy and because of the possibilities for errors in associating hosts and parasites during collecting or labeling. Wenzel et al. (1966) believe Artibeus jamaicensis and A. literatus are the primary hosts of M. aranea in Panama, although A. literatus, as interpreted by them, may represent a species complex (Wenzel, pers. comm.). Examination of collected lots of 14 bat species in Panama revealed that M. aranea was apparently restricted to three species, Artibeus jamaicensis, A. literatus, and Carollia perspicillata. Figure 3 shows the percentage of each of these species which was infested with M. aranea. Carollia per- spicillata is probably not a normal host of M. aranea because of its low frequency of parasitization and because no C. per- spicillata had more than one individual of M. aranea, suggesting a random transfer. The status of A. literatus as a host of M. aranea is uncertain. A. literatus and A. jamaicensis have been netted in the same collection lot, suggesting that they may forage together, but they have not been observed to roost together. They do, how- ever, share the same caves in Panama, such as the Chilibrillo Caves, and the possibility that M. aranea on A. literatus may have dispersed, if only temporarily, from A. jamaicensis cannot be ruled out. Of the several previously reported Panamanian hosts of M. aranea, only the vampire, Desmodus rotund us, was not adequately sampled. More than 200 vam- pire bats have subsequently been collected from the states of Veracruz, Oaxaca and Chiapas in Mexico, however, and M. No. Name Table 1. Names and coordinates of localities mapped in Figure 2. Coordinates'^ Description 1 Ancon, C.Z. 2 Balboa Heights, C.Z. 3 Barro Colorado Is., C.Z. 4 Frijolcs, C.Z. 5 Gamboa, C.Z. 6 Ft. Kobbe, C.Z. 7 Madden Forest, C.Z. 8 Ft. Sherman, C.Z. 9 Summit, C.Z. 10 David, Chiriqui 1 1 Aguadulce, Code 12 Anton, Code 13 Penonome, Code H Rio Hato, Code 15 Parita, Hcrrcra 16 Los Santos, Hcrrera 17 Arraijan, Panama 18 Calzada Larga, Panama 19 Cerro Campana, Panama 20 Chilibre, Panama 21 Pacora, Panama 22 Panama (city), Panama 23 Portobello, Colon 24 Chepo, Daricn 25 Limon, Costa Rica 8°57'N-79°34'W 8°57'N-79°34'W 9°09'N-79°5rw 9°10'N-79°49'W 9°06'N-79°42'W 8°54'N-79°36'W 9°05'N-79°39'W 9°21'N-79°57'W 9°03'N-79°40'W 8°26'N-82°26'W 8°14'N-80°33'W 8°24'N-80°16'W 8°31'N-80°22'W 8°22'N-80°16'W 7°59'N-80°32'W 7°45'N-80°21'W 8°57'N-79°41'W 9°10'N-79°34'W 8°41'N-79°56'W 9°08'N-79°38'W 9°04'N-79°18'W 8°58'N-79°32'W 9°41'N-79°4I'W 9°10'N-79°06'W 9°58'N-83°08'W under fig tree under fig tree forest clearing forest trail under fig tree abandoned bunker forest clearing wet forest Botanical Park near stream forest near docks under fig tree near Rio Zarati near shore forest boundary under fig tree forest near C.Z. abandoned bunkers cloud forest ChilibriUo Caves under fig tree under fig tree forest trail forest coffee finca ^Coordinates were compiled from several maps and from Fairchild and Handley (1966). Host-relations of the Batfly Megistopoda aranea aranea has not been found associated with any of them. M. aranea is found in these areas on Artibeiis, and its absence from Desmodits is significant. The results of the survey confirm the statement by Wenzel et al. (1966) that "this species appears to be a parasite pri- marily of Artibeits jatnaicensis'' and secon- darily of A. literatiis in Panama. Caution should be exercised in extrapolating these results to areas outside Panama, but collec- tions made in Costa Rica and tropical Mexico indicate that subspecies of A. ja- maicensis are the primary hosts of M. aranea in these countries also. Collections in the care of Dr. Wenzel (pers. comm.) show this to be true also in Venezuela, Surinam, Colombia and San Salvador. Megistopoda aranea can be considered monoxenous or, perhaps, oligoxenous. Field collections can shed light on the behavioral aspects of host selection. No difference was found in the average in- festation levels between males and females of Artibeits jamaicensis, indicating that the fly may not discriminate between the sexes of its host. This was confirmed in the laboratory with choice tests. The fre- quency distribution of infestation of M. aranea on A. jamaicensis follows the ex- pected Poisson (random) distribution 75r 50- z MJ 25- XIL ADTieEUS JAMAICENSIS (N-= 1I2) AITKEUS I llf HAtUS CADOUIA PtISPICILIAIA (N= 43) (N- 5 5) Fig. 3. Frequencies of infestation of Megistopod a aranea on three bats species in the Canal Zone. (Table 2 and Fig. 4), indicating that the flies do not preferentially segregate with one another and that hosts are nearly alike in their attractiveness. The frequen- cies of occurrence of both male and female flies on the same host are not significantly dififerent from the expected frequencies, and it may be assumed that flies do not pair with flies of the opposite sex in host selection, in contrast to the situation noted by Hurka (1964) in several European nycteribiids. No correlation of the para- site load with the weight or forearm length of the host was found in a sample of (S5 bats. Although the parasite is buffered from changes in environment by the homeo- stasis of the host, there were slight differ- ences in the average infestations of Artib- eits jamaicensis between the wet and dry seasons of 1971. Sixty-two bats caught dur- ing the wet season had an average of 0.843 flies per host, compared with 0.611 flies per host found on 57 bats caught during the dry season. That these differences could reflect a direct effect of climate on the parasite is only one possibility. It is also likely that there is an effect on the host, reflected in some significant altera- tion in its biology. Mares and Wilson (1971) have found a marked seasonality in the breeding cycle of several neotropical bats. Whether such a condition could in- fluence roost selection or sociality of the bats, in such a way that the host-parasite equilibrium would be disturbed, has yet to be explored. Observations on colonies of Artibeits jamaicensis in natural and man-made roosts indicate how the dynamics of host populations may affect the population lev- els of Megistopoda aranea. In several sit- uations bats roosted in close proximity to one another and the fur over which flies moved was effectively continuous from bat to bat. Host-to-host dispersal by this means was suspected and later confirmed. If the 8 The University of Kansas Science Bulletin proximity of roosting bats in a colony is assumed as an index of sociality, the possi- ble relationship of sociality to parasitism by Megistopoda aranea becomes clearer; dispersal to new hosts becomes dependent upon the sociality of the host. Colony size may also be important for the presence or absence of the parasite, since smaller col- onies may not provide sufficient refuge from the grooming activities of individual hosts. While there are no data to support this conjecture, it remains of interest for further investigation. Life Cycle of Megistopoda aranea The free-living, non-parasitic stages of Megistopoda aranea are reduced, as is the case throughout the Pupipara. The larval stadia last ten days, based on the mini- mum time between successive pupal depo- sitions by the same female. During this period the larva is nourished in the female by a uterine gland secretion (Hagan, 1951) and is only indirectly dependent upon a host bat. Attempts to demonstrate that gravid females ingested more blood than non-gravid females were inconclusive, and future investigations along these lines may require radioactive tracers to determine the amount of blood ingested. A gravid female ready to exf)el the prepupa can be distinguished by the degree of enlarge- ment of its abdomen. Such females were removed from hosts and placed in stop- pered vials for the collection of puparia. Within 10 min of its deposition, the Table 2. Frequencies of absence and single and multiple occurrences of Megistopoda aranea on Artibeus jamaicensis. Expected frequencies are based on a sex ratio of 1 male: 1.25 females and a Poisson distribution where the average is 0.843 flies per host for 412 hosts. All hosts were caught in the Canal Zone during June and July of 1971. No. of flies Sexes of flies Observed frequency Expected frequency 0 188 177.3 \$ 74 15 86 2$ 17 Ul$ 27 29 24 i$ 2 1522 5 2515 10 32 1 45 0 1532 1 2522 1 3512 0 42 1 .... 0 65 0 1552 0 2542 0 3532 1 4522 1 5512 0 62 0 133 68 18 66.4 83.1 12.4 31.1 19.5 1.6 5.8 7.3 3.0 0.15 0.73 1.36 1.14 0.35 0.001 0.006 0.017 0.026 0.024 0.013 0.003 149.5 63.0 17.7 3.73 0.63 0.09 Observed and expected frequencies arc not significantly different. (Chi-squared test.) Host-relations of the Batfly Megistopoda aranea 200r • - POISSON DISTRIBUTION 5 6 FLIES PER HOST Fig. 4. Frequencies of occurrence of Megistopoda aranea on Artibeus jamaicensis. subglobose larva darkens from a trans- parent colorless form to a brown puparium with evident segmentation. The ventral surface of puparia deposited on glass did not darken, suggesting that the process de- pends upon contact with air. In addition to the protection afforded it by the tough- ened larval cuticle, the larva is usually deposited in an indentation. The actual orientation of the puparium does not ap- pear to be important, and both vertical and horizontal surfaces were selected by M. aranea for larval deposition. A char- acteristic place in a stoppered vial was on the glass between the cork and the wall of the vial. In natural colonies of Artibeus jamaicensis, puparia were found in areas above the roosting bats. This pupa-posi- tion site differs from that of fully winged streblids which may deposit pupae in areas quite removed from the roosting bats, as in a different chamber of a cave, and from that of hippoboscids which may deposit pupae loosely in nesting materials or on the host itself. This last case is not a possibility for M. aranea, however, be- cause of the frequent and vigorous groom- ing by the host and the long develop- mental period of the pupa. The pupal stadium lasted 22 to 24 days at 100% R.H. and approximately 22°C. The mean developmental time for 25 pu- pae was 23 days. The puparium has two posterior spiracles and an oblong base by which it is attached to the substrate (see Hennig, 1952), The dimensions of 25 pu- paria of M. aranea averaged as follows: length — 1.6 mm; width — 1.1 mm; height — 0.8 mm. The teneral adult emerges through an anterior operculum and re- mains near the empty puparium for sev- eral hours. Table 3 shows the survival times of unfed teneral adults. The teneral fly is readily distinguished by its pale color 10 The University of Kansas Science Bulletin and crenulate abdomen. A ptilinum was not noted in newly emerged flies. As observed by Ross (1961) in Ari- zona, teneral individuals of Trichobiiis spp. cannot mate until they have fed, prob- ably because of the deformation or small size of the abdomen which may be me- chanically incapable of copulation. This could apply to M. aranea also because no unfed teneral flies were observed to mate. Mating occurs on the host bat and usually lasts one to five minutes. Females were sometimes seen to mate only a few min- utes after depositing a pupa. The presence of both sexes in nearly equal numbers (Table 3) suggests that mating is required for reproduction and that the storage of sperm by the female, if it occurs, is of little importance. Since six adult females separated from males produced maximally only one pupa each during the following three weeks and fe- males which had never been kept with males produced none, a separate mating is probably recjuired for each offspring produced. Attempts to mark individual flies for life -history studies were unsuccessful due to an apparent toxicity of the oil- and lacquer-based paints which were employed, and measurements of longevity and fe- cundity are therefore indirect. To meas- ure longevity, newly enclosed M. aranea were placed on ten caged Artibeiis jainai- censis hosts. The flies were counted at weekly intervals (Fig. 5) until the last fly had died or been lost. The average life- span of these 40 flies was 29 days, and the survivorship curve indicates that the bulk of the mortality was due to sources other than old age. Thirteen dead and damaged flies were recovered from the bottom of the bat cage. Certainly, both the groom- ing of the host (possibly involving pre- dation) and disassociation from a host should be considered as major causes of mortality. Adults of M. aranea were ac- cepted and eaten by A. jatnaicensis when offered in the laboratory. D. Howell (pers. comm.) has found streblids in the stom- achs of several species of Costa Rican bats. An acarine parasite of M. aranea adults, Monunguis streblida Wharton (Wharton, 1938; Linquist and Vercammen-Grand- jean, 1971), has been reported, but no hyperparasites were seen during this study. Pupal mortality was measured as the percentage of puparia which did not give rise to adults by the end of four weeks. Puparia were deposited on a piece of ply- wood on top of the outdoor bat cage over the dark corner in which the bats roosted. The board was then removed to the top of an empty cage, and the puparia were examined at the end of a month. Twelve per cent of the puparia (23 of 192) had not given rise to adults, although no rea- son for their failure could be seen. Two puparia had been damaged (probably in relocating the board) and had been at- tacked by fungus. The survival of pupae may be assumed to be a function of en- Table 3. Survival times of Megistopodu aranea separated from hosts. Both adults which had been allowed to engorge and unfed, newly eclosed tenerals were kept at 100% R.H. and 22 C. Survival times of engorged adult flies (in hours) Survival times of unfed teneral flics (in hours) 3 5 8 14 17 19 20 18 26 Z6 38+ Tenerals survived significantly longer than engorged adults. (P<.05) Mann-Whitney-t/ test.) Host-relations of the Batfly Megistopoda aranea 11 2 4 6 Fig. 5. Survivorship of Megistopoda aranea on ten caged Artibeus jamaicensis. vironmental conditions. The present ob- servations, therefore, may reflect more upon the conditions provided for pupal development than on the usual successes of the pupal stage. In Chilibrillo Caves and in bunkers at Ft. Kobbe, puparia which resembled those of M. aranea were found in prox- imity to colonies of Artibeus jamaicensis. Those removed at random in various seasons from the roosts were in a variety of developmental stages, suggesting that reproduction by the bat fly is asynchronous and continuous. The cave and bunker were similar environments with moderate temperatures and a relative humidity near saturation, but colonies of Artibeus ja- maicensis were also observed in foliage and in a hollow tree where conditions were much different. Pupae in the labora- tory developed at 18°C and those in out- door bat cages at temperatures 21 to 30°C, indicating that unsheltered colonies, if sufficiently permanent, may be para- sitized by M. aranea. Life history observations on M. aranea indicate a low reproductive potential. Ig- noring larval and pupal mortality and assuming adult age-specific fecundities to be ecqual, the assembled observations would indicate a net reproductive rate, Ro, of 1.45, .r where Ro^'^L-mx, L- is survivorship and m.r is age-specific fecundity. This figure, while derived from data which may in- clude the effects of predation and sub- 12 The University of Kansas Science Bulletin optimal conditions, indicates that a popu- lation of M. aranea would multiply 1.45 times in a generation, while under opti- mal conditions a greater net reproductive rate would be expected. It seems reason- able to assume that under natural condi- tions M. aranea populations are stable in the long run, although locally there may be extinctions, probably followed by rein- troductions and population growth. The low reproductive potential of M. aranea follows from its breeding biology which limits a single female to producing maxi- mally one pupa every ten days. Behavior of Megistopoda aranea The dominant aspects of the behavior of Megistopoda aranea are its blood-suck- ing habit and its dependence upon a host. Flies removed from hosts (Table 3) sur- vive for only a short time, in contrast to other groups of ectoparasites which may remain apart from a host for months. Megistopoda aranea, therefore, is found in contact with a host during all adult activi- ties with the exception of pupal deposition, which generally occurs close to a resting host during the day. In the terminology of Camin (1963), M. aranea can be said to be a "permanent" ectoparasite. On its host Megistopoda aranea can be found on the furred parts of the body, but prefers sites around the neck and shoul- ders where the fur is long and in the axillary regions below the wings. When disturbed, flies often run into the bat's ears. The fly moves on the host's fur by pushing itself with alternating extensions of the long hind legs. The thoracic ster- num is flat and sled-like, and the first two pairs of legs articulate to the sides. The fly can move with facility over the host pelage with its thorax at the level of the longer guard hairs and is capable of mov- ing forward, backward, or to the side. No fly was observed to jump. During host grooming, Megistopoda aranea was seen to retreat to the lower back of its host to a position just anterior to the uropatagium. This portion of its body could not be reached by the host, which groomed itself with its hind claws, thumb and mouth. Host bats were not observed to groom one another as do some birds. Although no pathological lesions were observed at the feeding sites of M. aranea, infested Artibeus bats were ob- served to groom more intensively and fre- quently when the parasite load was ex- perimentally increased. Flies rarely went onto the wings of the host and would not remain on newly born A. jatnaicensis, which are hairless, suggesting that thig- motaxis is important in host-recognition. This hypothesis was tested by offering flies a choice of substrates in a closed arena; the results are shown in Table 4, Flies consistently chose rougher substrates. Tac- tile stimuli are probably important in the fly's maintenance of contact with its host. Megistopoda aranea and other streblids do not engorge, but feed intermittently throughout the day. It is possible that more feeding is done during the daylight hours when the bats are roosting. This is based upon the frequency of defecation by the fly (Fig. 6). Feces were collected on paper at the bottom of a cage containing one Artibeus jatnaicensis and five M. aranea. The higher daytime feeding rates may be induced by a greater rate of desic- cation at the lower relative humidity pres- ent during the day, or they may be an adaptation to the resting habits of the host during the day. Favored areas for feeding on the host are around the ears and neck and in the axillary regions. Dissections showed that flies apparently ingest whole blood and do not concentrate blood cells. A liquid fecal drop is produced. Although no volumetric measure of the amount of blood ingested by a fly during a day was possible, it remains a possibility that sev- Host-relations of thl Batfly Megistopoda aranea 13 eral flies could seriously affect the health of their host. Megistopoda aranea does not appear to bite humans as do some species of Tricho- biits (Ross, 1%1). Flies were frequently handled and in two cases given access for several hours to the upper arm of the in- vestigator, but they did not bite. No ani- mals other than bats were accepted as hosts; mice and rats were rejected and flies placed on them left or died. The texture, odor and temperature of the host may be signal stimuli or feeding stimuli to the fly. Although Megistopoda aranea does not normally leave the body fur of its host except for pupal deposition, flies which have been removed from their hosts ex- hibit search behavior. Placed on a hori- zontal pane of glass under diffuse light, such a fly would move in a random fashion for several minutes before assuming an "alert" position in which it would stand on its fore- and hind-leg of one side and its mid-leg of the other side. This position was observed in over 50 flies. The legs not used to support the body were ex- tended above the body and waved about. This behavior suggests that tarsal recep- tors such as those of the Nycteribiidae, or some other type of sensory organ, may be present on the legs of M. aranea (Maa, 1971). This stance may also be adaptive for grasping hosts moving nearby. The fly can be made to abandon its "alert" stance and move rapidly if a sudden air current is directed at it, even in the ab- sence of host odor, as from an empty syringe. The fly will turn toward the source of the air current and run forward for several seconds or until the air current is stopped. Such behavior may be adaptive if a potential host were to cause an air current. In a second type of "search" pattern seen repeatedly in flies removed from hosts to a horizontal surface, the fly moves side- ways in a spiral, halting at intervals to raise one or more legs. How effective either of these "search" patterns would be in the more complex environment of a bat roost is not known. In a "T-tube" apparatus with an inter- nal diameter of about 8 mm, flies were seen to encounter, turn and walk against the flow of air (22 of 25 flies, P < .05, Chi-squared test). This may be related to their response to a current of air when standing still. Table 4. Preferences of Megistopoda aranea for type of substrate. Fifteen flies (five at a time) were placed in a petri dish with two substrate types, each covering half of the arena, and were allowed to move freely for 20 min before their selection was recorded. Comparisons were made using the Mann-Whitney-L^ test. (*=f<.05) Test Materials Trial Trial Trial Preferred no. as substrates ABC material 1 Glass 0 1 0 Paper 5 4 5 * 2 Glass 0 0 0 Cotton cloth 5 5 5* 3 Paper 4 3 5 Cloth 1 3 5 ns 4 Water 0 0 0 Glass 5 5 5* 5 Moist towel 0 3 1 Glass 5 2 4 ns 6 Cotton 0 1 - 0 Bat fur 5 4 5 • 7 Wet bat fur 0 1 0 Dry bat fur 5 4 5 * 14 The University of Kansas Science Bulletin 40|- (/) it! 30 20 10 04:00 08O0 12=00 1600 2000 2400 TIME OF DAY 40t- U li! 301- 20- 10 n 04«0 0800 12:00 16:00 2000 2400 TIME OF DAY Fig. 6. Defecation rate of five feeding Megistopoda aranea during a 48-hour period. Table 5 shows the results of chemo- sensory tests using a "T-tube" in which the various stimuli were allowed to diffuse from one end of the "T" for one-half hour before the flies were introduced into the stem. None of the stimuli was shown to have a marked attraction or other effect except the commercial insect repellent. This indicates that, while some olfaction is present, it may not be used in host- finding. This could be expected since the environment of the bat fly is probably saturated with the host odor, and no gra- dient of odor could be followed to find the host bat over the short distances the flies walk from their hosts. Host temperature is a valid clue for many ectoparasites in host-recognition and host-finding over small distances. It may also be a feeding stimulus. It appears that M. aranea is sensitive to substrate tempera- ture (Table 6). Flies consistently aggre- gated in the area of a temperature gradient which was between ?)i and 38 C. This temperature range includes the body fur temperature of resting Artibeits jamaicen- sis which was measured with a "banjo" probe and a Yellow Spring Instrument Host-relations of the Batfly Megistopoda aranea 15 Company tele-thermometer. These bats had a surface temperature of 32° =t 2°C, but temperatures rose rapidly as the bat struggled to free itself or bit the thermistor probe. Bats generally have fluctuating body temperature, and many resting bats allow their temperature to rise or drop to the ambient temperature (Henshaw, 1*^)70) . Artibeiis bats were not tested in an unstressed condition, and it is not known if this is true for them. Only to the extent that their temperature remains above ambient can temperature preference be dependable for host-finding by their parasites. Host-finding is initially important to Megistopoda aranea of both sexes after eclosion. The teneral adult can survive longer without a host than can a fed adult (Table 3). This result agrees with the findings of Marshall (1970^) and Leong and Marshall (1968) for two nyc- teribiid species. The initial advantage is probably greater for M. aranea, however, because no host-associated stimulus for eclosion seems to be involved which would indicate the presence of a host, such as has been reported for Basilia hispida (Nycteri- biidae) (Marshall, 1970a, 1970^). Even if such a stimulus were present, the roosting sites of Artibeiis are shared with other species, and the bat which may trigger eclosion could be one of a number of stenodermine or other bats. No doubt newly emerged flies could get onto irregu- lar hosts which have replaced an Artibeiis colony. Flies did not select a particular relative humidity in a gradient, nor were they excited or stimulated to move by low con- centrations of carbon-dioxide in the air. They did not preferentially select either light or dark areas in an arena, and their distribution in a vertical cylinder was without regard to gravity. Megistopoda aranea, like other streblids, can walk up- side-down on glass and could even adopt the "alert" stance in this orientation. Table 7 indicates that Megistopoda aranea has the ability to discriminate be- tween host species that are roosting to- gether or in close proximity. The sensory basis for this discrimination is unknown, although olfaction is suspected. Another Table 5. Results of chemosensory behavioral tests. For an explanation of the experimental design see text. Stimulus Locomotory response Host hair weak positive Host breath none Host faeces none Water washings of host none "OFF" (50%, N,N,-dimethyl-meta-toluamide) .... strong negative Number responding (20 flies) 1 0 0 0 19 Table 6. Temperature preference of Megistopoda aranea. Positions of flies on a warmed glass plate were recorded with reference to substrate temperature after 20 minutes of free movement. A shielded 100-watt light bulb was the heat source. Air temperature was 18°C. Expected num- bers of flies are based on relative areas of the substrate in each temperature range. Temperature (°C) No. of flies Expected number 18-20 2 12 21-23 4 9 24-26 4 9 27-29 3 8 30-32 9 6 33-35 18 4 36-38 7 2 Observed distribution of flies by area does not fit the expected (random) distribution. (P<.05, Chi-squarc test.) 16 The University of Kansas Science Bulletin factor observed which may bear on these results is that Carollia perspicillata, Glos- sophaga soricina, and Artibeus toltecus are small bats and groom themselves more vigorously and frequently than do the larger Artibeus jamaicensis and A. litera- tus. Batflies, therefore, may have been groomed off by the more active hosts. The fur of the smaller bats is also shorter and may be more difficult for the batflies to grasp. Thus, the association of parasite and host could result from factors other than sensory orientation. Similarly, the survival of Megistopoda aranea on various hosts may depend upon factors other than the nutritive require- ments of the fly or the presence of phago- stimulants. Of the bat species listed in Table 8, Carollia spp., Glossophaga sp., Uroderma bilobatum and Stirnira sp. did not adapt well to captivity. Individuals of these species showed signs of agitation and stress, did not adapt well to the diet pro- vided, and had to be released after about a week if they were not to be kept as voucher specimens. Three Artibeus cine- reus were kept with sheaths over hind claws, wings and mouth, and were force- fed for one week during which time two M. aranea were maintained on each with- out loss. It seems reasonable, therefore, that the survivorship of the bat fly on sev- eral hosts was dependent upon differences in host activities, including grooming. Ecological Considerations To the batfly the bat is both habitat and food, while the batfly causes a loss of Table 7. Host species selection by Megistopoda aranea. Four flies were placed in each cage containing a male and female of two bat species. Host-selection of flies was recorded after one, two, and three days. Host-selection by sex was non-significant in each species. (*P<.05; Mann-Whitney-Z7 test.) Cage Hosts Day 2 Day 3 Day 4 I Artibeus jamaicensis 13 1 Artibeus literatus 3 13 II Artibeus jamaicensis* 4 4 3 Carollia perspicillata 0 0 1 III Artibeus jamaicensis* 4 3 3 Glossophaga soricina Oil IV Artibeus jamaicensis 2 12 Phyllostomus discolor 2 3 2 V Artibeus jamaicensis* 4 3 4 Artibeus toltecus 0 10 VI Artibeus literatus* 3 4 4 Carollia perspicillata 1 0 0 Table 8. Survival times of Megistopoda aranea on various hosts. Two flies were placed on each of three bats of each listed species. Each bat was kept separately in a fine wire-mesh cage and checked daily. Number of flies surviving Host species Day 1 Artibeus j. jamaicensis 6 Artibeus literatus 6 Artibeus cincrcus 6 Carollia perspicillata 6 Glossophaga soricina 6 Glossophaga sp 6 Phyllostomus hastatus 6 Phyllostomus discolor 6 Uroderma bilobatum 6 Stirnira sp 6 Day 2 Day 3 Day 4 Day 5 6 6 6 6 6 6 6 6 5 2 2 1 4 1 0 0 5 5 5 5 4 0 0 0 6 6 4 4 5 3 1 1 4 3 3 3 3 1 1 0 Host-relations of the Batfly Megistopoda aranea 17 fitness to its host bat. The accommoda- tions between hosts and parasites are paral- leled on the level of their populations, for the relationship may be viewed as involv- ing a population of parasites and a popu- lation of hosts, the properties of neither being fully derivable from the properties of individual hosts or parasites. Although a single batfly can affect only one bat at a given time, it may parasitize several or even many host individuals dur- ing its lifetime. Table 9 shows the results of an inquiry into the ability of M. aranea adults to disperse from one bat to other bats in its colony. So rapid was this dis- persal and redistribution that the bat col- ony must be considered the effective host unit for a batfly for any but the shortest time scale. It is, however, premature to conjecture about the effect of the presence of additional bats in the colony upon the immediate environmental conditions af- fecting the parasite since the effective mi- croclimate of the batfly is not known, but speculation on the effect of host population densities or colony sizes on parasite inci- dence may now be fruitful. If, for exam- ple, the size of a colony governs its sta- bility or the permanence of its roost in some manner, it would greatly influence the reproductive success of its associated batflies, which require bats to be present for emergent adults following pupation. Likewise, a larger colony may afford a batfly more opportunity to abandon a dy- ing or unsuitable bat for a more suitable host. The potential of Megistopoda aranea to transmit diseases would seem great. Ser- gent and Sergent reported the transmission of Haemoproteus columbae by the pigeon- fly Pseudolynchia maura, a hippoboscid, in 1906 (Coatney, 1931); and O'Roke (1930) demonstrated that Haemoproteus lophortyx, a blood parasite of quail, is transmitted by another hippoboscid, Lyn- Table 9. Host-to-host transfers of Megistopoda aranea. Ten flies were introduced onto one of five Artibeus jamaicensis (A) in each of five cages to determine how the flies would distribute themselves among the caged hosts. Flies were not marked in order to avoid possible injury. The incidence of the flies on each of the hosts was checked daily for five days. Cage Host Day 1 Day 2 Day 3 Day 4 Day 5 I A 10 8 5 2 2 B 0 0 2 3 1 C 0 0 0 11 D 0 110 3 E 0 0 0 2 1 II A 10 6 3 2 3 B 0 3 2 2 2 C 0 0 12 1 D 0 12 2 1 E 0 0 0 0 0 III A 10 3 3 0 1 B 0 12 12 C 0 10 12 D 0 3 4 3 2 E 0 2 0 4 1 IV A 10 2 2 3 3 B 0 12 12 C 0 4 0 12 D 0 2 0 2 1 E 0 2 5 0 2 V A 10 5 2 0 3 B 0 0 15 3 C 0 2 12 2 D 0 0 4 10 E 0 3 2 0 0 18 The University of Kansas Science Bulletin chia hirsuta. Transmission of this sort is more complex than the mechanical trans- mission required for the bacterial or viral diseases to which Anibeus jamaicensis is known to be susceptible. Moving from one host individual to another, M. aranea would appear to pose a threat to the health of Artibeiis colonies if it serves as a vector for diseases of its hosts. Rabies may be spread as an aerosol, and investigations of the transmission of rabies in bats at the Trinidad Virus Laboratory did not impli- cate batflies as vectors (Greenhall, pers. comm.). One limitation to the vectorship- potential of M. aranea is its narrow host range, but further inquiry into this matter is warranted. Artibeiis jamaicensis is the host of other blood-sucking ectoparasites which would appear to compete with Megisto- poda aranea. Other parasites collected on A. jamaicensis during the course of this investigation included three streblids, Paratrichobiiis longicrus, Aspidoptera biisc- kji, and Metelasmits psettdopterits, and a spinturnicid mite, Periglischrus iheringi. Other parasites of Artibeiis jamaicensis are reported by Wenzel et al. (1966). While these three other streblids, especially, could compete with M. aranea, other parasites did not exclude M. aranea. It would seem that Artibeiis in Panama has not been "saturated" with parasites and that popu- lations of parasites of Artibeiis jamaicensis are not at their maxima. Competition be- tween ectoparasites would not be expected if their resources were not limiting. If, however, a host bat is considered as a limited habitat, it is possible that popu- lations of Megistopoda aranea arc at their maxima and that competition between ectoparasitic species for space, rather than food, may occur. The potential maximum population would be a function of the host's behavioral tolerance fcjr parasites, and parasites in excess of this limit would be removed by host grooming. The ad- vantage under such pressures would lie with those parasites which could best avoid being dislodged. M. aranea is by far the most common ectoparasite of A. jamaicen- sis and presumably has this ability to a greater extent than do other parasites. Summary Megistopoda aranea is the most fre- quent ectoparasite of Artibeiis jamaicensis in Panama where this study was con- ducted. Artibeiis literatiis is a much less frequent host. In spite of this, M. aranea is best considered as monoxenous, forming a stable system with its host. Megistopoda aranea disperses from host to host within a colony by direct transfer between adja- cent hosts. The frequency distribution of flies on Artibeiis jamaicensis resembles the expected Poisson distribution. The sexes of Megistopoda aranea are present in near- ly equal numbers. Copulation is required for each reproduction. The larval stages last about 10 days in the body of the mother, and the pupal stage lasts about 23 days. The adult Megistopoda aranea lives several weeks. Breeding occurs throughout the year. The reproductive potential of the species seems to be limited by its viviparous habit. A description of the general and orientation behavior of Megistopoda aranea is given. Host-speci- ficity of Megistopoda aranea may arise from ecological aspects of the host, rather than from the fly's sensory orientation. Resumen Megistopoda aranea es el ectoparasito mas frccuente en Artibeiis jamaicensis de Panama. Artibeiis literatiis es un huesped de segundo orden, por su escasez. Por lo que Artibeiis jamaicensis puede ser con- siderado como un huesped especifico para Megistopoda aranea. El metodo de disper- sion usado por Megistopoda aranea dentro de una colonia es por contacto directo cntre los hucspedes. La distribucion de Host-relations of the Batply Megistopoda aranea 19 frecuencia de Artiheus jamaicensis semcja la distribucion de Poisson. Hombras y machos de Megistopoda aranea se encuen- tran en igual numero. Cada vez que sc reproducen cs necesaria la copulacion. El estado larval dura 10 dias deiitro del cucr- po de la madre el estado pupal dura 23 dias. Los adultos de Megistopoda aranea viven varias semanas la reproduccion se realiza durante todo el aiio. El potencial reproductivo de la especie parece estar limitado por sus modo viviparo de repro- duccion. Descripcion acerca de la con- ducta de Megistopoda aranea es incluida. La especificidad de Megistopoda aranea esta possiblemente mas replacionada con la ecologia del huesped que con la orien- tacion sensorial del parasito hacia el hue- sped. Acknowledgments I wish to acknowledge the advice and assistance given me in Kansas and Pana- ma by Dr. George W. Byers and Dr. Rob- ert E. Beer of the University of Kansas. Dr. Rupert L. Wenzel, Curator of Zool- ogy of the Field Museum of Natural History, made helpful suggestions and provided specimens for comparison in the identification of the batfly. Bats were identified by Dr. E. Tyson, and by Dr. John Farney. Mr. Charles M. Keenan has been most generous with his time and advice in the field. I also wish to thank Dr. William Ramirez for advice and as- sistance in the field. I thank the Smithsonian Tropical Re- search Institute (Ancon, Canal Zone) for partial support and the use of the research facilities at Barro Colorado Island. 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In A Catalogue of the Diptera of the Americas South of the United States, Part 100. Museu de Zoologia, Universidade de Sao Paulo. Wenzel, R. L., and V. J. Tipton. 1966. Some re- lationships between mammal hosts and their ectoparasites, p. 677-723. In R. L. Wenzel and V. J. Tipton (eds.), Ectoparasites of Panama. Field Museum of Natural History, Chicago. Wenzel, R. L., V. J. Tipton, and A. Kiewlicz. 1966. The streblid batflies of Panama, p. 405- 675. In R. L. Wenzel and V. J. Tipton (eds.). Ectoparasites of Panama. Field Museum of Nat- ural History, Chicago. Wharton, G. W. 1938. Acarina of Yucatan caves. Carnegie Inst. Washington Publ. 491:137-152. W5S55*%::¥:::::WSft•:w^x•:•^^^^^^^^^^ THE UNIVERSITY OF KANSAS SCIENCE BULLETIN S s § s A COMPARATIVE ECOLOGICAL STUDY OF THE WOOD COCKROACHES IN NORTHEASTERN KANSAS s S 8 V. V. By Robert E. Gorton, Jr. W Vol. 52, No. 2, pp. 21-30 December 1, 1980 ANNOUNCEMENT The University of Kansas Science Bulletin (continuation of the Kansas Uni- versity Quarterly) is an outlet for scholarly scientific investigations carried out at the University of Kansas or by University faculty and students. Since its incep- tion, volumes of the Bulletin have been variously issued as single bound volumes, as two or three multi-paper parts or as series of individual papers. Issuance is at irregular intervals, with each volume prior to volume 50 approximately 1000 pages in length. Effective with volume 50, page size has been enlarged, reducing the length of each volume to about 750 pages. The supply of all volumes of the Kansas University Quarterly is now ex- hausted. However, most volumes of the University of Kansas Science Bulletin are still available and are offered, in exchange for similar publications, to learned societies, colleges and universities and other institutions, or may be purchased at $20.00 per volume. Where some of these volumes were issued in parts, individual parts are priced at the rate of 2 cents per page. Current policy, initiated with volume 46, is to issue individual papers as published. Such separata may be purchased individually at the rate of 3 cents per page, with a minimum charge of $1.00 per separate. Subscriptions for forthcoming volumes may be entered at the rate of $20.00 per volume. All communications regarding exchanges, sales and subscriptions should be addressed to the Exchange Librarian, University OF Kansas Libraries, Lawrence, Kansas 66045. Reprints of individual papers for personal use by investigators are available gratis for most recent and many older issues of the Bulletin. Such requests should be directed to the author. The International Standard Serial Number of this publication is US ISSN 0022-8850. Editor William L. Bloom Editorial Board Philip W. Hedrick Rudolf Jander Harvey Lillywhite Charles D. Michener Norman A. Slade Henry D. Stone George W. Byers, Chairman THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Vol. 52, No. 2, pp. 21-30 December 1, 1980 A Comparative Ecological Study of the Wood Cockroaches in Northeastern Kansas Robert E. Gorton, Jr. Department of Entomology, University of Kansas, Lawrence, Kansas 66045 CONTENTS Abstract 21 Introduction 2 1 Materials and Methods 22 Observations and Results 23 Habitat preferences 23 Vertical stratification 23 Species diversity and evenness of distribution 24 Feeding behavior 25 Predators 25 Interactions 25 Seasonal population cycles 25 Discussion 25 General patterns of species distribution 25 General patterns of species diversity 27 Competition, abundance, and community structure 29 Literature Cited 30 Acknowledgments 30 Abstract A field study of the ecology and natural history of the native cockroaches in Northeastern Kansas was under- taken in the vicinity of Lawrence, Kansas. Such parameters as vertical and horizontal distributions, relative abundances, food preferences, and population peaks were identified for each of the six species in the genera Parcoblatta and Ischnoptera, comprising the cockroach community in this area. The results are explained as mechanisms for co-existence and other adaptations to the biotic and abiotic environments encountered by the insects. Introduction Although the wood cockroaches in the genera Parcoblatta and Ischnoptera are common insects in much of the United States, the information available concern- ing the species in these genera is sparse and anecdotal (Gould and Deay, 1938, 1940; Rau, 1940). Blatchley (1920) gave general habitat descriptions for the various species, but most of his information was gathered in a nonrigorous manner. Law- son (1967) provided habitat descriptions of wood cockroaches in Kansas from trap data, but without quantification. No sys- tematic effort has been made to carefully examine the distribution, relative abun- dances, population dynamics, and general natural history of the several commonly sympatric species of wood cockroaches comprising a typical temperate cockroach community (but see Friauf, 1953; Dreisig, 1971). This study focused on si.x species of cockroach found in Douglas and Jefferson counties, Kansas: Parcoblatta pennsyl- vanica, P. lata. P. iihleriana, P. virginica, P. bolliana, and Ischnoptera deropelti- formis. The major objectives were to 22 The University of Kansas Science Bulletin elucidate the above-mentioned parameters, observe the behavior of the animals during their nocturnal activity periods, determine the community structure, and observe as many intraspecific and interspecific inter- actions as possible in a natural context. Materials and Methods The study areas were within the Uni- versity of Kansas Natural History Reser- vation and the Nelson Environmental Study Area, approximately 15 km north of Lawrence, Kansas, These areas repre- sent several discrete habitat types (Fig. 1) : forest — representative flora being basswood (Tilia americana), black walnut (Jttglans nigra), and red and bur oak (Qiiercus rubra and Q. macrocarpa, respectively), with an understory of poison ivy (Toxi- codendron radicans), rough-leaved dog- wood (Corntis drummondii), Buckbrush (Symphoricarpos orbicidattis), and big bluestem ( Andropogon gerardi); an eco- tone of thickets and shrubland — typical flora being red elm (Uhntis rubra), smooth sumac (Rhus glabra), white ash (Fraxinus americana), and bluestem (Andropogon spp.) ; and grassland — dominated by such plants as smooth brome (BromifS iner- mus), common milkweed (Asclepias syri- aca), switch grass (Panicuni virgatum), and tall goldenrod (Solidago altissima) (Martinko, 1976; Birdsell and Hamrick, 1978; J. L. Hamrick, pers. comm.). Observations were made at night from May to September, 1977 and 1978. Census- ing was done along approximately the same route each time for 3-5 h (Fig. 1). The time was about equally divided be- tween forest and nonforest areas. If less than 66.67% of the individuals of a given species were found in a general habitat (forest or nonforest), that species was con- sidered to have no particular habitat pref- erence. Most observations (>98%) were made at the Nelson Area. Each cockroach observed in the field Figure 1. Map of the primary study area at the Nelson Environmental Study Area. Legend: 1-2, forest; 3-4, thickets and shrublands (mostly woody plants less than 5 m tall) ; 4-6, grassland. Arrowed line denotes approximate path taken for observations. Scale: 1 cm =: 50 m. was identified as to sex and species, with- out disturbing the animal if possible. Sub- strate, vertical height, habitat, and activity were recorded for each sighting. Animals attracted to lights or collected nearby but well outside the study areas were counted in the seasonal abundance calculations only. Differences in intraspecific intersexual vertical distributions and shifts in these distributions as a function of habitat change were determined with the Mann- Whitney L^-test (Conover, 1971). Inter- specific comparisons of the vertical distri- butions of each sex were made with an ANOVA and an SNK test (Sokal and Rohlf, 1969). The Shannon-Weiner Index was used to measure species diversity be- tween forest and nonforest areas for both Ecological Study of Cockro.\ches 23 sexes and between the various major sub- strates (e.g., ground, grass, etc.) (Pielou, 1969). This index is given as: H' = -2Pj In Pi where S is the number of species in the assemblage and Pi is the rehitive abun- dance of the /th species. Evenness was calculated by /' = //'/In S, where H' is the Shannon-Wiener Index of species di- versity and S is the number of species present (Pielou, 1969). Observations and Results Habitat Preferences From the criterion for assigning gen- eral habitat preferences to a species Parco- blatta lata and Ischnoptera deropeltijormis were considered species of prairie and dis- turbed grassland, while P. hoUiana, P. vir- ginica, and P. iihleriana were considered forest species. Parcoblatta pennsylvanica was found in both general habitats, pri- marily on trees (Table 1). None of these species (especially females), however, was restricted to a particular habitat. In the disturbed areas, females were primarily foiuid moving about on the ground, while males were usually seen on or flying to blades of grass (Fig. 2). In the forest, temales of P. iihleriana and P. virginica were usually observed on the ground and the rare P. holliana females were found either on the ground or on forbes and bushes. Males of P. virginica were ob- served primarily on bushes and trees; males of P. bolliana almost entirely on bushes. Parcoblatta iihleriana males were primarily found on bushes, but also com- monly frequented the grass and ground. Males of this species were the most general in substrate choice. Parcoblatta pennsyl- vanica males were usually seen on the trunks of the larger trees, while the fe- males of this species were found on the ground as well as on trees. Vertical Stratification In five of the six species studied, males occurred at significantly higher points in the environment than females (Table 2). The exception, P. bolliana, showed no dif- ferences in mean height preferences be- Taele 1. Numbers and percentages of individuals of eacii species found in forest and nonforest habitats. Males Females Total Males Females Total Males 1 7 (12.5%) (87.5%) Females 1 5 (16.7%) (83.3%) Total 2 12 (14.3%) (85.7%) P. I 'ata P. virginica Nonforest Forest Total Nonforest 9 Forest Total 47 15 62 25 34 (75.8%) 31 (24.2%) 17 48 (26.5%) 4 (73.5%) 36 40 (64.6%) 78 (35.4%) 32 110 (10.0%) 13 (90.0%) 61 74 (70.9%) (29.1%) (17.6%) (82.4%) /. deropeltijormis P. pennsylvanica Nonforest Forest Total Nonforest Forest Total 46 10 56 41 33 74 (82.1%) 8 (17.9%) 5 13 (55.4%) 69 (44.6%) 49 118 (61.5%) 54 (38.5%) 15 69 (58.5%) 110 (41.5%) 82 192 (78.3%) (21.7%) (57.3%) (42.7%) P. bolliana P. uhlet •iana Nonforest Forest Total Nonforest Forest Total 8 22 37 59 (37.3%) (62.7%) 6 38 94 132 (28.8%) (71.2%) H 60 131 191 (31.4%) (68.6%) 24 The University of Kansas Science Bulletin tvveen the sexes. Parcoblatta pennsylvanica 13.3 cm: 20.32-33.02 cm (X=27.86±1.87; males were significantly higher up on the N=6). vegetation than all other males (ANOVA, F=316.53, P<.001; SNK test, P<.05), Species Diversity a^id Evenness and P. Pennsylvania females were signif- °/ Distribution icantly higher than all other females but When the species composition of forest those of P. bolliana (ANOVA, F=:39.11, and nonforest areas were compared, the F<.001; SNK test, P<.05). No other species diversity and evenness indices were differences in intrasexual mean height identical (Table 4). The male community, preferences were found for either sex. however, showed greater diversity in the The average height chosen by one or nonforest areas and the female community another of the sexes changed with general showed greater diversity in the forest habitat (forest or nonforest) in four of the areas. Comparisons by vegetation type re- six species studied (Table 3). In each case, vealed low species diversity on trees and forest "populations" were higher off the buildings and high diversity on forbs and ground than nonforest "populations." bushes. Males reflected this same general I saw ten copulating pairs during the trend. The female communities, on the study. The range of heights at which P. other hand, were most diverse on the pennsylvanica copulated was large: 53.34- ground and least diverse on trees and 365.76 cm (X=188.59±:71.97; A^=4). The buildings. The low diversity on trees and pooled heights of copulating pairs of the buildings was due to the preference of P. remaining five species ranged over only /7(°/;/7i-y/z/fl/7/cfl for these substrates (Fig. 2). Table 2. Mean heights (cm) at which males and females were found, with one standard error around the means. Heights were compared with the Mann-Whitney [/-test. Species Males Range Females Range U^ a-lcvel P. bolliana 59.36±17.51 0-121.92 67.72 ±52.96 0-304.80 33 >.10 P. uhleriana 62.06±11.71 0-609.60 7.02±2.35 0-187.96 7330 .0001 P. virginica 70.95±10.96 0-274.32 13.84±5.38 0-137.16 1240.5 .0001 P. lata 63.68±4.62 0-152.40 18.78±8.76 0-365.76 2790 .0001 P. pennsylvanica 120.42±13.02 0-548.64 92.56±10.59 0-609.60 6460 .0001 /■ dcropcltijonnis 51.02±4.29 0-150.00 1.92±2.09 0-25.00 717.5 .0001 Table 3. Differences in mean height preferences between forest antl nonforest areas (cm±l SE), compared with the Mann-Whitney t/-test. Only those species showing significant differences are given. Species Sex Height: forest Height: nonforest U^ a-level P. virginica Female 14.00±5.79 0.00±0.00 108 05 P. lata Female 45.72±23.89 5.31±4.05 375 0.01 P. uhleriana Female 7.12±2.96 4.51±3.09 3307.5 0.0001 P. uhleriana Male 70.96±17.9I 42.29±9.25 488 0.05 /. deropehiformis Male 59.13±16.95 49.37±7.37 311.5 0.05 Table 4. Species diversity and evenness indices of the cockroach community calculated by general habitat (for- est and nonforest) and substrate (grass, ground, etc.), with 1 SD. Habitat H' (bcjth sexes) / (both sexes) H' (male) / (male) H' (female) / (female) Forest 1.49±.04 ^83 1.51 ±.04 ^5 1.61±.05 ^90 Nonforest 1.49±.03 .83 1.64±.04 .92 1.39±.05 .78 Ground 1.43±.04 .80 1.46±.12 .81 1.76±.05 .98 Grass 1.27±.07 .71 1.27±.07 .71 1.10±.01 .61 Forbs and bushes 1.71±.03 .95 1.68±.05 .94 ].47±.ll .82 Trees and buildings .... 0.90±.08 .50 1.03±.ll .57 0.76±.20 .42 Ecological Study of Cockroaches 25 Feeding Behavior Of 45 nocturnal observations of adults feeding, 22 fed on mushrooms (both sexes of P. pennsylvanica and P. virginica, and females of P. uhleriana). Both sexes of P. pennsylvanica fed on mammalian, prob- ably coyote, fecal matter (Ar=8), and a P. uhleriana female ate bird feces. Parco- hlatta lata males and both sexes of P. penn- sylvanica fed on sap from wounded trees (N^S), and a /'. lata female ate the cam- bium layer from a wood chip. A P. penn- sylvanica male consumed spittle from a cercopid nymph, and a P. lata female ate flower petals. A P. uhleriana male ate moss off of a tree trunk and a female of the same species fed on mammalian carti- lage. The most unusual feeding observa- tion was that of a P. pennsylvanica female attacking a weakened but upright and mobile Camponotits pennsylvaniciis work- er. The cockroach repeatedly bit the ant, sometimes picking it up and carrying it a short distance. The roach left and returned to the ant several times before moving away from the still living animal the final time. Rau (1940) noted an instance of predation by a female P. pennsylvanica on a Polistes larva. Predators Nine acts of predation on wood cock- roaches were recorded, all but one on males. The predator was invariably a spider of the Thomisidae, Araneidae, or Agelenidae. On two occasions, mice (Peromyscits lencopits) were seen on a tree trunk with many cockroaches on it, possibly preying on them. Interactions Five P. uhleriana females feeding on a mushroom engaged in biting, kicking, lunging toward, and posturing when one contacted another. A P. pennsylvanica male pursued and drove off a P. lata male from flowing sap in a tree wound. An- other P. pennsylvanica male drove off a P. uhleriana male that had antennated it. Once, a P. pennsylvanica male was forced away from a tree wound by foraging ants. Seasonal Population Cycles Adult male populations appeared at about the same time as adult female pop- ulations, but they were nearly extinct long before the female populations (Fig. 3). Population peaks for the two most com- mon species, P. pennsylvanica and P. uhleriana, were distinct for both sexes. The bi modality of /-*. bolliana and /. de- ropeltiformis females was probably due to sampling error in rare populations. Discussion General Patterns of Species Distribution Mobile animals arrange themselves in space by preferentially occupying particu- lar parts of the environment. The pattern of distribution for a species is determined by such factors as the number of compet- ing species encountered in a given area, the population densities of those species, availability of resources (food, refugia, etc.), and the number of predators, both absolute and relative. Insect populations may also have subtle temperature and hu- midity requirements for optimal growth rates (Edney et al., 1978). The component species of the cock- roach community in question showed con- siderable geographic variation in habitat choice. Lawson (1967) maintained that P. bolliana is "strictly a grassland species in Kansas," but 85.7°o of the individuals I censused were found in the forest. The small numbers of this species observed, however, suggest that the area is subopti- mal for this species. Both Lawson (1967) and Blatchley (1920) describe /. deropelti- jormis and P. lata as forest species, but the demes I studied were primarily non- forest populations. I concur with both authors that P. virginica and P. uhleriana are forest species. Parcoblatta virginica was the most habitat-specific of the five 26 The University of Kansas Science Bulletin 50 0 zz P lata ^ .£5L J/) §100 C o S 50 E 3 0 ^ 0 p. uhleriana a cz 50 r 0 Ground IZL P. virginica t^ Z3 Grass Forbs & Bushes Trees Human Structures Figure 2 . Bar graphs of substrates upon whicli cockro;Klus were fouml. Numbers of females arc represented by open bars, males by hatched bars. Ecological Study of Cockroaches 27 50 0 P bolliana o I 50 c E 23 P. pennsylvanica a EZl 50 r 0 za I I. deropeltiformis FA Ground Grass Forbs & Bushes Trees Human Structures most common species, suggestmg a species closely adapted to the forest environment. Parcoblatta ithleriana was the most abun- dant of the forest species, due either to higher fecundity or survival. Perhaps this species affects resource availability, thereby limiting forest populations of the other species. Parcoblatta pennsylvanica is a habitat generalist and a patch specialist. occurring wherever trees were found. General Patterns of Species Diversity Species diversity and evenness are char- acters of a community. These concepts derive from the fact that individuals are distributed among species differently in different parts of the environment. In the cockroach community, the same assem- blage of species occurs in several different 28 The University of Kansas Science Bulletin CD E 40 - 15-29 30 May- 13-26 27 June- May 12 June June 10 July II July 8-21 7 August August Figure 3. Seasonal population fluctuations of cockroaches. Above: females, Below: males. P. boUiana ▲--▲ P. lata P. pennsyhanica ZW^ P. tihlciiana P. virgimca I. dcropcltijortnis |—)_r-| oo Ecological Study of Cockroaches 29 combinations as a function of habitat and patch types (Tabic 4). Implicit in these measurements is the assumption that while individuals undoubtedly move freely from, say, the grass to a bush, the average over- all species composition on all grasses and bushes is constant over time. (Also im- plicit, of course, is the assumption that the environmental partitions made by the in- vestigator reflect those made by the organ- isms. Familiarity with the species assem- blage investigated minimizes this source of error.) One of the most remarkable results of this analysis is the perfect correlation be- tween the species diversity found in the forest and nonforest habitats. A second very high correlation is that of the species diversity found on various patches when only species (not sexes) were considered and that found for the same patches for males. Males are, therefore, a good pre- dictor of general species diversity in this community. Patterns of diversity vary in a systematic way between male and female components. These patterns differ by hab- itat and patch of maximum diversity, demonstrating preference for these param- eters by species and sex. Competition, Abundance, and Community Structure Competition is a fundamental concept in ecology, yet ticklishly difficult to dem- onstrate directly, i.e., to actually see com- petition taking place at the population level (but see Roubik, 1978). Because the evidence for competition is often circum- stantial, it is sometimes incorrectly invoked to explain population parameters (Wiens, 1977). Nevertheless, I believe there are several lines of circumstantial evidence implicating competition as an important determinant of cockroach community structure. In the presence of competition, natural selection should operate to perpetuate those phenotypes displaying characters fa- cilitating co-existence, resulting in non- random species arrangements in time and space. In the cockroach community, some species were much more abundant than others. The two commonest species, P. pennsylvanica and P. uhleriana, tended to occur in dilTcrcnt parts of the environment and their population peaks (Fig. 3) were temporally staggered. Parcoblatta penn- sylvanica was also the topmost species ver- tically and the most patch-specific species (Fig. 2). While patch choice did not dif- fer for any species between forest and nonforest areas, there were five instances of different vertical distributions of at least one sex of a species as a function of gen- eral habitat (niche shifting). Females out- lived males in each species, and intrasexual adult populations tended toward simul- taneous extinction. In 2/3 of the species in this community, males were significantly higher on the vegetation than females. Finally, species showed overlap in food choices and some instances of direct be- havorial interference. These similarities in food preferences and life expectancies and the differences in vertical and horizontal spatial distributions argue for intersexual, interspecific, and probably inter-lifestage competition as a factor influencing com- munity structure. The nymphs were impossible to dis- tinguish to species in the field, so they could not be included in this study. They did, however, become increasingly abun- dant throughout the spring and summer until they were the only life stage ob- served. Nymphs were seen in the same places adults were observed, both at night and during the day when they were in- active, i.e., moving about on the vegetation and on the ground, and hiding under bark, manure, rocks, boards, and in and under rotting logs. Generally, the adult cockroach commu- nity seems to exist at two distinct levels: females on or near the siround and males 30 The University of Kansas Science Bulletin above them on the vegetation, with copu- lating pairs in between. Parcoblattu penn- sylvanica represents a third level above and rather isolated from the other two. Individuals at each level probably interact much more with each other than with those on other levels. The role of the nymphs in the community is difficult to assess, but undoubtedly has some impact. Acknowledgments I thank C. D. Michener, K. B. Armi- tage, and especially W. J. Bell for constant encouragement and support. I also thank P. G. Decelles and L. M. Roth for review- ing the manuscript. Costs were defrayed by a Beamer Award for 1977 and again in 1978. This paper is part of a Doctor of Philosophy dissertation in Entomology from the University of Kansas. Literature Cited BiRDSELL, R., and J. L. Hamrick. 1978. The effect of slope-aspect on the coinposit'.on and density of Oak-Hickory forest in eastern Kansas. Univ. Kansas Sci. Bull. 51:565-573. Blatchley, W. S. 1920. Orthoptera of North-east- ern America. Indianapolis. CoNOVER, W. J. 1971. Practical Non-parametric Sta- tistics. John Wiley, N.Y. Dreisig, H. 1971. Diurnal activity in the Dusky Cockroach, Ectobitis Japponiais L. (Blattodea). Entomol. Scand. 2:132-138. Edney, E. B., p. Fraxco, and R. Wood. 1978. The responses of Arenivaga irivestigata (Dictyoptera) to gradients of temperature and humidity in sands studied by tagging with Technetium 99m. Physiol. Zool. 51:241-255. Friauf, J. J. 1953. An ecological study of the Dermaptera and Orthoptera of the Welaka area in northern Florida. Ecol. Monogr. 23:79-126. Gould, G. E., and H. O. Deay. 1938. Notes on the bionomics of roaches inhabiting houses. Proc. Indiana Acad. Sci. 47:281-284. . 1940. The biology of six species of cock- roaches which inhabit buildings. Bull. No. 451. Purdue Univ. Ag. Expt. Station. 31 p. Lawson, F. a. 1967. Ecological and collecting notes on eight species of ParcoUatta (Orthoptera: Blattidae) and certain other cockroaches. J. Kan- sas Entomol. Soc. 40:267-269. Martinkc, E. a. 1976. Seasonal changes in di- versity of three adjacent insect communities. Ph.D. Dissertation. Univ. of Kansas, Lawrence. PiELOu, E. C. 1969. An Introduction to Mathe- matical Ecology. John Wiley, N.Y. Rau, p. 1940. The life history of the wood-roach, ParcobJatta pennsylvanica De Gcer (Orthoptera: Blattidae). Entomol. News 51:4-9, 33-35. RouBiK, D. W. 1978. Competitive interactions be- tween neotropical pollinators and Africanized honey bees. Science 201:1030-1032. SoKAi,, R. R., and F. J. Rohlf. 1969. Biometry. W. H. Freeman, San Francisco. WiExs, J. 1977. On competition and variable en- vironments. Amer. Sci. 65:590-597. •.•-•.•.•.•.• MUS. COMP. ZOOl_. LIBRARY ^^:^♦x«•:•^^^:r^:^?^^^:♦x♦:•^^%^^^ i^ THE UNIVERSITY OF KANS'A^^^^'^^ SCIENCE BULLETIN I S s s ft: :!: TWO NEW SPECIES OF INQUILINOUS MONOMORIUM FROM NORTH AMERICA (HYMENOPTERA: FORMICIDAE) By Mark B. DuBois g S g i Vol. 52, No. 3, pp. 31-37 June 12, 1981 ANNOUNCEMENT The University of Kansas Science Bulletin (continuation of the Kansas Uni- versity Quarterly) is an outlet for scholarly scientific investigations carried out at the University of Kansas or by University faculty and students. Since its incep- tion, volumes of the Bulletin have been variously issued as single bound volumes, as two or three multi-paper parts or as series of individual papers. Issuance is at irregular intervals, with each volume prior to volume 50 approximately 1000 pages in length. Effective with volume 50, page size has been enlarged, reducing the length of each volume to about 750 pages. The supply of all volumes of the Kansas University Quarterly is now ex- hausted. However, most volumes of the University of Kansas Science Bulletin are still available and are offered, in exchange for similar publications, to learned societies, colleges and universities and other institutions, or may be purchased at $20.00 per volume. Where some of these volumes were issued in parts, individual parts are priced at the rate of 2 cents per page. Current policy, initiated with volume 46, is to issue individual papers as published. Such separata may be purchased individually at the rate of 3 cents per page, with a minimum charge of $1.00 per separate. Subscriptions for forthcoming volumes may be entered at the rate of $20.00 per volume. All communications regarding exchanges, sales and subscriptions should be addressed to the Exchange Librarian, University OF Kansas Libraries, Lawrence, Kansas 66045. Reprints of individual papers for personal use by investigators are available gratis for most recent and many older issues of the Bulletin. Such requests should be directed to the author. The International Standard Serial Number of this publication is US ISSN 0022-8850. Editor William L. Bloom Editorial Board Philip W. Hedrick Rudolf Jander Harvey Lillywhite Charles D. Michener Norman A. Slade Henry D. Stone George W. Byers, Chairman THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Vol. 52, No. 3, pp. 31-37 June 12, 1981 Two New Species of Inquilinoiis Mojiomorittm from North America (Hymenoptera : Formicidae) Mark B. DuBois Department of F.ntdtnolfiuy, Univcrsit\' of Kan'^as, T^avvrcncc, Kansas 66045 Contents Abstract 31 Introduction 31 Afonomorium talhotac new species 31 Monomoriiim inqtiiliniim new species 34 Monomoritiryi (z=Epoeciis) pergandci Icctotype designation 36 Key to North American inquilinous Mononwritim 36 acknowledcmfnts 37 Literature Cited 37 Abstract The ant, Monomorium talbotae, new species, is described from Michigan. This species is compared briefly with Monomorium pcrgandei. Dealate and alatc females and males were found in three colonies of a host species which is the M. minimum of Talbot (1975). This new ant is apparently a workerless, permanent social parasite. Similarly, another workerless, permanent social parasite, M. inquilinum, is described from Mexico and compared with M. pergandei and M. talbotae. One female was discovered in a nest of what is, or is closely related to, M. cyaneum. Both inquilinous species are compared with their hosts and a key is provided for the separation of the inquilinous Monomorium. The taxonomic relationships between hosts and inquilines are also discussed. A lectotype is designated for M. pergandei. Introduction Although several species of parasitic Monomorium have been described from around the world, only two species have been described as inquilines of Mono- morium (minutitm group) in North America. Since Ettcrshank (1966) synony- mized M. metoecus (Brown and Wilson, 1957) with M. minimum because of the existence of a worker caste (Wilson and Brown, 1958), M. pergandei (Emery, 1892, described further 1895) has been the only known inquilinous Monomorium in North America. Recently, I received specimens of two new species of inquilinous Monomorium: M. talbotae from Michigan (courtesy of Dr. Mary Talbot) and M. inquilinum from Mexico (courtesy of Ms. Margaret Thayer at the Museum of Comparative Zoolo<:v). Monomorium talbotae is known from the holotype, allotype, and eight paratypcs. Monomorium inquilinum is known only from the holotype. Monomorium talbotae New Species Holotype female and allotype male. — MICHIGAN: Livingston Co.; E. S. 32 The University of Kansas Science Bulletin George Reserve; 30 June 1966; M. Talbot (66-13). Holotype and allotype are de- posited in the Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts. Holotype lacks left fore- wing and right hindvving. Paratypes are in the following collections: M. B. DuBois personal collection (2 fem.ales) ; Snow Entomological Museum, University of Kansas, Lawrence (1 female); M. Talbot personal collection (4 females, 1 male). Fefjiale Diagnosis. — Similar to M. pergandei in general morphology and lack of a worker caste, but differing in the following fea- tures: clypeus scarcely emarginate, lack- ing teeth; propodeum, petiole and post- petiole as in Figure 1; no depression on first gastral tergum. Similar to host in general morphology but differing in the following features: worker caste lacking; clypeus scarcely emarginate; mandible with two teeth; length less than 3 mm. Description. — Habitus, in general, like queens of other Monomorium (Fig. 1). Total length including mandibles 2.28 mm in alcohol. HEAD: Length 0.45 mm; width 0.43 mm; surface smooth and shin- ing with pits (representing insertions of setae) prominent; mandible with two teeth; maxillary palp 1-segmented; labial palp 1-segmented; clypeus scarcely emargi- nate, lacking teeth; antenna 12-segmented with a weakly developed 3-segmented club. Head and mandibles light brown, anten- nae light yellow. ALITRUNK: Length 0.68 mm; pronotal width 0.33 mm; surface smooth and shining with pits prominent on pronotum; propodeal spines absent. Alitrunk light brown, legs light brown except tibiae and tarsi which are light yellow. Wings as in Figure 1. PETIOLE: Length 0.21 mm; width 0.16 mm; height 0.23 mm; pedunculate; dorsal surface of node evenly convex with no trace of emargi nation; both anterior and posterior surfaces of node evenly covered with erect setae of uniform height; sides of petiole smooth. Color light brown. POST- PETIOLE: Length 0.13 mm; width 0.23 mm; height O.IS mm; dorsal surface of node evenly convex with no trace of emargination; sides of postpetiole punctate. Color light brown. CASTER: No de- pression on first gastral tergum; surface of gaster smooth and shining with pits visible on first tergum. Color light brown. Wo^-}{er Unknown and apparently non-existent, as in M. pergandei. Male Description. — Habitus, in general, like males of other Monomorium. Total length including mandibles 2.18 mm in alcohol. HEAD: Length 0.43 mm; width 0.43 mm; antenna with 12 segments; sculpture, palps and color as in holotype. Mandible re- duced, similar to female. ALITRUNK: Length 0.68 mm; pronotal width 0.29 mm; sculpture, propodeum, and color as in holotype. Wings as in Figure 1. PETIOLE: Length 0.20 mm; width 0.15 mm; height 0.16 mm; pedunculate; dorsal surface, pilosity, sculpture and color as in holotype. POSTPETIOLE: Length 0.13 mm; width 0.23 mm; height 0.15 mm; pilosity, sculpture and color as in holotype. Variation The following measurements, in milli- meters, were taken from the holotype, seven paratype females and one paratype male. All measurements were made in alcohol. Variation within each character measured lor the females is expressed as the mean (range). The actual measure- ments for the male specimen are given in Two New Species of Tnquilinous Monomorium from North America 33 brackets following the measurements of the females. HEAD: Length 0.44 (0.40- 0.4S) [0.39]; width 0.40 (0.39-0.42) [0.42]. ALITRUNK: Length 0.68 (0.63-0.72) [0.63 1 ; pronotal width 0.32 (0.27-0.33) |0.30|. PETIOLE: Length 0.18 (0.10-0.21) 1 0.16]; width 0.16 (0.15-0.18) [0.18]; hc\o\M 0.20 (0.18-0.23 [0.19]. POST- PETIOLE: Length 0.11 (0.09-0.14) [0.10]; width 0.20 (0.19-0.23) [0.21]; height 0.16 (0.15-0.21) [0.15 1 . TOTAL LENGTH: 2.32 (2.10-2.55) [2.33]. Derivatio nominis This species is named after Dr. Mary Talbot, who discovered it. Habitat From notes by M. Talbot: "A high, dry held where vegetation was not dense and small patches of bare sandy soil were numerous. Grassess were mostly Poa compressa and Aristida sp. A variety of scattered forbs included Lespedeza capitata, Liatris aspersa, Riimex acetosella, and Soli dago spp. Patches of Polytrichitm pilifenim moss were large and frequent, and the red-tipped lichen, Cladonia crista- lella var. vestita, was characteristic." Comparison of M. talbotae ifit/i its Host Wilson (1971, p. 374-5) cites a set of morphological and behavioral character- istics which are found in some combina- tion in all species of inquilinous ants. Several of these traits can be recognized in M. talbotae (numbers of traits corre- spond to those of Wilson) : 1. worker caste is lost; 4. female and male are reduced in size; 8. although the arrangement of veins is similar to that of the host, many of these "veins" are represented as the absence of microtrichia (dashed lines in figures), in- dicating a reduction in venation; 9. mouth- parts are reduced, with the mandible losing two teeth and the labial and maxillary palps losing one segment each; 13. petiole and postpetiole are thickened; 14. a spine is formed on the ventral surface of the post- petiole; and 16. cuticular sculpturing is re- duced. One additional character which is probably associated with small size is the reduction in size and complexity of the male genitalia. In s[iitc of the above modifications, M. talbotae appears to be closely related to its host for the following reasons. 1. Although the wing veins are reduced in M. talbotae, their placement is nearly identical with the placement of veins in the host's wings. 2. The reduced male genitalia are more similar to those of the host than to any other North American Monomorium. The structure of sternum VIII is correspond- ingly similar. Sternum IX is also similar, although it has been reduced to such an extent that it resembles the corresponding structure in most North American Mon- omorinm. 3. The metanotum is present as a small bump in side view (Fig. 1). This is true for both M. talbotae and its host. However, the metanotum is shaped differently in M. inquilinitm (Fig. 2). 4. The propodeum is of a similar rounded shape in both A/, talbotae and its host (Fig, 1). This can be compared with the flattened outline of the propodeum in M. inqiiihniim (Fig. 2). 5. The suture across the mesothorax is evenly curved and slightly thickened at the posterior end in both M. talbotae and its host (Fig. 1). 6. The pilosity of M. talbotae is more similar to its hcjst than to M. inqiiilinum. Measurements of host queen. — The fol- lowing measurements were taken trom a queen of the M. minimum oi Talbot (1975) collected from the E. S. George Reserve. Total length including mandibles 5.20 mm in alcohol. HEAD: Length 0.75 mm; width 0.8 mm. ALITRUNK: Length 1.45 mm; pronotal width 0.7 mm. PETIOLE: Length 0.50 mm; width 0.30 34 The University of Kansas Science Bulletin mm; height 0.38 mm. POSTPETIOLE: Length 0.25 mm; width 0.38 mm; height 033 mm. Measurements of host male. — The fol- lowing measurements were taken from a male of the M. minimimi of Talbot (1975) collected from the E. S. George Reserve. Total length including mandibles 4.86 mm in alcohol. HEAD: Length 0.72 mm; width 0.81 mm. ALITRUNK: Length 1.59 mm; pronotal width 0.78 mm. PETIOLE: Length 0.51 mm; width 0.27 mm; height 0.36 mm. POSTPETIOLE: Length 0.21 mm; width 0.31 mm; height 0.33 mm. Monomonitm inqiiilinum New Species Holotype label data. — Estado de Mexico : Highway 57 (between Mexico City and Queretaro), km 127 (measured from Mex- ico City); 9 August 1965; High Desert; Cornell University Mexico Field Party. W. L. Brown (pers. comm.) indicated that his field notes record the locality as 83 km south of Queretaro. Holotype is de- posited in the Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts. Wings lacking on holotype although specimen appears to have pos- sessed them when younger. Figure 1. Monomoiinm talhotae, female. A. Side view showinj,' oiiiline ami major sutures (legs omitted). Stippling indicates membranous areas. B. Wings. Stippling indicates most heavily sclerotized veins. Dashed lines indicate veins represented by lack of microtrichia. Both A and R drawn to same scale. Line equals 1 mm. Two New Species of Inquilinous Monomorium from North America 35 Female Diagnosis. — Similar in general morphol- ogy to M. pergandei but differing in the following features: clypeus emarginate but lacking distinct carinae or teeth; anterior edge of pronotum rounded in side view vs. angular projecting edge in M. pergandei (Figs. 2 and 3) ; dorsum of petiolar node emarginate. Similar to M. talbotae but differing in the following features: first gastral tergum with slight basal depres- sion; metanotum with outline as in Figure 2; dorsum of both petiole and postpetiole emarginate; outline of propodeum as in Figure 2; and dorsum of alitrunk with pilosity of uniform distribution and length. Description. — Habitus similar to that of M. pergandei (Fig. 2). Total length in- cluding mandibles 2.16 mm dry. HEAD: Length 0.45 mm; width 0.39 mm; surface smooth and shining; mandible with two teeth; maxillary palp 1-segmented; labial palp 2-segmented; clypeus emarginate but lacking distinct teeth; frontal groove ex- tends to level of anterior ocellus; antenna 12-segmented with weakly developed 3- segmented club. Head and mandibles dark brown, antennae yellow. ALI- TRUNK: Length 0.66 mm; pronotal width 0.30 mm; surface smooth and shin- ing; propodeal spines absent. Alitrunk dark brown, tibia and tarsi yellow. PETIOLE: Length 0.20 mm; width 0.16 mm; height 0.21 mm; pedunculate; dorsal surface of node emarginate; both anterior and posterior surfaces evenly covered with erect setae; sides of petiole smooth to weakly punctate. Q)lor brown. POST- PETIOLE: Length 0.09 mm; width 0.24 mm; height 0.18 mm; dorsal surface of node emarginate; sides of postpetiole punctate. Color brown. G ASTER: Small depression at base of first gastral tergum; surface of gaster smooth and shining. Color brown. Derivatio nominis This name is used in reference to the inquilinous way of life of this ant as in- dicated by the association with M. cyaneum and the lack of a worker caste. Habitat From notes by V/. L. Brown, Jr.: This specimen came from a populous colony of M. cyaneum (?) under a rock in plateau scmidesert with low shrubs. Comparison of M. inquilinum with its Host Monomorium incjuiliniim has many of the morphological characters found in inquilinous ants (Wilson, 1971) which were discussed with the previous species. Since the holotype lacks wings, it is un- known whether the wing venation is re- duced. In spite of these modifications, M. in- quilinum appears to be closely related to its host for the following reasons (num- bers correspond to previous comparison of M. talbotae with its host). 3. The outline of the metanotum, in side view, is similar between M. inquilinum and its host. 4. The shape of the propodeum is similar between these two species. 6. The pilosity of both species is similar. One difference between M. inquilinum and its host is that M. inquilinum presum- ably possessed wings as a young female while M. cyaneum (?) queens did not (based on the fusion of sclcrites). This is difficult to interpret since there is such a small sample of queens of both species from this locality. Measure?nents of host queen. — The fol- lowing measurements were taken from a queen of what is, or is closely related to, M. cyaneum collected at the same locality as M. inquilinum. Total length including mandibles 4.7 mm dry. HEAD: Length 36 The University of Kansas Science Bulleti N 0.85 mm; width 0.75 mm. ALITRUNK: Length 1.30 mm; pronotal width 0.55 mm. PETIOLE: Length 0.45 mm; width 0.35 mm; height 0.40 mm. POSTPETIOLE: Length 0.20 mm; width 0.40 mm; height 0.35 mm. Monomorium pergandei Lectotype Designation Originally described as Epoecus pergan- dei by Emery (1892, 1(S95), Ettershank (1966) placed this species in Monomoriittn. Although several museums contain syn- types of this species, the lectotype was selected from specimens in the National Museum of Natural History, since Per- gande collected this species, and most of his collection is deposited there. Lectotype female.— DlSTKlCi: OF CO- LUMBIA: Washington ;7T2/ [Next label illegible, ending in 206]/ No. 53581 U.S.N.M. Cotype/ [Handwritten label! Epoecus pergandei (type) Emery. Lectotype is in good condition possessing all legs, antennae and wings. In addition to this type, the U.S.N.M. contains four paralectotypes (1 badly damaged) and the Museum of Comparative Zoology contains two additional paralectotypes (1 badly damaged). All specimens examined are females and now possess red, handwritten labels indicating lectotype or paralectotype. Key to North American Inquilinous Monomorium The following key will provide for the separation of all three species of inc|uili- nous Monomorium occurring in North America. All these species would key to Epoecus pergandei in Creighton (1950). 1. Base of first gastral tergum with dorsal impression 2 Figure 2. Motiotnoiiimi nuiiiiliniini , female. Side view showing outline .ind major sutures (legs omitted). Stippling indicates membranous areas. Line equals 1 mm. Figure 3. Monomoritini pergandei, fem.ile. Si>K<«o>:.:«.:•;•.•Aw:w^:♦:•:•:•:•^^^^x•^^ THE UNIVERSITY OF KANSAS SCIENCE BULLETIN SOME EFFECTS OF SELENIUM, VANADIUM AND ZIRCONIUM ON THE SWIMMING RATE OF TETRAHYMENA PYRIFORMIS: A BIOASSAY STUDY a I ss By Eugene C. Bovee and Timothy L. O'Brien Vol. 52, No. 4, pp. 39-44 December 21, 1982 ANNOUNCEMENT The University of Kansas Science Bulletin (continuation of the Kansas Uni- versity Quarterly) is an outlet for scholarly scientific investigations carried out at the University of Kansas or by Universty faculty and students. Since its incep- tion, volumes of the Bulletin have been variously issued as single bound volumes, as two or three multi-paper parts or as series of individual papers. Issuance is at irregular intervals, with each volume prior to volume 50 approximately 1000 pages in length. Effective with volume 50, page size has been enlarged, reducing the length of each volume to about 750 pages. The supply of all volumes of the Kansas University Quarterly is now ex- hausted. However, most volumes of the University of Kansas Science Bulletin are still available and are offered, in exchange for similar publications, to learned societies, colleges and universities and other institutions, or may be purchased at $20.00 per volume. Where some of these volumes were issued in parts, individual parts are priced at the rate of 2 cents per page. Current policy, initiated with volume 46, is to issue individual papers as published. Such separata may be purchased individually at the rate of 3 cents per page, with a minimum charge of $1.00 per separate. Subscriptions for forthcoming volumes may be entered at the rate of $20.00 per volume. All communications regarding exchanges, sales and subscriptions should be addressed to the Exchange Librarian, University OF Kansas Libraries, Lawrence, Kansas 66045. Reprints of individual papers for personal use by investigators are available gratis for most recent and many older issues of the Bulletin. Such requests should be directed to the author. The International Standard Serial Number of this publication is US ISSN 0022-8850. Editor William L. RIoom Editorial Board Philip W. Hedrick Rudolf Jander Harvey Lillywhite Charles D. Michener Norman A. Slade Henry D. Stone George W. Bycrs, Chairman THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Vol. 52, No. 4, pp. 39-44 December 21, 1982 Some Effects of Selenium, Vanadium and Zirconium on the Swimming Rate of Tetrahymena pyriformis: a. Bioassay Study* Eugene C. Bovee and Timothy L. O'Brien^ Department of Physiology and Cell Biology, University of Kansas, Lawrence, Kansas 66045 Abstract By stroboscopic photography, the swimming rate of Tetrahymena pyriformis strain HSM was measured in aqueous solutions of each of three metallic substances: Selenium as sclenous acid (II2SEO3); vanadium as vandyl sulfate (VOS04'2HaO) ; and zirconium as zirconsl chloride (ZrOClii-GH^O). Swimming rate was meas- ured over -15 min periods, monitored each 15 min, in concentrations of each metal, singly, at 0, 5, 10, 20, and 30 and 50 ppm. Significant decrease in swimming rate was recorded for each metal at concentrations of 20 ppm or more. The method provides a useful bioassay to determine the presence of pollutants in freshwater, quickly and easily. Some recent studies (Cooley et al., 1972; Carter and Cameron, 1973; Bergquist, 1974; Bergquist and Bovee, 1976; Bovee et al., 1977; Bovee et al., 1979) indicate that measurement of the swimming rate and/or growth rate of Tetrahymena pyri- formis- can be used to bioassay the amount of a pollutant in water if the pollutant is known, or to identify the generally toxic level of pollutants in a freshwater source. Other species of Pro- tozoa may also serve as indicators of toxic levels of pollution, including Amoeba proteus, Euglena gracilis and Paramecium caiidatiim (Mills, 1973), Stentor coeruleus and Vorticella campanula (Burbanck and * Supported by Kansas Water Resources Institute Matching Grants A-058 and B-040 and, in part, by NSF Grant GB-16616. ^ Present address: Elcctronmicroscopy Center, Uni- versity of Illinois, Urbana, Illinois 61801. ° The specific identity of this strain is not known in relationship to the species of Tetrahymena named by Nanney and McCoy (Trans. Amer. Microscop. Soc. 95:664-682 [1976]). It is generally considered to be, as indicated, a strain of Tetrahymena pyriformis. Spoon, 1967), and populations of Protozoa (Lackey, 1938, 1968; Bovee, 1965; Sunder- asen et al., 1965; Bick, 1973; Cairns, 1973; Ruthven and Cairns, 1973; Small, 1973). Selenium, vanadium and zirconium are becoming important pollutants in water sources, due to their escape from industrial processes, from the burning of fossil fuels and, more recently, from their appearance in insecticides destined to replace the polychlorinated hydrocarbons and organo- phosphates. There is evidence that those metals are toxic to vertebrates, including humans, in amounts greater than 20 parts per million in water or in foodstuffs (Hathcock et al., 1964). Therefore, we measured the swimming rates of Tetrahymena pyriformis in var- ious concentrations of those metals in water, singly, to determine if such meas- urements might serve as a bioassay to detect toxic and subtoxic amounts of those metals. 40 The University of Kansas Science Bulletin Materials and Methods Tetrahymena pyrijormis (strain HSM) were grown at 22° C in 2% proteose- peptone (Difco), 0.1% sodium acetate (CH.sC00Na-3H:>0) and 0.1% sodium phosphate (Na2HP04) dissolved in Chalkley's solution (Chalkley, 1930) at pH 6.9-7.0. Freshly prepared test solutions were made up in Chalkley's solution and were used for each day's experiments, so that organisms were exposed in one of the following concentrations during a period of 45 min; 0, 5, 10, 20, 30 or 50 ppm of each metal. Selenium solutions were pre- pared from selenous acid (H2Se03) ; vanadium from vanadyl sulfate (VOSO4'- 2H-0) ; and zirconium from zirconyl chloride (ZrOCl2-6H:iO). The pH of each solution was adjusted for each ex- periment so that in the testing it was maintained at pH 7.0 ± 0.1. For stroboscopic photography the ap- paratus was a binocular, Zeiss-Nomarski, research model, phase-contrast microscope with phototube on which a 35 mm Zeiss camera was mounted. An aluminum disc with two one-inch holes 180° from one another at the periphery was mounted on the chuck of a variable-speed laboratory stirring motor and was set so that the disc rotated between the light source (a re- search model microscope lamp) and the EFFECTS OF SELENIUM AS SELENOUS ACID (H2Se03) ON SWIMMING RATE OF TETRAHYMENA PYRIFORMIS 3 o o 0 ppm TIME /MIN Fig. 1. EfTccts of selenium on the swimming rate of Tetrahymena pyrijormis. Effects of Se, Va and Zr on Swimming of Tetrahymena 41 O 3 c/i O O EFFECTS OF VANADIUM AS VANADYL SULFATE (VOSO4.2H2O) ON SWIMMING RATE OF TETRAHYMENA PYRIFORMIS. log Y = a + bX 5 ppm Y 10 ppm Y 20 ppm Y 2.72 + 0.005X 2.69 + 0.003X 2.60 + (-0.196)X 50 ppm 30 ppm Y = 2.69 + (-0.607)X 50 ppm Y = 2.68 + (-0.779)X 10 15 10 25 30 35 40 45 TIME/MIN Fig. 2. Effects of vanadium on the swimming rate of Tetrahymena pyrijormis. reflecting prisms of the microscope. The rotating disc thus dehvered pulsed Hght at a preset rate/sec to maximize the dis- tance between images on the film. The apparatus delivered pulsed light with an accuracy to 0.01 sec. Photographs were taken of groups of Tetrahymena swimming at 63X or lOOX magnification under darkfield illumina- tion, achieved by adjusting the phase con- densor system. The first was taken as soon as the Tetrahymena were placed on the microscope and the focus adjusted. This is designated as "zero" time and five photographs were taken within one min- ute. The light was turned off, then on again at 15 min and another five photos taken. The procedure was repeated at 30 min and 45 min. Distances between the images were measured from the rear of one to the rear of the next forward. Meas- urements were made on projections of the images onto a viewing screen at total magnification of 1200X to eliminate as much error of measurement as possible. Five measurements were made for each organism and ten organisms were meas- ured for each time period with mean rate calculated from the 50 measurements. Re- gression line plots were determined for each control and experimental set of data acquired for each concentration (Sokol and Rohlf, 1969). Prior to experimentation, about 10 ml of a 1-2 day old culture of Tetrahymena in mid log-phase growth (about 200,000 42 The University of Kansas Science Bulletin cells per ml) was concentrated by centrif- ugation at 1550 rpm for 15 min, the super- natant discarded and the cells resuspended in Chalkley's solution and washed, then reconcentrated by centrifugation and transferred to Chalkley's solution in a holding chamber (Bergquist and Bovee, 1976) where the solution was stirred con- tinuously. The pH of the solution in the holding chamber was checked at intervals and, if necessary, was adjusted with either HCl or NaOH solution to pH 7.0. Organisms to be measured for swim- ming rate were pipetted from the hold- ing chamber into a drop of the test solu- tion on a clean glass microscope slide and were covered with a glass cover slip sealed peripherally with medically pure petro- leum jelly. They remained thus during the experiment. The amount of fluid pipetted with the organisms was taken into consideration, so that the organisms were exposed to the desired concentration of the metal being used. Results of Experiments Swimming rates of controls. — The swimming rates of controls were un- affected by the Chalkley's solution or by changes of temperature from 22° to 27°C during continuous exposure to the light of the microscope. The pH did not aflect swimming rate in the range of 6.9 to 7.1. Control rates of swimming were close to EFFECTS OF ZIRCONIUM AS ZIRCONYL CHLORIDE (ZrOCl2'6H20) ON SWIMMING RATE OF TETRAHYMENA PYRIFORMIS. u c/1 o o TIME /MIN Fig. 3. EfTccts of zirconium on the swimming rate of Tetrahymcna pyrifonni^. Effects of Se, Va and Zr on Swimming of Tetrahymena 43 2700 ^m/sec without significant variation, during the 45 min of testing. Swimming rates in selenittyn. — Regres- sion Hne plots of the data for effects of selenium on the swimming rate showed a decrease of rate from 2700 jxm/sec to 2200 jxva/stc (18.5%) after 45 min in either 5 or 10 ppm. At 20 ppm the rate decreased to ~ 1000 ^m/sec (62.9%) ; at 30 ppm the rate decreased to ~ 35 jum/sec (98.6%); and at 50 ppm the organisms stopped swimming after ~ 40 min (Fig. 1). Swimming rates in vanadium. — Vana- dium had no effect on swimming rate within 45 min at 5 or 10 ppm. At 20 ppm the rate decreased to ~ 2000 jxm/stc {29.6%) ; at 30 ppm it decreased to ~ 520 ^m/sec (80.7%); at 50 ppm, to ~ 75 /xm/sec (97.2%) (Fig. 2). Swimming rates in zirconium. — At 5 ppm zirconium did not affect swimming rate. At 10 ppm the rate decreased to ~ 2200 ^m/sec (18.5%); at 20 ppm, to ~ 1000 ^m/sec (62.9%) ; at 30 ppm, to ~ 70 yum/sec (97.4%). The organisms stopped swimming in ~ 24 min at 50 ppm (Fig. 3). From these results it is clear that the swimming rates are significantly aflected by any concentration of these metals greater than 10 ppm under the conditions of experiment. Discussion Trace amounts of selenium (Cunja, 1973) or of vanadium (Weast, 1970) in the diet are stimulatory to the growth of vertebrates, as well as to Tetrahymena pyrijormis (Tippitt, 1975; Schreiber, 1978). Amounts greater than 10 ppm are detrimental to vertebrates if continued over a period of time (Muhler, 1957; Niimi and Leham, 1976) or to the growth of Paramecium (Stanton et al., unpubl.) or of Tetrahymena (Tippitt, 1975; Schreiber, 1978;' Makindc, 1979). Either may become concentrated in vertebrate tissues to toxic levels, especially in kidney. liver and spleen (Hathcock et al., 1964; Oehm, 1972). Zirconium is also toxic in like amounts in small mammals (Kang et al., 1977) and is potentially carcinogenic in small mammals (Turk and Parker, 1977) and has been noted as toxic to the human fetus (Scanlon, 1975). Therefore, the easy and rapid detection of toxic levels of these metals in a water source known to be polluted by one or another of them or a quantitative indica- tion of pollution at a toxic level, of un- known origin and of whatever nature, is a necessary tool in the evaluation of the safety of water for animal and human consumption. Our previous studies by these methods of measurement show that they are ef- fective and accurate to determine toxic and subtoxic levels of other metals (Ag, Cd, Fe, Ni, Pb, Cu; Bovee et al., 1977), providing an effective bioassay. These studies on Se, Va and Zr also indicate its utility. Further, it is not necessary to use the detailed photographic procedures. Visual observation is sufficient. If the organisms have stopped, or nearly stopped swim- ming in 30 to 40 min in the water being tested, there is a significant level of pollutant (s) in the water. Although the present method will not identify the pol- lutant (unless it is already known) nor indicate the quantity of it present in the water, it can serve as a means of assessing the relative safety of a water source and can determine the presence of potentially dangerous pollutants that should be quantitatively assessed by more critical methods. Literature Cited Bergquist, B. L. 1974. The effects of cadmium upon j^rowth, locomotion and ultrastructurc of Tetrahymena pyrijormis. PhD. Dissertation. Univ. Kansas, Lawrence. 147 p. Herc.ouist, B. L. and E. C. Bovee. 1976. Cadmium: quantitative methodology and study of its effect upon the locomotor rate of Tetrahymena pyri- jormis. Acta Protozool. 15:471-483. 44 The University of Kansas Science Bulletin BiCK, H. 1973. Population dynamics of Protozoa associated with the decay of organic matter in fresh water. Amer. Zool. 13:149-160. BovEE, E. C. 1965. An ec(jlogical study of amoebas from a small stream in northern Florida. Hydro- biologia 25:69-87. BovEE, E. C, B. L. Bergquist, E. Wyttenbach, D. J. Sternshein and A. Tippitt. 1977. Effects of heavy metals on Teterahymena pyriformis: a summary. Abst., 5th Intern. Congress Protozool., New York, June 1977. p. 318. BovEE, E. C, D. A. Kegley, D. Sternshein, E. Wyttenbach and B. L. Bergquist. 1979. Allevia- tion of the toxicity of copper to Tetrahymena pyriformis by nontoxic iron. Kansas Sci. Bull. 51:679-684. BuRBANCK, U. D. and D. M. Spoon. 1967. Use of sessile ciliatcs collected in petri dishes for rapid assessment of water pollution. J. Protozool. 4:739-744. Cairns, J., Jr. 1973. Factors affecting the numbers of species in freshwater Protozoan communities, pp. 219-247. In J. Cairns (ed.), The Structure and Function of Fresh-Water Microbial Com- munities. Res. Div. Monogr. #3. Va. Polytcch. Inst. State Univ., Blacksburg, Va. Carter, J. W. and I. L. Cameron. 1973. Toxicity assay of heavy metals in water, using Tetrahy- mena pyriformis. Water Res. 7:951-961. Chalkley, H. W. 1930. Stock cultures of Amoeba proteus. Science 71:442. CooLEY, N. R., J. M. Keltner, Jr. and J. Forester. 1972. Mirex and Achlor 1254: effect on and accumulation by Tetrahymena pyriformis. J. Protozool. 19:636-639. CuNjA, T. J. 1973. The value of selenium in animal feeding. Feedstuffs 45:21-27. Hathcock, J. N., C. Hill and G. Matrone. 1964. Vanadium toxicity and distribution in chicks and rats. J. Nutrition 82:106-111. Kang, K. Y., D. Bice, E. Hoffman, R. D'Amato and J. Salvaggio. 1977. Experimental studies of sensitization to beryllium, zirconium and alumi- num compounds in the rabbit. J. Allergy Clin. Immunol. 59:425-436. Lackey, J. B. 1938. Protozoan plankton as indicators of pcjllution in a flowing stream. Publ. Health Repts. 53:2037-2042. . 1968. The Ecology of Eiiglena, p. 28-44. In D. Buetow (ed.). The Biology of Euglcna, Vol. I. Academic Press, N.Y. Makinde, C. a. 1979. The toxic effect of vanadyl sulfate upon the growth of Tetrahymena pyri- formis. M.A. Thesis. Univ. of Kansas, Lawrence. 85 p. Mills, W. L. 1973. Some toxicity studies on selected Protozoa in certain toxicities of mercury. PhD. Dissertation. Univ. of Kansas, Lawrence. 128 p. MuHLER, J. C. 1957. The effects of vanadium pcn- toxide, fluoride and tin compounds on the dental experience of rats. J. Dental Res. 36:787-794. NiiMi, A. J. and Q. N. Laham. 1976. Relative toxicity of organic and inorganic compounds of selenium to newly-hatched zebra fish. Can. J. Zool. 54:501- 509. Oehm, F. W. 1972. The mechanism of heavy metal toxicities. Clin. Toxicol. 151-167. Ruthven, J. A. and J. Cairns, Jr. 1973. Response of Protozoan freshwater artificial communities to metals. J. Protozool. 20:127-129. ScANLON, J. W. 1975. Dangers to the human fetus from certain heavy metals in the environment. Rev. Environ. Health 2:39-64. Schreiber, D. E. 1978. The effects of vanadium upon the growth of Tetrahymena pyriformis. M.A. Thesis. Univ. of Kansas, Lawrence. 85 p. Small, E. B. 1973. A study of ciliate protozoa from a small polluted stream in east central Illinois. Amer. Zool. 13:225-230. SoKOL, R. R. and F. J. Rohlf. 1969. Biometry. Free- man, San Francisco. Sundaresen, B. B., E. C. Bovee, D. E. Wilson and J. B. Lackey. 1965. Effects of wood reduction waste pollution on the microbial ecology of a small creek. J. Water Poll. Contr. Fed. 37:1536- 1544. Tippitt, A. D. 1975. The effects of selenium upon the growth and locomotion of Tetrahymena pyri- formis. M.A. Thesis. Univ. of Kansas, Lawrence. 110 p. Turk, J. L. and D. Parker. 1977. Granuloma for- mation in normal guinea pigs injected intra- dcrmally with aluminum and zirconium com- pounds. J. Invest. Dermatol. 68:336-340. Weast, R. C. 1970. Handbook of Chemistry and Physics, 51st ed., p. B-36. Chemical Rubber Co., Cleveland. 1 1 ,v •!• .y 1 i i ,v •y A ^ s* .> •!• •v Iv S '^i X •A • •: % 1 i S' 'A A" S" ■A K 1 1 i .V ?• .V g ^ ^i s $ § g s i i 1 1 !«.!«.:•!••-•-• • .•.••.•.■.•.•.e .•.«.■ I. •,•.••.•.•.< THE UNIVERSITY OF KANSAS SCIENCE BULLETIN LIBRARY l-r ;rjtVFRSlTY ETHOMETRICS OF FIVE SYMPATRIC COCKROACH SPECIES IN NORTHEASTERN KANSAS .y ^ ft; i I s; i5 ^ I ••• ••• I S s By Robert E. Gorton, Jr. and W. J. Bell § i S Vol. 52, No. 5, pp. 45-58 January 21, 1983 ANNOUNCEMENT The University of Kansas Science Bulletin (continuation of the Kansas Uni- versity Quarterly) is an outlet for scholarly scientific investigations carried out at the University of Kansas or by University faculty and students. Since its incep- tion, volumes of the Bulletin have been variously issued as single bound volumes, as two or three multi-paper parts or as series of individual papers. Issuance is at irregular intervals, with each volume prior to volume 50 approximately 1000 pages in length. Effective with volume 50, page size has been enlarged, reducing the length of each volume to about 750 pages. The supply of all volumes of the Kansas University Quarterly is now ex- hausted. Howe\er, most volumes of the University of Kansas Science Bulletin are still available and are offered, in exchange for similar publications, to learned societies, colleges and universities and other institutions, or may be purchased at $20.00 per volume. Where some of these volumes were issued in parts, individual parts are priced at the rate of 2 cents per page. Current policy, initiated with volume 46, is to issue individual papers as published. Such separata may be purchased individually at the rate of 3 cents per page, with a minimum charge of $1.00 per separate. Subscriptions for lorthcoming volumes may be entered at the rate of $20.00 per volume. All communications regarding exchanges, sales and subscriptions should be addressed to the Exchangl Librarian, University OF Kansas Libraries, Lawrence, Kansas 66045. Reprints of individual papers for personal use by investigators are available gratis for most recent and many older issues of the Bulletin. Such requests should be directed to the author. The International Standard Serial Number of this publication is US ISSN 0022-8850. Editor William L. Rloom Editorial Board Philip W. Hedrick Rudolf Jandcr Harvey Lillywhiic Charles D. Michcncr Norman A. Slade Henry D. Stone George W. Hyers, Chairman THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Vol. 52, No. 5, pp. 45-58 January 21, 1983 Ethomctrics of Five Sympatric Cockroach Species in Northeastern Kansas Robert E. Gorton, Jr.-^ and W. J. Bell Department of Entomology, University of Kansas, Lawrence, Kansas 660-15 Contents Abstract 45 Introduction 45 Materials and Methods 46 Animal maintenance and observational procedures 46 Definition of acts 46 Analysis of encounters as units 47 Informational analysis 47 Diads 50 Results 50 Comparison of the behavioral repertoires 50 Parameters of agonistic behavior 50 Informational analyses 52 Diad analysis 55 Discussion 55 Acts and encounters 55 Informational analyses 57 Acknowledgments 58 Literature Cited 58 Abstract Five species of cockroaches in the genera Parcohlatta and Ischnoplera were collected in the field and ob- served under laboratory conditions. The behavioral repertoires of each species were determined, analyzed, and compared. Frequencies and sequences of acts, agonistic intensity, and lengths of encounters were con- sidered. Informational analyses were employed to determine the constraints the acts of one organism placed on the following acts of another and to determine the amount of information transmitted from one animal to another. Ecological and evolutionary implications of the results were discussed. Introduction willingness or unwillingness of individuals ....... to defend, usurp, or share those resources Agonistic behavior is a common way . , , .,..,,, , __ _. _ _ ^^ ^ , _ with other individuals. In general, ago- nistic behavior is the mechanism by which organisms efTect their social environments. for organisms to affect access to and con- trol of resources. Such behavior signals the It, . ,, r^ ■ • . rr 1 t^ Usiug Certain techniques, especially Present address: Division of Biology, Kansas " . . ., ■ State University, Manhattan, Kansas 66506. Markovian analyses, it is possible to quan- 46 The University of Kansas Science Bulletin tify and predict acts, sequences of acts, and the dependence of the behavior of one animal on the behavior of another (Stein- berg and Conant, 1974; Steinberg, 1977; Bell et al., 1979). With these techniques in mind, a comparative study was under- taken of live species of sympatric cock- roaches to determine, define, and com- pare the behavioral repertoires, frequencies and sequences of acts, the intensity and duration of agonistic encounters, and the interdependence of acts and runs of acts for conspecific encounters in Parcoblatta pennsylvanica, P. lata, P. virginica, P. uhlenana, and Ischnoptcra dcyopeltijormis. From these data the adaptive significances of the patterns derived are suggested. Materials and Methods Anijyial Maintenance and Observational Procedures Six field-captured animals were placed in each of three clear plastic cages 30X19X12.5 cm, covered by see-through plastic tops and placed on a 12:12 h photo- cycle at 26" C and 40% RH. After 24 h the animals were observed at various times during the dark period of the photocycle under red light (invisible to the cock- roaches; Walther and Dodt, 1957) until a minimum of 250 encounters had been ob- served and recorded on cassette tape for later transcription. When an individual died it was replaced and observations were continued after waiting another 24 h. Food and water were supplied ad lib. Definition of Acts The acts comprising the behavioral repertoires utilized in encounters with con- specific members of the same sex by /. deropeltijormis and Parcoblatta spp. were defined as follows: Approach (A) : moving toward and contacting another individual. This act began all encounters by defini- tion, Antennate (AN) : contacting an indi- vidual with the antennae, without the act being coupled with "ap- [Troach" above. Bite (B) : grasping another individual with the mandibles; a dangerous act sometimes resulting in lost body parts. Kick (K): kicking an animal, usually with the hind leg, occasionally with the middle leg. Stilt (S) : raising the body off the sub- strate by extension of the legs and often pointing the tip of the abdo- men vertically. Considered a warn- ing or threat posture. Stilt-kick (S-K) : kicking is executed as the body is lifted off the substrate (a composite act of the two acts above) . Climb (C) : non-sexual mounting and walking on the back of another in- dividual. Jerk (J) : rapidly vibrating the body one or more times in an anterio- posterior direction without locomo- tion. Ltinge (L) : rapidly moving toward an- other individual. Does nothing (DN) : remaining mo- tionless, without changing from nor- mal posture; the absence of an ob- servable response. Retreat (R): rapidly moving away from an individual (running); usu- ally a terminating act. Fly (F) : taking flight (a terminal act). Withdraw (W) : slowly moving away from another individual (walking) ; usually a terminal act. Truce (T) : remaining in contact with another individual with no further interaction for at least 5 sec (a ter- minal act). Back (BA) : backing toward an ap- Cockroach Buiavior 47 preaching animal (only in /. dero- peltifornii^). Butt (15U): ramming another indi- vidual with the proiiotiun (onl\ iii /. (leropcltijormis) . Lean (LE): tilting the dorsum side- wise in the direction of an approach- ing individual by flexing the legs on one side and extending the legs on the other (onK in /'. pennsyl- vanica) . The frequencies and probabilities of acts as a function of whether they were performed by the initiator of an encounter (that animal performing the first act of an encounter) or the responder (that ani- mal performing the second act of an en- counter) were calculated (Table 1). Analyses of Encounters as Units Agonistic intensity of each encounter was determined using an ordinal scale of one to five, depending on the intensity of the acts comprising the encounter (Table 2). The frequencies of the numbers of encounters falling into each agonistic level and the overall mean for each species is given in Table 3. Statistical significance of the ditiferences in mean levels of ago- nistic intensity of encounters between males and females of the same species was determined using the Smirnov 2-sample test (Table 4). Statistical significance of difTerences in the frequency distributions of agonistic levels (Table 3) of males of a species with males of each other species and females of a species with females of each other species was determined using the 2X2 contingency test (Table 5). Means and ranges of encounter lengths in numbers of acts were calculated (Table 6) and compared with ANOVA and SNK tests. Informational analyses Information theory provides a means of dissecting the dependence of one ani- mal's acts on another through the course ol an interaction. It provides a mechanism by which one can investigate the prob- abilistic constraints placed on the behavior of one animal by the acts of another, the restrictive value carried by specific acts, and the im[iortance of an animal's pre- ceding acts in determining its own subse- c]uent behavior (Steinberg, 1977). The measure of the diversity of acts chosen by the responder as influenced by the initiatcjr is H(x), such that where P^j,-^ is the probability of the ;th act executed by the responder given the /th act by the initiator has just occurred, and similarly for the diversity of the acts chosen by the initiator as influenced by the responder. When this index is normalized tor the number of acts in the repertoire of a species, it provides a means of inter- specific comparison. Normalized diversity varies from zero (complete certainty about the response to a given act) to one (com- plete uncertainty about the act that will follow the act in question, i.e., statistical independence) (Tables 7 and 8). Transmission, 7'(A^;y), is a measure of the information transferred. It is a measure of relatedness, equaling zero if the acts of the two animals are statistically independent: T(X;Y)=H{y)-H{Y/X). This index is derived from the Shannon- Wiener index of diversity: where V symbcjlizes the categories of acts and /', is the probability of the /th value of V, and from the conditional uncertainty index: H{Y/X)=-tPiM-2Pu/i and similarly for H(X) and H(X/Y). The value by which T{X;Y) approaches 48 The Univkrsity of Kansas Science Bulletin Table 1. Frequencies and probnbilitics of acts in the behavioral repertoires of each species. Animal A is the initiator of the encounter, Animai B the responder. (See Materials and Methods for explanation of symbols.) /. dcropi chijonnis ?. virgi nica Males (N=924) Females (N=923) Males (N=965) Females (Ar=1039) A B A B A B A B A 262 1 272 1 281 7 274 9 (.2835) (.0011) (.2'H7) (.0011) (.2912) (.0073) (.2637) (.0087) AN 44 21 37 23 52 61 69 63 (.0476) (.0227) (.0401) (.0249) (.0539) (.0632) (.0664) (.0606) DN 6 17 2 35 9 52 7 39 (.0065) (.0184) (.0022) (.0379) (.0093) (.0539) (.0067) (.0375) B 1 — 0 2 22 23 49 28 (.0000) (.0022) (.0228) (.0238) (.0472) (.0269) S-K 4 (.0043) 86 (.0931) 0 (.onno) 39 (.0422) ~~' " ~~~ ■ R 4 3 20 34 11 80 52 83 (.0043) (.0032) (.0217) (.0368) (.0332) (.0829) (.0500) (.0799) C 11 0 45 3 14 1 14 4 (.0119) (.0000) (.0487) (.0033) (.0145) (.0010) (.0135) (.0038) J 2 31 1 30 1 10 0 22 (.0022) (.0335) (.0011) (.0325) (.0010) (.0104) (.0000) (.0212) L 2 5 6 12 17 26 22 40 (.0022) (.0054) (.0065) (.0130) (.0176) (.0269) (.0212) (.0385) S 9 57 5 78 1 37 1 35 (.0097) (.0617) (.0054) (.0845) (.0010) (.0383) (.0009) (.0337) W 133 7 111 33 91 41 64 30 (.1439) (.0076) (.1203) (.0357) (.0943) (.0425) (.0616) (.0289) T 90 12 44 12 19 13 25 19 (.0974) (.0130) (.0477) (.0130) (.0197) (.0135) (.0241) (.0183) K 7 108 3 69 13 61 11 79 (.0076) (.1169) (.0033) (.0747) (.0135) (.0632) (.0106) (.0760) F ttt^ itt 0 (.0000) 1 (.0010) ~ ~ BA 2 (.0022) 0 (.0000) "^ ^"~ "" BU 5 (.0054) 1 (.0011) p. penmyhanica P. tihlcriana Males (N=z907) Females (iV=1049) Males (yV=869) Females (iV=863) A B A B A B A B A 282 0 276 6 262 1 268 4 (.3109) (.0000) (.2631) (.0057) (.3015) (.0011) (.3105) (.0046) AN 44 51 47 29 31 60 31 41 (.0485) (.0562) (.0448) (.0276) (.0357) (.0690) (.0359) (.0475) DN 2 65 4 15 1 28 2 16 (.0022) (.0717) (.0038) (.0143) (.0011) (.0322) (.0023) (.0185) B 2 0 51 36 10 10 9 4 (.0022) (.0000) (.0486) (.0343) (.0115) (.0115) (.0104) (.0046) S-K 0 3 3 22 1 11 2 21 (.0000) (.0033) (.0028) (.0209) (.0011) (.0126) (.0023) (.0243) R 11 81 33 47 42 81 43 69 (.0121) (.0893) (.0315) (.0448) (.0483) (.0932) (.0498) (.0799) C 44 5 20 7 22 3 9 6 (.0485) (.0055) (.0191) (.0067) (.0253) (.0034) (.0104) (.0069) J 2 14 0 16 1 15 0 13 (.0022) (.0154) (.0000) (.0153) (.0011) (.0173) (.0000) (.0151) Cockroach Behavior 49 Table 1. (concluded) p. pen nfylvanica P. uhleriana Males (N=z907) Females (iV=1049) Males (Ar=869) Females (iV=863) A B A B A B A B L 4 3 14 9 26 29 21 38 S (.0044) 3 (.0033) 54 (.0133) 12 (.0086) 129 (.0299) 3 (.0334) 50 (.0243) 7 (.0440) 78 W (.0033) 104 (.0595) 55 (.0114) 98 (.1229) 30 (.0034) 97 (.0575) 30 (.0081) 80 (.0904) 27 T (.1147) 14 (.0606) 3 (.0934) 39 (.0286) 9 (.1116) 7 (.0345) 3 (.0927) 38 (.0313) 3 K (.0154) 0 (.0033) 53 (.0372) 12 (.0086) 87 (.0081) 1 (.0034) 42 (.0440) 2 (.0035) 31 F (.0000) 0 (.0584) 1 (.0114) (.0829) (.0011) 2 (.0483) 0 (.0023) (.0359) LE (.0000) n (.0011) 3 (.0033) (.0023) (.0000) (.0000) P. lata Males (iV=807) Females (N=879) A B A B A 269 3 277 4 AN (.3333) 38 (.0037) 42 (.3153) 36 (.0046) 74 DN (.0471) 1 (.0520) 20 (.0409) 0 (.0842) 18 B (.0012) 1 (.0248) 0 (.0000) 3 (.0205) 2 S-K (.0012) 0 (.0000) 11 (.0034) 0 (.0023) 17 R (.0000) 30 (.0136) 122 (.0000) 29 (.0193) 103 C (.0372) 6 (.1512) 1 (.0329) 17 (.1172) 0 I (.0074) 0 (.0012) 8 (.0193) 0 (.0000) 9 L (.0000) 23 (.0099) 13 (.0000) 30 (.0102) 19 S (.0285) 3 (.0161) 62 (.0341) 4 (.0216) 65 W (.0037) 84 (.0768) 15 (.0046) 97 (.0739) 28 T (.1041) 7 (.0186) 1 (.1103) 11 (.0318) 2 K (.0087) 2 (.0012) 44 (.0125) 4 (.0023) 30 F (.0025) 0 (.0000) (.0545) 1 (.0012) (.0046) (.0341) Not part of behavioral repertoire. " Part of behavioral repertoire, but not observed in e.xpcriments. H{Y) is a measure of the influence of the H{X). Tran.smission can be normaHzed act of the initiator on the act of the re- to decrease the influence of the number spender, and similarly for T{Y;X) and of behavioral categories chosen by a par- 50 The University of Kansas Science Bulletin ticular investigator for a given system in the follow in£r manner: t{X;Y) = T(X;Y) H(Y) allowing cross-species comparisons (Table 9). Variances of transmission indices were calculated as described in Steinberg (1977) : Var T(X;Y) = (tS(Pij(\og2{Pij/PiPjr) -T(X;Y))/N. Diads The most common 2-act sequences for each species were determined and dia- grammed in Figure 1. From these diads, it was possible to contrast and compare typical encounters. Results Comparison of the Behavioral Repertoires The behavioral repertoires of the Parco- blatta spp. and Ischnoptera were virtually identical in the acts employed (Table 1), with the following exceptions: "back" was uiiic]uc to /. cleropeltijormis males, while "butt" was unic|ue to females of this species. "Lean" was unique to P. pennsylvanica males. "Stilt-kick" was never observed in either sex of P. virginica, and "bite" was never observed in Ischnoptera males. There was variation in the relative pro- portions of the acts observed within and Table 2. Levels of agonistic behavior. Level 1: encounters in which the major elements arc an- tcnnation, retreat, withdraw, or truce. 2: encounters in which the major elements are threat or intention movements (stilt, jerk, climb, lunge). 3: encounters in which the major elements arc bit- ing or kicking, demonstrated by one animal. 4: encounters in which the major elements are biting and/or kicking, demonstrated by both animals. 5: encounters in which the major elements are biting and/or kicking while both animals turn in a tight circle while engaging in these behaviors. between species, e.g., "retreat" was rare in Ischnoptera males, but comparatively com- mon in females of that species, and "stilt" was proportionately nearly four times as common in P. virginica females as in P. pennsylvanica females. Ischnoptera and P. pennsylvanica were often on one extreme or other in relative proportions of acts chosen, e.g., "stilt-kick" was common in Ischnoptera but uncommon in the Parco- blatta species, while "lunge" was least com- mon in the two aforementioned species. An interesting finding in all species was that 46-50° 0 of the total numbers of acts in the repertoires comprised 77.5% (P. pennsylvanica females) to 92.9% (P. lata males) of all acts used in intraspecific encounters. Some acts were characteristic of the initiator of an encounter, determined by that animal perpetrating at least 66.7% of the frequency of a given act: "approach" and "climb." Other acts, "jerk," "stilt," and "does nothing," were characteristic of the responder. "Truce" was characteristic of the initiator in four of the five species studied, while "kick" and "stilt-kick" were acts chosen by the responder in four of the five species studied. The remaining acts showed variability within and between sexes and species (Table 1). Parameters of Agonistic Behavior When the frequencies of encounters falling into the five levels of agonism (Table 2) were determined (Table 3), the following patterns emerged: males and females had different mean levels of ag- onism in three of the five species (P. penn- sylvanica. P. virginica, and /. deropelti- formis) (Table 4). When Parcoblatta females overall were compared with Par- coblatta males, there was a strong trend for females to be more intensely agonistic than males (sign test; P < .06). Males of Ischnoptera were significantly more ag- onistic than anv of the Parcoblatta males CoCKRO.\CH P.IHAVIOR 51 Table 3. Frequencies of the levels of agonism in male-male and female-fcm.il'- intrniction*;. P. lata P. t t/ileriana Level Males (A/=250) Females (A'=250) Males (N=250) Females (Nz=250) 1 2 3 4 5 X = 110 92 47 1 0 1.756 (44.00%) (36.80%) (18.80%) ( 0.40%) 99 109 38 4 0 1.788 (39.60%) (43.60%) (15.20%) ( 1.60%) 91 104 51 3 1 1.876 (36.40%) (41.60%) (20.40%) ( 1.20%) ( 0.40%) 78 120 48 3 1 1.916 (31.20%) (48.00%) (19.20%) ( 1.20%) ( 0.40%) P. t'irginica P. Pennsylvanica Level Males (N=263) Females (W=256) Males (N=250) Females (N=250) 1 2 3 4 5 X = 114 85 53 5 6 1.874 (43.35%) (32.32%) (20.15%) ( 1.90%) ( 2.28%) 72 105 56 16 7 2.144 (28.12%) (41.02%) (21.87%) ( 6.25%) ( 2.74%) 101 100 47 2 0 1.800 (40.40%) (40.00%) (18.80%) ( 0.80%) 32 107 81 26 4 2.452 (12.80%) (42.80%) (32.40%) (10.40%) ( 1.60%) /. dcrc ipchijormis Level Males (.V=250) Females (iV=250) 1 2 3 4 5 X = 12 84 150 4 0 2.584 ( 4.80%) (33.60%) (60.00%) ( 1.60%) 37 115 96 2 0 2.252 (14.80%) (46.00%) (38.40%) ( 0.80%) (Table 5). Females were significantly dif- ferent in afTonistic intensity from each other with two exceptions: P. uhleriana and P. lata were statistically identical, as were P. pennsylvanica and /. deropelu- jormis (Table 5). Males and females showed significant intrasexual differences in the mean num- Taele 4. Analysis of differences in agonistic levels comparing males and females (Smirnov 2-sample test). (T = max |5i(a-) - 52(.r) |, where S\{x) and Siix) are empirical distribution functions of the random samples.) Species T a-le\el r. pennsyh'anica 0.276 0.01 P. virginica 0.152 0.01 P. uhleriana 0.052 n.s.^ P. lata 0.044 n.s. I. deropeltijorniis 0.224 0.01 ' Not significant. bers of acts comprising an encounter, de- fined as the sum of the acts employed by both of the individuals involved. Parco- blatta lata males had significantly shorter encounters than all species except P. uhleri- ana (ANOVA, P < .001; SNK, P < .05) (Table 6). This difference was the only significant difference observed in males re- garding encounter length. Likewise, P. lata females had, on the average, fewer acts per encounter than all other females (Table 6). Parcoblatta uhleriana and /. deropelti- jonnis females had significantly fewer acts per encounter than P. pennsylvanica and P. virginica females, and more than P. lata females. Parcoblatta virginica and P. pennsylvanica females had significandy higher mean encounter lengths than all other species, but were not significantly different from each other. Ranges were 52 The University of Kansas Science Bulletin Table 5. Comparison of the frequencies of agonistic levels in Parcohlatta and Ischnoptera. Numbers in the matrix indicate T-values of pairwise 2X2 contingency tests (Class 1: agonism scores of 1-2; Class 2: ag- onism scores of 3-5. See Table 2 for definition of agonistic levels). Values for males are given above the diagonal, values for females, below. /. P. P. virginica P. lata P. tihlcriana P. pennsylranica J. deropeltijormis peltiforrnis pennsylvanica P. iihleriana P. lata P. virginica 72.8== 1.8^ 0.4' 2.3' 93.3== 0.01' 0.6^ — 13.7= 80.6== 0.4' — 2.6' 6.7= 46.9= — 31.7= 44.8= 9.9* 1.4' 20.8= 31.1= 3.7" 'not significant; =P<.001; ='P<.05; *P<.005; 'P<.015. Table 6. Analysis of mean encounter lengths in numbers of acts in Parcohlatta and Ischnoptera. Underlines indicate lack of significant differences (ANOVA and SNK test; P < .05). Numbers above species names in- dicate mean numbers ejf acts per encounter and numbers below indicate ranges of encounter lengths. 3.67 /. deropeltijormis (2-8) Males 3.47 P. tihlcriana 3.68 P. virginica P. pennsylvanica (2-7) 3.23 P. lata (2-12) (2-9) (2-11) 4.00 P. virginica Females 3.45 P. Iihleriana 4.20 P. pennsylvanica 3.70 /. deropeltijormis 3.12 P. lata (2-18) (2-13) (2-13) (2-S) (2-9) Table 7. Normalized diversity of acts in males of Parcohlatta and Ischnoptera. P. Iihleriana P. lata P. pennsylvanica p. vi) ■ginica /. derope A to B Itijormis Act A to B B to A A to B B to A A to B B to A A to B B to A B to A A .90 xxx' .80 xxx .81 xxx .86 xxx .80 xxx AN .81 .66 .77 .88 .90 .66 .86 .79 .48 .88 DN XXX .83 XXX .82 xxx .70 xxx .83 xxx .84 S XXX .78 xxx .78 xxx .72 xxx .64 xxx .68 S-K XXX xxx xxx xxx xxx xxx xxx xxx xxx .69 J XXX .82 xxx xxx xxx .94 xxx xxx xxx .72 K XXX .79 xxx .71 xxx .85 .94 .81 xxx .70 B XXX xxx xxx xxx xxx xxx .80 .87 xxx XXX C .81 xxx xxx xxx .82 xxx .81 xxx xxx xxx L .30 .32 .43 .49 xxx xxx .48 .73 xxx XXX 'Occurring with a frequency of less than 12. similar in all species except P. pennsyl- vanica females, in which an encounter could consist of up to 18 discrete acts. Informational Analyses The diversity of acts resulting from the perpetration of a given act (Tables 7 and S) dilTered on that act and on which ani- mal, the initiator (A) or the responder (H) perpetrated it. The effect of the acts of the initiators on the following acts o£ the responders showed that "approach" Cockroach Biiiavior 53 had the least constraining effect on the following acts in four of the five species in both sexes (P. ithleriana, P. lata, P. virginica, and /. deropeltijonnis), while "antennatc" had comparatively little ef- fect in at least one sex of all of the species studied (P. ithleriana, P. lata, P. virginica, I. cleropeltijormis females, and P. pennsyl- vanica males). Conversely, "lunge" was the most determining act in three of the five species studied in both sexes (P. Ithleriana, P. lata, and P. virginica'). "Climb" was second only to "lunge" in determining power in females of two spe- cies (P. lata and P. virginica), and showed the highest degree of determining power in females of /. dcropeltijormis. The effect of the responders on the following acts of the initiators was least for "does nothing" in at least one sex of all five species (P. Ithleriana, P. lata, P. virginica, P. pennsyl- vanica females, and /. cleropeltijormis males), "l.unge" was again a highly deter- mining act in at least one sex in four spe- cies (/'. lata, P. Ithleriana. P. virginica, and /. dcropeltijormis females). "Stilt," "kick," or "stilt'kick" were of little determining clTcct in males of P. pennsylyvanica and in all iemalcs, but rather high in /. dcropelti- jormis and in males of P. virginica. "An- tennatc" was among the least determining acts in males of P. lata and in /. dcropelti- jormis. but among the most determining acts in P. ithleriana, P. pcnnsylvanica, and in females of P. lata. "Jerk" and "bite" were rather non-determining acts in males of /*. Ithleriana. P. pcnnsylvanica, and P. virginica and in /. dcropeltijormis females, Table 8. Normalized diversity of acts in females of Parcoblatta and hchnoptera. P. tihlc ■viana P. lata P. pcnnsyt ! ran lea P. vii ■ginica /. dcropeltijormis Act A to B B to A A to B B to A A to B : ii to A A to B B to A A to B B to A A .85 XXX^ .88 XXX .75 XXX .91 XXX .89 XXX AN .91 .71 .93 .72 .74 .76 .86 .85 .84 .75 DN XXX .92 XXX .96 XXX .91 XXX .92 XXX .69 S XXX .79 XXX .81 .87 .89 XXX .79 XXX .77 S-K XXX .81 XXX .86 XXX .88 XXX XXX XXX .82 J XXX .73 XXX XXX XXX .86 XXX .85 XXX .89 K XXX .81 XXX .75 XXX .84 XXX .86 XXX .80 B XXX XXX XXX XXX .80 .76 .85 .78 XXX XXX C XXX XXX .52 XXX .85 XXX .81 XXX .77 XXX L .45 .35 .00 .00 .83 XXX .52 .54 XXX .71 "■Occurring with a frequency of less than 12. Table 9. Information transferred (bits) in encounters of Parcoblatta and hchnoptera, transmission (see text) is given in parentheses for each category. 2 SD. Normalized Species Male: A to B Male: B to A Female: A to B Female: B to A P. uhleriana 0.45±.04 0.83 ±.02 0.43±.04 0.86±.03 (0.14) (0.30) (0.14) (0.30) P. lata 0.29±.04 0.69±.03 0.42±.04 0.75±.03 (0.11) (0.27) (0.15) (0.28) P. pctinsylranica 0.20±.04 0.56±.05 0.58±.03 0.49±.03 (0.07) (0.23) (0.19) (0.16) P. virginica 0.49±.04 0.77±.02 0.43±.04 0.83±.03 (0.16) (0.27) (0.14) (0.28) I. deropeltijormis 0.46±.05 0.30±.04 0.37±.04 0.47±.04 (0.18) (0.13) (0.12) (0.18) 54 The University of Kansas Science Bulletin DN<- 19 OS' 14 K .-..: 05 OB Fig. 1. Diagrams of common dlads in Parcohlatta and hchnoptera. A: P. uhleriana. B: P. virginica. C: P. lata. D: P. pcnri..•.•-•.•.•.< '.•.•••.•-•.•.•.< >.•.•.•-•.•.•.< :: THE UNIVERSITY OF KANSAS SCIENCE BULLETIN •:• .*. V •:• .•• V •:• .% V •;• ••. V •!• .*. V ? g ? g S 5 § ft ¥ :5 ft ¥ * $ •:: V • • • •!• .*. •!• .*. V ♦ ,: •!• .V •!• .v V '> •:•; •: :•: •:• y •:•:::::: :::>:? •^ •:•; :•: :•: ft: ? •:• •:• > THE CRANE FLY GENUS CHIONEA IN NORTH AMERICA By George W. Byers ft w> v •'. •!• ••• V •!• V •!• I Vol. 52, No. 6, pp. 59-195 March 18, 1983 ANNOUNCEMENT The University of Kansas Science Bulletin (continuation of the Kansas Uni- versity Quarterly) is an outlet for scholarly scientific investigations carried out at the University of Kansas or by University faculty and students. Since its incep- tion, volumes of the Bulletin have been variously issued as single bound volumes, as two or three multi-paper parts or as series of individual papers. Issuance is at irregular intervals, with each volume prior to volume 50 approximately 1000 pages in length. Effective with volume 50, page size has been enlarged, reducing the length of each volume to about 750 pages. The supply of all volumes of the Kansas University Quarterly is now ex- hausted. 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Such requests should be directed to the author. The International Standard Serial Number of this publication is US ISSN 0022-8850. Editor William L. Bloom Editorial Board Philip W. Hetlrick Rudolf Janilcr Harvey Lillywhitc Charles D. Michener Norman A. Slatle Henry I). Stone George W. Bycrs, Chairman ...us. COMP. Z LIBRARY THE UNIVERSITY OF KANSAS APR "^ i^oo SCIENCE BULLETIN ^^.^J^^^^^y Vol. 52, No. G, pp. 59-195 March 18, 1983 THE CRANE FLY GENUS CHIONEA IN NORTH AMERICA' George W. I^ers Contents Introduction 60 Acknowledgments 60 The Genus C/iionea 61 North American Chionca 62 Historical Review 64 Materials and Methods 64 Adults — Exi ernal Morphology 66 Adults — Internal Morphology 78 Natural History of Chionca 94 Ecological Distribution 94 Geographical Distribution 98 Seasonal Distribution .. 99 Longevity of Adults 100 Behavior 100 Mating Behavior 104 Parasites and Predators 105 Immature Forms of Chionca 1 08 Eggs 108 Larvae Ill Pupae ; 120 Key lo Males of North Amtrican Chionca 122 Key 10 Females of Norih American Chionca 124 The C. scita Group 1 26 C. alhcrtenHs Alexander 126 C. mucnahcana Alexander 131, 195 C. nivicola Doane 133 C. scita Walker 138 C. wiUoni, new species 142, 191 The C. valga Group 1 42 C. hybrida, new species 1 42 C. obttisa, new species 1 45 C. stoncana Alexander 149 C. ralga Harris 154 The C. alcxandriana Group 1 62 C. alcxandriana (iarrett 162 C. caroliif Byers 169 C. diirbim, new species 1 72 C. cxcarata, new species 1 75 C. jellifoni, new species 1 ''9 C. lyrata, new species ' '^^3 C. nigra, new species I 86 Literature Cited „ 189 Appendix 1: C. wihoni, new species 191 Appendix 2: Male of C. dhk iiabcana 195 'Contribution No. 1780 from the Department of Entomology, The University of Kansas, Lawrence, Kansas 66045, U.S.A. 60 The University of Kansas Science Bulletii Abstract This is primarily a taxonomic revision: li.uht new species of ncarctic Chionca arc descriheil: diirhini, ex- cavata, hyhrida, jcUisoni, lyrata, nigni, ohtttsa and wiliotn. Chionca nniihoracensts Alexander is placeil in synonymy of C. valga Harris. These changes result in recognition of 16 North American species. The earlier grou[)ing of species based on number of flagcllomcres is rciected, and three species groups based on genital and other characters of both sexes are recognized. Separation of Nipliadohata Enderlein from Chionea is rejected. Morphological adaptations of Chionca fur loss of wmgs and a largely subterranean lite are examined. Ex- ternal anatomy and internal anatomy ot the adult ilies are described. The eggs, larvae and pupae are briefly described, including internal anatomy of the larva. Ecological relationships of Chionca to the physical environment and to other organisms are discusseil. In particular, the temperature range within which Chionca is actne on the snow and the flies" survi\al in extreme cold are explored. Various species of Chionea are shown to transport phoretic nematodes anil to ser\e as in- termediate hosts of mouse tapeworm. Geographic distribution of the species is correlated with latitude, eleva- tion and plant cover. Introduction Among the more curious of the thou- sands of known species of Tipulidae are the Httle wingless snow flies that comprise the genus Chionea. Their pecuHar, spider- like appearance and unusual occurrence as adults in uintcr have long made them subjects of comment by entomologists; however, the fact that they are abroad when most people are inclined to remain indoors has discouraged the collection of many specimens or of detailed informa- tion about their life histories and activities. Several species have been recognized and described, and occasional field observations of the living insects have been made, yet our fundamental knowledge of the genus remains fragmentary. Nearly every paper published on Chionea in North America has been based upon the chance discovery of one or two specimens; in fact it was precisely such a discovery oi a mere five individuals that prompted my own investi- gation of these insects. This study has been rather unsatisfying in certain respects, largely because of the difficulty of finding Chionca cither alive in its natural surroumlings or preserved in collections. Only about 1100 adult s[icci- mens could be brought together from all sources, and the distribution of these among the various species is exircmeb' uneven. Moreover, I have rarely h.ul the opjKjrtunity to observe living adults. The immature instars of Chionea have been encountered only two or three times in Europe, and only twice in North America. Because of this rarit\' of specimens, intra- specific variation has not been well under- stood in the past, with the result that the taxonomy of Chionea has been in certain instances confused. The long-standing misinterpretation of certain details of their external and internal anatomy may also be due to this rarity of specimens for study. In view of these circumstances, it seems to me worthwhile to brini: together and reorganize the accumtilated facts about Chionea, adding to them such things as I have been able to learn about the morphol- ogy, natural history and taxonomy of the genus. Perhaps what follows also will serve to stimulate further interest in these Hies, which may some day lead to a more complete treatment of them. Ackn()WLEIk;ments Several people have contributed to this undertaking in various ways, and the help of all is appreciated. For early encourage- ment in the study of Chionea, I am in- debted to the late Prof. J. Speed Rogers of the Univ. of Michigan; and for helping in many ways, especialK' bv making avail- able the types of his species of Chionea, I thank Prof. Emeritus Charles P. Alexan- der ot .\mherst, Massachusetts. I am in- liebted to eight hardy entomologists, in [■•articular, for making the major collec- tions of Chionca studied: Dr. M. S. Bris- The Crank Fly Gini's Chionea in North Amlrica 61 coe, Howard Univ.; Dr. John A. ('hap- maii, then ol Montana State Univ.; Mr. David \'. Durbin, San I'rancisco. (Cali- fornia; Mr. Roger A. Maiek, then ol the Univ. of Montana; Dr. W'lHiam L. Jelli- son, Hamilton, Montana, torinerK' ot the Rocky Mountain Laboratory, U. S. Public Health Service; Dr. William S. Osburn, then of the Univ. ol C'olorado; Mr. Davitl S. Potter. Univ. of Montana; and Mr. Ciary A. Shook, Ijoise, Idaho. 1 wish to thank the following curators and others for the loan of specimens: William F. Barr, Univ. ot Idaho; Sytlney G. Uannings, Univ. ol British C^olumbia; P. J. Darling- ton, Museum of C^om[\irative Zoology, Harvartl Univ.; Henry Dietrich and L. L. Pechuman, C'ornell Univ.; Paul Freeman. British Museum (Natural History); Mau- rice T. James and W'illiam }. Turner. Washington State Univ.; Edward L. Kes- sel and Paul H. Arnaud. Jr., California Academy of Sciences; Hugo G. Rodeck, Univ. of Colorado; Herbert H. Ross and Donald W. Webb. Illinois State Natural History Survey; Herbert J. Teskey and Guy E. Shewell. Entomology Research Institute, C'anada Department of Agricul- ture; Alan Stone. USD A, U. S. National Museum; and John A. Wilco.x, New York State Museum. I am particularly grateful to Dr. Ryuichi Matsuda and Dr. Herbert J. Teskey for constructive criticism of re- spectively the morphcjlogical and all other parts of this paper. Finally. I thank all others who have sent specimens of Chi- onea during this study, for every specimen has been a substantial addition. Generous support for part of this work by National Science Foundation grants G-9236 and DEB-77-15S6S is gratefully acknowledged. TiieC^enus Chionea Chionea is a small genus of only about two dozen species, widely scattered about the Holarctic region, 16 occurring in North America and the others in Europe, far eastern U.S.S.R., and Japan. Although these species e.xhibit c(jnsiderable variation in anatomical details, they have an overall simdariix' that involves a number of con- s[iicuous specializations (such as extreme modihcation ol the thorax), imparting to the genus the ap[iearance of structural homogeneity and at the same time so setting it ap.irt Irom other genera that it is not eas\' to conceive of a more "natural" group. In spite of this, the most recent re- vision ol the Etiropean species (Burghele- Balacesco, Y)i-f)) recognizes separate gen- era. Chionea and Niphadobata, for species groups differing in structure of the aedea- gus of the male and in details of the antennae. Enderlein (1*1^6: 1^) had based Niphadobata on the number of antennal segments (6-7, compared to 9-10 in typical Chionea). Kratochvil (1936: 250) found that most tlagellomeres in the species with longer antennae {longieornae group) have four verticillar hairs while those in the other group {hiaehycornae) have three. He also noted that in the lon'^icornae group the aedeagus is shorter than the gonapophyses and has small basal para- meres, in contr.ist to the condition in the brachy comae, in which the aedeagus is longer than the gonapophyses and has no basal parameres. Some North American species of Chionea bridge the gap in number of antennal segments, as discussed below, and the di (Terence in number of verticillar hairs does not hold for our species. Three verticillar hairs is the usual number in nearctic species with short antennae, for example, C. alexandriana, which has three flagellomeres. But three is also the usual number in C. scita and C. ulbertensis, which have respectively eight and ten fla- gellomeres, on the average, and also in those species having an intermediate num- ber of Hagellomeres. The number of such hairs is not constant within a species, however, and one or two flagellomeres 62 The Univlrsity of Kansas SciENcii Bulletin may have four hairs while others of the same antenna have three. None of the nearctic species belongs to the typical European group on the basis of short aedeagus with basal finger-like parameres; thus, there is no correlation whatsoever oi aedaegal structure with length of antennae. Furthermore, I am inclined to reject the division of Chionea into two genera on this basis because to do so ignores the great number of struc- tural specializations of the thorax, legs, wings, halteres and abdomen that all the species have in common. I have yet to see a key to the Diptera that will identify a specimen of Chionea as a tipulid. Its winglessness (actually it has tiny vestiges of wings) and stout body render it an unlikely candidate for offhand inclusion with other crane Bies, and it lacks the V-shaped transverse mesonotal suture by which supposedly all Tipulidae may be known from similar flies in other fami- lies." Its broad sternum separates the coxae widely, and when the antenna are greatly reduced, as happens in certain species, the inexperienced entomologist might suspect that Chionea belongs among the Pupipara. The genus was, in fact, placed in a sepa- rate family for a time during the middle 1800's by such eminent dipterists as Ron- dani, Zetterstedt and Bigot (e.g., Rondani, 1856: 38, 189). By 1869, however, Osten Sacken showed the eriopterine affinities of Chionea and moved the genus near to its currently accepted systematic position. Al- lowing for the moment that Chionea belongs somewhere among the Tipulidae, it must further be placed in the subfamily Limoniinae, on the basis of the short terminal segment ot the maxillary palps and the structure of the ovipositor, which ^ Marchanil (1917: 151) (iiscovcrcd ;in indication of taxonomic affinity that nearly cMryonc else seems to have overlooked: "One of ttu- . . . specitncns (of Chionea) . . . was found to have lost all its legs during transportation, showing ... in this [leculiar- ity Its relationship to the Tipululs." Sec also .Mnslie (1906). is unlike the form found in the Tipulinae or C'ylindrotominae. The placement of thv! genus in the tribe Eriopterini is based u[X)n the ftjllowing combination of char- acters: 1. eyes without short hairs interspersed among the facets, such as are found in the Pediciini, some of which are wingless. 2. tibiae without spurs. 3. pretarsal claws not toothed. 4. empodia well developed in form of arolia. Certain genera of the Eriopterini ex- hibit some of the unusual structural char- acteristics of Chionea, as, for example, the fusion segment at the base of the antennal flagellum or the undivided dististyles on the male genital segment. Accordingly, Alexander (1916: 530) regarded Chionea as a specialized relative of these genera, Cladiira and Pterochionea (now regarded as a synonym of Cladura), in a subtribe later designated the Claduraria, to which group the European Cryptena and Ameri- can Keolimnophila also belong. Subse- c]uent comparison of the immature forms of Chionea and Claditru has strongly sup- ported Alexander's placement of the genus near to Cladiira. North American Chionea Although aware of some differences between Old World and American Chio- neas, I do not believe the North American species can vet be distiiT^uished as a group from their Eurasian relatives. I have stud- ied the palearctic species only sufficiently to say that little more is known about them than about the American forms, ex- cept that their taxonomy has recently been revised in a detailed and excellently illus- trated publication by Anca Burghele- Balacesco (1969). Thirteen North American species of (Chionea have been described: C. alberten- Till. Crank Fly Genus Chionca in North Amirica 63 sis Alexander, C. alexandriana Garrett, C. iispcra. Walker, C carohts lUers, C. gracilis Alexander, (.. macnabcana Alex- ander, C. nivicola Doane, C novcboraccn- sis Alexander, C primitiva Alexander, C. scita Walker, C. stoneana Alexander, C. valga Harris, and C ivaiighi ('urran. Chionea aspera has lonu; been recognized as a synonym of C. valga. .md although C. scita was also regarded as a synonym of valga tor many years (Alexander, 1^42: 42')) I have considered it a valid name, with C. pnmitiva its synonym (Byers, 1%3: 160). On the basis of my opinions ex[')resscd in corres[X)ndence with him, Alexander (1%5: 72) placed C. gracilis and C. tvaughi in the synon\my of valga. These rearrangements in the Catalog of the Diptera of America North of Mexico (Stone et al., 1965) should have been noted as new synonymy. I further regard C. no- vehoracensis as a synonym of valga. To the remaining list I shall here add eight [previously undescribed species, bringing the total to 16. Species heretofore described have been dilTerentiated largely on the basis of color, size, and number of antennal segments. In the following revision, however, I shall trv to show that these characters, while useful for the dilTerentiation of species groups, have only limited utility in the diagnosis or recognition of the species. The most consistently useful characters for spe- cies recognition are the shapes of the gonapophyses in the male, and of the ovi- positor and associated sclerites in the fe- male; less commonly, another character may by some peculiarity prove to be whol- ly reliable. Many tipulid genera have been divided into subgenera — a practice widely adopted in the taxonomv of nematocerous Diptera. Although this has not been done in the case of Chionea, two major species groups were proposed by Kratochvil (1936: 250) for the European species: Chioncac longi- cornae, including those species with nine or ten antennal segments; and Chioneae brachycornac (emended to brachyccrae by Alexander, 1940: 101), including species that have six or seven antcnnomcres. En- derlein (1936: 19) gave the brachycomae generic status, as Niphadobata, but this name has not been consistently applied in the more recent literature (cf. liurghelc- 15alacesco, 1969). In classifying the North American ('hioneas, Alexander (1941: 212) adopted species group names, respec- tively aratu'oides grotip for the longicornae and valga group for the brachyccrae. While I have retained the valga grou[-) in my classification, it is based upon a combi- nation of characters as set forth below. In the North American species there is continuous variation in number of an- tennal segments from four (i.e., scape, pedicel, fusion segment and one additional Bagellomere) to 12, the number for any species varying often by one, rarely by two, exceptionally by more than two. I have therefore abandoned the older classifica- tion in favor of the following, applicable to the North American species only: 1. scita group — general body color yel- lowish brown or brown; antennae of 10- 12 segments; gonapophyses of male blade- like, without upturned apical hook; eighth abdominal tergum of female short (much less than half length of seventh tergum, measured front to rear); includes scita, albertensis, nivicola, tvilsoni n. sp., and macnabeana. 2. valga group — general body color red- dish brown, grayish brown or dark brown; antennae of 5 (rarely 4) to 9 segments; gona[)ophyses of male in form of a hori- zontal blade with a vertical keel, without apical hook; eighth abdominal tergum of female longer, about half length of sev- enth tergum; includes valga. stoneana, hvbrida n. sp., and obtitsa n. sp. Inclusion of stoneana and hybrida here is based pri- marily on the structure of the gonapo- plnses. The f)thcr species as a group have 7-9 antennal segments, as contrasted with 64 The University of Kansas Science Bullet IN 4 or 5 in stoneana and hybrida, and the eighth tergum in females of stoneana and hybrida is lengthened only at the sides, appearing short in dorsal aspect. 3. alexandriana group — general body color light brown, brown, dark brown or black; antennae of 5 (rarely 4) to 7 seg- ments; gonapophyses of male broad at base, slender, upturned and hook-like at apex; eighth abdominal tergum of female much shorter than half leniith of seventh tergum; includes alexandriana, carohts, durbtni n. sp., excavata n. sp., jellisoni n. sp., lyrata n. sp., and nigra n. sp. Historical Review The genus Chionea was established in 1816 by J. W. Dalman for a northern European species, C. araneoides, which is type species of the genus by monotypy. Thaddeus Harris described the first North American species, C. valga, in 1S41, though as early as 1833 he had mentioned the occurrence of the genus in Massachusetts and in 1835 named the species without describing it. Two North American spe- cies were shortly afterward described by Francis Walker, Chionea aspera (a syno- nym of valga) and C. scita (Walker, 1848). No additional North American species were discovered then for over 50 years. In 1900 R. W. Doane named the first species from western North America, (.. nivicola. Charles P. Alexander in 1917 described C. gracilis and C. noveboracensis (both here regarded as synonyms of valga). and C. primitiva (a synonym of scita). il. W. D. Garrett named the second species from the West, C. alexandriana, in 1922; and in 1925 ('. H. (Airran described C. waughi (synonymous with valga). Subseciuent additions to the list of American species have been the work oi C. P. Alexander and include C. stoneana (1940), C.albcrtcnsis (I'MI), and C. mac- nabeana (1946). References to Ncjrth Ameiican Chionea, other than these of a purely taxonomic nature, have been in general limited to a few brief notes on casual observations of the adults. Nothing has so far been pub- lished on the biology or life history of these Hies except by John Chapman (1954: 360-363), who discussed the ecological and seasonal distribution of some western spe- cies and described their activities at various temperatures. Mention should be made of the work of Jacques l^itsch (1955), which deals with European species but ofTers the only mod- ern treatment tjf the external anatomv of Chionea, and of Anca Burghele-Halacesco (1969), concerning taxonomy of the Euro- pean species based upon careful study of the genitalic anatomy. Ik'cause of its extreme structural modi- fication, Chionea has been accorded vary- ing degrees of special taxonomic status by various dipterists. Rondani (1841 : 284) early regarded it as in a separate "lamilia," C^hioneinae, and Zetterstedt (1842: 104) gave the genus apparent family status as C'hioncides. Bigot (1854: 466) also placed the genus in a separate family, C^hioneidae. However, Osten Sacken (1869: 136) noted the many similarities between Chionea and certain other crane flies and arranged it with the Eriopterina (now tribe Eriop- terini), where it remains today. More re- cently, nevertheless, Williston (1908: 86) separated all the Limnobiinae (now Li- moniinae) into tribes, except ior Chionea, left standing alone as a genus, liurghele- Balacesco (1969) recognized two genera, Chionea and Kiphadobata, for the Euro- [lean species, following Enderlein (1936) but with firmer morphological basis. Materials and Methods Taxonomic and morphological descrip- tions which follow are based upon the study of somewhat over 1100 specimens of adults of Chionea — all that could be lo- cated by a fairly thorough search of mu- The Crane Fly Gi;nus Chionea in North America 65 F.c. 1. Chiomu alcxandnunu Garrett, male, dorsal aspect (most of rij;ht legs omitted), to show general habitus. 66 The University of Kansas Science Bulletin seums and other collections. These have included the holotypcs or type series of all species except C. noi'cboraccnsis (t\pe lost) and C. alexatulriinni (paratypes seen) • The only immature forms studied were those of CInonea valga and C. stoneana. These will he described in detail in a separate paper, and only the latter are dealt with here in a 'icneral way. Although numer(jus flics pinned dry have been seen, a greater number have been preserved in alcoiioK the latter by far the more satisfactory s[iecimens. C'hioneas generalk make [iretty wretched pinned specimens because of their small size and tendency to become shriveled and distorted when dried, such as in extreme telescoping of the abdominal segments. Not only does preservation in fluid keep the insects in more or less their original shape and di- mensions, but it also permits subsequent study of the internal structtires and greatly facilitates identification, for the gona- pophyses of the male are readily exposed by pressing the membranous tenth abdom- inal segment forward beneath the ninth tergum, and the ovipositor of the female is easily opened to reveal the shape of the ninth abdominal sternum and the genital furca. Specimens mounted on microscope slides were hrst cleared in 10% potassium hydroxide, then washed in increasing con- centrations of ethyl alcohol (S0%, 95%, absolute), then in xylene, and finally cleared somewhat lurther in clove od. All were examined "in the round" bctore be- ing moinued in Euparal mediLun. Morphological studies were based upon ilies preserved in 70-!S0°o etlul alcohol. There .ire, of course, better lixing solutions for muscle and internal organs, but alcohol suffices; besides, few C^hioneas have ever been collected specifically lor mi)r|ihologi- cal examination. v\i)iLTs — External Moind it diminishes corresjiond- mglv trom *^^-l. 1 his first segment is accordingb' known ,ts the tusion segment, and although it cannot clearly be seen to comprise more than one segment in Chionea (but sec C'. W. Johnson's com- ment in Alexander, 1917: 202), its struc- ture in the closely allied Cladiira (formerly Ptcrochioneu) bradleyi (Alexander), show- ing five distinct verticils, seems to sub- stantiate this interpretation. I have com- jxired the antennae of Chionca scita and Clad lira bradleyi (as Pterochionea) in another paper (Byers, 196^: 190-191). The tusion segment is densely pitted and clothed with short hairs except on its base. Occasionally, especiall\- in species of the alexundnana group in which there is great- est reduction of the antennae, there will be touiul a long seta or two attached near the apex of the tusion segment. In the scita grouf") the second flagellomere (first beyond the tusion segment) is covered c.vtj» Chionca alcxuiidriaiia , drt.iiK of IkmiI. 2, IkmiI, left l.ittr.il aspect. 3, ri.ulit antenna, tiorsoiatcral luad, frontal or facial aspect (antennae removed). 5, head, ventral aspect. Abbreviations: ans — antennal M)cket. atp — anterior tentorial pit, cc — compound eye, dp — clypeus, cs— cervical sclcrite, es — espisto- mal suture (sulcus), hph — hypo|iharynx, lali — labial pal|i, or labellum, Ibr— lahrum, mxp— maxillary palp, ocd — occipital cond\le, of — occipital foramen, osl — or.il surface of JalH-llum, jic — pedicel of antenna, p.t;r — palpigcr of labial p. lip, |imt — prementum, ptp — posterior tentori.il pit, Fu;s. 2-5. aspect. -1, sti- stipes of m.ixill.i. 68 The University of Kansas Science Bulleti N with short hairs similar to those of the fusion segment. The other flagellomeres have three, or rarely four, long setae ar- ranged in irregular whorls. In general, these verticils increase in length as the number of flagellomeres is reduced, as if to compensate for the reduction and main- tain antennal length (e.g., compare Fig. 3 with 97). This may be an instance of what Matsuda (1979: 220) has called material compensation, such compensatory develop- ment otten being seen in insects with reduced wings (p. 224). The compound eyes are black in the living flies and often retain this color in preserved specimens. They are composed of about 80-200 ommatidia, the number varying generally according to species but also somewhat among individuals within a species, without any evident sexual di- morphism. They occupy a comparatively smaller portion of the head surface than in most Tipulidae, a reduction in size that may be related to the largely subterranean habits of Chioiu-a. It has been observed that subterranean and cavernicolous in- sects often have not only smaller com- pound eyes than their nearest relatives living above ground but also larger omma- tidia, which [K-rhaps are more efficient in gathering light in parts of the environment where some light is available. To see how this might a[i[)ly in the case of Chionea, 1 compared the com[i(iund eyes of male specimens cjf C.hionca valga .uul the related Claditra jJdVf)\evyitginca ot a[i[M(i\imately ecjual body size and head dimensions. In the slightly larger eye of Chionea there were nine iacets (3 X 3) within a sample Luut sc|uare, whereas the Cladiiru had 1() (4 X 4), a 78% greater number of omma- tidia [ler unit area in the day-flying Cla- ditru. The size and number of ommatidia in most other species of Chionea are rather close to those in C. valga. Ability to per- ceive lorm clearly, it seems, would be of little use to a slowly moving, non-flving insect such as Chionea: on the other hand, living as it does in darkened recesses be- neath the ground and coming onto the surface primarily at times ot low light in- tensity. It would derive scjme benelit irom eyes adapted to making the most (jf what light there was. Surface areas of the head such as genae. Irons, vertex and occijnit are indistinguish- ably lused, the only useful "landmarks" other than eyes and antennae being the tentorial pits, of which the posterior pair may easily be seen at about 125 X magnifi- cation (Figs. 2, 5). The anterior tentorial pits (Fig. 4) are smaller and concealed in the epistomal (tronto-cKpecd) suture, close beneath the eyes. The vertex bears a group of stout hairs, the longer ones of which are subet]ual in length to the scape. The short, thick rostrum is made up of the strongly arched clypeus in tront and a posterior (ventral) sclerotized area in uhich the stipites oi the maxillae are em- bedded (Fig. 2) and which is slightly in- serted under the antero-ventral marijin of the head, to which it has a membranous connection. /Vlthotigh ioi ned b\' this sclero- tization, the stiiiites are not ttised medially as m ti[Hiline crane Hies, such as Dolicho- peza (Byers, 19(,1^: (,S4, Fig. 9). Except for the stipes, the maxilla consists only of a five-segmented [xflp, of which the first segment is a[')[ir()\imatel\' spherical, the second ekjngate and somewhat curved, the third enlarged cUid having a more roughly Figs. 6, 7. Chionea alcxandriatia, external anatomy. 6, adult male, Id't Literal aspect (legs mostly omitted); as — anal (tenth) segment, bs — basist>le, cxi — coxa of metathorax, iK — tlististyle, fm — femur (cut shortly beyond trochanter), h — haltere, mxp — maxillary palp, pis — pleural suture of mesothorax, pn — pronotum, pst — pseudo- sutural fovea, sc — scape of antenna, sp— spiracle (of mesothorax), s9 — ninth abdominal sternum, t'J — ninth ab- dominal tcrgum, w — wing rudiment. 7, adult female, right lateral aspect (legs mostly omitted); ce — compound eye, cr — ccrcus, cs — cervical sclerite, hv — hypovalvc, ms — microsternite, mt — inicrotergite, sp — spiracle (of meta- thorax), si — 'first abilominal sternum, s8 — eighth abdoniin.d sternum, tS, t9, tlO — eighth, ninth and tenth abdominal tcrga. Tiiu Cram-. Fly Genus Chiontui in North Amurica 69 70 The University of Kansas Science Bulletin pitted surface (sensory structures') than the others, and the fourth and fifth seg- ments more elongate, as sh(n\n in Figures 2 and 6. Partially telescoped beneath the maxil- lary bases is the strongly sclerotized pre- mentun of the labium. It is forked at its inner (proximal) end, which extends into the head as an a[X)deme, and is slightly expanded dist.illy where it adioins directly the bases of the first labellar segments (Fig. 5). Over this junction lies a slightly sclerotized, setiferous lobe which I take to be a modification of the sclerite that in Tipulinae is interposed between the pre- mentum and the first labellar segment. This sclerite has been called the palpiger by Rees and Ferris (1939: 147, Fig. 73) and, where divided, the "kappa" and "sig- ma" sclerites by Peterson (1916: 45-46, Plate XVII, Fig. 288). The labellar lobes are broad and dorsally clearly show divi- sion into two segments. Their ventral surfaces are remarkable in being quite smooth; that is, there are no pseudo- tracheae whatsoever (Fig. 5). At high magnifications, a row of tiny denticles can be seen near the meso-ventral margin of each lobe. These possibly are the equiva- lent of the prcstomal teeth of certain higher Diptera (cf. Crampton, 1942: 36). The number of these denticles varies from 4-15, generally according to species, although it is not constant within a species. Closely appressed to the dorsal surface of the labium, in the dee[i notch between the labella, is the hyfiopharynx, which in Chionea extends lar beyontl the short labrum and, in the absence of labellar pseudotracheae, probabK is used to take ti[) lic]uid nutriment directU'. The hy[io- pharynx may be recognized by the com- pact brush ot fine, golden hairs at its ti[i. Thorax. — Loss of wings in Chioncu has resulted in a corresponding diminution ot tlie mesothorax, which, however, is still the largest thoracic segment. Wide and rather Hat on the ventral side, the thorax is narrowed and arched above as shown by serial cross sections (Fig. 13, 14, 15) and increases in breadth from front to rear. Since students of insect morphology cannot themselves agree on the interpreta- tion of the thoracic sclerites, perhaps a mere taxonomist will be excused for mak- ing a somewhat arbitrary selection from among the available names for parts. Iden- tification of thoracic areas in Chionea is complicated by the extensive fusion of dor- sal and lateral sclerites and obliteration of many oi the sutures that are fairly easy to discern in other tipulids. Using the coxae as reference points, however, it is not difficult to distinguish the three pleural sutures (Fig. 6, pis). The first of these (prothoracic) is in alignment with the junction between pronotum and mesono- tum and terminates dorsally in a deep im- pression, which internally is the attach- ment of the sternal apophysis; and the second and third extend from the meso- thoracic and metathoracic coxal processes, respectively, to the membranous areas about the bases of the vestigial wing and the haltere. Sclerotized areas before and behind these sutures may be identified as episternal and epimeral, respectively, but have no definite subdivisions (anepister- num, katepisternum, etc.), such as may be seen in the nearest relatives of Chionea. Crampton (1925: 201, 203) has commented on the reduction in Chionea ot the meso- thoracic meron, which in other genera of Eriopterini is large. The [ironotum, separated from the mesoncjtum by a fairly distinct suttire, is crossed from side to side by an indistinct row of bristles. Two transverse, shallowly im[)ressed, darkened spots on the meso- iiotiim just above the slit-like mesothoracic s[iiracles (Fig. 6) appear to be the pseudo- sutural foveae. There is no dilTerentiation ot mesoiKJtal prescutum, scutum, or scu- tellum, and the scjiaration of mesothorax trom metathorax is onlv weakly indicated dorsally by a shallow, transverse indenta- Tiiii Crane Fly Gunus Chionea in North America 71 tion and laterally by the position of the round metathoracic sjiiraclcs. Likewise, the nota grade impereeptibly into the [ilcu- ral sclerites along the entire length oi the thorax, the extensive, membranous para- tergal areas seen in other li[Uilid genera being absent altogether (cf. Byers, VHAb: 6S7,Fig. 14). In most crane flies, Gnophomyia being one exception, the coxae are set close to- gether ventrally, and the mesosternum and metasternum are almost wholly in- folded. l)y contrast, there are relatively broad ventral sclerites in Chinnca; but the cjuestion arises — what is then' identity.'' Are the sternal apo[-)hyses ol the ptero- thorax ". . . carried inward u[K)n a me- dian inflection ot the sternum to lorm the Y-shaped endosternal apodeme . . ." as stated by Snodgrass (19.^5: 171), or is it true that, as countered by Ferris (Rees and Ferris, YW: 149), ". . . there is ab- solutely no sternal element visible exter- nally on the thorax of any Dipteran . . ." app stn cs I app 3 &>V.S. abs I ex 3 E E in" d o-J Figs. 8-12, Chionea alcxandriatm, details of thorax. 8, thoracic venter (legs mostly omitted): abs 1— first abdom- inal sternum, app 1— apophyseal pit of prosternum, app2— apophyseal pit of mcsosternal furca, app3— apophyseal pit ot metasternal turca, cs— cervical sclerite, cx3— coxa of mctathorax. fcsl— furcasternum of prothorax, stn— sternum (combined basisternum and furcasternum) of metathorax. 9, haltcrc, posterior aspect. 10, detail of base of haltere, to show probable senv.ry structures. 11. haltere, lateral aspect; area of sensory pejjs greatly en- l.ir.urd. 12, Chionea stoncunu, fem.ile, portion of hind tibia showing three kinds of setae (see text for cx- pl.inati(jn) . Scale: Figs. 9, 11. 72 The University of Kansas Science Bulletin and that the apparent sternum is actually the subcoxal portion of the generalized insect leg, the endosternal apophyses (branches of the Y-shaped apodeme) being all that remain of the true sternum •" It seems to me that Snodgrass and Ferris arc in agreement that the ventral condyles ar- ticulating with the coxae belong to the subcoxal ring and that the sternal apoph- yses belong to the true sternum, so that what they are disputing is the location of the line ol fusion between these two scle- rites (cf. Snodgrass, 1935: 169, Fig. B, and Rees and Ferris, 1939: 165, Fig/ 80c). In the cleared specimens of Chionea I have examined, as apparently also in many other Diptcra, the line of fusion is not visible, its position therefore being hypo- thetical; consec]uently, there seems little to be gained by the argument. Rees and Ferris concede that what appears to be prosternum in Tipiila is in tact the pro- sternum. If the same point of view is accepted for Chionea, then I would say that on the basis of structural similarity (Fig. 8) the meso- and metathoracic ven- tral sclerites are also sternal and that the Y-shaped sternal apophyses have been formed in these segments by the reduction of the furcasterna toward the ventral mid- line. It is problematical why, in a diptcran that is altogether flightless, the halteres should be so well developed, but neither observation of the living flies nor micro- scopic examination of the structure of the halteres has led to an explanation. Except for the basal portion, or scabellum, the haltere is membranous and bladder-like (Figs. 9, 11) and covered with extremely fine hairs. The pedicel and capitellum, or knob, are not clearly divided. The capi- tellum seems to be hollow and in the living fly is probably filled with hacmolymph. The scabellum comprises an incomplete cup, curving around the pedicel ventrally, and a longer dorsal lobe, both merging proximally on a slender, membranous at- tachment without anv evidence of axillarv sclerites. It is likcK' that the halteres in Chionea have scjme sensory function, for each has a fairh' conspicuous nerve leading to it (but no visible tracheole), and on the base of the pedicel, mostly concealed with- in the scabellum, there are a dorsal and a ventral row of nodules, variable in number but most numerous dorsally, which prob- ably are groups of sensilla (Fig. 10). There is also, at the edge of the episternum adin— gonapophysis, Ml — muscle (see text for ori.i^'ins, nisertions and actions of numbered muscles). 17, gonapophyscs, vesica, penis ,ind .interior seminal duct, dors.il aspect. IS, .yenital structures, to show musculature: cmca— compressor muscle of compressor apodcme, gon— right gonapophysis, la— lateral apodcme, mg— muscle of gonapophysis, p — penis, rca— retractor muscle of compressor apodeme. sd— seminal duct, s 10— tenth sternum (membranous anal segment), vgl— vesicular gland, wgc— wall of genital chamber (mem- br.inous). Scale: Fig. IS. 76 The University of Kansas Science Bulletin fine hairs. Each basistvle bears a sintrle elongate, rigidly sclerotized dististyle, rath- er finger-like in general aspect. It is thick- est and irregularly shaped at the base and usually more or less evenly curved and somewhat tapered toward the Apex. There is a prominent, sometimes darkened, pos- teromesal tooth near the base (Figs. 155, 184), also a very small, inconspicuous, an- teromesal tooth and an additional black- ened ventral tooth of variable shape and size, most conspicuous in larger males, of- ten rudimentary in smaller males and ap- parently invariably absent in certain species. The dististyle is sparsely covered with setae along its outer (dorsal) surface but is bare on its inner curvature, except for a small patch of stiff hairs near the base. Ordinarily the gonapophyses (cf. Crampton, 1942: 92) are not easy to see because they are concealed from below and behind by the ninth sternum and basistyles and from above by the membranous tenth segment (Fig. 16). However, with the tip of a needle the tenth segment may, in a softened s[-)ecimen, be pushed forward be- neath the ninth, exposing the gonapoph- yses. Since this is the way they are most readily seen, I have illustrated the dorsal aspect for each species in the taxonomic treatment which follous. The gonapoph- yses are hardened and may be blade-like or hooked at the tip, as brietlv discussed earlier, but they are hollow basalK-, where they are immovably fused to the sclero- tized wall of the genital chamber and more flexibly attached laterally to the un- derside of the ninth tergum where it ad- joins the basistyles (Fig. IS). These pro- jections in (Jiioncd could be described morphologically as par.imeres and prob- ably are not homologous with the gona[iophyses ot some other tipulids, Doli- chopcza for exam[)le (Hyers, 1961^: 68(S); however, I am retaining the term because of its past use in taxonomv .uid because the structures seem to conlorm to a gen- eral definition of gonapophyses. Emerging through the anterior wall of the genital chamber between the bases of the gonajiophyscs. the penis, a short, non- extensible intromittent organ, curves down- ward to the floor of the genital chamber and then upward again (Fig. IS). In most species the distal end of the penis is un- modified, but in C. ulbertoisis it is pro- duced into two sharp points and in C. vuliia and C. ohtiisa into two sinuate fla- gella; in C. scita there is a spoon-shaped extension at the U\). Behind the ninth segment, or h\po- pygium, there is only the membranous tenth, or anal, segment. It could be ar- gued that morphologically this must be the fused tenth and eleventh segments, but in Chionea there is no indication of a division into two parts. In some species the sides of this segment are pigmented and lightly sclerotized (Fig. 6). Ill the female the eighth and succeeding abdominal segments are modified for copu- lation and oviposition. Elsewhere (Byers, 19()l/7: 690-692), I have compared the tipu- lid ovipositor in some detail with that of a typical pterygote insect, and since the fe- male terminalia in Chionea do not dilTer greatly from those of a tipuline crane fly. the comparison ncctX not be repeated here. Unlike the terga before it, the eighth ter- gum extends completely across the pleuron, and althoui^h it is not fused with the eighth sternum the two are joined by a strong, flexible attachment. In all species the eighth tergum is very short, about half the length of the seventh in scjme species (Figs. 112, 147) and reduced to a slender, transverse straji (Fig. 7) in others. It is bare except tor one or a tew hairs at each side. 1 he sternum, in contrast, is not only Ioniser than an\' other but is [Molonged l:.ick\\ard as two ^•^^'•2^ guides, the hypo- valves, or ventral valves, of the ovipositor (Fig. 7). There is a deep cleft at each side between the sternum [iroiuT and the hypo- valves, and in this region the connection is membranous, allowing vertical move- Tin: Ckank Fi.y (]iNi-s Chionca in Noktii America 77 mciit of the valves. Xuiiicrous loiiu;. stout, Ncllowish setae are t!ee[il\' set into the thickeueel, seleroti/.ed a[iieal and inlolded dorsal mariiiiis (Imu. 2-)). These a[i[iear to he immovahle or onK' slightly movable setae and are |)resLimed to he nistruniental in the [ilaeement ol ei;L!,s. In the menihrane between the eighth and ninth tcrga, at about the level of the lower edge of the ninth tergum on each side, there is a small, sclerolized spot about the size of a spiracle. These small sclerites are most prominent in species of the alex- jfuhi.ina grou[i. They are larger and more elongate dorso-vent rall\' in dlhcrtcnsis and icitd but inconspicuous in these species be- 57 58 60 59 lovd egg G.vvB Fic.s. 19-22. Chionca alcxandnaiui, 23, Chionca niricola. dft.iils of tcnialc rt-protluctivc system. 19, internal repnKkictivc orj^ans. (h.rsal aspect: acj: — accessory .ulanil, he— bursa opulatrix, ejij; — eg): or ovum, fspth functional spermatheca (anterior end of bursa), lovd — lateral ovuluct (some ovarioles cut away, their points of attachment indicated by elongate ovals on drawing), spd— spermathecal duct. s|)th— spermatheca (inner lines represent outlines of masses of sperm), s9— ninth abdominal sternum, tc— trophocyte. 20, composite longi- tucHnal section to show musculature of ovipositor: numbers indicate muscles (see text lor origins, insertions and actions of numbered muscles: note ventral end of m-S2 cut away to reveal cross-section of m-5-l). 21, bases of ducts of accessory glands enclosed in common covering. 22, sagittal section of posterior ends of bursa copulatrix and — muscles of furcal base, oacg— opening of accessory glands, pch— pouch between genital furca and upper wall of oviduct, spd— spermathecal duct, s9— ninth sternum. 23, hy- po\alves of eighth abdominal sternum, dorsal aspect; gp — gonopore. 78 The University of Kansas Science Bulletin cause of their pale yellowish color, and they appear to be absent in species of the valga group. I have not determinccl their significance. Segments behind the eighth arc reduced in diameter, their terga occurring above the elongate eighth sternum and their sterna partially or completely concealed. The ninth tergum is broadly fused dorsally with the tenth, from which it is separated lateral!}' by a clecji, narrtjw incision, below which the lateral extensions turn backward and inward to join the ninth sternum. The connection to the ninth sternum looks tenuous but is tough and not easily severed with dissecting needles. As used here the term ninth sternum means the sclerotized portion that is more darkly colored than surrounding membranous areas and is not cleared by potassium hydroxide in speci- mens prepared ior mounting on micro- scope slides. Of course, this may not con- stitute the entire morphological ninth sternum, the limits of which have not been determined. This sclerotized portion may comprise simply two convergent bands, but more often a median plate, of variable shape depending on the species, is formed. Between the ninth sternum and the upper surface of the eighth is the genital chamber into which the separate divisions of the reproductive system open (Fig. 22). Embedded in the membranous wall of the chamber just beneath the ninth sternum is the genital furca (see Byers, \%\b: 693), the arms of which join the ninth sternum and the base of which is attached to the roof of the commcjn oviduct. Situated me- dially below the ninth sternum and brack- eted by the arms of the turca is the aper- ture of the bursa copulatrix. The median or common oviduct rises sharply from the floor of the eighth sternum so that the gonopore lies not far below the opening to the bursa (Fig. 22). A deep [louch is formed between the furca and the lightly sclerotized upper wall ot the oviduct. Less modified than the ninth, the tenth or pygidial segment consists of a well sclerotized, saddle-like tergum and a weak- ly sclerotized, sparsely hairy sternum, above which is the anal opening. Flexibly joined to the posterior end of the tenth tergum, supposedly arising between the tenth and eleventh, are the cerci (Fig. 7) which typically are elongate, sabre-shaped and strcjiigly sclerotized, being especially thickened along their dorsal edges. A trace of sclerotization between the cereal bases may represent the eleventh segment. The cerci are often referred to as the dorsal or upper valves of the cjvipositor. Scattered over their outer surfaces but particularly concentrated at the tips are a number of microscopic pegs set in circular bases. Al- though their histology has not been ex- amined, these are presumed to be sensilla related io the detection of appropriate sites for oviposition. Adults — Internal Morphology Head. — Within the head, the most con- spicuous structure is a large, compact mass of nervous tissue inade up of the brain and suboesophageal ganglion (Figs. 24, 30). Seen from above (Fig. 29), the brain is clearly divided into right and left lobes, which appear to belong to the protocere- brum. These are curved downward at the sides (Fig. 30), concealing the poorly defined deutocerebral lobes from which the antennal nerves extend directly into the bases of the antennae. The optic lobes (Fig. 29) are remarkably small in relation to the large protocerebrum, probably due to the reduced number of ommatidia in the compound eye and hence fewer enter- ing nerve fibers. Only a slight constriction at each side between the brain and the large suboesophageal ganglion indicates the ["losition of the circumoesophageal con- nectives, and the ganglion and brain to- gedier make up an essentially continuous, broad, thick ring through which the pharynx passes (Fig. 30). From the lower surface oi the suboesophageal ganglion, Tin; Crane Fly Gi-nus Chionea in Noktii Amlrica 79 nerves were traced to the maxillary palps and to the labella. No nerves other than these ant! the antenn.il nerves were exam- ined in an\ detail. The pumping a[i[xiratiis ot the anterior digestive liihe, tc)i;ether with its mtiscula- tnre, occupies most ot the remaining sjiace within the head, h'rom the h\'[io[iharyn.\ the digestive tube [iroceeds ii(n\-ard in the form of the rigid hasi[-)harynx, the sides of which are strongly sclerotized (Fig. 24). Ihis jcjins the less strongly sclerotized pharyngeal [)ump, which is roughly tri- angular in cross-section (Fig. 30), con- cealed for most of its length by the brain and suboesophagcal ganglion, and en- larged at the ends, where the muscles that o[K'rate it att.ich. F'rom a membranous potich at the junction of basipharvnx and h\|iophar\'n\, where it enters the preoral cavity, the common salivary duct extends back along the ventral side of the head ca[isule, dividing just belore the neck, one branch passing up along each side of the nerve cord as it comes off the suboesopha- gcal ganglion (Fig. 24). F"or ease of reference frtjm text to fig- ures, I have numbered the major muscles 05 mm m-i7 Fics. 2-4-27, Chionea alcxcindriana, details of head and in.uithparts. 24, internal structures of head, parasagit- tal section; bph— basipharynx, br— brain, ejih— epipharynx, hph— hypopharynx, Ibr— labruni, m— muscle (sec text for origins, insertions and actions of nuiniiercd muscles), oes— oesophagus, php — pharyngeal pump, pmt — prementum, sgn— suboesophagcal ganglion, ski— salivary duct. 25, basipharynx and hypopharynx, ventral aspect; dp— dorsal plate in membranous wall, for attachment of muscle 12. eph— epipharynx, heph— hinge of cpiphar- ynx, hph— hypopharnyx, m— muscle, ski— salivary duct. 26, maxilla and basal segments of maxillary palp, an- terior (inner) aspect; in— muscle, sti — stipes. 27, lalMum, in part, anterior aspect; lab — labial palp (labellar lobe), m — muscle, pmt — prementum (prementum and mentum-). Scale: Fig. 2-1. 80 The University of Kansas Science Bulletin found in the head capsule and m()Uth[xirts and have Hsted them below by number, giving for each the origin, insertion and presumed action. Each numbered muscle or group of muscles is paired except where otherwise indicated. 1. labruni to oblong sclerotized plate in the thin, almost transparent epi- pharynx; elevates epipharynx, distend- ing cibarium (Fig. 24). 2. posterior clypeus alongside mid- line to end of oblong sclerotized plate of epipharynx; elevates epipharynx and draws it backward. 3-4. stipes (or sclerite in which it is embedded) to first segment of maxil- lar\- palp; respectively elevate and de- press palp (Fig. 26). 5. head capsule below eye to distal end (jf stipes; adducts palp. (Abduction may be brought about by resilience of the stipes and associated sclerite.) 6. dorsal and ventral inner surfaces of first palpal segment to proximal rim m-i7 m-i8 29 antn G.W.B. m-20 31 m-2i Fics. 28-31, Chioticu alcxiinihiunu, (Itl.iiK of head and anterior (.cntral nir\ous system. 28, internal (postcro- vc-ntral) as|)cct of frons; atp — anterior tentorial pit, ce — ennipound e\e, cs — ciiistonial suture, ice — inner surface of compound eye, m — muscle (see text for origins, insertions and actions oi nuinheretl muscles), sc — scape, tn — tentorium. 29, brain and anterior nerve cord, dorsal aspect; ol — optic lobe ot brain. 30, brain (br) and sub- oesophageal ganglion (sgn), posterior as|H-ct; nc — position ot attacliimnt ot ntr\c cord, php — phar\ngi-.il pump, in cross-section. 31, suboesoph.igeal ganglion and brain, \entrai aspect; antn — antennal or antenn.uy nerve; ice — inner surlacc of compound eye, m — muscle. Till-: Crane Fly Gincs Chionea in North America 81 m-29 m ■31 dv vn fb nipt nn-48 Fig. 32, Chionca alcxaiulriaiia, internal stnuturts of thorax and alHlonicn of adult male, composite of parasagit- tal sections; agnS — abdominal ganglion of fifth segment, as — anal (tenth) segment, at — aorta, cp — crop, tiv — dorsal vessel (heart), fli — fat body (|)ortion along dorsal vessel), hg — hind gut, or intestine, Ism — longitudinal sternal muscle, m — mu--cle (see text for origins, insertions and actions of numbered muscles; ni-28 originates on mesothoracic furca; m-il originates im metathoracic turca), m|)t — malpighian tubule, ocs — oesophagus, pm — pleural muscle from ninth sternum to ninth tergum (largely concealed by m-45; its outline indicated by dotted line), rp — rectal papilla, rs — reproductive system (enclosed in irr}>. dorsal surface of coxa to spatulate tendon attached to anterodorsal td^'^^o. of base of trochanter; elevates trochanter. 34. posterodorsal surface of coxa to small, disk-shaped tendon just above posterior condyle of trochanter; elevates trochanter. 35. posteroventral surface of coxa to ventral rim of base of trcnrhanter; de- presses trochanter. 36. anteroventral surface of coxa to cedv mpt spth egg ,. fD ^905 Fig. II, Chionea akxandrianu. inttrn.il Mructures of nclult fcm.ilc. composite of parasagittal sections; acg — accessory j^land, at — aorta, ccdv — canli.ic enlargement of dorsal vessel, covcl — common or median oviduct, cii — crop, i usually in the anterior sec- (jnd segment, the last in the seventh seg- ment, and the others spaced approximately as shown in Figures 32 or 3^. Each of the first h\'c gives rise to a posterior pair of major nerves and an anterior pair of very slender nerves, all extending laterally and branching onto the body wall. From the back of the metathoracic ganglion, a pair of nerves resembling the larger ones from an abdominal ganglion arises medioventral- ly and extends into the first abdominal seg- ment, suggesting fusion of the first abdom- inal and last thoracic >ranglia. The terminal abdominal ganglion, which varies from two to three times as broad as the others, gives rise not only to slender anterior nerves but two [xiirs of larger nerves extending back- ward to the muscles of the genitalia, again suggesting the fusion of two or possibly three ganglia. In females (Fig. M) the larger nerves from the last abdominal ganglion are of aj^proximately ec]ual diam- eter, but HI males (Fig. 32) the first or outer \^^\r are about twice the diameter of the second [xiir aiul lead into the basi- stvles. None of the nerves was traced be\()nd its basal [lortion. This description of the central nervous system is in fair agreement with that given by kJrauer et al. (1S54: b'lg. 7) except in the structure of the brain and suboesophageal ganglion. Till, Cranl Fly Ckni-s Chionea in North Amurica 87 Extending backward through the tho- rax from the [ihar\iigcal pump is the s'ender oesophagus, which joins the \ent- triculus, or mid-gut, near the anterior end of the abdomen (Fig. ^T) or farther ceph- alad. Beneath the anterior eiul ot the oeso[ihagus, in the lower prothorax near the tirst thoracic ganghon, are the two irreguhirlv shaped, notkilar sahvary ghinds (sgl). The ventriculus (vn) is thick- walled and muscular. It is usually some- what constricted near its mid-length, but in various individuals other constrictions ma\- also occur, the shape of the organ [probably reflecting its physiological condi- ticMi at the time of death and preservation. In a female whose abdcjmen is nearly filled with eggs, the ventriculus ma\' be displaced forward and the hind-gut drawn more (jr less straight (Fig. })}>). Just before the junction of the oesophagus and ventriculus, the crop joins the digestive tube ventrally. It is a large, membranous, rather amor- phous {jrgan, extending backward beneath the ventriculus, the posterior cwA t)tten bent back cephalad. Although it is not easy to see how Chionea can take any food other than litjuids, I have sometimes found rust-colored or greenish particles of solid material in the crop, as well as in the oesophagus. These mav be precipitates, however, as they seem to have no cellular structure even at high magnifications. Immediately behind the ventriculus, at the [M'octodeal valve, four Malpighian tu- bules (mi)t) are given o(T, two dor.solater- ally and two with their bases close together ventrally. In [ireserved specimens these have a laint puriilish color throughout most of their length but are whitish in the basal one-fourth, which portion of the tubule is also more slender than the rest. 1 ney are subetjual in length to the thorax and abdomen together and are usually in- tertwined among the viscera, although the a[iical portions may often be found King atop the other organs, adjacent to the rectum or the dorsal vessel. The hind-gut (hg) is a fairly simple tube, enlarged only slightly at its anterior c\u\ but constricted just behind the attachments of the Mal- pighian tubules. Less thick-walled than the mid-gut, it may have a slightK annu- lated .i[ipearance due to the muscle arrangement. The rectum is situated dor- sallv, close beneath the seventh and sue- dtt i /rnilliiiitiinmihiiintniiimiith | 38 CivS. Fk;.s. 3S, 39, Chionea alcxanilnaiia, details of tr.iclical system of .ulult. ?>>^, diagram of major tracheae in typical abdominal scj^nicnt, left side (anterior at right), interior aspect; ilc— tlorsal commissure, dtt — dorsal tracheal trunk, s— sternum, sp— spiracle, t— tcrgum. 39, detail . f tracheae adjoining an abdominal spiracle (positional relationships distorted by mounting on microscope slide); dtt — dorsal tracheal trunk, Itt— lateral tracheal trunk, sp— spiracle, v— valve (scleroti/.ed porti;m only), vlv— ventral branch (to sternum). 88 The Univirsity of Kansas Scikn-cl Bulletin ceeding terga. It is membranous and semi-transparent, with four conical rectal papillae (rp) attached near the anterior end, their vertexes usualK' directed some- what caudad. The luimher of these pa- pillae has been reported as varying with sex in some nematocerous Diptera, but in Chionea it is four in both males and females. I have not studied the tracheal system in detail, but the general pattern of it can be described (Figs. 3S, 39). From the thoracic spiracles (sp) a large dorsal tracheal trunk (dtt) forms on each side and extends back\\ard to the seventh seg- ment, where it narrows abruptly and branches into small tracheae. Dorsolateral in general position within the abdomen and closely applied to the body wall, it bends downward above each abdominal spiracle to receive an incoming trachea from the spiracle, then rises again near the intersegmental line, thus forming a zig-zag pattern. The smaller lateral tra- cheal trunk connects the spiracles directly but bends downward between them, giv- ing off a branch at the lowest point (Fig. 38). There is also a ventral branch trachea, so that in all there are four major branches entering from each abdominal spiracle (Fig. 39). These give oil lesser tracheae, but the detailed pattern of branching varies even from one spiracle to the next in the same individual. C'ommissures (dc) con- nect the dorsal tracheal trunks in each segment. One trachea enters each leg, that of the front leg coming rather directly trom the mesothoracic spiracle, that of the hiiul leg from the metathoracic, and that of the middle leg apparently lormed by convergence of branches from both spir- acles. In the tibia the tracheal diameter is as great as in the femur, but it dimin- ishes in the tarsus. Two pairs ol tracheae enter the head, supplying air to the an- terior central nervous system and the tced- ing apparatus. Each ot the abdominal spiracles opens into a short atrium, which is lined with hairs and which may be closed internally by a C'-shaped sclerotized valve (Fig. 39). Mide Rcproditctive System. — Ikauer ex- amined the reproductive anatf)my of Chi- onca araneoides more than a hundred years ago (Brauer et ak, 1S54: Figs. S, 9), but in his illustrations he omitted the sclero- tized structures and misidentified the sf)fter parts. As far as I a:n aware, no one has since st tidied the internal structure of Chionea, so these errors have never been corrected. In another paper (Byers, 1*^61/7: 696-700) I have described the reproductive system in a tipuline crane fly, Dohchopeza, and although this at first appears to difTer extremely from the arrangement in Chi- onea there is actually a part-for-part corre- spondence, except for certain muscles. Accordingly I shall make occasional com- [xirisons of the two genera in the following discussion. Perhaps the internal reproductive struc- tures in Chionea can best be understood by proceeding from the external genitalia already described. The genital chamber (Fig. IS) is the space between the basi- st\les, bordered above by the venter of the tenth segment (slO) and below by the ninth sternum (wgc). Its ventral wall or tloor is membranous, but the anterior wall is strongly sclerotized, except at the edges adjoining the basistyles. The dorsal por- tion oi this sclerotic region, from just be- neath the tenth segment down to the base of the penis, is formed by the fusion of the bases ot the gonapophyses. In Chionea, the vesica, a small sclerotized bulb from which the penis originates, lies at the sur- face and contributes to the anterior wall of the genital chamber. Extending down- waicl and to the sides irotn the vesica are its two immovable lateral apodemes (la) (cf. 15yers, 1961^: 699). Fusion of these apodemes along the mid-line is complete in some species ot Chionea, such as alexan- dnana, but there may be a partial separa- HI- ("rani. Im.y (]i.Nr.s Chionca in Noktii Ami.kica 89 tion leaving; a membranous area just hclow the base ot the penis, as in stoneana. On the lateral apodemes originate the thick muscles that activate the compressor apodemc (FIl^. IS, cmca). in Do'.ichopeza 1 called this compressor apodeme of the vesica the dorsal a[x)deme (Hyers, VXAb: d''''), but in that mentis the vesica antl all its a[iodemes have rotated inward, main- taiiuni; their clcjse association \\ ith the wall oi the genital chamber oid\- by way ol a thin, membranous [iouch that is an exten- sion cjf the chamber. 1 he motle of action of the compressor apodemc is discussed in the reterence just cited. There is at cither side (jf the median keel of this movable apodcmc a broad extension, variously curved according to the species, which serves tor the insertion oi the compressing mtiscle on one side, and oi the retracting muscles that expand the vesica on the other. '1 he latter muscles originate on a second pair oi relatively fixed apodemes that diverge ceiihalad from the vesica in Chionca (but are directed posteriorly in Dolichopezu, on account of the rotation G.kVfl 40 41 Figs. M). 41. Chionca alcxandiiaiia, r(-i)r()(lucti\ c system of m.ik-. 40, dorsal aspect. 41, ripht lateral aspect, or.uans of ri.u'ht side removed. /\hbre\ iations: ac.t;— accessory gland, gon — ponapophysis, p — penis, sd— semmal duct, sv — seminal vesicle, ts — testis, vgl — vesicular gland, vd — vas deferens. 90 The University of Kansas Science Bulletin described). These apodemes are connected b\' muscles (mg) to the adjacent ninth tergum and to the dorsal interior of the gonapophyses near their bases, and each receives a muscle originating at the side of the ninth tergum. The gonapophyses are only slightly movable. Continuing inward from the vesica is the short, thick seminal duct (sd) (appar- ently the morphological ejaculatory duct), which leads to a small, thin-walled seminal vesicle (Fig. 41, sv), more or less median in position, in the seventh abdominal seg- ment or thereabouts. At the sides of the vesica there are small, flattened pouches (vgl) connected around the anterior edge of the vesica to the seminal duct. While I could not find any evidence that these pouches are glandular in Chionea, their position suggests that they are homologous with the vesicular glands of DoUchopeza. There are two voluminous, dull whitish accessory glands (Figs. 40, 41, acgl), one lying at either side of the seminal vesicle and emptying into the vesicle by an ex- tremely short tube behind the gland's mid- length. From their points of attachment the accessory glands may extend posteriorly far into the ninth segment and anteriorly as far as the fourth or even the third abdominal segment. Their shape and size are extremely variable and probably fluc- tuate according to their physiological state and the volume of their contents. The glands may be elongate and thrown into one or two loops, or compartively short and thick, and the two in one individual are not apt to have even approximately the same shape (cf. Fig. 40). Their diameter also varies greatly, constrictions a^ipearing here and there, so that the glands may seem to be divided into a number of com- partments. No doubt this is what led Brauer to identify the anterior ends as the testes and the posterior ends as "p, tired appendages" of the scmin.il vesicle. The vasa deferentia enter the seminal vesicle a little cephalad from the openings of the accessory glands. Although these tubes are quite slender where they join the vesicle (Fig. 41), they enlarge greatly in diameter as they curve upward over the accessory glands and remain distended for a major portion of their length. This enlarged part of the vas deferens (vd) is usually found packed with bundles of sperm, giving it a glistening, silky appear- ance, and prcjbably is the functional semi- nal vesicle (cf. Snodgrass, 1935: 56S) for, as mentioned earlier, that organ is very small. The thickened portion of the vas deferens ordin.irily extends far into the ninth segment and often well into the basistyle. At its caudal extremity the vas deferens is abruptly narrowed, becoming a thread-like tube, which then reverses direction and passes back out of the ninth segment, terminating in a small, mem- branous sack, the testis. In his study Brauer overlooked the thin apical portions of the vasa deferentia and did not identify the sperm-laden enlargements. The rela- tionship of parts of the male reproductive system is not easy to describe, but Figures 40 and 41 may make the above description somewhat more comprehensible. Most of the muscles in the ninth seg- ment are those of the genital appendages. On each side, however, there is a super- ficial sheet of muscle (Fig. 32, pm) con- necting what might be termed the imagi- nary lateral margin of the ninth sternum with the equally imaginary lateral margin of the ninth tergum. Since the ninth seg- ment is a continuous, well sclerotized ring, the function of these "pleural" mus- cles is not clear. Their action would be to draw the ninth tergum downward, some- what flattening (depressing) the entire segment. The origins, insertions and ac- tions of the other major muscles (Figs. 16, il) are 44. anterolateral margin of ninth tergum to anterior base of basistyle; draws basistyle upward (forward). The Crane Fly Genus Chioncu in North Amirica 91 45. anterolateral margin and pleural region of ninth segment to anterolateral base of basistyle; abducts basistyle and draws it forward. 46. ventral and lower lateral ninth seement (sternum) to (Posterior base of basistyle; draws basistyle downward (backward). 47. mesoventral ninth sternum to mesal margin of base of basistyle; ad- ducts basistyle. Note: this is a large muscle, apparently responsible for the fact that preserved specimens of Chi- onea nearly always have the basistyles closed together, that is, drawn inward. 48-49. anterior and anteromesal basi- style to anterior base of dististyle; draw dististyle inward and forward, or "close" dististyles. 50. posterior basistyle to posterior base of dististyle; moves dististyle out- ward and backward, or "opens" disti- style. In most respects the male reproductive system in Chioncu is simjder than that in the Tipulinae, exemplified by DoUclwpeza. Examjiles of its presumably less specialized design are the undivided dististyles, the simpler basistyles, which are not lused to the sternum, the non-rotated vesica, the straighter, shorter seminal duct and the wholly separate vasa delerentia {d. layers, l%l//:Figs. 17, IS, 27). Female Reproductive System. — As in other crane tfies I have studied, the repro- tluctive system of females of Chionea con- sists of two separate parts: the bursa copulatrix and associated organs above, for reception and storage of sperm, and the ovaries and their ducts below. There is no internal connection between these two parts, but the external opening of each enters the poorly delimited genital cham- ber, as described earlier under external morphology, and to that extent they com- prise a unified system. The common or median oviduct is rather short, extending from the gonopore to just cephalad of the point where the /?^ Fk.s. 42-4S, co.,iparat>vc ckta.ls of rcprcluctivc system of fcmaU-. -12, Chwnca .toncana^ ^cmlA sternum, bursa copuhunx and spcrmatluval ducts, antcrodnrsal (mtcTH.r) aspect (sc.lc line 0.1 Chwnca lyrata. bursa copulatnx and bases of spermatlucal ducts, anterodorsal aspect AA. Unor^ca Kenital furca, nmtfi sternum, bursa copulatrix and apparently glandular structures (gl (cf. "ig. dorsal (mtenor) aspect. 45, Chwnca mvicola, burvi copulatr.x and bases of spermatlucal ducts, jett Scale a: Figs. 43-45. G wB. furca, ninth mm). 43, alcxandnana, 22), antero- lateral aspect. 92 The University of Kansas Science Bulletin base of the furca is anchored in the dorsal wall of the oviduct (Fig. 22, covd). Al- though thin-walled, it is muscular, liaving mostly transverse but also longitudinal muscles. The lateral oviducts (lovd) are extremely short as ducts alone, but each is continuous with the central lumen of its respective ovary, and the wIkjIc may be regarded as an elongate lateral oviduct giving ofif ovariolcs in a radiating pattern along almost its entire length (Fig. 19). A female that has not commenced oviposi- tion may contain as many as 200 eggs, the ovaries extending into the first abdominal segment and often even crowding into the thorax. A female of Chionea nivicola, for example, was found to contain 126 appar- ently mature eggs, of which eight were within the thorax (Fig. 48), extending even into the proihorax. The ovaricjjes are short, membranous and transparent, and each contains but a single egg. The older, more mature ovarioles are rounded api- cally, conforming more or less to the shape of the egir within, but toward the anterior end of the ovary most ovarioles contain in addition to the egg one or two smaller clusters of trophocytes, so that they are tapered at the apex (Fig. 19). In some other tipulids I have dissected, the dis- tended ovaries had pushed the digestive tube aside, against the body wall. Perhaps it is sometimes so in Chionea. Inii in the females examined the digestive tube was found in the median plane, somewhat com- pressed between the ovaries, which being full of eggs bulged outward but were flattened medially. Within the ovariole the egg lies with its anterior end (i.e., the end bearing the micropyle) away from the lumen of the ovarv. The egg moves into the oviduct in this same position and thus is deposited posterior end first. Eggs in the dorsal part of the ovary are inclined downward, those in the ventral part upward, and so on, suggesting an inclined radial arrangement of the ovarioles around the lumen of the ovary (lateral oviduct). The eggs in both live females and those preserved in alcohol are whitish to very pa'e tan, those toward the anterior end of the ovary palest. They will be described later in more detail. From its opening below the ninth ster- num, the bursa copulatrix descends along the inner surface of the membrane between the branches of the furca (Figs. 22, 42, 44). It is only slightly curved cephalad near the base of the furca in some species (e.g., Chionea alexandriana, Fig. 22) but makes one complete loop in others (as C. scita. C. nivicola, Fig. 45). Approximately above the base of the furca there is a sclerotized thickening of the cuticular lining of the bursa, at which point the spermathecal ducts are given ot?. The lining of the bursa may be more extensively sclerotized and darkened, the degree depending upon both the species and the individual. In C. valga, for example, the bursa is very darkly sclerotized for most of its length. Unexpectedly, the number of sperma- thecae was found to be either two or three. I would have expected a uniform number within the genus, but in females of C. scita, C. nivicola and C. stoneana there are three, while in C. alexandriana there are only two. The number of spcrmathecae could not be determined for every species because of a lack of properly preserved specimens in some cases; however, the bursa is ordinarily sufficiently sclerotized that it can be examined in cleared, slide- mounted specimens. It appears from such examinaii(jn that the arrangement found in alexandriana (i.e., two spcrmathecae) occurs also in lyrata (Fig. 43), nigra, ex- cavata, and other species of the alexandri- ana grouji. W hen there are only two spcrmathecae, the ducts open dorsolaterally on the burs,i; when there are three, the third is medioventral in position. Also, when there are but two spcrmathecae, the ducts are large and thick-walled, while in species having three spcrmathecae they are thin-walled and slender. The ducts are The Crani; Fly Gknts Ch'ionea in North Ami.rica 93 cither well sclerotized hasally, as in stone- ana (Imi;. 42), or contain spiral, ctenidia- likc rcintorccincnts near the base, as in the ah'xandnana i;roiip (Imi;. 4.-!). The spermathecae are not sclerolizccl and of hxed size as in some other crane flies, such as Dolichopcza (Byers, 1%!/^: 701), hut have nienihr.moiis or semi-mein- hranoLis (i.e., somewhat thickened yet translucent) walls and varv in size accord- ui>; to species, as well as with the volume ot sperm contained. The female illustrated in Figure 1'^ for example, had distended spermathecae which were packed with sperm, Init in some other females of the same species the spermathecae were small and wrinkled. The position ot the sperma- thecae is highly variable, and the ducts are usually irregularly curved or even looped. From the point at which the sperma- thecal ducts enter, the bursa copulatrix continues cephalad, terminating in a [louch (Fig. 1*), fspth) that, like the spermathe- cae, has membranous to semi-membranous walls. It is large in females of the alex- andriana group (the actual extent deter- mined only in alexandriana but interred from the part visible in slide-mounted specimens of the other species) but small to minute in other species. This appears to be the morphological equivalent of the "functional spermatheca" of Dolichopeza (Hyers, 1961/?: 703). In Chionca. however, I have never found any s[X"rm in this pouch, while the spermathecae may be filled with them, as in the specimen illus- trated (Fig. 19). The walls of this pouch are finely rugose, with abundant hair-like filaments extending into the lumen. The contents in preserved specimens are sparse aggregations of minute particles. Since the concentration of spermatozoa in the sper- mathecae is as dense as that in the vasa deferentia of the male, leaving virtually no space for any fluid carrier such as is probably produced in the male's as- cessory glands, it may be that the fluid portion of the semen received is stored in the [louch at the end of the bursa. Whatever its 1 unction, it is in some spe- cies a structure of considerable capacity situati'd where it is likeK' to be involved in the reception and storage of some part ot the semnial lluid. There are two accessory glands which em[ity b\' slender ducts (sometimes envel- oped in a common sheath, basally, as Fig. 21) into the bursa just inside the aperture. These var\ in size and general ap[iearance from one individual to the next, being sometimes turgid and consjiicuous (Fig. 19), other times so small and wrinkled as to be easily overlooked. Such difTerences are probabh' indicative ot the physiological state of the organs at the time of preserva- tion, the glands possibly being distended shortly before oviposition and collapsed after a batch of eggs has been laid. Closely app'ied to the membrane between the f ureal arms and lying one at either side (jf the bursa are a pair of elongate, sub- hyaline, seemingly hollow structures (Figs. 22, 44, gl) of unknown tunction. These are usually very difficult to see. They ap- pear to have no attachment to the bursa but may open to the outside near the ends of the arms of the furca. I have designated them "glands" (gl) with doubt. Nearly all the major muscles in the eighth through tenth abdominal segments of the female are concerned with the depo- sition of eggs. Since I have not observed oviposition in Chionca and consec]uently do not know how the valves of the ovi- positor move, I can only speculate on the function of certain of these muscles. Thei. origins, insertions and actions (see Fig.' 20,33) areas follows: 51. dorsolateral region of eighth tcr gum to anterodorsal corner of eighth sternum; elevates anterior c\\<\ of eighth sternum. 52. lateral ends of eighth tergum to side of eighth sternum, below margin and beside gonopore; elevates eighth 94 The Univkrsity of Kansas Science Bulletin sternum. Ncjtc: it cannot he surely said which (jf the described attachments is the orii;in and which the insertion. 53. anterior eighth sternum to Literal portion of ninth sternum; draws ninth sternum dcnvnward. Note: muscle pairs 51-53 probably function to bring the gonopore and the opening of the bursa copulatrix into proximity so the eggs can be fertilized at the time they leave the oviduct. 54. side of eighth sternum to oppo- site side, passing just beneath and be- hind common oviduct; draws sides of sternum and therefore bases of hypo- valves together. Note: this unpaired muscle together with pairs 51 and 52 may be involved in extrusion of the egg from the posterior end of the oviduct. 55. posterodorsal eighth sternum to anteroventral hypovalve; elevates hypo- valve. 56. dorsal ninth tergum to base of cercus; depresses cercus (see also m-5S). 57. dorsolateral ninth tergum to lat- eral ninth tergum near attachment of ninth sternum; elevates ninth sternum. 58. dorsal tenth tergum to base of cercus; depresses cercus (see also m- 56). 5'^. transverse, infolded ridge on tenth tergum to upper base of cercus; elevates cercus. Note: the elevators of the cerci are very small, while the de- pressors are large, indicating that the cerci are probably drawn down against the hypovalvcs duruig oviposiiion. 60. lateral tenth tergum to lateral tenth sternum; elevates tenth sternum. There are some lesser muscles associated with the turca, including those between the furcal base and the oviduct (Fig. 22, mfb). The membranous body wall at each side of the pouch (b'ig. 22, pch) between the furca and the distal roof oi the oviduct also contains a wide, thin sheet of muscle attached to the lurca. Natural History of Chionea Ecological Distribution Although there is a slowly growing bcjdy of knowledge about the underground habitats of Chionea, the fact remains that most of what we know about these flies is based upon their occurrence on the surface of snow. Against the white back- ground of sikjw even the palest Chionea is conspicuous, especially when moving. In contrast, the flies would be easily over- looked in the winter surface vegetation, which is largely dead and dried to dull browns and grays. On the surface of snow, species of Chionea may be found together with such other winter insects as Boreiis sp[-). (Meccjptera: Boreidae), various apter- ous, brachypterous or even fuMy winged stone-flies (Plecoptera: Nemouridae, espe- cially Capniinae), gall wasps (Hymenop- tera: Cynipidae) of the agamic generation, several kinds of snow-fleas (Collembola: Poduridae and Isotomidae), certain Chiro- nomidae, Anthomyiidae, and a few taxa of non-insect arthropods, notably spiders. By "found together," I mean they occur in the same general environment. Insects on the snow are usually widely dispersed, and I have never seen any direct interaction of a Chionea with any of the other kinds of arthropods mentioned. Activity of winter insects on the snow seems to be limited mainly by temperature and to a somewhat lesser extent by wind and blowing snow. Light intensity may also be an imjiortant factor in the case ot (Chionea, as will be discussed later, but its elTect is not readily separated from that ot tem[)erature. While the relationship of such physical factors of w inter environment to the distri- bution and abundance of various insects has been examined bv several authors, few ot these studies have direct bearing on the biology of Chionea. ('old hardiness in some Euro[Kan winter insects, including Chionea arancoides Dalman, was studied The Crane Fly Genus Chionca in North America 95 by Somme and Ostbye (1969). They found this Chionea active chiefly at air tempcra- lurcs of 0 to -4°C', occasionally at -7°C (near 20"F). They determined that it is able to avoid the lethal efTects of freezing of its hacmoKmph by sLipercooling to -7.5°(1 b'rom his held observations of C. arant'oides, Hagvar (1971) determined -6°C to be the lower limit of activity and found the flies most commonly on the snow at air temperatures between -4 and -5°C' when there was no appreciable \\ind. He seldom saw this species on the snow at even slightly higher temperatures (-3 to 0°C') and concluded that their activity is very narrowly limited by temperature. Wojtusiak (1950) found Chionea (Iiites- cens and " hraher) active primarily around -3.5°C. Hezzi (1919) reported Chionea spp. active at temperatures down to -6°C. Chapman (1954) similarly found some North American Chioneas (mostly C. alexandriana) active on snow at air tem- peratures in the range of -7 to 0°C but mainlv between -4 and O'^C. He also found C. alexandriana in cold stupor at -7.5°C (19.5"?). His observations are in close agreement with those of Scimme and Ostbye. Throutihout the range of Chionea, win- ter temperatures far below this tolerable level of approximately -7°(^ are frequent and often of many days' duration. It is accordingly imperative that the Hies have ready means f)f escape from severe cold. Routes for movement trom the snow sur- face to the soil are easy to {\n^\ when one is in the natural habitats of Chionea. They may result from accumulation of gently falling snow on low vegetation, otten leav- ing adequatelv large openings to the soil. Others occur around large trees and rocks, where surface wind may arrange the snow in such a way as to leave large, tunnel-like depressions leading to the soil or nearly so. Winds also move brush, saplings and other vegetation protruding above the snow sur- face from side to side, with the result that around the stems of such j^lants there are narrow chaimels leading down to the ground. 'I he insulating efTect ot snow cover on the scjil cannot be overemphasized. Mail (1930: 579) showed that temperature just below the soil surface (habitat of larval Elatcridae, (jr wireworms) Huctuatcd only about 4°C: (-1 to 3°C) over a month of observations (January), while air tempera- ture above the snow varied from -2 to -23"(- around a mean of about -14^C. He also demonstrated (p. 577) the cocjling elTect of air temperature on the soil in the absence of sncnv cover. Coulianos and John- els (1962) recorded similar results for the subnivean environment ot small mammals. (The subnivean air space results when snow is held away from the soil surface by grasses and other low vegetation; it may be enlarged by the activities of mam- mals or perhaps by heat generated by bacterial decomposition of leaf litter or other dead vegetation.) These investiga- tors found that ewer a 14-day period in March in Sweden, the temperature in a 5 cm subnivean stratum beneath 20 cm of snow varied only from 0 to 2^C, while during the same period daily ma.ximum temperatures fluctuated between -6 and 0.5°C and niiihtlv minima between -4 and -23.5°C. In another study of micro- climates to which small mammals are subjected, Hayward (1965) found that in burrows of Peromysciis mice in al[iine, sub- alpine, mesic antl arid habitats, winter temperatures never fell below O'^C^. (.Al- though it is not [xirtictilarly relevant here, Hayward further found that in summer, when air temperatures during a day might fluctuate 15.2 degrees, from 4.6 to 19.(S°C, the temperature in representative mouse burrows remained very close to 10 C.) 1 have matle similar comparative meas- urements ot temperature in the soil, in the snow cover, at the snow surface and in the air several centimeters above the snow surface. Mv findings, while onlv made 96 The University of Kansas Science Bulletin on certain days and in various winters, not over a span of many days, agree with those presented here. By seeking refuge in soil cavities, in rodent burrows, or in the subnivcan air space, Chioneu can not only escape severe cold at the snow surface but can in fact en- joy a habitat where the temperature is rela- tively stable, probably near the optimum for the adults, and where the humidity is relatively high. In view of these efTects of snow cover, one wonders if Chionea stone- ana, our least boreal species, might not be also our most cold-hardy Chionea, living as it does in a mid-continent region where winter temperatures are often very cold but where snow cover is frequently absent. Where Chioneas are and what they are dcjing when they are not on the snow sur- face is only sketchily known. Chance obser- vations by a number of authors have ap- peared chiefly in the European literature. Bezzi (1919) summarized ()7 collections of Chionea made up to 1917 in Europe and Asia. He recorded six collections from caves (two from France, four from Jugo- slavia), two from vacant underground wasp nests in IV'lgium and the Nether- lands, and one from the burrow of a meadow vole {Arvicola). Bezzi did not regard Chionea as a cavernicolous insect but supposed that the flies had only acci- dently wandered into caves in search of smaller cavities in which to conceal them- selves. There are sufHcient records from caves in the United States, however, to convince me that Chionea, while perhaps not a cavernicole in the strict sense, is more than an accidental visitor in caves. There have been a few subsequent discoveries of Chionea in European caves (e.g., Strinati and Aellen, 1967). In a study of cave insects, Dr. M. S. Briscoe of Howard Univ. set a number of pitfall traps (glass jars) in the floor of a cave near Harpers Ferry, West Virginia. These were baited with decom[X)si ng meat and were intended to attract primarily cave-dwelling Coleoptera. In addition to the beetles and various other insects. Dr. Briscoe trapped 29 specimens of Chionea sctta, a larger c(jllection of this species, by far, than any ever made from the snow surface. The traps in which C. scita was found. Dr. Brisccje informed me, were set lUO ft (31 m) or more inward beyond the border of total darkness in the cave. During summer months crane flies are not uncommonly found in the entrances to caves, that is, in those parts of the cave that alTord shade from the sun and in- creased humidit)' but are not isolated from the more normal environment of the flies; but such crane flies rarely venture into a cave beyond the zone of deep twilight. Cjraham {YH)()) has given an account of this, as well as of the occurrence of the trichocerid flies Diazosma sitbsinitata (/Mexander) and Trichocera columbiana Alexander in the zone of total darkness. I have occasionally found large numbers (it moscjuitos and fungus gnats hibernating in parts of caves that were totally dark yet not a great distance from the opening. It is likely that some fungus gnats (Myceto- philidae) can breed in caves, where such organic debris as bat guano accumulates on the floor to provide adequate substrate for the growth of fungi which in turn nourish the larvae. While the diet of larvae of Chionea scita is not yet known, there is a strong likelihood that it is comprised in [xirt of organic debris of various sorts. It is thus not improbable that the popula- tion sampled by Briscoe had become estab- lished in the cave. Mr. Roger Sperka (pers. comm.), then of the Univ. of Kentucky, collected a fe- male of C. scita in Wind C^ave, Pulaski (>)., Kentuck}'. He estimated that the fly was found 700 ft (215 m) from the en- trance ami in a lateral tunnel, thus far beyond the darkest crepuscular zone and in total darkness, where the temperature was 51.5°F (11 °C). However, the single individual had fallen into a small stream Thi: Cranf. Fly Ginus Chionea in North America 97 in llic cave, had clanihcrcd out ai;ain and "seemed disoriented," stj there is reason to suppose it may have strayed irom some area of the cave where a [lopulation of its species mii^ht have hcen estahhshctl, il not from outside the cave. Other records Irom caves involve \sx\- marily western species taken considerably closer to the cave entrances. The flics in these instances could have been accidental visitors. A collection of special interest is that of Chionea alexandriana from an ice cave in Union Co., Orei^on. The single male in this collection of IS individuals has gonapophyses of an atypical form not seen from elsewhere in the range of the species. Underground habitats of Chionea other than caves consist almost wholly of bur- rows constructed by other kinds of ani- mals. These vary in size from burrows of earthworms (Kryger, in Edwards, 1936) to nest burrows of wasps (Bezzi, 1^19) (conceivably enlarged from rodent nests) to burrows and nests of mice and voles. Hackman (10(,1, 1963a, \%^b) has made the most detailed and systematic study of Diptera in excavations made by small rodents. Among some 2000 indi- vidual flies of 240 species, he found onlv one Chionea, a female of C. lutescens. This fly was in a burrow of Clethrionomys glareohis beneath a tree stum[i in southern Finland. Hackman (1963t/) regarded C. lutescens as a xenocoenic species, that is, one that is foreign to the community in which it has been found (in this instance the burrow community) and actually adapted to life in some other kind of habi- tat. Trichoceridae, or winter gnats, were common in his winter samples; he found four species of these (also two of the crane ily genus Ovmosia, but not in win- ter). These he regarded as tychocoenic, generally adapted to the burrow environ- ment but with a wider ecological range enabling them to thrive in other, similar biotopes as well. Hackman concluded that Chionea merely utilizes the vole burrow as a shelter against unfavorable weather in winter, although he pointed out that the dcc[i burrows communicate with runways in the subnivcan air space. In his interpre- tation ol the presence of Chionea and 1 nchoccra in burrows ot Clethrionomys, he [lossibly overlooked (jne aspect of the biolcjgy ot these tlies, that is, larval diet. Larvae of Tnchocera develop in decom- posing organic debris such as may accumu- late in rcjdcnts" excavati(jns, particularlv nests. There is also reason to believe that such nest debris (e.g., feces of mice) is involved in the diet of larvae of Chionea. That is, I am suggesting that Trichocera and Chionea may spend much of their respective life cycles in rodent burrows. The extent to which rcjdent burrows or nests are utilized by North American spe- cies of Chionea is not known. We may make some inferences, however, from cer- tain facts about C. stoneana and C scita. Investigation of arthropods in nests of the prairie vole, Microtiis ochrogaster Wagner, by Philip C. Stone of the Univ. of Illinois, led to the discovery of the specimens upon which Alexander (1940) based the descrip- tion of C stoneana. Fourteen adult males and five females of C. stoneana were collected from cavities beneath large logs, apparently abandoned mouse nests (probabh' ot I'eromyscas leii- copiis), on 2 January, in a sparse oak- hickor\' woodland in northern Franklin Co., Kansas. The nest material upon which the adult Hies were walking, and from which numerous larvae were subse- tjuently recovered, consisted of finely shredded, decayed wood, small fragments of dead leaves and grass stems, and some loose soil [xnticles. Arthropods associated with die ('hioneas included small, active staphylinid beetles and some hibernating, inactive wood cockroaches, ants, lygaeid bugs and a machilid. Further details are found in the account of C. stoneana fol- lowing. A habitat in scjuthern Michigan from 98 The University of Kansas Science Bulletin which I obtained several individuals of C. scita was in many ways similar to that just described. The tlies were in cavities beneath two old, rotting stumps, where the accumulated material comprised shredded or crumbled decayed wood and bits of dead leaves and grasses. Ikjth stumjis were about 3 m from the marshy edge of a lake, about 50-60 cm above the water level. It has been noticed from the time of Bezzi's 1919 monograph onward that Chionea species are found only in environ- ments that are to some extent forested. It is furthermore clear that the particular kind of forest is not a factor in the eco- logical distribution of Chionea at the ge- neric level, nor does it seem to be for any species. Trees, of course, may play a dual role in the environment of Chionea, shading the terrestrial habitat in summer to provide a more nearly uniform tempera- ture the year around, and serving as wind- breaks in winter when Chionea is on the snow surface. In eastern North America Chionea oc- curs in deciduous woodlands, such as the relatively dry oak-hickory or the more moist, boreal-montane beech-maple asso ciations, also in mixed deciduf)us and ever- green forests of eastern Canada. In the western mountains. Chionea mav be lound from the rather low and dry forests of ponderosa [line (western \ellow [line), through the higher zone of lodgepole pine, antl in ihe high alpine forests of spruce antl In-. Within these various kinds of for- ests, Chionea often is found ni cle71) supposed that narrow limits of tol- erance oi temperature and wind led to the simultaneotis appearance of numerous in- dividuals of C. aruncoides on the snow Fig. -46, Chionea jcllisoni, terminal abdominal segments of male (scleritcs shaded) and female, to show relative positions during copulation. Fig. 47, cysticercoid of tapeworm, probably Hymcnokpis .sp., from abilominal cavity of adult female of Clnoncu itoticana. Fu.. -18, Chionea niiicola, female, to show position of eight eggs m thorax in si'"-"<-' '"■"'<■■ available by absence of indirect flight muscles. Scale: Fig. M. 104 The UniviiRsity of Kansas Science Bulletin surface. Repeated observation of mating of the flies on the snow would support the suggestion of varifjus authors that the reason is the bringing together of the sexes. It has also been suggested (Mar- chand, 1917: 150) that Chioneas are visu- ally attracted to each other and find one another more readily because of the con- trasting background afforded by the snow. It is true that they are conspicuous when seen from above against a white back- ground, but if one scans the snowy terrain from a millimeter or so above the undu- lating surface — that is, adopts the insect's view of things — the idea of visual attraction becomes somewhat less convincing. More- over, as discussed earlier under external morphology, the eyes of Chionea probably do not afford much visual acuity. To say that the flies come out onto the snow surface so that males can find females is not a very thorough explanation of their presence there. Where Chionea has been found anywhere except on snow, both males and females have ordinarily been taken together in restricted microhabitats such as mouse nests, cavities beneath logs or rocks, in caves, rodent burrows, aban- doned wasp nests, etc. Surely they are able to locate one another in such confined places. It occurs to me, however, that since Chionea is an insect of low vagility as compared to winged tipulids, and since it does live in groups (probably the progeny of one female) in partially isolated or restricted pockets, there might develop a degree of inbreeding detrimental to the species unless there was some device for bringing about a mixing of genes. Among winged Tipulidae reaching the adult stage in warmer weather, this may be accom- plished by mating swarms (in which, of course, there still may be some inbreed- ing), or by more or less random flight by one or both sexes prior to mating. I regard the appearance of Chionea on snow as partly, at least, a means of effecting genetic recombination. Certainly, the relatively smooth surface of snow is less of an im- pediment t(j the movement of strictly pedestrian insects in search (jf mates than would be vegetation at the soil surface whether snow-covered or not. In addition, it is known that Chionea is a strong walker and may cover substantial distances; ac- cordingly, wandering (jver the snow is probably important to the distribution of the flies into new microhabitats. Mating Behavior Several entomologists have commented on the rapid reaction of males and females of Chionea to each others' presence when placed in the same container. Lucszer (1S96), for example, observed that when a male and female (probably of C. stone- ana) were caged together "... the snow- flies immediately mated, notwithstanding it was several degrees below the freezing [loint." When I placed two males and two females of C. scita in a large vial lying on its side, in the field, they formed two mat- ing pairs almost at once. These pairs sepa- rated during the return trip to the labora- tory, but S(jon after the flies were put into rearing dishes they mated again. During the first three days after capture these same pairs of flies mated at least once daily, again on the fifth and sixth days, and once more on the eighth day. No mating behavior was seen thereafter; however, I was observ- ing the flies only occasionally and probably failed to note all matings. Duration of mating was about 30-70 min but mav be much longer. On the basis of my observations of four species of Chionea, I can state that mating behavior is essentially the same in all. The male approaches the female ordinarily from behind, rising high upon his legs as he does so, with his body ori- ented almost vertically. His abdomen is then curved forward so that the hypo- pygium can grasp the female's ovipositor. Hut instead of facing in opposite directions during copulation, as is usual among crane Tm- Crane Fly Gi-Nus Chionca in North America 105 flies that mate uhilc standing on a surface, with one or both partners' abdomen ro- tated, both Hies face in the same direction with the male's abdomen curved so that the hvpo[n\^iLim is inverted to achieve the usual ti[Hilid arrangement, that is, with the male genitalia rotated ISO degrees in relation to those of the temale (I'ig. 4*)). Mik's description of the mating behavior of a European species of Chionca agrees with this (Osten Sacken, 1SS7: 1Q6)/ The male's dististyles secure a grip in the pleural membrane of the female's ab- domen just ce[-)hr— hypostomal bridge, hph— hypopharynx, md— mandible, mxp — maxillary palp. 118 The University of Kansas Science Bulleti N bars. The maxillae are large, fleshy (i.e., thick but lightly sclerotized), trilobed, and covered with minute pubescence (Fig. 67). The more dorsal lobe is the pal[i and has at its apex what a["(pears to be a sen- sory apparatus, and the narrow, sc'erotic band around its base is possibly the rudi- mentary stipes. The smaller, lower lobes very likely represent the galea and lacinia, and the sclerotized band traversing their common base may be all that remains of the cardo. Ventrally, the mouth cavity is bounded primarily by the lower lobes of the max- illae (Fig. 71) and is closed medially by the combined hypopharynx and labium. The hypopharynx itself is well sclerotized and is connected laterally to the inner ends of the hypostomal bars. It is pilose on its dorsal surface, which slopes downward to a rounded anterior margin. Near the anterior end of each lateral plate and behind the mandibular attach- ment, there is a subcircular clear area in the plate, bordered above, before, and be- low by intense sclerotization. In the living larva and in some preserved specimens, a dark spot may be seen beneath this clear area. This pigmented spot is the eye, which in life has the characteristic purple color and which can be moved about slightly beneath its cuticular covering. Alexander (1920: 952, Fig. 445) drew attention to a peculiar structure on the dorsal surface of the head capsule illus- trated by Brauer et al. (1854: Fig. 5). He interpreted this as a pair of "appendages which are toothed on the outer face" and concluded that "these probably represent part of the mentum, but Ironi Brauer's figures they would appear to lie above the labrum." Brauer's illustration is contusing in certain respects but seems to mc to be purely a dorsal view, the struct tires in question probably being the two halves of the tronto-clypeus. Internal Anatomy.— 0\\\)' the larva of Chionea stoneana was explored internally. so comparisons with other species are not possible at this time, except for a few minor ccjmpariscjns with Brauer's figures of C. araneoides (Brauer et al., 1854: Figs. 3, 4). The larva illustrated (Fig. 61) was immersed in hot water, which resulted in its outstretched condition. In a living larva most of the organs are more con- tracted, thus are wider and not so linearly arranged. Digestive system: The most conspicous structures inside the larva are those com- prising the digestive tube (oesophagus, proventriculus, ventriculus, and intestine) and organs associated with it (salivary glands and Malpighian tubules). The oesophagus is a relatively straight, muscu- lar tube of narrow diameter, leading from the mouth t(j the anterior end of the short, thick-walled proventriculus. Above and beside the proventriculus and oesophagus lie the twcj salivary glands (Figs. 61, 64), which have a large lumen; they are widest anteriorly, tapering and recurved cephalad [Posteriorly, with thin u'alls, formed of large, flattened, polygonal cells containing nuclei of such great diameter that they can easily be seen with the dissecting micro- scope at 130X magnification. In most of the larvae dissected the salivary glands were larye and distended, and I have never seen salivary glands of such comparatively great size in any other crane fly larva. At each side of the anterior end of the ven- triculus there are two elongate gastric caeca. These are twice as long in some individuals as in others, their length possi- bly varying with physicjlogical activity, for in those specimens having the longer caeca there was a greater amount of food in the \entriculus. More than 40% of the length of the digestive tube is the ventriculus which is muscular and relatively smooth-walled, its anterior halt approximately three times the diameter of the posterior half. There .ire four Malpighian tubules attached two at either side of what appears to be the TiiE Crani-; Fly Gunus Chionea in North Amirica 119 [losterior end uf the vciitriciilus, although they are anatomically behind the procto- deal valve. Sometimes these tubules ajipear to be arrans^ed randomly in the haemocoel, but one or two may have their distal ends closely applied to the wall ot the rectum, supposedly to facilitate the excretory func- tion by fairly direct transport of recovered water. The small intestine and large in- testine are poorly dcHmited, short and thin- walled as contrasted with the ventriculus. Notablv absent from the hind gut is the intestinal caecum, which is conspicuous in all herbivorous tijuilid larvae 1 have in- vestigated. The rectum is longer than the rest of the hind gut and of greater diam- eter; it appears somewhat striated because of small bands of muscle in its nearly membranous walls. Tracheal system: The dorsal tracheal trunks are of surprisingly small diameter (Fig. 72), which may relate to the rate of metabolic activity of the larva. Since the distribution of branch tracheae and other parts of the respiratcjry system in Chionea follows rather closely that described for Dolichopeza (Byers/ 1961^; 740-741, Fig. )., 5-6 mi E of Chinook Pass, on siKJW about 2.5 ft deep, 17 Dec. 1977, P. J. Johnson (19); 15 mi E of Chinook Pass, on snow about 2.5 ft deep, 17 Dec. 1977, P. J. Johnson (1 6 ); Yakima Co., Sand Lake, 2 mi N of White Pass, Snoqualmie National Forest, 5295 ft, walk- ing on snow 30-45 cm deep, in coniferous forest, 25 Nov. 1979, W. Kaltcnthaler (16). Chionea macnabeana Alexander Chioncu macnabeana Alexander, 1946: 156; Alexander, 1954: 69; Alexander, 1965: 72. Diagnosis. — A large west coast species of yellowish brown color, with antennae of 12 segments, scape and pedicel thick; female cerci very long and slender; sixth and seventh abtlominal terga bearing sev- eral thick, blackened spines, es[K'cially on posterolateral corners; male gonapophyses appearing broadly rounded with inner apical [loint, in dorsal aspect. Description . — ( General appearance: Head, thorax, legs and abdomen yellowish brown to light brown, head slightly darker than thorax; sclerites not much darker than membranous areas; cerci of female long, amber colored, palest at base. Length of female, exclusive of antennae, about 8.2-9.4 mm. Head: Antennae (Fig. 83) consisting of thick, subcylindrical scape, short, club- shaped pedicel, and ten flagellomercs; fu- sion segment short-conical, or turbinate, shorter than its basal diameter, partially 132 Thl University of Kansas Science Bulletin ^^y^^ 0.5 mm G w.fl Figs. 83-85, Chionea macnahcana. S3, antenna of female holotype (from slide in C. P. Alexander collection). 84, terminal abdominal segments, female from Washin.i^ton, left lateral aspect. 85, ninth sternum and genital furca, female, ventral aspect. Scales: a — 83, 85; b — 84. sulxlivitlcd in one specimen seen; first flagellomcre beyond it short, cylindrical, bearing many short, fine hairs in addition to verticil of three hairs which are shorter than those of succeeding flagellomeres. Antennal length about 1.5 mm. Setae of vertex stout, long, black. Eyes with about 200 ommatidia each. Thorax: Haltcrcs tan, not contrasting conspicuously with color of thcjracic dor- sum. Legs slender (femora not incrassate), yellowish brown, tarsi only a little darker brown. Setae of legs long, stout, black; those in dorsal rows sHghtly larger and thicker than ventral setae. Setae of coxae, thoracic dorsum, and sternal sclerites stout, black. Abdomen of male: See Appendix 2. Abdomen of female: Eighth tergum short (i.e., slender) dorsally with lightly sclerotized anterior expansion at each side (Fig. (S4), posterior margin more darkly sclerotized. liypcjvalves of eighth sternum long, slender, evenly tapered toward nar- rowly rounded apex. Ninth tergum with three short setae near posterolateral margin on each side; tenth tergum mostly bare with two setae on each side near posterior margin. Tenth tergum somewhat humped upward near mid-length, and anteriorly its sides extended and curved ventrad. Arms of genital furca of approximately unitorm thickness throughout, smoothly curved outward to points cjf attachment on ninth sternum; basal portion a little longer than wide, most strongly sclerotized at sides. Ninth sternum (Fig. 85) formed by two slender, well sclerotized bands with expanded tips which almost meet medially. Cerci long, slender, about eight times as long as greatest width, evenly curved both dorsall)' and ventrally to acute tips, noticeably longer than hypovalves (about 1.17 times as long as hypovalves measured from notch to tip). Sixth and seventh tcrga with thick, blackened setae along posterolateral margiiis, most con- spicuous on seventh tergum; sternal setae The Crane Fly Glnus Chionca in North America 133 blackened but generally shorter and more slender than those on dorsum. Types. — Holotype, female, Wilson Riv- er, Tillamook C.o., Oret^on, elevation 2125 ft, S I'ebruary 1^M6, James A. Macnab collector. The tvpe is mounted on a micro- sco[ie slide in the collection of C'. P. Alex- ander, Amherst, Massachusetts (trans- ferred in I'^Sl to the U.S. National Museum, Washington, D.C.). Relationships. — That Chionea macna- hcana belongs to the scita group is indi- cated by the pale body, the relatively large number of antennal segments, and the similarity in structure of the female geni- tal segments between this species and C. albertensis and particularly C. scita. A dis- cussion of the relationships of macnabeana and albertensis will be ft)und under the heading of the latter species. The most useful differences for separating these two western species are the greater number of flagellomeres and shorter, thicker scape and pedicel m macnabeana and the pres- ence of thick, black setae on the sixth and seventh abdominal terga, where in females of albertensis the setae are more slender, not as darkly colored, and more sparsely distributed. Females of macnabeana and scita, both of which may have 12 antennal segments, may be differentiated by the detailed structure of the genital furca and ninth sternum. The furca in scita is some- what more lyriform and the ninth sternum interrupted at the mid-line, with a longi- luduial bar at each side of the break (Fig. •^H). Also, as pointed cjut by Alexander (1946: 156, as primitiva), females of scita are on the average smaller, with "more delicate vestiture" on the legs. Geographic Distribution. — For 25 years this species was known only from the type locality in northwestern Oregon. The sec- ond known sjiecimen was collected 30 De- cember 1971, about a mile east of the eastern end oi Lake Crescent, Clallam Co., Olympic Peninsula, Washington, between 900 and 1300 ft (277-400 m), by N. D. Penny. Both these localities are low on the outer slopes of the northern Coast Ranges (Ma[i 1). Both are in somewhat isolated mountainous regions separated trom I he higher Cascade Range by the Willamette Vallev in Oregon and by Pu- get Sound in Washington. The species ma\ occur noiihwaid into British Colum- bia ant! .ilmost surely southward in Ore- gon. One male was recently collected on snow at 5200-5500 ft, in the Sentinel Mills, Mt. Jefferson Wilderness, Linn Co., Ore- gon, on 22 March I'iSl, by Dr. L. K. Russell. Seasonal Distribution. — The holotype of macnabeana was collected in early Febru- ary, which is about mid-season for the appearance of western Chioneas generally. However, the specimen was taken at a lower elevation than most, which probably relates to its seasonal occurrence, the arrival of winter weather being later there. Habitats. — The type locality is the "Til- lamook burn," a forested area which was swept by fire in 1933, 1939, and again in 1945, only a few months before the speci- men of Chionca macnabeana was collected. The fly was tound on snow about 20 in deep, during a snow storm. The female from the Olympic Peninsula was also found on snow. Chionea nivicola Doane Clnonea nnncola Doane, L^OO: 188-189; Alexander, 1949: 319; Alexander, 1965: 72. Diagnosis. — A western species with brown coloration, antennae of eight or nine segments (6 or 7 flagellomeres); male gonapophyses blade-like, approximately vertically oriented, female cerci slender, slightly upturned in apical half. Description. — G e n e r a 1 a p p e a r a n c e : Head dark brown, thorax light brown dor- sally, brown at sides; abdominal .sclerites light brown to brown, except ninth seg- ment of male darker brown, cerci of fe- male amber colored; pleural membranes pale grayish brcnvn. Length of male about 134 The University of Kansas Science Bulletin 5 mm, female about 4-8 mm, exclusive of antennae. Head: Antennae (Figs. 86, 87) com- prising scape, pedicel and six or seven flagellomeres; fusion segment asymmetri- cally short-conical, about as long as greatest diameter, with apical portion sometimes resembling additional segment. Setae on vertex short, curved, stout. Eyes with about 80-100 ommatidia each. Thorax: Color of thoracic dorsum light brown to dark yellowish brown cen- trally, darkening to brown laterally on pronotum and on posterior metanotum; pale grayish brown spot on dorsal mid-line of metanotum in some individuals. Hal- teres pale grayish brown, contrasting some- what with color of thoracic dorsum. Meso- notal setae sparse, short, stout, often in transverse patch interrupted at mid-line. Legs long, hind femora extending far be- yond end of abdomen, femora light brown, with seven (4 dorsal, 3 ventral) longitudi- nal rows of short, curved thick and dark- ened setae; tibial setae longer, more slender. Tarsomeres beyond basitarsi dark- ening to brown, dark brown on last two. Abdomen of male: Setae on terga very short, sparse, occurring mostly along pos- terior and lateral margins. Sternal setae mostly short, on posterior margins, longer and more dense on posterolateral corners of htth through seventh sterna and on posterior edge of eighth sternum. Postero- dorsal margin ot ninth tcrgum (Fig. 88) bearing two prominent, setiferous lobes; median space between these projections broadly U-shaped; 6-11 straight, thick setae on each lobe. Gonapo[)hyses seen from above (Fig. 89) blade-like, almost vertically oriented, curved outward laterally, concave on mesal surfaces with bluntly rounded tips almost tcjuching dorsally, curving apart ventrally; space between gonapophy- ses broadest basally, usually narrow pos- teriorly; apex of penis flattened and slight- ly expanded, l^asistyles. elonj^atc, slender lor genus, sometimes paler than ninth sternum; posteromesal lobes inconspicuous, pilose, evenly rounded. Dististyles (Figs. 91, 92) dark yellowish brown, broad at base, bowed slightly laterad, then more or less evenly curved to acutely rounded apex; mesal tooth near base prominent, subacute; ventral tooth absent in all specimens seen, but small dorsomesal tooth in a few indi- viduals (Fig. 92, cf. C. scita). Tenth seg- ment broad at base, evenly narrowed, pigmented and lightly sclerotized laterally. Abdomen of female: Tergal and ster- nal setae more numerous and more evenly distributed over sclerites than in males. Eighth tergum at sides about half as long as seventh, shorter dorsally, with three to five posterolateral setae. Hypovalves of eighth sternum slender, elongate, with up- per margin gradually and evenly curved downward, lower margin straight for most of its length, then curved upward to tip (Fig. 95). Arms of genital furca tapering rather evenly from base to tip, more strongly divergent in basal half than in apical half, with noticeable bend near mid- length (Fig. 96); basal portion wider than long, not densely sclerotized medially; en- tire structure lyriform. Mid-ventral portion of ninth sternum (Fig. 96) lightly sclero- tized, produced caudad as short, acute or subacute point. Cerci elongate, slender, about eight to nine times as long as great- est width, slightly curved upward in apical half, in some specimens widened beyond mid-length, noticeably longer tlian hypo- valves (about 1.3-1.4 times as long as hy- povalve measured from notch to tip; Fig. 95).^ l\pcs. — Four iemale syntypes, Pullman (Whitman C'o.), Washington, published without further data. I have seen three ot these, which are now in the United States National Museum, Washington, !).(>. Two (pinned) are labelled Pullman, Washington, on snow, R. W. Doane col- lector, and dated 9 January and 12 January IS'i*;; both have been given U.S.N.M. cata- logue number 7025. The third I have The Crane Fly Genus Chionea in North America 135 0.5 mm 88 r-----,:yW^w--^. 93 89 90 b°L. 1 mm 92 G ►V5 95 94 96 Fk;s. 8f)-96, Chionea nivicola. 86, rij^lit antenna of pinned female, lateral aspect. 87, ri.u-ht antenna, female troin Oregon, in fluid, iiiesal aspect, showing six flagelloinercs be\ond fusion sej^ment. 88, ninth abdominal seument of male troin \\'a^hin>;ton, dorsal aspect (left hasist>le and chstist\ie omitted). 89, typical pona- pophyses and penis, dorsal aspect. 90, left gonapophysis and penis, rigiu lateral aspect: cf. Fig. 94. 91, right tlististsle, ventral aspect. 92, left distist\!e, dorsomesal aspect. 93, postericjr margin of ninth tergum, small male from Idaho, dorsal aspect. 94, atypically widely separated gonapophyses, dorsal aspect (same male as Fig. 90). 95, terminal abdominal segments, female syntype (USNM), left lateral as])ect. 96, ninth sternum and genital furca, female, ventral aspect. Scales: a— 86-87, 89-92, 94, 96; b— 88, 93, 95. 136 The University of Kansas Science Bulletin cleared and mounted on a microscope slide. It was labelled ''Chionea — on snow, 12-5- 99." The remaining syntype is in the collection of Washington State Univ., Pullman. Variation. — Among the specimens ex- amined, about half have five flagellomcres beyond the fusion segment, the others six, with the first being shortest (Fig. 87). Some individuals have a dark brown me- dian longitudinal stripe on the vertex and occiput. One female was seen in which the halteres contrasted conspicuously with the thoracic dorsum by reason of its un- usually dark color. The cerci of some females are slightly but noticeably widened in the apical half, which increases the curvature of the ventral margin and gives the cerci the appearance of being more abruptly upturned. In a few of the larger, more robust males, the basistyles are thick- er than illustrated, but the ventral tooth of the dististyle is not developed as it is in robust males of some other species in which it is more often absent. Although the tips of the gonapophyses are usually strongly convergent (Figs. 88, 89) and rarely are seen in contact, they may be widely separated (Fig. 94). The penis is ordinarily more upcurved apically than as shown in Figure 90. Occasional pres- ence of a small tooth on the inner surface of the dististyle and more dorsal than the large mesal tooth (Fig. 92) has been men- tioned in the description. Size variation, as usual lor the genus, is striking. In a population from Latah Co., Idaho (Moscow Mountain), sampled on 9 December 1971, the shortest male was only .67 times as long as the largest male, and its hind femur was only .53 times as long as that of the largest male. Al- though the basistyles of the smaller males were considerably more slender in lateral aspect than those of more robust males, their length — probably of more functional importance in grasping the tcmale — was m a ratio of about .82 to 1, Other genitalial measurements also showed less size varia- tion: width of ninth tergum .76 to 1, length (jf dististyle 1 to 1, length of gona- pophyses approximately 1 to 1. Certain parts of the original descrip- tion, notably the references to concave areas on the femora and tibiae and to the abdomen's being only six-segmented, are somewhat misleading and are based upon Doane's observation of dried, rather shriv- eled specimens. Relationships. — While antennal struc- ture, the eighth abdominal tereum of the female and perhaps body color suggest C. niuicola belongs with the valga group, the form of the male gonapophyses does not support this placement. The gonapophy- ses are in fact most like those of C. alber- tensis. The long, slender cerci and hypo- valves of the female also resemble those of females in the sata group. Presence of a prominent mesal tooth and occasional dorsomesal tooth at the base of the disti- style is similar to the condition seen in C. scita. The long and, even in males, not greatly thickened femora in nivicola re- semble those of albertensis. Within the sata group niuicola is easily recognized by its body color and number of anten- nomeres. In the field it may occur together with dlbotensis but also frequently with ulexandriana and obtiisa, which it resem- bles in general coloration. From these it may readily be differentiated by having Icjnger antennae than ulexandriana (6 or 7 flagellomeres in nivicola, only 3 or 4 in ulexandriana) and tergal lobes {$)■, slen- der cerci ( 9 ) and short femoral setae, contrasted with absence of tergal lobes, short, thick cerci and conspicuously long femoral setae in obtiisu. The short, thick setae of head, thoracic dorsum, abdomen ami legs are found elsewhere among North American species only in C. stoneana. Geographic Di stribiition. — As now known, the range of C. nivicola includes the C'olumbia Basin and the mountains surrcjunding it: the eastern Cascade Range Th[£ Crane Fly Gknus Chionea in North America 137 to the west, the Blue Mountains to the south, and the Sahnon Mcjuntains, Clear- water Mountains and Bitterroot Range to the east and northeast, hut also over the last of these to the western slopes of the Rocky Mountains in Montana (Ma(i 2). 1 exjx'ct the species may ran^^e into south- western Canada and perha[-»s may occur in the eastern Cascade Range of Oregon. Seasonal Distribution. — Records for mid-November are from mountainous areas at higher elevations than the Wash- ington and Idahcj localities at which C. nii'ico'.a was taken in December and Janti- ary. This seems to conform to the general pattern of earlier appearance of Chionea in areas where cold winter sets in first and subsec]uent appearance at lower latitudes or elevations. This pattern of seasonal distribution suggests a second generation of nivicola may occur in March and April in the colder parts of the range. Habitats. — Most specimens were found on snow, in areas of fairly open forest, at elevations from about 740 to 1850 m, or 2400 ft (type locality) to 6000 ft or more (Oregon). The Oregon specimens taken beneath a log may well have been in an excavation made by a small mammal. There have been too few observations or collections made to justify any other gen- eralizations about the habitats. Collection Records. — ID AHO: Be- newah Co., Crystal Creek, on snow, 22 Dec. 1968, M. Furniss (U , 2$); Boise Co., Idaho City, 14 Dec. 1974, G. A. Shook (19); Clearwater Co., S mi N of Elk River, 14 Nov. 1961, E. H. Dustman (Id); Idaho Co., Clearwater National Forest, Mud Creek Jet., 25 Nov. 1978, R. C. liiggam and M. A. Maurer (1 ^^ , 1?); Mud Creek, Lolo Creek Canyon, 14 Nov. 1975, R. C. Biggam (U); Latah Co., Moscow Mountain, 9 Dec. 1971, G. W. Urich (10 6 , 6 9 ) ; 3 mi S of Helmer, Little I^oulder Camp, 4 Dec. 1971, W. J. Turner (1 6 , 2 9 ); 8 mi E of Moscow, Robinson Lake Road, 6 Dec. 1972, Dunn and Mich- ta (3 i , 3 9); Big Meadows Rec. Area, 4.5 mi NNW of Troy, 9 Dec. 1973, M. C. Hunter (2 9 ) ; Moscow, on snow, 8 Dec. l')6l, R. E. Stecker (55,49); Latah Co., on snow, 14 Dec. 1960, W. F. Barr (2 o., St. Regis, on snow, 28 Dec. 1957, H. R. Dodge (U ); Missoula Co., Missotila, on tresh snow, about 32''F, 16 Nov. 1952, ]. A. Cha[Mnan (1 v ). ORE- CiON: Umatilla Cx). (southeast). Bowman Meadows to Fly C'reek, on snow, 16 Nov. l''()2, J. Schuh and E. Hansen (U , 3 9 ); Fly Creek valley, under log, 15 Nov. 1961, J. Schuh (U , 19); Wallowa Co., Wal- lowa-Whitman National Forest, Eagle Cap Wilderness, 6000-7000 ft, weather clear, 24 Nov. 1972, A. Greene (19). WASHING- Map 2. Ranj;e of Chionea niricola. Each spot rep- resents one or more collections within a county. 138 The University of Kansas Science Bulletin TON: Chelan Co., Stevens Pass, near 4100 ft, 17 Feb. 1973 (19); Garfield Co., Saddle Butte, Blue Mountains, 9 Nov. 1970, J. Eves and R. I). Akrc (3ci', 2?); Whitman Co., Pullman, 5 Dec. 1.S99 (19), 9 Jan. 1899 (1 9 ), 12 Jan. 1899 (1 9 ), all collected by R. W. Doane (types); Pull- man, 15 and 16 Dec. 1970, W. J. Turner (1 5 , 2 9 ); 7 mi NW of Colton, 21 Nov. 1973, L. Richards (1 i ). Chionea scita Walker Chionea scita Walker, 1848: 82-83; Byers, 1963: 160; Alexander, 1965: 72. Chionea primitiva Alexander, 1917: 204-205, Figs. 1, 4; Alexander, 1919: 805, 903; Leonard, 1928: 694; Alexander 1942: 428-429. Diagnosis. — A northeastern species hav- ing yellowish brown color, antennae of 12 (rarely 11 or 13) segments, male gona- pophyses blade-like, vertically oriented, ninth abdominal tergum entire, without projecting lobes; female cerci evenly curved, not greatly elongated; hypovalves long with nearly straight ventral margins. Description. — G e n e r a 1 appearance: Head dark yellowish brown to li^rht brown, thorax yellowish brown; abdom- inal sclerites yellowish brown faintly mot- tled with brown; ninth segment of male dark \cll(n\ish brown, cerci of female light amber colored; pleural membranes dull, sordid yellowish brown. Length of male about 5-8 mm, female about 6-8 mm, exclusive of antennae. Head: Antennae (Fig. 97) comprising scape, pedicel and usually ten Hagellomeres, rarely only nine or as many as 11; fusion segment elongate, conical, longer than its basal diameter, covered with short, fine hairs; second flagellomere also covered with short hairs, third and succeeding seg- ments verticillate, with three, sometimes four, setae. Eyes with about 180-200 om- matidia each. Thorax: Halteres dull, [mIc xcllowish brown, not contrasting sharply with color of thoracic dorsum. Femora, tibiae and proximal three-fourths of basitarsi yellow- ish brown, tarsi otherwise dark brown. Femoral setae arranged in four dorsal, three ventral rows, smaller hairs sometimes not aligned with setae. Mesonotal setae long, erect, conspicuous. Abdomen of male: Posterodorsal mar- gin of ninth tergum (Fig. 99) entire, with- out projecting lobes, broadly and evenly rounded, arched upward and projecting slightly caudad (appearing broadly emargi- nate when seen in posterodorsal aspect), bearing about a dozen marginal setae sometimes in two weakly defined groups. Gonapophyses (Figs. 101, 102) slenderly triangular in dorsal profile, with thickened dorsal margins, apices turned slightly in- ward; form in lateral aspect blade-like (Fig. 92), lower margin curving smoothly upward to meet irregular dorsal margin at upturned, narrowly rounded apex; space between gonapophyses narrow, rcjunded basally, slightly constricted near base; penis evenly curved, its membranous apex spatu- late, with slender ventral sclerotized re- inforcement (Fig. 100). Basistyles thick, their posteromesal lobes pilose, simply rounded. Dististyles (Fig. 98) evenly curved beyond base, yellowish brown, with strongly sclerotized, blackened mesal tooth, trom which a darkened ridge extends along inner, lower surface of dististyle toward ti[\ in iiKJst specimens; dorsomesal tooth low, blunt, rarely rudimentary or absent. Tenth segment as broad basally as space between basistyles, narrowed even- ly toward a[iex, lightly sclerotized and [uibescent on Icjwer, outer surfaces (pushed into ninth segment and not shown in Fig. 99). Abdomen of female: Eighth tergum short (i.e., slender) throughout (Fig. 104), most densely sclerotized along anterior margin, with narrow, \cllowish postero- hiteral edges. H)povalves of eighth ster- num elongate, slender, tapered toward narrowly rounded apex, knver margins The Cranl Fly Genus Chionea in North America 139 more or less straight, upper margins curving slightly downward. Arms of geni- tal furca divergent from basal plate, then subparallel lor ahoLit half their length until apicalh' curved outward to join ninth sternum (h'ig. 105); basal plate not densely sclerotized medially except tor narrow, transverse bar connecting arms. Ninth sternum formed by two converging, slen- der, well sclerotized bands which arch up- ward and backward, ending in small longitudinal bars, parallel and separated by a narrow, clear zone at mid-line (Fig. 105); a lightly sclerotized area with round- ed margin above terminal portion of each band of sternum. Ccrci elongate, about seven to eight times as long as greatest width, noticeably longer than hypovalves (about 1.20-1.36 limes as long as hypo- valves measured Irom notch to tip). Types. — Holotype, female, labelled "North America" but without further Fk;s. 97-1(15, Chinrica sata. 97, Uft nntcniKi, rnalc^ holotype of C. pnrmtiia, mcsal aspect; IcnRth 1.55 mm. 98, riKlu clististvlc, vcntro.ncsal aspect. 99, ninth alnlonunal scumcnt of male tr<.m .Michigan, dorsal aspect (left basistyle and liististyle omitted). 100, apex of penis, dorsal aspect. 101, g-napophyses and penis, dorsal aspect. 102, same, rii,'ht lateral aspect. 103, terminal abdominal segments, male, left lateral aspect. 104. termi- nal abdominal segments, female, left lateral asixct. 105, ninth sternum and genital furca, female, ventr- . aspect. Scales: a— 97-98, 101-102, 105; b— 99, 10-1. 140 The University of Kansas Science Bulletin data. The type was originally in the British Museum (Natural History) hut has been Icjst sometime since 1953. In that year I sent sketches of diagnostic features of fe- males of Chionea valga and what I then called C. primitiva to Dr. Paul Freeman, Curator of Diptera in the British Museum. He compared my sketches with the type and confirmed that primitiva is the same as scitu. Subsequently, however, the holo- type of scita became misplaced, and neither Dr. Freeman nor I was able to locate it when we searched briefly in 1955 and again more thoroughly in I960. The holo- type of C. primitiva Alexander is a male from Cascade, Owasco Lake, Cayuga Co., New York, collected on 15 November 1915 by Bishop and Crosby. It is mounted on a microscope slide in the collection of Dr. C. P. Alexander, Amherst, Massachusetts (transferred in 1981 to the U.S. National Museum, Washington, D.C.). Variation. — Among the many speci- mens examined, only two had eight flagel- lomeres beyond the fusion segment, and onK' one had ten; thus, it can be said that the total number of antennal segments is ordinarily 12. Walker described scita as having antennae with black flagella. Since, at the time Dr. Freeman compared the holotype with my drawings, both its fla- gella were missing, I cannot explain this except to surmise that Walker had viewed the specimen against a pale or illuminated background. The description of the ninth abdominal tergum ol the male, above, per- tains to the dorsal aspect; because the tergum is somewhat arched upward, it may appear broadly emarginate in postero- dorsal aspect. The ventral tooth at the base of the dististyle, which is absent or minute and not darklv colored in some species and even in s(jme males of scita, is usually both enlarged and darkly sclero- tized. The "tuft of stout black hairs" re- ferred to in the original dcscriiuion ot C. primitiva (Alexander, 1917: 205) is a group of short, inconspicuous hairs on the mesal face of the dististyle, near the base, occurring in this and all other species. Alexander also described the legs as having all the fetiKjra enlarged, which is true of males but not females. Variation in the female cerci and hypovalves has been noted above. In most females the pleura of the tenth abdominal segment hang down at each side in a fleshy, semicircular fold, partially concealing the ninth sternum in lateral view. These folds are more prom- inent in scitu than in most other species (Fig. 104). The specific name is derived from Latin scita, meaning beautiful or handsome. Relationships. — This species is one of four in North America that have yellow- ish brown overall coloration, antennae with more than nine segments, the eighth ab- dominal tergum of the female short, and the male gonapophyses blade-like and more or less vertically oriented. These species, together with nivicola, comprise the scita group. Within this group, Chio- nea scita is set somewhat apart from the other three yellowish species, C. alberten- sis, C. macnabeana and C. ivilsoni, on the basis of such characters of the fe- male as the ninth sternum, the furca and the setae on the abdominal scle- rites. Both scita and macnabeana have 12 antennal segments, as compared to 11 in albertensis, but in view of the known indi- vidual variation in this character and the tact that macnabeana is based upon so few sjiecimens, I would place more reliance in the genitalial characters as indicators of relationship. Males of scita difTer from those of albertensis in having the ninth abdominal tergum entire instead of bi- lobed, the penis spatulate at the tip instead (it torked, the surface of the gonapophyses less tnululating than in albertensis and the tips less acute. In scita there is usually a darkened ridge extending distad from the darkened mesal tooth of the dististyle, while in albertensis this tooth is ordinarily not ccjnspicuously blackened, and if a ridge The Crane Fly Geni's Chionca in North America 141 extends distad from it, the ridije is not blackened. Moreover, scita lacks the ckirkly sc'erotized ventral tooth seen in albertensis. Some otlur, less oiitstandiuL; dilTerences will he toiiiid in the discussion ot the rela- tionships of C. albcrtensis. G cographic Distribution. — As now known, the range of Chionca scita extends from Massachusetts and northern Vermont westward across New York to southeastern Michigan, southwestward to eastern Ken- tucky, and southward along the Appala- chian Mountains to northern Georgia, pre- senting a rather distinct and coherent pat- tern, except for the Georgia record (Map 1). \'ery likely the species ranges further northward, throughout New England and well into southern Ontario and Quebec and the Maritime Provinces. It will cer- tainly be found more widely along the lower slopes of the Appalachian Moun- tains. Seasonal Distribution. — Dates of collec- tion of scita extend from the middle of September to the middle of February, but the one September record is from the northernmost locality from which the spe- cies is known (northern Vermont). The one October record and the single one from January are both from cave environments, while the February collection is from a southern but montane environment. All other records are in November and Decem- ber, and even at the West Virginia locality most specimens were collected in Decem- ber. From this evidence I am inclined to believe that there is only one generation per year. Habitats. — This is the only species in which most of the specimens seen were taken from places other than the surface of snow. The most outstanding collection of Chionea scita is that made by Dr. M. S. Briscoe of Howard Univ., Washington, D.C. He set a number of traps in the floor of a cave near Harpers Ferry, West Vir- ginia. These were baited with decompos- ing meat and were designed primarily to attract cavern icolous Coleoptera; however, he took 29 specimens of Chionea scita in addition to the (jther insects. This collec- tion has been discussed earlier under the natural history of Chionea. Mr. Roger S[-ierka ot the Univ. ol Kentucky similarly lound one female sjiecimcn in a cave in Kentucky. It wouKl seem, therefore, that the occurrence of scita in fair numbers and well back from the cave entrance was no chance [ihenomenon and that having wan- dered into caves, these Chioneas might become at least semipermanent cave in- sects. Where I collected scita in southeast- ern Michigan, the habitat was a sparsely wooded shore of a marsh and the micro- habitat a series of cavities in the loose soil and finely divided, decayed wood beneath a stump in the advanced stages of decay. The debris included, besides soil and wood particles, various other bits of plant matter, fragments of dead insects, etc., and al- though I found no sure evidence that the microhabitat uas or had been a rodent nest, I would not rule out this possibility. The holotype of C. primitiva also was not found on snow but in leaf mold. The occurrence of scita at light in Pennsylvania is particularly interesting and has been dis- cussed elsewhere under behavior of the adult flies. Collection Records.— GEOV^GW: Un- ion Co., Vogel State Park, in pit-fall trap, elevation about 3200 ft, 10 Feb. 1972, J. Howell (1 ? ). KENTUCKY: Pulaski Co., Wind Cave (37°03'N, 84°3rW), elevation 840 ft, 24 Oct. 1970, R. Sperka (19). MASSACHUSETTS: Hampshire Co., Amherst, 13 Nov. 1933 (1$); same, 14 Nov. 1933 (U ); both collected by A. B. Gurney. MICHIGAN: Oakland Co., Highland Recreation Area, 4 mi E of Highland, S Nov. 1952, P. B. Kannowski (1 0 ); same locality, 12 Nov. 1952, G. W. Byers, D. Hynes, P. B. Kannowski (2 >., Urbana, in nests of mouse, Microtiis ochrogastcr, 11 Dec. 1938 (19, holotype), 16 Dec. 1939 (1 5 , allo- type), 2 Feb. 1940 (U , 1 9 ), all collected by P. il. Stone; Champaign Co., 2 mi N of H(jmer and 1.5 mi W of Route 49 bridge on Salt Fork of Vermilion River, 7 Nov.-2 Dec. 1972, in pitfall trap in oak- hickory woods, J. D. Unziker (85,99); Clark C'o., Rocky Branch, NE of Clarks- ville, 5 Feb. 1965, Rotrammel and Braasch (1 O ; DeWitt Co., Allerton Park, 27 Dec. 1969, J. Bouseman and R. Selander (3 .5 ) ; Jackson Co., Carbondale, on lighted win- dow, 11 I'.M., temperature 30-32°F, 24 Dec. 19(.0, J. C. Downey (U ); Randolph Co., Fort Kaskaskia State Park, on snow, 15 Jan. 19f,4, L. J. Stannard (U )• INDL ANA: Clay Co., Posey Twp., in pitfall trap, old field, 23-26 Dec. 1974, J. R. Mun- see (U, 19). IOWA: Boone Co., Ledges State Park, in oak-hickory leaf lit- ter on ground near Pease Creek, 7 Nov. 1967, H. D. Blankespoor, L. C. Lewis and R. L. Jacques (3 6, 19, all teneral); Ledges State Park, 19 Jan. 196S, D. J. Kotzin (U, 19). KANSAS: Douglas Co., Breidenthal Reserve, Univ. of Kansas, 2 mi N of Baldwin, alive in pitfall trap, 20 Jan. 1973, C. W. Byers (Id); Franklin C^o., 14.5 mi S of Lawrence, from old mouse nests, 1.M.-7 A.M., 21-22 Dec. 1961, F. D. Parker (U , 19); St. Francois Co., Wo- mack, 16 Jan. 1964, on snow, L. J. Stan- nard (19). OKLAHOMA: Cleveland Co., Norman, 11 Nov. 1926, M. J. Brown (\S). WISCONSIN: Iowa Co., Clyde Twp., Sec. 12, from nest box CH3-1, 23 Dec. 1968, J. O. Jackson (IS). Chionea valga Harris Chionea valga Harris, 1835: 595 (no- men nudum). Chionea valga Harris, 1841: 404-405; Harris, 1852: 482; Harris, 1862: 601, Fig. 260; Osten Sacken, 1869: 168-170; Osten Sacken, 1878: 29; Lugger, 1896: 230-231 (in part C stoneana); Ainslie, 1906: 275- 276; Johnson, 1907: 41-43, with figs.; Wash- burn, 1907: 103; Needham, 1908: 207; Johnson, 1912: 102-104; Alexander, 1917: 207; Alexander, 1919: 805, 903; Treherne, 1920 (misidentified — C. alexandriana) ; Crampton, 1925: 204 ff.. Fig. 9; Leonard, 1928: 694; Alexander, 1942: 429; Alexan- der, 1956: 177; Byers, 196k: 190, Fig. 1; Byers, 1963: 160; Alexander, 1965: 72. Chionea aspera Walker, 1848: 82; Alex- ander, 1919:903. Chionea gracilis Alexander, 1917: 206, Fig. 2; Alexander, 1919: 903; Leonard, 1928: 694; Alexander, 1942: 428-429. Chionea novehoracensis Alexander, 1917: 205-206; Alexander, 1919: 903; Leon- ard, 1928: 694; Alexander, 1^42: 429 (new synonymy). Chionea waiighi Curran, 1*'25: 24-26; Alexander, 1942:429. Diagnosis. — A northern and eastern spe- cies of dull reddish brown to grayish brown color, with antennae of seven or eight segments; male g()na[ii;[")hyses, seen from above, broad, shallouly concave, blade-like, with concealed, deep \ential keel; female cerci slightly widenetl and upturned beyond mid-length, then tapered to acute apex; median j^ortinn of ninth sternum weakly sclcrotized, truncate api- cally. Description. — General appearance of specimens preserved in alcohol: head dark reddish brown, thorax and abdominal scle- rites reddish brown to brown, except ninth segment of male dark reddish brown; membranous areas paler brown; cerci of female dark amber-brown. Specimens pre- served dry (on pins) more grayish brown in color. Length of male about 3.5-6.0 mm, female about 4-8 mm, excluding antennae. Head: Antennae (Fig. 137) with pedi- cel slightly longer than cylindrical scape; fusion segment subconical, asymmetrical, about as long as its basal diameter; four or five additional flagellomeres. Setae on ver- tex dark brown, stout. Eyes each with about 100-130 ommatidia. Thorax: Dorsum mostly reddish brown to brown, darkening unevenly to sides, on pronotum and on metanotum. Mesonotum in both sexes conspicuously "humped" in lateral aspect, with anterior slope nearly perpendicular to pronotum (Figs. 139, 140). Setae on mesonotum numerous, long, erect. Halteres grayish brown, con- trasting with color of thoracic dorsum but not conspicuously so. Legs of moderate length, with hind femora extending only approximately to end of abdomen; femora light reddish brown to dark yellowish brown, their setae long, attenuate at tip, brown, not in readily discernible rows in males, usually in four dorsal, three ventral rows in females. Tibiae and basitarsi same color as femora, with almost equally long, slender setae; tibial setae of females and of fore and middle legs of males generally in longitudinal rows, more randomly dis- tributed on hind legs of males. Tarsomeres beyond basitarsus darkening to brown. Abdomen ol male: Tergal setae long, [xi!e, sparse, chicHy near posterior margins of ters^a. Sternal setae loniiest and most dense at sides ol sterna, with a tew along posterior margins. Posterodorsal margin of ninth tergum (Fig. 141) with widely U-shaped emargination, without lobes but with scattering of submarginal setae; emar- The Cran'i: Fly Ginus Chionca in North Ami-rica 155 Fkis. 137-149, Chionca valga. 137, riulit antenna of male, approximately dorsal aspect (slide-mounted). 138, hasal portion of left antenna, male trom Michigan (J. S. Rogers slide 28-18), showing subchvision of basal flageliomcre. 139, thf)racic dorsum of male, left lateral aspect, to show abrupt slope of mcsonotum. 140, thorax of female, left lateral aspect; compare with C. obtiisu, Figs. 123, 124. 141, ninth abdominal segment of male, dorsal aspect (left basistylc and dististyle omitted, tenth segment pushed inward to exjiose gonapoph- yses). 142, right gonapophysis and penis, male from Wisconsin, right lateral aspect. 143, gonapophyscs and penis, male from New Hampshire, dorsal aspect. 144, gonapophyscs, male from Minnesota, dorsal aspect. 145, left dististyle, male from Wisconsin, ventral aspect (somewhat foreshortened because of curvature). 146, left dististyle, ventromcsal aspect. 147, terminal ;:bdominal segments, female from Michigan, left lateral aspect. 148, ninth sternum and genital furca, female from Vermont, posterovcntral aspect. 149, genital furca, left lateral aspect. Scales: a— 137-138, 142-146, 148-149; b— 139-141, 147. 156 The University of Kansas Science Bulletin gination so wide and shallow as to give impression of entire margin in certain as- pects. CJ()na[X)physes in dorsal aspect (Fig. 144) brcjad, nearly oval, narrowest at rounded apex, with upturned mesal and lateral margins producing central concav- ity; mesal margin noticeably thickened; in lateral aspect (Fig. 142) deeply keeled, posterior end of keel dee[iest. Apex of penis with two slender, elongate iilaments (Fig. 143). Hasistyles each with rounded, pilose postercjmesal lobe; dististyles (Figs. 14"^, 146) dark reddish brown, with promi- nent, acute, blackened mesal tooth; ventral tooth absent. I'roctiger (tenth segment) membranous, [)ale, evenl)' narrowed to- ward rounded apex, extending approxi- mately to tips of gonapophyses. Abdomen of female: Setae of terga and sterna [lale, more tiispersed over scle- rites than in male. Eighth tergum (Fig. 147) expanded at sides to more than half length of seventh tergum, shorter dorsally, with darkened, transverse line or zone near mid-length and with three cjr four marginal setae. Ihpovalves extending to about mid length ot cerci, with upper mar- gin slightl}' sinuate, lower margin straight near base, then curved u[n\ard toward narrowly rounded a[iex. Arms of genital lurca (Fig. 14S) relatively thick at base, slender in apical half, often more strongly divergent basally than apicallv, with nearly angular deflection near niid length; base with divergent a|)otleincs; darkened median portion oi bursa copulatrix adja- cent to lurca visible through nieinbranc in posterior aspect (or ui cleared, slide- moup.ted s[ucimens). Mid ventral [lortion ot ninth stenuim (big. 14S) not strongly sclerotizcd, somewhat trapezoidal in shape. Cerci about 1.3-1.4 times length of hypo- valves (measured Irom notch to ti[)), up- ttirnetl and slightly widened beyond mid- length (Fig. 147), with sharply rounded or acute apex. Types. — Holotype, female, labelled onlv "174. 9" and "M.C.Z. Type 2639S," in the Museum of Comparative Zoology, Harvard Univ. This pinned specimen is considerably shriveled, with the abdominal sclerites telescoped. A label near the speci- men reads "Chionea valga Harris Type." Mr. Norman Woodley found a catalogue of Thaddeus Harris' collection in the ar- chives of the Museum of Comparative Zoology, and after the number 174 the entry "Dublin, N. H., Mr. Leonard." He also found a list of specimens entitled "Insects for Mr. Say, from T. W. Harris," on page 20 of which the number 174 is followed by '^Chionea valga, mi. Mss. N. Hamp. 4." The type, therefore, although reccjrded fnjm Massachusetts (i.e., in "A Report on the Insects of Massachusetts"'), is apjxu'cntly 70. D. Hansen (15,19); Iowa Co., Clyde Tup., Sect. 12, on snow, north-facing slope, Dec. 1967 (15); Price C^o., walking on snow, 21 Jan. 1979 (1 5 , 2 9); Sauk Co., Otter Creek, on snow along stream bank, 8 Feb. 1969, R. P. Narf (19); same, but 7 Jan. 1967 (15); same, but 27 March 1967 (19); Parfrey's Glen, TUN, R7E, Sect. 25, 7 Jan. 1967, R. P. Narf (39); same, but Sect. 22-23, 13 March 1965 (19); same, but 14 March 1965, J. K. Lawton (19); Waushara Co., Bird Creek, T18N, RIOE, Sect. 3, Hwy. 21 bridge, 9 Feb. 1967 (19). 152 The University of Kansas Science Bulletin The C. alexandriana Group A western montane group of species. Body color brown, with evident pattern on abdominal terga in some species, dark brown or black; antennae with two to four flagellomeres beyond fusion segment; gonapophyses of male with upturned, often hook-like apical point, usually broad at base and conspicuously narrowed toward apex, in dorsal aspect; eighth abdominal tcrgum of female short, much less than half length of seventh icrgum; median portion of ninth sternum a sclerotized plate, often with three posterior lobes or teeth. Included species: alexandriana G.xr- rett, carolus Byers, diirbini new species, excavatu new species, jellisoni new species, lyrata new species and nigra new species. Chionea alexandriana Garrett Chionca alexandriana Garrett, 1922: 62-64, Figs. 6-11; Alexander, 1%5: 72. As Chionea valga: Trchcrne, 1920: 201-202 (misidcntihcation). Diagnosis. — A dark bruuii, western, primarily montane species; antennae with three to four flagellomeres; male gona- pophyses slender and hooked at apex, disti- styles smoothly rouiukd dors, illy; female cerci slender, upturned in distal luilf, no- ticeably longer than hypovalves. Description. — General appearance: Head dark grayish brown, thorax dark brown; abdominal sclerites light brown to brown, terga darker brown anteriorly, along mid- line and at lateral edges; ninth segment of male dark grayish brown, cerci of fe- male amber-hrcjuii; pleural membranes grayish tan. Length of male about 4-8.3 mm, female abcnit 4-S mm, exclusive of antennae. Head: Antennae (Figs. 150-152) com- prising scape, pedicel, and usually three, not uncommonly four flagellomeres; fu- sion segment short, turlyinate to subconical, its apical portion sometimes resembling an additional short segment. Eyes with about 110-120 ommatidia each. Thorax: Halteres pale grayish brown (in alcohol) to yellowish brown (dried), contrasting with color of thoracic dorsum. Legs dark brown to dark reddish brown; seven (4 dorsal, 3 ventral) tcj 11 longitudi- nal rcnvs of setae on femora, in males sometimes additional hairs not in rows; second and succeeding tarsomeres blackish brown to black. Abdomen of male: Posterodorsal edge of ninth tcrgum (Fig. 153) with a broad, shallow median emargination; a small, thick lobe bearing six to eight setae at each side of emargination. Gonapophyses (Figs. 156-159) broad at base, narrowing abrupdy near mid-length; apical portion hook-like, with prominent, angular ventral process, slender, curved upward to acute tip; space between gonapophyses narrowly U-shaped, or slightly narrowed caudally, less often constricted near posterior end; apex of penis unmodified. Basistyles moderately thick for genus to thick in large males; [Kjsteromesal lobe of each pilose, some- times with two rounded protuberances. Dististyles (Fig. 155) dark brown, thick, with prominent, blackened, subacute mesal tooth at base, then abruptly narrowed, more or less circular in cross-section, and evenly tapered to apex; ventral tooth usu- ally small,- in some large males darkly sclerotized but not strongly developed. Tenth segment broad at base, narrowed near mid-length, pigmented and lightly sclerotized laterally. Abdomen of female: Eighth tergum short (i.e., slender) throughout. Hypo- valves of eighth sternum comparatively slender, dorsal margin sloping downward toward a[K\\, ventral margin curving up- ward, producing subacute apex. Arms of genital furca slender apically, slightly thickened near base (Fig. 161); basal por- tion about as wide as long, weakly sclero- tized medially. Mid-ventral portion of ninth sternum (Figs. 161, 162a,b) well Thh Crank Fly (n nus Chionca in North America 163 Fir.s. 150-162, Chionca alexandriana. 150, left antenna of male, mcsal aspect. 151, 152, antennal fla,£;cila of females from Alberta, showint; variation in number ol ilauellonieres (in 152, one of the four setae arises on the tusion segment). 153. ninth .ilxlominal ter^um, male, dorsal aspect. 15-4, uenit.i! se>:ment of robust male, lelt l.iter.il aspect. 155, left tlististyle, ventral aspect. 156, gona|X)|)hyses and penis, dorsal aspect. 157, same, left lateral aspect. 158, 159, tjonapophyses of atypical male from ice cave in Oregon. 160, terminal alxlominal segments, female, left latir.d aspect. 161, ninth sternum and genital furca, female, ventral aspect. 1 62a, b, variations in median portion of ninth abdominal sternum. Scales: a — 150-153, 156-159, 161-162; b — 154- 155, 160. 164 The University of Kansas Sciilnce Bulletin sclerotized, with three bhiiit teeth, mickile one longest; a rounded, H_L;htly sclerotized area above and to each side ot darkened portion. Cerci elongate, about 6-6.5 times as long as greatest width, curved upward in apical half, noticeably longer than hypo- valves (about 1.2-1.4 times as long as hypo- valve measured from notch to tip). Garrett's (1922: 63) illustrations do not adequately portray the structural details he intended to show; they are, in fact, rather grotesque caricatures. Types. — Holotype, male, Cranbrook, British Cokimbia, elevation 3500 ft, 22 Feb- ruary 1921, C ]). 1). (kirrett collector. Allotype, female, ami two female para- types, same data as holotype. Additional paratypes: 1 9, type locality but collected 13 February 1920; 1 9 , Canal Flats, Koote- nay Valley, British Columbia, March or April 1921. Holotype, allotype and Koote- nay Valley paratype in collection of Mr. C Ik D. (larrett, Cranbrocjk, Ikitish Co- lumbia. One paratype each sent to Dr. C]. P. Alexander, Amherst, Massachusetts, United States National Museum, Wash- ington, D.C., and (kanadian National Ck)l- lection (Canada Department of Agricul- ture), Ottawa, Ontario. Variation. — In a series of 15 males and 17 females from the type locality, ten males and ten females had two flagellar segments beyond the fusion segment, and five males and seven females had three. In individ- uals from other localities, also, the number most often seen is two in addition to the fusion segment. Rarely, there is only one (Fig. 152). Variation in the number of rows of femoral setae, in the ventral tooth of the iTiale dististyle, and certain other characters has already been noted. The lobes of the ninth tergum ot the male vary from low and broad to prominent and longer than wide. Seen from above, the space between the gonapophyses may be narrowly U-shaped or a little constricted near the bases of the apical hooks. A single male in a collection of 18 individuals from an ice cave in Union Co., Oregon, has this sjiacc strcjiigly constricted caudally (Fig. 15S); the gonapophyses are atypical in other respects as well (Fig. 159), with rela- tively short, thick apical hooks. Compres- sion by the basistyles during preservation mav distort the gonapophyses by partial rot.ition to the sides or constriction of the space between them. The outline of the gonapophyses (in this and all other spe- cies) can also be made to vary by com- pression on a microscope slide, but this kind ot distortion should not in interpreted as variation. In the genital furca of the female the deflected apices of the arms vary in the degree of sclerotization and may not be as dark as shown in the illus- tration. There is considerable variation also in the relative sizes — particularly the lengths — of the three teeth of the ninth sternum (ct. Figs. 161, 162). Relationships. — The gonapophyses and antennae of the male and the combination (jf short eighth tergum and abbreviated antennae in the female show the close relationship of Chionea alexandriana with nigra and jellisom, and suggest relationship with excavata and lyrata and a somewhat more distant relationship with carolits and diirhini. This grouping of species (the alcxdndriana group) is reinforced by sev- eral other similarities, such as the structure of the ninth sternum in the female and the internal reproductive systems of both sexes. Within the group, alexandriana most close- ly resembles C. nigra n. sp., from which it dilTers most conspicuously in color. These two species may also be differentiated by the more prominent ventral process on the apical hook of the male gonapophysis and by the tridentate ninth sternum in the female of alexandriana. These is a differ- ence— though not always apparent — in the shape ot the tusion segment of the an- tenna, longer in alexandriana, short and broadly turbinate in nigra. Geographic Distribiitio n. — Ch io n ea alexandriana has been found from the Thi-: Crank Fly Ghnus Chionca in North America 165 niiiiiiUaiiis (it western Alberta and south- ern Ihitish ('c)lunibia southward through Washington, Oregon, Idaho antl western Montana, to the east of the Great I'asin in the Wasatch Mountains ol Utah and along the Sierra Nevatla to western Ne- vada and east-central (California. Only recently (I'^Sl) has this range been ex- tended significantly northwestward, to the vicinity of Whitehorse, Yukon Territory, over SOO miles from the nearest known lo- calities in southwestern Canada. (This localitN' is not included on the distribution map. Map 4). The species is generally montane, but occurs from near sea level to 3000 meters (9S00 ft). It coexists with several other western species, both in its general range and in its particular habi- tats. It is the most widely distributed of the western group of species (Map 4). Seasonal Distribution. — There are col- lection records from some part of the range for every ten-day period from 1 October through 20 April, with isolated collections in mid-September and mid- May. Concentrated local collections, such as those made by Garrett and Woods in British Columbia and Chapman in Mon- tana, show a peak period of abundance of the adults in November and December and an almost et]ually large but more pro- longed peak in February and March. There may be two generations per year. As for some other species, the largest number of separate collections is in late December and early January, probably as a result of academic holidays. The snow cover was sometimes fresh, other times older and firm, and varied from 2.5-3 ft in thickness. Temperatures recorded at times of collection ranged from 21-32°F. Al- though active at 22°F and slightly so at 21 °F, C. alexandriana was in cold stupor at 19.5°F (Chapman, 1954: 362). Near Mara in the northern Okanagan Valley, British Columbia, Mr. H. J. Blurton, a trapper, observed Chionea alexandriana on the snow in open stands of spruce and bal- sam (but not in dense stantls) near timber- Inie, between 5000 and 6500 ft. He saw the insects occasionally during January through April antl noted that they were active on the snow surlace on stormy days in A[iril but were not seen when the tem[ierature was so high as to make the suriace moist (Treherne, 1920). Data from many collectors are less complete and do not indicate whether the specimens were taken on snow or not; probably most were, as Chiotica is ordinarily not otherwise likely to be seen. However, in the Lava Ikxls National Monument, Modoc Co., California, Mr. Joe Schuh collected a male of alexandriana from an abandoned nest of a pack rat (Neototna sp.). Occurrence Map -1. Range ot Chionea alexandriana. Each spot represents one or more collections within a county (in the United States) or at a locahty. A locality near Whitehorse, Yukon Territory, is not indicated on the map. 166 The University of Kansas Science Bulletin of alexandriana in an ice cave has been mentioned earlier. Habitats. — The clearest picture oi the ecological distribution of Chioneu alexan- driana comes from the studies of winter insects made in the IJittcrroot Mountains in Missoula Co., Montana, by Dr. John A. Chapman (1954). He found the flies always on snow and invariably in forested areas, although sometimes several yards from the nearest tree. Chapman's collec- tions were made at elevations from 3600- 8000 ft, most of the tlics being taken be- tween 6000 and 7000 ft. The forest at these elevations is composed largely of lodgepole pine (Piniis contorta) and Douglas fir {PseitdotSLiga taxi folia) . Collection /^c-ro/v/y.—ALUERTA: Banff National Park, Boulder Pass, 7490 ft, on 12 in of snow, 14 Oct. 1%S, K. W. Rich- ards {2 6, 19); Banff National Park, Castle Mountain (Mt. bjsenhower), on snow, 5400 ft, 0°F, sunny, 31 Jan. 1956 (2 6); Banlf vicinity, 20 March 1930, Owen I)r\ant (1 i ); same, but 15 April 192(S (id, 1$); same, but on snow, 23 Oct. 1932 (16); same, but C^ascade Moun- tain, 6000 ft, 26 Jan. 192S (19); also, a tew specimens without tlatc or collector's name from vicinity of Banlf; Jasper Na- tional Park, Tup.' 43, Range 1, 6200 ft, Sta. 7, 2-4 Oct. 1964 (36,49); same, but 7600 ft, Sta. 8 (2 9); Athabasca (Glacier area, 13 Oct. 1970, (]. G. Y.. Scudder (3 i , 19); \V of Ya-Ha Tinda Ranch (about 20 mi X of lianlf), 27 Jan. VHA) (56,69, inchuluig 1 mating [xiir). IIRITISH CO- LUMBIA: Cranbrook (49"31'N, 115°46' W), elevation about 3000 ft, on snow, 1 Nov. (2 6, 19), 2 Nov. (16), 4 Nov. (3 9), 10 Nov. (1 6 ), 14 Nov. (1 9), 17 Nov. (4 6 , 3 9 ), 21 Nov. (1 9 ), 23 Nov. (1 6 ), 3 Dec. (1 9 ), 13 Feb. (1 9 ), 22 Feb. (16, 3 9, types), 15 March (M), 16 March (1 6 , 2 9 ), 12 April (46,29), 19 April (4 9), all collected by C. B. D. Garrett, with no years recorded; (ilacier National Park, 7 Dec. 1979 (1 6 ), 24 Dec. 1979 (5 6, 3 9), 13 Feb. 1977 (16, 19), 20 Feb. 1980 (16, 2 9), 24 Feb. 1979 (4 6 , 3 9 ), 16 March 1980 (46,59), all collected by John G. Woods; Gold- stream River, 118°45'W, 52°45'N, 26 Dec. 1979 (116,99), John G. Woods; Hunt- er's Range, E of Mara (50°4rN, 118°45' W), elevation 5000-6500 ft, on snow, Feb. 1927 (4 9), March 1920, H. J. Blurton (46,29, including a mating pair) ; Koo- tenay Valley, March-April 1921 (19); Manning Prov. Park, near 0°C, snowing heavily,' 5000-5700 ft, 10 Feb. 1979 (4 6, 3 9); same locality, near 0°C, on deep snow, 4000 ft, 11 Feb. 1978 (1 6 ); same, Cambie Creek, -1°C, about 4300 ft, 16 Mar. 1980 (36,79); same -8°C, 17 Feb. 1980 (16); same but about 0°C, after sun- set, 4500-5800 ft, 16 Feb. 1980 (5 6, 89); same but -5 to 4-5 °C, in dappled shade or bright sun, on snow, 28 Feb. 1981 (66,6$); all collected by S. G. Cannings; Manning Park, in shade on snow, 4000 ft, 3°C, 1 Mar. 1981, L. Vasington (16); same but on snow at dusk, about 0°C, 4000 ft, 9 Feb. 1980, R. J. Cannings (16); Gari- Ixddi Prov. Park, Diamond Head Trail, about 4000 ft, -1 to 3"C, overcast, 5 Apr. 1981, S. C;. Cannings (19); Mount Revel- stoke National Park, 28 Nov. 1976 (1 9 ), 12 Dec. 1979 (19), 25 Dec. 1979 (19), 28 Dec. 1976 (1 6 , 1 9 ), 13 Jan. 1980 (2 6 , 5 9 ), 1 I'eb. 1977 (1 9 ), 3 Feb. 1980 (19), 9 Feb. 1980 (2 6 , 3 9 ), 10 Feb. 1980 (8 6 , 6 9), 23 Feb. 1978 (1 6 , 1 9 ), all collected by John G. Woods; Salmon Arm (50° 39'N, 119°16'W), 2 Jan. 1926 ("1-2-1926") (2 6 ); Silver Star Prov. Park, Silver Star Mt., on snow in sunshine, noon, 12 Jan, 1981, A. I'.rvan (16, 19); same but snownig lightly, near -3°C, 1100-1300 hrs, 29 J.,,,. P«l, A. Bryan (16, 4 9 ); Van- couver, on snow, 21 Jan. 1933, H. II Leech (IV); V'ancouver vicinity, Holly- k-urn Mt., HOO ft, on snow, 1 Apr. 1978, P. Withler (L! ); -^^inie but 4000 ft, on sncjw, -3 C, 1 Mar. 1979, S. G. ('annings (1$); Crown Mt., 3 Nov. 1935, J. Poole The Cranu Fly Gknus Chionea in North America 167 (1 '',19); Ml. Seymour (15 mi NE Van- couver), 1000 m, overcast, snowing:;, 18 March 1980, A. C^rass (19); Wells '(^,ray Provincial Park (about 50°30'N, 120^00' W), aloiiL; willow flats, Stillwater, 1 Jan. 19()2, P. Josliii (19); same area, on snow, 11 Jan. 19S1, T. Goward (2?); Yoho National Park, fir-pine habitat. Lake O'Hara, 12 Feb. 1^77, C. W. Aitchison (1 I , 1 V ); Yoho Park, () Feb. 1977 (1 9 ), 25 Feb. 1980 (1 9 ), both John G. Woods; Apex Mountain, Similkameen Dist., 6500 ft, on snow, 22 Dec. 1977, R. \. Gainiini^s (19); Vaseux Lake, Okanagan River, 1900-2500 ft, on snow, 0-3°G, 23 Dec. 1980, R. J. Cannings (24); Mineral King Mine, 4500 ft, Toby Creek, near Lake Winder- mere, snowing, -1 to 0°C, coniferous for- est, 8 Feb. 1960, G. O. M. Stewart (44); Whistler Mountain, 5000 ft, 10 Oct. 1970, 1. ISntt (19); Paul Ridge, 5000 ft, shady N slope, 0-l°C, 11 Jan. 1981, S. G. Can- nnigs (1 9 ). CALIFORNIA: Alpine Co., Carson Pass, off C:alif. Hwy. 88, 8600 ft, collected dead on snow 3-4 ft deep, 15 Dec. 1974, D. Durbin (1 6 ); Modoc Co., Lava IJeds National Monument, Crystal Cave, in nest of dead pack rat, 9 Feb. 1962, J. Schuh and E. Hansen (14); Moncj Cf)., Carnegie Inst. Exper. Sta., 9800 ft, on snow, 28 Oct. 1973, D. Durbin (14); Nevada Co., Sagehen Creek at Univ. Calif. Field Station, 16 March 1972, R. A. Haick (14); Plumas Co., near Caribou, 24 Jan. 1922, H. S. Barber (19). IDAHO: Benewah Co., Crystal Creek, on snow, 22 Dec. 1968, M. M. Furniss (24,39); Boise Co., 0.25 mi E of Idaho City, on snow in pine forest, 7 Dec. 1974, G. A. Shook (19); Boundary Co., Round Prairie, 2-3 mi W ot Addie (Jii U.S. Hwy. 95, Hall Mt., walking on snow on old logging trail in mixed coniierous lorest, in light rain, about 35"F, 11 Nov. 1973, A. Cjreene (14 ); Clearwater Cx)., East Fork of Pcjtlatch River, 5.5 mi E of Bovill, on snow, 11 Dec. 1978, R. (". Biggam, T. F. Hansen and P. C. Schroeder (3 4 , 19); Idaho Co., Clearwater National Forest, Mud Creek Jet., 25 Nov. 1978, R. C. Biggam and M. A. Maurcr (1 4 ); Warm Springs Creek, about 20 mi SW of Lolo Pass, on U.S. Hwy. 12, 28 Dec. 1975, A. (heene (4 4, 2$); Latah Co., 4 mi NE of Harvard, Lost Creek, on snow, 6 Dec. 1975, R. C. P)iggam (19); 3 mi 5 ot Helmer, Little lioulder Camp, 4 Dec. 1971, W. J. Turner (14, 2 9); 6-7 mi NW of Troy, along l)ig Meadow Oeek, overcast, still, 20"'h\ 14 Dee. 1975, A. (Jreene (!:■); Big Meadows Rec. Area, 4.5 mi NNW of Troy, 9 Dec. 1973, M. C. Hunter (3 4 ); 4 mi E of Emida Summit, Alt. Hwy. ')5, 11 Feb. 1978, D. M. Kirk- bride (14, 19); Shoshone C^o., C>larkia, on snow, 14 Dec. I960, W. F. Barr (14); Gold Hill, on St. Joe Divide, S of Wal- lace, on snow, 6000 ft, 23 March 1957, D. Fellin (14). MONTANA: Glacier Naticjiial Park, near foot of Lake Mc- Donald, walking on snow, near 22°F, 31 Dec. 1966 (19); Lake Co., Lion Creek, Swan Valley, 4 March 1972. R. A. Haick (19); Lincoln (>)., Leigh Lake, 20 mi S of Libby, 5200-5400 ft, on fresh snow on rocky ground with moss, bear grass and sparse shrubs, 17 Sept. 1976, L. Wright (14, 2 9); Mineral Cx)., St. Regis, on snow, 28 Dec. 1957, H. R. Dodge (3 5, 2 9); Missoula Co., Lolo vicinitw Bitter- root Mountains, on snow, 6000-7500 ft, 14 Feb. 1953 (14 4 , 12 9 , including 2 mating pairs); same, but 3800 ft (1 4 , 1 9 ); near Lolo, on fresh snow, about 6500 ft, 14 March 1953 (2 4,19); same, but 6700 ft (14, 19); Lolo, (Ml fresh snow, Nov. 1950 (14,19); near Lolo, on fresh snow, about 6500 ft, 8 Apr 1953 (74,69); same, but 7800 ft, P. May 1953 (14,19); Mis- soula vicinity, on snow, 6 Feb. 1953 (2 4); same, but about 3600 ft, 9 March 1952 (14); same, but about 32''F, 21 Dec. 1952 (3 4); all collected by J. A. Chap- man; Missoula Co., on snow 26 Feb. 1957, H. R. Dodge (14, 19); Elk Creek, Lubrecht Forest, 4000 ft, 22 Jan. 1972, 168 The UNivERsiTi' OF Kansas Science Bulletin R. A. Haick, D. D. Jones and A. L. Costa (3 6, 7 9); Grant Creek, GC-3, 13 Feb. 1972, R. A. Haick (2S, 1?); Grant Creek, 13 Dec. 1971, R. A. Haick (1 6 ); l^utler Creek, Snow Bowl, 15 Apr. 1972, R. A. Haick (19); Ravalli Co., Tin Cup Canyon, 3 mi W of Darby, on snow, near 4100 ft, 29 Nov. 1972 (2 6, 19); same, but 1 Dec. 1970 (26); same, but 13 Dec. 1973 (5i, 2 9); same, but 16 Jan. 1972 (2 c5 , 1 9 ) ; all collected by W. L. Jellison: Lake Como, on snow, 31 Dec. 1969 (IS); same, but 27 Nov. 1965 (19); Gird's Creek, vermiculite mines, on snow, 15 Oct. 1954 (li); all collected by W. L. Jellison; Tin ('up Canyon, 3 mi W of Darby, 4100 ft, on snow, 1 Jan. 1971, G. W. Byers (I 6 , 19); West Fork Ranger Sta- tion, on snow, 28 Nov. 1962, W. L. Jellison (26,19). NEVADA: Washoe Co., NE of Hunter Lake, Mt. Rose, 7200-S200 ft, 19 Jan. 1923, H. S. Barber (4 6 ). ORE- GON: Benton Co., Mary's Peak, Woods Creek Road, 7 March 1976, G. L. Peters (19); Clackamas Co., Mt. Hood, on snow, 32°F, 2 Jan. 1976, J. D. Lattin (19); Deschutes Co., Mt. Bachelor, on snow, 10°C, overcast, 26 Dec. 19S0, L. Russell (2 9); same, 8000 ft, on snow, 3 Jan. 1969, K. Goeden (4 6, 8 9); 22 mi SW of Bend, near Todd Lake, 6300 ft, 21 March 1973, P. Lattin (19); Grant Co., Anthony Lakes Ski Area, on snow, 18 Feb. I'i74, K. Goeden (7 6, 13 9); Hood River Co., 30 mi S of Hood River, 3 mi E of Bennett Pass, 20 Jan. 1974, P. Johnson (16,39); JefTerson (>)., Square Lake Trail, 1.5 mi E of Santiam Pass sum- mit, on snow, 28 Dec. 1975, G. L. Peters (16); Klamath Co., Crater Lake National Park, on snow, park headquarters area, near 6500 ft, 15 Dec. 1957, D. H. Hunt- zinger (16); same, but 1 1 Feb. 1958, R. M. lirown (16); Lane Co., IVngra Pass, 22 mi SE of Oakridge, 5100 fi, 4 Jan. 1975, J. D. Lattin (16); Linn Co., Mc- Kcnzie Pass, 5100 ft, 28 Dec. 1977, R. S. Zack (4 6, 4 9); 24 mi NW of Sisters, 4600 ft, 30 Dec. 1974, J. D. Lattin (3 6, 19); Lost Prairie, Willamette National Forest, on snow, 11 Jan. 1975, G. L. Peters (2 6, 19); 2 mi E of jet. of Oregon Hwys. 20 and 26, on snow, 18 Jan. 1975, D. C. Carlson (16, 19); Parish Lake Road, 6 mi S of Marion Forks, 3500 ft, 27 Jan. 1973, J. D. Lattin (16, 5 9); Lost Lake, on snow, 12 Jan. 1976, G. L. Peters (16); Potato Hill Road, near Lost Lake, on snow, 18 Jan. 1976 (l9); 0.25 mi W of jet. Oregon Hwys 20 and 22, 4600 ft, on snow, 30 Dec. 1974, J. D. Lattin (16 ); 2.5 mi E of jet. Oregon Hwys. 20 and 22, 4500-5200 ft, near 0°C, 1 Feb. 1975, P. Johnson (2 9 ); Iron Mt., on snow, 31 Jan. I'JSl, L. Russell (5 6, 3 9); Pacific Coast Trail to N Santiam Pass, dead on snow, 20 Dec. 1980, L. Russell (26,39); Sentinel Hills, Mt. Jefferson Wilderness, 4200-5500 ft, on snow, 22 Mar. 1981, L. Russell (16, 19); Marion Co., 1 mi W of Cheatcheek, about 3000 ft, 22 March 1975, R. J. Bury (16); Whitewater Creek, 14 mi NE Detroit, on snow, 3800 ft, 29 Mar. 1981, L. Russell (19); Polk Co., Laurel Mountain, 3000 ft, 21 Jan. 1978, L. Russell (2 6); Umatilla Co. (south- east). Bowman Meadows to Fly Creek, on snow, 16 Nov. 1962, J. Schuh (26); Union Co., from ice caves 10 mi S of Union, no date, R. Zack et al. (1 6 , 17 9 ) ; Wallowa Co., Wallowa-Whitman National Forest, Eagle Cap Wilderness, 6000-7000 ft, weath- er clear, 24 Nov. 1972, A. Greene (26); same, but Lostine River, about 6000 ft, 27 Dec. 1974, no collector (19); same, but 3 Jan. 1975 (2 6 ); Wasco Co., 15 mi SE of (jovernment Camp, Frog Lake Butte, 31 Dec. 1975, P. J. Johnson (4 6, 5 9). UTAH: Wasatch C'o., Cascade Springs, Little Deer Creek, on snow, 29 Jan. 1966, E. C. Devenport (2 9). WASHING- TON: Chelan C^o., near Wenatchee, No. 2 C:yn., 16 Dec. 1973, M. C. Hunter (19); Kittitas C'o., near Lost Lake, S of Sno- c]ualmic Pass, 3200 ft, on snow, about 1 pcv niile along logging road, 26 Nov. 1971, Till-. Cran'i; Fly Ghnus Chionca in North America 169 J. S. Edwards (1 c5 , 2?); Snoqunlmic Pass, 1 Jan. \^l(^ (2 o ) ; Spokane C:.)., Mt. Spokane State Park, 5000 ft, 9 Nov. P)75, A. Greene (5 6,69); Yakima Co., 15 mi E of C^hinook Pass, on snow about 2.5 It deep, 17 Dec. 1977, P. J. Johnson (3 5, 6 9); near Yakima, 5000-6000 ft, on snow, April P>(,3 (Is)- YUKON TERRI- TORY: Evelyn Creek, near Whitehorse, on snow, 7 Jan. lOSl, W. (]. Johnston (2 6 ), .same hut 21 Eeh. 1981 (2 9). Chionea carolus 1)VErs Chionca carolus livers, 1979: 719-722. Diagnosis. — A sinaH, dark hrown Chio- nca ot the lujrthern and central Sierra Figs. 163-170, Chionea carolus. 163, left antenna of male, lateral aspect. 164, pona- pophyses, penis, vesica and its apodemes, riKht lateral aspect. 165, ninth abdominal tergum, left hasistyle and left dististyie (tenth segment pushed inward to show gona- pophyses), dorsal aspect. 166, left hasistyle and dististyie, ventral aspect. 167, left hind femur, male, dorsolateral aspect. I6.S, right front femur and tibia, female, anterior aspect. 169, ternimal alulominal segments, female, left lateral aspect. 170, ninth sternum and genital furca, female, ventral aspect. Scales: a — 163-166, 170; h — 167-169. 170 This University of Kansas Science Bulletin Nevada; antennae with four or five fla^^el- lomeres; male gonapophyses compressed, bowed outward near mid-length, conver- gent before acute apices, mcjre than half length of basistyles; female cerci short, about 1.1 times length of hypovalves; ninth sternum formed of two subapically wid- ened, apically bifurcate bands converging but not or barely in contact medially. Description. — General appearance: Scle- rites dark hrown td brown, hcail blackish brown; membranous areas pale grayish brown; genital segment ol male blackish brown; cerci of female amber brown. Femora in both sexes conspicuously thick- ened except slender near trochanteral at- tachments; hairs of legs relatively k)ng. Length of male about 3-5 mm; female about 3.5-5 mm. Head: Antennae (Fig. 163) compris- ing cylindrical scape, distally enlarged pedicel, long-conical fusion segment and three or tour atklitional flagellomeres, the apical one very short. Setae of vertex stout but few and scattered. Eyes with about 70-90 ommatidia. Thorax: Halteres pale gra\ish brown, conspicuously paler than dark brown tho- racic dorsum. Femora slender and light brown to yellowish bniwn near attach- ments to troch. inters, thickened and darker brown in both sexes through most of their lengths, especially noticeable in hind legs of male (Fig. 167) and fore legs of female (Fig. 16S); seven or eight longitudinal rows of setae on femora, as well as a few scattered setae ncjt in rows. Setae of tibiae nearly three times as long as tibial diam- eter, giving legs an unusually iiairy ap- pearance. Tarsi only about as dark as tibiae and femora. Abdomen of male: Terga 1-S with sparse, thick hairs along posterior margins, otherwise generally bare. Ninth tergum short, its posterodorsal emargination deep, U-shaped, extending nearly to base of ter- gum; no distinct lobes beside emargination but four to six long setae along margin at each side. Gonapophyses (Figs. 164, 165) proportionately large, more than half to nearly two-thirds length of basistyles, compressed, divergent near base but con- vergent just before slender, upturned, out- wardly curved apices; penis curved down- ward at base, without other prominent curves, apex unmodified (Fig. 165) ; me- dian, membranous pouch below gonapoph- yses with acute posteromedian margin. Basistyles only moderately thickened; pos- teromesal lobe darkly sclerotized, smooth (not pilose), with acute upper point and rounded lower one. Dististyles a little nar- rowed and most strongly curved near mid- length, with subacute but inconspicuous mesal tooth at base; ventral tooth not present. Ninth sternum with dense brush of dark yellowish brown setae near pos- terior median edge. Tenth (anal) segment membranous, small, protruding only to about mid-length of gonapophyses, of nearK' uniform width throughout. Abdomen of female: Terga 1-8 only sparsely hairy except for thick hairs along [Posterior margins; sterna more hairy and with scattered darkly pigmented spots. Eighth tergum darkly sclerotized through- out, short, slender, with slight ckirsolateral thickening at each side (Fig. 169); lateral tips attenuate. Hyixn'alves of eighth ster- num \x\\c with darkened borders in apical two-thirds,- brown basally; dorsal margin curved slightly ventrad toward acute apex, ventral margin smoothly upcurved, with a thni, translucent blade-like cdii^c below mai'ginal dark band. Genital furca (Fig. 170) lyre-shaped, narrowly joincLl to ninth sternum; latter in form of two convergent, strap-like extensions of lowermost corners ot luiuh tergum, each abruptly expanded before its shallowly bifurcate tip. Cerci short, approximately 1.1-1.2 times length of hypovalves measured from notch to tip, both dorsal and ventral margins evenly upcurved in lateral aspect; color dark amber brown. Types. — Holotype, male, 1 mi S of The Crane Fly Genus Chiotiea in North America 171 Hobart Mills, on snow, 5S00 ft (1770 tn), Nevada Co., California, 20 Dec. 1971, N. D. Penny. Allotype and 1 6 , 1 9 para- tvpes, same data as holotype. Additional paratyjKs: California, Placer (^o., 3 mi N of Tahoe Vista, on snow, 7000 ft (2154 m), 20 Dec. 1971, N. D. Penny. Entire type series preserved in alcohol, in Snow Entomological Museum, Univ. of Kansas. Variation.— Over A\ length of males varies from l.i) and 2.9 mm in two ex- tremely contracted specimens (i.e., with abdomens strongly "telescoped") to 4.9 mm in a robust individual. Most males, however, are in the range of 3-4.5 mm. Females seen measure 3.1-5.1 mm, with nearly a normal distribution of lengths. As noted in the description, the terminal llagellomere is short. Two individuals (1 (^ , 1 9 ) from Mariposa Co., California, have four unusually short flagellomeres be- yond the fusion segment. Another male from the same locality has four flagello- meres (including fusion segment) on one side, five on the other. Although carolits is generally a hairy-looking Chionea, this is due largely to the setae of the legs; and individuals with shorter, less dense hairs than usual are not rare. Considerably less variation in robustness is found in males of C. carolus than in most species; that is, the smaller males are not characterized by slender femora and relatively slender basi- styles. There exists some variation in the proportionate length of the ninth tergum, which is shorter than in most species. The tlegree of development and sclerotization of the mesal tooth of the dististyle differs considerably among males; and size varia- tion in the gonapophyses has been noted in the description. Relationships .—i\.\\\\ovi^\\ in general ap- pearance C. carolus does not very closely resemble other members of the alexandri- ana group, I have placed it among them mainly on the basis of the male gona- pophyses. Antennal structure supports this placement; so also does the short (narrow- ly transverse) eighth abdominal tergum of the female. All characters considered, carolus is probably most like C. lyrata, geo- graphically its nearest neighbor, with which it shares part of its range. It may be readily differentiated from lyrata (and from other species of the alexandriana group in the Sierra Nevada) by its small size, overall hairiness and dark color, by the expanded femora with slender bases, by the shape of the male ninth tergum (deep emargination and lack of projec- tions) and gonapophyses, and by the ab- sence of the ventral tooth of the dististyle. Geographic Distribution. — A species largely of middle elevations in the north- ern and central Sierra Nevada, C. carolus has to date been collected in Alpine, Mari- posa, Nevada, Placer and Tuolumne coun- M.\p 5. Range of Chionea caioltis. Each spot repre- sents one or more collections within a county. 172 The UNivERsiTi- OF Kansas Science Bulletin ties in California and Washoe Co. in Nevada (Map 5). This is rou^^^hly from just north of Lake Tahoc to Ycjsemite National Park. In his detailed study of winter insects in Yosemite National Park, David Durbin found fairly consistent alti- tudinal separation of carohts from the higher montane C. ditrbini and other, less common members of the alexandnana group. Elevations of collection sites of C. carohts range from 5800-8600 ft (1770-2646 Seasonal Distribution. — Most specimens were found in the period 8-25 December, yet one male was collected on 4 November and another on 2 February. Probably there is a single, prolonged emergence of the adults in late autumn and early winter. Habitats. — The habitats of caroliis are chiefly in the lower lodgepole pine zone or upper ponderosa pine-sugar pine zone of the Sierra Nevada. Durbin (1975: 8) described the habitat at Crane Flat, Yo- semite National Park, as follows: "The dominant trees are Abies niagnifica (red fir), Finns larnbertiana (sugar pine), A. concolor (white fir) and /'. miirrayana (= P. con tor ta, lodgepole pine) ; other prominent parts of the biota include Libo- cedrus decnrrens (incense cedar), Arcto- staphylos sp. (manzanita), Ceanothits sp. (mountain lilac), Pniniis sp. and Phyllo- doce bretveri (mountain heather)." (Com- mon names in parentheses added by me.) Collection Records. — CALIFORNIA : Alpine Co., Carson Pass, 8600 ft (2646 m), on fresh snow 3-4 ft deep, overcast, light snowfall, 12:30-1:30 i-.m., 14 Dec. 1974, D. Durbin (U , 19); Mariposa Co., Crane Flat Meadows, Yosemite National Park, 6200 ft (near 1900 m), from log in snow, 25 Dec. 1972 {I $)\ same, but on snow, 2 Feb. 1973 (1 6 ); same, but 8 Dec. 1973 (2(5, 3$); same, but during snow- fall, 21 Dec. 1973 (19); same, but on S-10 in of powdery snow, 23 Dec. 1''73 (3 i , 7$ ); all collected by David Durbin; Ne- vada Co., 1 mi S of Hobart Mills, on snow, 5800 ft (1770 m), 20 Dec. 1971, N. D. Penny (2 6, 2 9, including types); Placer Co., 3 mi N of Tahoe Vista, on snow, 7000 ft (2154 m), 20 Dec. 1971, N. D. Penny (2 6, 19, paratypes) ; Tuolumne Co., Crane Flat Meadows (area is partly in Mariposa Co.), Yosemite National Park, 6190 ft, 4 Nov. 1972, D. Durbin (U ). NEVADA: Washoe Co., 3 mi E of Tahoe Vista, on snow, 20 Dec. 1971, N. D. Penny (l6,29). Chionea durbini, new species Diagnosis. — A brownish species of the central, high Sierra Nevada of California, with two or three flagellomeres; legs long and slender, tarsi contrastingly darker; male gonapophyses compressed but curved to present a widened dorsal aspect, trun- cate at apex and bearing a small postero- dorsal tooth; cerci slender, extending well beyond hypovalves, upturned slightly in apical half. Description (based on 17 (5 , 17 9 , pre- served in alcohol). — General appearance: Overall color of sclerites dark yellowish brown to brown, membranous surfaces dull pale brown; genital segment of male dark brown in most specimens; ovipositor of female amber brown; legs conspicuously long, tarsi much darker than tibiae. Length of male normally contracted, about 4.7-7.7 mm; length of hind leg 8.3-13.2 mm; fe- male, 5.9-9.6 mm, hind leg 7.8-10.2 mm. Head: Antennae (Fig. 171) compris- ing subcylindrical scape about three times as long as its diameter, pedicel not quite as long as scape, broadly conical or turbi- nate fusion segment, and usually two addi- tional flagellomeres (if three, most proxi- mal one very short as in Fig. 171). Setae on vertex and occiput long, slender, brown. C'omfiound eyes with 110-120 ommatidia each, more in males than in females. Thorax: Halteres pale brown, contrast- ing noticeably with thoracic dorsum in most individuals. Hairs of mesonotum, pronotum and coxae numerous, long, slen- TnK Cran'i: Fly Ghnus Chionea in North America 173 clcr, brown (not dark), loiiqer on males than on tcmalcs. Lcljs \cllo\\ish brown; hintl tcmora of males onlv modcralcK' incrassatc, especially a little betore mitl- len^th, those of females not noticeably en- lart;ed. Femoral setae in seven longitudi- nal rows in females; setae much longer and with attenuate lips in males, in four rcjws on dorsal side of femur but very numerous and not in distinct rows on lower side, l^iasitarsi yellowish brown proximally, darkening to brown in apical half; other tarsomeres dark, blackish brown, in conspicuous contrast to leg color. Figs. 171-18], Chionca dwinni. 171, right antenna of large male, lateral aspect. 172. two alKlominal tcrpa (anterior at top), to show color pattern. 173, ninth abilominal tergum, male, dorsal aspect. 17-1, left clisti- style and apex of basistvle, r.ihust male (iiolotype), posterior (ventral) aspect. 175. left dist.style of more slender male, postcromesal aspect. 17(., gonapophyses and penis, dorsal aspect. 177, gonapophyscs, posterior aspect. 17S, same as 176, riudit lateral aspect; structure below gonapophysis is dorsally concave portion ot floor of genital chamber. 17'), termmal aJKl-.minal segments, female allotype, left lateral aspect. ISO ninth sternum and genital furca, allotype, posteroventral aspect. 1«1, same, left lateral aspect. Scales: a— 1/1, 1/1- 178, I8U-181; b— 172-173, 179. 174 The UxivERsiTi- of Kansas Science Bulletin Abdomen of male: Terga 1-8 brown along mid-line and posterior and lateral margins, otherwise lighter brown uiih scattered small brown spots (Fig. 172); setae longest along posterior margins, others arising from some of scattered spots; corresponding sterna more uniformly brown. Ninth tergum (Fig. 173) with broadly U-shaped posteromedian emargi- nation; at each side of emargination a lobe about as long as its basal width, bearing approximately ten setae. (Jonapophyses (Figs. 176-178) actually compressed blades but having undulating surface sloping downward and inward from upper, outer margin, such that they appear both in dorsal and posterior aspect to have con- siderable thickness; on upper margin of each, near posterior end, a small point or tooth directed upward. Penis sinuate, with unmodified apex (Fig. 178, dashed line). Basistyles with broad, pilose postero- mesal l(;be; dististyles (F'igs. 174, 175) with subacute mesal tooth at base, ventral tooth rarely present (absent in holotype). Tenth segment lightly sclerotizcd laterally, nar- rowed near mid-length to about half basal width. Abdomen of female: Terga 1-7 with median brown stripe and brown lateral margins, otherwise pale brown with scat- tered brown spots; sparse setae arising from spots. Eighth tergum (Fig. 179) short (i.e., slender) throughout, with tliin, darkened, sclcrotized line near mid-length, from side to side. C'erci long, slender, evenly upcurved in apical half to acute tip, extending well beyond ends of hypovalves. Hypovalves nearly straight along lower edge, dorsal edge gradualK curved down- ward toward narrowly rounded a}ie\; blades mostly translucent, with amber- brf)wn borders; dark yellowish hairs along outer half to two-thirds of dorsal margin. CJenital furca lyre-shaped, its slender arms bowed both outward (Fig. 180) and dor- sad (Fig. 181). Ninth sternum formed of convergent strap like prolongations of lowermost corners of ninth tergum, each widened near attachment of furca and e.\[xuided slightly and abruptly just before lightly sclerotized median portion. Types. — Holotype, male, near Carnegie Institute Experimental Station, Inyo Na- tional Forest, Mono Co., California, on 2-6 in of snow, 9800-10,300 ft, 14 October 1973, collected by David Durbin. This locality is in the high Sierra Nevada west of Lee \'ining, at approximately 37°58'N, 119° 17'W. All()l\pe, female, and 2 6,79 para- types, same data as holotype. Paratopo- types: 13 c5 , 8 9 , 28 October 1973, at 9800 tt (3015 m). One i paratype, Yosemite National Park, Tuolumne Co., California, on snow in Dana Meadows, '^900 ft, 22 October 1972, David Durbin (locality is SW of Lee Vini ng, about 2 mi W of the Sierra crest, which is the county line). One 9 paratype, California, Mariposa Co., Yosemite National Park, Crane Flat Campground, 6200 ft (1908 m), on snow, 9 December 1973, David Durbin. Holo- type, allotype, and most paratypes in the California Academy of Sciences, San Fran- cisco; 36, 2 9 paratypes in the Snow Entomological Museum, Univ. of Kansas, Lawrence. One additional female, badly damaged, is recognized as belonging to this species but is not made a paratype; it is from 10,350 ft (3185 m) on Mt. Dana, Tuolumne Co., California, 29 Oct. 1972, collected by David Durbin. Rdiitionslups. — Chioneu durbini gener- ally resembles C. alexandrianu. its size, color and short antennae contributing to the similarity. On the basis of the gona- pojihyscs and dististyles of the male and the eighth abdominal tergum and ninth sternum ot the female, however, I judge this species to be most closely related to C. carolits, geographically its nearest neigh- bor. It may be difTerentiated from the smaller C caruliis by antennal structure, shape of femora, and details of male and female genital segments. Specifically, the usual number cjI tlagellomeres in durbini The Crane Fly Genus Chionca in North America 175 is one fewer than in caroUis. and the fu- sion segment is much shorter; the femora are much less incrassate in relation to their length; the gonapophyses of the male have an utterly ditferent shape (cf. Figs. 176 and U)4), and the ccrci of the female of diirbini are relatively much longer and more slender than in carolus. This species is named for its collector, Mr. David Durbin. who has contributed greatly to our knowledge of Chionea and other winter insects of the Sierra Nevada through his collections and field observa- ticjns. Geographical Distribution. — This spe- cies is known only from the central Sierra Nevada of California, from about 1900- 3200 m, in Mono, Tuolumne and Mari- posa counties, which are contiguous (Map 6). Its range broadly overlaps that of C. carolus, although durbini generally occurs at higher elevations. It overlaps to a lesser extent the ranges of excavata and lyrata of the more northern Sierras and of the widespread alcxundriana. Thus, C. durbini is the southernmost of the montane species of western North America. Seasonal Distribution. — Specimens from high elevations (above 3000 m) were col- lected from mid to late October; those collected in December are from the lower elevations recorded. It seems probable that the weather at the type locality and others in the high Sierras is too severe in Decem- ber for Chionea to be active. Habitats. — This is a species of high ele- vations in the Sierra Nevada, where the subalpine forest of lodgepole pine (Pinus contorta) and white-bark pine {P. albi- caulis) is patchy, giving way here and there to small willows {Salix spp.) and Sierra junipers (funiperus occidentahs) just below tree-line. At the Carson Pass locality, mountain hemlock (Tsuga 7ner- tensiana) and red fir {Abies magnifica) were additional fcjrest trees. While the trees are, of course, not the habitat of Chionea, they characterize the general hab- itat. Chionea excavata, new species Diagnosis. — A western montane species of brown to grayish brown color, with antennae of four (rarely three) flagello- meres; male gonapophyses hooked at tip, dististyles shallowiy concave on outer (up- per) curvature; female cerci evenly curved, not much longer than hypovalves; ninth stenumi mostly lightly sclerotized, with narroNN'ly triangular darker median struc- ture. Description (based on 3 c^ , 7 9 , pinned, and 2 i , 19, originally pinned but re- moved to microscope slides). — General ap- pearance: Head dark brown, thorax brown to gr.i\ish brown; abdominal sclerites brown, except ninth segment of male dark Mm- (>. Ranpc of Chionea durbini Each sjiot rep- resents one or more collections within a county. 176 The University of Kansas Science Bulletin brcnvn, cerci of female yellowish; pleural membranes light brown. Length of male about 4-6 mm, female about 6-7 mm, ex- clusive of antennae. Head: Antennae (Fig. 1S2) comprising scape, pedicel and lour flagellomeres (three in one specimen); fusion segment short- conical, its apical portion constricted at base in some specimens, resembling addi- tional short segment. Eyes composed of about (SO-100 ommatidia each. Thorax: Haltcrcs [xile, contrasting with color of thoracic dorsum. Legs dark yellowish brown; seven (four dorsal, three ventral) longitudinal rows of setae on fem- ora, sometimes an indication of fourth ventral row in male; tarsi brown. Abdomen of male: Posterodorsal edge of ninth tergum (Fig. LS3) with broadly V-shaped median emargi nation; at each side of emargination a low, blunt lobe bearing about six setae. Gonapophyses (Figs. 1S7, LSS) broad at base, abruptly narrowed in distal half; apical portion hook-like, slender, curved tipward, with blunt ventral hcel-likc process and acute tip; space between gonapophyses narrowly U-shaped; apex of penis unmodified. Rasistyles thick, each with a bluntly rounded, pilose, posteromesal lobe (Fig. 184). Dististyles (Figs. 184-186) yellowish brown to reddish brown, thick at base, with blunt mesal tooth, somewhat flat- tened and shallowly excavated dorsally in apical half; excavated area largely pilose, also bearing scattered hairs as on remain- der of outer (dorsal) surface of dististyle; ventral tooth small and blackened (holo- type) to rudimentary. Tenth segment broad at base, narrowed near mid-length, darkly pigmented and pubescent laterally. Abdomen of female: Eighth tergum short (i.e., slender) throughout. Hypo- valves of eighth sternum deep, blunth' rounded at tii^ (Fig. 189), with lower mar- gin more curved than dorsal margin. Arms of genital furca slender throughout; basal portion small, subrectangular, not much l(jnger than wide. Ninth sternum with strongly sclerotizecl, narrowly subtriangu- lar, projecting mid-ventral porti(jn and a subquadrate, more lightly sclerotized and dillusely pigmented area above and to each side (Fig. 190). Cerci of medium length for genus, about 4.5-5 times as long as greatest width, evenly curved on both dorsal and ventral margins, about 1.2 times as long as hypovalves. Types. — Holotype, male, 8 mi SW of Reno, Washoe Co., Nevada, elevation 7200- 8200 ft, 19 January 1923, H. S. Barber col- lector. Allotype, female, saine data as holo- type. Parat\pes: lo, 19, same data as holotype; 2i, 4 9, northeast of Hunter Lake, Mount Rose, Washoe Co., Nevada, 7200-8200 ft, 19 January 1923; 29, near Caribou, Plumas Co., California, 27 De- cember 1922; \i, near Caribou, Plumas Co., California, 24 January 1922 (1923.= ); all collected by H. S. Barber; 1 9 , Washoe Co., Nevada, 3 mi E of Crystal Bay, 7300 ft, 20 Dec. 1971, N. D. Penny. Holotype, allotype, three male and six female para- types in United States National Museum, Washington, D.C.; one male and one fe- male paratypes (on slides) in Univ. of Michigan Museum of Zoology, Ann Ar- bcjr; one female paratype in Snow Ento- mological Museum, Univ. of Kansas, Lawrence. Several female specimens from farther south in the Sierra Nevada apparently be- long to this species. Since there are no associated males, however, these have not been made paratypes. All are in the col- lection of the California Academy of Sci- ences, San Francisco. The collection rec- ords are: CALIFORNL\: Alpine Co., C'arson Pass, ofT California Hwy. 88, on snow, 8600 ft, 8:15 p.m., clear night, near 32°F, 14 Jan. 1975, D. Durbin (19); same place, but specimen found dead on snow, 1 I'.M., clear day, 55°F, 15 Jan. 1975, D. Durbin (19); Mono Co., Inyo Na- tional Forest, Tioga Pass, Dana Meadows, 9900 ft, 26 Oct. 1968, R. P. Papp (29); The Crani; Fly Gi-nus Chionea in North America 177 liu'o National Forest, ("lariiei^ic Inst. Ex- pcr. Sta., 2S Oct. F-^73, U. Durbiii (19); Tuolumne C^o., Yosemite National i'ark, Mt. Dana, 11,000 ft, 2') Oct. 1972, D. Dur- ban (1 ? ); same general are.i, Dana Mead ows, 10,000 ft, 15 .April 1973, D. Durhin (19); same, but 16 April 1973 (19); same area, but (iaylor Peak, 11,000 ft, collected on snow, "() VI 1973,"" I). Durbin (19) (Mr. Durbin states tbat the June date is correct but that (he specimen may not have been alive when found). V anatiou . — Femoral setae in two of the m,iles are very attenuate at the tip, many 184 185 CWh 187 188 186 ' mm _J Figs. 182-190, Chionea excavata. 182, right antenna of male, lateral aspect. 183, posteromedian portion of ninth abdominal tcrgum and tenth segment, male holotypc, dorsal aspect. 18-4, left dististyle, robust male (holotype), posterior (ventral) aspect; depression in outer, upper surface indicated by dashed line. 185, left dististyle of more slender male, posterolateral aspect (large hairs omitted). 186, same, dorsolateral aspect (ven- tral surface to right). 187, gonapophyses, dorsal aspect. 188, right gonapophysis and penis, right lateral aspect. 189, terminal abtlominal segments, female paratypc, left lateral aspect. 190, ninth sternum and genital furca, composite from two females, ventral aspect. Scales: a — 182-188, 190; b — 189. ■8 The University of Kansas Science Bulletin tips slightly curled. The cerci of the fe- male vary from 1.1-1.2 times as long as the hypovalves (measured from notch to tip). Variation in color and size have already been mentioned. The holotype is a larger male than the others and is some- what more darkly colored hut was selected because both antennae were intact. It has a dried nematode capstile around its neck. Relationships. — Chiotiea cxcavata be- longs to the alexandriana grou^) and with- in this group seems, on the basis of the male genital structures, particularly the gonapophyses, most nearly akin to alex- andriana and nigra. From both these spe- cies it dilTers in general coloration and number of flagellomeres. Males of exca- vata can be distinguished from those of Map 7. Ranges of C.hiotica cxciirala (circles) and C. jellisoni (triangles). I'.ach symbol represents one or more collections within a county. all other species by the shallowly concave tips of the dististyles; the specific name (Latin excavata = hollowed out) refers to this unique characteristic. There is a close resemblance between females of ex- cavata and those of lyrata n. sp. in the structure of the ninth sternum; however, excavata has relatively shorter cerci, a smaller number (;f flagellomeres, and a less distinct color pattern on the abdominal terga. In lateral aspect, the ventral mar- gins of the hypovalves in excavata are more curved than the dorsal margins, while in lyrata both margins curve about ec]ually toward the bluntly rounded apex. Geographic Distribution. — This seems to be a species of the upper slopes (2215- 3385 m) of the northern and central Sierra Nevada of California and westernmost Nevada (Map 7). So few specimens are known that the actual range is still un- clear. This is the "species 1" recorded by Alexander (1%7: 121), who through a misunderstanding reported the holotype from California (because I had sent him only the California records for the species). Seasonal Distribution. — The very wide span of dates on which adults have been found, from late October to at least mid- April, suggests there may be two genera- tions per year. The June collection record is unusual,_yet not particularly to be doubt- ed; apparently, suitable environmental con- ditions still prevailed at the time the specimen was collected. Habitats. — Elevations in the Sierra Ne- vada at which C. excavata has been col- lectetl are mainlv subalpine, characterized b\- mixed coniferous forests of lodgepole pine (Ptnus contorta), white-bark pine (/'. alb/caulis), red lir (Abies magnifica), mountain hemlock {Tsuga niertensiana), and western or Sierra iunijier (funiperus occidcntalis). The higher recorded sites are near local tree line, where the white- bark pine and small jumpers and willows arc characteristic vegetation. Tiu; Crank Fly Gknus Chionea in North America 179 Chionea jcllisoni, xfav species Diagnosis. — A western montane sjxxies with antennae haviiii; three flai^ellomercs; male ^onaiiophyses broad at base witli slender tips turned upward and outward; female cerei curved eveiiK' tqiward in api- cal half, very short (not quite as long as h\p()valves) but wide in dtjrsal aspect, especialK' in basal half. Description (based on 31 (5 , 50 9 pre- served in alcohol, 76,69 pinned). — Gen- eral ap[x\u-ance: Head reddish brown to dark brown, antennae and mouthparts slightly paler; thorax tlark yellowish brown to brown, a little paler dorsally between halteres; abdominal sclerites yel- lowish brown to light brown except ninth segment oi male reddish brown to dark brown; pleural membranes pale, dull yel- lowish brown; cerci and apical half of hypovalves of female translucent yellow- ish. Length of male, exclusive of antennae, about 5.0-5. S mm (holotype 5.S mm); length of female about 6.5-7.4 mm (allo- type 7.2 mm). Head: Antennae (Fig. 191) comprising scape and pedicel of about equal length, and three flagellomeres; fusion segment broadly conical, slightly asymmetrical, wider than long, with apex somewhat pro- longed, giving ap[iearance of another short flagellomere; terminal hairs of flagellum a little longer than all antennal segments together. Setae on vertex pale yellowish brown to light brown. Com[iound eyes with about 140-150 ommatidia each. Thorax: Color abruptly paler than that of head, evenly dark yellowish brown or brown, except for paler area on meta- thoracic dorsum. Halteres yellowish to brownish gray, slightly darker than ab- dominal pleura but not conspicuously paler than thoracic sclerites generally. Setae of mesonotum long, stout, not darkly sclero- tized, in two loiigitudinal patches. Legs yellowish brown to light brown; tarsi only slightly darker thaii femora and tibiae. Femora with six or seven well defined rows ot dark brown setae; tibial setae more slender th.in those of femora, slightly longer, also in readily discernible rows. Abdomen of male: Terga and sterna evenlv pigmented, without evident color pattern, except ditltisc paler spots on terga of some individuals; tergal setae sparse, short, mostly along posterior margins of terga; sternal setae longer, more dense, especially at sides of sterna. Posterodorsal margin ot ninth tergum (Fig. 192) with two prominent lobes separated by broadly V-sha[ied incision; api^roximately six setae of various lengths on or near apex of each lobe; setae laterad from lobes often longer than those on lobes. Gonapophyses (Figs. 193, 195, 196) broad in basal half, with slightly concave dorsal surface irregularly and shallowly emarginate at posterior edge; apical half becoming abruptly nar- rowed to elongate, slender, hook-like tip curving upward and outward; gonapophy- ses nearly in contact slightly beyond mid- length; apex of penis unmodified. Basi- styles thick, their posteromesal lobes blunt- ly rounded or with two low prominences, with scant pilosity. Dististyles (Fig. 194) slightlv paler than basistyles, evenly curved on outer surface, ridged along most of inner curvature except near mid-length, somewhat expanded slightly beyond mid- length. Mesal tooth of dististyle promi- nent, broadly subconical; ventral tooth also prominent, darkly sclerotized, usually with short, chisel-like apex (more nearly conical in holotype). Tenth segment only lightly sclerotized and pigmented laterally. Abdomen of female: Terga and sterna tmitormlv [ligmented; setae sparse, not much d.nker than sclerites, distributed about as in male. Eighth tergum a nar- row, transverse strap, at widest (i.e., long- est) only about one third length of seventh tergum, without setae. Hypovalves deep, onl\' slightly longer than basal portion of eighth sternum, with both ti[iper and lower margins curving gently toward tips. Cerci 180 The University of Kansas Science Bulletin slightly shorter than hypovalvcs (ratio of 0.94 to 1.0), depressed, appearing of about uniform width throughout in lateral as- pect (Fig. 199) but actually conspicuously widened in basal half (Fig. 197b), tapering toward acutelv rounded apices. Ninth ter- gum without posterolateral setae; tenth tergum short, wide, with only slight indi- cation of transverse sulcus near posterior margin (Fig. 197b). Basal portion of geni- tal furca longer than wide, weakly sclero- tized medially; arms of furca strongly bowed outward near mid-length, thick- ened near bases (Fig. 200). Median por- tion of ninth sternum more densely sclero- lized marginally than centrally, slightly concave ventrally, with bluntly rounded median tooth and irregular, lightly pig- mented areas in adjacent membrane (Figs. 200,201). Types. — Holotype, male, Bcjulder Creek, Ravalli Co., Montana, on snow, 9 Decem- ber 1956, collected by W. L. Jellison and R. Scott. Allotype, female, and 1 (5 , 4 9 parat)pes, same data as for holotype. Ad- ditional paratypes: MONTANA: Ravalli Co., West Fork Boulder Creek, on snow, 29 Jan. 1966, W. L. Jellison (1 9 ); Lake Como Road, on snow, 27 Nov. 1965, W. L. Jellison (2 9 ) ; Tin Cup Canyon, 3 mi W of Darby, on snow, 1 Dec. 1970, W. L. Jellison and J. E. Keirans (2 S , 3 9, in- cluding a mating pair); Missoula Co., Elk Creek, Lubrecht Forest, 4000 ft, 22 Jan. 1972, R. A. Haick (14^, 25 9, including one mating pair). IDAHO: Boise Co., 0.25-0.5 mi E of Idaho City, 21 Dec. 1974 (45); same, but on snow in pine forest, 7 Dec. 1974 (U ) ; H mi N of Idaho City, on snow in pine forest, 7 Dec. 1974 (19); same locality, 30 Nov. 1974 (29); Idaho City, 14 Dec. 1974 (1 6 ) ; same locality, on snow, 16 Jan. 1976 (1 ;e ot central Utah, where nigra contacts tlie similar C jcllisoni. Scdsoudl Distribution. — Ihere are plain- 1\- two separate periods of occurrence of adults on the snow in the type locality, one in October ami November, the other in late bY'bruary, March and April. Since Osburn's study of the area was continuous and since he \\'as watchini; for Chionea particularly during the entire 1952-1953 season, I believe that the interval between these peaks is actually a time when adults of Chionea are not present and that the peaks may represent two generations. Habitats.— \ am indebted to Mr. Os- burn for several very useful observations on the environments in which this species occurs and for searching beyond the areas in which the insects were found, so that the pattern of ecological distribution was made clear. Chionea nigra was collected on 12 dilTerent dates in three years. In every instance the specimens were found on snow, although the extent of the snow cover varied from shallow and patchy in distribution to deep and completely cover- ing the ground, the condition of the snow from freshly fallen and soft to several days old, with an icy crust at the surface. Col- lections made between 9500 and 11,200 ft elevation were in lodgepole pine (Finns contorta) forest, except where as a result of fire these trees had been temporarily replaced by aspen, (jr, in one case, limber [line. Above 11,200 ft the forest is spruce {Picea engebnanni) and fir {Abies lasio- carpa) to timberline, which is near 11,500 ft in that vicinity. Chionea nigra was never found more than 100 ft from trees projecting above the snow level, while a species of Boreiis often found in asso- ciation with the tipulids was taken as high as 700 vertical feet above timberline (see B\ers, 1955: 4). Four of the 12 collections, C()m[-»iising nearly half the specimens, were made when the temperature was between 1••••• W^ • • • • • 4 '-•.•-•-•.•-•-< •-•.•-•.•.•>*••-••-•.•-•.•.< ^ s s THE UNIVERSITY OF KANSAS SCIENCE BULLETIN co^ UNI^ i I NEOR THOL OMUS, A NEW GENUS OF ORSILLINI (HEMIPTERA— HETEROPTERA: LYGAEIDAE: ORSILLINAE) .♦. g f.< By Steven W. Hamilton ,v v i ifi S g Vol. 52, No. 7, pp. 197-234 April 8, 1983 g ANNOUNCEMENT The University of Kansas Science Bulletin (continuation of the Kansas Uni- versity Quarterly) is an outlet for scholarly scientific investigations carried out at the University of Kansas or by University faculty and students. Since its incep- tion, volumes of the Bulletin have been variously issued as single bound volumes, as two or three multi-paper parts or as series of individual papers. Issuance is at irregular intervals, with each volume prior to volume 50 approximately 1000 pages in length. Effective with volume 50, page size has been enlarged, reducing the length of each volume to about 750 pages. The supply of all volumes of the Kansas University Quarterly is now ex- hausted. However, most volumes of the University of Kansas Science Bulletin are still available and are offered, in exchange for similar publications, to learned societies, colleges and universities and other institutions, or may be purchased at $20.00 per volume. Where some of these volumes were issued in parts, individual parts are priced at the rate of 2 cents per page. Current policy, initiated with volume 46, is to issue individual papers as published. Such separata may be purchased individually at the rate of 3 cents per page, with a minimum charge of $1.00 per separate. Subscriptions for forthcoming volumes may be entered at the rate of $20.00 per volume. All communications regarding exchanges, sales and subscriptions should be addressed to the Exchange Librarian, University OF Kansas Libraries, Lawrence, Kansas 66045. Reprints of individual papers for personal use by investigators are available gratis for most recent and many older issues of the Bulletin. Such requests should be directed to the author. The International Standard Serial Number of this publication is US ISSN 0022-8850. Editor William L. Bloom Editorial Board Philip W. Hedrick Rudolf Jander Harvey Lillywhite Charles D. Michcncr Norman A. Slade Henry D. Stone George W. Byers, Chaivmun S\7J\ THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Vol. 52, No. 7, pp. 197-234 April 8, 1983 NeortJioIomus, A New Genus of Orsillini (Hemiptera — Heteroptcra: Lygacidac: Orsillinac)' LIP Steven W. Hamilton^ niiA /. c Department of Entomology, University of Kansas, Lawrence, Kansas 66045 riARVA.KI ' UNIVElf Contents Abstract 197 Introduction 197 Materials and Methods 198 Phylogenetic Relationships in Neort/iolomus 199 Genus Neortholomits, New Genus 201 Key to the Species of Neortholomus 202 Neortholowus arphnoides (Baker) 203 Neortholomus gibhijer (Berg) 206 "'Neortholomus jamaicejisis Species Complex" 208 Neortholomus ]amaicensis (Dallas) 208 Neortholomus rubricatus (Berg) 212 Neortholomus \oreshanus (Van Duzee) 215 Neortholomus nevadensis ( Baker) 217 Neortholomus procerodorus, New Species 219 Neortholomus scolopax (Say) 222 Neortholomus usingeri (Ashlock) 226 Acknowledgments 228 Table 1 and Figures 230 Abstract A new genus, Neortholomttf, is erected to include only New World species of Ort/iolo?)uis. This refinement is based primarily on characters of the male and female genitalia. Nine species arc recognized in this genus, including a new species, Neortholomtis proccrodonif. These species arc described, and external and internal characters illustrated. .\ key to the species is provided. A proposed phylogeny is presented and three species groups arc recognized. The scolopax group includes arphnoides janiaicensis l^orcslianiis rtihricattis, and scolopax; the gibhifcr group is comprised cjf gibhijer, procerodorus, and usingeri; nevadensis is the only member of the third species group. Introduction kind used to delimit other genera of Orsil- In a revision of the Orsillinac, Ashlock ,^,^r~\ 1 I- I 1 1 1 (L 'Contribution number 1851 from the Department ( 1%/) CStahhshed the modern concepts Ot ^^ Entomology, University of Kansas, Lawrence, Kan- thc included tribes and described genera. sas 66045. TT -J r J /^ 1 1 -11 1 "Present address: Department of Entomology, He identihed Urtholornus as iiossibly poly- ,- , ■ i w? \ n t n\ „, ,, tT,.;v,.r.;tv ri,-m^nn 1 J I J Fisheries and Wildlire, Clemson Univcrsit), Clcmson, phyletic because it lacks characters of the South Carolina 29631. 198 The University of Kansas Science Bulletin lini, such as femoral spines, mesopleural overlap of the propleuron, elongate anten- nal segments, modified bucculae, and so forth. Ashlock stated that "species are as- signed to Ortholomus if they have the characters of the Orsillini but lack the dis- tinctive features of the other nine genera in the tribe." Baker (1906) noted that the Nysitts subgenus Ortholomus was "a perfectly valid genus, much better founded than are numerous others of the family Lygaeidae." Three new species and a variety of O. longiceps Stal 1874, were very briefly de- scribed by Baker in this same paper. Although Baker (1906) recognized that the straight costal margin distinguished Stal's (1872) subgenus Ortholomus from other Nysitts in North America, this char- acter is not sufficient to separate Ortholo- mus from many other closely allied genera. He was unaware that the two most widely distributed species, O. scolopax (Say) 1831, and O. jamaicensis (Dallas) 1852, were al- ready described. Two of Baker's proposed taxa, cool{ii and uhleri, respectively, are a common color variant and an abnormal specimen of O. scolopax. His descriptions were extended couplets in the keys, insuf- ficient for the recognition of these species. Thus, leaker's revision of American Or- tholomus, the largest and most recent treatment of the genus, is inadecjuate. No comparisons of the New and Old World species of Ortholomus have been undertaken. Wagner (1958) included Or- tholomus in his revision of European or- sillines, but he did not compare the three named Old World sp>ecies with those in the Americas. The cvoluticjnary relationships of Or- tholomtis, Neortholomtis, a new genus recognized herein, and the other genera of the tribe Orsillini will be presented in a separate paper. Materials and Methods Nearly 4,800 specimens, most of which were borrowed from the institutions and individuals listed in the acknowledgments, were examined. Type specimens to be c(jmpared with one another and with other determined and undetermined specimens were borrowed when possible. A few field and laboratory observations were made on Neortholomus scolopax, a widespread North American species, to supplement the study of museum speci- mens. The remainder of the biological data used herein has been gleaned from the literature and from label data on indi- vidual specimens. The male and female genitalia of all species were dissected and examined for characters useful in species separation or phylogenetic analysis. In males, the para- mere, aedeagus, and sperm reservoir were examined, and in females the spermatheca was studied. For study, the aedeagus was removed from the pygophore, or male genital cap- sule, and inflated using osmotic pressure (Ashlock, 1957, 1967). For the most part Ashlock's 1967 method was used in this research although a mild solution of potas- sium hydroxide (KOH) was used as the initial wetting and macerating agent as in the 1957 method. The technique for dissection of the spermatheca has been outlined by Ashlock (1967) and is more fully presented by Hamilton (19S0). Akhough Ashlock (1967) notes that "no inflation is neces- sary" to study the structure of the sperma- theca, care must be taken because inflation can occur if it is removed from KOH and placed in water. The central [lortion of the spermathccal pump is weakly sclero- tized so that it can expand and contract with changes in osmotic pressure. For this reason, the overall length of the spermathccal pump is a very poor charac- Nfortholomus, A New Gknus of Orsillini 199 tcr and was not used in this study. This pressure ean also cause the terminal hulh t(j collapse or to expand and crack. The synonymies include valid names and their synonyms, and all relerences since VHA. C>)mplete lists of references can be found in Slater (1%4). Descriptions are based on the primary ty[ie in each species. Parentheses around data in the description indicates characters that could not be studied on the type speci- mens owiiiiT u) obstruction or loss. These values or characters were obtained from other specimens, either in the type series or amoiiL:; the other available specimens, which were judi^ed similar to the type. The summaries of species distributions are based on the data gathered from the specimens I have seen. Detailed data from every specimen have been left with Dr. Peter D. Ashlock of the Department of EntomoloiTv at the University of Kansas. The dimensions used in the descrip- tions are illustrated in Figure 2. The height of the mesopleural evaporative area was measured by comparing the distance from the dorsal apex of the scent gland auricle to the upper margin of the evap- orative area with the distance from the dorsal apex of the auricle to the ventral margin of the metapleural callosity (Fig. 6d). The distances above the apex of the auricle were measured on a line perpen- dicular to the plane of the dorsal surface cjf the specimen and not parallel to the su- ture between the meso- and metapleura. The evaporative area has a granular and nonglossy appearance, which may be ob- scured on dirty or greasy specimens. The callosity is a smooth, often glossy, elongate area near the dorsum of the metapleuron. The length of the scent gland auricle was determined by comparing the distance from the ventral tip of the mesepimeron to the dorsal apex of the auricle with the distance from the ventral tip of the mese- pimeron to the ventral margin of the meta- [ileural callosity (Fig. 6c). These distances were taken along a line from the tip of the mesepimeron to the apex of the auricle, w hich is roughly perpendicular to the dor- sal surlace ot the specimen. The mese- pimeron is the area of the mesopleuron posterior to the jileural suture (Fig. 6d). (characters used in the analysis of phy- logeny are either complex derived charac- ters or unic|ue evolutionary innovations in the group, that is, apomorphies. When such derived characters are shared with other taxa, these synapomorphies demon- strate various phylogenetic relationships of the taxa. Such characters have been used here to infer the phylogenetic (cladistic) relationships of the species of Neortholo- nuis. Phylogenetic Relatioxships ix Neort/wlomiis The proposed pihylogeny of the in- cluded species of Neortholomus is pre- sented in Figure 1. The characters used in this phylogeny, the apomorphous and plesiomorphous states of these characters, and references to illustrations of the char- acter states are summarized in Table 1. The genera Ortholotyms, Oysillns, and SinorsiUns have been used as out-groups for comparison with Neortholomus. The synapomorphy establishing the holophyly of the four-genus Orsilliis group is the presence of complex lobes on the vesica of the aedeagus (Fig. 4b; Ashlock, 1967 Figs. 3a, 4a) between the sperm reservoir and the pigmented lobe. This character and others will be discussed in another paper on the phylogeny of the Orsillini. Based on this phylogeny the N eortholo- 7niis progenitor had a female with an elongate duct between the spermathecal [lump and the terminal bulb, and a short, stout sclerotized basal portion of the pump, while the male had the paramere shank without a carina and the gonoporal process of the aedeagus was surrounded by little 200 The University of Kansas Science Bulleti N or no inOatable membrane. The sperma- thecal pump of the Neortholomiis ancestor completely lacked the dorsal flange and had a reduced basal flange, conditions unic]uc in the Orsillus branch of the Orsil- lini. I have recognized three species groups in Neortholomiis, the scolopax, gihbifer, and nevadensis species groups. The carina on the shank of the paramere and the lack of any flange at the base of the spermathe- cal pump clearly delimits the scolopax species group, which includes arphnoides, jamaicensis, }{oreshaniis, riibricatits, and scolopax. All of these species except scolo- pax have the elongate basal sclerotized area and a large evaporative area. The size of the evaporative area has not been used in this analysis because it is somewhat variable in the various species and is neither unicjue nor complex. The gibbifer species group comprises gibbijer, procerodonts, and iisingeri. These three species have a well developed gono- poral membrane. In gibbijer and iisingeri the basal flange of the spermathecal pump is further reduced but not completely lost. 1 he greatly shortened duct between the spermathecal pum[-) and the terminal biilh is not found in nevadensis but is shared by the gibbifer and scolopax species groups indicating these two are sister groups. Nevadensis shares no apomorphies with the other Neortholomiis s[iecies, ex- cept those delimiting the genus, and thus represents the most primitive lineage in it. A very short shank of the paramere (Fig. 4a) represents an auta|iomorphy uint]ue to nevadensis among all species of the Orsillus branch of the Orsillini. Other characters which may be increas- ingly used in the future are the chromo- some number, shape, and behavior. In a recent study of the cytotaxonomy of ly- gaeids, chromosome studies of four species of Neortholomiis were reported (Ueshima and Ashlock, 19(S0). The chromosomes are holocentric in Heteroptera which means that the centromere is not localized (mo- noccntric) but instead occurs along the entire length of the chromosome. If one of the chromosomes is broken, the frag- ments will behave normally and "perpetu- ate themselves during meiosis" (Ueshima, V)7')). Ueshima and Ashlock (1980) found that nevadensis and iisingeri have a diploid chromosome number of 14 with "five pairs of autosomes, an m-chromosome pair, and an XY sex pair," and arphnoides and scolopax have 16 chromosomes with "six pairs of autosomes, an m-chromosome pair, and an XY sex pair." They suggest that the diploid number of 16 is derived from the ancestral number of 14 chromo- somes. Evidence of this conclusion is an intermediate-sized autosome pair in the 14-chromosome species that is not found in the 16-chromosome species. The 16- chromosome species have two small auto- somes which are probably the fragments of the broken intermediate autosome. The large autosome is always present in Neor- tholomiis. In the Nysiim all species have 14 chro- mosomes (except one species with 22) while in the Metrargini 16 chromosomes is the rule except in the apjiarently primi- tive Darii'inysius, which has a diploid number of 14 (Ueshima, 1979; Ueshima and Ashlock, 19(S0). They point out that the metrargine change from 14 to 16 chro- mcjsomes apparently involved the frag- mentation of the large autosome and not the intermediate autosome. Therefore, the increase in chromosome number is an in- dejKMident event in Neortholomiis, unre- lated to the seemingly similar change in the metrargines. Interestingly, both of the derived 16- chromosome Neortholomiis species are members of the well-delimited scolopax s[iecies gnnip and the 14-chromosome spe- cies are found among the other groups. Although more species of Neortholomiis Neortholomiis, A Ni:w Genus of Orsillini 201 need to be studied cytogenetically, the presence of 1() chromosomes may be an- other s\ na[iomorphy for this entire species L;rou|">. This prediction can be made since it is Linhkely, i;iven the synapomorjihies above, that scolopax and aiphnoides share an ancestor with l6-chromosomes which is not also shared with the other three members ot the scolopax species group. A new genus, Neortholomits, is pro- posed herein to comprise the New World s[iecies of Ortholomiis Stal scnsti lato. Ortholoniiis scnsu stncto will consist of the three Old World species: pitnctipennis (Herrich-SchaefTer), carinatiis (Lindberg), and jordani Hobertlandt. I have been able to studv specimens only of piinctipcnnts, the tvpe species, but I assume that these three species are very closely related. In light of the substantial variation known in widelv distributed Neortholomiis, it is pos- sible that carinatiis and jordani are junior synonyms of piinctipennis. In his study of the Palaearctic Nysius complex, Wag- ner (195.S) recognized these as species, but it seems clear from his characters that they are, at least externally, very close. For these reasons, as well as their overlapping Old World distributions, I have assumed that the genitalic characters for these three species will be similar, if not identical. Ashlock (1%7) gave characters to dis- tinguish Ortholomiis s. I. from other gen- era of Orsillinae. In a paper on the phylog- eny of New World Orsillini to follow this [laj^T more characters of this nature will be given for Ortholomiis s. s. and it will be redescribed in keeping with the eleva- tion of the New Wcjrld members to ge- neric rank. No single external character has been found which distinguishes all of the spe- cies of Neortholomiis from Ortholomiis piinctipennis. These two genera can read- ily be separated by difTerences in charac- ters of the male and female genitalia. The spermatheca of Ortholoiyiiis has well-developed flanges at both ends of the spermathecal pump, as well as a con- voluted duct of mf)derate length to the terminal bulb and a long unsclerotized [Portion (Ashlock, 1967 Fig. 8c). In Ne- ortholomiis, the spermathecal pump has no distal llange and a greatly reduced or lacking basal flange (b'ig. 3). The duct to the terminal bulb is much shorter in Neortholomiis than in Ortholomiis (ex- cept in N. nevadensis, Fig. 3c), as is the basal unsclerotized duct. Other useful characters that distinguish Ortholomiis and Neortholomiis are found on the aedeagus. The gonoporal process of Ortholomiis (Ashlock, 1967 Fig. 4a) is a long spiral with no attached inflatable membrane. Neortholomiis has a relatively short gonoporal process with varying amounts of attached membrane extending to near the distal qx\<\ (Figs. 4b-d, i, j). In Neortholomiis the sperm reservoir of the aedeagus (Figs. 5c, d) appears to be in a relatively primitive condition for the Orsillini. The body of the sperm reservoir is reduced or lacking, and the wings are fused dorsally over the ejaculatory duct. The lateral [Portions of the wings may or may not extend ventrally for a short dis- tance. In Ortholomiis (Figs. 5a, b) the lateral portions of the wings extend ven- trally and fuse beneath the ejaculatory duct to form a complete ring or collar ab(;ut the duct, a derived condition. The uniqueness and homogeneity of the spermathecae and the other genitalic characters of Neortholomiis indicate that this group of species is a monophyletic grouj) deserving full generic status. Geims Neortholomiis, New Genus Type species: Lygaeiis scolopax Say, 1S31. Cieneric description: Form moderately elongate to elongate, not depressed; body covered with short, pale, appressed pu- bescence, in some species mixed with 202 The University of Kansas Science Bulletin sparse, short to moderately long, erect, pale hairs. Head moderately stout to elongate, produced before eye for distance slightly greater than length of eye to about two times length of eye; width of head including eyes less than width at posterior margin of pronotum; antennifer- ous tubercle not extending beyond point of insertion of antenna. Buccula impunctate, high anteriorly, tapering to near base of head or to point short of this (usually about mid eye level) and continuing to near base of head as a low carina; labium extending to point between middle and hind coxa (procerodorits) or nearly to apex of abdo- men ( I^oreshanits). Vertex low, when viewed laterally, level with high point of eye or extending above eye one-fourth height of eye, frequently more flattened between eyes, sometimes strongly convex (arphnoides), with carina extending an- teriorly from ocellus paralleling margin of eye to base of antenniferous tubercle; eye sessile to substylate, always prominent. Scutellum with a moderate to pro- nounced and swollen Y-shaped elevation, apex of scutellum acute and slightly up- turned. Evaporative area small, only sur- rounding scent gland auricle on meta- pleuron, or larger, including mesepimeron along posterior edge of mesopleuron and nearly attaining lateral callosity on meta- plcuron. Scent gland auricle typically short and broad, about one-third height of meta- pleuron, sometimes elongate and narrow being about one-half height of meta- pleuron. Hemelytron complete, partly ex- posing connexivuin laterally, terminating short of or slightly beyond tip of abdo- men, corium iinininctate; costal margin straight, coincident with vein R + M at least to level of apex of scutellum; branch M of R + M intersecting apical margin of corium at a pf)int closer to intersection of vein Cu than oi vein R with apical margin of corium, branches often obscure, corium nearly clear to opat^uc light-brown between veins; vein lA of membrane fre- quently turned toward vein Pcu, cell never completely closed, membrane with numer- ous transverse wrinkles. Spermatheca lacking distal flange on [Himp, distal edge rounded, basal flange very reduced or lacking, duct between pump and bulb usually short, occasionally longer ( ncvadensis). Aedeagus with com- plex lobes on vesica between sperm reser- voir and pigmented lobe, gonoporal proc- ess moderately inflatable to lacking; sperm reservoir narrow, simple platelike structure dorsally positioned over sperm duct, con- nected by narrow duct distally on reser- voir, sides occasionally turned ventrally, hut never joining below duct. Characteristics to separate this genus from Ortholomns are in the genitalia, as outlined above. Included species: arphnoides (Baker), 1906, Ortholomns; gibbijer (Berg), 1892, Nysiiis; jamaicensis (Dallas), 1852, Ny- siits; /{oreshanns (Van Duzee), 1909, Be- lonocliiUis; nevadensis (Baker), 1906, Or- tholomns; procerodoriis Hamilton, new species; rubric at us (Berg), 1879, 'Nyshis; scolopax (Say), 1831, Lygaeus; usingeri (Ashlock), 1972, Ortholomus. Etymology: Neos, Greek, meaning new. In reference to the elevation of the included species to a new genus near Or- tholomus and to the New World distribu- tion ol the genus. Key to the Species of Neortholomns 1 Height of evaporative area from apex of auricle to margin of evap- orative area directly above at least half distance from apex of auricle to metapleural callosity (Figs. 6c, d) ., 2 — Height of evaporative area from apex of auricle to margin of evap- orative area directly above less than hall distance from apex of auricle Neortholomus, A Nr.w Genus of Orsillini 203 to metaplcural callosity (Figs. 6a, h) 6 2(1) Third labial segment reaching be- yond hind coxae; head elongate, width oi head including eyes sub- ec]ual to length of head before ocelli (Figs. 7a, d) ; Florida, Isle of Pines N. \oreshanus — Third labial segments sometimes attaining abdomen between hind coxae, but never distinctly exceed- ing coxae; head not so elongate, head width including eyes at least 1.2 times length of head before ocelli (Figs. 7b, c, e-h) 3 3(2) Scent gland auricle elongate, dis- tance from ventral tip of mes- epimeron to apex of auricle about two-thirds distance from ventral tip of mesepimeron to metapleural callosity (Fig. 6c) ; Peru N. procerodonis, n. sp. — Scent gland auricle not elongate, distance from ventral tip of mes- epimeron to apex of auricle about half or less distance from ventral tip of mesepimeron to metapleural callosity (Figs. 6a, b, d) 4 •1(3) Second antennal segment shorter than interocular distance, eyes small and sessile, vertex strongly convex betwxxn eyes (Figs. 7b, e) ; California N. arphnoides — Second antennal segment longer than interocular distance, eyes mod- erately large and substylate, head flat or slightly convex between eyes (Figs. 7c, f) 5 5(4)''Third labial segment usually short- er than anteocellar head length; California, Arizona, and Florida south to Argentina, Caribbean Is- lands, some Pacific islands N. jamaicensis ■' Debatable species distinction — see text. — Third labial segment usually great- er than anteocellar head length; South America N. rubricatus 6(1) Evaporative area reduced, extend- ing posterior of auricle a distance less than width of auricle, covering only small part of mesepimeron (Fig. 6b) ; Galapagos .... N. usingerl — Evaporative area extending pos- terior of auricle by a distance great- er than width of auricle, covering mesepimeron (Fig. 6a) 7 7(6) Scutellum swollen, in lateral view, slightly higher than pronotum (Fig. 6e) ; Argentina, Brazil, Chile N. gibbijer — Scutellum not swollen, in lateral view, level with pronotum (Fig. r.f) 8 (S(7) Viewed with vertex of head level, ocelli on or before imaginary line drawn through posterior margins of eyes, head margin behind eye nearly perpendicular to imaginary median longitudinal line of head; eyes large and substylate (Fig. 7h) ; western United States .. N. nevadensis — Viewed with vertex level, ocelli posterior to imaginary line drawn through posterior margins of eyes, head margin behind eye oblic]ue to imaginary median longitudinal line of head; eyes smaller and only slightly substylate (Fig. 7g) ; south- ern Canada south to Guatemala .... N . scolopax Neortholomus arphnoides (Baker) Ortholomns arphnoides Baker, 1906, 140; Slater, 1964, 335; Ashlock, 1967, 6, 15, 34. Nysiiis arphnoides; Banks, 1910, 62. The acute head, strongly convex vertex and relatively small, sessile eyes easily dis- tinguish this small, somewhat elongate Californian species from all other species 204 The University of Kansas Science Bulletin of Neortholomus. The hemelytra are well developed, the membrane clearly exceed- ing the apex of the abdomen. The anten- nae are short and stout with the scape distinctly not exceeding the tylus. Speci- mens of A', jamaicensis from California may be confused with specimens of N. urphnoides but can readily be separated when the characters of the head and an- tennae are carefully considered. Structure: Head strongly convex be- tween eyes, obscurely punctate; clothed densely with short appressed hairs about twice as long as an ocellar diameter; eyes small and sessile, in dorsal view slightly less than hemispherical, highest point of eye level with head at base of eye and dis- tinctly lower than vertex; head length 0.85, width 0.85, anteocular length 0.44, interocular space 0.54, interocellar space 0.34, eye length 0.22, eye width 0.17, eye height 0.27. Ikiccula elevated anteriorly, gradually narrowing posteriorly to mid- eye level without abrupt change in width, extending anterior of jugum a distance equaling one-third length of scape; labium just attaining third visible abdominal seg- ment, first segment nearly reaching thorax, second segment ot labium reaching just be- yond fore coxae, third segment attaining point between hind coxae, segment lengths from base 0.58, 0.58, 0.61, 0.43. Antenna with sparse, fine erect hairs, shorter and more dense distally, scape not exceeding tylus, shorter by distance equaling scape diameter, segment lengths from base, 0.26, 0.37, 0.37, 0.44. Pronotum with pubescence as on head, erect hairs on front lobe longer by about one-hall and more erect, [nuictation mod- erate and irregular, distance between [uinc- tures 0.25 to 1.0 X diameter of a [)uncture; pronotum evenly rounded and slightly de- pressed at callosities, posterior lobe less convex, sides slightly swollen at callosities, remaining margin straight to posterior an- gle; length 0.71, width 1.16. Scutellum with vestiture as on head, and punctation as on pronotum; Y-shaped carina evident, up[-)er arms moderately swollen, stem of "Y" very low and outlined by punctures, length 0.54, width 0.70. Metapleuron slightly swollen at callosity; distance be- tween highest point of scent gland auricle to upper edge of evaporative area directly above about one-half distance from auricle to callosity directly above, evaporative area f)n mese[-)imeron, extending dorsally along intersegmental suture nearly to dorsal edge oi pleuron, evaporative area with 20-dis- tinct p/unctures each with one thin hair. Hemelytron exceeding abdomen; co- riuni moderately clothed with short, curved, appressed hairs, basal one-third of costal margin with erect laterally project- ing hairs, longest anteriorly, decreasing steadily in length posteriorly, length of claval commissure 0.46, length of corium 1.5S. Membrane with veins indistinct; basal length to level of corial apex 0.75, apical length from corial apex 1.02. Abdomen with moderate vestiture of sh(jrt ap[-)ressed hairs and sparse vestiture of shcjrt erect hairs; connexivum narrowlv ex[X)sed lateral to corium, spiracles ob- scure. C'olor: Head with brownish-yellow broad lons^itudinal band from vertex be- tween ocelli to tylus and laterally onto jugum; buccula, distal and ventral anten- niferous tubercle, and ring about base of eye brownish-yellow, remainder of head blackish-brown. Antenna pale basally with distal segments darker, fourth seg- ment brown. Pronotum generally yellow, [umctures and scattered areas slightly darker, cal- losities blackish-brown with surrounding area dark, wide area laterally and narrow anterior margin light; yellow area of pos- terior lobe contiguous with triangular area in posterodorsal corner of [iropleuron. Acetabulum and posterior propleural area pale yellow to yellow, anterior prosternum Neortholomus, A Ni:w Genus of Orsillini 205 vcllow to brown, remainder of projileuron and prostcrnuin dark brown to black. Meso- and nK'ta[-)lcurc)n similarly dark; acetabula, scent inland auricle, and narrow anterior [lortion of eva[iorative area aloni; interscynit iilal sulure pale. Scutcllum dark brown, slightly lighter posteriorly on stem of \ -sha[ied carina. C'orium and cla\us light brownish-yel- low, clawil commissure and irregular areas along veins slightly darker, corial apex red to pale reddish-brown; membrane slightlv milkv, nearly hvalinc, without dark markings. Abdomen ventrally with basal lour seg- ments and dorsolateral area of sixth seg- ment blackish-brown, posterior segments \ellowish-brown. Legs generally yellowish-brown, coxae slightlv darker basallv; femur with irregu- lar, slightly contrasting spots, tarsi slightly darker distally. Size: Male, length 3.6-4.2, pronotum width 1.0-1.2; female, length 4.0-4.9, pro- ncjtum width 1.1-1.4. Types: 1 have seen the types of Ortho- lonuis arphnoidcs Baker through the kind- ness of Dr. David A. Guthrie of the Claremont College, C^aremont, California. One of these specimens has been labeled as the holotype although Baker failed to designate a holcjtype and his specimens therefore represent a syntypic series. The Icctotype here selected is a female in Ikiker's collection which bears the follow- ing three labels, the uppermost first: "Mts. near/Claremont/Cal. Baker," "3213" and on a red label '^Ortholumus/arphnoides/ I>aker." The specimen is point-moimted and is now preserved at the C>alifornia Academy of Science, San Francisco, (Cali- fornia. Specimens with Baker's original labels, "Mts, near Claremont Cal. Baker" or "Claremont Cal. Baker" arc considered paralectotypes for these specimens were probably before Baker when he described the species. There arc three such speci- mens (1 male, 2 females) in the Baker ('ollcction with the lectotype, others arc in the Snow F.ntomological Museum, Univer- sity ot Kansas (1 male, 1 female), the Caii- lornia Insect Survey at the University of California ui Berkeley (1 male) and the Los Angeles C'ounty Natural Blistory Mu- seum (1 male, 1 female). Variation: This s[-)ecies usually has a dark overall a[')pearance. Fret]uently the head, anterior lobe of the pronotum, and venter are nearly totallv black but occa- sional individuals are [xiler, having brown in these same areas. Most specimens are intermediate in color, being similar to the lectotype described above. The beak length varies somewhat. Shorter beaks only reach the third seg- ment of the abdomen while the Icjngcst attain the htth abdominal segment. Specimens examined: 148 males and 191 females. Distribution: All known specimens of this species were collected in California. Specimens have been examined from the following counties: Alameda, Contra Costa, Inyo, Kern, Kings, Los Angeles, Modoc, Monterey, Orange, Riverside, San Benito, San Bernardino, San Diego, San Luis Obispo, Santa Barbara, Santa Clara, Santa (auz, Sonoma, Tulare, and Ventura. Several of the more peripheral records, from Modoc (Buck Creek just east of Wil- low Ranch), Inyo (Big Pine), and San Diego (jacumbo) counties, indicate that the range of N. arphnoidcs may extend outside of C'alitcjrnia intcj Oregon, Nevada, and northwestern Mexico. Food Plants: Of the dO specimens of N. arphnoidcs with plant association data, 35 in four series were labelled Salvia incl- lifcra, two in a single series were labelled Salvia spathacca, two in another series were labelled Salvia and 17 in another series were labelled "sage," the common name for Salvia. Single specimens were 206 The University of Kansas Science Bulleti N reported on Rosa woodsii var. gratissima (Rosacea), Atriplex semibaccata (Cheno- podiacea), Baccharis (Compositae), and Eriogoninm (Polygonaceae). Van Duzee (1914) reported this species as being "found commonly on the flower heads of the black sage (Salvia mellijera) during May and June." Torre-Bueno (1946) also recorded black sage as the food plant and Rlatchley (1934) found N. arphnoides the "most common winter bug about Los Angeles, occurring in numbers on dead weeds . . ." It appears that Salvia mellijera, black sage, is the most common food plant. Several specimens, however, have been collected outside the range of S. mellijera — Coast Range from Contra Costa and western Stanislaus counties in northern Lower California (Munz and Keck, 1959) — especially those specimens from Tulare, Tulare County and Buck Creek, Modoc County. In these peripheral localities, other species of Salvia are likely to be the primary food plant. Like mcxst seed-feed- ing lygaeids, N . arphnoides is undoubtedly opportunistic, feeding on the seeds of other plants wjicn the seeds of the [irnnary food [ilant are not available. In the laboratory this species has been reared on sunflower seeds and water (Ashlock, 1967). Neortholomits gibbijer (pjcrg) Nysiiis gibbijer Berg, 1892, 155; Slater, 1964, 280. Ortholomiis gibbijer; Ashlock, 1967, 34. Ortholomus gibbits (sic) ; Ashlock, 1972, 93. The greatly swollen scutellum distin- guishes this species from others of Neor- tholomits. The strongly declivate head and pronotum, moderate sized substylate eyes, flattened vertex, broadly exposed con- nexiva and longitudinal pale line on the head, pronotum and scutclhun are uselul in its identification. Structure: Head nearly flat between eyes, slightly convex between ocelli, ob- scurely punctate, clothed densely with shcjrt silver hairs and very sparsely with long erect hairs about one and one-half times diameter of an ocellus; eye large, hemis[-)hcrical, appearing slightly stalked above and apj^ressed to head below, high- est point of eye higher than head at base (jf eye, just below level of vertex in lateral view; head length 1.11, width 1.21, ante- f)cular length 0.44, interocular space 0.77, interocellar space 0.24, eye length 0.34, eye width 0.22, eye height 0.31. Buccula ele- vated anteriorly gradually narrowing pos- teriorly to mid-eye level and ending with abrupt curve to head, extending beyond anterior of jugum, only slightly, the dis- tance less than apical diameter of scape; labium short (reaching to point between middle and hind coxae), first segment not attaining thorax, second segment attaining point between fore coxae (third segment just reaching anterior edge of middle coxae) ; segment lengths from base 0.54, 0.56 (0.53, 0.36); (antenna with fine erect pubescence as on head), sparse on basal segments (dense on fourth segment), scape just exceeding tylus; segment lengths from base 0.31, 0.70 (0.65, 0.63). Pronotum with pubescence as on head, punctation moderate and irregular, dis- tance between punctures from 0.5 to 1.5 X a diameter of a puncture; strongly de- pressed across callosities, anterior and pos- terior lobes convex transversely, posterior lobe strongly convex longitudinally, lateral margin moderately swollen laterally at cal- losities, remaining margin straight to pos- terior angle; length 1.07, width 1.77. Scu- tellum vestiture as on head and prothorax, [Hinctation dense on posterior one-fourth, distance between punctures 0.5 X or less a puncture diameter, Y-shaped carina [Tfomincnt, ujiper arms greatly swollen covering anterior three-fourths of scutel- lum and projecting dorsally higher than Ncortholomus, A Ni;\v Genus of Orsillini 207 [Xjstcrior lohc of pronotum, stem of "Y" short aiul low, outlined by punctures, leni;tli D.SO. width 1.00. Mctapleuron swollen at callosit\'; distance between high- est point ot auricle and dorsal margin of eva[iorative area directly above distinctly less than one-hall distance trom .luricle to callosity directly above, evajiorative area on mesepimeron and extending dorsally along intersegmental suture to level of meta[ileural callosity; evaporative area with about 20 distinct pimctures each with a thin hair. Hemelytron exceeding abdomen; co- rium and clavus clothed densely with ap- pressed hairs and sparsely with erect hairs, shorter than on scutellum, erect laterally directed hairs on basal one-third of costal margin becoming progressively shorter pos- teriori}; length of claval commissure 0.53, length (jf corium 2.41; membrane with vems distinct, basal length to level of corial a[K\\ 1.00, apical length from corial apex 1.19. Abdomen with moderately dense vesti- ture of short appressed hairs interspersed with occasional erect hairs; connexivum broadly exposed lateral to corium, spiracles obscure. Color: Head ground color light brown, dark around ocelli and along carina to mid-eye level, pale median longitudinal band on posterior one-half of vertex, pale around base ot eye and on tylus; dark reddish-brown on lorum anterior to an- tenniferous tubercle, laterally on tubercle, at base of head posterior to buccula; pale yellcnv on gena below eye and on buccula, antenna yellow, apical one-half pedicel brown. Pronotum ground color yellow to light brown with pale median longitudinal band mediosteriorlv; moderately dense vestiture with very short pale erect hairs and occa- sionally with anteriorly directed erect hairs; eyes large and hemispherical, ap- pearing slightly St. liked above and ap- pressed to head below, to[i of eye slightly highei- than Jiead at base of eye and at vertex; head length 0.'-*'', width 1.05, an- teocular length 0.4'*. interocular space 0.63, intenjcellar space 0.31, eye length 0.31, eye width 0.20, eye height 0.31. Buccula ele- \ated anteriorly, gradually narr(n\'ing pos- teriorly tcj mid eye level without abrupt NcortholoniHs, A Ni;w Genus of Orsillini 209 chani^'c in witlth, extending beyond an- terior ot JLi^uni ior distance equaling about one-tbird lengtb ol sca[ie; hibium reaebing second abdominal segment, first segment not attaining tborax, second seg- ment reaebing posterior edge of lore coxae, third segment attaining posterior edge of middle coxae, segment lengtbs from base 0.63, n.61, 0.61, 0.40. Antennae witb erect distally directed short hairs, hairs sparse on scape and pedicel, slightly more dense on third segment, dense on fourth, scape nearly attaining ti[i of tylus; segment lengths from base 0.31, 0.71, 0.68 (0.70). Pronotum with pubescence as on head, moderately punctate, distance between punctures 0.5 to l.OX diameter of a punc- ture, punctation more dense anteriorly especially along mid line; surface nearly Bat and slightly depressed at callosities, posterior lobe convex; lateral margins dis- tinctly swollen at callosities, slightly con- cave between callosity and [posterior angle; length 1.02, width 0.92. Scutellum vesti- ture as on [injnotum and head, punctation nicjre dense, distance between punctures 0.5 X or less the diameter of a puncture, each puncture also appearing slightly larger; Y-shaped carina distinct, arms moderately swollen, stem very low, out- lined by punctation; length 0.70, width 0.92. Metapleuron swollen at callosity, dis- tance between highest point of auricle to upjier edge of evaporative area directly above approximately two-thirds the dis- tance from auricle to callosity directly above, evaporative area on mesL-pimeron extending dorsally along intersegmental suture to level (jf callosity, evaporative area having about 25 distinct punctures, each with a single thin hair. Hemelytron exceeding abdomen; co- rium and clavus clothed moderately with short, curved, api^ressed hairs, erect hairs lacking, costal margin with laterally [pro- jecting curved hairs cjn basal one-ball be- coming progressively shorter posteriorly; length ol claval commissure 0.54, length of corium 2.43. Membrane with veins in- distinct; basal length to level of corial apex 0.95, a[")ical length Irom corial apex 1.24. Abdomen with dense vestiture of very short, appressed hairs and sparse vestiture of erect, slightly longer, posteriorly di- rected hairs; connexivum narrowly ex- posed laterally, spiracles obscure. C>olor: I lead ground color pale brown, darker brown area around cjcellus extend- ing forward along carina to mid-eye level and outlining base of tylus, narrow mar- gin about base of eye and median stripe on p,osterior one-half of vertex pale; buc- cula pale yellow, dark brown on venter of head surrounding buccula from lorum to near base of head, pale to thorax, lateral area of antenniferous tubercle from tip to eye brown, pale area below eye to dark venter. Antenna brown, slightly darker near joints, fourth segment darker. Proncjtum ground color pale brown, callosities blackish-brown, posterior lobe with indistinct checkerboard pattern formed by irregular, slightly dark areas, posterior angles dark; [iropleuron with callosity dark brown, circular area between callosity and base of acetabulum yellowish- brown, remainder pale yellow; prosternum dark brown between acetabulae. Meso- pleuron pale with small dark spot on up- per one-third and large spot on lower one- third, dark brown ventrally; metapleuron brown laterally with callosity and small area in u[ij-)er evaporative area darker, auri- cle slightly lighter than evaporative area; second and third acetabula and posterior metapleural llange pale, nearly white. Scu- tellum generally brown to dark brown, [lale })osteriorly and along posterior one- third of sides. ('orium and clavus generally very pale, brown on some veins, especially posteriorly, irregular brown spots on claval commis- sure and (Ml clavus, apex with irregular markings of red and reddish-brown; mem- 210 The University of Kansas Science Bulletin brane very slightly milky with faint, ir- regular infuscations near center. Abdomen brindled with brown, ventral one-third slightly darker, no distinct pat- tern. Legs pale yellow overall, femora with brown spots, some fused, trochanter pale, coxae pale mesally and distally, darker lat- erally, second and third tarsomeres darker. Size: Male, length 3.9-4.8, pronotum width 1.1-1.3; female, length 3.8-5.8, pro- notum width 1.1-1.7. Type: Mr. W. R. Dolling of the British Museum (Natural History), Lon- don, lent me the type of Nysiits jamaicen- sis Dallas for study. It is a single female with a small round white label with an orange border and the word "Typus," an- other small round white label with hand- written numerals about the edge reading in the clockwise direction "40, 3, 324, 4," and an elongate rectangular white label with "15. Nysius Jamaicensis," printed on it. Types of the synonymous names are discussed in a separate section below. Variation: Considerable structural vari- ation in N. jamaicensis makes the recogni- tion of species limits difficult. Substantial color variation occurs in the species as well, but none stands as a distinct cluster, instead the variants grade from one to the next. Four specimens from Clipperton Island off the Pacific Coast of Mexico have a dark brown pattern on a pale, almost cream, ground color. In many other speci- mens the same mottled pattern occurs but on a light brown background. The other extreme in color of this species is the evenly colored, light-brown individuals which are similar to the type of Nysius spiircus Stal. Overall, no pattern of dis- tribution in these color patterns was noted. A fascia similar to that lound on most N. nibricatiis is also occasionally lound in N. jamaicensis. It varies from obvious to barely discernible. The antenna is somewhat shorter in the northwestern part of its range (west- ern and southwestern U.S.), being slightly shcjrter than those described on the type but not as short as those of N. arphnoides. This character, like the others, is not clear- cut, and all intermediate forms can be found. Although the color and antennal char- acters are quite variable, they are only slightly more so than in N. scolopax and do not seem beyond the limits of species variation. But the variability of the beak in the N. jamaicensis species complex ap- [lears to be beyond these limits. In the northern part of the range, the United States, the Caribbean, and Mexico, little problem is found, the beak length agree- ing relatively closely with the type of N. jamaicensis. In southern Mexico and Cen- tral America longer beaks are more com- mon. In regions north of Panama this increase in beak length is not great, but from Panama south, specimens with beaks similar in length to those on the types of Nysius rnbricatus Berg are more common. In N. nihricatus, the length of the third labial segment is greater than the length of the head before the ocelli, whereas in N . jamaicensis the third labial segment is shorter than the anteocellar head length. In the discussion of N . rnbricatus I have compared the types of the species to N. jamaicensis and a description is included below. Specimens examined: 1255 males and 1103 females. Distribution: Coast Range counties of California (Humboldt County and south), southern Arizona (Santa Cruz County), southern Texas (Cameron, Hildago, Starr, and Willacy counties) and southern Flor- ula (Dade and Monroe counties), south through the Caribbean islands, Mexico, and Central American to Peru, Bolivia, Paraguay, northeastern Argentina (Cor- rientes and Formosa provinces), and south- Neortholomus, A New Genus of Orsillini 211 crn ilrazil (Santa Catarina, state). Not know II from Chile and Uruguay. Records known Ironi Clipperton, Clarion, Socorro, and Tahiti islands in the Pacific. Speci- mens arc known trom coastal regions up to 10,000 feet in one example from Mex- ico. Food Phints: I have seen specimens with the following pkint data (number of collections : number of specimens) : Cleome spinosa (Capparidaceae) (1 : 1), Baccharis sp. (Compositae) (1 : 3), Eiipa- torium adenophontm (Compositae) (2 : 14), Cncnrbita moschata (Cucurbitaceae) (1 : 1), Ocinutm saiive (Labiatae) (1 : 1), Stachys calijonucits (Labiatae) (1 : 5), Salvia splendens (1 : 3) and S. xalapensis (1:1) (Labiatae), Ribes hesperia (Saxi- fragaceae) (1 : 2), Solanum tuberosum (= famesii, Solanaceae) (1 : 4), Lippia sp. (Verbenaceae) (1 : 1), Verbena pro- strata (Verbenaceae) (1 : 1), and Vitex pyramidata (Verbenaceae) (1 : 1). One collector included on a note with the speci- men, "On a species of Hyptis (Labiatae) growing in damp and somewhat shady place. Others were seen on same plant but escaped, they conceal themselves on the flower head." The habit of hiding on the flower head has been observed in N . scolo- pax and may be characteristic of all Neortholomus. Synonymized species: Nysius spurcus Stal has presented some minor problems primarily concerning distribution. Usinger (1941) synonymized N. spurcus with Or- tholomus jamaicensis (Dallas). I have seen Stal's type specimens from the Swed- ish Museum of Natural History through the courtesy of Dr. Persson. I have com- pared these specimens with the type of the Dallas species and agree with Usinger. In the type series sent to me by Dr. Persson were two females from "Taiti" and a male from Honduras. Stal (1859) in the original description, also included "Rio Janeiro" as a locality. Dr. Persson has been unable to locate such a specimen or specimens. He has suggested that Stal, after publishing the original description of N . spurcus, may have removed the "Rio Janeiro" specimen from the type series thinking it was not conspecific. In the Enumeratio Hemipterorum IV (1874), Stal mentions only "Insula Taiti" as the locality for Nysius spurcus. No Icctotype has been selected for Nysius spurcus Stal. Although the two female specimens are marked as "Typus" and "Paratypus," I do not believe these labels were placed by Stal. As lectotype of Nysius spurcus, I have chosen the female specimen with the labels "Taiti," "Kinb.," "spurcus Stal" in Stal's handwriting, "Typus" on a red label, "50 75" on a red label and "Riksmuseum Stockholm" on a light green label. Included as paralecto- type are a female specimen with the labels "Taiti," "Kinb.," "spurcus" (a recent la- bel), "Paratypus" on a red label, "51 78" on a red label, "Riksmuseum Stockholm" on a light green label; and a male speci- men labeled "Honduras" and "Hjalmar- son" both in Stal's handwriting, "52 78" on a red label and "Riksmuseum Stock- holm" on a light green label. Although the male specimen was not marked as a type, it is in the type series and Stal men- tioned a male specimen in the original description. Stal stated that he had sent a specimen from Honduras to "Dom Hjal- marson." Judging from these facts, I be- lieve this male belongs in the type series. The major difference between the types of spurcus and jamaicensis is the paleness of the Stal types compared to that of Dal- las. This pale coloration is not unusual in Neortholomus jamaicensis and I consider this individual variation. The major problem with N. spurcus is the question of the accuracy of the "Taiti" locality data on the two female specimens. Tahiti is an unlikely locality for an otherwise New World species. The 212 The University of Kansas Science Bulletin only other Pacific islands on which this species has been found are Clarion, Clip- perton, and Socorro islands, all within 800 miles of the west coast of Mexico. Usinger (1941) wrote, "Locality records for the Eugenies Resa expedition have proved to be notoriously inaccurate and several competent collectors . . . have failed to turn up any Orsillini in the So- ciety Islands." For reasons (jf this sort, Stal's record from "Taiti" has been thought erroneous and the specimens considered mislabeled. In material from the United States National Museum of Natural His- tory, however, I have seen a single female specimen of Neortholomus jamaicensis from near the Fantana River in Tahiti collected in 1961 by J. F. G. Clarke. I have compared this specimen with the types of N. jamaicensis and N. spitrcus. The Tahiti specimen has a slightly paler head and slightly darker pronotum than the N . jamaicensis type and is generally darker than the N. spitrcus type. No other differences could be found. The color dif- ferences are easily within the limits of species variation. It is possible that the species was intro- duced to the islands by early traders who sailed between the New World and the South Pacific islands. Natural introduc- tion to the islands seems unlikely over these distances. There is no evidence of populations between the New World and Tahiti on islands that might have acted as "stepping stones." I have seen the holotype of Nysiiis providus Uhler at the U.S. National Mu- seum of Natural History. This specimen is clearly a Neortholomus jamaicensis. This species was synonymizcd by Distant (1901). Although I have not seen the entire type series, it is likely that it in- cludes specimens of both A/, scolopax and N. jamaicensis judging from the distribu- tion given by Uhlcr in the original descrip- tion. His statement of distribution for providus was "North American from Que- bec to Arizona, from thence it spreads into Mexico and Central America, and follow- ing south it is found on the Isthmus of Panama, and in Colombia and northern Brazil. In the West Indies it occurs in Trinidad, Grenada, St. Vincent, Porto Rico, San Domingo, and Cuba, and from thence it extends through Florida into all of the eastern states as far as Maine" (Uhler, 1894). Neither of the above-men- tioned species has such an extensive dis- tribution but the combination of the two would. Neortholomus rubricatus (Berg) Nysius rubricatus Berg, 1879, 102-103. Belonochilus rubricatus; Kiritshenko, 1931, 16; Slater, 1964,236. Ortholomus rubricatus; Ashlock, 1967, 34. Neortholomus rubricatus is most simi- lar to A*', jamaicensis and N . \oreshanus. The lenirth of the third labial segments is greater than the preocellar head length in N. rubricatus whereas the preocellar head length is greater in N . jamaicensis. Neor- tholomus rubricatus usually has a pale fascia along the midline of the head, pro- notum, and scLitellum. Neortholomus \oreshanus has a much longer head and labium than does N. rubricatus. For a more detailed discus- sion of the diagnostic characters see the section on variation for A^. jamaicensis above and for N . rubricatus and N . \ore- shanus below. Structure: Head slightly convex be- tween eyes, obscurely punctate; (moder- ately dense vestiture of very short, pale, ajipressed hairs and very few erect, an- teriorlv directed hairs); eyes moderate sized, less than hemispherical, appearing slightly stalked above and appressed to head below, top of eye level with head at base of eye and lower than vertex; head length 1.02, width 1.00, anteocular length 0.55, interocular space 0.62, interocellar Neortholomus, A Ni:\v Genus of Orsillini 213 space 0.30, eye Icni^th 0.30, eye width 0.21, eye height 0.30. Ikiccula elevated an- teriorly, gradually narrowing posteriorly to mid eve level without abrupt change in width, extending beyond anterior of ju- gum for distance equaling about one- third length ol scape; labitnn nearly at- taining third abdominal segment, first segment not attaining thorax, second seg- ment reaching posterior edge of fore coxae, third segment attaining mid level of hind coxae, segment lengths from base 0.f)4, 0.75, 0.90, 0.55. Antenna with scat- tered distally directed hairs on scape, (hairs sparse on pedicel, slightly more dense on third segment, dense on fourth), scape not attaining tip of tylus; seginent lengths from base 0.26 (0.73, 0.70, 0.70). I^ronotum (with pubescence as on head), moderately punctate, distance be- iwccn punctures 0.5 to l.OX diameter of a puncture, impunctate along mid line, punctation more dense along callosities and along anterior portion of fascia; de- pressed across callosities, posterior lobe ccjnvex; lateral margins slightly swollen at callosities and straight between callosity and posterior angle; length 1.02, width 1.56. Scutellum (vestiture as on pronotum and head), punctation more dense, dis- tance between punctures 0.5 X or less the diameter of a puncture, each puncture also appearing slightly larger; Y-shaped carina distinct, arms moderately swollen, stem very low, outlined by punctation; length 0.76, width 0.91. Metapleuron swollen at callosity, distance between highest point of auricle to upper edge of evaporative area directly above approximately two- thirds the distance from auricle to callosity directly above, evoporative area on mes- epimeron extending dorsally along inter- segmental suture to level of callosity, evaporative area having about 26 distinct punctures, each with a single thin hair. Hemelytron exceeding abdomen; co- rium and clavus clothed moderately with short, curved, ajipressed hairs, erect hairs lacking (costal margin with laterally pro- jecting curved hairs on basal one-half be- coming [progressively shorter posteriorly); length of claval commissure 0.46, length of c(jriLun 2.32. Membrane with veins in- distinct; basal length to level of corial apex 1.17, apical length Irom corial apex 0.94. Abdomen with dense vestiture of very short, appressed hairs (and sparse vesti- ture of erect, slightly longer, posteriorly directed hairs); connexivum narrowly ex- posed laterallv, s|-»iracles obscure. C'olor: Head ground color tawny to browiush-ycllow, slightly darker around ocellus, slightly paler along mid line, near base of head and in margin around base of eye; buccula pale yellow, slightly darker on venter of head surrounding buccula from IcMum to base of head and on lateral area of antenniferous tubercle from tip to eye, \x\\c area below eye to darker venter. Scape pale brown apically, [laler near base (remainder of antenna pale brown, darkest near joints). Pronotum ground color brownish-yel- low, callosities and punctures slightly darker causing impunctate median fascia to appear as a pale line; propleuron ground color and puncture color similar, pale reddish-brown area at base of acetabulum; presternum slightly darker than ground color between and before acetabula an- tericjr margin slightly paler. Mesopleuron pale yellowish-brown with posterior mar- gin and acetabula more pale, darker brown ventrally; meta[ileuron pale yellowish- brown dorsally increasingly pale ventrally, very pale on dorsal margin, metapleural flange and acetabula. Scutellum pale yel- lowish-brown, more pale along mid line and along posterior one-third of sides. Corium and clavus pale brownish-yel- low, ajiex with irregular reddish-yellow markings; membrane hyaline. Abdomen ground color pale brownish- yellow, slightly darker dorsally and an- 214 The University of Kansas Science Bulletin teriorly, few scattered reddish markings, ovipositor, in repose, very dark brown. Legs pale yellow overall, femora with pale brown spots, some fused, apically tarsomeres slightly darker. Size: Male, length 3.6-4.5, pronotum width 1.0-1.3; female, length 4.0-5.2; pro- notum width 1.2-1.5. Types: I have studied three specimens that are syntypes of Nysius riibricatus. All three have green labels with the name "Misiones" printed on them. The two fe- male specimens have small rectangular labels bearing the word "Typus." The other specimen, a male, does not have a label indicating it is a type. Because Berg indicated in his original description that he had seen a male specimen and since the locality label on this specimen is iden- tical to those on the female specimens, I assume the male is a syntype of Nysius rtibricatus. I have chosen the smaller of the two females as the lectotype. The specimen is mounted on a new point and the pin bears the following labels: a small white label with " 9 ," a small white label with a red border and the word "Typus" in red, and the green locality label with "Misio-/nes:" printed in black. The fe- male paralectotype is also on a new point and the specimen bears the following la- bels: an identical green locality label, an identical "Typus" label, a small white la- bel with "150" on the underside, a large white label with "Nysius rubricatus Berg" in Berg's handwriting and a smaller white label with "C.J. Drake/Coll. 1956" printed on it. The male paralectotype is mounted on an old, yellowed point, below this is a green locality label identical to those on the other specimens, and a white label with "C.J. Drake/Coll. 1956" as on the female paralectotype. These specimens are de- posited in the Drake Collection at the U.S. National Museum of Natural History in Washington, D.C. The lectotype is the most distinct indi- vidual in the type series and was selected because it is most likely to represent a species distinct from ISI . jamaicensis (Dal- las). The Nysius rubricatus type differs from the Dallas type in being much paler, almost yellow, having an even paler im- punctate fascia originating on the scutel- lum, crossing the pronotum, and ending on the posterior vertex, and having an elongate head and labium, the labium clearly exceeding the hind coxae. The third labial segment of the lectotype clearly longer than the head length before the ocelli. The third segment of the beak of N . jamaicensis is shorter than the preocel- lar head length. The paralectotypes are not so distinct in this character, with the third labial segment only slightly longer than the preocellar head length. The problem lies in specimens which appear either to be ambiguous for crucial characters or to have intermediate charac- ters. Some specimens of N. rubricatus have the dorsal fascia and short beak while other specimens have an indistinct fascia or a beak with the third segment equal in length to the head before the ocelli. There appears to be no correlation between the color, the fascia, and the beak length. The genitalia of these species seem identical, but the types represent such distinct condi- tions that I feel it is counterproductive to synonymize N. rubricatus at this time. When more information concerning the biology or cytology of this species is avail- able, this problem may be solved. Variation: As indicated above, none of the characters are as stable as one would wish. Most individuals have a mottled pattern, dark brown on a pale background, although other specimens are like the type, having an even light brown to light red- dish-brown coloration. The very pale lon- gitudinal fascia is usually evident on the pronotum and scutellum though occasion- Neortholomus, A Ni:w Genus of Orsillini 215 ally the fascia is not present or, at most, ill defined. The size is cjuite variable although not unusually so for species of Neortholomus. The third segment of the labium may be only slightly longer than the anteocellar head length or it may be very clearly longer although not as long as in N. kpreshauiii. Specimens examined: 41 males and 29 females. Distribution: From Panama north of the Canal Zone south to central Peru (Huanuco, department), central Bolivia (Santa Cruz, department), and northern Argentina (Formosa and Misiones prov- inces) with scattered records from through- out Brazil (Amapa, Golias, Mato Grosso, Minas Gerais, Pernambuco, and Rio Grande do Sul, states). No records from Chile, Ecuador, French Guiana, Guyana, Uruguay, and Venezuela. Specimens have been found from coastal locations to ele- vations of approximately 1600 meters. Food Plants: No data available. Neortholonnts \oreshanus (Van Duzee) Belonochilus koreshaniis Van Duzee, 1909, 165-166. Ortholonms koreshaniis; Barber, 1947, 62; Slater, 1964, 337; Ashlock, 1967, 29, 34; Alayo, 1973, 22. Neortholomus koreshaniis is distin- guished from other Neortholomus by the long head with small eyes, the very long beak which exceeds the mid level of the abdomen, and the pale, longitudinal fascia on the scutellum, pronotum, and posterior part of the vertex. The antenna is long but the scape does not exceed the very elongate tylus. Neortholomus koreshaniis specimens with slightly shorter beaks and correspond- ingly short heads look much like N. rubri- catus, especially those N. rubricatiis with longer than average beaks. Despite the similarity, the beak of N. \oreshanus is always longer and the eyes are slightly smaller. There is no distributional overlap (jf these species. Structure: Head elongate, especially tylus, vertex slightly convex between eyes and more strongly between ocelli, ob- scurely punctate; clothed densely with short appressed hairs and sparsely with short erect hairs about as long as an ocellar diameter; eyes moderate in size and ap- pearing slightly stalked from head, highest point of eye slightly higher than head at base of eye, vertex distinctly higher; head length 1.21, width 0.99, anteocular length 0.70, interocular space 0.60, interocellar space 0.29, eye length 0.29, eye width 0.19, eye height 0.32. Buccula elevated anteri- orly and gradually narrowing posteriorly to level of posterior eye margin without abrupt change in height, extending beyond anterior of jugum by one-half the length of scape; labium long, extending posteri- orly to mid level of sixth abdominal seg- ment, mid level of ovipositor, first segment attaining thorax, second segment of labium reaching anterior edge of middle coxae, third segment just reaching fourth ab- dominal segment, segment lengths from base 0.92, 0.92, 1.24, 0.85. Antenna with sparse vestiture of partially erect, distally directed, short hairs (more dense on fourth segment), scape not reaching tip of tylus, short by one-half its length, segment lengths from base 0.34, 0.70, 0.68 (0.66). Pronotum with pubescence as on head, punctation moderate, distance between punctures from 0.5 to l.OX diameter of a puncture, narrow median longitudinal fascia impunctate and slightly raised, pro- notum moderately depressed at callosities, lateral margins moderately swollen at cal- losities, remaining margin to posterior an- gles slightly convex, length 1.02, width 1.41. Scutellum vestiture and punctation as on pronotum, Y-shaped carina evident but very low, stem of "Y" outlined by 216 The University of Kansas Science Bulletin punctures, arms not so distinct; length 0.66, width 0.71, metapleuron sHghtly swol- len at callosity; distance from highest point of auricle to dorsal margin of evaporative area directly above greater than one-half distance from auricle to callosity directly above, evaporative area on mesepimeron and extending dorsally along interseg- mental suture to level just above meta- pleural callosity, evaporative area with about 25 distinct punctures each with a thin hair. Hemelytron exceeding abdomen; co- rium and clavus densely clothed with short appressed hairs and scattered erect hairs of same length, erect hairs projecting later- ally along basal one-third of costal margin, becoming progressively shorter posteriorly; length of claval commissure 0.53, length of corium 2.21. Membrane with veins mod- erately distinct; basal length to level of corial apex 0.92, apical length from corial apex 1.21. Abdomen densely clothed with short appressed hairs, moderate vestiture of short, erect, posteriorly directed hairs; con- nexivum narrowly exposed laterally, spira- cles obscure. Color: Head ground color yellow to yellowish-brown; dark reddish-brown about ocellus, anteriorly along lateral sides of carina to mid level of eye, and medially along carina as narrow band to anterior edge of eye; two parallel light brown lines at base of tylus, most pale about base of eye, on tylus, and between ocelli at base of head; very pale below eye from antcn- niferous tubercle to base of head; lorum brown, darker ventrally towards buccula, base of head around buccula brown, buc- cula pale yellow, dark brown spot behind eye next to thorax. Antenna yellowish- brown, more pale at joints, scape more pale basally. Pronotum generally pale yellow, each puncture appearing slightly darker, cal- losities very dark reddish-brown with sur- rounding area slightly darker than ground color, [iropleuron colored as on dorsum, shcjrt band at base of acetabulum dark reddish-brown with surrounding area slightly darker than ground color, an- terior edge of propleuron behind eye dark. Meso- and metapleuron yellow to yellow- ish-brown; acetabula, evaporative area, and pcjstericjr meta[ileural flange very pale; mesopleuron with short central longitudi- nal band, middle portion of anterior edge, and spot on dorsal one-half dark reddish- brcnsn; metaiileural callosity very dark reddish-brown. Scutellum light brown along anterior margin and becoming more yellow to pale yellow posteriorly, punc- tures slightly darker. Corium and clavus very pale, claval commissure slightly darker, corial apex red to reddish-yellow; membrane very slightly milky, nearly hyaline, slightly in- fuscated centrally. Venter of abdomen yellow to yellow- ish-brown, one brown spot dorsally on hrst three visible segments, some brown medially on hrst visible segment and on ventral one-third of first two intersegmen- tal sutures. Legs pale yellow, femora of all legs with light brown spots in no distinct pat- tern, some spots fused, near distal end spots fusing to form band, narrow basal band on tibia, last tarsomere dark brown. Size: Male, length 4.()-5.4, pronotum width 1.2-1.4; female, length 5.3-5.9, pro- notum width 1.3-1.6. lyjie: Van Duzee's type series of Belotiochdits /(orcs/uiniis consists of two female specimens individually point mounted on a single pin. The upper white label reads "Estcro, Fla My. 6-12. 08 Van Duzee," below is a white label reading "C>)-type." A red U.S.N. M. label below this reads "C:o.Type No. 12244 U.S.N.M." 'TJelonochilus Koreshanus V.D." is hand [•;rinte(l on a white label below all of the other labels. The upper specimen is here Neortholoinns, A Nuvv GtNus of Orsillini 217 designated as the lectotype of Belonochilus /(urc-s/h//jiis Van Duzee, 1%'^. The speci- mens are in the U.S. National Museum of Natural History. V' ariation : 1 he lectotype and paralecto- tvpe are pale specimens, while the ma- iority of the other specimens are much darker, esjiecially in those areas on the ty[K' which are described above as dark. The pronotum, corium, and membrane inav have dark brown markings and large portions ot the venter of the head and abdomen may be dark brcjwn to blackish- brown. Nevertheless, this species is gener- allv paler than N. scolopax, N. ncvadensis, or A', arphnoides. Beak length varies from reaching the third abdominal segment to attaining the [losterior edge of the sixth, or occasionally in small males it may reach the tip of the abdomen. Specimens examined: 41 males and 49 lemales. Distribution: Found throughout Flor- ida from Pensacola and Jacksonville south to Key Largo. Also recorded from Isles of Pines but not from Cuba (Alayo, 1973) or any other Caribbean islands. Food Plants: Only two plant associa- tions have been made for this species. Van Duzee (1909), in the original description of N. l^ureshaniis, reported finding it "abundant ... in all stages of growth" on the seed heads of "a low branching hirsute labiate [^lant locally called 'jiennyroyal.' " Needham (194S) referred to the plant as Florida Pennyroyal, Pycnothywits rigidiis. He has recorded N. koreshanus as having been reared once from the seed head of Bidens pilosa (Compositae). This appar- ently is an uncommon occurrence since it was found in only one out of 1000 flower heads he had placed in jars. P)latchley (1926) reported taking "small numbers by beating water oriutn with anterior one-half pale with some light brown, irregular spots; clavus similar with anterior margin and cla\al ccjmmissure more brown, posterior one-half of corium generally brown and dark brown with cjbscure pale markings, corial a[xx red; membrane slightly milky, irregular brown markings between veins. V^enter f)f abdomen yellowish-brown, dark brown to black markings on ventral one-third of all segments, anterior three segments nearly all black ventrally, spots scattered on more posterior segments. Femora pale yellow with rows of dis- tinct brown s[iots, coxae and tibiae simi- larlv pale, coxae brown basally, tibiae pale at joint lollowed by pale brown ring, pale central section, and a light brown distal one-third, tarsi light brown, dark distally. Size: Holotype, male, length 4.3, pro- noturn width 1.4; paratype (same locality as holoty[-)c), male, length 4.3, pronotum width 1.4; paratype (Huancabamba), male, length 4.1, pronotum width 1.4; [xiratype (Panao), female, length 4.1, pro- notum width 1.3. Holotype: Male, Peru, Dept. Ama- zonas, vicinity of Chachapoyas, Andes, altitude 2000 m, 7 August 1936, F. Woyt- kowski collector, no. 3674, brook, in the Snow Entomological Museum, University of Kansas, Lawrence. Paratypes: 1 male, same locality as holotype, 6 August 1936, no. 3673, boggy pond (genitalia dissected), in the Snow F.ntomological Museum. 1 female, Peru, Dept. Huanuco, Panao, altitude 8000 ft, 26 January 1947, J. (-. Pallister collector (geni- talia dissected), in the American Museum of Natural History, New York. 1 male, Peru, department unknown (see discus- sion below), Huancabamba, 13 August 1945, P. A. Ikrry collector. South Ameri- can Parasitology Lab 1316-8, 46-4503 (geni- t)unty, Kansas. It was collected on mature seed heads of Pyc- nanthciim tcniiijoUitm. This locality is approximately 150 miles south of Cedar Bluff, Iowa where one of Say's types was collected. The neotype is in the Snow Entomological Museum at the University of Kansas. The types of synonymous spe- cies are discussed below. In agreement with Barber (1923) and IMatchley (1926), I consider Nysiiis (Or- tholonius) longiccps Stal (1(S74) a syno- nym of Lygaeiis scolopax Say 1(S31. I have seen specimens from the Swedish Museum of Natural History, Stockholm, through the kindness of Dr. Persson, which appear to be the types of StaTs N . (O.) longiceps. There is no label that in- dicates these are the types, nor is there any indication of this status at Stockholm according to Dr. Persson. Despite this lack of absolute identification as types of N. longiccps, the circumstantial evidence is strong. This series of 17 specimens fol- low a large label with the genus Nysiiis Tablk 2. Size ranges for N. scolopax from extremes in distribution. Oaxaca, Mexico Dallas Co., Iowa Males len.i;lh 4.10-4.50 4.40-5.05 width l.lS-1.35 1.33-1.53 Females len,L;th 4.55-5.05 4.90-6.10 width 1.40-1.55 1.40-1.70 Neortholonius, A New Genus of Orsillini 225 writlcn on it in StdTs handwriting. The series contains only specimens irom all of the localities i;iven by Stal in the original descri|ition oi A. longiceps. Dr. Persson has been unable to locate any specimens in the collection at Stockholm that . Ashlock)." (Ashlock, 1972). Paratypes. 96 males, 95 females, 4 nymphs collected on Santa Cruz and San Cristobal Islands in the Galapagos Archipelago. All of the types were collected in 1964 during the Galapagos International Scientific Proj- ect. The holotype is in the California Academy of Sciences, San Francisco, and the majority of the paratypes are in Peter D. Ashlock's personal collection, Univer- sity of Kansas, Lawrence. Variation: Neortholomus tisingeri is relatively uniform in shape although the size varies considerably. The labium ends between the middle and hind coxae, the uniformity of this character being unusual in Neortholomus. The color is somewhat variable with a range from pale, uniformly colored specimens to the darker specimens with dark brown at the apices of the clavus and/or corium and on the pronotal callosities. Darker specimens also have scattered, slightly contrasting brown areas on the [ironotum. 'ihcre is also variation in the degree ot mottling on, and the whiteness of, the membrane. Specimens examined: 89 males, 94 fe- males, 4 nymphs (paratypes); 2 males, 1 female, 7 nymphs (not types). 228 The University of Kansas Science Bulletin Distribution: Galapagos Archipelago; Santa Cruz, San Cristobal, Santa Maria and Pinzon islands. Ashlock (1972) recorded specimens from Hypericum pratense (Hypericaceae), Verbena (Verbenaceae), and Cordia (Bo- raginaceae). Acknowledgments I am grateful to Dr. Peter D. Ashlock who suggested this project and who has been an unending source of help through his personal library, the loan of specimens from his extensive orsilline collection, and his patient advice. I also wish to thank Dr. George W. Byers for his suggestions and advice and Marjorie Rothschild Ham- ilton for assistance in rect^rding much of the label data from the nearly 5,000 speci- mens examined, and for proofreading of the manuscript. Thanks are also extended to Dr. Charles D. Michener of the Uni- versity of Kansas, Dr. James A. Slater of the University of Connecticut, and Dr. John C. Morse of Clemson University for their suggestions concerning the manu- script. Thanks also to Jolene Walker for the typing of this manuscript. My thanks go to the following indi- viduals and institutions who helped in my research by generously allowing me to borrow specimens: P. H. Arnaud, Jr., Cal- ifornia Academy of Sciences; H. Brailov- sky A., Instituto de Biologia, Universidad Nacional Autonoma de Mexico; P. J. Clau- sen, University of Minnesota; W. R. Dolling, British Museum (Natural His- tory), London; R. L. Fischer, Michigan State University; R. C. Froeschner, U.S. National Museum of Natural History; D. A. Guthrie, Claremont Colleges, Cali- fornia; W. J. Hanson, Utah State Univer- sity; C. L. Hogue, Los Angeles County Museum of Natural History; J. D. Lattin and 11 B. Frost, Oregon State University; T. E. Moore, University of Michigan; D. Otte, Academy of Natural Sciences of Philadelphia; L. L. Pechuman, Cornell University; I-*. I. Persson, Naturhistoriska Riksmuseet, Stockholm; J. A. Powell, University of California, Berkeley; G. H. Rosado Neto, Universidade Federal do Parana, Curitiba, Brasil; J. C. SchafTner and S. C. Merrit, Texas A & M Univer- sity; R. T. Schuh, American Museum of Natural History; R. O. Schuster, Univer- sity of California, Davis; J. A. Slater, Uni- versity of Connecticut (personal collec- tion); C. A. Triplehorn, Ohio State University; H. V. Weems and F. W. Mead, Florida State Collection of Arthro- pods, Gainesville; D. A. Young, North Carolina State University. Final!)', I would like to express my sincere appreciation to Virginia Ashlock whose critical editing of the Master's thesis, from which this paper is taken, has uiidcjubtedly added to the clarity. I, of course, am responsible ftjr any hick of clarity which remains. Literature Cited Alayo D. p. 1973. Los Hcmiptcros dc Cuba, Parte XI, Familia Lypacidae. Torreia, no. 25, 19 p. Ashlock, P. D. 1957. An investigation of the taxo- nomic value of the phallus in the Lygacidae (Hemiptcra-Heteroptera). Ann. Entomol. Soc. Amcr. 50:407-426. . 1967. A generic classification of the Orsil- linac of the world (Hemiptcra-Hctcroptcra: Ly- gacidae). Univ. California Publ. Entomol. 48. 82 p. 1972. The Lygaeidae of the (;a!a[)agos Is- lands (Meiniptcra: Iletcroptera). Proc. Califor- nia Acad. Sci. 39:87-103. Hakik. C. 1-". 1906. Notes on the Nyntis and Or- tholoniiK of America. Invert. Pac. 1:133-140. Banks, N. 191 (I. Catalogue of the Neartic Hemip- tcra-Heteroptera. Amcr. Entomol. Soc. Phila- delphia, P. O. Stockhausen. 103 p. Barbkr, H. G. 1923. Family Lygaeidae, p. 708-737. In E. B. Britton (ed.). Guide to the Insects of Connecticut, Part IV: The liemiptera or Sucking Insects of Connecticut. Bull. Connecticut Geol. Nat. Hist. Surv. ■ . 1947. The family L\'gacidac (Hcmiptera- Heteroptera) of the island of Cuba and the Isle of Pines. Mem. Soc. Cubana Hist. Nat. 19:55-75. Berg, C. 1879. Hemiptera Enumeravit Specieque Novas. Pauli E. Conni, Bonariac. 316 p. Neortholomus, A Ni:\v Clnus of Orsillini 229 . 1 S92. Nova Hcniiptera faunarum Argcnti- nac- ct Uru,i;a\cnsis, l.ycacidac. Ann. Sco. Cicnt. Ar.ucniina .•?3:1 ^ 1 -1 d^. I'.i.Arcm.i Y, W. S. l'>26. Ilctcroptera or true bups of eastern North America, with special reference to the faunas of Indiana and l^iorida. Nature Puhl. C":o., Indianapolis. 1116 ji. . 1934. Notes on a collection of F hteroptera taken in winter in the vicinity of Los Angeles, California. Trans. Amcr. Entomol. Sfic. 60:1-16. Dailky, p. J., R. C. Graves, and J. L. Herring. 1978. Survey of Heniiptera collected on coinmon milkweed, Asclepiaf syviaca, at one site in Oliio. Entomol. News 89:157-162. Dallas, \V. S. 1 852. List of the Specimens of Herniiuerous Insects in the Collection of the British Museum, Part II, p. 369-592. Taylor and Francis, Inc., London. Distant, W. L. 1"'()1. Rhynchotal notes, X: Meterop- tcra, Family Ly.uaeiilae. Ann. Mag. Nat. Hist. 7:531-541. ' Gillette, C. P., and C. F. Baker. 1895. A pre- liminary list of the Hcmiptera of Colorado. Bull. Colorado Agr. Exp. Sta. 31:1-137. Glovi.r, T. 1876. Manuscript notes from my jour- nal or illustrations of insects, native and foreign. Ortler Ilemiptera, suborder Heteroptera or plant bugs. J. C. Entwisle, Washington, D.C. 132 p. Hamilton, S. W. I'^Sll. Revision of the New World Orsillini, with special reference to Ncorthnloniiis, n. gen. (Hemiptera-Hcteroptera: Lygaeidae). Mas- ter's Thesis, Univ. Kansas, Lawrence, Kansas. 113 p. Herrich-Schaeffer, G. H. W. 1838. Die Wan- zenartigen Inseckten. C. H. Zch'schen Buchhand- lung: Nurnberg. 4:33-92. Johnson, C, and R. Ledig. 1918. Tentative list of Hemiptera from the Claremont-Laguna region. J. Entomol. Zool. 10:3-8. KiRirsnENKO, A. N. 1931. Bemerkungen uber einige Gattungen dcr Astacopinae. N. Beitr. syst. Insect. 5:16. MuNz, P. A., AND I). 1). Keck. 1959. A Californi:i Flora. Univ. California Press, Berkeley. 1681 p. Neediiam, J. D. l'M8. Ecological notes on the insect population of the flower heads of Bidc/K pilosa. Ecol. Monogr. 18:433-446. Provancher, L. 1872. Description dc plusieurs Ilcmipteres nouveaux. Nat. Canada 3:73-79. Say, T. 1831. Descriptions of new species of heterop- terous Hcmiptera of North .'\merica. Reprinted 1859 In J. L. LcConte (ed.), The Complete Writ- ings of Thomas Say on the Entomology of North America, Vol. 1, p. 310-36S. Baillicre Brothers, N.Y. Slater, J. A. l')64. A Catalogue of the Lygaeidae of the World. Univ. of Connecticut, Storrs. 2 vols., 1688 p. St.al, C. 1859. Ilemiptera species novas dcscriptsis. Konglika Svenska Fregattens Eugenies resa om- kring jorden. 111: Zoologi, Insckter 1859:219-298. ■ . 1 872. Genera lygaeidarum Europae disposuit. {)fvers. Vetensk.Xkad. Forh. Stockholm 29:37-62. . 1874. Enumeratio 1 lemipterorum, Part 4. K. Svenska VetenskAkad. Hand!. 12:1-186. ToRRi-BuENo, J. R. l'H6. A synopsis of the Hcmip- tera-Heteroptera of America north of Mexico, 111: Family XI, Lygaeidae. Entomol. Amer. 26:1-141. Ueshima, N. 1979. Hemiptera, II: Heteroptera. hi Animal Cytogenetics, 3, Insecta 6. Gebrijder Borntraeger, Berlin. 117 p. Ueshima, N., and P. D. Ashlock. 1980. Cytotax- onomy of the Lygaeidae (Hemijitera-Ileterop- tera). Univ. Kansas Sci. Bull. 51:717-801. Uhlkr, p. R. 1894. On the Hemiptera-Heleroptera of the island of (jtenada. West Indies. Proc. Zool. Soc. London 1894:167-224. Usincer, R. L. 1941. The jiresent status and synon- ymy of some orsilline species (Hemiptera, Lygaei- dae). Bull. Brooklyn Entomol. Soc. 36:129-132. Van Duzee, E. P. 1907. Notes on Jamaican Hemip- tera. Bull. Buffalo Soc. Nat. Sci. 5:1-79. . 1909. Observations on some Hemiptera taken in Florida in the spring of 1908. Bull. Buffalo Soc. Nat. Sci. 9:149-230. 1914. A preliminary list of the Hemiptera of San Diego County, California. Trans. San Diego Soc. Nat. Flist. 2:1-57. Wagner, E. 1958. Der A^yy/'/rc-Komplex (Hemiptera- Heteroptera: Lygaeidae) in der Palaearktis. Com- ment. Biol. Helsing. 19:1-54. 230 The University of Kansas Science Bulletin 00 CO CEi 09 c/r: CO ^^ o u CO 3 CO o a o ca u cs o z ^ ^J CQ sz ^ s < > o BQ o a. S CQ Cd q:: CO u tf «< S z \ C5 y CO \ < QC2 / \ \^ CspPBF, \ \ \ / . CO Z s CO W o£ U3 ,AeGM 'SpPBFj >PC - -SpPDF --SpPBF, Fig. 1. Proposed phylogeny for Neortholomiis. Character abbreviations explained in Table I. Table 1. Characters supporting the phylogeny of Neort/iolomtis (Fig. 1). Character Apomorphous Reference* Plesiomorphous Reference* abbreviation Sex Character state and Fig. # state and Fig. # SpPDF 2 Distal flange of sperm pump absent H— 3 present A— 8a, c SpPBFi 2 Basal flange of sperm pump partially reduced H— 3c, d complete A— Sa, c SpPBFo 9 Basal flange of sperm pump gready reduced, present H— 3b reduced (SpPBFx) H— 3c, d SpPBFs ? Basal flange of sperm pump completely lost H— 3a, e reduced (SpPBF,) H— 3c, d SpDDu 9 Distal duct of spcrmatheca short H — 3a, b, d, c long A— 8a, c H— 3c SpPSR 2 Sdcrotized basal ring of sperm pump long and narrow H— 3e short & stout H— 3a-d PC $ Shank of with H-4g, h without W— 15 paramere carma carma 1 1— 4a, e, f AcGM $ Gonoporal process well developed membrane H-4c, j litde membrane A— 3a, 4a H— 4b, d, i •A = Ashlock (1967), H = this paper, W = Wagner (1958). Neortholomus, A New Genus of Orsillini 231 Fig. 2. Ncort/iolomiis scolopax (Say), a == pronotum width, b = pronotum length, c ^ scutellum width, d = scutellum length, e = claval commissure length, f ^ corium length, g ^ basal membrane length, h ^ apical membrane length, i = interoccUar space, j =: interocular space, k = head width, 1 z= eye width, m := head length, n = anteocular head length, () = eye height, p = eye length. 232 The University of Kansas Science Bulletin procerodorus nevadensis jamaicensis Fig. 3. Spcrmathecac of named species. Scale =^ 0.15mm. jamaicensis procerodorus Fig. 4. a, c-h, paramcrcs of named species; b-d, i, j, acdeagi of named species. Arrows on g and h indicate carina on shank of paramere. Scale = 0.15mm for a, c-j and 0.30mm for b. NeorthoJomKs, A New Genus of Orsillini 233 0. punctipennis N. scolopax Fig. 5. Sperm reservoirs for named species, a, c, right lateral aspect; b, distal aspect; d, dorsal aspect. Scale = 0.016mm. procerodorus arphnoides nevadensis Fig. 6. a-d, meso- and ineta- pleural areas of named species (dimensions: a/b = scent gland auricle length; c/d =: evaporative area height), c-f, lateral aspect showing relative scutcUum height (right ar- row) to pronotum height (left arrow) of named species. Scale ^ 0.-40mm for a-d. 234 The University of Kansas Science Bulletin koreshanus arphnoides jamaicensis scolopax nevadensis Fig. 7. a-c, head lateral aspects of named species; d-h, hcati dorsal aspects of named species. Scale := 0.1 1mm. »•••■•••••••••< *_•-•-•-•-• ' » • •_•»•.•»•.•.•.•.•_•_• »••_•_••••* »_• _•_•_•_•_•_* .•-•-•-•.•.•.•J i >••••_• •.•.•.•-•-•-•-•-•.•-•-•.••.•-•• ► •.•••.•••• • > •_• •_• • < > * * * *-*-< THE UNIVERSITY OF KANSAS SCIENCE BULLETIN g i V .JO. tw L(l UNIV ■y S REVISION OF THE BEE GENUS LIPHANTHUS (HYMENOPTERA: ANDRENIDAE) S I By Luisa Ruz and Haroldo Toro V V x y 5 VoL 52, No. 8, pp. 235-299 December 23, 1983 ANNOUNCEMENT The University of Kansas Science Bulletin (continuation of the Kansas Uni- versity Quarterly) is an outlet for scholarly scientific investigations carried out at the University of Kansas or by University faculty and students. Since its incep- tion, volumes of the Bulletin have been variously issued as single bound volumes, as two or three multi-paper parts or as series of individual papers. Issuance is at irregular intervals, with each volume prior to volume 50 approximately 1000 pages in length. 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All communications regarding exchanges, sales and subscriptions should be addressed to the Exchange Librarian, University OF Kansas Libraries, Lawrence, Kansas 66045. Reprints of individual papers for personal use by investigators are available gratis for most recent and many older issues of the Bulletin. Such requests should be directed to the author. The International Standard Serial Number of this publication is US ISSN 0022-8850. Editor William L. Bloom Editorial Board Philip W. Hedrick Rudolf Jandcr Harvey Lillywhite Charles D. Michener Norman A. Sladc Henry D. Stone George W. Byers, Chairman THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Vol. 52, No. 8, pp. 235-299 December 23, 1983 Revision of the Bee Genus Liphanthus (Hymenoptera: Andrenidae) LuisA Ruz* "^ AND Haroldo Toro* Contents Abstract 237 Introduction 237 Acknowledgments 237 Materials and Methods 238 Natural History 239 Genus Liphanthus Reed 240 Key to the species and subgenera 241 Subgenus Liphanthus Reed, sensu stricto 245 L. sabulosus Reed 246 L. hrevicornis n. sp 247 L. barbatiis n. sp 251 L. chillanensis n. sp 252 Subgenus Pseudoliphanthus n. subgen 253 L. rozeni n. sp 254 L. spiniventris n. sp 255 L. andinus n. sp 257 L. unifasciatus n. sp 258 Subgenus Xenoliphanthus n. subgen 260 L. parvulus (Friese) 260 L. tofensis n. sp 263 L. moldenkei n. sp 264 L. micheneri n. sp 265 Subgenus Tricholiphanthus n. subgen 267 L. leucostomus n. sp 268 L. pilifrons n. sp 268 L. tarsalis n. sp 270 Subgenus Melaliphanthus n. subgen 271 L. atraius n. sp 271 L. penai n. sp 272 ■^Department of Entomology, University of Kansas, Lawrence, Kansas 6604-5, U.S.A. and *Laboratorio de Zoologi'a, Universidad Catolica de Valparaiso, Av. Brasil 2950, Valparaiso, Chile. Subgenus Neoliphanthus n. subgen 274 L. bicellularis n. sp 275 Subgenus Leptophanthus n. subgen 277 L. nitidus new name 277 L. hreviceps (Friese) 280 L. anacanthus n. sp 280 L. australis n. sp 282 L. coquimbensis n. sp 283 L. cerdai n. sp 285 L. alicahue n. sp 287 Species Not Assigned to Subgenus 289 L. fnesellus n. sp 289 Phylogenetic Considerations 290 Distribution 296 Literature Cited 299 The Bee Genus Liphanthus 237 Abstract The genus Liphanthus is interpreted in a wider sense than that of Reed (1894) and subsequent authors. The following new subgenera, in addition to Liphanthus sensu stricto, are described: Pseudoliphanthus, Xenoliphanthus, Tricholiphanthus, Melaliphanthus, Neoliphanthus, and Leptophanthus. The subgenus Liphanthus sensu stricto includes: L. sabulosus Reed 1894, L. brevicornis, L. barbatus, and L. chillanensis n. spp.; subgenus Pseudoliphanthus: L. rozeni, L. spiniventris, L. andinus, and L. unifasciatus n. spp.; subgenus Xenoliphanthus includes: L. parvulus {Fnesc, 1916), /.. tofensis, L. rnoldenkei, and L. micheneri n. spp.; subgenus Tricholiphanthus: L. leucostomus, L. pilifrons, and L. tarsahs n. spp.; subgenus Melaliphanthus: L. atratus and L. penai n. spp.; subgenus Neoliphanthus: L. bicellularis n. sp.; subgenus Leptophanthus: L. nitidus new name; L. breviceps (Friese, 1916), L. anacanthus, L. australis, L. coquimbensis, L. cerdai, and L. alicahue n. spp. For L. friesellus n. sp. no subgeneric status is given. An analysis showing the probable cladistic relationships among the subgenera is also provided. All the species are described and illustrated. Introduction Liphanthus was originally proposed by Reed (1894) to include only L. sabulosus. Like other species of the genus, L. sabulosus presents such a re- markable sexual dimorphism that Reed could not associate the sexes; as a result he only described the male. The genus Liphanthus was recognized by Ashmead (1899) who correctly placed it among the Andrenidae and clearly differenti- ated it from the other genera of this family. In 1908 Friese redescribed the male of L. sabulosus and considered Liphan- thus as a synonym of Psaenythia. This placement was followed by Ducke (1912), Herbst (1921), Holmberg (1921) and Jaffuel and Pirion (1926). Again, Friese in 1916 described five more closely related species in Psaeny- thia: P. rufiventris, P. parvula, P. pygmea, P. nigra, and P. breviceps; these species are considered as Liphanthus in the present paper. In 1922 Herbst recognized the sex- ual dimorphism in L. sabulosus and L. parvulus and placed P. rufiventris as a synonym of the former and P. pygmaea as a synonym of the latter. Four years later, Claude-Joseph (1926), in biolog- ical studies, confirmed Herbst's sex association for L. sabulosus, an associa- tion that was also indicated by Jaffuel and Pirion in the same year. Contemporary to Friese's studies, Cockerell in 1916, following Reed's conceptions, considered Liphanthus as a clearly distinct genus. The genus was also considered distinct by Michener (1944), Rozen (1967, 1970) and Ehren- feld and Rozen (1977). The genus as here understood con- tains seven subgenera and 26 species. Since several of the species are known from very few specimens, it is nearly certain that more species exist. Acknowledgments We wish to express our profound thanks to the following persons: Dr. C. D. Michener of the University of Kan- sas for his useful suggestions and advice which contributed to the prepa- ration of this paper and for the use of material in the Snow Entomological Museum, University of Kansas; Dr. F. Koch and the kite Dr. E. Konigsmann, both (jl' the Museum fiir Naturkunde der Humboldt-Universitiit zu Berlin, for lending Friese's types; Padre J. S. Moure of the Universidade Federal do 238 The University of Kansas Science Bulletin Parana (Curitiba, Brazil) for study specimens; Dr. R. J. McGinley of Harvard University for sending type material and specimens from the Reed collection; Dr. P. H. Arnaud of the California Academy of Science for study material; Dr. M.J. Vianaofthe Museo Nacional Bernardino Rivada- via (Buenos Aires, Argentina), Senor A. Camousseight of the Museo Nacio- nal dc Historia Natural de Santiago (Chile), Dr. Jorge Artigas of the Uni- versidad de Concepcion (Chile), and Dr. L. Campos of the Facultad de Agronomia de la Universidad de Chile (Santiago, Chile) for permitting study of the material from collections under their care; Professor J. Solervicens of the Universidad Valparaiso (Chile), Seiior H. W. Sielfeld of the Museo de la Patagonia (Punta Arenas, Chile), and Dr. M. Cerda of the Ministerio de Salud (Santiago, Chile) for providing study specimens. Very special thanks to Dr. J. G. Rozen, Jr. of the Ameri- can Museum of Natural History for his valuable suggestions and all the help that he provided for study in that mu- seum. Special thanks to Messrs. Jon K. Gelhaus and Robert W. Brooks of the Department of Entomology, Univer- sity of Kansas for their help. The drawings were made by Sefior Enrique Peralta and the typing of the manuscript was by Mrs. Joetta Weaver. This study was possible thanks to the financial aid of the American Mu- seum of Natural History arranged through Dr. J. G. Rozen, the Univer- sidad Catolica de Valparafso (Chile), and especially the National Science Foundation Grant DEB 77-23035 (C. D. Michener, principal investigator). Materials and Methods This paper is based on material from the collections whose curators are mentioned in the Acknowledgments above. The descriptions of species and sub- genera have been written with charac- ters in the same sequence to facilitate comparisons. The term areolate in the key and descriptions is used as indi- cated by Harris (1979) for very minute reticulate patterns. The specific de- scriptions of females consider only the characters different from those men- tioned in descriptions of males. Sexual characters of females such as antenna, similar in all species, are omitted (see generic characters). The prestigma and pterostigma are about the same color as nearby veins; in the descriptions only vein color is mentioned. The space between the mandibular socket and the transverse part of the hypostomal ca- rina is called the postmandibular area. The section on natural history is based primarily on previously pub- lished information. The data about geographical distribution have been obtained only from the material stud- ied. The measurements were taken with a Wild 5 stereoscopic microscope equipped with a 10 X micrometer in the manner indicated by Michener (1965). The length of the propodeal triangle laterally is measured two thirds of the distance from the midline to the lateral extremity of the triangle and compared to the length of the meta- notum measured at the same distance from the midline. The thickness of the clypeus (in side view) is measured from its tangent to its lateral extremity. The drawings were made using a camera lucida, comparable structures being drawn at about the same size regardless of dilierences in size of the bees. The genitalia were cleared in KOH (10%) for 24 h (cold solution) or for 30 min at 70° C. The Bee Geni's Liphanthus 239 Most holotypes and allotypes have been deposited in the Toro collection, now at the Universidad Catolica de Valparaiso, Chile. The paratypes will be broadly distributed in ditlerent col- lections as shown by the abbreviations at the end of each description. These abbreviations are as follows: AMNH, American Museum of Natural His- tory, New York; NMNH, National Museum of Natural History, Washing- ton, D.C.; CAS, California Academy of Sciences, San Francisco; KU, Uni- versity of Kansas, Lawrence; MCZ, Museum of Comparative Zoology, Harvard University, Cambridge, Mas- sachusetts; MIZT, Museo ed Istituto di Zoologia Sistematica della Univer- sita di Torino, Italy; BMNH, British Museum (Natural History), London, England; MNHUB, Museum fiir Na- turkunde der Humboldt-Universitat zu Berlin, Germany; MNHNP, Musee National d'Histoire Naturelle de Paris, France; MLP, Museo de la Plata, Ar- gentina; MNBR, Museo Nacional Ber- nardino Rivadavia, Buenos Aires, Argentina; CONG, Universidad de Concepcion, Chile; FAUCH, Facultad de Agronomia de la Universidad de Chile, Santiago, Chile; UCV, Univer- sidad Catolica de Valparaiso, Chile; HT, Toro collection, Valparaiso, Chile. Natural History Little information about the natural history of Liphanthus exists. The avail- able published data refer only to L. sabulosus and L. parvulus. Claude- Joseph (1926) indicates that L. sabulosus nests in clayey soil, describing briefly at the same time the shape and direc- tion of the burrows and the egg location relative to the mass of food. Rozcn (1967) notes that the linear disposition of the nest cells is unusual among the Panurginae. Rozen (1970), studying the biology of Kelita chilensis, points out that this nomadine bee is a parasite of L. par- vulus and describes in detail the place- ment of the parasite egg in the cell. Some years earlier, Friese (19 1()) had also suggested a similar association be- tween Kelita and L. sabulosus ( = rufiven- tris) but apparently without any proof of the association. Moreover, Ehrcn- feld and Rozen (1977) indicate that a small undescribed species of Liphanthus here described as L. alicahue is probably associated with another nomadine bee, Kelita tuberculata. Moldenke and Neff (1974) indicate that L. sabulosus and L. parvulus are associated with the following plants: L. sabulosus: Chaetanthera incana, Pleurophora presilla, Chaetanthera sp., Adesmia sp., Hoffmannseggia falcaria; and L. parvulus: Adesmia arborea, Adesmia angustifolia, Adesmia sp., Calandrinia sp., Aristolochia sp. Personal observations are that cer- tain species of Liphanthus visit the fol- lowing plants: L. parvulus: Oxalis sp.; L. chillanensis n. sp.: Oxalis sp.; and L. coquimbensis n. sp.: Gutierrezia taltalensis, Mesembrianthemum cristalinum, Baccharis paniculata. Biogeographic Considerations. — Ac- cording to Moldenke (1976) Liphanthus is endemic to Chile. It is now known that it is primarily restricted to Chile. Its species range from Antofagasta to Chile Chico (between about 22° S and 46.5° S). It also extends, however, to Argentinean Patagonia from Chubut (40.3° S approx.) to Santa Cruz near Los Antiguos (47° S approx.) (Maps I and II). Two undescribed species are fromjujuy, Argentina. In the scarce material available, little specific diversity has been ob- served in the arid zones of the north, but the number of species increases greatly in central and southern Chile. 240 The University of Kansas Science Bulletin The altitudinal distribution of the spe- cies is principally below 2,500 m, especially in areas of abundant vegeta- tion. The distributions of the sub- genera are considered after the cladistic analysis. Genus Liphanthus Reed Small bees (3 to 7 mm long), non- metallic black, sometimes with red metasoma; yellow areas conspicuous in males, sometimes gready reduced in females; pilosity reduced, not forming metasomal hair bands; usually with three submarginal cells. Coloration. Head of male with yellow marks on mandibles, labrum (generally), clyp- eus, lower paraocular area (except in Melaliphanthus), and legs, sometimes forming one or two metasomal bands; marks reduced and more variable in female. Pubescence. Yellowish white, brown on last metasomal terga. Hairs, in general, short and sparse, mostly simple, with some briefly branched hairs mixed with simple ones. Distal margin of clypeus, at each side of midline, with two hairs so close to each other as to appear fused, and generally longer than the rest. Propodeal triangle laterally pubescent (scarcely hairy in L. fnesellus, umjasciatus, and atratus) . Meta- somal terga with scarcely visible hairs; sterna with sparse hairs. Integument. Generally areolate, with some longitu- dinal striation on propodeal triangle (apparendy absent in females of L. nitidus and coquimbensis). Punctation rather sparse and shallow, usually coarse on clypeus, absent on middle ot basal area of propodeal triangle, fine and generally dense on metasomal terga but coarser on the last ones, absent on broad, usually depressed, marginal areas of terga, sparse on sterna and absent on very narrow smooth marginal areas. Labrum smooth proxiinally. Structure. Head somewhat broader than thorax. Episto- mal suture below outer subantennal suture distinctly angulate, resulting an- gle of lower paraocular area smooth and usually swollen. Tentorial pit gen- erally puncture-like in shape. Su- praclypeal and interalveolar areas generally prominent. Flagellum longer than head in male, shorter than head and unmodified in female. First flagel- lar segment longer to much longer than broad in male, less than twice as long as second one in female. Facial fovea usually more distinct and longer in female than in male. Labrum in female less than twice as broad as long, with well marked transverse preapical ca- rina, distal margin convex. Mandible without tooth (expanded on upper margin before apex in L. rozeni and umjasciatus). Glossa elongate. Maxil- lary palpus six-segmented, first three segments of similar length, last three progressively shorter. Labial palpus with four thin segments, first one about as long as rest of segments together. Hypoepimeral area trapezoidal. Scu- tellum about twice as long as meta- notum. Marginal cell with truncation strongly slanting. Pterostigma narrow, slightly broader than prestigma, with sub-parallel sides, stigmal margin within marginal cell straight. Generally with three submarginal cells, one sub- genus with two. Tibial spurs microser- rate. Basitibial plate with marginal rim, somewhat widened distally and rounded in female, narrower at apex than base in male. Male with bifurcate claws. Second metasomal tergum prox- imally with strong transverse groove; lateral fovea elongate and well marked, wider and longer in female than in male. Pygidial plate well developed in female; in male, if present, without median longitudinal ridge. Metasomal sternum VII X-shaped, sheet-like lat- eral expansions small. Metasomal Thp: Bee Genus Liphantiujs 241 sternum Vlll longer than broad, ar- rowhead-shaped with rounded apex. Genital capsule without gonobase, gonocoxal apodeme narrow; gonocox- ite rather elongate; gonostylus articu- lated, more than half of length of gonocoxite; penis valve usually reach- ing gonostylar apex; ventral basipenial plate rather broad, sclerotized, fused with penis at its base; volsella without special modifications. Discussion: Liphanthus, among the Panurginae, could be close to: Meta- psaenythia, Protandrena, Pterosarus, He- terosarus, and Pseudopanurgus. From Metapsaenythia, Pterosarus, Heterosarus, and Pseudopanurgus it usually is sepa- rated easily by having three submargi- nal cells. From the above genera including Protandrena it is differentiated by its narrow pterostigma and usually by its sparse punctation. The recognition of Liphanthus is fa- cilitated by the head which is broader than the thorax, and by the groove on the second metasomal tergum which differs from that of other terga. Etymology: The derivation of the name Liphanthus is not explained in the original description. According to Dr. Michael H. Shaw, Professor of Classics at the University of Kansas, the word Liphanthus cannot be easily broken down into Greek elements. Dr. Robert L. Lind, also Professor of Classics (re- tired) at the University of Kansas, sug- gests that Liphanthus may have been derived from the Greek leipo which means lack and anthos which means flower. This is perhaps related to the fact that the males often fly in dried areas without flowers. Dr. George W. Byers of the Department of Entomol- ogy of the University of Kansas postu- lates that Liphanthus is an anagram of Philanthus, a genus of small sphecids of black and yellow color (as in Liphan- thus). This explanation is strongly sup- ported by the fact that Rec-d included L. sabulosus in Philanthidae. Key to riiK Species and Subgenera 1. Males 2 Females 27 2(1). Submarginal cells two (Fig. 108). Propodeal triangle laterally less than half length of meta- notum (laterally). Subgen. Neo- liphanthus L. bicellularis Submarginal cells three (Figs. 5 and 53). Propodeal triangle lat- erally more than half length of metanotum (laterally) or almost same length 3 3(2). Hind tibial spurs with api- ces curved like claws, subequal in length (Fig. 56). Vertex con- cave or almost straight in frontal view. Subgen. Xenoliphanthus . ... 4 Hind tibial spurs with apices only slightly curved, inner some- what longer than outer (Fig. 4). Vertex convex in frontal view ... 7 4(3). Clypeus almost 4 times as broad as long, yellow without black spots 5 Clypeus about 3 or little more than 3 times as broad as long, yellow with 2 basal black spots; if there are no spots, integument of mesonotum areolate and with homogeneous punctation 6 5(4). Hypoepimeral area scarce- ly punctate. Marginal cell very slightly longer than distance from apex to wing tip . . L. parvulus Hypoepimeral area densely punctate. Marginal cell conspic- uously longer than distance from apex to wing tip L. tofensis 6(4). Second metasomal sternum with 2 strong protuberances side by side in the middle. Head more than 1.5 times as broad as long; metasoma black. . L. micheneri Second metasomal sternum 242 The University of Kansas Science Bulletin without protuberances. Head less than 1.5 times as broad as long; metasoma reddish L. moldenkei 7(3). Distance between alveolus and inner orbit shorter than in- teralveolar distance or similar ... 8 Distance between alveolus and inner orbit longer than inter- alveolar distance 16 8(7). Flagellar segment 1 about 3 times as long as broad. Clypeus flattened basally. Subgen. Melali- phanlhus 9 Flagellar segment 1 less than twice as long as broad. Clypeus convex. Subgen. Leptophanthus . . 10 9(8). Sternum III with posterior margin projecting mesally. Fa- cial fovea almost half as long as scape. Clypeus yellow on distal third L. atratus Sternum III with posterior mar- gin almost straight. Facial fovea less than one-third as long as scape. Clypeus yellow only on distal margin L. penai 10(8). Sternum II with projection broadly truncate apically, less than twice as long as broad (width measured at apex) 11 Sternum II with projection acute or slightly truncate apically, two or more times as long as broad (width measured at apex) 12 11(10). Sternum II with projection little longer than its apical width, posterior distal margin clearly sinuate lateral to projection (Fig. 117) L. breviceps Sternum II with projection not salient, distal margin scarcely sinuate lateral to projection (Fig. 123) L. anacanthus 12(10). Labrum with transverse ca- rina at distal fifth and distal pubescent band almost linear (Fig. 128 and 135) 13 Labrum with transverse carina at distal third and distal pubes- cent band wide (Figs. 110, 141, and 147) H 13(12). Sternum II with posterior margins laterally gradually con- vergent toward projection. Ster- num III with weak median protuberance (Fig. 129) . L. australis Sternum II with posterior mar- gin distincdy sinuate lateral to acuminate projection. Sternum III flat (Fig. 139) . . L. coquimbensis 14(12). Face above level of alveoli slightly areolate and shiny. Sub- antennal area generally black. . L. nitidus Face above level of alveoli strongly areolate and opaque. Subantennal area generally yel- low 1-^ 15(14). Flagellum distinctly longer than width of head; flagellar seg- ments 9 and 10 distinctly longer than broad (Fig. 140). Meso- notum strongly areolate, with punctures sparse and weakly marked L. cerdai Flagellum scarcely longer than width of head; flagellar segments 9 and 10 almost as broad as long (Fig. 146). Mesonotum almost smooth between punctures, if areolate, always with well marked and dense punctures . . L. alicahue 16(7). Metasomal sterna III and IV with posterior margins usu- ally clearly concave (if slightly concave, with median, de- pressed, triangular, transparent area between apical thickenings). Subgen. Tricholiphanthus 17 Metasomal sterna III and IV with posterior margins convex or straight (without thickenings) . . 19 17(16). Pronotum with slightly protuberant dorsolateral angle. Pubescent margin of second ster- Thf. Bee Genus Liphanthus 243 nal projection as broad (side to side) as concavity on posterior margin of metasomal sternum III (Fig. 86) L. pilifrons Pronotum with distinctly protu- berant dorsolateral angle. Pubes- cent margin of sternal projection narrower (side to side) than con- cavity on posterior margin of metasomal sternum III (Figs. 78 and 89) 18 18(17). Hind basitarsus with base broader than apex, second tarsal segment somewhat longer than broad L. tarsalis Hind basitarsus parallel-sided, second tarsal segment twice as long as broad L. leucostomus 19(16). Clypeus, in side view, con- cave basally and thickness (due to great protuberance) greater than length (Fig. 153) . . L. friesellus Clypeus, in side view, convex basally and thickness similar or smaller than length (Fig. 2) ... 20 20(19). Pygidial plate present. Head in ventral view with latero- ventral area of clypeus widened mesally (Fig. 3). Frontal line in a very long and well marked groove. Subgen. Liphanthus. ... 21 Pygidial plate absent. Head in ventral view with lateroventral area of clypeus widened laterally (Fig. 45). Frontal line scarcely marked, not in groove. Subgen. Pseudo liphanthus 24 21(20). Flagellum gradually en- larged distally. Laterodistal mar- gin of clypeus in frontal view strongly concave (Fig. 16) and with long hairs L. barbatus Flagellum not enlarged distally. Laterodistal margin of clypeus almost straight (Fig. 1), with short hairs only 22 22(21). Flagellum longer than head; flagellar segments longer than broad, last one about two and a hall times as long as broad (Fig. 11) L. brevicornis Flagellum generally distinctly longer than head; flagellar seg- ments about twice as long as broad or more, last one three or more times as long as broad ... 23 23(22). Last flagellar segment dis- tally about half as broad as me- dian segments, strongly curved (Fig. 1). Scape with yellow spot L. sabulosus Last flagellar segment about as broad as median segments, al- most straight (Fig. 21). Scape without yellow spot . . L. chillanensis 24(20). Metasomal sternum II without postero-median projec- tion. Outer subantennal suture slightly arcuate laterally. Meta- soma with one yellow transverse band L. unifasciatus Metasomal sternum II with post- ero-median projection. Outer subantennal suture strongly ar- cuate laterally. Metasoma with 2 yellow transverse bands 25 25(24). Projection of sternum II acute (Fig. 35). Flagellum longer than head width . . . . L. spiniventris Projection of sternum II truncate (Figs. 40 and 30). Flagellum as long as head width or less .... 26 26(25). Metasomal sternum III with postero-median protuber- ance. Posterior margin of ster- num II with median projection much broader than long (Fig. 30). Mandible with preapical widened plate (similar as in Fig. 154) L. rozeni Metasomal sternum III unmodi- fied. Posterior margin of ster- num II with median projection about as long as apical width (Fig. 40). Mandible not widened preapically L. andinus 244 The University of Kansas Science Bulletin 27(1). Submarginal cells two. Propodeal triangle laterally about half length of metanotum (laterally). Subgen. Neoliphanthus L. bicellularis Submarginal cells three. Pro- podeal triangle laterally slightly more than half length of meta- notum (laterally) or almost same length '. 28 28(27). Hind tibial spurs with api- ces curved like claws, subequal in length (Fig. 56). Vertex con- cave or almost straight in frontal view. Subgen. Xenoliphanthus ... 29 Hind tibial spurs with apices slightly curved, inner somewhat longer than outer (Fig. 4). Ver- tex convex in frontal view .... 32 29(28). Claws apparently simple. Clypeus almost 4 times as broad as long 30 Claws bifurcate. Clypeus scarcely more than 3 times as broad as long 31 30(29). Hypoepimeral area almost smooth. Marginal cell scarcely longer than distance from apex to wing tip L. parvulus Hypoepimeral area densely punctate. Marginal cell clearly longer than distance from apex to wing tip L. tofensis 31(29). Mesonotal integument dis- tinctly areolate. Metasoma black L. micheneri Mesonotal integument almost smooth. Metasoma red L. moldenkei 32(28). Lateral ocelH, in frontal view, above upper orbital tan- gent. Inner orbits divergent dor- sally. Subgen. Leptophanthus ... 33 Lateral ocelli, in frontal view, at same level or slightly below up- per orbital tangent. Inner orbits convergent dorsally or ahnost subparallel 37 33(32). Integument above aheoli smooth and shiny. Pronotal lobe black \ 34 Integument above alveoli strongly areolate and dull. Pro- notal lobe generally with distal yellow spot 35 34(33). Base of mid tibia with yel- low spot about as long as last flagellar segment. Clypeus gen- erally black L. coquimbensis Base of mid tibia with yellow spot much shorter than last flagellar segment. Clypeus gen- erally with distal yellow spot . . L. nitidus 35(33). Mesonotum areolate, scarcely punctate. Clypeus gen- erally with small yellow spot dis- tally. Length about 4 mm . L. cerdai Mesonotum almost smooth be- tween punctures, punctation moderately dense. Clypeus black or with longitudinal yellow band. Length about 5 mm or more 36 36(35). Facial fovea somewhat shorter than scape. Clypeus with longitudinal yellow band L. alicahue Facial fovea longer than scape. Clypeus black L. anacanthus 37(32). Outer subantennal suture almost straight (Figs. 90 and 96). Clypeus, inferior paraocular area and base of mandible with- out yellow. Subgen. Melaliphan- thus 38 Outer subantennal suture dis- tinctly arcuate laterally (Figs. 1 and 26). At least clypeus with yellow 39 38(37). Frontal line, in lower half, hardly visible. Propodeal tri- angle sparsely striate. Tibial scopa less dense than hairs of fore femur L. penai Frontal line, in lower half, a nar- row groove. Propodeal triangle densely striate. Tibial scopa The Bee Genus Liphanthus 245 dense, similar to hairs of fore femur L. atratus 39(37). Frontal line almost imper- ceptible. Lower paraocular area and tegula without yellow. Sub- gen. Pseudoliphanthus 40 Frontal line distinct, in a groove. Lower paraocular area and tegula with yellow. Subgen. Li- phanthus 41 40(39). Clypeus (central area) de- pressed distally, more than 3 times as broad as long. Suban- tennal area and basal half of mandible yellow. Metasomal terga yellowish L. rozeni Clypeus slightly convex, less than 3 times as broad as long. Subantennal area black. Mandi- ble mostly dark mahogany. Met- asomal terga black . . . . L. andinus 41(39). Marginal cell generally longer (or similar in length) than distance from apex to wing tip. Second submarginal cell with an- terior margin about as long as posterior or somewhat shorter. Paraocular spot extending ven- trally to level of tentorial pit or slightly below it L. sabulosus Marginal cell usually shorter than distance from apex to wing tip. Second submarginal cell with anterior margin distinctly shorter than posterior. Paraocu- lar spot generally extending ven- trally farther below level of ten- torial pit 42 42(41). Integument of thorax al- most smooth. Facial fovea sin- uous, widened ventrally L. brevicornis Integument on thorax strongly areolate. Facial fovea almost straight, not widened ventrally 43 43(42). Facial fovea subparallel to inner orbit. Clypeus 3 times as broad as long. Outer surface of mid basitarsus with small basal yellow spot L. barbatus Facial fovea ventrally farther from inner orbit than dorsally. Clypeus less than 3 times as broad as long. Outer surface of mid basitarsus with large yellow area extending beyond middle . L. chiUanensis Subgenus LiPU.\.\'ruL's Reeu, SENSU STRICTO Figs. 1-25 Liphanthus Reed, 1894: 645. Type species: Liphanthus sabulosus Reed, 1894 (monobasic). Both sexes: Vertex convex. Lateral ocelli at same level as or below upper orbital tangent. Interalveolar area prominent. Supraclypeal area convex. Frontal line in a groove above alveoli. Clypeus convex, lateral area trans- versely elongate, sloping upward to- ward central area, widened mesally in ventral view. Inner orbits almost straight medially, divergent below. Distance between alveoli less than dis- tance from alveolus to inner orbit and longer than alveolus diameter. Outer subantennal suture arcuate laterally (Fig. 1). Pronotum without protuber- ant dorsolateral angle. Submarginal cells three. Propodeal triangle laterally more than half length of metanotum (laterally). Tibial spurs yellow or pale testaceous. Hind tibial spurs with api- ces slightly curved, inner slightly longer than outer. Male: Head and legs with yellow marks, metasoma with two yellow transverse bands. Lateral area of clyp- eus with hairs sparse (dense in L. bar- batus) and shorter than clypeal length. Flagellum generally much longer than head; first flagellar .segment approx- imately three times as long as broad. Facial fovea short, clearly separated 246 The University of Kansas Science Bulletin from inner orbit, half as long as scape or less. Middle of paraocular area with transverse swelling below facial fovea; lower part of paraocular area rather flat. Postmandibular area mesally much less than half as long as width of base of mandible. Sterna unmodified. Pygidial plate present. Female: Lower paraocular area and tegula each with yellow spot. Tibial scopa somewhat dense, similar to or less dense than hairs of fore femur. Third flagellar segment broader than long. Liphanthus (Liphanthus) sabulusus Reed Figs. 1-10; Map I, A Liphanthus sabulosus Reed, 1894: 646, 652; Cockerell, 1916: 428; Rozen, 1967: 5, 9; Rozen, 1970: 147; Ehrenfeld and Rozen, 1977: 3. Psaenythia sabulosa, Friese, 1908: 32, 39; Herbst, 1922: 183; Jaffuel and Pi- rion, 1926: 369; Claude-Joseph, 1926: 213. Psaenythia rufiventris Friese, 1916: 166, 169. Male: Length about 6 m, forewing length 4 mm, head width 2.5 mm, thoracic width 2 mm. Coloration. Head and thorax black, the following parts yellow: face below median ocellus and facial fovea, labrum, mandible (apex mahogany), under side of scape (upper surface brown), flagellum (sometimes brown apically and with upper surface or segmental spots brown), pronotal lobe, small spot on tegula, distal mar- gins and under surfaces (variable) of trochanters, ventral edges and apices of fore and middle femora, apex of hind femur, tibiae and tarsi (except middle distitarsus and small segments of hind tarsus which are light brown). Wings almost hyaline; veins and tegula testa- ceous. Metasomal terga black, terga I and II each with transverse yellow- band (wider on II). Metasomal sterna brown. Pubescence. In general short and sparse. Labrum pubescent on distal margin. Lateral areas of clypeus scarcely pubescent, with hairs shorter than those of labrum. Integument and Punctation. Head and thorax lightly to strongly but finely areolate and shiny, smooth on gena and yellow parts of face, with some striation on propodeal triangle. Facial punctation weak and sparse, deeper and denser in orbito- ocellar area (strongly marked in other specimens) and dorsal half of gena, and coarsest on lower central part of clyp- eus. Dorsal part of thorax finely punc- tate, sparsely so on discal scutal area, punctures fine and sparse on hypo- epimeral area, denser and deeper on rest of mesepisternum. Structure. Head approximately 1.5 (rarely 1.3) times as broad as long. Flagellar segments much longer than broad, last one more slender than others, three or more times as long as broad and distinctly curved. Facial fovea about one-third as long as scape, oval, generally narrower near dorsal end. Inner subantennal suture with upper part strongly angu- late toward mid-line of face (almost straight in some specimens) and shorter than width of subantennal area. Clypeus about 4 times as broad as long, projecting beyond lower orbital tangent for about one-third of its length, with longitudinal protuber- ance. Labrum flat, about twice as broad as long or less, distal margin slightly convex. Mandible unmodified. Marginal cell about as long as distance from apex to wing tip. Second sub- marginal cell with anterior margin about as long as posterior or somewhat shorter. Legs and sterna unmodified. Genitalia and associated sterna as illus- trated. Female: Length about 7 mm, fore- wing length 4.5 mm, head width 2.5 mm, thoracic width 2 mm. Coloration. The Bee Genus Liphanthus 247 Head and thorax black, antennal flagellum light mahogany beneath, the following parts yellow: area lateral to outer subantennal suture extending down to tentorial pit or slightly below it, subantennal area, T-shaped spot on upper part ofclypeus, base of" mandible (apex mahogany), pronotal lobe, small spots on apices of fore and mid femora and bases of fore and mid tibiae (tarsi dark brown). Wings and tegula as de- scribed for male but somewhat darker. Metasoma red (somewhat darker in some specimens) except base of seg- ment I, lateral fovea on tergum II, margin of tergum V and pygidial plate which are black (the last sometimes dark mahogany). Pubescence. As de- scribed for male but denser and longer on head, mesepisternum and legs; tibial scopa somewhat densely hairy, but less dense than hair of fore femur. Integument and Punctation. Head and thorax lightly, fmely (sometimes more strongly) areolate, smooth on lower parts of face and gena. Densely and coarsely punctate on head, sparser and coarser on clypeus, finer and denser above antennal sockets and on genal area. Thorax with medium-sized punc- tures, sparse in discal part of scutum, deeper and sparse on scutellum, dense on mesepisternum. Pygidial plate slightly areolate. Structures. As de- scribed for male except: facial fovea shorter than scape and slightly sinuate (almost straight in some specimens). Clypeus approximately 3 times as broad as long, with weak longitudinal elevation on upper half. Lower part of frontal line carinate, apex minutely grooved. Marginal cell generally longer than (or equal to) distance from its apex to wing tip. Second submargi- nal cell with anterior margin somewhat shorter than posterior. Pygidial plate almost flat apically. The specimens, especially males, show wide variability in some charac- ters, such as size, sculpturing (from smooth to areolate) and curvature of the subantennal suture. Color varia- tions also exist, from yellow to brown on the labrum, different spot sizes on the legs, and sporadic presence oi a yellow spot on gena. Material Studied (all from Chile): 7 males and 2 females, Coquimbo Prov., diverse localities, from October to Jan- uary; 2 males and 5 females, Acon- cagua Prov., January and February; 87 males and 22 females (2 more with- out data), Valparaiso Prov., diverse localities, October to February; 28 males and 5 females, Santiago Prov., diverse localities, December to Janu- ary. Altitudes range from sea level to at least 2200 m. Among the specimens collected by P. Herbst, 2 males labeled as Type were found in the Museum of Com- parative Zoology, Harvard University. One of them (with broken head) has a label that indicates it was collected in Valparaiso in 1900. The other is head- less, with an additional red label which reads "Type coll. Edw. C. Reed," but no date, locality or collector name is provided. The latter specimen (unfor- tunately headless) is the only known syntype, and is here designated as lec- totype and has been so labeled. Liphanthus (Liphanthus) brevicorms new species Figs. 11-15; Map I, B Diagnosis: Similar to L. sabulosus but male with flagellum not distinctly longer than head, last flagellar segment as broad as median segments and about 2V2 times as long as broad, fore and mid femora sometimes with short yellow spot on ventral edge; female with marginal cell generally shorter than distance from its apex to wing tip, yellow spot on paraocular area usually 248 The University of Kansas Science Bulletin Figs. 1-10. Liphanthus (Liphanlhus) \ahitlosus Reed, male: 1, 2, 3, frontal, lateral and \entral \-iews of head; 4, hind tibial spurs; 5, wings; 6, inaiulihie; 7, labrum; 8, 9, seventh and eighth inetasomal sterna; 10, dorsal and ventral views of genitalia. Figs. 11-15. Liphanthus {Liphanthus) hrevicomis n. sp., male: 11, head frontal view; 12, labrum; 13, 14, seventh and eighth metasomal sterna; 15, dorsal and ventral views of genitalia. extending farther below tentorial pit or touching epistornal suture, second sub- marginal cell with anterior margin dis- tinctly shorter than posterior. Male: Length about 6 mm, {bre- wing length 4 mm, head width 2 mm, thoracic width 1.5 mm. Coloration. Head and thorax black, distal margins of clypeus and labrum brown, the fol- lowing parts yellow: face below median ocellus and facial fovea, mandible (apex mahogany), middle part of under side of flagellum (sometimes extending to most of its ventral surface), pronotal The Bee Genus Liphanthus 249 lobe, small spot on tcgula, apex and part of ventral edge of tore femur, tibiae (except brown spot on both sur- faces of tibiae I and II), tarsi (except small segments of hind tarsus which are brown). Wings slightly testaceous, veins brown. Tegula testaceous. Meta- somal terga black with transverse yellow bands on terga I and II (wider on II). Metasomal sterna brown. Pubes- cence. In general short and sparse. La- brum pubescent on distal margin. Lateral areas of clypeus scarcely pubes- cent, with hairs shorter than those of labrum. Integument and Punctation. In general almost smooth, shiny on gena, and yellow parts of face, finely punc- tate on yellow parts, punctures some- what denser and coarsest on clypeus, deeper on dorsal part of head, fine on dorsal part of thorax apd sparse on hypoepimeral area, denser on rest of mesepisternum. Structure. Head ap- proximately 1 .4 times as broad as long. Flagellar segments somewhat longer than broad, last one about 2'/2 times as long as broad. Facial fovea about one- third as long as scape, usually oval (somewhat variable). Inner subanten- nal suture with upper part distinctly arcuate toward mid-line of face and shorter than width of subantennal area. Clypeus about 4 times as broad as long, projecting beyond lower orbital tangent for about one-third of its length, with a weak longitudinal pro- tuberance. Labium flat, a little more than twice as broad as long, distal margin varies from slightly convex to almost straight. Mandible unmodified. Marginal cell shorter than distance from apex to wing tip. Second sub- marginal cell with anterior margin dis- tinctly shorter than posterior. Legs and sterna unmodified. Genitalia and asso- ciated sterna as illustrated. Female: Length about 5.5 mm, fore- wing length 3.5 mm, head width 2 mm, thoracic width 1 .5 mm. Coloration. Head and thorax black, fiagellum brown beneath, the following parts yellow: area lateral to outer subanten- nal suture extending down below ten- torial pit, usually touching epistomal suture, subantennal area, T-shaped spot on upper part of clypeus, base of mandible (apex mahogany), pronotal lobe, small spots on apices of fore and mid femora and bases of fore and mid tibiae and of basitarsi (the latter some- times absent) (small segments of tarsi brown). Wings and tegula as described for male. Metasoma red, except most of segment I, lateral fovea on tergum II and distal margin of tergum V which are black and pygidial plate dark ma- hogany. Sterna II and III each with central area brown. Pubescence. As de- scribed for male, but denser and longer on head, mesepisternum and legs; tibial scopa less dense than hair of fore femur. Integument and Punctation. Head and thorax lightly, finely areolate, smooth on lower parts of face, gena, and pygidial plate apically, densely and coarsely punctate on head, sparser and coarser on clypeus, finer and denser above antennal sockets. Thorax with medium-sized punctures, sparse on discal part of scutum, deeper and sparse on scutellum, fine and sparse on hypoepimeral area, coarser and denser on rest of mesepisternum (sparser than L. sabulosus). Structure. As described for male except facial fovea about as long as scape, usually sinuate, widened dis- tally. Clypeus somewhat more than three times as broad as long. Lower part of frontal line minutely grooved. Marginal cell usually shorter than dis- tance from its apex to wing tip. Pygidial plate with rounded longitudi- nal median ridge on apex. Type Material: Holotype male. Val- paraiso Prov., El Granizo, Chile, XI-1964 (F. Rojas). Allotype female. 250 The University of Kansas Science Bulletin Valparai'so Prov., El Belloto, 30-X-66 (H. Toro), both in Toro collection. Paratypes (all from Valparaiso Prov., Chile, except as indicated). 16 males, Olmue, X-1917 (3 of them without abdomens) and 4 females, XI-1900 (P. Herbst); 1 male. El Salto, 2-II-1968 (H. Toro); 4 males, Granizo, XI-1964 (F. Rojas); 1 male, Granizo, XI-1964 (V. Cabezas); 5 females, Belloto, 22- X-1967 (H. Toro); 5 females, Belloto, 30-X-1966 (H. Toro); 1 female, Bel- loto, 19-XI-1966 (H. Toro); 1 female, Colliguay, 18-X-1964 (H. Toro); 3 females, El Salto, 29-X-1967 (H. Toro); 4 males. El Roble, 2100 m, 7- XI-1971 (H. W. Sielfeld); 1 female, Santiago Prov., road to Maitenes, Rio Colorado, 31-X-1971 (J. G. Rozen, L. Peiia); 2 females, O'Higgins Prov., Rancagua, 12-X-1964 (L. Marnef); 1 male, Cautin Prov., Temuco, XI-1900 (P. Herbst). Paratypes are deposited in the following collections: KU, AMNH, MNHN, CAS, MCZ, and UCV. Etymology: The specific name is based on brevis (short) and cornu (horn) with reference to the short antenna. Figs. 16-20. Liphanlhus (Liphanlhui) barbalm n. sp., male: lb, head IroiUal \icw; 17, labium; 18, eighth metasomal sternum; 19, dorsal andventral views of genitalia; 20, seventh metasomal sternum. Figs. 21-25. Liphanlhus (Liphanlhus) chillanrnsis n. sp., male: 21, head frontal view; 22, labrum; 23, dorsal and ventral views ol genitalia; 24, 25, seventh and eighth metasomal sterna. The Bee Genus Liphanthus 251 Liphanthus (Liphanthus) barbatus new species Figs. 16-20; Maj^ I, I Diagnosis: Close to L. sabulosus but male with antenna gradually enlarged distally, lateral areas of clypeus with long pilosity and distal clypeal margin concave; female with longitudinal yel- low band on clypeus, metasoma black, and punctures on mesepisternum shal- low. See also the characters given in the kev. Male: Length about 5.5 mm, fore- wing length 3.5 mm, head width 2 mm, thoracic width 1.5 mm. Coloration. Head and thorax black, the following parts yellow: face below median ocellus and facial fovea (except lateral areas of clypeus which are mostly brown and supra-alveolar area which is black), labrum (with a dark spot in some speci- mens), mandible (apex mahogany), under side of most of flagellum (but with an oblique, dark spot on flagellar segments 2, 3, and 4), pronotal lobe, small spot on tegula, apices of femora (ventral edge of femur I), tibiae and tarsi (except a dark spot on internal side of tibia III and in some specimens distitarsi dark). Wings slightly testa- ceous, veins brown. Tegula testaceous. Metasomal segments black, terga I and II each with transverse yellow band (much wider on II). Pubescence. In gen- eral short and sparse. Labrum pubes- cent on distal margin. Lateral areas of clypeus densely pubescent, with hairs longer than those of labrum. Integument and Punctation. Head and thorax areo- late, smooth between punctures on ventral part of gena and yellow parts of face. Punctation rather coarse and moderately sparse on clypeus, denser on orbito-ocellar area and upper half of gena. Dorsal surface of thorax finely punctate, punctures fine and dense on hypoepimeral area, coarser on rest of mesepisternum. Structure. Head about 1.5 times as broad as k^ig. Flagellar segments 1-8 longer than broad, 9-11 as long as broad, last one truncate. Facial fovea small, about one-third as long as scape. Inner subantennal su- ture with upper hcilf slightly arcuate toward mid-line of face, about as long as width of subantennal area. Clypeus somewhat more than 4 times as broad as long, convex, projecting beyond lower orbital tangent for about one- third of its length. Labrum flat, some- what broader than long, distal margin with central part emarginate. Mandi- ble unmodified. Marginal cell dis- tinctly shorter than distance from apex to wing tip. Second submarginal cell with anterior margin shorter than pos- terior. Legs and sterna unmodified. Genitalia and associated sterna as illus- trated. Female: Length about 6 mm, fore- wing length 4 mm, head width 2 mm, thoracic width 1.6 mm. Coloration. Head and thorax black, the following parts yellow: area lateral to outer sub- antennal suture extending down below tentorial pit (somewhat variable in size), subantennal area, longitudinal median spot on clypeus, mandible (apex mahogany), under side of flagel- lum (except most of basal segments and sometimes the distal ones, brown), pro- notal lobe, spot on tegula, small spot on apices of fore and mid femora, bases of tibiae (somewhat extended on exter- nal side of tibia I). Wings and tegula as described for male. Metasomal terga black, pygidial plate dark mahogany; sterna brown. Pubescence. As described for male but much longer and denser on head, mesepisternum and legs; tibial scopa somewhat densely hairy but less dense than hair of fore femur. Integument and Punctation. Integument strongly areolate, almost dull between punctures. Punctures rather dense and coarse on clypeus, finer on rest of head. 252 The University of Kansas Science Bulletin Thorax with rather small, shallow punctures. Hypoepimeral area and rest of mesepisternum with punctation less dense and less deep than in male. Pygidial plate apically smooth. Struc- ture. As described for male except: head 1.2 times as broad as long; facial fovea almost as long as scape and narrow. Clypeus three times as broad as long. Lower part of frontal line a narrow groove. Pygidial plate with narrow ridge on apex. Type Material: Holotype male and al- lotype female. Malleco Prov., Icalma, Chile, 11-1-1979 (O. Martmez), in Toro collection. Paratypes. (All from Malleco Prov., Chile.) 5 males and 2 females, Galletue, Lonquimay area, 9-12-1-1962 (L. Peha) (2 of the males parasitized by Strepsiptera); 4 males, Galletue at source of Bio-Bi'o, near Argentine border, 1-1-1968 (L. Pefia); 3 females, Lago Icalma, Cordillera de Lonquimay, 15-L1962 (no collector in- dicated); 1 male and 1 female, Lago Icalma, Cordillera Lonquimay, 7- 1-1962 (both parasitized by Strepsiptera) (no collector indicated); 3 males, Icalma, 11-1-1979 (O. Martinez, E. Balart, L. Ruz); 1 male and 1 female, Icalma, 11-1-1979 (H. Toro); 5 fe- males, Malleco, Lago Icalma, 1-1962 (E. Reed). The paratypes are deposited in: AMNH, KU, MLP, MNBR, MNHN, FAUCH, UCV, and HT. Comments: Among the paratypes parasitized by Strepsiptera, the female has retained normal characters. How- ever, one male shows some features as if it were female, that is, reduced yellow marks on the face (wider than in female) and legs, broad metasoma without transverse yellow bands, smaller head, and short flagellum and not enlarged distally. The two other parasitized males, from approximately the same locality, show similar femi- nization, and also a narrow yellow band on the supraclypeal area. Etymology: The specific name is a Latin adjective meaning bearded, re- ferring to the dense and long pilosity on the latero-distal area of clypeus. Liphanthus (Liphanthus) chillanensis new species Figs. 21-25; Map I, F Diagnosis: Near to L. sabulosus but males smaller, the last flagellar seg- ment with almost straight lateral mar- gins and facial fovea about three times as long as broad; females with clypeus less than three times as broad as long, ocello-occipital distance similar to ocel- lar diameter, much shorter than in L. sabulosus. It differs from other species by the characters given in the key. Male: Length about 4.5 mm, fore- wing 2.9 mm, head width 1.5 mm, thoracic width 1.2 mm. Coloration. Head and thorax black, the following parts yellow: face below median ocellus and facial fovea (except most of lateral area of clypeus which is almost black), labrum (proximal area) (brown dis- tally), mandible (apex mahogany), un- der side of flagellum, pronotal lobe, spot on tegula, apices of femora, tibiae and tarsi (except distitarsi mostly brown and a brown spot on internal face of hind tibia and basitarsus). Wings slightly dusky, veins brown. Tegula testaceous. Metasomal terga black, terga I and II each with trans- verse yellow band (wider on II). Meta- somal sterna dark brown. Pubescence. In general short and sparse. Labrum pubescent on distal margin. Lateral areas of clypeus scarcely pubescent, with hairs shorter than those of la- brum. Integument and Punctation. Head and thorax strongly but finely areolate between punctures, yellow parts of face and most of gena smooth and shiny. Punctation, in general fine and sparse The Bee Genus Liphanthus 253 on head and thorax, very fine and sparse on hypoepimeral area, deeper and denser on rest of mesepisternum. Structure. Head about 1.5 times as broad as long. Flagellar segments much longer than broad. Facial fovea about half as long as scape, almost straight. Inner subantennal suture with upper part arcuate toward mid-line of face, shorter than width of subantennal area. Clypeus broader than long, pro- jecting beyond lower orbital tangent for somewhat less of half of its length, convex with median longitudinal pro- tuberance, labrum fiat, somewhat broader than long, distal margin con- vex. Mandible unmodified. Marginal cell shorter than distance from apex to wing tip. Second submarginal cell with anterior margin distinctly shorter than posterior. Legs and sterna unmodified. Genitalia and associated sterna as illus- trated. Female: Length about 5.5 mm, fore- wing length 3.4 mm, head width 1.5 mm, thoracic width 1.2 mm. Coloration. Head and thorax black, flagellum light brown beneath, the following parts yellow: area lateral to outer subanten- nal suture extending slightly below tentorial pit, subantennal area, median longitudinal spot on clypeus, mandible (apex mahogany), pronotal lobe, spot on tegula, small spots on apices of fore and mid femora and bases of fore and mid tibiae (spot on ventral edge of tibiae I), external spot on basitarsi I and II (small segments of tarsi brown). Wings and tegula as described for male. Metasomal segments usually dark brown, sometimes reddish, pygid- ial plate mahogany. Pubescence. As de- scribed for male but longer and denser on head, mesepisternum and legs; tibial scopa somewhat densely pubes- cent but less dense than hairs of fore femur. Integument and Punctation. As de- scribed for male but densely and coarsely punctate on face, with smooth intervals between punctures below fa- cial fovea. Pygidial plate arcolate (or smooth) apically. Structure. As de- scribed for male except: head 1.3 times as broad as long, facial fovea half as long as scape, straight. Clypeus almost three times as broad as long. Lower part of frontal line unmodified except narrowly grooved apex. Pygidial plate with a narrow, median ridge apically. Type Material: Holotype male and al- lotype female. Nuble Prov., Chilian, Las Trancas, Chile, 11-1977 (P. Toro), in Toro collection. Paratypes. (All from Nuble Prov., Chile, except as other- wise indicated.) 3 females, Linares Prov., Castillo, 11-1976 (H. Toro); 2 males and 1 female, Chilian, Trancas, 11-1977 (P. Toro); 4 males, Chilian, Trancas, 11-1977 (H. Toro); 2 males, Termas de Chilian, 11-1976 (H. Toro); 1 male, Termas de Chilian, 15-III- 1977 (E. Tosti); 2 males, Termas de Chilian (H. Toro); 1 male and 3 females, Termas de Chilian, 28-1-1967 (Schlinger). The paratypes are depos- ited in: KU, AMNH, CAS, UCV, and HT. Etymology: The specific name refers to the area in which most of the speci- mens were collected. PSEUDOLIPHANTHUS NEW SUBGENUS Figs. 26-50 Type species: Liphanthus rozeni new spe- cies. Both sexes: Vertex seen in frontal view convex. Lateral ocelli at about same level as or slightly below upper orbital tangent. Interalveolar area gen- erally slightly protuberant and similar to supraclypeal area. Frontal line weak (except minutely grooved apex in male, sometimes carinate). Clypeus convex, lateral area sloping upward toward central area (widened laterally 254 The University of Kansas Science Bulletin in ventral view in male). Inner orbits not emarginate, divergent below. Dis- tance between alveoli less than distance from alveolus to inner orbit, about two or more times alveolar diameter. Outer subantennal suture strongly arcuate laterally as in Figs. 26, 32, and 38 (less arcuate in L. unifasciatus). Pronotum without protuberant dorsolateral an- gle. Submarginal cells three. Propodeal triangle laterally more than half length of metanotum (laterally). Tibial spurs yellow or pale testaceous. Hind tibial spurs with apices slightly curved, inner slightly longer than outer. Male: Head and legs with yellow marks, metasoma with one or two transverse yellow bands. Lateral area of clypeus with hairs sparse and shorter than clypeal length. Flagellum much longer than head, first flagellar seg- ment approximately three times as long as broad. Facial fovea linear, shorter than scape, distance from its external margin to inner orbit similar to its width. Middle of paraocular area con- vex beside inner orbit, depressed to- ward alveolus; lower part of paraocular area sloping upward toward central area of face. Postmandibular area mes- ally much less than half as long as width of base of mandible. Sternum II produced into a spine (except in L. unifasciatus). Pygidial plate absent. Female: Lower paraocular area and tcgula without yellow. Tibial scopa about as dense as hairs of fore femur. Third flagellar segment broader than long. Etymology: The subgeneric name is based on pseudo-idihc, plus the generic name, to indicate that the species of this subgenus look like those of the subgenus Liphanthus s. str. Liphanthus (Pseudoliphanthus) rozeni new species Figs. 26-31; Map I, E Diganosis: Similar to L. sabulosus but male with flagellar segments 2-10 a little longer than broad; metasomal sternum II with distal projection and sternum III with premarginal pro- tuberance; female with upper half of face strongly areolate, dull and with punctures almost invisible. Male: Length about 5 mm, fore- wing length 3.2 mm, head width 1.8 mm, thoracic width 1.4 mm. Coloration. Head and thorax black, flagellum mostly testaceous beneath, the follow- ing parts yellow: face below facial fovea as in Fig. 26, labrum, mandible (apex mahogany), pronotal lobe, small basal spot on tegula, apices of femora, most of tibiae and basitarsi, small segments of fore and middle tarsi (brown on hind tarsus). Wings slightly testaceous, veins brown. Tegula testaceous. Meta- somal terga black, I and II each with transverse yellow band (longer and wider on II). Metasomal sterna brown. Pubescence. In general short and sparse. Labrum pubescent on distal fifth. Lat- eral area of clypeus scarcely pubescent, with hairs slightly longer than those of labrum. Integument and Punctali on. Head and thorax strongly areolate, very dull on ocellar area, smooth and shiny on ventral paraocular area, clypeus and gena, with some striae in propodeal triangle. Metasoma smooth. Puncta- tion in general shallow and fine, deeper on gena, coarser on clypeus, sparse on hypoepimeral area, well marked on rest of mesepisternum. Structure. Head 1.4 times as broad as long. Flagellar segments longer than broad, especially first and last one distinctly so. Facial fovea a little more than half as long as scape, convergent above. Inner suban- tennal suture slightly arcuate toward mid-line of face and distinctly shorter than width of subantennal area. Clyp- eus somewhat more than 4 times as broad as long, projecting beyond lower orbital tangent somewhat more than The Bee Genus Liphantmus 255 half of its length, with median pro- tuberance. Labrum flat, somewhat broader than long, distal margin broadly emarginate in the middle. Mandible with preapical widened plate (similar to that in Fig. 154). Marginal cell somewhat shorter than distance from apex to wing tip. Second sub- marginal cell with anterior margin about as long as posterior. Legs un- modified. Metasomal sternum II with postero-median projection much broader than long and hairy at apex (Fig. 30). Sternum III with premargi- nal protuberance on the midline; ster- num IV slightly protuberant on the middle. Genitalia and associated sterna as illustrated. Female: Length about 5.6 mm, fore- wing length 3.2 mm, head width 1.6 mm, thoracic width 1.4 mm. Coloration. Head and thorax brown, the following parts yellow: spot on subantennal area, T-shaped spot on basal part of clypeus, mandible (apex mahogany), pronotal lobe, very small and weak spot on tegula, small spot on apices of fore and mid femora and bases of fore and mid tabiae (extending to external face on tibia I). Wings and tegula as described for male. Metasomal terga I-IV mostly yellowish (the rest testaceous), or in some specimens I and II each with distinct transverse yellow band (some- times also on III); III and IV usually light testaceous (though somewhat var- iable in extent), remaining terga light brown (or darker). Pygidial plate ma- hogany. Metasomal sterna testaceous or brown. Pubescence. As described for male; tibial scopa somewhat densely pubescent, as dense as hairs of fore femur. Integument and Punctation. Integ- ument in general areolate, but strongly areolate between punctures on upper half of face (mostly dull from alveolar level to vertex) and thorax, smooth on gena and metasomal terga; metasomal sterna almost smooth. Punctation as described for male but much denser and deeper on clypeus. Structure. As described for male except: head 1.3 times as broad as long, facial fovea somewhat more than half as long as scape and extremely narrowed on up- per part. Clypeus depressed distally, somewhat more than three times as broad as long. Marginal cell about as long as distance from its apex to wing tip. Second submarginal cell with ante- rior margin shorter than posterior. Pygidial plate with median, longitudi- nal ridge on apex. Type Material: Holotype male and al- lotype female. Talca Prov., Alto de Vil- ches, Chile, 15-XI-1969 (Gonzalez), m Toro collection. Paratypes. 3 females, Talca Prov., Alto de Vilches, Chile, 30-X-1969 Q. G. Rozen, L. Pefia). Paratypes are deposited in AMNH col- lection. Etymology: This species is dedicated to Dr. Jerome G. Rozen, Jr., who has contributed greatly to the knowledge of the biology of Chilean insects and kindly helped us in several ways during this study. Liphanthus (Pseudoliphanthus) spiniventris new species Figs. 32-37; Map I, G Diagnosis: Similar to L. sabulosus but males with outer subantennal suture strongly arcuate and metasomal ster- num II with a distal and median spine- like projection. See also the characters given in the key. Male: Length about 5 mm, fore- wing length 3.2 mm, head width 1.8 mm, thoracic width 1 .4 mm. Coloration. Head and thorax black, the following parts yellow: face below level of facial fovea, labrum, mandible (apex ma- hogany), most of flagellum, pronotal lobe, small spot on tegula, apices of femora (continued along ventral edge 256 The University of Kansas Science Bulletin Figs. 26-31. Liphanthus (Pseudoliphanthns) rozeni n. sp., male: 26, head I'rontal view; 27, labrum; 28, eighth metasomal sternum; 29, dorsal and ventral views of genitalia; 30, ventral view ofmctasoma; 31, seventh metasomal sternutn. Figs. 32-37. Liphanlhus (Pseudoliphanthus) spiniventris n. sp., male: 32, head frontal view; 33, labrum; 34, eighth metasomal sternum; 35, ventral view of'metasoma; 36, seventh metasomal sternum; 37, dorsal and \entral views of genitalia. of femur I), tibiae and basitarsi (small segments ol mid and hind tarsi brown, small segments of fore basitarsus not observed). Wings slightly testaceous, veins and tegula dark testaceous. Met- asomal terga black, 1 and II each with transverse yellow band (longer and wider on II). Metasomal sterna brown. Pubescence. In general short and sparse. Labrum pubescent on distal fifth. Lat- eral area of clypeus scarcely pubescent, with hairs nearly as long as those of labrum. Integument and Punctation. Body in general strongly areolate, very dull between punctures in ocellar area, smooth and shiny on lower paraocular The Bee Genus Liphanthus 257 area close to inner orbit and lower gena, with some striation on propodeal triangle. Punctation in general sparse and shallow, denser in orbito-ocellar area and gena, very line on hypo- epimeral area, denser and well marked on rest of mesepisternum. Structure. Head 1.5 times as broad as long. Flagellar segments much longer than broad. Facial foveae somewhat less than half as long as scape, convergent above. Inner subantennal suture with upper part angulate and distinctly shorter than width of subantennal area. Clypeus 4 times as broad as long, projecting beyond lower orbital tan- gent for somewhat less than half of its length, with basal and median pro- tuberance. Labrum flat, a little broader than long, distal margin slightly exca- vated in middle. Mandible arcuate. Marginal cell shorter than distance from apex to wing tip. Second sub- marginal cell with anterior margin dis- tinctly shorter than posterior. Mid basitarsus a little less than 3 times as long as broad. Metasomal sternum II with median spiniform projection on posterior margin (Fig. 35). Genitalia and associated sterna as illustrated. Type Material: Holotype male. Nuble Prov., Chilian, Las Cabras, Chile, XII-1954 (L. Peha), in Toro collec- tion. Etymology: The species name refers to the acute spine on metasomal ster- num II. Liphanthus (Pseudoliphanthus) andinus new species Figs. 38-43; Map I, L Diagnosis: Closely related to /.. spin- iventris but male with short and trun- cate projection on posterior margin of metasomal sternum II and flagellum somewhat longer than width of head; female recognizable by characters given in key. Male: Length about 4.4 mm, fore- wing length 3 mm, head width 1.8 mm, thoracic width 1 .6 mm. Coloration. Head and thorax black, most of flagel- lum brown, the following parts yellow: face below level of facial foveae, la- brum (brown in center), mandible (apex mahogany), median part of un- der side of flagellum, pronotal lobe, very small spot sometimes on tegula (right side only in type), apices of femora (and ventral edge of femur I), tibiae and tarsi (except tibia and basitarsus III which are testaceous and small segments of tarsi I-III which are brown). Wings light testaceous, veins brown. Tegula testaceous. Metasomal terga black, terga I and II each with transverse yellow band (longer and wider on II). Metasomal sterna brown. Pubescence. In general short and sparse. Labrum pubescent on distal margin. Lateral area of clypeus scarcely pubes- cent, hairs longer than those of labrum. Integument and Punctation. Integument in general strongly areolate, very dull be- tween punctures on ocellar area, more smooth and shiny on paraocular area from below facial fovea to its internal lobe beside clypeus, clypeus, and lower half of gena; with some striation on propodeal triangle. Punctation in gen- eral weak, fine and sparse, coarser on clypeus and denser on upper half of gena, very fine and sparse on hypo- epimeral area, rest of mesepisternum with punctures somewhat bigger but weak. Structure. Head 1.5 times as broad as long. Flagellar segments much longer than broad. Facial fovea about half as long as scape, with upper part obliquely directed toward mid-line of face. Inner subantennal suture slightly arcuate toward mid-line of face and much shorter than width of suban- tennal area. Clypeus more than 4 times as broad as long, projecting beyond lower orbital tangent for about one- 258 The University of Kansas Science Bulletin third of its length, with a protuberance at base of central area. Labrum flat, somewhat broader than long, distal margin convex with median emargina- tion. Mandible, in frontal view, with distal fourth curved (curvature varies from slight to strong). Marginal cell shorter than distance from apex to wing tip. Second submarginal cell with anterior margin slightly shorter than posterior. Mid basitarsus somewhat less than 3 times as long as broad. Metasomal sternum II with postero- median projection about as long as apical width, apically truncate and densely hairy (Fig. 40). Genitalia and associated sterna as illustrated. Female: Length about 4.6 mm, fore- wing length 3 mm, head width 1.4 mm, thoracic width 1.4 mm. Coloration. Head and thorax black, the following parts yellow: T-shaped spot on upper part of clypeus, very small and weak spot on base of mandible (apex dark mahogany), spot on pronotal lobe, spot on tegula almost invisible (or absent), small spot on bases of fore and mid tibiae (prolonged to most of anterior border of tibia I). Wings lightly testa- ceous, veins and tegula brown. Meta- soma black; pygidial plate sometimes mahogany. Pubescence. As described for male; tibial scopa as dense as hairs of fore femur. Integument and Punctation. In general finely and slightly areolate and shiny, a little more strongly areolate between punctures on upper part of face and thorax, somewhat opaque around ocelli. Pygidial phite smooth apically. Punctation as described for male but denser and deeper on face, absent around ocelli, coarser on clyp- eus, sparse on hypoepimeral area. Structure. As described for male except: head 1.4 times as broad as long, facial fovea straight, parallel to inner orbit, clypeus somewhat less than 3 times as broad as long, not protuberant. Type Material: Holotype male and al- lotype female. Malleco Prov., Cordillera, Nahuelbuta, Chile, IX-1980 (H. Toro), in Toro collection. Paratypes. 3 males and 1 female, Malleco Prov., Cordillera Nahuelbuta, Chile, IX-1980 (H. Donoso, H. Toro, H. Burgos); 1 male, Neuquen Prov., Lago Lacar, 650 m, Argentina, 15-1-1953 (Sen- kute). Paratypes are deposited in AMNH, KU, and UCV. Etymology: The species name indi- cates the mountainous area where most of the specimens were collected. Liphanthus (Pseudoliphanthus) unifasciatus new species Figs. 44-50; Map I, C Diagnosis: Similar to L. sabulosus but male with frontal line carinate distally, face black from below alveolar level to vertex and metasoma with only one yellow transverse band. See also char- acters given in the key. Male: Length about 4.6 mm, fore- wing length 3.4 mm, head width 1.7 mm, thoracic width 1.4 mm. Coloration. Head and thorax black, the following parts yellow: lower part of paraocular area, clypeus, mandible (apex ma- hogany), labrum (darkened in the mid- dle), pronotal lobe, weak spot on apices of femora, external surfaces of tibiae and basitarsi, hind tibia with dark area in middle; small segments of tarsi brown. Wings lightly testaceous, veins brown. Tegula testaceous. Metasomal terga black, with short and narrow transverse yellow band on II. Metaso- mal sterna dark brown. Pubescence. In general short and sparse. Labrum pubescent on distal margin. Lateral area of clypeus scarcely pubescent, hairs as long as those of labrum. Integu- ment and Punctation. Head and thorax strongly areolate, lightly areolate be- tween punctures on metasomal terga, smooth on metasomal sterna, with The Bee Genus Liptianthus 259 Figs. 38-43. Liphanthus (Pseudoliphanlhui) andinus n. sp., male: 38, head frontal view; 39, labrum; 40, ventral vieu' of metasoma; 41, seventh metasomal sternum; 42, dorsal and ventral views of genitalia; 43, eighth mctasomal sternum. Figs. 44-50. Liphanthus (Pseudoliphanthus) unijascialus n. sp,, male; 44, 45, 46, head frontal, ventral and lateral views; 47, labrum; 48, dorsal and ventral views of genitalia; 49, 50, seventh and eighth metasomal sterna. some striae on propodeal triangle. Punctation in general sparse and weakly marked, almost undistinguisha- ble on face, coarser on clypeus, ex- tremely fine on hypoepimeral area, somewhat coarse but weak on rest of mesepisternum. Structure. Head about 1.3 times as broad as long. Flagellar segments distinctly longer than broad. Facial fovea somewhat less than half as long as scape, narrow and straight. Inner subantennal suture slightly arcu- ate toward outer part of face. Clypeus more than 3 times as broad as long, projecting beyond lower orbital tan- gent for less than half its length. La- brum flat, less than twice as broad as long, distal margin slightly convex with 260 The University of Kansas Science Bulletin median shallow emargination. Mandi- ble widened forming preapical plate (similar to Fig. 154). Marginal cell shorter than distance from apex to wing tip. Second submarginal cell with anterior margin distinctly shorter than posterior. Legs and sterna unmodified. Genitalia and associated sterna as illus- trated. Type Material: Holotype male. Val- paraiso Prov., Valparaiso, Chile, 12- X-1971 (H. W. Sielfeld), in Toro col- lection. Paratypes (all from Chile). 3 males, Valparaiso Prov., Valparaiso, 12-X-1971 and 1 male, Valparaiso, 27 VIII-1972 (H. W. Sielfeld); 3 males without information about date, lo- cality, and collector; 4 males, Val- parai'so, IX-1920 (P. Herbst) and 4 males, Valparaiso Prov., Marga- Marga, from September to December, 1920 (P. Herbst). Paratypes deposited in: MCZ, CAS, and UCV. Etymology: The specific name indi- cates that there is only one metasomal yellow band. Xenoliphanthus new subgenus Figs. 51-73 Tvpe species: Psaenythia parvula Friese, 1916. Both sexes: Vertex generally concave except almost straight in L. michenen and L. moldenkei. Lateral ocelli at same level as upper orbital tangent. Inter- alveolar area usually convex, similar to supraclypeal area. Clypeus flat with longitudinal median groove; lateral area slightly sloping upward toward central area, widened mesally in ven- tral view. Inner orbits straight and parallel medially, divergent ventrally. Distance between alveoli similar to dis- tance from alveolus to inner orbit and about twice as long as alveolar diame- ter. Outer subantennal suture arcuate laterally. Pronotum without protuber- ant dorsolateral angles. Submarginal cells three. Propodeal triangle laterally about as long as metanotum (laterally) or slightly shorter. Tibial spurs testa- ceous or brown. Hind tibial spurs with apices strongly curved, inner and outer subequal in length. Male. Head and legs with yellow marks, metasoma red to black, without yellow bands. Lateral area of clypeus with hairs sparse and shorter than clypeal length. Frontal line in shallow concavity basally. Flagellum a little longer than head; first flagellar segment 1.5 to 2.5 times as long as broad. Facial fovea linear, distance from it to inner orbit longer than its width. Middle of paraocular area convex beside inner orbit, slightly concave toward alveolus; lower part of paraocular area almost flat toward mid-line of face. Postmandibular area mesally much less than half as long as width of base of mandible. Sterna un- modified except male of L. micheneri has sternum II prominent mesally. Pygid- ial plate present. Female. Lower para- ocular area without yellow spot. Tibial scopa less dense than fore femoral hairs. Third flagellar segment broader than long. Etymology: The subgeneric name is from the Greek xenos meaning stranger plus Liphanthus. Liphanthus (Xenoliphanthus) parvulus (Friese) new combination Figs. 51-58; Map II, E Psaenythia parvula Friese, 1916: 166; Herbst, 1922: 183; Jaffuel and Pi- rion, 1926: 369; Rozen, 1970: 146-147; Ehrenfeld and Rozen, 1977: 3. Psaenythia pygmea Friese, 1916: 166; Herbst, 1922: 183. Psaenythiafriesei Herbst, 1922: 183 (new synonymy). Male: Length about 5 mm, fore- wing length 4 mm, head width 1.7 mm, thoracic width 1.4 mm. Coloration. The Bee Genus Liphanthus 261 Head and thorax black except flagel- liini mostly testaceous beneath, the fol- lowing parts yellow: clypeus, lower part of paraocular area extending up beyond alveolar level, subantennal area, supraclypeal area, labrum (some- times black or testaceous distally), mandible (apex mahogany), spot (sometimes absent) or under side of scape, pronotal lobe, spot on tegula, apices of femora, tibiae and most of basitarsi (small segments of tarsi brown). Wings slightly dark brown, \eins brown, tegula testaceous. Meta- soma red, black or basal and apical segments black, central segments red- dish. Pubescence. In general short and sparse. Labrum pubescent on distal fourth. Lateral area of clypeus scarcely pubescent, with hairs shorter than those of labrum. Integument and Puncta- twn. Integument in general smooth and shiny between punctures, except strongly areolate on upper half of face, slightly areolate on hypoepimeral area and metasomal sterna, scarcely striate on propodeal triangle. Punctation in general well marked, fine and dense on face, but coarse and sparse on lower part of paraocular area, clypeus and between inner orbit and lateral ocellus, fine but sparse on thorax, sparse on hypoepimeral area, coarser and some- what denser on rest of mesepisternum. Structure. Head 1.6 times as broad as long. Flagellar segments somewhat longer than broad, first one more con- spicuously longer. Frontal line carinate distally. Facial fovea elongate, straight, more than half as long as scape. Inner subantennal suture arcuate toward mid-line of face and slightly shorter than width of subantennal area. Inter- alveolar area protuberant. Clypeus somewhat less than 4 times as broad as long, projecting beyond lower orbital tangent for one-fourth of its length. Labrum strongly convex, more than twice as broad as long, distal margin with median, shallow emargination. Mandible unmodified. Marginal cell a little longer than distance from apex to wing tip. Second submarginal cell with anterior margin somewhat shorter than posterior. Legs and metasomal sterna unmodified. Genitalia and associated sterna as illustrated. Female: Length about 6 mm, fore- wing length 4.1 mm, head width 2.2 mm, thoracic width 1 .7 mm. Coloration. Head and thorax black, llagellum tes- taceous beneath, mandible mosdy ma- hogany, the following parts yellow: small spot on pronotal lobe, apices of fore and mid femora, and bases of fore and mid tibiae. Wings and tegula as described for male, veins darker. Met- asoma red-orange (sometimes with dark areas especially on last segments varying in extent), except lateral fovea on tergum II black and pygidial plate mahogany or black. Pubescence. As de- scribed for male, but tibial scopa less dense than hairs of fore femur. Integu- ment and Punctation. Integument as de- scribed for male, but much smoother on face and mesepisternum. Hypo- epimeral area scarcely punctate. Pygid- ial plate smooth or areolate. Structure. As described for male but facial fovea longer than scape and claws apparently simple. Pygidial plate with longitudinal ridge on apex. Material Studied (all from Chile): 3 females, Aconcagua Prov., Rio Blanco, XII-1970 (M. Pino); 180 males and 61 females, Valparaiso Prov., diverse localities and collectors from September to January; 1 male, Santiago Prov., Santo Domingo, X- 1966 (RamiVez). Several specimens from Valparaiso have been collected on Adesmia sp. Comments: Herbst (1922) recog- nized the synonymy of pygmaea and 262 The University of Kansas Science Bulletin parvula but unnecessarily erected a new U3imc, friesei, for the species. Tiie type of Psaenythia parvula Friese, 1916, has apparently been lost. In the Berlin Museum, according to the information sent to Ruz by Dr. I. Wegener, there is only one female la- beled as P. parvula Fr. It is from Val- paraiso, Chile, X-1900 (P. Herbst), Friese determination, 1915; there is no type label on it. Since the authors have not seen the specimen from Berlin, a female from Valparaiso, Laguna Verde, Chile, 1-11-1978 (L. Ruz) is here designated as the neotype, which will be deposited at the Universidad Catolica de Valparaiso, Valparaiso, Chile. Figs. 51-58. Liphanlhus (Xenoliphanlhus) parvulus (Friese), male: 51, 52, head frontal and lateral views; 53, forewing and hindwing; 54, labrum; 55, dorsal and ventral views of genitalia; 56, hind tibial spurs; 57, 58, seventh and eighth metasomal sterna. Figs. 59-63. Liphanlhus (Xenoliphanlhus) lojensis n. sp., male: 59, head frontal view; 60, labrum; 61, 62, seventh and eighth metasomal sterna; 63, dorsal and ventral views of genitalia. The Bee Genus Liphanthus 263 Liphanthus (Xenoliphanthus) tofensis new species Figs. 59-63; Map II, C Diagnosis: Similar to L. parvidus but marginal cell distinctly longer than dis- tance from apex to wing tip and hypo- epimeral area with anterior part well punctate; female with frontal line end- ing in narrow groove apically instead of carina. Male: Length about 5.4 mm, fore- wing length 3.5 mm, head width 1.6 mm, thoracic width 1.4 mm. Coloration. Head and thorax black, flagellum brown with under side mostly testa- ceous, the following parts yellow: clypeus, lower part of paraocular area extending near inner orbit up beyond alveolar level, subantennal area, su- praclypeal area (sometimes black), la- brum (testaceous distally), mandible (apex mahogany), spot (sometimes ab- sent) on under side of scape, pronotal lobe, spot on tegula, apices of femora, tibiae and most of basitarsi (small seg- ments of tarsi dark-brown). Wings slightly dark brown, veins brown, tegula testaceous. Metasomal segments reddish, tergum I and most of 2 or 3 last segments dark (brown or black areas varying in extent). Pubescence. In general short and sparse. Labrum pubescent on distal fourth. Lateral area of clypeus scarcely pubescent, with hairs about as long as those of labrum. Integument and Punctation. In general smooth and shiny between punctures, face with upper half strongly areolate and mesepisternum slightly areolate and dull, propodeal triangle scarcely striate. Punctation in general well marked, rather fme and dense on face, but coarse and sparse on lower part of paraocular area, clypeus and between inner orbit and ocellus. Thorax rather coarsely punctate, punctures dense on anterior part of hypoepimeral area and very dense on rest of mesepisternum. Structure. Head 1.3 limes as broad as long. Flagellar segments about as long as broad, first and last longer. Frontal line a small groove apically, not cari- nated. Facial fovea straight, narrow, slightly shorter than half of scape. In- ner subantennal suture arcuate toward midline of face and as long as width of subantennal area. Interalveolar area slightly prominent. Clypeus almost 4 times as broad as long, projecting be- yond lower orbital tangent for about half its length. Labrum slightly convex, somewhat more than twice as broad as long, distiil margin almost straight. Mandible unmodified. Marginal cell distinctly longer than distance from apex to wing tip. Second submarginal cell with anterior margin shorter than posterior. Legs and sterna unmodified. Genitalia and associated sterna as illus- trated. Female: Length about 6 mm, fore- wing length 4.5 mm, head width 2.2 mm, thoracic width 2.1 mm. Coloration. Black, flagellum testaceous beneath, brown above, the following parts yel- low: small spot on pronotal lobe, spot on tegula, small spots on apices of fore and mid femora and bases of fore and mid tibiae (hind tibia and tarsi I-III brown). Wings as described for male. Metasoma red-orange, lateral fovea on tergum II black (sometimes with dark- ened areas on base of tergum I), pygidial plate mahogany. Pubescence. As described for male, tibial scopa less dense than hairs of fore lemur. Integu- ment and Punctation. As described for male but pygidial plate smooth. Struc- ture. As described for male except head 1.5 times as broad as long, facial fovea somewhat narrower dorsally and al- most as long as scape. Clypeus project- ing beyond lower orbital tangent for less than half of its length. Claws ap- parently simple. Pygidial plate with no ridge apically. 264 The University of Kansas Science Bulletin Type Material: Holotype male and al- lotype female. Coquimbo Prov., El Tofo, Chile, X-1971 (H. Toro), in Toro col- lection. Paratypes (all from Coquimbo Prov., Chile). 2 males and 9 females. El Tofo, X-1971-72 (H. Toro); 1 female. El Pangue, X-1972 (H. Toro); 2 males and 2 females, Llano de la Higuera, N. ofTofo, 15-X-1971 Q. G. Rozen, L. Pefia). Paratypes are de- posited in the following collections: AMNH, KU, MNHN, and UCV. Etymology: The name of the species is based on the type locality. El Tofo. Liphanthus (Xenoliphanthus) moldenkei new species Figs. 64-68; Map II, H Diagnosis: Closely related to L. par- vulus, but vertex almost flat. Male with clypeus little more than 3 times as broad as long, with large black spot on each side basally. Female with claws cleft, inner ramus well developed. Male: Length about 5.2 mm, fore- wing length 3.5 mm, head width 1.7 mm, thoracic width 1.5 mm. Coloration. Head and thorax black, flagellum tes- taceous beneath, labrum testaceous distally, the following parts yellow: clypeus (with black spot on each side basally), lower part of paraocular area extending nearly to inner orbit up to alveolar level, subantennal area (in- completely), labrum (except distal area), mandible (apex mahogany), small spot on under side of scape, pronotal lobe, small spot on tegula, apices of femora, tibiae and most of basitarsi (small segments of tarsi brown). Wings dark brown, veins mostly brown, tegula testaceous. Meta- soma reddish, first two terga (especially lateral fovea of second), distal margin of fifth and pygidial plate darkened. Pubescence. In general short and sparse. Labrum pubescent on distal fourth. Lateral area of clypeus with hairs mod- erately long and sparse, about as long as those of labrum. Integument and Punc- tation. In general smooth and shiny between punctures, areolate on upper half of face and slightly so on meso- notum, hypoepimeral area and sterna; propodeal triangle scarcely striate. Punctation in general well marked, coarse and sparse on lower part of paraocular area, clypeus and orbito- ocellar area. Thorax with punctures similar to those on upper face, moder- ately dense on scutal area, fine and dense on hypoepimeral area coarser, deeper on rest of mesepisternum. Struc- ture. Head 1.4 times as broad as long. Flagellar segments almost as long as broad, first and last ones clearly longer. Frontal line evanescent distally. Facial fovea slightly sinuous, about as long as scape. Inner subantennal suture slightly arcuate toward mid line of face and about as long as width of subanten- nal area. Interalveolar area slightly prominent. Clypeus little more than 3 times as broad as long, projecting be- yond lower orbital tangent for some- what less than half of its length. Labrum convex, about twice as broad as long, distal margin slightly convex. Mandible unmodified. Marginal cell clearly longer than distance from apex to wing tip. Second submarginal cell with anterior margin somewhat shorter than posterior. Legs and sterna un- modified. Genitalia and associated sterna as illustrated. Female: Length about 6.2 mm, fore- wing length 4.4 mm, head width 2 mm, thoracic width 1.8 mm. Coloration. Head and thorax black, the following parts yellow: inost of under side of antennal flagellum, mandible (weakly yellow, apex dark mahogany), pronotal lobe, small spot on tegula, small spots on apices of femora and bases of fore and mid tibiae (somewhat elongate on anterior edge of tibia I). Wing, veins The Bee Genus Liphanthus 265 and tegula as described for male. Met- asoma orange-reddish, lateral fovea on tergum II black, metasomal terga I and II with darkened areas and pygidial plate mahogany. Pubescence. As de- scribed for male, but tibial scopa less dense than hairs of fore femur. Integu- ment and Punctation. As described for male, but pygidial plate slightly areo- late and hypoepimeral area sparsely punctate. Structure. As described for male except about half of frontal line in rather wide groove apically, clypeus somewhat more than 3 times as broad as long and pygidial plate with longitu- dinal ridge apically. Type Material: Holotype male. Talca Prov., N. Laguna del Maule, 2,200 m, Chile, 16-1-1968 (Pena-Barros). Al- lotype female. Santiago Prov., La Ollita, Cantillana, 2,000 m, Chile, 1-8-XII- 1969 (L. Pena), both in Toro collec- tion. Comments: The association of sexes may not be correct since the male was collected at a different locality from that of the female. Etymology: This species is named for Dr. Andrew Moldenke in recognition of his contribution to the study of the Chilean bees. Liphanthus (Xenoliphanthus) micheneri new species Figs. 69-73; Map II, K Diagnosis: Near to L. parvulus but male with black spot on each side of clypeus dorsally and 2 median pro- tuberances on metasomal tergum II. Females with vertex flat and metasoma black. Male: Length about 5.2 mm, fore- wing length 3.7 mm, head width 1.8 mm, thoracic width 1.5 mm. Coloration. Black, antennal flagellum testaceous beneath, the following parts yellow: clypeus except black spot on each side (very reduced and weak in some speci- mens, absent in others), lower part of paraocular area (sometimes extending up beside inner orbit to level of al- veolus), part of subantennal area, small spot on supraclypeal area in some spec- imens, labrum (brown spot in the cen- ter in some specimens), mandible (apex mahogany), pronotal lobe, small spot on tegula, spots on apices of femora, tibiae and most of basitarsi (small segments of tarsi brown, except fore tarsal segments II-IV testaceous). Wings dark brown, Veins brown. Tegula testaceous. Pubescence. In gen- eral short and sparse. Labrum pubes- cent on distal margin. Lateral area of clypeus with hairs sparse, about as long as those of labrum. Integument and Punc- tation. In general almost smooth and shiny, except clearly areolate on dorsal half of face and thorax, coarsely and sparsely punctate in lower part of para- ocular area, clypeus and orbito-ocellar area, scarcely so on striate propodeal triangle. Punctures weakly marked on scutal area and scutellum, finer and rather dense on hypoepimeral area. Structure. Head 1.4 times as broad as long. Flagellar segments about as long as broad, first and last longer. Frontal line in weakly marked apical groove. Facial fovea slightly S-shaped or straight, somewhat shorter than scape. Inner subantennal suture almost straight and about as long as width of subantennal area. Interalveolar area slightly prominent. Clypeus a little more than 3 times as broad as long, projecting beyond lower orbital tan- gent for somewhat less than half of its length. Labrum flat, almost twice as broad as long, distal margin straight. Mandible unmodified. Marginal cell a little longer than distance from apex to wing tip. Second submarginal with an- terior margin somewhat shorter than posterior. Legs unmodified. Metaso- mal sternum II with 2 strong median 266 The University of Kansas Science Bulletin 70 Figs. 64-68. Liphanthus (Xenoliphajilhm) moldcnkei n. sp., malt": 64, head tVnntal view; 65, labium; 66, 67, eighth and seventh metasomal sterna; 68, dorsal and ventral views of genitalia. Figs. 69-73. Liphanthus (Xenoliphanlhus) tmchenen n. sp., male: 69, head frontal view; 70, labrurn; 71, dorsal and ventral views of genitalia; 72, 73, seventh and eighth metasomal sterna. protuberances side by side. Genitalia and associated sterna as illustrated. Female: Length about 6 mm, fore- wing length 3.8 mm, head width 1.9 mm, thoracic width 1.8 mm. Coloration. Black, llagellum mostly testaceous be- neath, brown above, the following parts yellow: weak spot on pronotal lobe, small spot on bases of fore and mid tibiae. Wings and tegula as de- scribed for male. Pygidial plate ma- hogany, with ridge black. Pubescence. As described for male but tibial scopa less dense than hairs of fore femur. Integu- ment and Punctation. As described for male but pygidial plate almost smooth and punctures in general denser, es- pecially on face except clypeus. Meta- somal sterna coarsely punctate, punctures less deep and sparser on The Bee Genus Liphanthus 267 metasomal terga, especially sparse on two first terga. Structure. As described for male, but head 1.6 times as broad as long, clypeus projecting beyond lower orbital tangent for a little more than half of its length. Pygidial plate with longitudinal ridge apically. Type Material: Holotype male and al- lotype female. Nuble Prov., Chilian, Las Trancas, Chile, XII-1977 (L. Ruz), in Toro collection. Paratypes (all from Nuble Prov.), Chilian, Las Trancas, Chile, collected on 15-XII-1977). 60 males and 3 females (H. Toro), 73 males and 16 females (H. Flores), 52 males and 9 females (L. Ruz), 21 males and 1 female (E. Tosti-Croce), 44 males and 4 females (O Martinez), 34 males and 4 females (E. Chiappa), 12 males and 1 female (E. Peralta). Para- types are deposited in the follow- ing collections: AMNH, CAS, KU, BMNH, MNHUB, IMZT, MNHNP, MNHN, MCZ, CONC, UCV, and HT. Etymology: This species is named in honor of Dr. Charles D. Michener for all his suggestions in relation to this paper and his great contribution to the study of bees. Discussion: A few more specimens, 4 males, Arauco Prov., Nahuelbuta, Chile, 1-1979 (H. Toro, M. Cerda); 4 males and 2 females, Malleco Prov., Nahuelbuta, Chile, XI-1980 (diverse collectors) have also been examined. They are not paratypes because of the red metasoma of the females. This difference may indicate either intra- specific variation, or two closely related species. More material should be col- lected in the future. Tricholiphanthus new subgenus Figs. 74-89 Type species: Liphanthus leucostomus new species. Both sexes (based on males only): Vertex slightly convex. Lateral ocelli at about same level as upper orbital tan- gent. Interalveolar area generally pro- tuberant. Supraclypeal area variable. Clypeus usually slightly convex, with- out longitudinal median groove; lateral area slightly sloping upward to central area, widened mesally in ventral view. Inner orbits slightly emarginate in dor- sal half (more broadly in L. leuco- stomus), generally divergent above, except subparallel in L. tarsalis. Dis- tance between alveoli somewhat less than distance from alveolus to inner orbit and somewhat greater than diam- eter alveolus. Outer subantennal su- ture arcuate laterally. Pronotum with dorsolateral angle generally protuber- ant but less conspicuously so in L. pilifrons. Submarginal cells three. Pro- podeal triangle laterally almost as long as metanotum (laterally). Tibial spurs pale testaceous. Hind tibial spurs with apices slightly curved, inner slightly longer than outer. Male: Head and legs with yellow marks, metasoma completely black. Pubescence in general rather long and dense. Lateral area of clypeus with hairs as long as or longer than clypeal length. Frontal line weakly marked, generally without groove distally but L. pilifrons with a minute one. Flagellum much longer than head; first flagellar segment about 3 times as long as broad. Facial fovea narrow, arcuate laterally, slightly shorter than scape and very close to inner orbit (more separated in L. pilifrons dorsally). Mid- dle part of paraocular area flat, lower paraocular area slightly sloping toward center of face. Postmandibular area mesally much less than half as long as width of base of mandible. Sterna III and IV with shallow median emargina- tion on posterior margin (barely con- cave on L. pilifrons). Pygidial plate absent. Etymology: The prefix of the sub- 268 The University of Kansas Science Bulletin generic name is from the Greek trichos meaning hair, to indicate rather dense pilosity in the species here included. Liphanthus (Tricholiphanthus) leucostomus new species Figs. 74-79; Map I, Ch Diagnosis: Close to L. pilifrons but clypeus and lower paraocular area whitish and sterna III and IV with posterior margins clearly concave; in- teralveolar area depressed. Male: Length about 4.6 mm, fore- wing length 3.2 mm, head width 1.4 mm, thoracic width 1.2 mm. Coloration. Black, except flagellum mostly testa- ceous, the following parts yellowish white: clypeus, lower part of paraocu- lar area extending up to level of lower margin of alveolus, labrum, mandible (apex mahogany), apex of pronotal lobe, spot on tegula, apices of femora, tibiae (except a brown spot on III) and most of basitarsi (small segments of tarsi brown). Wings scarcely testa- ceous, veins and tegula testaceous. Pubescence. Labrum pubescent on distal fourth. Clypeus with hairs much longer than those of labrum. Integument and Punctation. Dorsal part of head and thorax clearly areolate between punc- tures; lower half of face smooth and shiny; propodeal triangle striate; meta- soma almost smooth. Punctation fine, shallow and sparse on upper head, discal area of scutum and scutellum, dense and fine in lower part of face and upper gena, very sparse and minute on hypoepimeral area, sparse and weak on metasomal sterna. Structure. Head 1.3 times as broad as long. Flagellar seg- ments much longer than broad, last one strongly curved. Inner subanten- nal suture almost straight, a little shorter than width of subantennal area. Interalveolar area fiattened. Supra- clypeal area with median protuber- ance. Clypeus about 3 times as broad as long, projecting beyond lower orbi- tal tangent for somewhat less than half of its length. Labrum little convex, broader than long, distal margin slighdy convex, almost straight. Man- dible unmodified. Pronotum with dor- solateral angle protuberant, acute. Marginal cell about as long as distance from apex to wing tip. Second sub- marginal cell with anterior margin slightly shorter than posterior. Legs unmodified. Metasomal sternum II with wide and truncate median projec- tion on posterior margin (Fig. 78). Sterna III and IV posteriorly with broad, shallow median emargination. Genitalia and associated sterna as illus- trated. Type Material: Holotype male. Santi- ago Prov., Cantillana, Chile, XII-1969 (L. Peiia), in Toro collection. Paratypes. 9 males, same data; 1 male Santiago Prov., Cantillana, La Ollita, XII-1969 (L. Peha). The paratypes are deposited in the following collections: AMNH, KU, MNHN, UCV, and HT. Etymology: The species name indi- cates the whitish color of clypeal area. Liphanthus (Tricholiphanthus) pilifrons new species Figs. 80-86; Map I, H Diagnosis: Close to L. leucostomus but lower paraocular area and distal band of clypeus yellow, and posterior emar- gination of metasomal sternum III fee- ble and narrower; posterior margin of sternum IV almost straight; interalveo- lar area prominent. Male: Length about 5.3 mm, fore- wing length 3.2 mm, head width 1.6 mm, thoracic width 1.3 mm. Coloration. Black, except flagellum mostly testa- ceous; the following parts yellow: para- ocular area below level of alveolus, distal area on clypeus, labrum, mandi- ble (apex mahogany), spot on scape distally, pronotal lobe, small spot on The Bee Genus Liphanthus 269 tegula, apices of femora, tibiae and most of basitarsi (small segments of tarsi brown). Wings slightly testaceous, veins and tegula brown. Metasomal sterna dark brown. Pubescence. Lab rum pubescent on distal fifth. Clypeus with hairs much longer than those of la- brum. Integument and Punctation. Head and thorax strongly areolate between punctures, almost smooth on gena and Fi(;s. 74-79. Liphanthus ('I'richoliphanthus) lemoslomus. n. sj)., iiialc: 74. head frontal virw; 7.'), labrwm: 76, eighth metasomal sternum; 77, dorsal and ventral views of genitalia; 78, ventral view of nictasoma; 79, seventh metasomal sternum. Figs. 80-86. Liphanthus (Trtckoliphanthus) piltfrom n. sp., male: 80, 81 , head Ironial and lateral views; 82, labrum; 83, dorsal and ventral views of genitalia; 84, 85, seventh and eighth metasomal sterna; 86, ventral view of metasoma. Fios. 87-89. Liphanthus (Tricholtphanthus) larsalis n. sp., male: 87, head frontal view; 88, labrum; 89, ventral view of metasoma. 270 The University of Kansas Science Bulletin metasoma, striate on propodeal tri- angle. Punctures in general scarcely visible, more distinct in yellow areas of face, on gena and mesepisternum, very dense on metasomal terga. Structure. Head about 1.3 times as broad as long. Flagellar segments distinctly longer than broad, last one strongly curved. Inner subantennal sutures almost straight, convergent above and slightly shorter than width of subantennal area. Interalveolar and supraclypeal areas prominent. Clypeus flattened, more than 3 times as broad as long, project- ing beyond lower orbital tangent for about half its length. Labrum convex, distal margin convex. Mandible un- modified. Pronotum with dorsolateral angle slightly protuberant. Marginal cell scarcely longer than distance from apex to wing tip. Second submarginal cell with anterior margin slightly shorter than posterior. Legs unmodi- fied. Metasomal sternum II with broad postero-median projection (Fig. 86). Posterior margin of sternum III with shallow median emargination, that of IV almost straight. Genitalia and asso- ciated sterna as illustrated. Type Material: Holotype male. Nuble Prov,, Termas de Chilian, Chile, 18- 1-1953 (L. Pena), in Toro collection. Paratype. 1 male, Nuble Prov., Las Cabras, Chile, 6-31-1-1953 (L. Pena), deposited at KU. Etymology: The species name is re- lated to the moderately dense pubes- cence on the face. Liphanthus (Tricholiphanthus) tarsalis new species Figs. 87-89; Map II, L Diagnosis: Closely related to L. leu- costomus but fore and mid tarsi with third segment more than twice as long as broad; upper part of clypeus flat- tened. Male: Length about 4.8 mm, fore- wing length 3.6 mm, head width 1.4 mm; thoracic width 1.2 mm. Coloration. Black, antennal flagellum mostly testa- ceous; the following parts pale yellow: lower area on clypeus, lower part of paraocular area (narrow and transverse band), labrum, mandible (apex ma- hogany), spot on apex of scape be- neath, pronotal lobe, spot on tegula, apices of femora, most of tibiae and tarsi (except distitarsi brown) of fore and mid legs, tibia III (except median dark area), first two segments of hind tarsus (rest of small segments of tarsi light brown). Wings almost hyaline. Tegula and veins dark testaceous. Pubescence. Labrum pubescent mesally on distal fifth. Clypeus with hairs much longer than those of labrum. Integument and Punctation. Head and thorax areo- late and dull between punctures but almost smooth and shiny on yellow parts of face and gena as well as on metasoma, striate on propodeal tri- angle. Punctures scarcely visible except more distinct on yellow areas of face and on metasoma. Structure. Head 1.2 times as broad as long. Flagellar seg- ments much longer than broad, last one strongly curved. Inner subanten- nal suture slightly arcuate toward mid- line of face and a little shorter than width of subantennal area. Interalveo- lar area distinctly prominent, su- praclypeal area slightly protuberant. Clypeus ilat, more than 3 times as broad as long, projecting beyond lower orbital tangent for about half of its length. Labrum slightly convex, broader than long, distal margin con- vex. Mandible unmodified. Pronotum with dorsolateral angle protuberant, acute. Marginal cell somewhat longer than distance from apex to wing tip. Second submarginal cell with anterior margin slightly shorter than posterior. Second segment of fore and mid tarsi narrow, about twice as long as broad The Bee Genus Liphanthus 271 and similar to third. Hind basitarsus broadest at base, second tarsal segment almost as long as broad. Metasomal sternum II with wide truncate projec- tion posteriorly (Fig. 89). Sterna III and IV each with broad median emar- gination on posterior margin. Geni- talia and associated sterna missing in only specimen. Type Material: Holotype male. Mal- leco Prov., Curacautin, Termas Tol- huaca, Chile, 20-M959 (L. Peha), in Toro collection. Etymology: The species name refers to the modified tarsi. Melaliphaxthl's new subgenus Figs. 90-101 Type species: Liphanthus atratus new species. Both sexes: Vertex convex. Lateral ocelli at same level as upper orbital tangent (male) or below (female). Interalveolar and supraclypeal areas protuberant. Frontal line weakly devel- oped. Clypeus flattened basally (male) or slightly convex (female). Lateral area of clypeus abruptly sloping up- ward toward central area, widened mesally in ventral view. Inner orbits broadly emarginate in dorsal half and divergent above in male, almost straight and subparallel in female. Dis- tance between alveoli similar to dis- tance from alveolus to inner orbit and greater than alveolar diameter. Suban- tennal sutures almost straight. Prono- tum without protuberant dorsolateral angle. Submarginal cells three. Pro- podeal triangle laterally more than half as long as metanotum laterally (slightly longer in female of L. atratus). Tibial spurs pale testaceous or brown. Hind tibial spurs with apices slightly curved, inner somewhat longer than outer. Male. Head and legs with yel- low marks, metasoma without yellow bands. Lateral area of clypeus with hairs sparse and shorter than clypeal length. Flagellum much longer than head; first flagellar segment about 3 times as long as broad. Facial fovea straight, length variable, fairly close to inner orbit. Middle of paraocular area convex, lower paraocular area abruptly sloping upward toward mid-line of face. Postmandibular area mesally half as long as width of base of mandible or more. Metasomal sternum II with postero-median projection. Pygidial plate absent. Female: Lower paraocular area without yellow. Tibial scopa sparse, density similar to or less than that of fore femur. Third fiagellar segment as long as broad or longer. Etyynology: The prefix comes from the Greek melas, black, since this is the predominant color in these species. Liphanthus (Melaliphanthus) atratus new species Figs. 90-95; Map I, K Diagnosis: Similar to L. penai but male with facial fovea almost half as long as scape and propodeal triangle distinctly striate; female with mandible mahogany only on distal third, frontal line in lower half a groove. See also characters given in the key. Male: Length about 5.0 mm, fore- wing length 3.2 mm, head width 1.5 mm, thoracic width 1.3 mm. Coloration. Black, the following parts yellow: distal third of central area of clypeus, la- brum, mandible (apex mahogany), an- terior and outer surface of fore tibia and most of fore basitarsus, small spot on base and on apex of mid tibia, base of hind tibia, and small and weak spot on base of mid and hind basitarsus (small segments of fore tarsus and most of tarsi II and III brown or almost black). Wings almost hyaline, tegula and veins dark brown. Pubescence. In general short and sparse. Labrum 272 The University of Kansas Science Bulletin pubescent on distal margin. Lateral area oiclypeus with hairs rather dense, as long as those of labrum. Integument and Punctation. Areolate between punc- tures, except smooth on yellow part of clypeus and on ventral paraocular area; propodeal triangle distinctly striate. Punctures hardly visible and sparse, better marked and denser on inferior paraocular area and lateral area of clypeus. Structure. Head 1.3 times as broad as long. Flagellar seg- ments much longer than broad, the first one almost 3 times as long as broad. Facial fovea almost half as long as scape. Inner subantennal suture al- most straight and distinctly longer than width of subantennal area. Clypeus about 3 times as broad as long, project- ing beyond lower orbital tangent for about one-third of its length. Labrum smooth, slightly convex, less than twice as broad as long, distal margin with median part convex. Mandible un- modified. Marginal cell shorter than distance from apex to wing tip. Second submarginal cell with anterior margin shorter than posterior. Legs unmod- ified. Metasomal sternum II with acuminate postero-median projection extending forward in sternum as a ridge with a groove at its base, pos- terior margin laterally as indicated in Fig. 92. Metasomal sternum III pos- teriorly produced in the middle. Genitalia and associated sterna as illus- trated. Female: Length about 5.6 mm, fore- wing length 3.7 mm, head width 1.6 mm, thoracic width 1.6 mm. Coloration. Black, except yellow and mahogany spot on distal third of mandible, weak yellow spot on anterior edge of fore tibia. Wings testaceous, veins and tegula dark brown. Pygidial plate ma- hogany. Pubescence. As described for male, tibial scopa rather sparse, den- sitv similar to that of hairs of fore femur. Integument and Punctation. In general areolate between punctures, clypeus with medium-sized punctures arranged in irregular longitudinal lines. Pygidial plate almost smooth api- cally. Punctation fine and somewhat dense on subantennal and supraclypeal area. Thoracic punctation sparse; weak and rather fine on metasoma. Structure. Head 1.2 times as broad as long. Frontal line, in lower half, a narrow groove. Facial fovea about three- fourths as long as scape. Subantennal suture as described for male. Clypeus less than 3 times as broad as long, projecting beyond lower orbital tan- gent for about half its length. Mandi- ble, legs and marginal cell as described for male. Pygidial plate with slightly prominent longitudinal ridge on apex. Type Material: Holotype male. Mal- leco Prov., Nahuelbuta, Chile, 9- 1-1979 (L. Ruz). Allotype female. Same data but (H. Toro), both in Toro col- lection. Paratypes. Same locality and date, 2 females (E. Chiappa, H. Pas- ten), 4 females (P. Toro), and 1 male (E. de la Hoz); 1 female, Arauco Prov., Caramavida, Cordillera Na- huelbuta, 13-1-1967 (Schlinger), and 1 male, Arauco Prov., Nahuelbuta, 1-1978 (M. Cerda). Paratypes are de- posited in the following collections: AMNH, KU, CAS, UCV, and HT. Etymology: The species name refers to its almost completely dark colora- tion. Liphanthus (Melaliphanthus) penai new species Figs. 96-101; Map I, J Diagnosis: Closely related to L. atratus but male with facial fovea very small, less than one-third as long as scape and propodeal triangle sparsely striate; female with mostly mandible yellow-mahogany, frontal line with lower part not a groove. The Bee Genus Liphanthus 273 94 95 101 Figs. 90-95. Liphanthus (Melaliphanthus) atratus n. sp., male: 90, head frontal view; 91, labrum; 92, ventral view of rnetasoma; 93, 94, seventh and eighth metasomal sterna; 95, dorsal and ventral views of genitalia. Figs. 96-101. Liphanthus (Melaliphanthus) penai n. sp., male: 96, head frontal view; 97, labrum; 98, seventh metasomal sternum: 99, ventral view of meiasoma, 100, eighth metasomal sternum: 101. dorsal and ventral views of genitalia. Male: Length about 4.4 mm, fore- wing length 2.8 mm, head width 1.4 mm, thoracic width 1.1 mm. Coloration. Head and dorsal part of thorax black, rest of the body brown, the following parts yellow: weak line on distal mar- gin of clypeus, labrum, mandible (apex mahogany), spots on apices of femora, inner and outer surfaces of fore tibia, most of fore basitarsus, spot on base of mid and hind tibiae (small segments of fore tarsus and tarsi II and III brown). Wings lightly testaceous, veins and tegula dark testaceous. Pubescence. In 274 The University of Kansas Science Bulletin general short and sparse. Labrum pubescent on distal margin. Lateral area of clypeus with hairs rather abun- dant, slightly longer than those of labrum. Integument and Punctation. Strongly areolate and somewhat dull between punctures, except lighdy areo- late and shiny in propodeal triangle (also sparsely striate) and metasoma. Punctation weak, deeper on inferior paraocular area, very sparse on hypo- epimeral area. Structure. Head 1.3 times as broad as long. Flagellar segments much longer than broad, the first one somewhat more than 3 times as long as broad. Facial fovea small, less than one-third as long as scape. Inner subantennal suture almost straight, slightly shorter than width of subanten- nal area. Clypeus more than twice as broad as long, projecting beyond lower orbital tangent for about one-fourth of its length. Labrum slightly convex, broader than long, distal margin lightly produced in middle. Mandible un- modified. Marginal cell shorter than distance from apex to wing tip. Second submarginal cell with anterior margin shorter than posterior. Legs unmodi- fied. Metasomal sternum II with very acute postero-median projection (with median longitudinal groove at base), posterior margins of sternum II later- ally gradually convergent toward pro- jection (Fig. 99). Sternum III with posterior margin almost straight. Geni- talia and associated sterna as illus- trated. Female: Length about 4.8 mm, fore- wing length 3.7 mm, head width 1.6 mm, thoracic width 1.6 mm. Coloration. Brown, almost black, except: mandible mostly yellow-mahogany, weak yellow band on anterior edge of fore tibia, weak yellow spot on base of mid tibia. Wings lightly brown, veins and tegula darker than in male. Pygidial plate dark mahogany. Pubescence. As de- scribed for male; tibial scopa less dense than hairs of fore femur. Integument and Punctation. In general areolate between punctures, with some striae in pro- podeal triangle. Clypeus with medium- sized punctures irregularly arranged, shallow in interalveolar area, sparse on thorax, rather fine in metasoma. Py- gidial plate smooth. Structure. Head 1.2 times as broad as long. Frontal line, in lower half, hardly visible. Facial fovea little more than three-fourths as long as scape. Inner subantennal suture as de- scribed for male. Clypeus little less than 3 times as broad as long, project- ing beyond lower orbital tangent for somewhat more than half of its length. Mandible and marginal cell as de- scribed for male. Pygidial plate with longitudinal ridge on apex. Type Material: Ho lo type male. Mal- leco Prov., Rio Blanco,' Chile, 11-1964 (L. Peiia), in Toro collection. Allotype female. Malleco Prov., Rio Blanco, Chile, 11-1959 (L. Pena), in FAUCH collection. Paratypes. 3 females, Malleco Prov., Rio Blanco, Curacautin, Chile, 1-5-II-1959 (L. Peiia) deposited in the following collections: UCV and HT. Etymology: This species is dedicated to Sefior Luis Peiia in recognition of his contributions to the knowledge of the insects of Chile. Neoliphanthus new subgenus Figs. 102-108 Type species: Liphanthus bicellularis new species. Both sexes: Vertex convex. Lateral ocelli above upper orbital tangent. In- teralveolar and supraclypeal areas pro- tuberant. Frontal line weakly marked, without groove on lower end. Clypeus convex basally, flat distally, without median longitudinal groove, lateral area slightly sloping upward toward central area, widened mesally in ven- The Bee Genus Liphanthus 275 tral view. Inner orbits slightly emargi- nate in dorsal half (almost straight in female), divergent above. Distance be- tween alveoli somewhat greater than distance from alveolus to inner orbit and longer than length of alveolar di- ameter. Outer subantennal suture almost straight. Pronotum without protuberant dorsolateral angle. Sub- marginal cells two. Propodeal triangle laterally less than half length of meta- notum laterally (about half in female). Tibial spurs pale testaceous. Hind tibial spurs with apices slightly curved, inner somewhat longer than outer. Male: Head and legs with yel- low marks, metasoma without yellow bands. Lateral area of clypeus with hairs about as long as clypeal length and sparse. Flagellum much longer than head; first flagellar segment about twice as long as broad. Facial fovea short, well defined, close to inner orbit. Middle of paraocular area convex, lower part of paraocular area slightly sloping upward to central area. Post- mandibular area mesally much less than half as long as width of base of mandible. Sterna II and III each with median apical projection. Pygidial plate absent. Female: Lower paraocular area without yellow spot. Tibial scopa sparse, with hairs as dense as those of fore femur. Third llagellar segment longer than broad. Etymology: The subgeneric name is from the Greek neos meaning new plus Liphanthus. Liphanthus (Neoliphanthus) bicellularis new species Figs. 102-108; Map II, J Diagnosis: Similar to L. nitidus but both sexes with 2 submarginal cells, head about 1.3 times as broad as long, clypeus projecting beyond lower orbital tangent for less than hall' of its length, supra-antennal area distinctly areolate and dull. Male llagellum much longer' than head. Male: Length about 3.3 mm, fore- wing length 2.3 mm, head width 1.2 mm, thoracic width 0.8 mm. Coloration. Black, except most of llagellum testa- ceous beneath, the following parts Figs. 102-108. Liphanthus (Neoliphanthus) hicellularis n. sp., male: 102, lOS, head frontal and lateral views; 104, labrum; 105, dorsal and ventral views of genitalia; 106, 107, seventh and eighth inetasomal sterna; 108, forcwing and hindwing. 276 The University of Kansas Science Bulletin yellow: clypeus, lower part of paraocu- lar area extending above alveoli, sub- antennal area, labrum, mandible (apex mahogany), under side of scape, pro- notal lobe, apices of femora, tibiae, basitarsi and most of 2nd tarsal seg- ments (rest of segments of tarsi brown). Wings almost hyaline, veins and tegula brown. Pubescence. In general short and sparse. Labrum pubescent on distal third. Lateral area of clypeus scarcely pubescent, with hairs somewhat longer than those of labrum. Integument and Punctation. Head strongly areolate and dull, except weakly areolate or smooth and brilliant below alveolar level and lowest part of gena. Thorax slightly areolate, with some striae on propodeal triangle, smooth on metasoma. Punc- tures in general weak, fine and sparse, denser on most of gena and lower part of paraocular area, coarser on clypeus, very fine and sparse on hypoepimeral area, shallow on rest of mesepister- num. Structure. Head about 1 .3 times as broad as long. Flagellar segments much longer than broad. Facial fovea elongate oval, less than half as long as scape. Inner subantennal suture almost straight, longer than width of suban- tennal area. Clypeus more than twice as broad as long, projecting beyond lower orbital tangent for one-fifth of its length. Labrum with transverse carina, pubescent at distal area, somewhat broader than long, distal margin lightly convex. Mandible unmodified. Margi- nal cell shorter than distance from apex to wing tip. Second submarginal cell with anterior margin distinctly shorter than posterior. Legs unmodified. Ster- num II with posterior margins laterally gradually convergent toward median spine-like projection (similar to Fig. 142, but projection very acute, straight, not hairy at apex). Sterna III and I\' with posterior margins dis- tinctly produced medially; sternum V with posterior margin slightly convex. Genitalia and associated sterna as illus- trated. Sterna III and IV with posterior margins distinctly produced medially; sternum V with posterior margin slightly convex. Genitalia and associ- ated sterna as illustrated. Female: Length about 4.4 mm, fore- wing length 3 mm, head width 1.4 mm, thoracic width 1.3 mm. Coloration. Brown, almost black, the following parts yellow: T-shaped spot on upper part of clypeus (size somewhat vari- able), labrum (rarely), subantennal area (sometimes absent or asymmetri- cal), inner lobe of lower paraocular area (sometimes absent or asymmetri- cal), weak spot on mandible (apex ma- hogany), pronotal lobe, basal spot on each fore and middle tibia. Wings, veins and tegula as described for male. Pygidial plate mahogany. Pubescence. As described for male; tibial scopa sparse, hairs as dense as those of fore femur. Integument and Punctation. Integ- ument as described for inale except face more weakly areolate and shiny. Py- gidial plate smooth apically. Puncta- tion as described for male but deeper and denser on face, gena and hypo- epimeral area. Structure. As described for male but facial fovea sinuous, more than half as long as scape, clypeus about 3 times as broad as long, project- ing beyond lower orbital tangent for one-fourth of its length. Pygidial plate with longitudinal ridge on apex. lype Material: Holotype male and al- lotype female. Linares Prov., Castillo, Chile, 11-1976 (P. Toro), in Toro col- lection. Paratopes. Same locality and date, 14 males and 4 females (H. Toro), 2 males (H. Toro, Jr.), 6 males and 3 females (P. Toro); same locality, 10-11-1977, 3 males and 15 females (H. Toro), 2 males and 4 females (P. Toro). From Linares Prov., Bullileo, 15-XII-1977, 14 females (O. Mar- The Bee Genus Liphanthus 277 tmez), 2 females (E. Chiappa, H. Toro), 4 females (E. Peralta) and 1 male (L. Ruz). Paratypes are deposited in the following collections: AMNH, KU, MNHN, CAS, FAUCH, and UCV. Etymology: This species is called L. bicellularis because its forewings have only 2 submarginal cells. Comments: The allotype, like some of the other female paratypes, lacks yellow on the face except for the mark on the clypeus. Leptophanthus new subgenus Figs. 109-151 Type species: Psaenythia nigra Friese, 1916. Both sexes: Vertex convex. Lateral ocelli above upper orbital tangent. In- teralveolar area in general little pro- tuberant (more prominent in L. cerdai). Supraclypeal area in general pro- tuberant. Clypeus protuberant without median longitudinal groove, lateral area abruptly sloping upward toward central area, widened mesally in ven- tral view (similar to Fig. 3). Inner orbits divergent above, slightly con- cave in upper halves, more distinctly concave in L. nitidus and L. coqunnbensis (almost straight in females). Inter- alveolar distance longer than alveolor- bital distance and about twice as long as alveolar diameter. Outer subanten- nal suture slightly arcuate laterally. Pronotum with dorsolateral angle not protuberant. Submarginal cells three. Propodeal triangle laterally more than half length of metanotum laterally (al- most same length in L. nitidus and L. alicahue). Tibial spurs light yellow. Hind tibial spurs with apices slightly curved, inner slightly longer than outer. Male. Head and legs with yel- low marks. Metasoma without yellow bands. Lateral area of clypeus with hairs sparse and shorter than length of clypeus. Frontal line generally well de- fmed, in a wide depression basally (except L. anacanthus and L. australis). Flagellum little longer than head; first flagellar segment less than twice as long as broad. Facial fovea oval, shorter than scape, separated from inner orbit by a variable distance, borders some- times not clearly defined. Median paraocular area generally slightly de- pressed toward mid-line of face; lower paraocular area strongly sloping up- ward toward mid-line of face. Post- mandibular area mesally much less than half as long as width of base of mandible. Metasomal sternum II gen- erally with median distal projection. Pygidial plate absent. Female. Inferior paraocular area without yellow (except in some specimens of L. alicahue). Tibial scopa of variable density. Third flagellar segment broader than long. Etymology: The subgeneric name is from the Greek leptos, meaning thin, small (a common feature for the species here included) plus the major part of the generic name. Liphanthus (Leptophanthus) nitidus new name Figs. 109-114; Map II, G Psaenythia nigra Friese, 1916: 167 (not P. bizonata var. nigra Friese, 1908: 41); Jaffuel and Pirion, 1926: 369. Male: Length about 4 mm, fore- wing length 2.8 mm, head width 1.2 mm, thoracic width 1.0 mm. Coloration. Black, flagellum mostly testaceous, the following parts yellowish white: in- ferior paraocular area extending up to alveolar level, clypeus (sometimes black basally), labrum, mandible (apex mahogany), pronotal lobe, small spot on tegula, outer surface of fore tibia, bases and apices of mid and hind tibiae and most of basitarsi (rest of tarsi brown). Wings hyaline, veins and tegula testaceous. Pubescence. In general 278 The University of Kansas Science Bulletin short and sparse. Lab rum pubescent on distal half. Lateral area of clypeus sparsely pubescent with hairs about as long as those of labrum. Integument and Punctation. Head and thorax almost smooth, shiny between punctures ex- cept areolate, dull on vertex; propodeal triangle striated. Punctures in general shallow, denser on lower half of face, somewhat coarse on clypeus, very fine and somewhat dense on hypoepimeral area, coarser and sparser on rest of mesepisternum. Structure. Head about as broad as long. Flagellar segments in general little longer than broad. Facial fovea in a smooth, shiny, long and wide groove, little more than half as long as scape, sometimes difficult to see. Inner subantennal suture almost straight and longer than width of sub- antennal area. Clypeus about twice as broad as long, projecting beyond lower orbital tangent for a little more than half of its length. Labrum convex, little less than twice as broad as long, with transverse carina at distal third, distal margin lightly concave. Mandible un- modified. Marginal cell longer than distance from apex to wing tip. Second submarginal cell with anterior margin about as long as posterior. Legs un- modified. Metasomal sternum II with postero-median spiniforni projection, curved at apex; distal margin of ster- num II strongly sinuous lateral to pro- jection (Fig. 111). Sternum III with posterior margin slightly produced mesallv. Genitalia and associated sterna as illustrated. Female: Length about 4.9 mm, fore- wing 3.4 mm, head width 1.4 mm, thoracic width 1.2 mm. Coloration. Black, usually most of llagellum testa- ceous (sometimes with darker areas beneath, the rest brown, the following parts yellow: small spot (sometimes absent) distally on clypeus, preapical spot on mandible (mahogany) toward the apex, and small proximal spot on fore and hind tibiae. Tegula without yellow spot. Wings as described for male but veins darker. Pygidial plate brown. Pubescence. As described for male, but tibial scopa much denser than hairs of fore femur. Integument and Punctation. About as described for male but much coarser, with well marked punctures on clypeus, hypoepimeral area more densely punctate, propodeal triangle smooth and shiny, without striae. Pygidial plate smooth, shiny. Structure. As described for male except: head 1 . 1 times as broad as long, facial fovea about as long as scape, variable in shape (varies from wide to narrow, and from straight to sinuous), clypeus projecting beyond lower orbital tan- gent for more than half of its length, marginal cell shorter than distance from apex to wing tip. Pygidial plate with longitudinal ridge apically. Material Studied (all from Chile): 3 males and 4 females, O'Higgins Prov., Termas de Cauquenes, 30-1-1972 Q. Leon) (1 female parasitized by Strep- siptera); 2 males and 2 females, Curico Prov., Rio Teno, Los Quenes, 14- 11-1965 (L. Peha); 5 males and 19 females, Linares Prov., Longavi, IX- 1980 (diverse collectors); 1 male and 1 1 females, Curico Prov., Los Queries, IX- 1980 (diverse collectors); 1 female, Concepcion Prov., Concepcion, 1903 (P. Herbst); 1 male and 30 females, Bfo-Bio Prov., Negrete, 31-XII-1950 (Ross and Michelbacher). Etymology: The new name is the Latin word nitidus, meaning shiny. Discussion: The type of Psaenythia nigra Friese, 1916, is perhaps lost. In the Berlin Museum there is a type labeled as P. nigra Fr., from Salta, Argentina, III- 1905 (Steinbach). How- ever, its correct name is P. bizonata var. nigra Friese, 1908, which is a real Psaenythia. According to the original The Bee Genus Liphanthus 279 description of P. nigra Fr., 1916, males and females were collected in Concep- cion, Chile. One female from this local- ity (as mentioned in Material Studied) was found in the Museum of Compara- tive Zoology. It is here designated as the neotype. The material studied was compared with the neotype and seems to be con- specific. However, females from Termas de Cauquenes show some dif- ferences, especially in the facial fovea, which is wider with poorly defmed edges, and in the weaker punctation of the face. On the other hand females from Linares also show some variation in size and shape of the fovea. Since all the males studied look alike, all the specimens recorded in that paper have been considered conspecific. More collections and especially males from Concepcion should help to decide whether there is intraspecific variation or ii closely related species are involved. y|^fS^\'^'^ pf^^ 119 Figs. 109-114. Liphanthus (Leptophanthus) nitidus n. name, male: 109, head frontal view; 110, labrum; 111, ventral view of metasoma; 112, 113, seventh and eighth meta.soinal sterna; 114, dorsal and ventral views of genitalia. Figs. 115-120. Liphanthus (Leptophanthus) breviceps (Friesc), male: 115, head frontal view; 116, labrum; 117, ventral view of metasoma; 118, 119, seventh and eighth metasomal sterna; 120. dorsal and ventral views ol genitalia. 280 The University of Kansas Science Bulletin Liphanthus (Leptophanthus) breviceps (Friese) new combination Figs. 115-120; Map II, D Psaenythia breviceps Friese, 1916: 167- 168; Jaffuel and Pirion, 1926: 369. Male: length about 4.2 mm, fore- wing 3 mm, head width 1.3 mm, tho- racic width 1.1 min. Coloration. Black, flagellum mostly testaceous beneath (rest of flagellum brown), the following parts yellowish white: most of clypeus, lower paraocular area (sometimes ex- tending up to alveolar level), lower half (approximately) of subantennal area in some specimens, labrum, mandible (apex mahogany), pronotal lobe, api- ces of femora, tibiae (except tibiae II and III with brown spot on outer sur- face), most of basitarsi (small segments of tarsi brown). Wings dark-brown, veins light brown, tcgula testaceous. Pubescence. In general short and sparse. Labrum pubescent on distal margin. Lateral area of clypeus scarcely pubes- cent with hairs little longer than those of labrum. Integument and Punctation. Head and thorax slightly areolate and shiny, smooth on face below alveoli level and most of gena, propodcal tri- angle with some striae. In general sparsely punctate, with punctures very sparse in hypoepimeral area above scrobe, deeper, coarser and denser on rest of mesepisternum. Structure. Head little broader than long. Flagellar seg- ments little longer than broad. Facial fovea straight, narrow, about half length of scape. Inner subantennal su- ture almost straight, longer than width of subantennal area. Clypeus about twice as broad as long, projecting be- yond lower orbital tangent for about half of its length or somewhat more. Labrum broader than long, flat bas- ally, with transverse carina at distal fifth, distal margin almost straight. Mandible unmodified. Marginal cell slightly shorter than (sometimes equal to) distance froin apex to wing tip. Second submarginal cell with anterior margin almost as long as posterior. Legs unmodified. Metasomal sternum II with short postero-median projec- tion, little longer than its apical width, pubescent and curved downward at apex; distal margin of sternum II slightly sinuous lateral to projection (Fig. 117). Genitalia and associated sterna as illustrated. Material Studied: 6 males, Acon- cagua Prov., Rio Blanco, Los Andes, Chile, 17-1-1969 (C. Vivar); 3 males, Aconcagua Prov., Las Coimas, 24- XI-1974 (L. Ruz); 1 male, Ri'o Hornillos, Chile, 7-XII-1917 and 1 male same locaHty, 29-XII-1917 (P. Herbst). Discussion: The type of Psaenythia breviceps Fr. has*probably been lost. We have examined one male (mentioned in Material Studied) from the Berlin Mu- seum labeled Rio Hornillos, 7-XII- 1917 (P. Herbst), "Typus" and Psae- nythia breviceps Friese (1909, sic). How- ever, this could not be the type, since its date of collection (1917) is later than the time of publication (1916). More- over in Friese's original description males were reported from Victoria, Chile. We have not seen specimens from Victoria and have not been able to find the whereabouts of Rio Hor- nillos. The specimen labeled Rio Hor- nillos is here designated as the neotype, because it bears the "Typus" and identification label written by Friese. Liphanthus (Leptophanthus) anacanthus new species Figs. 121-126; Map II, F Diagnosis: Similar to L. nitidus but male without spine-like projection on metasomal sternum II and labrum pubescent on distal fifth; female with Thk Bee Genus Liphanthus 281 integument strongly areolate and opaque above alveoli. Male: Length about 4.4 mm, tbrx'- wing 2.8 mm, head width 1.4 mm, thoracic width 1.2 mm. Coloration. Black, flagcllum brown, the following parts light yellow: clypeus (sometimes dorsal area black), spot on lower para- ocular area extending up, close to inner orbit, to alveolar level, subantennal area (lower part), labrum, mandible (apex mahogany), small spot on tegula pronotal lobe, apices of femora, tibiae (except median dark spot on II and III), basitarsi, and second segment of hind tarsus (rest of tarsi brown). Wings slightly dark brown, veins brown, tegula testaceous. Pubescence. In general short and sparse. Labrum pubescent on distal fifth. Lateral area of clypeus sparsely pubescent, with hairs longer than those of the labrum. Integument and Punctation. Head and thorax areolate, more distinctly areolate and opaque on face above alveolar level, smooth and shiny on lower half of face and gena, propodeal triangle with some striae. Metasoma smooth. Punctation in gen- eral fine and rather sparse, somewhat denser on face distally, coarser on clypeus, rather sparse and very fine on hypoepimeral area, coarser on rest of mesepisternum. Structure. Head little broader than long. Flagellar .segments longer than broad, first and second only slightly so. Frontal line, weakly distinguishable. Facial fovea in a some- what shiny area beside inner orbit, straight, narrow, about half as long as scape. Inner subantennal suture straight and longer than width of subantennal area. Interalveolar area slightly prominent, clypeus about twice as broad as long, projecting beyond lower orbital tangent for about half (or less) of its length. Labrum less than twice as broad as long, flattened basally, with transverse carina at distal fifth, distal margin straight. Mandible unmodified. Marginal cell shorter than distance from apex to wing tip. Second submarginal cell with anterior margin distinctly shorter than posterior. Legs unmodified. Metasomal sternum 11 without well-developed projection, mid-line posteriorly with short ex- panded and slightly bifurcate area, pubescent on margin; posterior margin of sternum II scarcely sinuous lateral to projection (Fig. 123). Genitalia and associated sterna as illustrated. Female: Length about 5.4 mm, fore- wing 3.6 mm, head width 1.6 mm, thoracic width 1.5 mm. Coloration. Brown, almost black, flagcllum testa- ceous beneath, the following parts yellow: mandible (apex mahogany), la- brum, pronotal lobe, bases of tibiae. Wings, veins, and tegula as described for male. Pygidial plate dark ma- hogany. Pubescence. As described for male but tibial scopa as dense as hairs of fore femur. Integument and Punctation. As described for male, but pygidial plate slighdy areolate and punctures deeper on lower half of face. Structure. As described for male except head L3 times as broad as long, facial fovea widened distally, longer than scape; metasoma unmodified, pygidial plate with longitudinal ridge. Type Material: Holotype male and al- lotype female. O'Higgins Prov., Termas de Cauquenes, Chile, 30-1-1972 (J. Leon), in Toro collection. Paratypes (all from Chile). 9 males and 5 females, O'Higgins Prov., Termas de Cau- quenes, 30-1-1972 (J. Leon); 21 males, O'Higgins Prov., Pangal, 11-1978 (H. Flores); 3 females, Valparaiso Prov., Cuesta La Dormida, 30-XII-1971 (H. Toro); 1 male, Aconcagua Prov., Las Coimas, XI-1974 (L. Ruz). Paratypes are deposited in the following collec- tions: AMNH, KU, MNHN, and HT. Etymology: The specific name refers 282 The University of Kansas Science Bulletin to the absence of a long spine on metasomal sternum II. Such a spine is present in most species of this sub- genus. Liphanthus (Leptophanthus) australis new species Figs. 127-132; Map II, N Diagnosis: Similar to L. nitidus but male with projection of metasomal ster- num II shorter and wider, labrum pubescent on distal fifth. Male: Length about 4.1 mm, fore- wing 2.6 mm, head width 1.3 mm, thoracic width 1.1 mm. Coloration. Black, tlagellum mostly testaceous be- neath (rest of flagellum brown), the following parts light yellow: clypeus, lower paraocular area extending up to alveolar level (or a little further), sub- antennal area, labrum, mandible (apex mahogany), pronotal lobe, spot on tegula, apices of femora, tibiae, basi- tarsi and most of second segment of hind tarsus. Wings slightly testaceous, veins and tegula testaceous. Pubescence. In general short and sparse. Labrum pubescent on distal fifth. Lateral area of clypeus scarcely pubescent, hairs somewhat longer than those of labrum. Integument and Punctation. Areolate and somewhat opaque on head above al- veolar level and on mesepisternum; rather smooth on lower half of face, 123 124 Figs. 121-126. Liphanthus (Leptophanthus) anacanlhus n. sp., male: 121, head frontal view; 122, labrum; 123, ventral view of metasoma; 124, 125, seventh and eighth metasomal sterna; 126, dorsal and ventral views of genitalia. Figs. 127-1.32. Liphanthus (Leptophanthus) australis n. ^y. male: 127, head frontal view; 128, labrum; 129, ventral view of rnetasoma; 130, 131, seventh and eighth metasomal sterna; 132, dorsal and ventral views of genitalia. The Bee Genus Liphanthus 283 gena and dorsally on thorax, propodeal triangle distinctly striated. Metasoma smooth. Punctation in general fine and not dense, deeper on lower paraocular area, coarse on clypeus, sparse on scutum, very fine and sparse on hypo- epimeral area, coarser and denser on rest of mesepisternum. Structure. Head about as broad as long. Flagellar seg- ments about as long as broad, first and last litde longer. Frontal line weak toward base, with weakly grooved apex. Facial fovea elongate oval, less than half as long as scape. Inner suban- lennal suture almost straight and longer than width of subantennal area. Clypeus little more than twice as broad as long, projecting beyond lower orbi- tal tangent for less than half of its length. Labrum less than twice as broad as long, with light transverse carina at distal fifth, distal margin al- most straight. Mandible unmodified. Marginal cell longer than distance from apex to wing tip. Second sub- marginal cell almost rectangular, with anterior margin almost as long as pos- terior. Legs unmodified. Metasomal sternum II with short postero-median projection, about twice as long as broad (width measured at apex), pu- bescent at apex; posterior margins of the sternum gradually convergent to- ward projection (Fig. 129). Genitalia and associated sterna as illustrated. Type Material: Holotype male. Santa Cruz Prov., 2 km E Los Antiguos, 280 m, Argentina, 21-XI-1966 (Irwin and Schlinger), in Toro collection. Para- types. 1 male, Santa Cruz, Argentina, 21 -XI- 1966 (Irwin and Schlinger), in UCV collection. Etymology: The species name al- ludes to the austral distribution of this species. Liphanthus (Leplophanthus) coquimbensis new species Figs. 133-139; Map II, A Diagnosis: Closely related to L. nitidus but male with transverse carina of labrum at distal fifth and metasomal sternum III with posterior margin straight mesally; female with clypeus generally black, basal yellow spot on mid tibia longer than last fiagellar seg- ment. Male: Length about 3.6 mm, fore- wing 2.7 mm, head width 1.2 mm, thoracic width 1.0 mm. Coloration. Black, most of flagellum testaceous, the following parts light yellow: clypeus (dorsal part black in some specimens), inferior paraocular area usually ex- tending up to alveolar level (sometimes reduced), usually lower half of suban- tennal area (approximately), labrum, weak basal spot on mandible (apex mahogany), pronotal lobe, apices of femora, most of fore tibia, bases (con- tinued along dorsal edges) and apices of mid and hind tibiae (small segments of tarsi brown). Wings slighdy brown, veins, tegula and last metasomal ster- num testaceous. Pubescence. In general short and sparse. Labrum pubescent on distal fifth. Lateral area of clypeus sparsely pubescent with hairs some- what longer than those of labrum. In- tegument and Punctation. In general shiny, slightly areolate between punc- tures on upper part of head. Rest of body in general smooth, but hypo- epimeral area areolate; propodeal tri- angle shiny, apparently without striae, sometimes somewhat areolate. Head and thorax with rather fine and sparse punctures, denser and coarser on clypeus and mesepisternum, very fine and sparse on hypoepimeral area. Structure. Head scarcely broader than long. Flagellar segments little longer than broad, last one more distinctly so. Frontal line weak toward base, apically 284 The University of Kansas Science Bulletin in well developed groove. Facial fovea narrow, poorly defined. Inner suban- tennal suture slighdy arcuate toward mid-line of face and little longer than width of subantcnnal area. Clypeus little more than twice as broad as long, projecting beyond lower orbital tan- gent for less than half of its length. Labrum less than twice as broad as long, with transverse carina at distal fifth, distal margin slightly concave. Mandible unmodified. Marginal cell about as long as distance from apex to wing tip. Second submarginal cell with anterior margin slightly shorter than posterior. Legs unmodified. Metaso- mal sternum II with posterior margin sinuous at each side and with median projection more than twice as long as broad (width measured at apex), curved downward at apex; posterior margin of sternum II distinctly sinuous lateral to projection (Fig. 139). Geni- talia and associated sterna as illus- trated. Female: Length about 4.6 mm, fore- wing 3.6 mm, head width 1.5 mm, thoracic width 1.4 mm. Coloration. Black, small basal yellow spot on man- dible (apex mahogany), rarely with yellow spot on clypeus. Wings, veins and tegula as described for male. Pygidial plate mahogany. Pubescence. As described for male; tibial scopa about as dense as hairs of fore femur. Integument and Punctation. Shiny as in male, but i)unctures much denser and much deeper on lower half of face, fine around alveoli, coarse and dense on clypeus. Pygidial plate smooth. Struc- ture. As described for male but head 1 .2 times as broad as long, facial fovea well developed, little longer than scape. Clypeus slightly more than twice as broad as long. Pygidial plate with lon- gitudinal ridge on apex. Type Material: Holotype male. Co- quimbo Prov., Los Choros, Chile, 2- 11-1972 (L. Ruz). Allotype female. Same locality and date as holotype (H. Toro), both in Toro collection. Para- types (all from Chile). Coquimbo Prov. 2 females, 8 km W of La Junta, 4 females, 16 km W of La Junta and 1 female, 15 km W of La Junta, all 7- XII- 1950 (Ross and Michelbacher); 5 females, 8 km N of Laguna Dam, 2.438 m elev., 6-XII-1950 (Ross and Michelbacher); 3 females, 32 km E of La Serena, 3-XII-1950 and 1 female, 40 km E of La Serena, 5-XII-1950 (Ross and Michelbacher); 36 males and 8 females, Rio Los Choros, 6- XI- 1956 (R. Wagenknecht); 5 males and 4 females, Rivadavia, 28-X-1957 (L. Peha); Choros Bajos, 12-X-1977, 4 males and 4 females (H. Toro), 8 males and 16 females (J. C. Magunacelaya), 12 males and 3 females (L. Ruz), 3 males and 1 female (E. de la Hoz), 2 males and 2 females (E. Balart), 1 male and 1 female (V. Cabezas), 9 males and 2 females (H. Flores), 1 male and 1 female (M. Rojas); Quebrada Los Choros, 12-X-1977, 4 females (V. Cabezas), 1 female (L. Ruz); Los Choros, 2-IM972, 13 males and 7 females (H. Toro), 3 males and 7 females (H. W. Sielfeld), 7 females (M Rojas), 19 males and 40 females (L Ruz), 25 males and 40 females (M Pasten), 2 males and 23 females (E Montenegro), 1 female, 6-XI-1956 (R Wagenknecht); 23 males, Alcones, 29- XI-I97O (M. Pino); 2 females, Rio Choapa, 1-1970 (H. Toro); 6 males and 2 females. La Serena, 1-III-1970 (M. Cerda); 3 males and 2 females. Hacienda Illapel, 19-X-1966 (Irwin, Schlinger, Peha); 1 male, Socos, 22- IX-1966 (Irwin); 1 male. Fray Jorge, 28-XII-1966 (Irwin); 1 male, Illapel, Huintil, 19-X-1966 (L. Peha); 2 males, Huanta, 6-1-1966 (L. Pefia); 1 female, Buenos Aires, 2-XI-1954 (R. Wagen- knecht). Atacama Prov. 2 females. Las The Bee Genus Liphanthus 285 Juntas, IX-1968 (E. Montenegro); An- tofagasta Prov. 1 female, Chiu-Chiu, 24- 1-1972 (H. Toro). Paratypes are de- posited in the following collections: AMNH, KU, NMNH, CAS, BMNA, IMZT, MUPA, CONG, MNHN, UCV, and HT. Comment: The females of this spe- cies are difficult to separate from those of L. nitidus; the main differences are indicated in the key and diagnosis. Etymology: The species is named for the Coquimbo region where most of the specimens have been found. Liphanthus (Lcptophanthus) cerdai new species Figs. 140-145; Map II, M Diagnosis: Close to L. nitidus but male with head and thorax strongly areolate and frontal line almost cari- nate toward lower part and in minute groove; female strongly areolate and very finely punctate on face above al- veolar level. See also characters given in the key. Male: Length about 3.2 mm, fore- wing 2.4 mm, head width 1.1 mm, thoracic width 0.9 mm. Coloration. Black, flagellum mostly testaceous, the following parts yellow: clypeus, lower paraocular area extending up slightly beyond alveolar level, subantennal area (lower half), labrum, mandible (apex mahogany), pronotal lobe, api- ces of femora, outer surface of fore tibia, most of mid and hind tibiae except brown in the middle, spots on bases of fore and mid basitarsi and most of hind basitarsus (small segments of tarsi brown). Wings almost hyaline, veins and tegula brown. Pubescence. In general short and sparse. Labrum pubescent on distal fourth. Lateral area of clypeus sparsely pubescent, hairs somewhat longer than those of labrum. Integument and Punctation: Head and thorax areolate between punctures, more strongly areolate and opaque above alveolar level, propodeal triangle with some striation. Metasoma smooth and shiny. Punctation on clypeus, coarser, denser, and deeper than on rest of face, shallow and sparse on thorax. Structure. Head 1.2 times as broad as long. Flagellar segments longer than broad, the second one slightly so. Facial fovea distinct, about one-third as long as scape. Inner sub- antennal suture almost straight, longer than width of subantennal area. Clyp- eus little more than twice as broad as long, projecting beyond lower orbital tangent for almost half its length. La- brum twice as broad as long, with weak transverse carina at distal fourth, distal margin slightly concave laterally. Man- dible unmodified. Marginal cell shorter than distance from apex to wing tip. Second submarginal cell rectangular, with anterior margin as long as pos- terior. Legs unmodified. Metasomal sternum II with postero-median acu- minate projection, curved downward and hairy at apex, posterior margins of sternum laterally gradually convergent toward projection (Fig. 142). Sterna III and IV with posterior margins slightly produced mesally (more dis- tinct on III). Genitalia and associated sterna as illustrated. Female. Length about 4 mm, fore- wing 2.7 mm, head width 1.3 mm, thoracic width 1.2 mm. Coloration. Black, flagellum mostly testaceous, the following parts yellow: small apical spot on clypeus, weak spot on base of mandible, pronotal lobe, apices of femora, basal parts of mid and hind tibiae. Wings slightly dusky. Veins and tegula about as described for male. Pygidial plate dark mahogany. Meta- somal sterna brown. Pubescence. As de- scribed for male; tibial scopa about as dense as hairs of fore femur. Integument and Punctation. As described for male 286 The University of Kansas Science Bulletin 134 135 136 jffW^^\ Figs. 133-139. Liphanthus (Leplophanthus) coquimbensis n. sp., male: 133, 134, head frontal and lateral views; 135, labrum; 136, dorsal and ventral views of genitalia; 137, 138, eighth and seventh metasomal sterna; 139, ventral view of metasoma. Figs. 140-14.'). Liphanthus (Leptophanlhus) cerdai n. sp., male: 140, head frontal view; 141, labrum; 142, ventral view of metasoma; 143, dorsal and ventral views of genitalia; 144, 145, seventh and eighth metasomal sterna. Figs. 146-151. Liphanthus (Leplophanthus) alicahue n. sp., male: 146, head frontal view; 147, labrum; 148, seventh metasomal sternum; 149, dorsal and ventrcd views of genitalia; 150, eighth metasomal sternum; 151, ventral view of metasoma. The Bee Genus Liphanthus 287 but pygidial plate smooth and face less distinctly areolate and more densely punctate in its lower hall, punctures very shallow on thorax. Structure. As described for male but head 1.3 times as broad as long, facial fovea elon- gated, little shorter than scape, inner subantennal suture as long as width of subantennal area, clypeus little less than 3 times as broad as long. Second submarginal cell with anterior margin slightly shorter than posterior. Pygidial plate without ridge apically. Type Material: Holotype male. Mal- leco Prov., Nahuelbuta, Chile, 11-1978 (P. Toro). Allotype female. Same locality and date (H. Toro), in Toro collection. Paratypes (all from Chile). 5 males and 4 females, Malleco Prov., Nahuelbuta, 11-1978 (H. Toro); 2 males, same lo- cality and date (P. Toro) and 2 males, without collector's name; 2 females, Malleco Prov., Nahuelbuta, 9-1-1979 (P. Toro) and 2 females, same locality and date (E. Balart); 8 males and 4 females, Malleco Prov. Angol, IX- 1980 (E. de la Hoz, H. Toro, F. Rodri- quez). Paratypes are deposited in the following collections: AMNH, KU, MNHN, and UCV. Etymology: This species is dedicated to Dr. Miguel Cerda from the Institute de la Patagonia (Chile), in appreciation of his contribution to the knowledge of the Chilean entomofauna. Liphanthus (Leptophanthus) alicahue new species Figs. 146-151; Map II, B Diagnosis: Close to L. nitidus but male with posterior margins of metaso- mal sternum II laterally gradually con- vergent toward median apical spine- like projection, face strongly areolate and dull above alveoli; female with longitudinal yellow band on clypeus. Male: Length about 4.3 mm, fore- wing 2.8 mm, head width 1.3 mm. thoracic width 1.1 mm. Coloration. Black, flagellum mostly testaceous, the following parts yellow: clypeus, in- ferior paraocular area extending up beyond alveolar level, most of suban- tennal area, labrum, mandible (apex mahogany), pronotal lobe, weak spot on tegula, apices of femora, tibiae (but dark area in middle of tibiae II and III) and most of basitarsi (small segments of tarsi brown). Wings hyaline, veins and tegula testaceous. Metasomal sterna brown. Pubescence. In general short and sparse. Labrum pubescent on distal third. Lateral area of clypeus sparsely pubescent, with hairs some- what longer than those of labrum. In- tegument and Punctation. Head on upper half strongly areolate and opaque, with punctures very fine, not dense and not distinct; lower half of head smooth, shiny and finely punctate except coarser punctures in central area of clypeus. Thorax almost smooth, shin- ing, with well distinct and fairly dense punctures on scutum; hypoepimeral area areolate, with fine and sparse punctures, coarser and denser on rest of mesepisternum, propodeal triangle somewhat striated. Structure. Head slightly broader than long. Flagellar segments little longer that broad. Frontal line weakly defined on upper part, in a groove at lower apex. Facial fovea distinct, short, about one-fourth as long as scape. Inner subantennal suture almost straight, little less than twice as long as width of subantennal area. Clypeus little more than twice as long as broad, projecting beyond lower orbital tangent for half of its length. Labrum with weak transverse carina at distal third, little less than twice as broad as long, distal margin slightly concave. Mandible unmodified. Mar- ginal cell about as long as distance from apex to wing tip. Second submarginal cell with anterior margin almost as long 288 The University of Kansas Science Bulletin as posterior. Legs unmodified. Meta- somal sternum II with postero-median spine-like projection curved downward at apex; posterior margins of the ster- num gradually convergent toward spine (Fig. 151). Metasomal sternum III with posterior part produced mes- ally. Genitalia and associated sterna as illustrated. Female: Length about 5 mm, fore- wing 3.4 mm, head width L6 mm, thoracic width 1.4 mm. Coloration. Black, flagellum mostly testaceous, la- brum testaceous, the following parts yellow: longitudinal median band on clypeus, small spot (sometimes absent) on inner lobe of lower paraocular area, mandible (apex mahogany), pronotal lobe, small spots on apices of femora and bases of tibiae. Wings slightly tes- taceous, veins and tcgula testaceous. Pygidial plate mahogany. Pubescence. As described for male, though in gen- eral denser; tibial scopa as dense as hair of fore femur. Integument and Punc- tation. As described lor male but pro- podeal triangle scarcely striated, shiny in middle; pygidial plate slightly areo- late. Structure. As described for male except: head 1.2 times as broad as long; facial fovea somewhat wider, elongated, little shorter than scape; clypeus little less than 3 times broader than long. Marginal cell somewhat shorter than distance from apex to wing tip. Pygidial plate with wide lon- gitudinal ridge on aj^ex. Type Material: Holotype male. Ata- camaProv., Paipote, Chile, 2()-X-1971 (J. G. Rozen, L. Pefia). Allotype female. Same locality and collectors, 11 -X- 1971, both in AMNH. Paratopes (all from Chile). Atacama Prov. 23 males and 11 females, Paipote, 20-X-1971 Q. G. Rozen, L. Pefia); same locality and collectors, 1 female, ll-X-1971, 14 males and 17 females, 12-X-1971; 13 males and 13 females, vicinity of Freirina, 14-X-1969 and 1 female, same locality, 15-X-1969; 1 female, 16-32 km S Copiapo, 18-X-1969; 6 males and 1 female, between Paipote and Lajunta, 15-X-1969 (all the above collected by J. G. Rozen, L. Pefia); 1 female, Vallenar, X-1972 (H. W. Siel- feld); Lajunta, IX-1969, 1 female (E. Montenegro), 2 females (H. Toro), and 1 female (L. Ruz). Coquimbo Prov. Rio Laguna, 3,000 m, 1-1970, 1 male (L. Ruz), 1 male and 1 female (E. de la Hoz), 3 males (H. Toro). Para- types are deposited in: AMNH, KU, NMNH, CAS, MNHN, UCV, and HT. Etymology: The species name is de- rived from the Mapuche word alicahue which means "dried out place." Comments: This species is men- tioned by Ehrenfeld and Rozen (1977) as a small undescribed species of Liphanthus and a possible host of Kelita tuberculata. Discussion: Besides the material mentioned above, the following series of other specimens, also from Chile, apparently belonging to the same spe- cies, have been examined: 6 females, Valparai'so Prov., Cuesta La Dormida, 21-XI-1971 (H. Toro); 4 males and 2 females, Aconcagua Prov., Riecillo, 20-1-74 (H. Toro, H. W. Sielfeld); 2 males and 1 female, Santiago Prov., Caj6n del Yeso, 20-XI-1967 (Ramf- rez); 1 female, Santiago Prov., El Romeral, 9-XI-1967 (Rami'rez); 1 female, Santiago Prov., Farellones, 1- 1-1973 (A. Moidenke); 1 female, Santi- ago Prov., El Canelo, 9-1-1967 (Irwin); 1 female, same collector, Santiago Prov., Las Condes, 19-III-1966; 8 males and 8 females, Santiago Prov., Farellones, 8-1-1981 (H. Toro); 2 males and 2 females, same data (L. Ruz); 1 female, same locality, 27- XlI-1979 (M. T. Arroyo); 1 male, Colchagua Prov., San Fernando, Ter- The Bee Genus Liphanthus 289 mas del Flaco, 11-1967 (H. Toro); 1 female, Curico Prov., Los Quencs, Ri'o Teno, 7-14-II-1965 _(L. Pena); 5 males and 4 females, Nuble Prov., Chilian, Las Cabras, 3-1-1963 (L. Pefia); 2 females, Nuble Prov., Las Trancas, from January to February (diverse collectors); 1 male and 3 females, Nuble Prov., Termas de Chilian, IM976 (H. Toro); 1 female, Malleco Prov., Icalma, 11-1973 (H. Toro); 1 male, Santiago Prov., Ce- rros de Tiltil, 1700 m, 16-1-1919 (P. Herbst). These specimens from central and southern Chile were not considered as paratypes. Assuming that all the specimens listed are conspecific, the geographic distribution of the species is from Mal- leco (south, central Chile) to Atacama in the north (letter B on Map II). Species Not Assigned to Subgenus Liphanthus friesellus new species Figs. 152-158; Map I, D Diagnosis: Similar to L. (Pseudoli- phanthus) unijasciatus but male with clypeus, in side view, concave basally, its thickness (due to great protuber- ance) greater than length; upper and median paraocular areas distinctly de- pressed; interalveolar area slightly pro- tuberant. Male: Length about 4.4 mm, fore- wing 3.3 mm, head width 1.6 mm, thoracic width 1.2 mm. Coloration. Head and thorax light brown, the fol- lowing parts yellow: clypeus distally, lower paraocular area below alveolar level, labrum (brown on median part), mandible (apex mahogany), small spot on pronotal lobe, spots on apices of femora (extending on ventral edge of femur II), most of tibiae (internal sur- face of each tibia and median area of external surface of II brown) and basitarsi (small segments of tarsi brown). Wings slightly testaceous, veins and tegula testaceous. Mctasoma light brown, black distally, tergum II with transverse median yellow band. Pubescence. In general short and sparse. Labrum pubescent on distal margin. Lateral areas of clypeus somewhat pubescent, with hairs longer than those of labrum. Integument and Punctation. Slightly areolate, smoother and shiny on face; propodeal triangle scarcely striate. Punctures in general shallow and sparse, extremely fine on meta- soma. Structure. Vertex slightly convex. Lateral ocelli at same level as upper orbital tangent. Head 1.3 times as broad as long. Inner orbit slightly concave medially. Interalveolar area prominent, somewhat less than dis- tance from alveolus to inner orbit and about twice as long as diameter of alveolus. Supraclypeal area slightly convex. Flagellum distinctly longer than head, flagellar segments conspic- uously longer than broad; first flagellar segment about 3 times as long as broad. Frontal line carinate, apex a minute groove. Facial fovea straight, narrow, about half as long as scape and about as broad as distance between its outer margin and inner orbit. Inner subantennal sutures converging above, each angulate in upper part and about as long as width of subantennal area; outer subantennal suture lightly arcu- ate laterally. Clypeus more than 3 times as broad as long, projecting be- yond lower orbital tangent for more than hall' its length, in side view con- cave basally with weak longitudinal median ridge; lateral area abruptly sloping upward toward central area, widened laterally in ventral view. Labrum flat with marginal carina, broader than long, distal margin al- most straight. Mandible curved, with widened preapical plate (Fig. 154). 290 The University of Kansas Science Bulletin 155 Fit;s. 152-158. Liphanthus friesellus n. sp., male: 152, 153, head frontal and lateral views; 154. mandible; 155, labrum; 156, seventh metasomal sternum; 157, dorsal and ventral views of genitalia; 158, eighth metasomal sternum. Median paraocular area depressed; lower paraocular area strongly sloping upward toward mid-line of face. Post- mandibular area mesally much less than half as long as width of base of mandible. Pronotum with dorso lateral angle not protuberant. Marginal cell shorter than distance from apex to wing tip. Second submarginal cell with anterior margin distinctly shorter than posterior. Propodeal triangle laterally more than half as long as metanotum laterally. Legs unmodified. Hind tibial spurs with apices slightly curved, inner slightly longer than outer. Metasomal sterna unmodified. Genitalia and asso- ciated sterna as illustrated. Type Material: Holotype male. Col- chagua Prov. (Pichilemu), Chile, X- 1929 (F. Ruiz), in Toro collection. Etymology: This species is dedicated to H. Friese who apparently recog- nized it as new. Discussion. Specimens of Liphanthus friesellus have been found identified in collections with a Friese manuscript name (in the genus Psaenythia) referring to the long antennae. This name was listed by Jaffuel and Pirion (1926). However, it seems that it has never been validly published. Phylogenetic Considerations Considering the groups proposed by Rozen (1951), Liphanthus is clearly not in the Panurgus group (Panurgus, Camptopoeum, Panurginus, and Epime- thea) or the Nomadopsis group No- madopsis, Acamptopoeum, Hypomacrotera, Spinoliella, and Callonychium). It is dis- tinguished from them both by the gen- eral shape of genital capsule and by the associated sterna. From Xenopanurgus and Perdita it is separated by the slender gonocoxites and comparatively small gonostyli. On the other hand, as Michener (1952) points out, Xenopanurgus differs from the other genera of Panurginae by nu- merous external morphological charac- ters. Perdita also differs from Liphanthus by its markedly short submarginal cell and broader pterostigma. Liphanthus was considered as a syn- onym of Psaenythia for several years. The Bee Genus Liphanthus 291 However, the significant dilTerences that the latter genus presents in its genitalia, such as non-articulated gono- styh with specialized hairs, pecuhar penis valves and large volsellac with many small teeth, show its considerable divergence from Liphanthus. Liphanthus is included in a group of genera having the alternatives to the above characters: articulated gonostyli without specialized hairs, simple penis valves, smaller vosellae with fewer teeth. These genera are Protandrena, Metapsaenythia, Pseudopanurgus, Hetero- sarus, and Pterosarus. Although the wing venation, and particularly the ptero- stigma, present important differences, the relation of Liphanthus to the genera listed above, and especially to Pro- tandrena, seems probable. A better knowledge of the genera of Neotropical Panurginae, however, is needed to clarify relationships between Liphanthus and the other genera. The plesiomorphic and apomor- phic characters have been determined by comparison within the genus and with other Panurginae. The genera listed in the preceding paragraph are, together, considered as the outgroup for cladistic analysis. Their resem- blances to Liphanthus are plesiomor- phies. The characters used in the cladistic analysis of Liphanthus are mainly those of males. Females, in general, are uniform within this group, being plesiomorphic for most of their characters. The apomorphies that define Li- phanthus as a holophyletic group are: pterostigma (measured from distal end of prestigma to base of vein r) narrow, similar to prestigma in width and as long as prestigma or slightly longer (0.6:0.6, 0.9:0.7) and with subparallcl sides; margin of pterostigma within marginal cell straight; basal transverse groove of metasomal tergum II (es- pecially in males) deeper than in terga III to VII. In the outgroup the con- trasting characters are: pterostigma broader than prestigma, about twice as long as prestigma or somewhat longer (0.8:0.45, 1.2:0.6, 1.15:0.5, 1.9:0.8), sides before vein r rather divergent toward apex of wing; pterostigmal margin in marginal cell convex; basal groove of metasomal tergum II as well developed as in terga III to VII. Cladistic relationships are postu- lated on the basis of 26 characters listed in Table 1. Their distribution among the subgenera is shown by numbered line segments in Figure 159. Initially, 30 characters were considered, but four of them were eliminated because thev may show convergence rather than synapomorphy. One of the discarded characters was the relation between the inter- alveolar distance and alveolar diame- ter. In Neo/iphanthus, Tricholiphanthus, and Leptophanthus these values are simi- lar; therefore a relation between the three subgenera seem possible. How- ever, as shown in Figure 159, on the basis of other characters these sub- genera are well separated and this char- acter must have arisen at least twice. Another character, also eliminated, was the presence of a mesal projection on the posterior margin of metasomal sternum II. This is a typical character of Neoliphanthus and Leptophanthus, but also present, in a similar way, in some species of Pseudoliphanthus and Tri- choliphanthus. The diverse positions of these sub- genera in Figure 159 suggest polyphy- letic origin of this feature. Moreover, the projection in the last subgenus looks somewhat different and could therefore be either another variable or a modification of the first variable. Sinuous inner orbits occur in Lep- tophanthus, Neoliphanthus, Melaliphan- 292 The University of Kansas Science Bulletin Table 1 . List oi Apomorphies for Subgenera of Liphanthus. 1. Vertex concave or almost flat. An autapomorphic character for Xeno liphanthus. The outgroup for Liphanthus and all the other Panurginae examined have the vertex convex, a character here considered as plesiomorphic. 2. Male with lateral ocelli at same level as upper orbital tangent. The outgroup and most of the Panurginae observed present the plesiomorphic condition, that is, male with the lateral ocelli above the upper orbital tangent. 3. Male with first flagellar segment 2 or more times as long as broad. This character separates the 6 remaining subgenera from Leptophanthus which retains the plesiomorphic alternative from the whole outgroup, that is, first flagellar segment less than twice as long as broad. 4. Male with flagellum distinctly longer than width of head. Only Leptophanthus has retained the plesiomorphic condition. Xenoliphanthus apparently has lost the derived condition. All the outgroup and most of the other Panurginae examined are also plesiomorphic in this character, i.e., with the first flagellar segment short and broad. 5. Male with inner orbits divergent ventrally and not sinuous. The outgroup presents the plesiomorphic condition, that is, inner orbits in males divergent dorsally and weakly sinuous. 6. Male with inner orbits strongly divergent ventrally. This is a progression of character 5. It is found in Liphanthus s. str. and Pseudoliphanthus, although in one species (L. unifasciatus of the later subgenus) it is not so conspicuous, being more as in Xenoliphanthus. The whole outgroup presents the plesiomorphic condition. Xenoliphanthus has its inner orbits slightly divergent ventrally (character 5). 7. Male with outer subantennal suture strongly arcuate laterally, almost angulate. An autapomorphic character of Pseudoliphanthus, although it is not so strongly curved in one of its species. The inner subantennal suture is almost straight or moderately arcuate in many bees including most Panurginae; this condition is here considered as plesiomorphic. 8. Male with median paraocular area with transverse swelling below the level of the facial fovea. This character is an autapomorphy for Liphanthus s. str. The absence of this character in the outgroup and other Panurginae is considered as plesiomorphic. 9. Clypeus with well developed long, longitudinal median groove (as long as clypeus). Some species of the outgroup have a groove but it is usually weak. If it is well developed, it is short and different from that o^ Xenoliphanthus. In one of the species of Tncholiphanthus a poorly developed groove is also present. Most bees do not have a well developed median longitudinal groove; this lack is here considered plesiomorphic. 10. Male clypeus with upper part transversely prominent (Fig. 103), the rest flat and directed backward. This is an autapomorphy for Neoliphanthus. None of the genera in the outgroup presents this character. Most bees have the whole clypeus slightly convex or more or less flattened proximally. In Pseudoliphanthus and Liphanthus s. str. there is a median proximal protuberance (character 15), but the clypeus is not transversely prominent as in Neoliphanthus. 1 1 . Male with the upper part of clypeus below the level of adjacent integument. This is an autapomorphic for Melaliphanthus . Among all the genera examined, none The Bee Genus Liphanthus 293 showed the upper area of the clypeus depressed. Therefore the condition of being at same level or somewhat above the adjacent integument seems to be plesiomorphic. 12. Male with lower part uj clypeus transversely convex, and shiny, rest of clypeus tending to be flattened. This character seems to be synapomorphic for Tricholiphanthus and Melaliphanthus, even though it is less distinct in L. (Tricholiphanthus) leucostomus. The outgroup and many other bees have the distal part of the clypeus more or less convex (but not shining and distinctly transversely convex) which is considered as plesiomorphic. 1.'3. Upper part of clypeus in males dull, this character appears as synapomorphic for Melaliphanthus and Tricholiphanthus. Similar integument of the clypeus is present in one species of Pseudoliphanthus and also in Neoliphanthus; however the shape of the clypeus in these last subgenera is completely different from that of the other 2 subgenera mentioned above. Probably a complex of clypeal characters is demonstrating their relationship. 14. Male with lateral area oj clypeus (ventral view) widened laterally. Autapomorphic for Pseudoliphanthus. The outgroup and other Panurginae present the plesiomorphic condition, that is, clypeus with lateral area widened mesally (ventral view). 15. Alale clypeus with a rounded protuberance, more or less developed, in mid line of its upper part. This character is synapomorphic for Pseudoliphanthus and Liphanthus s. str. In some species it looks like a part of the median ridge, but it seems to be independent of it. It is not found in the outgroups or other Panurginae examined. 16. Male with lower paraocular area and lateral area of clypeus sloping upward to mid part of clypeus and almost at right angles to it. This character is autapomorphic for Leptophanthus . Most of the genera of the outgroup and many other Panurginae have the areas mentioned above at an angle much less than 90°; this is considered plesiomorphic. 17. Male lower face with moderately long and dense pubescence. This feature is au- tapomorphic for Tricholiphanthus. The outgroup and most other Panurginae show very short and sparse pubescence, the plesiomorphic condition. 18. Male with postmandibular area mesally about half or more as long as base of the mandible. This character is autapomorphic for Melaliphanthus. All the other Panurginae examined have this area much shorter than the base of the mandible, a feature here considered plesiomorphic. 19. Male with pronotum protuberant at dorso-lateral angle. This feature is autapomorphic for Tricholiphanthus. The protuberant area is acuminate apically, much less conspicuous in L. pilifrons. In some species of the outgroup and of other Panurginae this area is carinate or a rounded, conspicuous, protuberance. However, in most of the genera the whole dorsal margin of the pronotum is slighdy rounded. In no case is it angulate as in Tricholiphanthus. 20. With 2 submarginal cells. Four genera of the outgroup and most other Panurginae present this character. It also appears in Neoliphanthus as an autapomorphy. The presence of 3 submarginal cells is widespread in bees and wasps; therefore this appears to be the plesiomorphic condition, still retained in one of the genera of the outgroup as well as in most Liphanthus. 21. Hind tibial spurs strongly curved at apices. This feature is autapomorphic for Xenoliphanthus. The whole outgroup and nearly all other Panurginae examined 294 The University of Kansas Science Bulletin have these spurs almost straight or very sHghtly curved. Only in Austropanurgus and in some species of Perdita are the spurs apically curved as in Xenoliphanthus . In one genus (Arhysosage) the entire spurs are more distinctly curved. 22. Male with propodeal triangle laterally less than half length of metanotum (laterally). This character is autapomorphic for Neoliphanthus. The plesiomorphic condition, found in other Panurginae, seems to be the propodeal triangle laterally longer than half of the length of the metanotum (laterally). These measurements are all made parallel to the long axis of the insect's body. 23. Male metasomal sternum III with posterior margin concave in the middle. This character appears as a wide concavity in 2 of the species of Tricholiphanthus, but is distinctly reduced in the third (L. pilifrons). Most of the outgroup and all of the other Panurginae examined have this sternal margin almost straight or somewhat convex, no doubt the plesiomorphic condition. 24. Male ?netasomal sternum IV with posterior margin concave in the middle. This character is autapomorphic for Tricholiphanthus. It is more reduced (almost straight) in L. pilifrons than in the other species of the subgenus. Most of the outgroup and several other Panurginae present the plesiomorphic condition, that is, posterior margin of sternum IV is convex or almost straight. 25. Pygidial plate present in male. This may be a weak character. The plate appears clearly defined by a carina in Xenoliphanthus and Liphanthus s. str. The rest of the subgenera show this structure similar to that of the outgroup and many other Panurginae, that is a bare and shiny area forming a triangle (between the hairy sides), poorly defined, not delimited by carinae, sometimes with a median longitudinal ridge, usually not reaching the posterior margin of the tergum. The pygidial plate in males is widespread in bees and wasps. Because of its poorly developed condition in nearly all Panurginae examined, it is here described as absent when merely represented by a bare area. The presence of this plate in other bees is plesiomorphic, but for Liphanthus is apomorphic. Reversal is possible since the genes for this structure must be retained for the females; the plate could reappear in males merely by turning on the functioning of these genes in males. This apomorphy may have appeared in the ancestor of Liphanthus s. str., Xenoliphanthus, and Pseudoliphanthus, but has been lost in the latter subgenus (Figure 159). It would also be possible for this structure to arise twice independently in Liphanthus s. str. and Xenoliphanthus; the same number of changes (2) would be required in either case. 26. Male metasoma with yellow bands. This is a synapomorphy for Liphanthus s. str. and Pseudoliphanthus. This character is widespread among bees; however is not present in the outgroup or in many other Panurginae. Therefore its absence is here considered as plesiomorphic. thus, and some Tricholiphanthus, which genera. The sinuosity is evidently a could indicate for them a common deri- plesiomorphy and its loss is associated vation. However, it seems that the with the divergence below the inner elongation of the first flagellar segment orbits (characters 5, 6, which are apo- is a meaningful apomorphic charac- morphies). A flat upper paraocular ter which separates Leptophanthus and area is present in Neoliphanthus and clearly groups the rest of the sub- Melaliphanthus. This character appar- The Bee Genus Liphanthus 295 ently is also correlated with the sin- uosity of the inner orbits, already discussed above, and is well marked in these two subgenera. The cladogram (Figure 159) was prepared by hand using the methods ol Hennig (1966). In addition, a com- puter program by J. S. Farris (1970) was used by Mr. Robert W. Brooks at the University of Kansas to prepare a cladogram (Wagner tree). The clado- gram obtained using the computer was like that in Figure 159 but less par- simonious since character 2 appeared Pseudo liphanthus Liphanthus Xenoliphanthus Tricholiphonthus Neollphanthus Meloliphonthus Leptophanthus Fig. 159. Cladogram showing the dadistic relationships of thr subgenera oi Liphanthus. The lengths ofthe lines show the amount of change indicated by the synapomorphies and autapomorphies listed in Table 1 . 1 he angles have n() meaning. X means the loss of an apomorphous character. 296 The University of Kansas Science Bulletin twice, once as a reversal. It is interesting to notice the posi- tion of Leptophanthus in Figure 159, since it presents numerous plesiomor- phic characters. This generahzed con- dition could be related to a broader geographic distribution, as will be indi- cated later. Neoliphanthus, Tricholiphanthus, and Melaliphanthus are separated by a num- ber of apomorphic characters which characterize them clearly. On the other hand, they share important plesiomor- phic characters that make them phe- netically highly similar. Tricholiphanthus and Melaliphanthus seem to be more closely related to each other than to Neoliphanthus (Figure 159). Xenoliphanthus stands out for its dis- tinctive autapomorphic characters, which set it phenetically apart from the rest, especially the strongly curved hind tibial spurs and the scarcely to distinctly concave (rather than convex) vertex. Liphanthus and Pseudoliphanthus are separated from the rest of the sub- genera by three apomorphic charac- ters, which at the same time are conspicuous and give them a general common appearence different from the other subgenera. UlSTRIBUTION The species oi Leptophanthus are dis- tributed from Atacama Province in the north to Arauco Province in the south, also crossing to Argentina (see Maps I and II). It is important to notice that this is the most diversified of the sub- genera, including seven species de- scribed in the present paper. Neoliphanthus is one of the most narrowly distributed groups, since only one species is known; it is from the precordillera of Linares Province. The two species of Melaliphanthus have been found only in the precor- dillera zone of Malleco Province, and like Neoliphanthus have a restricted geo- graphic distribution. Tricholiphanthus , which contains only three species, appears somewhat more diversified than the subgenera mentioned in the preceding paragraph; it is known only in Santiago and Mal- leco Provinces. Xenoliphanthus is represented by four species broadly distributed from Coquimbo Province in the north to Arauco Province in the south. Consid- ering the number of known species, this subgenus is restricted in distribu- tion. Pseudoliphanthus and Liphanthus are sympatric in most of their range. How- ever, Pseudoliphanthus is more broadly distributed to the south, extending from Valparaiso Province to Lacar Lake in southern Argentina. On the contrary, Liphanthus s. str. occupies more northern areas, from Coquimbo Province to Malleco (south-central Chile). The great number of new species described here, and the small number of specimens of some of them, suggest that more new species will be collected and that the distributions of the sub- genera summarized above will be mod- ified. The Bee Genus Liphanthus 297 ytHILE CHICO 72) |70p. '-^|M| LoslfmiU* en wto carto no tientrmf^cUf o ti i Vol. 52, No. t, pp. 301-481 (This concludes Vol. 52] Oct. 1, 1985 ANNOUNCEMENT The University of Kansas Science Bulletin (continuation of the Kansas University Quarterly) is an outlet for scholarly scientific investigations carried out at the University of Kansas or by University faculty and students. 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Byers, Chatrman THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Vol. 52, No. I, pp. 301-481 October 1, 1985 A Taxonomic Revision of the Lygaeinae of Australia (Heteroptera: Lygaeidae) Alex Slater' Department (if Kniomology, University of Kansas Contents Abstract 30 j Introduction 302 Materials and Methods 304 Taxonomic Characters 305 Acknowledgements 307 Key to the Australian genera of Lygaeinae 308 Arocatus Spinola 309 Astacops BoisDUVAL 326 Scopiastella Slater 330 Scopiastes Stal 345 Thunbergia Horvath 370 Woodwardiastes, gen. nov 378 Aspilocoryphus Stal 381 Graptostethus Stal 385 Pyrrhobaphus Stal 398 Melanerythrus Stal 401 Melanotelus Reuter 411 Spilostethus Stal 417 Oncopeltus Stal 426 References Cited 437 Abstract The lygaeid subfamily Lygaeinae is revised for Australia. Keys to genus and species are provided for all included taxa. Each genus and species is described or redescribed in detail. The external male and female genitalia are figured for most species. Drawings of the dorsum of many species arc also provided. Thirteen genera are recognized. The genus Woodwardiastes, gen. nov., is described. Scopiastella Slater is raised from subgeneric to generic rank. Brachylygaeus Gross is synonymized with Melanerythrus Stal. A total of 63 species are recognized. Three species are removed from synonymy: Alelanotelus rubiginosus (Walker), Oncopeltus purpurascens Distant, and Spilostethus pacificus (Boisduval). Six species are reduced to junior synonyms: Graptostethus parvus Distant oi Arocatus contmctus Distant; Lygaeus singularis Walker oi Arocatus rusttcus (Stal); and Lygaeus caslaneipes Dallas. Lygaeus innotatus Dallas, Oncopeltus sordidus var. castaneipes Horvath, and Oncopeltus consors Distant of Oncopeltus quadrigultatus (Fabricius). Twenty-five new species are described: Arocatus chiasmus, Arocatus Jastosus, Arocatus montanus, Graptostethus pubescens. Graptostethus varipictus, Melanerythrus Ignitus, Melanerythrus uptoni, Melanotelus geriae, Scopiastella htmertos, Scopiastella nigrtcoronatus, Scopiastella ruhricinctus, Scopiastella woodwardi. Scopiastes facetus, Scopiastes impeditus, Scopiastes kurandensis, Scopiastes obscurus, Scopiastes perplexus, Scopiastes scudden, Scopiastes taroomi, Scopiastes yorkensis. Thunbergia marcida. 'I hunhrrgia rufifemur. Thunbergia tumida, Woodwardiastes exilis, and Woodwardiastes monteithi. A ncotype is designated for Oncopeltus quadrtguttatus (Fabricius). Parasitization oi Arocatus rusticus (Stal) and Oncopeltus sordidus (Dallas) by the tachinid fly Alophora aureiventris CAirran is reported. Some food plant records are included. ' James A. Slater II. .Alex Slater is used to avoid confusion in the literature with the author's father. Dr. J. A. Slater. 302 The University of Kansas Science Bulletin Introduction The subfamily Lygaeinae may be characterized by: all abdominal spira- cles dorsal; sutures of abdominal ven- ter straight or nearly so, all reaching lateral margin of abdomen; apical mar- gin of corium straight; hemelytra im- punctate; membrane of forewing with distinct basal cell; hindwing with hamus and subcosta; hindwing lacking intervannals; no m-chromosome; phal- lotheca with lateral processes; vesica with ring sclerite; ovipositor with ex- tensive membranous areas; and nym- phal abdominal scent glands between terga IV and V and V and VI. Many members of the Lygaeinae are unusual among the Lygaeidae both in appearance and habitat. Where the "average" lygaeid is a small, cryp- tically colored ground dweller, many (if not most) lygaeines are of medium to large size, boldly patterned in red or orange and black, and are found on or in conspicuously placed plant seed pods. The association of many species with members of the plant family As- clepidaceae has earned the subfamily the common name of milkweed bugs. Many species feed on "poisonous" plants of the families Asclepidaceae, Apocyanaceae, and Solanaciae. This has led to the belief that the bright coloration is aposomatic. Scudder and Duffy (1972) reported the presence of cardiac glycosides in several genera. Avoidance by predators of brightly col- ored species has not yet been demon- strated. The last major classification of the subfamily is that of Stal (1874). Slater (1964a) raised many of Stal's subgen- era to generic level. Unfortunately, as pointed out by Slater (19646) and by Ashlock (1975), Stal's reliance on color characters to define his generic level groups resulted in an identification scheme which tends to obscure the phylogenetic relationships within the subfamily. As a result the generic clas- sification of the Lygaeinae is in need of a complete revision. In this paper I am fairly conservative in the use of generic names, preferring to await study of species from other areas before suggest- ing major changes in classification. As a result of their size, coloration, and habitat the Lygaeinae received more attention from early entomolo- gists than did the majority of lygaeids. The first members of the subfamily to be described from Australia were Mela- nerythrus biguttatus and Oncopeltus quadn- guttatus. Both species were described in the genus Cimex by Fabricius (1775) from material in the Banks Collection. A total of 40 additional species were described by Gmelin (1790), Boisduval (1835), Erichson (1842), Dallas (1852), Stal (1865, 1867, 1874), Walker (1872), Breddin (1900), Distant (1901fl, 1918), and Horvath (1902, 1914). Of these 40 species 25 are still considered distinct. Following Dis- tant's 1918 paper no new species were described from Australia until Scudder (1963) in his revision of the ''Astacops complex" described five species as new. The addition of six species de- scribed from other parts of the world but occurring in Australia and 25 spe- cies described as new in this paper brings the total for the continent to 63. Slater (1964a) listed 39 genera and 414 species in the Lygaeinae. Two genera, Lygaeus and Oncopeltus, are dis- tributed worldwide. The greatest diver- sity in both genera and species occurs in the tropics of both hemispheres. The Australian fauna consists of 63 species in 13 genera. This is roughly compara- ble to the 32 species in 14 genera from South Africa, 35 species in six genera from North America north of Mexico, and 41 species in 12 genera from the Palearctic reported by Slater (19646). The Lygaeinae of Australia 303 The tropical element present in the Australian fauna and largely absent in the other areas accounts for the greater number of species in Australia. The Australian fauna exhibits a high degree of endemism at the species level and virtually no endemism at the genus level. Of the 63 known species 51 are endemic while only one of the 13 known genera is restricted to the conti- nent. The chaotic state of the higher classification of the subfamily makes a detailed analysis of the Australian fauna impossible. However, a few gen- eral statements may be made. Only one genus found in Australia has a worldwide distribution. This is On- copeltus which is widespread in both hemispheres with the majority of its species in the tropics. A group of gen- era including Arocatus, Aspilocoryphus, Graptostethus, Melanotelus , and Spiloste- thus is widespread in the eastern hemi- sphere. Pyrrhobaphus is primarily Oriental and Pacific in distribution. Melanerythrus is centered in Australia with one species recorded from Fiji and Timor and another species from New Guinea and Timor. A final group in- cluding the genera Astacops, Scopiastella, Scopiastes, and Thunbergia reaches max- imum diversity in Australia, New Guinea, and the Wallacian region with some species in the Oriental and Pa- cific regions. Woodwardiastes, the only strictly endemic Australian genus, also belongs to this group. At the specific level one species, Graptostethus servus (Fabricius), is wide- spread in the eastern hemisphere. Two species, Spilostethus hospes (Fabricius) and Pyrrhobaphus leucurus (Fabricius), are widespread in the Oriental and Pacific regions. Arocatus continctus Dis- tant displays a curious disjunct dis- tribution, being found in western Africa, India, and Australia. The re- maining extralimital species are shared with New Guinea and/or the Wallacian region with the single exception of Arocatus rusticus (Stal) which is found on Lord Howe Island, Norfolk Island, and in New Zealand. Within Australia the most interest- ing genera geographically are Me- lanerythrus, Scopiastella, and Scopiastes. Melanerythrus contains five species, all pritnarily or entirely Australian in dis- tribution. Of the five, three (two mi- cropters and an extremely vagile macropter) are Eyrean, one micropter is restricted to high altitudes in south- eastern Australia, and one macropter- ous species is Torresian in Australia and also found in New Guinea and Timor. The other two genera are Tor- resian in their Australian distribution. Most species are associated with rain- forest. Both genera show evidence of recent speciation in the presence of several closely related pairs of species where one species is northern and the other southern in distribution. Despite their high visibility little is known about the life history and biol- ogy of the Australian Lygaeinae. Food plant records are very scarce, especially for the ''Astacops complex" which com- prises about half the fauna. The sub- family does not appear to be of economic importance in Australia de- spite reports of Oncopeltus sordidus (Dal- las) on cotton in New South Wales and of Graptostethus servus (Fabricius) on var- ious crops in northern and western Australia. Label data for the type series of Scopiastes perplexus, sp. nov. , indicates them as "playing havoc among potato plants" and a series of Melanerythrus higuttatus (Fabricius) was taken on maize and on "Graminae" in northern Queensland. This may indicate poten- tial for at least local damage to food crops. An interesting aspect of lygaeinc bi- ology is an apparent fondness for nee- 304 The University of Kansas Science Bulletin tar. Many specimens of several genera have been collected on eucalyptus flow^- ers by Dr. E. M. Exley and her assis- tants. It does not seem likely that they were feeding on other parts of the trees. Other records are from the flowers of various rainforest vines and the flowers of milkweed plants {Gomphocarpus sp.). Graptostethus varipictus, sp. no v., was collected on roses in northern Queens- land by R. I. Storey. Materials and Methods Specimens were examined from the following collections: Australian Mu- seum (AM), Australian National In- sect Collection (ANIC), University of Queensland (UQ), Queensland Mu- seum (QM), Musee d'Histoire Na- turelle de la Hongrie (Budapest), British Museum (Natural History) (BMNH), Naturhistoriska Riksmuseet (Stockholm), Institut fiir Pflanzen- schutzforschung, Abteilung Taxono- mie der Insekten (Berlin), Department of Primary Industries, Queensland (DPI), West Australian Museum (WAM), South Australian Museum (SAM), New South Wales Department of Agriculture (NSWDA), Tasmanian Museum (TM), National Museum of Victoria (NMV), and Bernice P. Bishop Museum (Bishop). Only general data on distribution are given in this paper. Complete infor- mation on label data and depository for all specimens examined is available from the author. Complete label data is supplied for all holotypes and for para- types of newly described species. Complete synonymies are not pro- vided for either genera or species. Original descriptions, alternate names to which the species have been referred in the literature, and references to the locations of the taxa in Slater's (1964a) catalogue are given for all taxa. Papers of nomenclatoral or biological signifi- cance published since 1964 are also included. As many of the taxa treated here are not common in collections, and were described at a time when a 12-line description was generally considered more than adequate, fairly detailed de- scriptions are included for both genera and species. To avoid repetition those characteristics common to all species within a genus are included in the generic description. As both generic and species descriptions are based on Australian species, they should be ap- plied to extralimital specimens only with great caution. Measurements were made with an ocular micrometer in a binocular dis- secting microscope. Unless otherwise indicated ten specimens of each sex were measured. All measurements are given in millimeters with the mean of each measurement followed in paren- theses by the range. In some cases, where very few specimens were avail- able, measurements for all specimens examined are included. All measure- ments except those of the antennae and labium were made with the hemelytra parallel to the plane of focus of the microscope. Where the head or pro- notum could not be completely in focus at one setting, focus was taken at the midpoint between the two extremes. As the head is often differently exserted on different specimens, this method re- sults in considerable variation in meas- urements of head length. Much less manipulation of the specimen is re- quired than with other methods and a good idea of the general shape of the insect is conveyed. This is especially true where both head and pronotum are declivent. Measurements of head and pronotal length made with the plane of focus parallel to their longest dimensions would give the impression The Lygaeinae of Australia 305 of a longer structure than appearance of the whole insect conveys. Both male and female genitalia were removed from the insects after they were relaxed in a solution modified from the formula for Barber's Fluid by substituting toluene for benzene. The genitalia were grasped with watch- makers' forceps and pulled free of the abdomen. They were then soaked in concentrated "Dekon 90," a liquid surfactant used to clean laboratory glassware, for from ten minutes to overnight before being transferred to distilled water for further dissection. Muscles and fatty tissue were removed from female genitalia using watchmak- ers' forceps, probes made from minu- ten pins, and a fine sablehair brush. The two valves were separated by gently pulling with watchmakers' for- ceps. The aedeagus of the male was inflated using Ashlock's (1957) osmotic method substituting "Dekon 90" for potassium hydroxide. It was found that the delicate membranes of the lygaeine aedeagus were less likely to break with this change. Drawings of genitalia were made with the aid of a squared grid in a binocular dissecting microscope. The pygophore was oriented so that the anterior opening was perpendicular to the microscope's plane of focus. Clasp- ers were drawn from three angles, pos- terior and interolateral views with the opening at the base of the clasper per- pendicular to the plane of focus and dorsal view with the basal opening parallel to the plane of focus. Complete ovipositor valves were prepared for drawing by tearing the membrane con- necting the valvifers only and then spreading and flattening the ovipositor, resulting in an approximately lateral view of valvifers and valvulae and a ventral view of the membrane connect- ing the valvulae and the median scle- rite. Ovipositor valves were pinned to a wax-bottomed dissecting dish with mi- nuten pins and small parts of the male genitalia supported on cotton wool im- mersed in glycerine in an excavated block. The excavated block allows a modified form of dark field illumina- tion obtained by shining a microscope light through the side of the block. This is very helpful in observing hairs and sculpture on smaller parts of the geni- talia. Dorsal views of entire insects were made either with the aid of a squared grid or a camera lucida. A scale line equivalent to one millimeter is in- cluded with each dorsal view. Taxonomic Characters Until now the identification of Ly- gaeinae has been based primarily on their color pattern. This approach has been reasonably successful in species identification as many species, even those with wide geographic ranges, have very consistent color patterns. However, other species are very var- iable in coloration and many struc- turally distinct species are very similarly patterned. There is some in- dication that the coloration of some species is affected by environmental factors. Specimens of Spilostethus hospes (Fabricius) from northern and central Australia resemble specimens from southern India while those from .south- eastern Queensland and New South Wales resemble specimens from Paki- stan. Specimens from hotter and/or drier areas are lighter colored than those from damper and/or cooler areas. In this paper I have tried to include both color and structure in keys to genera and species. Color characters are often easier to see and to explain while structural characters are gener- ally more reliable. In case of doubt reference should be made to the figures 306 The University of Kansas Science Bulletin of male and female genitalia as these structures are definitive at species level. The keys, as with the de- scriptions, are based on Australian specimens only and should be used cautiously if at all on extralimital speci- mens. Such structural characters as the rel- ative lengths of antennal or labial seg- ments, degree of development of the ostiolar peritreme, shape of head, punctation and shape of pronotum, and type and location of pubescence are useful in identification at both genus and species level. In addition to these characters I have found the struc- ture of both male and female genitalia to be of great use in identification of species and in delimitation of higher categories. The aedeagus is most useful at ge- neric level. Such features as the apical process of the secondary gonopore, de- velopment of the ring sclerite, and presence and shape of conjunctival and vesical lobes are usually consistent within a genus. The general shape of the ejaculatory reservoir and phallothe- cal processes remains constant within a genus while details of their structure vary between species. The claspers are perhaps the best means of confirming the identity of a male specimen. They are almost always unique to a species though the differ- ences between species are sometimes hard to describe or draw. Ideally, direct comparison between claspers should be used when dealing with closely related species. In light of the difficulties in portraying the true shape of the clasp- ers I have tried to avoid using them in the keys. I have attempted to show their shape by drawing each one from three different angles, the three angles related to one another by rotation of the clasper in one plane or another. The pygophore is generally similar within a genus, with structural details of the areas around the dorsal opening frequently varying from species to spe- cies. The shape of the lateral margins of this opening and the median projec- tion of the posterior margin provide especially useful characters. The shapes of these areas are often corre- lated with the structure of the claspers and the phallothecal processes. The female genitalia have been little used in the identification of Lygaeidae in general. For the most part only broad surveys of higher taxa have in- cluded information on their appear- ance, though Scudder (1963, 1965) used the spermatheca at both generic and specific levels. I found them to be of great taxonomic value in this study. The spermatheca especially provides excellent generic and often specific characters. Within the genus Scopiastes the structure of the spermatheca was very useful in determining relation- ships of several very similarly colored species and first led me to suspect that there might be a series of north-south species pairs in the genus. The ovipositor was also very useful taxonomically at both genus and spe- cies level. The general shape of the ovipositor and the shape and develop- ment of the median sclerite of the membrane connecting the first val- vulae is for the most part consistent within genera. The exact shape of the valvifers and valvulae and details of the shape of the median sclerite often vary between species. The shape of the apex of the second valvula was found to be an especially good character at the species level. In the difficult genera Scopiastes and Scopiastella this structure is sometimes of better diagnostic value than the claspers. In addition to their value in identifi- cation of genera and species I believe that the external genitalia will be of Tmk Lygaeinae of Australia 307 great value in understanding phyloge- netic relationships among the Lygaei- nae of the world. Scudder & Duffy (1972) reported the presence of a series of glands opening dorsolaterally on the niesothorax, metathorax and abdominal sterna II through VII of 28 lygaeine species be- longing to several genera. In the new world species Oncopeltus fasciatus, O. san- darachatus and Lygaeus kalmii the secre- tions of these glands were shown to contain higher concentrations of car- denolides than did the hemolymph. As species in some genera they examined did not possess these glands the charac- teristic should prove useful in phy- logenetic studies. The gland openings are most easily seen in specimens mac- erated in potassium hydroxide. Those of the abdominal sterna, which appear as short curved scars at the antero- lateral angles, are often visible on un- treated specimens. In this paper the presence or absence of these dorsolat- eral glands is noted in that part of each generic description concerned with the abdominal sterna. Acknowledgments This work was financed by a grant from the Australian Biological Survey to Dr. T. E. Woodward, Department of Entomology, University of Queens- land. I am indebted to the authorities of the following institutions who willingly lent specimens, many of them types: Australian Museum, Sydney; Austra- lian National Insect Collection, Can- berra; Bernice P. Bishop Museum, Honolulu; British Museum (Natural History), London; Department of Pri- mary Industries, Brisbane; Institut fiir Pflanzenschutzforschung, Abteilung Taxonomie der Insekten, Berlin; Mu- see d'Histoire Naturelle de la Hongrie, Budapest; National Museum of Vic- toria, Melbourne; Naturhistoriska Riksmuseet, Stockholm; New South Wales Department of Agriculture, Sydney; Queensland Museum, Bris- bane; South Australian Museum, Ade- laide; Tasmanian Museum, Hobart; University of Queensland, Brisbane; and West Australian Museum, Perth. I am especially indebted to Mr. W. D. Dolling of the British Museum (Natural History) for providing excel- lent color slides of many of the type specimens in that collection. Special thanks also to the members of the Department of Entomology, University of Queensland, who made my stay in Australia so pleasurable. Brian Cantrell of DPI kindly identi- fied the tachinid fly Alophora aureiventris Curran. Food plants were identified by the staff of the Queensland Herbarium with the exception of Craspedia mac- rocephala which was identified through the help of M. S. Upton of CSIRO, Canberra. Dr. P. D. Ashlock, Department of Entomology, University of Kansas, and Dr. T. E. Woodward, Department of Entomology, University of Queens- land, read the entire manuscript and provided helpful comments. Parts of the manuscript were read and com- mented on by Drs. G. W. Byers and C. D. Michener, University of Kansas. For the fine dorsal views of many of the species I thank J. Hartshoorn, J. E. McDonald, S. R. Monteith, G. T. Thompson, and G. Yeo. My very special thanks go to Dr. T. E. Woodward for giving me the chance to work on the Australian Lygaeinae and for his help and encouragement throughout the project. Lasdy but by no means least I would like to thank my wife Geri for compil- ing distribution data and making the distribution maps and for her support 308 The University of Kansas Science Bulletin and encouragement throughout the project. Key to the Australian Genera OF Lygaeinae 1 . Eyes carried on distinct stalks . . 2 r. Eyes not stalked, at most head swollen behind eyes 5 2. Posterior pronotal lobe with at most a few scattered punctures . . 3 2'. Posterior pronotal lobe at least moderately strongly punctate ... 4 3. Posterior pronotal lobe shiny; head across eyes much narrower than pronotum at posterior mar- gin Astacops Boisduval 3'. Posterior pronotal lobe dull except shiny posterior and lateral margins; head across eyes about equal in width to pronotum at posterior margin Scopiastella Slater 4. Antennal segment I elongate, at least V3 as long as width of head across eyes; bucculae acutely an- gulate anteriorly Woodwardiastes, gen. nov. 4'. Antennal segment I at most 'A as long as width of head across eyes; bucculae usually rounded, at most obtusely angulate anteri- orly Scopiastes Stal 5. Scutcllum tumid, at most with a distinct longitudinal median carina 6 5'. Scutellum not tumid, with a T- shaped median carina and well defined lateral foveae 7 6. Posterior margin of pronotum produced caudad on either side of scutellum; lateral pronotal margins covered with long, stout hairs Oncopeltus Stal 6'. Posterior pronotal margin nearly straight, not produced caudad to either side of scutel- lum; pronotum laterally without long, straight hairs Melanerythrus Stal 7. Lateral scutellar foveae com- posed of several confluent punc- tures 8 7'. Lateral scutellar foveae not composed of confluent punc- tures, floor of foveae smooth or with transverse striations 9 8. Profemora of both sexes with a stout subapical spine; antennal segment IV much longer than II Thunbergia Horvath 8'. Profemora unarmed; antennal segment IV at most slightly longer than II. . . . Arocatus Spinola 9. Metathoracic scent gland open- ing auriculate (ostiolar peritreme considerably raised above sur- rounding pleural surface) 10 9'. Metathoracic scent gland opening not auriculate (ostiolar peritreme scarcely differentiated from surrounding pleural sur- face) Spilostethus Stal 10. Clavus and corium covered with long upright hairs strongly curved apically Melanotelus Renter 10'. Clavus and corium usually without long upright hairs, if such hairs present then slightly and evenly curved from base to apex 11 11. Labial segment III extending beyond metacoxae; pronotum black except lateral margins be- hind calli Pyrrhobaphus Stal 11'. Labial segment III not extend- ing beyond metacoxae (labial segment IV sometimes extending that far); midline of pronotum behind calli red or orange .... 12 12. Membrane of forewing with a white discal macula; para- and mediotergites of abdominal seg- ments II to VI distinct Aspilocoryphus Stal 12'. Membrane of forewing with- out discal macula; para- and The Lygaeinae of Australia 309 mediotergites of abdominal seg- ments II to VI fused Graptostelhus Stal Arocatus Spinola Arocatus Spinola, 1837:257; Slater, 1964a: 18-19; Slater and Sperry, 1973:130; Hamid and Meher, 1973a:36. Tetralaccus Fieber, 1860-1861:44, 164. Xerophagius Kiritshenko, 1950:350. Type-species Lygaeus melanocephalus Fabricius 1798, monobasic. Diagnosis. — Antennal segment IV not or but slightly longer than II; eye separated from anterior pronotal angle; head swollen behind eye; paratergites II to VI distinct; female paratergite VII fused with mediotergite; male par- atergite VII fused with laterotergite; femora unarmed. The only Australian Lygaeinae with which Arocatus might be confused are Thunbergia, Astacops, Scopiastes, Scopia- stella, and Woodwardiastes. It differs from the first by lacking a subapical spine on the fore femur and from the latter four by not having distinctly stalked eyes. Hamid and Meher (1973a) credit the genus with 24 species of which 19 are from the Palearctic region and only two from Australia. Slater and Sperry (1973) consider it to be composed of 14 species, nine with a Palearctic distribu- tion. In this paper six Australian spe- cies are recognized. Four of these species are apparently endemic, one is also found in New Zealand and Lord Howe and Norfolk islands, and one has a curious disjunct distribution, occur- ring in western Africa and India as well as Australia. As presently constituted Arocatus is a rather heterogenous group. In most lygaeine genera such characters as the extent and placement of granulate dull areas, the shape of the abdominal ter- gites, and the basic appearance of the genitalia of both sexes are constant. This is not the case in Arocatus. The Australian members of the genus vary in these and other charac- ters. The six species may be split into four groups primarily based on gen- italic differences. Tables I and II illustrate the differences and interre- lationships between the groups through an analysis of the distribution of 28 characters. These characters were taken from Australian species within the groups. The list is biased towards differences rather than similarities be- tween groups and may give the impres- sion that they are less similar in appearance than is in fact the case. As three of the groups are represented by only one species some of the character states may be of only specific value. Despite these limitations I believe that at least the large difference between the combined groups I and II on the one hand and III and IV on the other indicate two distinct lines of evolution- ary development. Group I contains /I. rusticus (Stal), A. aenescens Stal, A. chiasmus, sp. nov. , and an undescribed species from New Cal- edonia which is very similar to rusticus and chiasmus in general appearance. Group II conidiins A. fastosus, sp. nov., and at least one undescribed species from New Guinea. Group III contains A. montanus, sp. nov., a very similar undescribed species from New Guinea, and probably A. rubromarginatus (Dis- tant) from New Caledonia. Group IV contains A. continctus Distant and a species from the Ryukyu Islands tenta- tively identified as A. fasciatus jakovlev. I have examined males of five Aroca- tus species from Africa and the Palearc- tic and find that they are most similar to group III. However there is consid- erable variation in many characters. As 310 The University of Kansas Science Bulletin Table I. Distribution of selected characters in species groups within the Austrahan members of Arocatus Spinola. Group Character I H III IV + + + + + + + + + + + + Dull pronotal area extending onto posterior lobe — — + + Median projection of pygophore short and nar- row Median projection of pygophore with a tablike process Membrane connecting first valvulae of ovipositor with median and sublateral sclerites + + — — First valvulae reaching apex of connecting mem- brane — — -I- -I- Conjunctiva with subapical dorsal lobe Secondary gonopore flared Membrane connecting second valvulae of ovipositor cleft medially Blade of clasper narrow, rounded in cross section Posterolateral metapleural angle distinctly pro- duced — — — + Posterior margin of abdominal sternum II convex laterally + + + — Vesica greatly expanded laterally at ring sclerite — — — -t- Vesica forming a cup distally into which retracted gonoporal process fits -I- -I- -I- — Spermatheca tightly coiled just basal to apical bulb ^ - - - + Gonoporal process coiled when aedeagus inflated -I- -f- -I- * — Length scutellum compared to length claval com- missure First valvulae notched subapically Labium reaching at least to metacoxae Dorsal opening of pygophore with depressed "shelf on lateral margins Head less than half as long as wide Posterior margin of pygophore evenly convex or nearly so — -I- -I- -I- Gonoporal process constricted subapically -I- — — — Tergum IX of male impressed medially -I- — — — Tergum IX of male distinct from surrounding membrane -I- — -f + Meso- and metapleura punctate -f- — + + Median hypandrial projection deeply excavated apically — -I- — — Median length female abdominal sternum VI compared to lateral length % 'A Ve 'A 1.25 1.25 2.5 2.0 -1- + * * _ * * * + + — + — — -1- — — — -1- — The Lygaeinae of Australia 3\ Tablk I. (Continued.) Character Caudolateral lobe of clasper present + - Group II III IV + + + + = character as stated. + + = character very strongly developed. H — = character much reduced. — = character not as stated. * longer and more tightly coiled than in groups II and III. ** unmodified. *** produced. Table II. Analysis of character distribution in species groups within the Australian members oiArocatus Spinola. Gi roup I II III IV I ^^^8 20 8 6 Number of II 12 ^^^^ 8 6 Number of characters ^^ characters restricted III 0 0 16 shared by to IV 0 0 10 V! females were not available for these species and as a detailed study of extra- Australian species is beyond the scope of this revision I have not attempted to formalize any phylogenetic conclusions based on analysis of the Australian species. Redescription. — Structure: Moder- ately elongate, parallel sided. Thoracic venter, extreme base of scutellum and often pronotum anteriorly finely gran- ulate giving a dull appearance. Entire head and body usually with inter- spersed semidecumbent, moderately long and longer upright hairs, the latter shortest on head, posterior pronotal lobe, and hemelytra, longest on venter. Head scarcely to moderately declivent, vertex nearly flat, produced laterally and at least slightly swollen behind eye; ocelli closer to eyes than to each other, about level with vertex; eye removed from anterior pronotal angle, directed slightly dorsolaterally; bucculae slightly to moderately produced ven- trally, at least slightly convex, not reaching anterior prosternal margin. Antennal segment I surpassing ante- clypeus by less than half its length, segment IT or IV longest. Labial seg- ment I thickest, each succeeding seg- ment slightly thinner than that preceeding it. Pronotum subquadrate; narrower anteriorly; punctate except callus and extreme base; impressed and constricted behind callus; anterior margin concave; posterior margin con- vex; lateral margin slightly convex an- teriorly, directed anteromedially and nearly straight posteriorly; posterior margin obliquely impressed lateral to scutellum; median carina at most faindy indicated behind callus; callus 312 The University of Kansas Science Bulletin moderately swollen, slightly oblique, reaching or almost reaching lateral margin at anterior angle; callar impres- sion oblique, sinuate, unbranched. Scutellum wider than long (occasion- ally very slightly); bluntly acute api- cally; T-shaped carina present; lateral fovea deeply, coarsely punctate. Hemelytra not completely covering lat- erotergites except apically; costal mar- gin nearly straight basally, slightly convex and produced on apical half; veins prominent; membrane surpass- ing tip of abdomen. Prosternum coarsely punctate, anterior margin slightly raised. Meso- and metasterna impunctate, posterior margins raised, mesosternum broadly, shallowly sul- cate medially. Propleuron punctate. Mesopleuron at least moderately punc- tate. Metapleuron at least moderately punctate; ostiolar peritreme well devel- oped; posterolateral angle not or slightly produced posteriorly; posterior margin straight or slightly concave, usually directed dorsoventrally, parallel to posterior mesopleural margin. Ab- dominal sterna without dorsolateral scent gland openings; II not covered by metapleuron; VI of female shorter me- dially than laterally; VII of female cleft to base, slightly concave laterally; VII of male concave. Abdominal terga with paratergites III to \T distinct, VII of male fused with laterotergites, VII of female fused with mediotergite; ap- odemes on paratergites when present. Pygophore subcircular in cross section; median projection present. Clasper variable. Aedeagus with moderately heavily pigmented phallothcca; phal- lothecal process curved, slightly pro- duced dorsad; conjunctiva elongate, tubular; ejaculatory reservoir com- plete, Y-shaped basal projection pres- ent; vesica basally moderately long, twisted, without lobes; ring sclerite elongate, incomplete; vesica distally at least slightly flattened, slightly pro- duced laterally, coiled, often forming cuplike receptacle for retracted gono- poral process; gonoporal process at least partly covered by visible mem- brane; secondary gonopore flared or oblique, without apical process. Ovi- positor laciniate, variously modified. Spermatheca with pigmented apical bulb and tube distad of sperm pump; sperm pump and basal tube unpig- mented. Key to the Australian Species o/" Arocatus 1. Meso- and metapleura at most indistinctly punctate; costal mar- gin of corium orange fastosus, sp . nov. r. Meso- and metapleura dis- tinctly punctate; at least center of costal corial margin black or brown 2 2. Labium not exceeding meso- coxae; hemelytra nearly unicol- ored; anterior pronotal lobe reddish orange . . montanus, sp. nov. 2'. Labium extending at least to metacoxae; if hemelytra uni- colored then anterior pronotal lobe brown or blackish brown ... 3 3. Scutellum about twice as long as claval commissure; anterior pro- notal lobe dark brown to black; posterior pronotal lobe orange medially and on lateral margins continctus Distant 3'. Scutellum about IV4 times as long as claval commissure; if an- terior pronotal lobe brown then posterior lobe also brown 4 4. Pronotum and hemelytra uni- colored (brown) aenescens Stal 4'. Anterior pronotal lobe reddish orange; posterior lobe dark brown to black 5 5. Labium reaching abdominal sternum IV; thoracic pleura completely orange . . . rusticus (Stal) The Lygaeinae of Australia 313 5'. Labium reaching only abdomi- nal sternum III; thoracic pleura almost completely brown .... chiasmus, sp. nov. Arocatus fastosus, sp. nov. Figures 1; 7a; 46c; 47g, h; 48k, 1; 49h, i; 50J-1; 5 If; 52c, d; 53j. Types.— Holotype i, "Heathlands, Y- junction, Cape York Peninsula, N. Qld., 5/12-II-1976. G.B. Mon- teith." in QM. Paratypes: \\ 2$, same data as holotype, in UQ^; l?, "Lockerbie area, Cape York, N. Qld., 13/27-IV-1973, G.B. Mon- teith." in UQ. Fig. 1. Arocatus foilosus, sp. nov., dorsal view. Scale = 1 mm. Description. — Color: Orange, heme- lytra and abdominal venter medially darkest. The following dark brown with brilliant greenish iridescence: pos- terior pronotal lobe except posterior margin; apex and fovea of scutellum; clavus except base; corium except base and broad costal margin; and hemely- tral membrane. The following dark brown to black: prosternum laterally; thoracic pleura except acetabulae, lat- eral and posterior margins of pro- and mesopleura; ostiolar peritreme; an- tenna; and labium. Legs except coxae light brown. Eye brown. Ocellus red. Abdominal venter mediolaterally and mesosternum laterally sometimes suf- fused with brown. Hemelytral mem- brane except base subhyaline brown, becoming paler apically. Structure: Elongate, narrow, flat- tened posteriorly. Hairs shorter than in other members of the genus, longer upright hairs almost absent except on callus where short semidecumbent hairs absent. Head scarcely declivent, moderately swollen behind eye; poste- rior margin of ocellus about level with that of eye; buccula moderately pro- duced, convex anteriorly, very low, slightly concave posteriorly. Antennal segment I surpassing anteclypeus by about 'A its length. Labial segment I slightly surpassing anterior prosternal margin, II slightly surpassing procoxa, III reaching middle of metacoxa, IV reaching slightly beyond middle of ab- dominal sternum III. Pronotum shal- lowly punctate, anterior lobe including callus finely granulate giving a dull appearance. Scutellum about 1 'A times as long as claval commissure, subacute apically. Propleuron coarsely, shallowly punctate anteriorly and posteriorly, impunctate centrally. Mesopleuron shallowly punctate posteriorly, im- punctate anteriorly. Metapleuron im- punctate; posterior margin slightly 314 The University of Kansas Science Bulletin concave, directed dorsoventrally; pos- terolateral angle not produced. Ab- dominal sternum II with posterior margin convex laterally; VI of female about Vs as long medially as laterally. Abdominal dorsum as in Fig. 46c; mediotergites IV and V elongate and narrowed to their common margin. Pygophore as in Figs. 47g, h; convex posteriorly; median projection concave apically, tablike apical process present; tergum IX indistinctly separated from surrounding membrane, not impressed medially. Clasper as in Figs. 50j-l; no caudolateral lobe; blade rounded in cross section, curved and twisted; shank very shallowly excavated. Ae- deagus similar to A. rusticus (Fig. 49e) but smaller in relation to body size; phallotheca and phallothecal process as in Figs. 48k, 1; ejaculatory reservoir as in Figs. 49h, i; ring sclerite elongate, strongly convex basally; gonoporal process not strongly constricted sub- apically; secondary gonopore slightly flared, slightly oblique. Ovipositor as in Figs. 52c, d, 53j; membrane of first valve shallowly notched medially. Sper- matheca as in Fig. 5 If; sperm pump very near base. Measurements: Holotype ^ . Length 11.0; abdominal width 3.1; head length 1.44; width across eyes 2.20; pronotal length 1.66; pronotal width 3.00; scutellar length 1.68; scutellar width 1.84; length antennal segments I .62, II 2.31, III 2.00, IV 2.48; length labial segments I 1.72, II 1.64, III 2.06, IV 1.20. Measurements paratypes. \S, 2$. Length S 10.1, v 10.8, 10.7; abdomi- nal width ' 2.8, V 2.9, 2.9; head length i 1.38, ? 1.54, 1.56; width across eyes ^ 2.08, ? 2.12, 2.20; pro- notal length ' 1.72, ■■ 1.70, 1.76 pronotal width ' 2.94, . 2.92, 3.04 scutellar length ' 1.50, i 1.52, 1.52 scutellar width t 1.66, V 1.68, 1.72 length antennal segments I t .60, $ .65, .65, II ^ 2.35, * 2.36, 2.42, III S 2.04, V 2.18, 2.11, IV $ 2.52, ? 2.62, 2.68; length labial segments I c? 1.48, ? 1.48, 1.68, II i 1.38, $ 1.60, 1.58, III $ 1.86, $ 2.04, 2.00, IV $ 1.14, ? 1.18, 1.14. Comments. — This species is easily separated from the other members of the genus by its large size and com- pletely orange costal hemelytral mar- gins. It is somewhat similar in appearance to Thunbergia marcida, sp. nov. , but lacks subapical femoral spines. There is at least one closely related species in New Guinea with completely dark hemelytra. The holotype and two paratypes were taken on the flowers of a rain- forest vine (G. B. Monteith, pers. comm.). The species is known only from the type locality (Fig. 7a). Arocatus montanus, sp. nov. Figures 2a; 3; 46d, g, h; 48i, j; 49k, 1; 50a-c; 51c; 52a, b; 53k. Types.— HoXoiy^Q $, "Bunya Moun- tains, Qld., 22-1-1938, N. Geary, 200 ft." in AM. Paratypes: \$, "Wallacia, III-1932"; 1', "Mt. Wilson, N.S.W., 1-1929, H.J. Car- ter"; li, "Lamington National Park, 14/20-11-1958, I.C. Yeo" in AM and UQ. Description. — Color: Orange. The following dark brown to black: head above; subquadrate postcallar macula on either side of pronotum; crossbar of scutellar carina; mesosternum; meta- sternum; abdominal sternum VII me- dially; antenna; labium; and legs. Most of posterior pronotal lobe and scutellum brown. Coriacious parts ol hemelytra purplish brown becoming almost black apically. Eye brown. Ocellus red. Hemelytral membrane The Lygaeinae of Australia 315 montanus continctus Fig. 2. a. Arocatus montanus, sp. nov, , dorsal view. b. A. coiitinctus UistaiU, dorsal view. Scale = 1 mm. subhyaline; dark brown basally becom- ing paler apically. Structure: Fairly broad, deep bod- ied. Head moderately declivent, mod- erately swollen behind eye; posterior margin of ocellus level with or slightly posterad of posterior margin of eye; buccula moderately prominent, con- vex, tapering gently posterad. Anten- nal segment I surpassing anteclypeus by about Vm its length. Labial segment I not or just reaching anterior prosternal margin, II reaching about middle of procoxa. III slightly surpassing pro- coxa, IV reaching mesocoxa. Prono- tum very coarsely punctate; scarcely impressed or constricted behind callus; entire anterior lobe and about anterior half of posterior lobe finely granulate giving a dull appearance. Scutellum about 2'/2 times as long as claval com- missure; subacute apically. Hemelytra mesad of radius finely granulate giving a dull appearance. Thoracic pleura deeply, coarsely punctate except ace- tabular regions. Metapleuron with pos- terior margin nearly straight, directed dorsoventrally; posterolateral angle not produced. Abdominal sternum II with posterior margin convex laterally; \'II of female about 'A, as long medially as laterally. Abdominal dorsum as in Fig. 46d; mediotergites not elongate or narrowed; VII of both sexes with para- tergites apparently fused with medi- otergite. Pygophore as in Figs. 46g, h; convex posteriorly; median projection scarcely produced, narrow, without tablike apical process. Claspcr as in Figs. 50a-c; caudolaterai lobe well dc- 316 The University of Kansas Science Bulletin veloped; blade flattened, broad basally, gradually narrowed to apex; shank shallowly excavated. Aedeagus some- what similar to A. rusticus (Fig. 49e); phallotheca and phallothecal process as in Figs. 48i, j; conjunctiva elongate, tubular, without lobes; ejaculatory res- ervoir as in Figs. 49k, 1; vesica basally short, thick; ring sclerite heavily pig- mented and sclerotized, incomplete, elongate, narrow, extremely convex basally, nearly forming a tube around vesica; vesica distally short, thick, scarcely produced laterally at base, dis- tally forming cup with pigmented outer margins; gonoporal process coiled, moderately elongate (about 3'/2 coils), visibly covered with membrane for about 'A its length, sperm duct narrow- ing gradually to apex; secondary gono- pore not flared or produced, oblique. Ovipositor as in Figs. 52a, b, 53k; first valvula reaching apex of connecting membrane, unmodified; connecting membrane of first valve broadly cleft apically, without sclerites; second val- vula narrow, subacute apically; con- necting membrane of second valve with median sclerite, not cleft. Spermatheca as in Fig. 51c; sperm pump much nearer apex than base. Measurements: Holotype 5. Length 6.9; abdominal width 2.2; head length .66; width across eyes 1.60; pronotal length 1.34; pronotal width 2.24; scu- tellar length 1.18; srutcllar width 1.20; length antennal segments I .36, II .91, III .88, 1\' 1.16; length labial segments I .66, II .64, III .36, IV .50. Measurements paratypes. 2S , 19. Length ^ 6.3, 6.7, > 6.8; abdominal width ' 1.9, 2.2, V 2.3; head length S .56, .64, J .64; width across eyes S 1.50, 1.60, ■■ 1.56; pronotal length ^ 1.16, 1.40, I 1.36; pronotal width S 2.04, 2.24, . 2.26; scutellar length S 1.08, 1.16, X 1.20; scutellar width S 1.08, 1.26, i 1.28;. length antennal segments I ' .34, .37, i .36, II S .92, .97, i .95, III _^ .93, J .90, IV S 1.20, $ 1.16; length labial segments I S .60, .66, ? .68, II S .58, .62, ^ .60, III S .34, .34, 2 .30, IV S .48, .56, ? .56. Comments. — The extremely short la- bium separates Arocatus montanus from all other Australian members of the genus. The dorsal color pattern is rem- iniscent of that of^. melanocephalus from the Palearctic region. A very closely related undescribed species from New Guinea differs from A. montanus in that it is larger, the hemelytral membranes are relatively longer and more acute apically, the punctations are sparser and deeper, the antennae are longer and comparatively thinner, the meso- and metapleura are infuscated, the ab- dominal venter is completely orange, and labial segments I and II are or- ange. From the few specimens examined A. montanus seems to be restricted to upland rainforest areas in New South Wales and eastern Queensland (Fig. 3). Nothing is known of its biology. Arocatus continctus Distant 1906 Figures 2b; 3; 46e; 48g, h, m, n; 49a, b, j; 50d-f; 51b; 52e, f; 531. Arocatus continctus Distant, 1906:410; Slater, 1964^:20, 1964/^:57; Hamid and Meher, 1973a:36; Slater, 1978: 856. Graptostethus parvus Distant, 1918:422; Slater, 1964a:55. syn. nov. Types. — Lectotype 2 of ^4. continctus, "Eppawela, N.C.P. 9-05, 1888, Dis- tant Coll., 1911-383" in BMNH. Paralectotypes of ^. continctus, 3^, same data as lectotype, in BMNH. Type i^ of G. parvus, "Townsville, Qld., Julv-03, F.P. Dodd." in BMNH. Redescription. — Color: Orange. The The Lygaeinae of Australia 317 ^ <^, • Acontinctus o A.montanus -L Fig. 3. Distribution of Arocatus continctus Distant and A. monlanus, sp. nov. in Australia. following dark brown to black: median longitudinal dorsal vitta of head; ante- rior pronotal lobe; broad longitudinal vitta on either side of posterior pronotal lobe, connecting with anterior lobe; scutellum except narrow median longi- tudinal vitta and apex; clavus except margin of claval suture; diffuse tri- angular macula along costal corial margin not reaching either base or apex, broadest opposite claval apex; presternum except anterior margin; meso- and metasterna; propleuron ex- cept anterior margin, posterior half of lateral margin and acetabula; meso- pleuron except lateral margin and ace- tabula; metapleuron except lateral and posterior margins, acetabula, and os- tiolar peritreme; abdominal sternum VII medially; genital segments medi- ally in females, completely in males; antennal segments I to III; labium; and legs except trochanters and distal margins of coxae. The following pale yellow: acetabulae; ostiolar peritreme; trochanters; and distal margins of coxae. Antennal segment W dark brown basally, light brown apically. Eye brown. Ocellus red. Hemclytral membrane subhyaline, brown basally becoming almost clear apically. Structure: Fairly broad, deep bod- ied. Head slightly declivent, strongly swollen behind eye; posterior margin of 318 The University of Kansas Science Bulletin ocellus level with or posterad of poste- rior margin of eye; buccula moderately produced, slightly convex, tapering gendy posterad. Antennal segment I surpassing anteclypeus by about 'A its length. Labial segment I not or just reaching anterior prosternal margin, II slightly surpassing procoxa, III slightly surpassing mesocoxa, IV reaching ab- dominal sternum III. Pronotum mod- erately coarsely punctate; postcallar impression interrupted on either side by a short submedial ridge; anterior lobe and about anterior half of poste- rior lobe finely granulate giving a dull appearance. Scutellum about twice as long as claval commissure; subacute apically; entire base finely granulate giving a dull appearance. Thoracic pleura coarsely, shallowly punctate an- teriorly, impunctate posteriorly; post- erolateral metapleural angle slightly produced; posterior metapleural mar- gin directed slightly anteroventrally. Abdominal sternum II about half cov- ered by metapleuron, posterior margin nearly straight; VI of female about Ve as long medially as laterally. Abdomi- nal dorsum as in Fig. 46e; medi- otergites IV and V elongate, narrowed laterally, most strongly in female. Pygophore as in Figs. 48m, n; convex posteriorly; median projection short, narrow, lacking tablikc apical process; tergum IX distinct, not depressed me- dially. Clasper as in Figs. 50d-f; caudolateral lobe moderately devel- oped; blade flattened, very broad ba- sally; shaiply curved and narrowed apically; shank moderately excavated interiorly. Aedeagus as in Fig. 49j; phallotheca and phallothecal process as in Figs. 48g, h; conjunctiva elongate, without lobes; ejaculatory reservoir as in Figs. 49a, b; vesica basally tubular, about as thick as conjunctiva, twisted, without lobes; ring sclerite complete or nearly so, broadest basally; vesica api- cally coiled, flattened, greatly pro- duced laterally especially at ring sclerite, grooved to receive retracted gonoporal process; gonoporal process very long, twisted, corkscrew-like sub- apically; secondary gonopore oblique, without apical process. Ovipositor as in Figs. 52e, f, 531; first valvula reaching apex of connecting membrane, pro- duced basolaterad subapically; con- necting membrane of first valve broadly notched medially, lacking scle- rites; second valvula produced ventrad subapically, scarcely thickened api- cally; connecting membrane of second valve reaching almost to apex of val- vula, slightly produced and pigmented medially. Spermatheca as in Fig. 51b: tighdy coiled immediately basad of api- cal bulb; cup shaped flange usually (not always) covering coiled section; sperm pump immediately basad of coiled sec- tion. Measurements: Length S 6.6 (6.1- 7.2), $ 7.4 (6.7-8.4); abdominal width S 2.'o (1.9-2.1), i 2.3 (2.0-2.5); head length S .90 (.86-. 93), 2 1.07 (.99- 1.18); width across eyes S 1.45 (1.32- 1.55), ? 1.59 (1.45-1.73); pronotal length S 1.13 (1.05-1.18), 2 1.28 (1.13-1.45); pronotal width S 1.97 (1.81-2.20), ? 2.25 (2.00-2.51); scutel- lar length S 1.08 (1.00-1.21), i 1.19 (1.05-1.35); scuteflar width S 1.11 (1.00-1.30), 2 1.29 (1.13-1.50); length antennal segments I ^ .35 (.30-. 38), ? .38 (.35-. 42), II S 1.09 (1.03-1.25), ? 1.15(1.05-1.29), III c?.94(.86-l.ll), $ 1.06(.86-1.13), IV S 1.26(1.16-1.38), ? 1.35 (1.23-1.44); length labial seg- ments I S .89 (.83-. 98), i 1.03 (.93- 1.13), II S .92 (.85-1.04), v 1.07 (.93- 1.18), III S .90 (.83-1.01), i 1.05 (.94-1.18), IV ^ .60 (.58-. 63), ? .65 (.60-. 71). Comments. — Arocatus continctus is eas- ily separated from other Australian members of the genus by the pronotal The Lygaeinae of Australia 319 markings and the orange median scu- tellar vitta. The dark hemelytral mark- ings are variable in extent, often covering the entire clavus and corium. The abdominal venter is sometimes nearly completely infuscated medially, approaching the appearance of Grap- tostethus species. Distant (1918) described a light col- ored specimen as Graptostethus parvus and stated that it was allied to G. cardinalis (Stal). The two species are somewhat similar in general appear- ance and somewhat atypical of their respective genera, at least considering only Australian species. They do, how- ever, conform to the basic generic char- acteristics. Arocatus continctus has a curious dis- junct distribution, occurring in western Africa, India and Australia. In Aus- tralia it is apparently almost restricted to the eastern coastal region of Queens- land (Fig. 3). It is often found in large numbers within and on the pods of "Wild Cotton" [Gornphocarpus spp.). I have also found adults and nymphs on Nerium oleander. I believe that this species is oriental and was introduced fairly recently to Africa and Australia. The male and female genitalia differ considerably from other members of the genus I have examined while a species from the Ryukyu Islands tentatively identified as A. fasciatus Jakovlev has genitalia very similar to those of A. continctus. Arocatus aenescens Stal 1874 Figures 4; 7a; 46f; 47c, d; 48e, f; 49m, n; 50p-r; 51e; 53c, d, i. Arocatus aenescens Stal, 1874:115-116; Slater, 1964a: 19. Scopiastes bergrothi Kirkaldy, 1903:16. Type. — Type +, "Australia" in Stock- holm. Redescription. — Color: Brown. The aenescens Fig. 4. Arocatus aenescens Stal. dorsal \iew. Scale = 1 iiitii. following orange: head except longitu- dinal median vitta dorsally; scutellar apex; thoracic pleura except acctabu- lae; posterior metapleural margin and ostiolar peritreme; abdominal sterna; and abdominal terga except VII poste- riorly. The following pale yellow: ante- rior prosternal and propleural margins; posterior margins of abdominal sterna; acetabulae; posterior metapleural mar- gin; ostiolar peritreme; apices of coxae; trochanters; and extreme bases of femora. Antenna and labium dark brown to black; antennal segment IV lighter apically. Eye brown. Ocellus red. Hemelytral membrane sub- hyaline, dark brown basally, lighter apically. Structure: Narrow, elongate. Head slightly declivcnt, moderately swollen behind eye; posterior margin of ocellus anterad of posterior margin of eye; 320 The University of Kansas Science Bulletin buccula moderately produced, slightly convex, tapering gently posterad. An- tennal segment I surpassing anteclyp- eus by about 'A its length. Labial segment I not reaching anterior pro- sternal margin, II reaching middle of procoxa, III reaching mesocoxa, IV reaching about middle of metacoxa. Pronotum coarsely punctate, finely granulate dull area restricted to ante- rior angles. Scutellum about I'A times as long as claval commissure; acute apically; finely granulate dull area re- stricted to basal angles. Pro- and mes- opleura except acetabulae coarsely punctate. Metapleuron except acetab- ula and narrow posterior margin coarsely punctate; posterior margin very slightly concave, directed dorso- ventrally; posterolateral angle not pro- duced. Abdominal sternum II convex laterally; VI of female about % as long medially as laterally. Abdominal dor- sum as in Fig. 46f; mediotergites IV and V somewhat elongate, slightly nar- rowed at common margin. Pygophore as in Figs. 47c, d; nearly truncate posteriorly; short tablike process at apex of median projection; tergum IX moderately broad, impressed medially. Clasper as in Figs. 50p-r; caudolateral lobe scarcely indicated; blade narrow, rounded in cross section; shank exca- vated, with moderately projecting overhang. Aedeagus similar to^. rush- cus (Fig. 49e); phallotheca and phal- lothccal process as in Figs. 48e, f; ejaculatory reservoir as in Figs. 49m, n. Ovipositor as in Figs. 53c, d, i; resembling A. rusticus; connecting membrane of first valve cleft medially, cleft surrounded by sclerotized area; apex of second valvula bluntly pro- duced dorsoapically. Spermatheca as in Fig. 51e; sperm pump near base. Measurements: Length ' 6.0 (5.6- 6.4), ; 6.6 (6.2-7.1); abdominal width S L8 (1,7-1.9), V 2.0 (1.8-2.2); head length ^ .86 (.72-1.00), 2 .96 (.90- 1.08); width across eyes ^ 1.44 (1.36- 1.48), i 1.56 (1.50-1.62); pronotal length ' 1.11 (1.00-1.16), $ 1.19 (1.08-1.30); pronotal width S 1.74 (1.64-1.84), . 1.97 (1.86-2.08); scutel- lar length ^ .82 (.76-. 90), v .92 (.80- .96); scutellar width ^ .90 (.84-1.00), $ 1.04 (.92-1.08); length antennal seg- ments I ' .39 (.36-. 41), i .40 (.36- .43), II ' .95 (.91-. 98), . 1.00 (.96- 1.05), III ' .86 (.80-. 91), $ .92 (.88- .97), IV ' 1.06 (.96-1.20), 5 1.14 (1.00-1.23); length labial segments I S .74 (.70-. 76), $ .82 (.76-. 86), II ' .75 (.72-. 86), i .82 (.76-. 88), III S .66 (.62-. 74), i .76 (.72-. 86), IV S .69 (.64-. 72), V .78 (.74-. 82). Comments. — Arocatus aenescens is the smallest Australian member of the genus and the only one with unicolored pronotum and hemelytra. In the field it could be mistaken for dark specimens of A. continctus from which it is easily separated by the orange thoracic pleura. The extent of the dark dorsal vitta on the head is variable, sometimes covering the entire vertex and occa- sionally confined to the apex of the anteclypeus. There is often an indis- tinct longitudinal vitta on either side of the abdominal venter. A single male from northern Queensland has dark thoracic pleura. This species is closely related to A. rusticus (Stal) and A. chiasmus, sp. nov. It differs from those two species in the placement of the median sclerite of the membrane connecting the first valvu- lae of the ovipositor and in the only slightly elongate and scarcely narrowed fourth and fifth mediotergites of the abdominal dorsum, approaching A. montanus, sp. nov., in the latter charac- teristic. Arocatus aenescens occurs in the eastern coastal part of Australia from New South Wales to the Torres Strait (Fig. 7a). The species appears to be The Lygaeinae of Australia 321 associated primarily with lowland rain- forest. It has been collected in large numbers under bark in New South Wales. In Queensland I have found it commonly on Parsonsia straminia and occasionally on Gomphocarpus spp. Arocatus rusticus Sthl 1867 Figures 5b; 6; 46a; 47a, b; 48a, b; 49e- g; 50g-i; 51a; 53a, b, g. Tetralaceus (sic) rusticus Stal, 1867:163. Astacops ? caligatus Walker, 1872:36-37. Lygaeus subjectus Walker, 1872:62. Lygaeus singulans Walker, 1872:63; Le- thierry and Severin, 1894:157; Dis- tant, 1901fl:541; Slater, 1964a:117. syn. nov. Lygaeus ruficollis Walker, 1872:64. Arocatus rusticus Stal, 1874:115; Slater, 1964a:28-29; Slater, 1978:857. Arocatus ruficollis B. -White, 1878:32; Tillyard, 1926:11, 146. 7>/>i?^. — Lectotype ? o{ rusticus, "Aus- tral, boreal. Thorey." in Stockholm. Paralectotype V of rusticus, same data as lectotype, in Stockholm. Type of L. singularis lost fide Distant, 1914a. Redescription. — Color: Reddish or- ange. The following dark brown to black: head dorsally including anten- niferous tubercle and swollen area be- hind eye; posterior pronotal lobe except extreme posterior margin; scu- tellum except apex; triangular macula along costal margin of corium from near base to about '/(, distance from apex, subparallel to claval suture and apical margin, widest opposite apex of clavus; meso- and metasterna except raised posterior margins; rounded cen- chiasmus rusticus Fig. 5. a. Arocatus chiasmus, sp. nov., dorsal view, b. Arocatus rusticus (Stal), dorsjil view. Scale = 1 mm. 322 The University of Kansas Science Bulletin tral macula near lateral margin of ab- dominal sternum VI; similar macula on sternum VII, sometimes expanded to cover entire sternum except lateral margin; genital segments; antenna; la- bium; and legs. The following pale yellow: anterior prosternal and pro- pleural margins; posterior margins of all thoracic sterna; posterior margins and acetabulae of all thoracic pleura; buccula; and sometimes posterior pro- notal margin. Eye brown. Ocellus red. Hemelytral membrane subhyaline; ba- sal angle whitish, otherwise dark brown basally becoming paler apically. Structure: Narrow, elongate. Head scarcely declivent, slightly swollen be- hind eye; posterior margin of ocellus anterior to posterior margin of eye; buccula moderately produced, slightly convex, tapering gently posteriorly. Antennal segment I surpassing ante- clypeus by about Vi its length. Labial segment I surpassing anterior proster- nal margin by about 'A its length, II reaching middle of mesosternum. III reaching anterior margin of abdominal sternum III, IV reaching abdominal sternum IV. Pronotum moderately coarsely punctate; anterior lobe includ- ing callus finely granulate giving a dull appearance. Scutellum about 1 '/t times as long as claval commissure; acute apically; entire base finely granulate giving a dull appearance. Propleuron coarsely punctate anteriorly and poste- riorly, indistinctly punctate centrally. Meso- and metapleura indistinctly punctate anteriorly, coarsely punctate posteriorly except impunctate posterior margins; posterior metapleural margin slightly concave, directed dorsoven- trally; posterior metapleural angle not produced. Abdominal sternum II with posterior margin convex laterally; VI of female about % as long medially as laterally. Abdominal dorsum as in Fig. 46a; mediotergites IV and V strongly narrowed to their common margin, most strongly in female. Pygophore as in Figs. 47a, b; concave posteriorly; median projection with tablike apical process; tergum IX narrow, impressed medially. Clasper as in Figs. 50g-i; caudolateral lobe slightly indicated; blade evenly curved to apex, rounded in cross section; shank excavated inte- riorly, overhang strongly produced, somewhat tentlike. Aedeagus as in Fig. 49e; phallotheca as in Figs. 48a, b; conjunctiva elongate, tubular, with notched, pigmented subapical lobe dorsally; ejaculatory reservoir as in Figs. 49f, g; vesica basally moderately long, twisted, slightly flattened and produced laterally; ring sclerite incom- plete, moderately elongate, scarcely convex; vesica distally tightly coiled, expanded laterally for entire length, when retracted forming cuplike recep- tacle for retracted gonoporal process; gonoporal process short, slightly con- stricted subapically, visibly covered by membrane to apex; secondary gono- pore flared, without apical process. Ovipositor as in Figs. 53a, b, g; first valvula not reaching apex of connect- ing membrane, ventral margin notched subapically; connecting mem- brane of first valve with median and sublateral sclerites, the latter reaching apex of membrane; second valvula slightly swollen subapically, a very acute curved projection at apex; con- necting membrane of second valve deeply cleft medially. Spermatheca as in Fig. 51a; sperm pump much closer to base than to apex. Measurements: Length ^ 8.3 (7.7- 8.9), V 9.0 (8.1-9.8); abdominal width S 2.2 (2.0-2.4), V 2.5 (2.2-2.7); head length ' 1.17 (1.08-1.24), $ 1.27 (1.14-1.44); width across eyes c^ 1.85 (1.72-1.96), 5 1.97 (1.80-2.16); prono- tal length c^ 1.38 (1.28-1.52), v 1.47 (1.26-1.68); pronotal width S 2.31 TnK Lygaeinae of Australia 323 (2.12-2.50), r 2.48(2.20-2.72); scutcl- lar length ' 1.13 (.92-1.26), i 1.22 (1.12-1.32); scutellar width ' 1.28 (1.14-1.36), . 1.41 (1.24-1.54); length antennal segments I '^ .50 (.48-. 54), V .53 (.48-. 59), II S 1.71 (1.66-1.84), 2 1.85 (1.63-2.03), III ^ 1.42 (1.33- 1.54), 9 1.53 (1.36-1.75), IV S 1.43 (1.29-1.55), . 1.50 (1.33-1.59); length labial segments I ^ 1.33(1.28-1.40), ? 1.50(1.33-1.59), II ? 1.35 (1.30-1.40), ? 1.57 (1.36-1.76), III ' 1.61 (1.50- 1.76), V 1.83 (1.52-2.08), IV ^ .97 (.92-1.00), ? 1.05 (.92-1.16). Comments. —Arocatus rusticus is very similar to A. chiasmus, sp. nov. , in general appearance but lacks dark markings on the thoracic pleura and has a longer labium. The extent of the triangular corial macula is quite vari- able, sometimes reduced to little more than a thin line along the corial margin and sometimes covering almost the whole corium. Dark specimens might be confused with ^. montanus, sp. nov., but are easily separated from that spe- cies by the black maculae on either side of the abdominal venter near its apex. In Australia A. rusticus occurs only in the east, its range extending from Tas- mania in the south as far north as the Tropic of Capricorn (Fig. 6). The spe- cies also occurs on Lord Howe and Norfolk islands and in New Zealand. It is apparently tolerant of more diverse climatic conditions than are most members of the genus and has a wider range of food plants. Adults and nymphs have been collected on Parson- sia straminia, Araujia hortorum, Nerium oleander and Gomphocarpus sp. Adults have also been taken on Asclepias cur- rasavica. A parasitic fly, Alophura aureiventns Curran (Tachinidae), emerged from a female collected in Brisbane and kept alive in the laboratory. The fly larva emerged at the base of the ovipositor. killing the host. This lachinid has been recorded from the pyrrhocorid Dysder- cus sidae Montrouzier from .several lo- calities in Queensland (B. Cantrell, pers. comm.). Arocatus chiasmus, sp. nov. Figures 5a; 6; 46b; 47e, f; 48c, d; 49c, d; 50m-o; 51d; 53e, f, h. 7>/?^.5.— Holotype ^, "Richmond, Queensland, VIII-1965, A. Fen- wick" in QM. Paratypes: 2', 15, same data as holotype, in UQ. Also: Western Australia — 15, 8 mi S of Lansdowne Homestead, 21-28. vi. 1964, R. Plumb; 1; , 8 mi S of Lans- downe Homestead, l.ix.l964, R. Plumb; 1 '", 8 mi S of Lansdowne Homestead, 4.ix.l964, R. Plumb; 19, 4 km ESE of Millstream, 21.36S, 117. 07E, 31.x. 1970, M. Upton and J. Feehan (ANIC); 19, Fortescue River Hammersley Range, W. D. Dodd (SAM); 19, 39481 mileage N of White Mountain S of Rosewood Station, 29. v. 1970, Lemley Expedition (WAM); North- ern Territory — 1^, Amadeus Basin, 3.vii.l962, P. Ranford, swept from foliage; 19, Burnside, Brock's Creek, 18. iv. 1929, T. G. Campbell; 19, 1 km N of Cahill's Crossing, East Alligator River, 12.25S. 132. 58E, 8-9.xi.l972, J. C. Cardale; 19, 7 km NW by N of CahiU's Crossing, 12.23S, 132. 56E, 12. xi. 1972, Upton and Feehan; 1 ', 1 km S of Cahill's Crossing, 12.26S, 132. 58E, 3.xi.l972, Upton and Bar- rett; 1 ; , 4 mi W of Coolibah Home- stead, 15.34S, 130. 54E, 6.vii.l968, M. Mendum: 1 . Koongarra, 12.52S, 132. 50E, 16. xi. 1971, V. E. Ingle; l^ 12 km NNW of Mt. Cahill, 12.46S, 132. 39E, 25.x. 1972, M. Upton (ANIC); 1 ^ \ ■ , Port Keats, ix-x.l973, R. Easton (NMV); 1', Darwin, xii.1963. \V. 324 The University of Kansas Science Bulletin • A.rusticus o A chiasmus _L ± _L Fig. 6. Distribution oi' Arocatus ruslicus (Stal) and A chiasmus, sp. nov. in Australia. P. Walsh (SAM); 19, Swim Creek, 13.x. 1971, Wein and Allwood (PIBNTH); 1 ; , 4.8 km S of Renner Springs, 8.iii.l966, N. McFarland, at UV light; H, 10 mi NW of Yuerduni Creek bed, 20. ii. 1968, at Hght (SAM); iv, Horn Islet, Sir Edward Pcllcw Group, 22-28. ii. 1968, B. Cantrell; 1., Horn Islet, Sir Edward Pellew Group, IS.viii. 1967, M. C. McKee (DPI); Queens- land—1., Clermont, 20. ix. 1928, Dr. K. K. Spence (AM); 19, Gap Creek, 6 mi N of Bloomfield River, 13-14. X. 1965, G. Monteith; IS, Lankelly Creek, Mcllwraith Range, near Coen, Cape York Peninsula, 28-31. X. 1969, B. Cantrell; 1^, 2?, Mica Creek, 8 mi S of Mt. Isa, 20.iii.l973, E. M. Exley, on Tris- tania grandiflora (UQ). Description. — Color: Orangey red. The following dark brown to black: apex of anteclypeus; posterior pronotal lobe except broad posterior margin; scutellum except apex; clavus apically along scutellar margin; triangular mac- ula along central half of costal corial margin, widest opposite apex of clavus; produced posterior part of propleuron; mesopleuron except acetabula; meta- pleuron except lateral, posterior, and acetabular margins and ostiolar per- The Lygaeinae of Australia 325 itreme; transversely ovate macula lat- erally on abdominal sterna VI and VII; genital segments; and antennal segments 1,11, III, and base of IV. The following pale yellow: anterior proster- nal margin; posterior margins of tho- racic sterna; anterior, posterior, and acetabular propleural margins; acetab- ulae of meso- and metapleura; and ostiolar peritreme. Eye brown. Ocellus red. Posterior margin of pronotum, apex of scutellum, antennal segment IV apically, labium, and legs brown. Hemelytral membrane subhyaline, ba- sal angle whitish, otherwise dark brown basally becoming paler apically. Structure: Narrow, elongate. Head slightly declivent, slightly swollen be- hind eye; posterior margin of ocellus level with or anterior to posterior mar- gin of eye; buccula moderately pro- duced, slightly convex, tapering gradually posterad. Antennal segment I surpassing anteclypeus by about 'A its length. Labial segment I surpassing anterior prosternal margin by about Ve its length, II surpassing anterior 'A of mesosternum, III reaching metacoxa, IV surpassing anterior 'A of abdominal sternum III. Pronotum moderately coarsely punctate; anterior lobe includ- ing callus finely granulate appearing dull. Scutellum about 1 'A times as long as claval commissure; acute apically; narrow band at base finely granulate appearing dull. Propleuron coarsely punctate anteriorly and posteriorly, moderately coarsely punctate centrally, acetabula impunctate. Mesopleuron moderately heavily punctate except im- punctate lateral margin and acetabula. Metapleuron moderately punctate ex- cept impunctate lateral and posterior margins and acetabula; posterior mar- gin slightly concave, directed dorso- ventrally; posterolateral angle not produced. Abdominal sternum II con- vex laterallv, VI of female about -'A as long medially as laterally. Abdominal dorsum as in Fig. 46b; mediotergites IV and V elongate, narrowed to their common margin. Pygophore as in Figs. 47e, f; posterior margin moder- ately concave; median projection with tablike apical process; tergum IX wide, expanded laterally at posterior margin, impressed medially. Clasper as in Figs. 50m-o; very similar to A. rusticus except margin of shank's excavation "rolled." Aedeagus similar to A. rusti- cus (Fig. 49e); gonoporal process longer, thicker, more strongly con- stricted subapically; secondary gono- pore more strongly flared; phallotheca and phallothecal process as in Figs. 48c, d; ejaculatory reservoir as in Figs. 49c, d. Ovipositor as in Figs. 53e, f, h; similar to A. rusticus except apical pro- jection of second valvula less curved. Spermatheca as in Fig. 51b; tube thick for genus; sperm pump slightly further from base than from apex. Measurements: Holotype .^ . Length 7.6; abdominal width 2.1; head length 1.02; width across eyes 1.88; pronotal length 1.40; pronotal width 2.24; scu- tellar length 1.06; scutellar width 1.08; length antennal segments I .43, II 1.44, III 1.20, IV 1.36; length labial segments I 1.02, II 1.00, III 1.20, IV .76. Measurements of paratypes. Length S 7.7 (6.9-8.0), ? 8.3 (7.5-9.2); ab- dominal width S 2.1 (1.8-2.2), $ 2.4 (2.1-2.8); head length ^ 1.01 (.92- 1.10), $ 1.02 (.84-1.26); width across eyes S 1.84 (1.66-1.94), . 1.97 (1.82- 2.18); pronotal length ^ 1.35 (1.16- 1.46), 9 1.45 (1.26-1.66); pronotal width .^ 2.15(1.84-2.32), . 2.42(2.14- 2.68); scutellar length S .99 (.84-1 . 10), $ 1.15 (1.00-1.30); scutellar width S 1.04 (.80-1.16), V 1.20 (1.04-1.36); length antennal segments I ' .42 (.40- .44), . .44 (.41-. 47), II S 1.44 (1.25- 1.53), i 1.53 (1.40-1.64), III ' 1.17 326 The University of Kansas Science Bulletin (1.06-1.24), 5 1.25 (1.14-1.36), IV S 1.31 (1.27-1.36), I 1.38 (1.27-1.52); length labial segments I ^ 1.02 (.90- 1.10), . 1.15 (1.06-1.24), II ' 1.00 (.86-1.10), 5 1.13 (1.04-1.20), III S 1.19 (1.10-1.24), $ 1.30 (1.12-1.40), IV (? .77 (.72-. 82), $ .80 (.72-. 84). Comments. — Arocatus chiasmus is very similar to^. rusticus with which it forms a north-south species pair. It may be separated from rusticus by the dark thoracic pleura and shorter labium. The reddish orange head is a very distinctive character when present but is frequently as dark as light examples of rusticus. This species is apparently restricted to northern Australia, occurring as far south as the Tropic of Capricorn (Fig. 6). It appears to be associated with drier areas than the other Australian members of the genus. Astacops BoiSDuvAL Astacops Boisduval, 1835:637; Scudder, 1963:319-324; Slater, 1964^:33; Blote, 1972:410-411. Montaltus Distant, 1888:483. Type-species Astacops australis Boisdu- val, 1835 (fixed by Distant, 1904). At one time Astacops contained all Lygaeinae with distinctly stylate eyes. The group was revised by Scudder (1963) who provided keys and diag- noses for the genera and species within the complex. He recognized four gen- era, two of which, Astacops and Scopias- • A aenescens ° A. fastosus -L • A. vindiventris o A. dodd Fic; a. Distribution ol .\rocatus aenescens Stiil and A. Jastosus. sp. nov. in Australia, b. Distribution o( Astacops . viridivenlris Stal and A. doddi Scudder in Australia. Thk Lyoaeinae of Australia 327 tes, occur in Australia. In this paper 1 am erecting the genus Woodwardiastes, and elevating the subgenus Scopiastella to generic level. This brings the total of Australian genera with stylate eyes to four. The generic divisions within the Astacops complex are difficult to define concisely. Genitalic characters are by far the best to use in doubtful cases. As Scudder (1963) has given a ge- neric diagnosis for Astacops I do not feel it necessary to provide one here. How- ever his specific descriptions, while generally adequate for ssparating spe- cies, are minimal — especially as re- gards structure. Further, I have found that I disagree with some of his color designations for the Australian species. I therefore give rather detailed species descriptions. As presently constituted Astacops contains about 50 species with the greatest number occurring in New Guinea. Scudder (1963) split the genus into three groups based on the struc- ture of the aedeagus and spermatheca. He placed the two Australian species in the Astacops dorycus group, charac- terized by an S-shaped portion of the spermatheca near the apical bulb and a lateral flange near the aedeagal ring sclerite. Both Australian species are restricted to northern Queensland in Australia. One is apparently endemic and the other also occurs in New Guinea. The two species are very simi- lar structurally and may be separated from other Australian Lygaeinae with stylate eyes by their large size and rather robust bodies. Key to the Australian Species o/ Astacops 1. Pronotum black; abdominal ven- ter red, tinged with orange, and without transverse fasciae .... doddi Scudder r. Pronotum pale yellow or or- ange; abdominal venter pale yel- low, often suffused with green, and usually with transverse fas- ciae on segments IV to VI ... viridiventris Stal Astacops viridiventris Stal 1874 Figures 7b; 8; 54f-j; 55a-i Astacops viridiventris Stal, 1874:100; Scudder, 1963:370; Slater, 1964a: 38. Astacops subochraceus Distant, 1918:416; Slater, 1964a:37. Types. — Lectotype :^ , "Cape York. Thorey. " in Stockholm. Paralecto- types: \S, 4$, same data as lecto- type, in Stockholm. Redescription. — Pale yellow. The fol- lowing orangey red: head except venter medially; base of antennal segment I; pronotal humerus; coreaceous part of hemelytron, becoming darker apically; often apical half of fore and mid fem- ora; and sometimes scutellum. The following dark brown to black: apex of anteclypeus; eye; antenna except base of segment I; labium; sometimes scu- tellum; legs except coxae and apical part of fore and mid femora; sometimes \ / viridiventris Fig. 8. Astacops viridiventris Stal, dorsal view. Scale = 1 mm. 328 The University of Kansas Science Bulletin abdominal terga II to VT; transverse fascia, broadest medially, on abdomi- nal sterna IV to VI, sometimes indi- cated on sternum III of male and VII of female; margin of cleft in female sternum VII; ovipositor except terga; and sometimes lateral sternal margins. The following infuscated to varying degree: apical Vj of clavus; corium excluding costal margin; scutellum; and sometimes distal half of fore and mid femora. Abdominal venter often suffused with green. Ocelli red. Hem- elytral membrane subhyaline, dark brown basally becoming lighter until almost clear at apex. Structure: Moderately robust; tho- rax finely granulate, subshining, rest of body and appendages shining. Head moderately declivent; narrower across eyes than is base of pronotum; vertex slightly depressed; a low subapical hump on anteclypeus; ocelli higher than vertex, about as far from eyes as from each other; eye on distinct slightly dorsally directed stalk; buccula sub- triangular, most prominent medially, reaching level of front of eyes. Anten- nal segment I surpassing anteclypeus by half its length. Labial segment I scarcely surpassing anterior prosternal margin, II reaching posterior procoxal margin, ill reaching middle of meso- coxae, IV reaching middle of meta- coxae. Pronotum much higher posteri- orly than anteriorly, scarcely con- stricted but distinctly impressed behind callus; anterior margin straight; poste- rior margin convex; lateral margin nearly straight; posterior lobe about three times as long as anterior lobe; callus slightly swollen; callar impres- sion oblique, slightly sinuate, un- branched; postcallar impression with row of punctures of varying size; me- dian carina obsolete, faintly indicated immediately behind callus; an oblique impression at posterior margin laterad of scutellum. Scutellum with well de- veloped T-shaped median carina; lat- eral fovea very coarsely punctate. Hemelytra not covering abdominal dorsum laterally; costal corial margin slightly concave basally, slightly convex apically; claval commissure about V4 length of scutellum; veins moderately raised; membrane greatly surpassing abdomen. Thoracic sterna impunctate; posterior margins raised. Pro- and mesopleuron with dorsoventral, indis- tinct row of punctures behind middle. Metapleuron impunctate; posterior margin slightly sinuate, directed slightly posteroventrally; posterolateral angle not produced. Legs relatively long; femora scarcely swollen. Abdom- inal sternum II not covered by meta- pleuron; posterior margin male sternum VII concave; VII of female cleft nearly to base, posterior margin slightly concave laterally. Abdominal dorsum with paratergites III to VI separate, VII aparently fused with lat- erotergites; mediotergite IV elongate; mediotergite VII with slightly concave posterior margin, indistinctly sepa- rated from fused para- and lateroter- gites. Pygophore as in Figs. 55b, c; median projection well developed, nar- row basally, broader apically, apex with narrow dorsal hump; well developed ridge on either side of median projec- tion separated from projection by chan- nel; tergum IX fused to pygophore. Clasper as in Figs. 54h-j; caudolateral lobe scarcely indicated; blade rounded in cross section, slightly curved, api- cally acute; shank slightly excavated, a sinuate ridge anterolaterally. Aedeagus as in Figs. 55e, f; phallotheca moder- ately sclerotized, dorsum slightly cari- nate medially; phallothecal process as in Fig. 55a, projecting dorsally; con- junctiva robust, a low lateral lobe on each side, a pair of small, conical dorsal lobes just before apex; ejaculatory res- The Lygaeinae of Australia 329 ervoir as in Figs. 541, g, without basal projection; vesica basally robust, di- \ ided into two vaguely defined lobes; ring sclerite complete, elongate, broader distally; vesica apically with produced lateral flange, gready pro- duced at ring sclerite, forming lightly pigmented cup at apex which holds gonoporal process when retracted, a short subapical sclerotized spike on in- ner magin; gonoporal process short, thick, visibly covered with membrane to apex, twisted subapically; secondary gonopore slightly flared, with an irreg- ularly bent apical process. Ovipositor as in Figs. 55d, g, h; first valvula nearly reaching apex of connecting membrane; connecting membrane of first valve pigmented, granulate, deeply notched medially, two rounded projections at base of notch; second valvula flattened, bluntly rounded api- cally; connecting membrane of second valve cleft nearly to base, sclerotized except at junction with valvula. Sper- matheca as in Fig. 55i; short, pig- mented distad of sperm pump, sinuate and with cup-shaped flange just basad of apical bulb; sperm pump bell- shaped, pigmented distally, closer to apex than base. Measurements: Length ^ 11.2 (10.8-11.7), 5 12.4 (12.0-13.0); ab- dominal width S 3.5 (3.2-3.8), ? 4.0 (3.7-4.2); head length S .80 (.72-. 96), ? .88 (.60-1.04); width across eyes S 2.66 (2.54-2.72), $ 2.86 (2.76-3.00); pronotal length S 1.90 (1.74-2.16), ^ 2.06 (1.80-2.18); pronotal width S 3.41 (3.30-3.62), 2 3.88 (3.62-4.10); scutellar length ^ 1.74(1.60-1.90), ? 2.08 (1.90-2.20); scutellar width ^ 1.80 (1.60-2.04), I 2.12 (1.96-2.28); length antennal segments I ? .89 (.84- .93), $ .92 (.90-. 95), II ^ 2.36 (2.19- 2.50), $ 2.33 (2.20-2.48), III ^ 2.06 (1.93-2.18), 2 2.08 (1.97-2.27), IV S 2.47 (2.31-2.56), 2 2.59 (2.46-2.78); length labial segments I ' 1.13 (1.06- 1.20), 9 1.31 (1.20-1.40), II ' 1.13 (1.04-1.20), V 1.33 (1.24-1.42), III S 1.06(.98-1.12), . 1.22(1.12-1.36), IV S 1.02 (.98-1.04), V 1.13(1.06-1.20). Comments. — Astacops viridiventris is easily separated from A. doddi, the only other Australian member of the genus by the characters given in the key. Scudder (1963) provides a key to the genus including the two Australian spe- cies and gives a short discussion of their diagnostic characteristics. Occasional individuals have the pronotum com- pletely orange or lack the transverse fasciae of the abdominal venter. Ten- eral specimens may be completely or- ange except the eyes and abdomen. Nothing is known of the biology or food preferences oi Astacops viridiventris. Its range in Australia is restricted to the Cape York Peninsula (Fig. 7b). Adults have been collected throughout the year. Astacops doddi Scudder 1963 Figures 7b; 54a-e. Astacops doddi Scudder, 1963:342. 7>/)r— Holotype $, "Cairns, 1920, A. P. Dodd, ex scrub" in ANIC. Redescription.— Color: Red, tinged with orange. The following infuscated: apical half of costal corial margin; gu- lar area; and antennal segment I dis- tally. The following black: pronotum except anterior margin; scutellum; tho- racic sterna except anterior prosternal margin; and thoracic pleura except an- terior propleural margin, lateral and posterior metapleural margins, acetab- ulae, and ostiolar peritreme. Posterior pronotal lobe, pro- and mesopleura suffused with red. The following or- ange: anterior prothoracic, propleural, and prosternal margins; base of costal corial margin; acetabulae; and lateral 330 The University of Kansas Science Bulletin and posterior metapleural margins. Legs except apical half of femora dark brown to black. Antennal segments II and III dark brown to black. Antennal segment IV and ostiolar peritreme light brown. Eye and labium dark brown. Ocellus red. Hemelytral mem- brane subhyaline, dark brown basally becoming progressively lighter until al- most transparent at apex, a pale mark at basal angle. Structure: Very similar to A. viridi- ventris, differs as follows. Robust, tho- rax and gular area pruinose. Head with prominent anteclypeal bump. Labial segment I surpassing anterior proster- nal margin, II reaching posterior half of procoxa. 111 reaching mesocoxa, IV reaching metacoxa. Pygophore with median projection narrower apically and with apical bump obsolete. Clasper as in Figs. 54c-e; shorter, more robust, more broadly rounded caudolaterally, apex of blade blunt. Aedeagus with phallothecal process slightly less pro- duced; ejaculatory reservoir as in Figs. 54a, b: gonoporal process slightly longer and thinner. Measurements: Holotype ^ . Length 11.4; abdominal width 3.7; head length .93; width across eyes 2.92; pronotal length 2.04; pronotal width 3.70; scutellar length 1.68; scutellar width 1.84; length antennal segments I .76, II 2.54, in 2.33, IV 2.45; length labial segments I 1.28, II 1.25, III 1.33, IV 1.04. Comments. — Astacops doddi is known only from the holotype male. It is easily separated from A. viridiventris by the characters given in the key. Scopiastclla Slater Scopiastes s.g. Xestonotellus Horvath, 1914:629 (nom. preocc). Scopiastes s.g. Scopiastella Slater, 1957: 36; Scudder, 1963:371-372; Slater, 1964fl:191-192; Blote, 1972:411. Type-species Scopiastes cruentus Horvath 1914 (fixed by Slater, 1957). Diagnosis. — Eye distinctly stalked; head across eyes at least as wide as base of pronotum; antennal segment I sur- passing anteclypeus by V2 to % its length; buccula convex anteriorly; pro- notum except posterior margin and humerus finely granulate; posterior pronotal lobe only slightly higher than anterior lobe; posterior pronotal lobe with at most scattered punctures; sper- matheca short, with small, basal, glob- ular sperm pump, tube connecting sperm pump and apical bulb gradually widened distally, and apical bulb large; clasper tilted anteriorly, a distinct dor- sal ridge at junction of shank and blade, caudolateral lobe well developed and subconical, and shank not or slightly excavated and bearing some- what dorsally directed shelflike projec- tion; conjunctiva with unpigmented lobe on each side at about middle; ejaculatory reservoir with Y-shaped ba- sal projection; vesica expanded lat- erally into short process forming continuation of ring sclerite and similar but larger process distal of and at 180° to first; and gonoporal process with transparent, cupped, lateral subapical process. Horvath (1914) erected Xestonotellus as a subgenus of Scopiastes Stal, separat- ing it from Scopiastes, sensu stricto, by the impunctate or very finely punctate as opposed to heavily punctate pos- terior pronotal lobe. Slater (1957) pointed out that Xestonotellus was preoc- cupied and established Scopiastella as a new name for the subgenus. Scudder (1963) discussed the difficulty of as- signing species to the subgenus on the basis of the single character on which it was based. He did not assign species to subgenera within Scopiastes on the ground that more than two groups The Lygaeinae of Australia 331 would be necessary to show relation- ships correctly. Blote (1972) provided a key to the genera of stylate eyed Ly- gaeinae based on the "classic" generic characters which included the two sub- genera of Scopiastes. While agreeing with Scudder that Scopiastes sensu lato contains more than two natural species groups I have found that the Australian members of the group fall into two quite distinct and well defined assemblages of species. One group includes Scopiastes degeen (Stal) the type species oi Scopiastes. The other group agrees in external struc- ture and in male and female genitalia with Scopiastes cruentus Horvath, the type species of Scopiastella. The differ- ences between the two groups in the structure of the aedeagus, claspers, and spermathecae are particularly striking. As these structures are usually consist- ent within genera in the subfamily I have no hesitation in raising Scopiastella to generic status. It is quite likely that Scopiastes should be split further but as those species differing significantly from S. degeeri do not occur in Australia such action is beyond the scope of this study. The following species belong to Sco- piastella: S. costalis (Horvath); S. cruentus (Horvath); S. elegans (Distant); S. himer- tos, sp. nov. ; S. melvillensis (Scudder); S. militaris (Distant); S. nigricoronatus, sp. nov.; S. plagiatus (Stal); S. rubricinctus, sp. nov.; S. walkeri (Distant); and S. woodwardi, sp. nov. As I have not seen all the species placed in Scopiastes sensu lato the list should not be considered to include all species belonging in Scopia- stella. The only genera of Australian Ly- gaeinae with which Scopiastella might be confused are those with distinctly styl- ate eyes, that is, Astacops, Scopiastes, and Woodwardiastes. Astacops differs in that its members are much larger and more robust, the posterior pronotal lobe is much higher than the anterior lobe, and the pronotum is much wider at the base than is the head across the eyes. Scopiastes differs in that the posterior pronotal lobe is densely and very coarsely punctate and that at most the anterior lobe and narrow anterior mar- gin of the posterior lobe of the pro- notum are finely granulate giving a dull appearance. Woodwardiastes differs in that the posterior pronotal lobe is moderately but very densely punctate, the first antennal segment is elongate, surpassing the anteclypeus by about 'V; of its length, and the bucculae are angulate anteriorly. The male and fe- male genitalia of all four genera are distinctive. Redescription. — Color: Variously, but usually brightly, patterned in red, yel- low, orange, brown, and black. Structure: Rather narrow and elon- gate, slightly to moderately constricted medially. Thorax except broad poste- rior margin and pronotal humeri and a narrow posterior metapleural margin finely granulate giving a dull appear- ance. Hairs except on finely granulate area a mixture of short, dense, decum- bent and a few longer, semiupright hairs, these latter longer and denser ventrally. Hairs of finely granulate ar- eas long, moderately dense, upright, longest and densest on calli. Head strongly declivent; vertex impressed in front of ocellus; eye on distinct stalk; ocelli slightly closer to eyes than to each other, higher than vertex; anteclypeus directed slightly posterad apically. sometimes with indistinct subapical bump; buccula convex and moderately prominent anteriorly, slightly concave and very low posteriorly, reaching 'h to 'A distance to anterior prosternal mar- gin. Antennal segment I surpassing anteclypeus by '/j to ~l^ its length, IV 332 The University of Kansas Science Bulletin longest, III often longer than II. La- bium not surpassing metacoxa; seg- ment I thickest, II to IV progressively thinner. Pronotum subquadrate, slightly wider posteriorly than anteri- orly, impressed and slightly constricted behind callar area; anterior margin straight or slightly concave; posterior margin straight or slightly convex; lat- eral margin slightly sinuate, slightly explanate centrally; posterior lobe im- punctate or with scattered at most moderate punctation; a shallow oblique impression at posterior margin laterad of scutellum; posterior lobe of- ten slightly impressed along posterior margin of finely granulate area in re- gion of humerus; callus moderately swollen, often restricted anteriorly by transverse medial impression; callar impression oblique, slightly sinuate, unbranched. Scutellum subacute api- cally; T-shaped median carina and lat- eral fovea at least moderately developed. Cla\us moderately rugose, coarsely granulate, not appearing dull. Corium moderately rugose, coarsely granulate, not dull; costal margin very slightly concave for basal '/2, very sliglitly convex for apical '/2; not com- pletely covering outer laterotergites ex- cept apically. Hemelytrai membrane surpassing tip of abdomen. Thoracic sterna impunctate, posterior margin raised. Thoracic pleura impunctate or punctate to varying degree; ostiolar peritreme well developed; posterior metapleural margin truncate, directed nearly dorsoventrally; posterior meta- pleural angle not produced, slightly rounded. Abdominal sterna without dorsolateral gland openings, II not cov- ered by metapleuron; female VII cleft to base, rather elongate, slightly con- cave laterally; male VII shallowly con- cave. Abdominal dorsum with paratergites III to VI distinct, VII in- distinctly separated from mediotergite VII; apodemes on paratergites III to VI. Pygophore subcircular in cross sec- tion; median projection well devel- oped; tergum IX almost completely fused with base of pygophore. Clasper with well developed, subconical, cau- dolateral lobe; blade curved, subcircu- lar in cross section basally, flattened and acute apically; shank canted ante- rad, with dorsal ridge and dorsally directed shelflike projection on interior side, flattened but not excavated on interior side. Aedeagus with lightly pigmented phallotheca ridged dorso- medially; phallothecal process low, di- rected dorsally; conjunctiva elongate, tubular, a dorsolateral lobe on each side at about midpoint, a low dorsal lobe over ejaculatory reservoir; ejacu- latory reservoir complete, Y-shaped basal projection present; vesica basally robust, slightly coiled, produced into broad apically directed lobe near ring sclerite; ring sclerite incomplete, nar- row basally, broader apically; vesica apically coiled, outer margin flanged and moderately pigmented, lateral pro- jection just beyond (almost a continua- tion of) ring sclerite, larger lateral projection distad of this on other side of vesica; gonoporal process short, sur- rounding membrane visible almost to apex; secondary gonopore simple, lacking apical process, bearing trans- parent, flanged subapical process. Ovi- positor with first valvula nearly reaching apex of connecting mem- brane; connecting membrane of first valve deeply cleft medially, pigmented but not sclerotized; second valvula rather flat, scarcely curved inwards apically; membrane connecting second valve almost entirely sclerotized, deeply cleft medially. Spermatheca with very short, unpigmented, basal tube; sperm pump small, rather globu- lar; rather short pigmented tube be- tween sperm pump and apical bulb. The LvGAEiNAK OF Australia 333 gradually widened to subapical flange; apical bulb large, pigmented. AV)' to the Australian Species o/Scopiastella 1 . Posterior pronotal lobe yellow, sometimes suffused with orangey red; a transverse dark brown to black macula just behind calli ... 2 r. Posterior pronotal lobe com- pletely or nearly completely dark brown to black 4 2. Head yellow dorsally, a subquad- rate dark brown to black macula at vertex . . . nigricoronatus, sp. no v. 2'. Head at least partly orangey red dorsally, no dark macula at ver- tex 3 3. Ostiolar peritreme black .... woodwardi, sp. nov. 3'. Ostiolar peritreme yellow . . . elegans (Distant) 4. Scutellum deep red 5 4'. Scutellum dark brown to black, sometimes light medially 6 5. Abdominal venter deep red; cor- ium mostly black with only a narrow red costal border; north- ern Queensland and New Guinea costalis (Horvath) 5'. Abdominal venter orange, sometimes very pale and suf- fused with green; corium with broad red costal margin; south- ern Queensland and northern New South Wales militaris (Distant) 6. Hemelytra completely dark brown to black walkeri papuas (Scudder) 6'. Hemelytra at least partly red. . . 7 7. Hemelytra dark brown to black with a broad transverse median fascia red; legs completely dark brown to black rubricinctus, sp. nov. T . Hemelytra with at least corium nearly completely red or orange; legs at least partly red or or- ange 8 8. Propleura dark brown to black except anterolaterally; labial seg- ment II red only at base; meta- femora red on only about basal half; abdomen not infuscated. . himertos, sp. nov. 8'. Propleura orange except on pro- duced posterior area; labial seg- ment II and metafemora completely orange; abdomen in- fuscated basally on each side . . melvillensis (Scudder) Scopiastella costalis (Horvath 1914) Figures 9; 12a; 56a, i-k; 57d, k. Scopiastes (Xestonotellus) costalis Horvath, 1914:630-631. costalis Fig. 9. Scopiastella costalis (Horvath), dorsed view. Sceile= 1 mm. 334 The University of Kansas Science Bulletin Scopiastes costalis Scudder, 1963:378. Scopiastes (Scopiastella) costalis Slater, 1964a:192. 7>/?^5.— Lectotype ?, no locality, in Budapest. Paralectotype v, no lo- cality, in Budapest. Redescription.— Color: Head and sometimes scutellum brown suffused with red, head darker dorsally. The following dark red: anterior pronotal lobe except sometimes anterior mar- gin; about central V3 of costal corial margin; sometimes scutellum; meso- and metasterna; thoracic pleura except narrow anterior margin, narrow lateral margin, broad posterior margins, and acetabulae; and abdomen. The follow- ing dark brown to black with purplish cast: posterior pronotal lobe except posterior margin and humerus; some- times anterior pronotal margin; clavus; corium except central Vi of costal mar- gin; margins of thoracic pleura; pro- sternum; antenna; labium except segment I; and legs except coxae. The following brown: posterior pronotal margin; humeral angle of pronotum; coxae; labial segment I; eye; and ostio- lar peritreme. Ocellus red. Hemelytral membrane subhyaline, dark brown ba- sally becoming paler apically. Dark pronotal areas except posterior margin and humerus of pronotum pruinose. Structure: Robust, rather strongly constricted at middle. Head very strongly declivent; anteclypeal bump only slightly produced. Antennal seg- ment I surpassing anteclypeus by about 7) its length. Labial segment I slightly surpassing anterior prosternal margin, II reaching posterior margin of pro- coxa, III reaching about anterior 7:i of mesosternum, IV reaching or slightly surpassing posterior margin of meso- coxa. Pronotum impressed near hu- merus at posterior margin of granulate area; shallowly, distinctly punctate an- terior of callus; postcallar impression moderately punctate; scattered punc- tures on posterior lobe. Scutellum with T-shaped median carina moderately developed, upright and crossbar about equally thick and high. Hemelytral membrane slightly surpassing apex of abdomen. Propleuron impunctate on margins; indistincdy punctate cen- trally; moderately but distinctly punc- tate immediately anterior and posterior of center. Mesopleuron indistinctly punctate except moderately but dis- tinctly punctate just behind center and impunctate along lateral and posterior margins and on acetabula. Metapleu- ron indistinctly punctate except im- punctate acetabula and evaporative area surrounding ostiolar peritreme. Pygophore very slightly concave poste- riorly; apex as in Fig. 56a, median projection subacute apically. Clasper as in Figs. 56i-k; dorsal ridge and lateral projection well developed. Aedeagus similar to that of 5. nigricoronatus (Fig. 58e). Ovipositor similar to S. nigricoro- natus (Figs. 57j, 1); apex of second valvula as in Fig. 57d. Spermatheca as in Fig. 57k. Measurements: 2^, 2i. Length S 7 A, 7.7, 2 8.3, 8.4; abdominal width S 2.7, 2.7, J 2.5, 2.8; head length S .50, .63, ? .65, .65; width across eyes S 2.28, 2.35, 2 2.50, 2.51; pronotal length ^ 1.18, 1.33, *" 1.38, 1.53; pronotal width S 2.23, 2.30, 2 2.61, 2.65; scutellar length ^ 1.11, 1.13, 2 1.23, 1.25; scutellar width ^ 1.25, 1.35, V 1.50, 1.50; length antennal segments I S .52, .52, $ .59, .61, II S 1.41, 1.42, V 1.49, 1.57, III ^ 1.40, 1.42, 2 1.48, 1.54, IV S 1.86, 1.87, $ 1.73, 1.90; length labial segments I S .71, .73, 2 .79, .83, II ^ .65, .68, 2 .78, III S .53, .55, ^ .63, IV ' .58, .60, 2 .68. Comments. — Scopiastella costalis is very similar to 6". militaris with which it The Lygaeinae of Australia 335 forms a north-south species pair. In addition to the characters given in the key the two species differ in the struc- ture of the clasper and the apex of the second valvula. Scopiastella militaris is more strongly punctate. The two spe- cies are more robust than other Aus- traHan members of the genus. The spermatheca figured has the margin of the subapical flange rolleci. As other species in this and related genera show considerable within spe- cies variation in development of this flange this condition may not be typical of the species. The range in Australia is restricted to the far north (Fig. 12a) and it also occurs in New Guinea. Scopiastella clcgans (Distant 1918) Figures 10; 56d, o-q; 57b. Scopiastes elegans Distant, 1918:418; Scudder, 1963:380-381 (part); Sla- ter, 1964a: 190; Slater, 1978:856. 7>/)f,r — Lectotype ., "Kuranda, F. P. Dodd. Mch. '04." in BMNH. Para- lectotypes: 2 ^T , IV, same locality as lectotype, "April 04." in BMNH. Redescription. — Color: The following yellow tinged with orange: head ven- • S. elegans * S. melvillensis «> S, nigricoronatus » S. woodward! o S. walkeri papuas _L -L Fig. 10. Distribution of Scopiastella elegans (Di.stant), .S'. melvillensis (Scudder), -S'. nigricoronatus . sp. nov. .sp. nov., and S. walkeri papuas (Scudder) in .Xusiralia. .S". woodwardi, 336 The University of Kansas Science Bulletin trally and sometimes laterally; anterior pronotal lobe; posterior pronotal lobe except transverse postcallar macula, humerus, and usually posterior mar- gin; prosternum; propleuron except rounded central macula; posterior margin and acetabula of mesopleuron; metapleuron except oblong posterior macula; abdomen; base of antennal segment I; coxae; and trochanters. The following light red: head dorsally and sometimes laterally; ocellus; humerus; usually posterior pronotal margin; clavus; corium; labial segment I; and profemur. The following dark brown to black: transverse, oblong, postcallar macula, not reaching lateral margin of pronotum; scutellum; meso- and meta- sterna; rounded central propleural macula; mesopleuron except posterior margin and acetabula; oblong macula near posterior margin of metapleuron, not including ostiolar peritreme; and meso- and metafemora. The following brown: antenna beyond base of seg- ment I, lightest on segment IV; labial segments II to IV; tibiae; tarsi; and eye. Clavus infuscated apically. Heme- lytral membrane subhyaline, brown basally becoming almost clear apically. Structure: Narrow, elongate, scarcely constricted medially. Head moderately strongly declivent; ante- clypeal bump moderately developed. Antennal segment 1 surpassing ante- clypeus by about V2 its length. Labial segment I surpassing anterior proster- nal margin by about 'A its length, II slightly surpassing procoxa, III reach- ing mesocoxa, IV reaching metacoxa. Pronotum with a few indistinct punc- tures anterior of callus; postcallar im- pression moderately punctate; posterior lobe with few scattered mod- erately coarse punctures, most numer- ous along posterior margin of postcallar macula. Scutellum with T- shaped carina moderately swollen, up- right and crossbar about equally high and thick; lateral fovea well developed, strongly punctate. Hemelytral mem- brane greatly surpassing tip of abdo- men. Propleuron indistinctly punctate centrally; moderately but distincdy punctate immediately anterior to and posterior of center. Mesopleuron indis- tinctly punctate except moderately punctate just behind center and im- punctate along lateral and posterior margins and on acetabula. Meta- pleuron indistincdy punctate except impunctate on acetabula and evapora- tive area. Pygophore convex posteri- orly; apex as in Fig. 56d; median projection blunt apically, thickened dorsally. Clasper as in Figs. 56o-q; dorsal ridge well developed, especially posteriorly; lateral projection well de- veloped, directed slightly posteriorly, close to dorsal ridge. Aedeagus similar to S. mgncoronatus (Fig. 58e). Oviposi- tor similar to S. nigricoronatus (Figs. 57j, 1); apex of second valvula as in Fig. 57b, apical projection short. Sperma- theca similar to S. costalis (Fig. 57k). Measurements: 3?, 4+. Length S 6.6 (6.4-7.0), 5 7.2 (6.8-7.6); abdomi- nal width S 1.8 (1.7-1.8), ? 2.0 (1.9- 2.1); head length i .56 (.48-. 65), $ .59 (.52-. 68); width across eyes S 2.13 (2.08-2.16), $ 2.32 (2.13-2.44); prono- tal length f 1.12 (1.08-1.21), 2 1.24 (1.13-1.34); pronotal width S 1.87 (1.80-1.94), ? 2.10(1.96-2.21); scutel- lar length S .87 (.84-. 90), J .96 (.90- 1.02); scutellar width $ 1.05 (1.00- 1.11), 2 1.16 (1.12-1.24); length an- tennal segments I S .43 (.41-. 47), 2 .45 (.42-. 47); II t 1.38 (1.34-1.41), ? 1.36 (1.29-1.41), III S \A1 (1.39- 1.54), $ 1.52 (1.41-1.72), IV S 1.66 (1.60-1.71), ? 1.68 (1.59-1.79); length labial segments I S .71 (.70-. 74), 2 .78 (.75-. 80), II S .67 (.65-. 69), $ .72 (.71-. 76), III S .57 (.56-. 58), v .60 The Lygaeinae of Australia 337 (.58-. 61), IV S .65 (.62-. 68), . .68 (.66-. 70). Comments. — Scopiastella elegans i.s very similar to both S. nigricoronatus , with which it forms a north-south species pair, and S. woodwardi in general ap- pearance. The three species differ from all known members of the genus in possessing a pale pronotum with a dark brown to black postcallar macula. Sco- piastella elegans may be easily separated from the other two species by the com- bination of yellow ostiolar peritreme and head red dorsally. The lectotype is listed as a male by Slater (1978) due to a typographical error. The specimens recorded by Scudder (1963) from Mt. Tamborine, Monteville, and National Park, Queensland, belong to S. nigricoronatus n. sp. Its range seems to be restricted to northern Queensland (Fig. 10) with most specimens known from the type locality. Scopiastella himertos, sp. nov. Figures 1 1; 12a; 57f. Type. — Holotype ?, "Koumala, Queensland, 18-V-1927" in QM. Description. — Color: The following orangey red: head except U-shaped dorsal mark; anterior pronotal lobe; corium except claval and apical mar- gins; abdomen; base of antennal seg- ment I; labial segment I; base of labial segment II; coxae; trochanters; and femora except apex of metafemur. The following dark brown to black: U- shaped dorsal mark of head; posterior lobe of pronotum; scutellum; clavus; claval and apical margins of corium; thoracic sterna except anterior and pos- terior prosternal margins; thoracic pleura except anterior margin of pro- pleuron and all acetabulae; antennal segment I except base; labial segments II and III except base of II; apex of metafemur; tibiae; and tarsi. The fol- lowing pale orange: anterior and pos- terior prosternal margins; anterior propleural margin; and acetabulae. Eye and ostiolar peritreme brown. Ocellus red. Hemelytral membrane subhyaline, basal angle whitish, other- himertos Fig. 11. Scopiastella himertos, sp. nov. Scale = I mm. dorsal view. 338 The University of Kansas Science Bulletin wise dark brown basally becoming al- most clear apically. Structure: Moderately robust, scarcely constricted medially. Head moderately strongly declivent; ante- clypeal bump moderately developed. Antennal segment I surpassing ante- clypeus by about V2 its length. Labial segment I surpassing anterior proster- nal margin by about 'A its length, II slightly surpassing procoxa. III reach- ing metacoxa, IV absent. Pronotum with a few coarse shallow punctures anterior of callus; postcallar impression finely, distinctly j^unctate; posterior lobe with few scattered punctures, strongest and most numerous near hu- merus at posterior margin of granulate area. Scutcllum with well developed T- shaped carina, upright and crossbar equally developed; lateral fovea well developed, strongly punctate. Hemely- tral membrane scarcely surpassing apex of abdomen. Propleuron mod- erately strongly punctate centrally, otherwise impunctate. Mesopleuron moderately punctate anteriorly, im- punctate posteriorly. Metapleuron with a few indistinct, scattered, punctures. Ovipositor similar to S. nigricoronatus (Figs. 57j, 1); apex of second valvula as in Fig. 57f, notched shallowly, pro- duced strongly dorsally. Spermatheca similar to .S". costalis (Fig. 57k). Measurements: Holotype ^. Length 7.7; abdominal width 2.3; head length .70; width across eyes 2.29; pronotal length 1.39; pronotal width 2.24; scu- tcllar length 1.10; scutellar width 1.31; length antennal segment I .48, II to IV missing; length labial segments I .78, II .73, III .63, IV missing. Comments. — ScopiasleUa himertos is similar to 5". melvillensis in coloration but is larger, darker, and the abdomen is not infuscatcd basally. \\\v apex of the second valvula is very distinctive. The holotype is the only specimen known to me. It lacks labial segment IV and all antennal segments except segment I of the right antenna. Scopiastella melvillensis (Scudder 1963) Figures 10; 57h. Scopiastes melvillensis Scudder, 1963:387. Type. — Holotype 5, "Mellville Island, Northern Territory, W. D. Dodd" in SAM. Redescription. — Color: The following bright orange: head; anterior lobe of pronotum; apex of clavus; corium ex- cept extreme base and apical margin; prosternum centrally; propleuron ex- cept anterior margin and produced posterior area; abdomen; base of an- tennal segment I; labial segments I and II; coxae; trochanters; and femora ex- cept extreme apex of metafemur. The following dark brown to black: poste- rior pronotal lobe; scutellum; base of clavus; extreme base and narrow apical margin of corium; meso- and meta- pleura except acetabulae; antennal seg- ments I and II except base of I; and labial segments III and IV. Extreme apex of metafemur, tibiae, and tarsi light brown. Produced posterior area of propleuron, meso- and metasterna, and abdominal venter on each side at base infuscated. Anterior and posterior prosternal margins, anterior propleural margin, and acetabulae pale orange. Eye brown. Ocellus red. Hemelytral membrane subhyaline, basal angle whitish, otherwise dark brown basally becoming paler apically. Structure: Rather short, moderately robust. Head moderately strongly de- clivent; anteclypeal bump scarcely developed. Antennal segment I sur- passing anteclypeus by slightly more than Vj its length. Labial segment I surpassing anterior prosternal margin The Lyoaeinae of Australia 339 by about V:? its length, II slightly sur- passing procoxa, III almost reaching mesocoxa, IV reaching metacoxa. Pro- notum with a few indistinct punctures along postcallar impression, otherwise impunctate. Scutellum with upright of T-shaped carina low, very broad, crossbar narrower and slightly higher; lateral fovea shallow, moderately coarsely punctate. Hemelytral mem- brane slightly surpassing apex of ab- domen. Thoracic pleura impunctate. Ovipositor similar to ^. nigricoronatus (Figs. 57j, 1; apex of second valvula as in Fig. 57h; very narrow, acute api- cally. Spermatheca similar to that of 5'. costalis (Fig. 57k). Measurements: Holotype $. Length 6.9; abdominal width 2.0; head length .65; width across eyes 2.24; pronotal length 1.21; pronotal width 2.05; scu- tellar length .88; scutellar width 1.08; length antennal segments I .48, II 1.21, III, IV and apex of II missing; length labial segments I .78, II .71. Ill .59, IV .69. Comments. — Scopiastes melvillensis is somewhat similar to .S". himertos but is smaller and generally lighter in color. The shape of the apex of the second valvula is very distinctive. The species is known only from the holotype female which lacks the left antenna, the right antenna except segment I and most of segment II, and the left hemelytron. The specimen is also slightly distorted by the pin. • Wmonteithi o W.exilis Fig. 12. a. Distribution o( Scopiastella himertos, sp. nov., S. mbncinclus, sp. nov.. S. cuslalis (Hoivaili). ^>">d S. ,ml,tS , 3$. Length S 7.1 (6.6-7.5), $ 8.1 (8.0-8.2); abdomi- nal width ' 1.8 (1.7-1.9), 2 2.1 (2.1- 2.2); head length ' .70 (.65-. 79), ? .80 (.71-. 84); width across eyes S 2.15 (2.04-2.26), l 2.40(2.36-2.45); prono- tal length ' 1.24 (1.14-1.36), x 1.44 (1.40-1.48); pronotal width S 1.95 (1.83-2.05), 2 2.34(2.25-2.36); scutel- lar length ' .92 (.90-. 93), $ 1.02 (.95- 1.05); scutellar width ^ 1.08 (1.01- 1.14), H 1.24 (1.20-1.28); length an- tennal segments 1 ^ .47 (.46-. 48), ? .49 (.48-. 51), II ' 1.50(1.46-1.53), 9 1.58 (1.53-1.64), III ' 1.58, ; 1.71, IV ^ 1 .45, i 1 .63 (III and IV based on one member of each sex); length labial segments I i .81 (.80-. 81), 5 .91 (.89- .93), II S .73, $ .80 (.78-. 83), III S .60 (.60-. 61), •■ .68 (.65-. 70), IV ' .70 (.68-. 71), > .74 (.73-. 75). Measurements holotype ^ . Length 7.5; abdominal width 1.9; head length .79; width across eyes 2.26; pronotal length 1 .36 pronotal width 2.05; scutel- lar length .93; scutellar width 1.14; length antennal segments I .48, II 1.53, III 1.58, IV 1.45; length labial segments I .81, II .73, III .60, IV .71. Comments. — Scopiastella nigricoronatus is the only species with a yellow pro- notum and dark postcallar macula oc- curring in southern Queensland. It is very similar to S. elegans and to S. woodwardi but may be separated from the former by the yellow head with dark vertex and from the latter by its yellow ostiolar peritreme. It is more robust than S. elegans and less so than S. woodwardi. Specimens from Monteville and Na- tional Park, Queensland were listed under S. elegans by Scudder (1963). Scopiastella rubricinctus, sp. nov. Figures 12a; 57i. Type. — Holotype +, "Queensland, Iron Range, Cape York Peninsula, 28-IV/4-V-1968, G. Monteith" in QM. Description. — Color: The following dark brown to black: head dorsally and laterally; posterior pronotal lobe except anterior half medially and at lateral margin; scutellum; clavus; corium ba- sally and apically; and legs. The fol- lowing pale orange: head ventrally, anterior pronotal lobe; anterior half of posterior pronotal lobe medially and at lateral margin; pro- and mesosterna; and thoracic pleura. Broad, transverse median corial fascia, metasternum, and abdomen reddish orange. Eye brown. Ocellus red. Hemelytral mem- brane subhyaline, dark brown basally becoming paler apically. Structure: Elongate, moderately ro- bust, slightly constricted medially. Head moderately strongly declivent; anteclypeal bump very strongly devel- oped. Pronotuin with moderately punctate postcallar impression; poste- rior lobe with few moderately coarse punctures near humerus along poste- rior margin of granulate area. Scu- tellum with well developed T-shaped carina, upright and crossbar thick, equally high, crossbar vaguely defined anteriorly; lateral fovea well devel- oped, moderately coarsely punctate. The Lygaeinae of Australia 343 Hemelytral membrane greatly surpass- ing tip of abdomen. Pro- and meso- pleura impunctate. Metapleuron with lew indistinct punctures slightly behind center. Ovipositor similar to S. nigricor- onatus (Figs. 57j, 1); ape.x of second valvula as in Fig. 57i, only slightly produced apically, with ventral hump. Spermatheca similar to S. costalis (Fig. 57k). Measurements: Holotype H. Length 8.9; abdominal width 2.5; head length .78; width across eyes 2.59; pronotal length 1.59; pronotal width 2.50; scu- tellar length 1.23; scutellar width 1.38; antennae and labium missing. Comments. — Scopiastella rubricinctus is named for the distinct hemelytral fas- cia. It is similar in general coloration to S. cruentus from New Guinea. That species is smaller, has red legs, has only a very narrow black apical corial mar- gin, and has an orange head. I have seen specimens of an undescribed spe- cies from New Guinea similar to S. rubricinctus in size, hemelytral colora- tion, and leg color but with a com- pletely orange head and black meso- and metapleura. The holotype female is the only known specimen. It is in very poor condition. The labium, both antennae, and the right proleg are missing; the head is nearly detached, and the thorax is badly ripped ventrally. It gives one the impression that it is slightly teneral in which case the coloration of the pronotum anteriorly and the thoracic venter may be lighter than in fully mature specimens. Scopiastella walkeri papuas Scudder 1963 Figures 10; 14; 56c, k, r-t. Scopiastes walkeri papuas Scudder, 1963: 396. Types. — Holotype "Kokoda, walkeri papuas Fig. 14. Scopiastella walkeri papuas (Scudder), dorsal view. Scale = 1 mm. (S. R. Monteiih del.) 400m., SE New Guinea, IV-1933, L. E. Cheesman" in BMNH. Para- types: from various localities in New Guinea in ANIC, BMNH, Bishop, Purdue, Scudder, and Hawaiian Sugar Planter's Association collec- tions. Redescription. — Color: The following yellow, often suffused with orange: head laterally and ventrally; anterior pronotal lobe; presternum; propleuron except produced posterolateral area; acetabular margins; and procoxa. The following dark brown to black: broad 344 The University of Kansas Science Bulletin median dorsal macula of head; poste- rior pronotal lobe; scutellum; clavus; corium; meso- and metasterna; pro- duced posterolateral area of pro- pleuron; meso- and metapleura except acetabular margins; antenna; labium; and legs except procoxa. Head above heavily infuscated orange except me- dian macula. Ocellus and abdomen reddish orange, abdomen infuscated basally. Eye brown. Hemelytral mem- brane subhyaline, dark brown basally becoming paler apically. Structure: Short, moderately robust, scarcely constricted medially. Head very strongly declivent; anteclypeal bump strongly developed. Antennal segment I surpassing anteclypeus by about -/i its length. Labial segment I slightly surpassing anterior prosternal margin, II slightly surpassing procoxa, III reaching mesocoxa, IV reaching metacoxa. Pronotum impunctate. Scu- tellum with low T-shaped carina, up- right broad, crossbar narrow; lateral fovea shallow, moderately punctate. Hemelytral membrane slightly sur- passing apex of abdomen. Thoracic pleura impunctate. Pygophore convex posteriorly; apex as in Fig. 56c; me- dian projection subacute apically. Clasper as in Figs. 56r-t; dorsal ridge low, broadly separated from lateral projection. Oxipositor as in ^S". nigricoro- natus (Figs. 57j, 1); apex of second valvula as in Fig. 57g, rather blunt, little produced apically. Spermatheca similar to S. costalis (Fig. 57k). Measurements: \i, 25. Length 5" 6.5, I 7.2, 7.5; abdominal width S 1.7, . 2.1, 2.3; head length ^ .58, ? .62, .68; width across eyes c^ 2.08, $ 2.39, 2.39; pronotal length ' 1.11, ? 1.32, 1.32; pronotal width ' 1.80, ? 2.16, 2.21; scutellar length ' .70, $ .98; scutellar width ' .88, . 1.30; length antennal segments I c^ .43, ? .47, .48, II S 1.22, $ 1.32, 1.34, III S 1.35, 2 1.41, 1.42, IV S 1.58, $ 1.81, 1.86; length labial segments I ? .66, 2 .79, .81, II S .61, . .71, .75, III ^ .45, ? .58, IV S .59, 2 .66. Comments. — Scopiastella walkeri papuas is the only Australian Scopiastella with completely dark hemelytra. The Aus- tralian specimens examined have less orange color in the light parts of the head and thorax than do those from Papua New Guinea. The blade and caudolateral lobe of the clasper are slightly more acute. It is possible that the two forms are not conspecific, but as the differences are minor, and as Scudder (1963) considered larger dif- ferences of only subspecific importance in the same species, it seems better not to split the two groups at present. Examination of a larger series from both Australia and New Guinea will be necessary to establish within species variation needed to resolve the situa- tion. The range in Australia includes only the Cape York Peninsula (Fig. 10). I have also examined specimens from various localities in New Guinea. Scopiastella woodwardi, sp. nov. Figures 10; 57a. Type. — Holotype 2, "Queensland, Forest Road, 8-15 mi. from highway, Ingham to Townsville, 20-III-1961, R. Straatman" in ANIC. Description. — Color: Head and pro- notum, except transverse postcallar macula, yellow irregularly mottled with orangey red. The following yellow: prosternum; propleuron except cen- trally; posterior margin and acetabula of mesopleuron; posterior and narrow lateral margin and acetabula of meta- pleuron; abdomen; base of antennal segment I; coxae; trochanters; and ex- treme bases of femora. The following The Lygaeinae of Australia 345 dark brown to black: transverse, ob- long postcallar macula not reaching lateral pronotal margin; scutellum; clavus except apically and along mar- gin of claval suture; meso- and meta- sterna; mesopleuron except posterior margin and acetabula; metapleuron except posterior and lateral margins and acetabula but including ostiolar peritreme; eye; antenna except base of segment I; labial segments II to IV; meso- and metafemora except extreme bases; tibiae; and tarsi. Corium, labial segment I, profemur, and ocellus or- angey red. Clavus apically and along claval suture and apical margin, and propleuron centrally inluscated. Hem- elytral membrane subhyaline, dark brown basally becoming paler apically. Structure: Elongate, narrow, nearly parallel sided. Head moderately strongly declivent; anteclypeal bump scarcely produced. Antennal segment I surpassing anteclypeus by about 7i its length. Labial segment I slightly sur- passing anterior prosternal margin, II slightly surpassing procoxa, III almost reaching mesocoxa, IV slightly sur- passing mesocoxa. Pronotum with dis- tinctly punctate postcallar impression, posterior lobe with a few indistinct punctures along posterior margin of postcallar macula. Scutellum with up- right and crossbar of T-shaped carina narrow, crossbar indistinct anteriorly; lateral fovea well developed, very coarsely punctate. Hemelytral mem- brane greatly surpassing apex of abdo- men. Propleuron lightly punctate centrally. Meso- and metapleura poste- riorly with a few very indistinct punc- tures. Ovipositor similar to S. mgricoronatus (Figs. 57j, 1); apex of sec- ond valvula as in Fig. 57a, deeply notched. Spermatheca similar to S. cos- talis (Fig. 57k). Measurements: Holotype ?. Length 7.6; abdominal width 2.1; head length .75; width across eyes 2.35; pronotal length 1.48; pronotal width 2.25; scu- tellar length .95; scutellar width 1.15; length antennal segments I .45, II 1.41 (1.79), III 1.49 (2.64), IV 1.55; length labial segments I .75, II .75, III .53, IV .64. Comments. — Scopiastella woodwardi is named for Dr. T. E. Woodward who provided me with the opportunity to undertake this revision. This species is very similar to S. elegans and 6". mgricoronatus. It differs from both in possession of a black ostiolar peritreme, being slightly more robust and in the distinctly notched apex of the second valvula. The holotype 2 is the only known specimen. This specimen exhibits oli- gomery of the right antenna. Only three segments are present with the apical two elongate (figures in paren- theses in the measurements section) and the basal segment unaffected. The left antenna appears normal. Scopiastes Stal Scopiastes Stal, 1874:98, 100; Scudder, 1963:371; Slater, 1964a: 189; B15te, 1972:411. Abgarus Distant, 1910:313. Astacops Breddin, 1901:78 (part); Ber- groth, 1918:56-57 (part). Type-species Astacops degeeri Stal 1865, monobasic. Diagnosis. — Eyes stalked; head at least as wide across eyes as is pronotum at base; first antennal segment surpass- ing anteclypeus by about half its length; buccula convex, highest anteri- orly, tapering posteriorly to about half distance to pronotal margin; pronotum with at most anterior lobe and narrow anterior margin of posterior lobe gran- ulate; posterior pronotal lobe slightly higher than anterior lobe, densely and 346 The University of Kansas Science Bulletin strongly punctate; spermatheca moder- ately long, sperm pump at least moder- ately elongate and semi-tubular, tube connecting sperm pump and apical bulb nearly parallel sided except api- cally, apical bulb moderately large; shank of clasper nearly at right angle to line through basal opening, caudolateral lobe well developed, often blunt or bent, lateral projection on mesal side overhanging distinct excava- tion, dorsal ridge connects overhang- ing projection with junction of blade and shank; conjunctiva of aedeagus with bifid, pigmented dorsal lobe medi- ally and conical subapical dorsal lobe; ejaculatory reservoir with Y-shaped ba- sal projection; vesica robust through- out, no lateral processes; and gonoporal process without apical or subapical processes. Scopiastes was erected by Stal (1874) who separated it from Astacops Boisdu- val by the strongly punctate pronotum, the less slender antennae, and the first tarsal segment being about equal in length to the other two segments to- gether. Since that time Lygaeinae with stylate eyes and with characters inter- mediate between Astacops and Scopiastes have been discovered. As a result au- thors have either united the two genera or separated them on combinations of characters other than Stal's. The latest attempt to delimit Scopiastes is that of Scudder (1963). In his key to the gen- era of the ''Astacops complex" (pp. 318-319) he lists a number of charac- ters separating the genera. Unfortu- nately almost every character listed under Scopiastes is qualified by "usu- ally." The only consistent character is tlic i)resence in Scopiastes of a " 'wish- bone-like' portion near entry of ducts" on the ejaculatory reservoir. At least part of the difficulty in defin- ing Scopiastes results from the inclusion within the genus of more than one group sufficiently distinct to be ac- corded generic status. One such group, Scopiastella Slater, is raised to generic status in this paper. The Australian members of Scopiastes, including the type species S. degeeri (Stal), form a homogenous and easily defined group. Scopiastes is unlikely to be confused with any Australian Lygaeinae except those genera whose members have styl- ate eyes. It may be separated from Astacops and Scopiastella by the strongly, densely punctate posterior pronotal lobe and from Woodwardiastes by the first antennal segment surpassing the anteclypeus by only half its length and the bucculae not angulate anteriorly. The genus contains several north- south species pairs and at least two very similar sympatric species. In contrast to the other genera with stylate eyes, some of the species appear from their distribution to be dry adapted. Redescription. — Color: Variously pat- terned in red, orange, yellow, brown, and black. Structure: Rather narrow and elon- gate, nearly parallel sided, very slightly constricted medially. Thoracic venter and sometimes anterior pronotal lobe finely granulate giving a dull appear- ance. Hairs except on callar region and vittae of abdominal venter (when pres- ent) a mixture of dense, short, semi- decumbent hairs and longer, sparser, upright hairs, the former shorter and the latter longer on venter. Short de- cumbent hairs absent on abdominal vittae. Hairs of callar region long, up- right, usually dense. Thorax and hem- elytra sometimes without mixture of short decumbent and long upright hairs but rather with moderately dense, moderately long upright hairs only, sparsest on thoracic pleura. Head mod- erately strongly declivent, at least as wide across eyes as is pronotum across base; vertex at least slightly impressed The Lygaeinae of Australia 347 anterior of ocellus; eye stalked; ocellus higher than \ertex, at least as far from eye as from other ocellus; anteclypeus directed slightly posterad apically, sub- apical bump present; buccula convex, moderately prominent anteriorly, slop- ing gently posteriorly, reaching about halfway to anterior prosternal margin. Antennal segment I surpassing ante- clypeus by '/i to V2 its length, II or IV longest. Labium not surpassing meta- coxa; segment I thickest, II to IV progressively narrower. Pronotum sub- quadrate, slightly narrower anteriorly than posteriorly, distinctly impressed and slightly constricted behind callus; anterior margin straight; posterior margin slightly convex; lateral margin slightly sinuate; posterior lobe densely, coarsely punctate, not impressed ante- romedial of humerus, a shallow oblique impression at posterior margin lateral of scutellum; callus moderately swol- len, impunctate; callar impression ob- lique, sinuate, unbranched. Scutellum bluntly rounded apically; T-shaped median carina sometimes low, always distinct; lateral fovea sometimes shal- low, always distinct. Hemelytra moder- ately rugose at least apically on corium; granulate; not dull; not completely covering outer latertergites medially; slightly surpassing apex of abdomen. Thoracic sterna impunctate except sometimes prosternum; posterior mar- gins raised. Thoracic pleura impunc- tate to strongly punctate; ostiolar peritreme well developed; posterior metapleural margin directed almost dorsoventrally; posterolateral meta- pleural angle not or slightly produced. Abdominal venter with segment II not covered by metapleuron; female ster- num VII cleft to base, slightly concave laterally; male sternum VH slightly concave posteriorly; sterna without dorsolateral gland openings. Abdomi- nal dorsum with paratergites III to Vl distinct; paratergites VII apparently fused with mediotergite and later- otergites VII; apodemes on jKira- tergites where these distinct. Pygophore subcircular in cross section; median projection well developed; tergum IX nearly completely fused with base of pygophore. Clasper with well developed caudolateral lobe some- times bent; blade curved, subcircular in cross section basally, flattened api- cally, acute; shank at right angle to base, interior side excavated with later- ally projecting overhang, overhang with dorsal ridge. Aedeagus with phal- lotheca moderately to strongly pig- mented, median dorsal ridge; phallothecal process simple, projecting slightly dorsally; conjunctiva tubular, moderately robust, at middle with bi- fid, pigmented dorsal lobe, connical dorsal lobe subapically, low dorsal lobe over ejaculatory reservoir; ejaculatory reservoir complete, with Y-shaped ba- sal projection; vesica basally robust, twisted; ring sclerite incomplete, broad basally, narrowed apically; vesica api- cally robust, twisted, lightly pigmented along lateral margin; gonoporal proc- ess moderately long, visibly covered by membrane nearly to apex; secondary gonopore simple, without apical proc- ess. Ovipositor with fust valvula al- most reaching apex of connecting membrane; connecting membrane ol first valve sometimes pigmented, never sclerotized, deeply cleft medially and sublaterally; second valvula produced, sometimes twisted apically; connecting membrane of second valve almost com- pletely sclerotized, cleft medially. Sper- matheca with pigmented apical bulb; tubular section connecting sperm pump and apical bulb pigmented, slightly wider and usually Hanged api- cally; sperm pump at least moderately elongate, usually tubular, sometimes 348 The University of Kansas Science Bulletin pigmented basally; tube basal of sperm pump moderately long. Key to the Australian Members of Scopiastes 1. Anterior pronotal lobe red, or- ange, or yellow 2 r. Anterior pronotal lobe brown to black 8 2. Hemelytra unicolored, brown to black 3 2'. Hemelytra at least partly red or orange 4 3. Second antennal segment long- est; scutellum at least moderately infuscated; posterior pronotal lobe as dark as hemelytra; south- ern Queensland and northern New South Wales . . bicolor Distant 3'. Fourth antennal segment long- est; scutellum yellow; posterior pronotal lobe slightly paler than hemelytra; northern Queensland yorkensis, sp. no v. 4. Corium completely red except sometimes extreme base and nar- row apical margin 5 4'. Corium if almost completely red then with a black triangular macula at apical margin 6 5. Abdominal venter with a dis- tinct longitudinal vitta on each side melampus (Bergroth) 5'. Abdominal venter without lat- eral longitudinal \ittae ajfinis Distant 6. Corium with triangular black macula at apical margin; abdom- inal venter with distinct longitu- dinal vitta on each side degeeri (Stal) 6'. Corium without triangular macula at apical margin; abdom- inal venter without lateral longi- tudinal vittae 7 7. Corium with an oblique, dark brown to black fascia roughly parallel to apical margin; meso- and metapleura dark brown to black obliquus Scudder 7'. Corium with a broad, trans- verse, infuscated fascia facetus, sp . nov. 8. Hemelytra completely brown to black 9 8'. Hemelytra with at least costal corial margins red or orange at base 10 9. Head yellow; longitudinal lat- eral vittae of abdominal venter becoming narrow and indistinct posteriorly scudden, sp. nov. 9'. Head orange; longitudinal lat- eral vittae of abdominal venter about as broad and distinct pos- teriorly as anteriorly turneri Distant 10. Basal half of hemelytra dark brown to black, apical half bright orangey red; legs red with at most bases of femora infuscated 11 10'. Hemelytra variously patterned but never with basal half dark brown to black and apical half orangey red; legs brown or black except sometimes bases of fem- ora 12 11. Abdominal venter completely dark brown to black; pro- and mesofemora lightly infuscated basally hackeri Scudder 1 1 ' . Abdominal venter red laterally at base; pro- and mesofemora not infuscated basally kurandensis, sp. nov. 12. Hemelytra red basally and api- cally, with broad transverse brown fascia medially; abdomi- nal venter with at most a longitu- dinal row of small dark brown to black spots on each side eylandtensis Scudder 12'. Hemelytra variously patterned but never as above; abdominal venter with a distinct, sometimes The Lygaeinae of Australia 349 interrupted, dark brown to black longitudinal vitta on each side . . 13 13. Anterior pronotal lobe finely granulate, appearing dull .... taroomi, sp. no v. 13'. Anterior pronotal lobe shiny except sometimes narrow lateral margins 14 14. Clasper (Figs. 60g-i) very deeply excavated laterally, lateral projection with a ventral ridge posteriorly; apex of second val- vula (Fig. 62b) very strongly pro- duced, very strongly curved mesad; northern Queensland. . perplexus, sp. nov. 14'. Clasper only moderately exca- vated laterally; lateral projection without a posterior ventral ridge; apex of second valvula at most moderately strongly produced and curved mesad; southern Queensland and northern New South Wales 15 15. Antennal segment IV longer than II; smaller {S 5.0-5.5, 2 5.6-6.3 mm long); clasper (Figs. 60m-o) with lateral projection not appearing notched apically impeditus, sp. nov. 15'. Antennal segment II longer than IV; larger (^ 5.6-6.5, ? 6.5-6.9 mm long); clasper with lateral projection appearing notched apically 16 16. Antennal segment II shorter (1.10 mm long); light coloration of hemelytra and legs yellowish orange; clasper (Figs. 60d-r) with caudolateral lobe not bent strongly dorsad, lateral projec- tion not swollen subapically; south-central Queensland .... obscurus, sp. nov. 16'. Antennal segment II longer (1.15-1.28 mm long); light color- ation of head and legs reddish orange; clasper (Figs. 60a-c) with caudolateral lobe bent strongly dorsad, dorsum of lateral projec- tion swollen subapically; south- ern Queensland and northern New South Wales laticeps (Breddin) Scopiastes affinis Distant 1901 Figure 20. Scopiastes qffims Distant, 1901a:533; Scudder, 1963:374; Slater, 1964a: 189. Types. — Lectotype ?, "Western Aus- tralia, Roebuck Bay, J.J. Walker" in BMNH. Paralectotype S, same data as lectotype, in BMNH. Redescription. — Color: The following bright orangey red: head; pronotum except broad posterior margin; clavus; corium; propleuron; abdominal venter except sternum VII and genital seg- ments; base of antennal segment I; and labial segment I. The following dark brown to black: broad posterior margin of pronotum; scutellum; meso- and metapleura; abdominal sternum VII; genital segments; eye; antenna except base of segment I; labial segments II to IV; and legs. Hemelytral membrane subhyaline, dark brown basally becom- ing paler apically. Structure: Moderately robust. Head very strongly declivent; anteclypeal bump strongly produced; ocelli slightly closer to each other than to eyes. An- tennal segment I surpassing anteclyp- eus by about % its length. Labial segment I reaching anterior prosternal margin. Pronotum with posterior lobe very coarsely punctate. Scutellum with upright of T-shaped carina rather nar- row; lateral fovea well developed, very coarsely punctate. Hemelytral mem- brane .slightly surpassing tip of abdo- men. Measurements: Length 4.8; width 350 The University of Kansas Science Bulletin across eyes 1.73; pronotal length 1.27; pronotal width 1.82; length antennal segments 1. 27, II. 82, III .73, IV 1.09. Comments.— I have not seen this spe- cies. The redescription is based on excellent color slides of the dorsum and venter kindly supplied by Mr. W. D. Dolling of the BMNH. The measure- ments are from Scudder (1963). Scopiastes a/finis is similar in appear- ance to S. melampus but is much smaller and lacks the lateral vittae on the ab- dominal venter characteristic of that species. In addition to the type series Scudder lists a male from Connexion Island, Groote Eylandt, in the South Australian Museum. Scopiastes bicolor Distant 1901 Figures 15; 17b; 58c, d; 60p-r; 611; 62g, m. Scopiastes bicolor Distant, 1 90 la: 533- 534; Scudder, 1963:374-375; Slater, 1964fl:189; Slater, 1978:855-856. Scopiastes vitticeps Horvath, 1902:603; Slater, 1964a: 191. Types. — Lectotype ? of bicolor, "Queensland" in BMNH. Paralec- totype $ of bicolor, "Gayndah" in BMNH. Type 2 oi vitticeps, "Rich- mond" in Budapest. Redescription. — Color: The following yellow or bright orange: head except broad dorsal vitta; anterior lobe of pronotum; pro- and metasterna; tho- racic pleura except apex of ostiolar pcritrcme, posterior margins, and ace- tabulae; and abdomen except row of maculae forming interrupted vitta each side of venter and genital segments. The following dark brown to black: broad longitudinal vitta from ocellus to apex of anteclypeus; posterior lobe of pronotum; clavus; corium; mesoster- num; apex of ostiolar peritreme and evaporative area; subquadrate macula bicolor Fig. 15. Scopiastes bicolor Distant, dorsal view. Scale = 1 mm. on each side of sterna III to VII, not reaching posterior margins, sometimes meeting or nearly meeting medially on sterna V to VII; genital segments; eye; antenna; labium; and legs. The follow- ing pale yellow: buccula; posterior pleural margins; acetabulae; and pos- terior margins of abdominal sterna. Ocellus red. Hemelytral membrane subhyaline, dark brown basally becom- ing almost clear apically. Structure: Moderately robust; clothed with intermixed short, decum- bent and long, upright hairs. Head with moderately strongly developed anteclypeal bump; ocelli slightly closer to each other than to eyes. Antennal segment I surpassing anteclypeus by slightly more than V^ its length. Labial segment I slighdy surpassing anterior The Lygaeinae of Australia 351 prosternal margin, II reaching middle of procoxa, III reaching anterior third of mesosternum, IV not or scarcely reaching mesocoxa. Pronotum moder- ately densely, moderately coarsely punctate except callar area and narrow posterior margin impunctate. Scutel- lum with upright of T-shaped carina lower than crossbar, equally thick; lat- eral fovea narrow, strongly punctate. Hemelytral membrane scarcely sur- passing tip of abdomen. Prosternum with a few coarse, shallow punctures. Thoracic pleura indistinctly to moder- ately punctate except posterior margins and acetabulae impunctate. Pygophore as in Figs. 58c, d; subcircular in cross section, convex posteriorly; median projection low, blunt apically, with very thin dorsal crest. Clasper as in Figs. 60p-r; caudolateral lobe broad basally, directed slightly dorsad. Ae- deagus similar to S. degeeri (Fig. 58f); phallotheca moderately pigmented. Ovipositor with first valve similar to S. laticeps (Fig. 62n); second valve as in Fig. 62m; apex of second valvula as in Fig. 62g, strongly convex laterally at apex. Spermatheca as in Fig. 611; sperm pump short, constricted medi- ally, pigmented basally; subapical flange variously developed, sometimes absent. Measurements: 4.^, 10?. Length S 6.8(6.6-7.0), . 7.8 (7.2-8.3); abdomi- nal width i 2.0 (1.9-2.1), V 2.3 (2.1- 2.7); head length S .66 (.62-. 74), ? .71 (.63-. 90); width across eyes ? 2.26 (2.22-2.30), V 2.43 (2.31-2.58); prono- tal length ' 1.30 (1.28-1.36), V 1.46 (1.24-1.65); pronotal width ^ 2.03 (1.99-2.09), V 2.34(2.10-2.52); scutel- lar length ' .90 (.83-. 96), ? 1.03 (.94- 1.20); scutellar width ' 1.09 (1.00- 1.16), i 1.27 (1.08-1.38); length an- tennal segments I i .50 (.47-. 52), $ .52 (.49-.56), II ' 1.54 (1.46-1.64), $ 1.58 (1.50-1.70), III ' 1.43 (1.33- 1.53), ? 1.42 (1.23-1.51), IV ' 1.31 (1.20-1.41), V 1.38(1.19-1.55); length labial segments I ' .61 (.59-. 63), ; .70 (.64-. 78), II / .59 (.58-. 60), . .64 (.61-. 69), III .f .42 (.40-. 45), v .48 (.44-. 51), IV S .59 (.56-. 63), . .64 (.60-. 68). Comments. — Specimens of .V. bicolor with yellow rather than orange mark- ings are very similar in appearance to 6". yorkensis. The two species may be separated by the characters given in the key. The type specimens of vitticeps and bicolor differ from the majority of speci- mens examined in that the light mark- ings are bright orange rather than yellow, the black abdominal maculae are larger, and they are more robust. The types of bicolor are only specimens examined with antennal segment IV longer than III. It is possible that the two forms are not conspecific but I can find no significant difference in the genitalia. In the absence of males with orange coloration I think it best to treat them as one species. A female from Fletcher, Queens- land, matches Scudder's (1963) de- scription of a female in the Bishop Museum collection from Macleay River, New South Wales, in that it has the head completely black dorsally. A specimen from Tweed River, New South Wales, lacks the dark macula of the head. These specimens may also not be conspecific with the types oi bicolor but again I can fmd no signifi- cant genitalic differences. All of these specimens are also females except pos- sibly the Tweed River specimen which is lacking the abdomen. Scopiastes degeeri (Sthl 1865) Figures 16a; 17b; 58f; 59f; 61g-i. m; 62k, n, o. Astacops degeeri Stal , 1 865 : 1 8 7 . 352 The University of Kansas Science Bulletin b degeeri kurandensis Fig. 16. a. Scopiasles degeeri (Stal), dorsal view. b. Scopiasles kurandensis, sp. nov. , dorsal view. Scale = 1 mm. Scopiastes degeeri Stal, 1874:100; Scud- der, 1963:379; Slater, 1964a: 190. Type. — Type ■■ , "Austral, boreal." "Thorey. " in Stockholm. Redescription. — Color: The following bright orangey red: head except broad dorsal vitta; anterior pronotal lobe; clavus except scutellar margin; corium except costal margin at base and trian- gular macula at apical margin; narrow anterior prosternal margin; propleuron except posteriorly and ventrally; ab- dominal venter except broad longitudi- nal vitta on each side from sternum II to VI, segment VII and genital seg- ments; abdominal dorsum except ter- gum VII and genital segments; coxae; trochanters; extreme bases of femora; ocellus; and sometimes base of anten- nal segment I. The following dark brown to black with purplish cast: pos- terior pronotal lobe; scutellum; scutel- lar margin of clavus; costal margin of corium basally; triangular macula at apical margin of corium; thoracic sterna except anterior margin of pro- sternum; propleuron except anteriorly and dorsally; meso- and metapleura; broad longitudinal vitta on each side of abdominal venter; abdominal segment VII; genital segments; eye; antennal The Lygaeinae of Australia 353 segment II (sometimes I and IV); la- bial segment IV (sometimes I-III); metafemur except extreme base; tibiae; and tarsi. Broad vitta from ocellus to apex of anteclypeus, sometimes anten- nal segments I and II, sometimes labial segments I to III, and occasionally meso- and metafemora infu seated brownish red. Ocellus orange. Heme- Ivtral membrane subhyaline, dark brown basally becoming paler apically. Structure: Robust. Clothed with moderately dense, moderately long hairs on thorax and hemelytron. Head with well developed anteclypeal bump; ocelli about 1.2 times as far from eyes as from each other. Antennal segment I surpassing anteclypeus by about 'A its length. Labial segment I slightly sur- passing anterior prosternal margin, II reaching about middle of procoxa, III reaching about anterior Vi of mesoster- num, IV reaching posterior margin of mesosternum. Pronotum very strongly punctate. Scutellum with T-shaped ca- rina rather broad, upright and crossbar equally high, lateral fovea narrow, shallow, heavily punctate. Thoracic pleura moderately, shallowly punctate, less distincdy so centrally. Pygophore convex posteriorly; apex as in Fig. 59f; median projection very strongly pro- duced, subacute apically. Clasper as in Figs. 61g-i; lateral overhang rather high, moderately produced. Aedeagus as in Fig. 44c. Ovipositor similar to S. laticeps (Figs. 62n, o); apex of second valvula as in Fig. 62k. Spermatheca as in Fig. 61m; sperm pump short, ro- tund; subapical flange developed to varying degree. Measurements: 2^, 6?. Length $ 6.9, 5 7.6 (7.0-8.0); abdominal width $ 2.1, 2.2, S 2.5 (2.2-2.7); head length $ .56, .78, V .78 (.64-. 89); width across eyes S 2.30, 2.41, ? 2.51 (2.28- 2.74); pronotal length S 1.34, 1.35, $ 1.54 (1.36-1.70); pronotal width S 2.15, 2.35, V 2.55 (2.26-2.85); scutel- lar length S .91, 1.00, V 1.10 (1.00- 1.24); scutellar width ' 1.05, 1.25, ? 1.37 (1.23-1.56); length antennal seg- ments I S .44, ; .46 (.44-. 48), II i 1.32, . 1.28(1.21-1.36), III $ 1.13, $ 1.19 (1.14-1.28), IV ' 1.20, V 1.35 (1.31-1.38); length labial segments I S .63, .64, V .66 (.63-. 70), II S .56, .58, 5 .59 (.53-. 63), III i .29, .36, i .39 (.35-. 44), IV t .45, .46, v .48 (.46- .53). Comments. — The hemelytral colora- tion oi S. degeeri is similar to that of light specimens of S. laticeps and S. impeditus but both those species are less robust and have the pronotum unicolorous. A single female from New South Wales has the base of the hemelytron com- pletely black almost to the level of the scutellar apex as does S. hackeri, but that species lacks the triangular macula at the apical corial margin. Scopiastes eylandtensis Scudder 1963 Figures 17a; 591, p-r. Scopiastes eylandtensis Scudder, 1963: 381. 7>/?^.— Holotype S, "Groote Eylandt, N. Territory, Australia, N. B. Tin- dale" in SAM. Redescription. — Color: The following reddish orange: head except broad me- dian vitta; clavus at base; corium at base and apical 'A; abdomen except lateral and posterior margins of sterna II to VII; and ocellus. The following dark brown to black with purplish cast: broad longitudinal vitta from ocellus to apex of anteclypeus; pronotum; scutel- lum; thoracic pleura except acetabulae; thoracic sterna; antennal segments I, II, and III; labium; legs; and row of small spots on each side of abdominal venter. Eye, clavus except base, and corium except base and apex brown. 354 The University of Kansas Science Bulletin <» S. obliquus " * S eylandtensis « S. facetus ® S melampus o S degeeri ® S hackeri » S. kurandensis * S. yorkensis Fig. 17. a. Distribution oi Scopiastes impeditus, sp. nov., S. turneri Distant, S. obliquus Scudder, S. eylandtensis Scudder, S. facetus, sp. nov., and 5'. melampus (Bergroth). b. Distribution of Scopiastes bicolor Distant. 5 degeeri {?>tk\), S. hackeri Scudder, S. kurandensis, sp. nov., and S. yorkensis, sp. nov. Acetabulae (narrowly) and lateral and posterior margins of abdominal sterna II to MI pale yellow. Antennal seg- ment IV light brown. Hemelytral membrane subhyaline, dark brown ba- sally becoming almost clear apically. Structure: Slender. Clothed with mixture of short decumbent and very long upright hairs. Head with moder- ately developed anteclypeal bump; ocelli slightly closer to each other than to eyes. Antennal segment I surpassing anteclypeus by about Vj its length. Labial segment I slightly surpassing anterior prosternal margin, II almost reaching posterior margin of procoxa, III reaching about middle of mesostcr- num, IV reaching about middle of mesocoxa. Pronotum with posterior lobe deeply, moderately coarsely punc- tate; anterior lobe finely granulate ap- pearing dull. Scutellum with well developed T-shaped carina; upright and cross bar equally high, equally thick; lateral fovea deep, strongly punctate. Hemelytral membrane mod- erately surpassing tip of abdomen. Thoracic pleura moderately punctate. Pygophore convex posteriorly; apex as in Fig. 591; median projection short, broad, excavated apically. Clasper as in Figs. 59p-r; caudolateral lobe curved dorsad; lateral excavation broad, shal- low, overhang short. Aedeagus similar to S. degeeri (Fig. 58f). Measurements: Holotype u. Length The Lygaeinae of Australia 355 5.7; abdominal width 1.7; head length .55; width across eyes 1.89; pronotal length 1.00; pronotal width 1.70; scu- tellar length .78; scutellar width .95; length antennal segments I .38, II 1.33, III 1.08, IV 1.19; length labial segments I .63, II .58, III .38, IV .45. Comments. — Scopiastes eylandtensis and S. facetus are the only Australian Scopi- astes with a broad dark median fascia on the hemelytra. They are easily sepa- rated by the coloration of the pro- notum, thoracic pleura, and legs. This species is known only from the holotype. facetus Fig. 18. Scopiastes facetus, sp. nov., dorsal view. Scale = 1 mm. (S. R. Monteith del.) Scopiastes facetus, sp. nov. Figures 17a; 18; 61o; 62f. Type. — Holotype ?, "Queensland, Morven District, IV-1941, N. Geary" in AM. Description. — Color: The following reddish orange: head except broad lon- gitudinal vitta; ocellus; anterior pro- notal lobe; prosternum medially; propleuron medially except acetabula; and base of profemur. The following pinkish red: scutellum; clavus; corium; metasternum; mesopleuron except ace- tabula; metapleuron except acetabula and posterior margin; and base of mes- ofemur. The following infuscated: callar area; scutellum; prosternum me- dially; propleuron medially; and broad transverse fascia across hemelytron in- cluding about central '/2 of corium and apical Vi of clavus. The following brown: posterior lobe of pronotum; mesosternum; produced posterior part of propleuron; posterior margin of metapleuron; eye; labium; coxae; and trochanters. The following dark brown to black with purplish cast: broad me- dian vitta from ocellus to apex of ante- clypeus; antenna; femora except base of pro- and mesofemora; tibiae; and tarsi. Anterior prosternal margin, an- terior propleural margin, acetabulae, and abdomen pale orange. Hemelytral membrane subhyaline, dark brown ba- sally becoming almost clear apically. Structure: Slender. Clothed with in- termixed short decumbent and very long upright hairs. Head with moder- ately produced anteclypeal bump; ocelli about 1.2 times as far from eyes as from each other. Antennal segment I surpassing tip of anteclypeus by about '/2 its length. Labial segment I surpass- ing anterior prosternal margin by about Vi its length, II slightly sur- passing procoxa. III nearly reaching 356 The University of Kansas Science Bulletin mesocoxa, IV reaching metacoxa. Pro- notum with posterior lobe deeply, densely, not coarsely punctate; anterior lobe finely granulate, appearing dull. Scutellum with broad T-shaped carina, upright slightly lower than cross bar; lateral fovea shallow; strongly punc- tate. Hemelytral membrane slightly surpassing apex of abdomen. Pro- sternum very indistinctly punctate. Thoracic pleura shallowly, distinctly punctate, deepest on proplcuron. Ovi- positor similar to S. laticeps (Fig. 62n, o); apex of second valvula as in Fig. 62f, produced and curved mesad. Sper- matheca as in Fig. 61o, rather short, thick; sperm pump short, tubular. Measurements: Holotype S. Length 5.7; abdominal width 1.6; head length .53; width across eyes 1.93; pronotal length 1.08; pronotal width 1.71; scu- tellar length .73; scutellar width .96; length antennal segments I .36, II 1.14. Ill 1.03, IV 1.19; length labial segments I .65, II .59, III .40, IV .54. Comments. — Scopiastes facetus is named for the elegance of its appearance. It is somewhat similar in appearance to S. eylandtensis but the latter has completely dark legs and pronotum and antennal segment II is much longer than IV. From the locality data of the only known specimen ^. facetus appears to be adapted to more arid conditions than most members of the genus. Scopiastes hackeri Scudder 1963 Figures 17b; 59j; 61d-f; 62i. Scopiastes hackeri Scudder, 1 963 : 38 1 . Type. — Holotype $, "Nanango Dist., Queensland, XI-1927, H. Hacker" in QM. Redescnption. — Color: The following bright scarlet: head except broad gular area and buccula; extreme apex of clavus; about apical 7:2 of corium except very narrow apical margin; ocellus; antennal segment I; metafemur; pro- and mesofemora apically; and tibiae. The following dark brown to black with purplish cast: gular area; buccula; pro- notum; scutellum; clavus except ex- treme apex; basal V2 and narrow apical margin of corium; thoracic sterna; tho- racic pleura; abdominal venter; anten- nal segments II and III; labium; coxae; and trochanters. Basal V2 of pro- and mesofemora (latter less heavily), tarsi, and sometimes apex of anteclypeus in- fuscated. Antennal segment IV light reddish brown. Eye brown. Hemelytral membrane subhyaline, dark brown ba- sally becoming slightly lighter apically. Structure: Short, moderately robust. Thorax and hemelytron clothed with moderately long, moderately dense up- right hairs, sparser on thoracic pleura. Head very strongly declivent; ante- clypeal bump very well developed; ocelli slightly closer to each other than to eyes. Antennal segment I surpassing anteclypeus by about % its length. Labial segment I slightly surpassing anterior prosternal margin, II reaching about middle of procoxa, III reaching about anterior 'A of mesosternum, IV just reaching mesocoxa. Pronotum with lateral margin of anterior lobe finely granulate, appearing dull; poste- rior lobe deeply, very densely, moder- ately coarsely punctate. Scutellum with crossbar of T-shaped carina very broad, upright slightly narrower and lower; lateral fovea narrow, deep, coarsely punctate. Hemelytral mem- brane almost reaching tip of abdomen. Prosternum indistinctly punctate. Tho- racic pleura moderately densely punc- tate, dorsolateral line of coarser, deeper punctures behind middle of each pleu- ron. Pygophore with posterior margin convex; apex as in Fig. 59j; median projection short, high, subacute, not produced apically, triangular appear- The Lygaeinae of Australia 357 ance if viewed anteriorly. Clasper as in Figs. 61d-f; caudolateral lobe subcon- nical, subacute apically; lateral excava- tion shallow, overhanging projection moderately produced. Aedeagus simi- lar to S. degeeri (Fig. 58f). Ovipositor similar to S. laticeps (Figs. 62n, o); apex of second valvula as in Fig. 62i, bluntly produced ventrally. Spermatheca simi- lar to S. bicolor (Fig. 611). Measurements: 1 c^ , 2i. Length S 6.0, S 5.9, 6.6; abdominal width S 2.1, i 1.9, 2.4; head length :' .61, $ .50-. 66; width across eyes S 2.13, ? 2.13, 2.25; pronotal length S 1.16, ? 1.09-1.33; pronotal width S 1.98, $ 1.95, 2.13; scutellar length i .83, $ .80, .83; scutellar width ' 1.03, 2 1.03, 1.09; length antennal segments I $ .44, .47, II V 1.23, 1.34, III ? 1.25, IV % 1.28; length labial segments I S .54, $ .56, .56, II { .53, i .50, .55, III S .35, i .33, .39, IV ^ .46, i .43, .48. Comments. — Scopiastes hackeri is some- what similar to S. degeeri in coloration but lacks a triangular macula at the apical corial margin and has a com- pletely dark abdominal venter. It is very similar to S. kurandensis with which it forms a north-south species pair. The two species are easily separated by the characters given in the key and by the genitalia. The type specimen is slightly teneral with the dark areas more brown than black. It is the only specimen examined with anything resembling useful lo- cality data. The two other specimens examined are listed only as from Queensland. It is an assumption on my part that its range is restricted to south- ern Queensland and probably northern New South Wales. Scopiastes impeditus, sp. nov. Figures 17a; 19; 59e; 60m-o; 61k; 62c. Scopiastes laticeps Scudder, 1963:382 (part, nee Breddin). impeditus Fig. 19. Scopiastes impeditus, sp. nov. Scale = 1 mm. dorsal view. Scopiastes turneri Scudder, 1963:393-394 (part, nee Distant). Types. — Holotype ;, "Queensland, Imbil State Forest, 5-XI-1957, T. E. Woodward" in QM. Paratypes: \$ , 3$, Same data as holotype, in UQ. Also: New South Wales — H, Gos- ford, 22.x. 1904, W.B.F.; 2^ IJ, Tweed River, 20.x. 1901, W.W.F. (NSWDA); 12, Ulong, East Dor- rigo, W. Heron (AM); 1 j , River- tree, ii.l930, E. Sutton (DPI); 1.^, Dorrigo National Park via Dorrigo, 9-10. iv. 1966, T. Weir (UQ); Queensland — 12, Binna Burra, 13. xi.l943, A. Blombery; \S , 32, Bunya Mountains, 2000', 22. i. 1938, N. Geary (AM); 2', 1 ^, Bunya Mountains, 10.xii.l925, H. Hacker; \S, National Park, 3000', iii.1929, A. J. Turner; 2/, Tambourine Mountain, 29. xi. 1925, H. Hacker (QM); 1^, 12, Fletcher, iii.1932, E. 358 The University of Kansas Science Bulletin Sutton; 12, Killarney Plateau, 4.xi. 1932, H. Hacker; IS, Tambourine, W.A.T.S. (DPI); 1^, IJ, Bunya Mountains, 14.xii.l937, F. A. Per- kins; \S, 15, Fletcher, 28.iii.1929, E. Sutton; 1^, Joalah National Park, 15. ii. 1960, P. R. Webb; 22, Lever's Plateau, 12.iii.l966, B. A. Frangmann; 3S, Lower Ballunjui Falls and Track, Lamington Na- tional Park, 30.x. 1955, T. E. Wood- ward; 3S, 22, Tamborine Mountain, 16. ii. 1960, F. A. Perkins (UQ). Description. — Color: The following reddish orange: head except sometimes broad median vitta; clavus usually at least basally along margin of claval suture; corium usually at least basally along costal margin; abdominal seg- ments II to VII except moderately broad, sometimes interrupted longitu- dinal vitta on each side of venter; coxae; trochanters; pro- and mesofem- ora ventrally except at apex; and meta- femora basally. The following brown to black: broad vitta from front of eye stalk to apex of anteclypeus; pronotum; scutellum; clavus variably; corium var- iably; thoracic sterna except posterior margins; thoracic pleura except ace- tabulae; apex of ostiolar peritreme; lat- eral longitudinal vitta of abdominal venter; genital segments; eye; antenna; labium; pro- and mesofemora dorsally and apically; metafemur except base; tibiae; and tarsi. Antennal segment IV and areas of pro- and mesofemora of- ten Hghtcst. Posterior margins of tho- racic sterna, acetabulae and base of ostiolar peritreme pale yellow. Hemely- tral membrane subhyaline, dark brown basally becoming lighter apically. Structure: Short, moderately slen- der. Clothed with mixture of short, decumbent and longer, upright hairs. Head moderately declivent; ante- clypeal bump only slightly produced; ocellus very slightly farther from eye than from other ocellus. Antennal seg- ment I surpassing anteclypeus by about 73 its length. Labial segment I reaching or slightly surpassing anterior proster- nal margin, II almost reaching poste- rior margin of procoxa. III reaching about anterior Vs of mesosternum, IV reaching about middle of mesocoxa. Pronotum with posterior lobe mod- erately coarsely, moderately densely punctate. Scutellum with crossbar of T-shaped carina higher and thicker than upright; lateral fovea well devel- oped, coarsely punctate. Hemelytral membrane reaching or slightly sur- passing tip of abdomen. Prosternum sparsely, finely, distinctly punctate. Thoracic pleura finely, densely, dis- tinctly punctate. Pygophore convex posteriorly; apex as in Fig. 59e; me- dian projection low, not expanded dor- sally. Clasper as in Figs. 60m-o; caudo- lateral lobe curved strongly dorsad; overhanging lateral projection slightly produced. Ovipositor similar to S. lati- ceps (Figs. 62n, o); apex of second valvula as in Fig. 62c, rather strongly produced, curved mesad apically, ven- ter moderately curved mesad subap- ically. Spermatheca as in Fig. 61k; sperm pump short, curved. Measurements: Length $ 5.3 (5.0- 5.5), 2 6.0 (5.6-6.3); abdominal width $ 1.6 (1.5-1.6), 2 1.8 (1.7-1.8); head length S .52 (.48-. 55), ^ .59 (.51-. 69); width across eyes $ 1.83(1.75-1.90), 2 l.S)8 (1.90-2.05); pronotal length S 1.04 (.99-1.09), 2 1.12 (1.10-1.19); pronotal width $ 1.65 (1.58-1.75), 2 1.88(1.75-2.05); scutellar length $ .72 (.59-. 78), 2 .82 (.74-. 90); scutellar width S .91 (.85-. 98), . 1.03 (.95- 1.15); length antennal segments I S .38 (.35-. 41), 2 .40 (.37-. 42), II ' .98 (.94-1.02), 2 1.01 (.97-1.09), III .^ .93 (.88-. 99), * .96 (.92-1.02), IV S 1.07 The Lygaeinae of Australia 359 (1.01-1.12), V 1.10 (1.02-1.14); length labial segments I ' .52 (.48-. 59), 'i ^59 (.54-. 62), II S .48 (.45-. 51), 9 .55 (.50-. 58), III S .30 (.29-. 32), V .30 (.30-. 31), IV S .42 (.40-. 45), ? .45 (.42-. 49). Measurements holotypc ^ . Length 5.2; abdominal width 1.5; head length .48; width across eyes 1.75; pronotal length 1.02; pronotal width 1.64; scu- tellar length .78; scutellar width .89; length antennal segments I .36, II 1.00, III .94, IV 1.11; length labial segments I .51, II .48, III .29, IV .40. Comments. — The name impeditus is derived from the Latin impedire which is what this species did to progress on this revision. Scopiastes impeditus is very similar to S. laticeps in both color and structure. Specimens with completely brown hemelytra are very similar to speci- mens of iS. turneri with which it forms a north-south species pair. In addition to the characters in the key, which are consistent and will separate most speci- mens easily, it differs from S. laticeps in that the dark markings are generally lighter and more extensive (all laticeps examined had at least some reddish orange on clavus and corium), it ap- pears relatively broader and less at- tenuate posteriorly, and the pilosity is slightly denser. If in doubt of an identi- fication one should refer to the geni- talia, particularly the claspers. That S. impeditus often resembles the sympatric S. laticeps more closely in coloration than it does the more closely related S. turneri suggests the possibility of Mullerian mimicry. Field studies would be necessary to explore this pos- sibility. The specimens listed as S. turneri from Imbil, Queensland, by Scudder (1963) and those from the following localities listed by Scudder (1963) as S. laticeps belong to this species: \6, Goodna; 2^, Tamborinc Mt.; 1 ', 29, Bunya Mts.; 1 '^ , Queensland National Park; and 1 ^ 1 ^ , Bunya Mts. The last two specimens in UQ the rest in QM. Scopiastes kurandensis, sp. nov. Figures 16b; 17b; 59h; 61a-c; 62j. Types. — Holotype ^, "Queensland, Kuranda, MV-1952, C. Oke" in NMV. Paratypes: 1 ^ , 1 v , same data as holotype, in NMV. Description. — Color: The following bright orangey red: head except gular area and buccula; extreme apex of clavus; about apical V2 of corium except narrow apical margin; abdomen later- ally and medially at base; ocellus; an- tennal segment 1: femora; and tibiae except extreme apices. The following dark brown to black: broad gular area; buccula; pronotum; scutellum; clavus except extreme apex; about basal '/2 and narrow apical margin of corium; thoracic sterna; thoracic pleura; ab- dominal venter except medially and laterally at base; and labium apically. The following brown: eye; labium ba- sally; coxae; trochanters; extreme api- ces of tibiae; and tarsi. Hemelytral membrane subhyaline, dark brown ba- sally becoming lighter apically. Structure: As in S. hackeri except: apex of pygophore as in Fig. 59h, median projection convex dorsally; clasper (Figs. 61a-c) with caudolateral lobe thick, blunt, overhanging lateral projection directed more anterad; apex of second valvula as in Fig. 62j, not produced ventrally. Measurements: Holotype .' . Length 5.9; abdominal width 2.0; head length .63; width across eyes 2.06; pronotal length 1.23; pronotal width 2.00; scu- tellar length .85; scutellar width .94; length antennal segment I .45; length labial segment I .58. Measurements of paratype S and $ . 360 The University of Kansas Science Bulletin Length $ 6.1, $ 6.0; abdominal width (? 2.1, $ 2.2; head length $ .70, J .53; width across eyes $ 2.11, ? 2.06; pro- notal length $ 1.25, ? 1.11; pronotal width I 2.00, 2 2.06; scutellar length $ .88, $ .89; scutellar width <^ 1.06, ? .98; length labial segments I ^ .61, ? .58, II 5 .55, III 5 .33, IV $ .45. Comments. — Scopiastes kurandensis is named for the type locality. This spe- cies is very similar in appearance to S. hackeri. The red coloration is more or- ange than in hackeri and the coloration of the profemora, mesofemora, and the abdominal venter is different. Both male and female genitalia of the two species are quite distinctive. The type series represents the only known specimens. The first segment of the right antenna of the holotype is all that remains of the antennae. Scopiastes laticeps (Breddin 1900) Figures 20; 58i-k; 59a; 60a-c; 62a, n, o. Astacops (Scopiastes) laticeps Breddin, 1900:170-171. Scopiastes laticeps Scudder, 1963:382; Slater, 1964a: 190. Type. — Lectotype S. Wales" in Berlin. ^New South taroomi « S. scudder ® S. affinis _L ± -L ■H.. 10. Disirihuiioii o( .Scopia.Ues laticeps Breddin. S. perplexus. sp. nov., A', ubuurus. sp. nov., ^ laroomi. sp. nov., S. scudderi, sp. nov., and .S'. affinis Distant. The Lygaeinae of Australia 361 Redescription. — Color: The following reddish orange: head except sometimes dorsal vitta; clavus at least basally along claval suture; corium at least basally along costal margin; abdomen except longitudinal lateral vittae and genital segments; coxae; trochanters; pro- and mesofemora ventrally except apices; and metafemur. The following brown to black: broad vitta from front of eye stalk to apex of anteclypeus; pronotum; scutellum; clavus to varying degree, never along base of claval su- ture; corium to varying degree, never basally along costal margin; thoracic sterna except posterior margins; tho- racic pleura except acetabulae; apex of ostiolar peritreme; longitudinal vitta laterally on abdominal venter; genital segments; eye; antenna; labium; pro- and mesofemora dorsally and apically; metafemur except base; tibiae; and tarsi. Antennal segment IV and pro- and mesofemora often lighter. Poste- rior margins of thoracic sterna, acetab- ulae, and base of ostiolar peritreme pale yellow. Hemelytral membrane subhyaline, dark brown basally becom- ing lighter apically. Structure: Narrow. Clothed with in- terspersed short, semidecumbent and longer, upright hairs. Head moderately strongly declivent; anteclypeal bump moderately produced; ocelli slightly farther from eyes than from each other. Antennal segment I surpassing ante- clypeus by about V2 its length. Labial segment I slightly surpassing anterior prosternal margin, II almost reaching posterior margin of procoxa. III reach- ing about middle of mesosternum, IV reaching about middle of mesocoxa. Pronotum with moderately coarsely, moderately densely punctate posterior lobe. Scutellum with upright of T- shaped carina lower than crossbar; lat- eral fovea narrow, deeply punctate. Prosternum indistinctly punctate. Tho- racic pleura finely, very distinctly punctate. Pygophore slightly concave posteriorly; apex as in Fig. 59a; me- dian projection high, narrow, widened dorsally. Clasper as in Figs. 60a-c; caudolateral lobe curved strongly me- diodorsad; overhanging lateral projec- tion projecting farther at anterior than at posterior end, slightly bifid, swollen subapically dorsally. Aedeagus similar to S. degeeri (Fig. 58f); phallothecal process as in Fig. 58k; ejaculatory res- ervoir as in Figs. 58i, j. Ovipositor as in Figs. 62n, o; apex of second valvula as in Fig. 62a, strongly curved mesad apically, ventral margin curved strongly mesad subapically. Sperma- theca as in Fig. 61j; sperm pump long, tubular, unpigmented. Measurements: Length ^ 6.1 (5.6- 6.5), $ 6.7 (6.5-6.9); abdominal width .^ 1.6 (1.5-1.7), 2 1.9 (1.8-2.0); head length ^ .54 (.40-. 59), i .62 (.50-. 85); width across eyes i 1.98(1.86-2.03), $ 2.15 (2.01-2.24); pronotal length $ 1.11 (.98-1.18), i 1.21 (1.13-1.28); pronotal width S \.ll (1.61-1.88), $ 2.00(1.91-2.09); scutellar length ' .82 (.71-. 89), ? .89 (.83-. 98); scutellar width $ .97 (.86-1.08), 5 1.13 (1.06- 1.20); length antennal segments I $ .41 (.38-. 44), S .42 (.38-. 44), II $ 1.21 (1.15-1.28). 5 1.20 (1.15-1.25), III i 1.10 (1.01-1.22), $ 1.15 (1.11-1.18), IV ^ 1.10 (1.07-1.20), 2 1.10 (1.05- 1.15); length labial segments I i .59 (.56-. 65), $ .64 (.61-. 68), II S .55 (.50-. 63), $ .61 (.58-. 63), III i .35 (.29-. 38), 2 .38 (.36-. 40), IV ' .50 (.45-. 56), 2 .53 (.50-. 55). Comments. —Scopiastes laticeps shows great variability in the extent of the dark hemelytral markings. Most speci- mens examined have the inner half of the clavus and the outer half of the corium red. Some specimens have no dark areas at all on the hemelytron and 362 The University of Kansas Science Bulletin some have the entire hemelytron, ex- cept the base along the claval suture and costal margin, dark. The light areas of the venter are usually more orange than those of the dorsum. This species is similar to S. impeditus, S. obscurus, S. perplexus, and S. taroomi in general appearance. Dark specimens are somewhat similar to S. turneri. Some individuals have hemelytral markings similar to those of S. degeeri. The characters given in the key will serve to distinguish most specimens. In doubtful cases one must resort to the clasper, especially the projecting lateral overhang. This is quite distinct from all other species except S. obscurus where the shape of the caudolateral lobe is markedly different. Scopiastes laticeps is apparently re- stricted to rainforest areas of south- eastern Queensland and northern New South Wales. It forms a north-south species pair with 6". perplexus. Some of the specimens listed as members of this species by Scudder (1963) are referable to S. impeditus (which see) and I believe that the specimens he lists from north- ern Queensland will prove to belong to 5". perplexus. Scopiastes melampus Bergroth 1918 Scopiastes nigriceps Distant, 190 la: 533 (preoccupied). Astacops melampus Bergroth, 1918:57 (new name for S. nigripes Distant). Scopiastes melampus Scudder, 1963:386- 387; Slater, 1964a: 190. Type.— Type V. "Gayndah, Queens- land" in BMNH. Redescription. — Color: The following scarlet: head; anterior pronotal lobe; clavus except extreme base and narrow scutellar margin; corium except ex- treme base and narrow apical margin; about anterior 7:5 of prosternum; pro- pleuron anteriorly and medially; abdominal sterna II to VI except longi- tudinal vitta on each side; ocellus; and extreme base of antennal segment I. The following dark brown to black: posterior pronotal lobe; scutellum; ex- treme base and narrow scutellar mar- gin of clavus; extreme base and narrow apical margin of corium; thoracic sterna except prosternum anteriorly; propleuron posteriorly; propleural ace- tabula; meso- and metapleura; longitu- dinal vitta on each side of abdominal venter, broader and darker posteriorly, nearly meeting medially on sternum VI; sternum VII; and genital seg- ments. Eye, antenna except extreme base of segment I, labium, and legs dark brown suffused with red. Hemely- tral membrane subhyaline, dark brown basally becoming paler apically. Structure: Moderately robust. Head moderately strongly declivent; ante- clypeal bump strongly produced; ocelli slighdy closer to each other than to eyes. Antennal segment I surpassing anteclypeus by about V2 its length. Labial segment I slightly surpassing anterior prosternal margin, II almost reaching posterior margin of procoxa, III almost reaching middle of mesoster- num, IV reaching posterior margin of mesocoxa. Pronotum with extremely coarsely, densely punctate posterior lobe. Hemelytral membrane moder- ately surpassing tip of abdomen. Measurements: Holotype $. Length 8.5; width across eyes 2.33; pronotal length 1.59; pronotal width 2.42; length antennal segments I .42, II 1.09, III .92. Comments. — I have not seen this spe- cies. The description is based on excel- lent color slides of the dorsum and venter kindly supplied by Mr. W. D. Dolling of the British Museum (Natu- ral History). The measurements are from Scudder (1963). Scopiastes melampus is a very distinc- The Lygaeinae of Australia 363 tive species. It is somewhat similar to S. degeeri but lacks the triangular macula at the apical corial margin and the partially red legs of that species. The dorsal coloration is similar to that of\V. affinis which lacks the lateral vittae of the abdominal venter. Scopiastes obliquus Scudder 1963 Figures 17a; 21; 59i, m-o; 61 n; 621. Scopiastes obliquus Scudder, 1963:389. Types. — Holotype ?, "Gayndah, Queensland, A. M. Lea" in SAM. Paratypes: Iv, same data as holo- type, in ANIC; 2?, IS, "Eidsvold, Queensland, 6-X-1929, M.J. Mac- kerras"; 1;^, "Eidsvold, Queens- land, 6-X-1929"; 1., "Wallaville, Queensland, T. L. Bancroft" all in ANIC. Redescription. — Color: Head except median vitta, base of clavus, and co- obliquus Fig. 21. Scopiastes obliquus Scudder, dorsal view. Scale = 1 mm. rium except irregular, oblique, median fascia reddish orange. The following orange: anterior pronotal lobe; hu- merus, sometimes lateral margin, sometimes posterior margin of poste- rior pronotal lobe; scutellum except extreme base; lateral and posterior metapleural margins; abdomen; coxae; trochanters; and ovate longitudinal medial macula ventrally on each fe- mur. The following dark brown to black: broad longitudinal vitta from level of ocellus to apex of anteclypeus; posterior pronotal lobe except humerus and sometimes posterior and lateral margins; extreme base of scutellum; clavus except base; oblique irregular corial fascia; thoracic sterna except an- terior and posterior prosternal mar- gins; thoracic pleura except acetabulae, posterior mesopleural margin, and lat- eral and posterior metapleural mar- gins; apex of ostiolar peritreme; eye; antenna; labium; femora except ven- tral maculae; tibiae; and tarsi. The following pale yellowish orange: ante- rior and posterior prosternal margins; acetabulae; posterior mesopleural mar- gin; lateral and posterior metapleural margins; and base of ostiolar peri- treme. Hemelytral membrane subhy- aline, dark brown basally becoming lighter apically. Structure: Robust. Clothed with in- terspersed short, decumbent and long, upright hairs. Head very strongly de- clivent; anteclypeal bump moderately produced; ocelli nearly twice as far from eyes as from each other. Antennal segment I surpassing anteclypeus by slightly more than Vj its length. Labial segment I slightly surpassing anterior prosternal margin, II reaching poste- rior procoxal margin; III slightly .sur- passing center of mesosternum, IV reaching about posterior margin of mesocoxa. Pronotum with densely, very coarsely punctate posterior lobe. 364 The University of Kansas Science Bulletin Scutellum with low, broad T-shaped median carina, crossbar sHghtly higher than upright; lateral fovea reduced, appearing as group of interconnected very coarse, deep punctures. Hemely- tral membrane moderately surpassing tip of abdomen. Thoracic pleura indis- tinctly punctate except coarsely, shal- lowly punctate slightly behind middle; posterolateral metapleural angle pro- duced slightly posteriorly. Pygophore truncate posteriorly; apex as in Fig. 59i; median projection blunt apically. Clasper as in Figs. 59m-o; robust; cau- dolateral lobe elongate, curving dor- sad; lateral excavation with sinuate ridges; lateral projection scarcely exca- vated, dorsal ridge concave. Aedeagus similar to S. degeen (Fig. 58f), phal- lotheca very heavily pigmented. Ovi- positor similar to .S*. laticeps (Figs. 62n, o); apex of second valvula as in Fig. 621, slightly produced, convex ven- trally. Spermatheca as in Fig. 6 In; robust; sperm pump ovate, heavily pig- mented basally. Measurements: 4^, 8^. Length S 8.0(7.9-8.4), . 8.4(7.7-9.0); abdomi- nal width ' 2.4 (2.3-2.5), ? 2.5 (2.2- 2.8); head length .' .68 (.64-. 81), 2 .66 (.53-. 75); width across eyes S 2.50 (2.48-2.56), ? 2.61 (2.45-2.68); prono- tal length ^ 1.49 (1.48-1.51), ? 1.52 (1.42-1.60); pronotal width S 2.40 (2.31-2.52), V 2.57 (2.31-2.78); scutcl- lar length ^1.10 (1.05-1.15), V 1.25 (1.05-1.36); scutellar width S 1.29 (1.25-1.34), $ 1.46(1.25-1.58); length antennal segments I -^ .54 (.53-. 56), ? .54 (.48-. 60), II f 1.53 (1.47-1.58), ? 1.44 (1.35-1.49), III ' 1.32 (1.25- 1.41), i 1.31 (1.29-1.34), IV / 1.38, ? 1.40 (1.31-1.51); length labial seg- ments I ' .73 (.72-. 75), . .83 (.80- .86), II ' .71 (.70-. 74), ; .75 (.71-. 81), III p^.— Holotype ^ "Queensland, Ta- room, XII-1930" in QM. 368 The University of Kansas Science Bulletin Description. — Color: The following orange, palest on abdomen: head ex- cept anteclypeus; clavus; corium; ab- domen except broad longitudinal vitta laterally on venter, sternum VII medi- ally, and genital segments; ocellus; coxae; trochanters; pro- and mesofem- ora; and basal 'A of metafemur. The following dark brown to black: eye; pronotum; scutellum; thoracic sterna and pleura except acetabular margins; genital segments; apical 'A of meta- femur; and metatibia. Anteclypeus, antenna, labium, pro- and mesotibiae, and tarsi brown, antennal segment IV lightest. The following infuscated: cla- vus apically; corium except costal mar- gin; and pro- and mesofemora apically. Hemelytral membrane dark brown ba- sally becoming lighter apically. Structure: Rather narrow and elon- gate. Clothed with interspersed short, decumbent and long, upright hairs. Short hairs absent on granulate, dull areas. Head moderately strongly decli- vent; anteclypeal bump moderately de- veloped; ocelli about 1 'A times as far from eyes as from each other. Antennal segment I surpassing anteclypeus by about % its length. Labial segment I reaching anterior prosternal margin, II surpassing about anterior third of pro- coxa, III slightly surpassing procoxa, IV reaching mesocoxa. Pronotum with anterior lobe and anterior margin of posterior lobe finely granulate, appear- ing dull; posterior lobe very densely, very deeply punctate. Hemelytral membrane slightly surpassing tip of abdomen. Prosternum moderately punctate. Thoracic pleura somewhat coarsely punctate. Pygophore convex posteriorly; apex as in Fig. 59g; me- dian projection short, blunt apically. Clasper as in Figs. 60v-x; caudolateral lobe short, subconical, curving dorsad; deeply excavated laterally with greatly produced projecting overhang. Aedea- gus similar to S. degeeri (Fig. 58f). Measurements. Holotype ^. Length 6.4; abdominal width 1.9; head length .61; width across eyes 2.28; pronotal length 1.18; pronotal width 1.94; scu- tellar length .93; scutellar width 1.08; length antennal segments I .48, II 1.19, III 1.12, IV 1.30; length labial segments I .60, II .53, III .36, IV .46. Comments. — Scopiastes taroomi is named for the type, and only known, locality. The species is similar to S. laticeps and S. perplexus in size and color- ation. It is easily separated from both by the granulate anterior pronotal lobe. From the type locality it appears that S. taroomi is adapted to drier condi- tions than the majority of the genus. Scopiastes turner! Distant 1918 Figures 17a; 62d. Scopiastes turneri Distant, 1918:416; Scudder, 1963:393-394 (part); Sla- ter, 1964^:191. Type. — Type $, "Kuranda, N. Queensland, 366 m, 21-VI/24-VII- 1913, R. E. Turner" in BMNH. Redescription. — Color: The following orange, darkest on head: head except apex of anteclypeus; abdominal sterna II to VI except longitudinal vitta on each side; abdominal sternum VII lat- erally; coxae; trochanters; meso- and metafcmora except apically; and basal third of metafemur. The following dark brown to black: apex of anteclypeus; pronotum; scutellum; clavus; corium except base of costal margin; thoracic sterna; thoracic pleura except acetab- ulae; apex of ostiolar peritreme; ab- dominal sternum VII medially; lateral longitudinal vitta on abdominal sterna II to VI; and genital segments. The following brown: eye; costal corial margin basally; antennal segments II The Lygaeinae of Australi/ 369 to IV, sometimes I; labium; apices of pro- and mesofemora; apical 7i of met- afemur; tibiae; and tarsi. Acetabulae, base of ostiolar peritreme, and some- times antennal segment I pale orangey yellow. Hemelytral membrane subhy- aline, dark brown basally becoming lighter apically. Structure: Small, slender. Clothed with interspersed short, decumbent and long, upright hairs. Head strongly declivent; anteclypeal bump moder- ately strongly produced; ocelli slightly closer to each other than to eyes. An- tennal segment I surpassing anteclyp- eus by about 7i its length. Labial segment I slightly surpassing anterior prosternal margin, II reaching middle of proco.xa. III surpassing about ante- rior V4 of mesosternum, IV reaching middle of mesocoxa. Pronotum with narrow lateral margin of anterior lobe finely granulate, appearing dull; pos- terior lobe moderately coarsely, very densely punctate. Scutellum with mod- erately developed T-shaped carina, crossbar much higher than upright; lateral fovea small, moderately coarsely punctate. Hemelytral membrane slightly surpassing tip of abdomen. Prosternum shallowly, distinctly punc- tate. Thoracic pleura shallowly, moder- ately coarsely punctate, more strongly on propleuron. Ovipositor similar to S. laticeps (Fig. 62n, o); apex of second valvula as in Fig. 62d, flattish, apical projection short, curving somewhat mesad. Spermatheca similar to S. im- peditus (Fig. 61k). Measurements: 2* . Length 5.5, 5.6; abdominal width L6, 1.7; head length .43, .50; width across eyes 1.93, 2.03; pronotal length .98, 1.04; pronotal width 1.70, 1.80; scutellar length .74, .81; scutellar width .91, .99; length antennal segments I .41, .43, II 1.15, 1.15, III 1.08, 1.13, IV 1.25, 1.30; length labial segments I .54, .58, II .53, .53, III .30, .34, IV .46, .46. Comments. — Scopiasles turneri is very similar to 5". scudderi and to dark speci- mens of S. impeditus, forming a north- south species pair with the latter. It differs from scudderi in size (smaller), in the lateral abdominal vittae being un- interrupted and about equally broad throughout, and in antennal segment IV being slightly longer in comparison to segment II. Scopiastes impeditus usu- ally has at least the base of the costal corial margin orange, is slightly larger, and has all antennal segments shorter than those of turneri. The structure of the apex of the .second valvula is dis- tinctive for all three species. I have not seen males of this species. The specimens from Imbil State For- est, Queensland, mentioned by Scud- der (1963) belong to S. impeditus. The species appears to be restricted in its range (Fig. 17a). Scopiastes yorkensis, sp. nov. Figures 17b; 59c; 60J-1; 62h. Types. — Holotype i, "Queensland, Dividing Range, 15 km W. of Cap- tain Billy Creek, Cape York Pen- insula, 4/9-VII-1975, G. B. Monteith" in QM. Paratype 2, "Queensland, Dividing Range, 15 km W. of Captain Billy Creek, Cape York Peninsula, 142°45' E, 11°40' S, 4/9-VII-1975, G. B. Monteith" inUQ. Description. — Color: The following yellow suffused with orange: head ex- cept longitudinal dorsal vitta; anterior pronotal lobe; scutellum; thoracic sterna except mesosternum laterally; thoracic pleura except acetabulae and posterior margins; and abdominal seg- ments II to VII except interrupted lateral vitta. The following dark brown 370 The University of Kansas Science Bulletin to black: broad dorsal vitta from ocellus to apex of anteclypeus; clavus; corium; apex of ostiolar peritreme; evaporative area; subquadrate macula laterally on abdominal sterna III to VII, not reaching posterior margins, forming interrupted vitta; genital seg- ments; eye; antenna; labium; and legs. Mesosternum infuscated laterally. Pos- terior pronotal lobe brown, darker at humerus. Acetabulae and posterior margins of thoracic pleura pale yellow. Ocellus orange. Hemelytral membrane subhyaline, dark brown basally becom- ing nearly clear apically. Structure: Very similar to S. bicolor except: long upright hairs of dorsum longer, sparser; apex of second valvula as in Fig. 62h; apex of pygophore as in Fig. 59c, median projection higher, slightly thickened dorsally; and clasper as in Figs. 60j-l, caudolateral lobe bent dorsad, lateral overhang more pro- duced, dorsal ridge lower. Measurements: Holotype ^, para- type S. Length i 6.9, + 8.1; abdomi- nal width ' 2.0, : 2.3; head length S .64, i .75; width across eyes S 2.48, ? 2.70; pronotal length $ 1.31, x 1.54; pronotal width ? 2.13, ? 2.55; scutel- lar length ^ .93, 5 1 .06; scutellar width S 1.13, ♦ 1.30; length antennal seg- ments I ^ .52, 'i .54, II S 1.48, ? 1.62, 111 ' 1.33, V 1.54, IV ' 1.65, $ 1.86; length labial segments I ' .60, ? .68, II i .55, i .60.' Ill ' .40, ; .48, IV $ .60, V .64. Comments. — Scopiastes yorkensis is named for the type locality. It may be separated from the very similar S. bi- color by the characters given in the key, the generally lighter dorsum, and the genitalia. The species is known only from the type series. Thunbergia Horvath Caenocons s.g. Thunbergia Horvath, 1914:633; Slater, 1964^:44. Thunbergia Slater, 1978:854. Type-species Alydus marginatus Thun- berg 1822, monobasic. Diagnosis. — Antennal segment IV much longer than II; head swollen behind eye; forefemur of both sexes with short stout subapical spine; ante- rior pronotal margin raised, forming distinct collar; meso- and metapleura impunctate or nearly so; aedeagus with conjunctiva greatly produced laterally, two lightly pigmented, laterally di- rected, pointed lobes and a lightly pig- mented basally directed lobe on either side, and two very small, basally di- rected, pigmented projections dorsally; and valvulae of ovipositor much re- duced, scarcely laciniate. Thunbergia and Achrobrachys were erected by Horvath (1914) as subgen- era of Caenocoris Fieber. Horvath pro- vided a key to subgenera but placed only the type species to subgenus. Sla- ter (1978) raised both of Horvath's subgenera to generic level and gave generic placements for most species previously contained in Caenocoris. The three genera Achrobrachys, Caenocoris, and Thunbergia are remarkable among the Lygaeinae in having the valvulae of the ovipositor reduced and scarcely lac- iniate. The conjunctiva of the aedeagus is partially sclerotized and at least mod- erately expanded laterally. In Achrobrachys and Thunbergia the con- junctiva is quite similar to that of some coreids and rhopalids. It seems likely that during copulation this expansion allows a firm union of the male and female genitalia in the absence of the long bearing surface normally provided by the laciniate ovipositor. The majority of Thunbergia species occur on islands between Asia and Australia. The generic range is from India in the west, the Philippines in the east and the Queensland-New South The Lygaeinae of Australia 371 Wales border in Australia in the south. Most but not all species are associated with rainforest. Redescription. — Color: Red or orange dorsally. Hemelytral membrane and scutellum mostly black. Thoracic ven- ter mostly black. Abdominal venter red or orange with black transverse vittae. The following often pruinose and so appearing lighter than their surround- ings: thoracic venter except posterior pleural margins; gular area; and base of scutellum. Structure: Large, elongate. The fol- lowing finely granulate appearing dull: thorax; scutellum; clavus; corium; base of head dorsally; and gular area. Abdo- men, remainder of head, antenna, la- bium, and legs shining. Body densely covered with moderately to very long hairs, these shortest on base of pro- notum and on hemelytra, finest on thoracic pleura, semidecumbent on ab- dominal venter. Dense fringe of short posteriorly directed hairs on posterior pronotal margin. Head scarcely decli- vent, convex; buccula moderately prominent and curved mesad anteri- orly, very low posteriorly, ventral mar- gin nearly straight, almost reaching anterior prosternal margin; eye consid- erably removed from anterolateral pro- notal angle; head behind eye moderately swollen; ocelli much closer to eyes than to each other. Antennal segment I scarcely surpassing ante- clypeus; IV considerably longest, blunt apically; I and IV slightly thicker than II and III. Labium surpassing meta- coxa; segment I slightly thicker than II; II slightly thicker than III and IV; IV slightly swollen subapically. Pronotum convex; much narrower anteriorly than posteriorly; coarsely deeply punctate behind callus except at posterior mar- gin; median carina faintly indicated; anterior margin concave, swollen, forming distinct collar; posterior mar- gin straight to slightly concave medi- ally, produced slightly to rear laterad of scutellum; lateral margin convex ante- riorly, concave medially, convex to- wards humerus; slightly but distinctly explanate at least anteriorly; callus moderately swollen, separated from an- terior margin by very short depressed punctate area, nearly at level of hemelytra dorsally; callar impression oblique, sinuate, unbranched; poste- rior lobe much higher posteriorly than anteriorly, separated from callus by interrupted deeply punctate impres- sion, a nearly transverse impression at posterior margin near humerus. Scu- tellum blunt apically; upright of T- shaped carina slightly swollen apically, strongly swollen basally, crossbar very strongly swollen; lateral fovea in angle of T deep basally scarcely indicated apically, coarsely and deeply punctate. Hemelytron with costal margin nearly straight, slightly constricted opposite claval commissure; veins slightly prom- inent; membrane surpassing tip of ab- domen. Prosternum with anterior margin raised, a few coarse punctures immediately behind anterior margin. Meso- and metasterna impunctate, swollen, longitudinally sulcate medi- ally. Propleuron coarsely punctate an- teriorly and posteriorly, impunctate centrally; anterior margin raised. Mesopleuron impunctate; a deep post- erolateral pit almost connected to lat- eral (dorsal) margin by a faintly defined curved impression perhaps analogous to the anterolateral gland openings of abdominal sterna of other lygaeines. Metapleuron impunctate; posterior margin truncate to slightly convex, directed dorsovent rally; ostio- lar peritreme well developed. Femora scarcely swollen; Ibrefemur of both sexes with short stout subapical spine. Abdominal sternum II almost com- pletely visible laterally; VI as long me- 372 The University of Kansas Science Bulletin dially as laterally; male VII concave posteriorly; female VII cleft to base, membranous medially, posterior mar- gin concave laterally; sterna without dorsolateral gland openings. Abdomi- nal dorsum with fused para- and medi- otergites; outer and inner laterotergites distinct except fused together and with median tergites on segment VII; poste- rior margin of median tergite I con- cave, of II and III moderately convex, of IV produced posteriorly at middle, of VI truncate, of VII concave; tergum \'ll laterally rounded and produced posterad. Pygophore broadly, shal- lowly excavated dorsally; posterior margin produced broadly posterad; median projection well developed, low; a lateral projection of lateral margin of dorsal opening just anterior to inser- tion of clasper. Clasper without caudolateral lobe; blade curved, twisted, subcircular in cross section basally, becoming flatter apically; shank variously excavated ante- rolaterally. Aedeagus with moderately sclerotized phallotheca; phallothecal process scarcely produced, C-shaped ridge dorsally; conjunctiva large, ro- bust, cylindrical at base, greatly ex- panded with four pigmented acute projections on each side; low lobe over ejaculatory reservoir; ejaculatory reser- \oir complete, with broad Y-shaped basal projection; vesica basally thin, twisted; ring sclerite at most indicated by hght sclerotization; vesica apically coiled and fused, forming pigmented cup; gonoporal process long, coiled, without visible surrounding mem- brane, contained in vesical cup when retracted. Ovipositor with very re- duced valvulae, rami very short; con- necting membrane of first valve lightly pigmented, cleft to about Vi distance from base of first valvula; second val- vulae broadly separated, connected only at base. Spermatheca with pig- mented apical bulb surmounting circu- lar llange; sperm puinp immediately basal to flange, unsclerotized, narrow- ing gradually to long thin unsclerotized basal tube. Key to the Australian Species o/Thunbergia 1 . Posterior pronotal margin pro- duced posteriorly into a short triangular "flap" on each side of scutellum; pronotum bright red with no black markings augur (Stal) 1'. Posterior pronotal margin scarcely produced on either side of scutellum (appearing slightly convex laterally, slightly concave medially); pronotum orange or purplish-red, usually with black marking anteriorly 2 2. Pronotum greatly swollen dor- sally at humeral angles tumidus, sp. no v. 2'. Pronotum not swollen dorsally at humeral angles, posterior lobe evenly convex 3 3. Dark purplish-red above; abdo- men greenish yellow torrida (Distant) 3'. Dorsum and abdomen orange . . 4 4. Femora concolorous with tibiae and tarsi; labium reaching or almost reaching abdominal ster- num V marcida, sp. nov. 4'. Femora red, tibiae and tarsi black; labium almost reaching abdominal sternum IV rufifemur, sp. nov. Thunbergia augur Stal 1865 Figures 23; 63a, d, g, j, k; 64c; 65b, d- f; 66a, b. Lygaeus augur ^ikX, 1865:187. Caenocoris augur Stal, 1874:116; Slater, 1964a: 40.' Thunbergia augur Slater, 1978:854, 855. The Lygaeinae of Australia 373 • T. augur o T. rufifemur « T. tumida » T. torrida J. -L Fig. 23. Distribution of Thunbergia augur {Sik\), /' rujijernur, sp. nov., T. tumida, sp. nov., and 7". torrida (Distant). Types. — Lectotype ?, "Moreton Bay." "Stevens." in Stockholm. Paralec- totype V, same data as lectotype, in Stockholm. Redescription. — Color: Bright blood red. The following deep black: basal % of scutellum; thoracic sterna; thoracic pleura except lateral margins; a trans- verse vitta on each side of abdominal sterna III to VI, not reaching lateral margin, usually narrowly connected medially on III to V, covering posterior -A of each sternum except narrow pos- terior margin, reduced on VI; an- tenna; labium; and legs. Eye brown. Ocellus red. Hemelytral membrane black with blue-green irredescence ba- sally; small whitish mark at basal an- gle; shading from dark brown through subhyaline brown to hyaline apically. Hairs dorsally dark brown to black; ventrally whitish except brown to black on and around transverse abdominal vittae; brown to black on legs, lighter on tarsal pads and antennal segment IV. Structure: Callus separated from pronotal collar by single row of punc- tures; punctures on posterior pronotal lobe often coalescent; pronotal emar- gination only slightly developed; pos- terior pronotal lobe evenly convex; 374 The University of Kansas Science Bulletin posteriorly directed projection of poste- rior pronotal margin well developed; transverse impression near posterior pronotal margin just anterior to pos- terior projection; labial segment I slightly surpassing anterior prosternal margin, II almost reaching mesocoxa, III at least reaching center of abdomi- nal sternum III, IV at least reaching center of abdominal sternum IV. Pygo- phore as in Figs. 63a, d. Clasper as in Figs. 65d-f. Aedeagus as in Figs. 63g, 64c, d; ejaculatory reservoir as in Figs. 63j, k. Ovipositor as in Figs. 66a, b. Spermatheca as in Fig. 65b. Measurements: Length ^ 15.2 (13.8-16.2), $ 18.0 (17.0-19.3); ab- dominal width ' 4.2 (3.5-4.5), 5 5.1 (4.7-5.4); head length ' 1.64 (1.44- 1.88), 2 1.82 (1.64-2.06); width across eyes ^ 2.23 (1.94-2.40), ? 2.43 (2.30- 2.54); pronotal length S 2.20 (1.84- 2.40), $ 2.59 (2.30-2.92); pronotal width ^ 3.87(3.41-4.22), 9 4.68(4.38- 4.97); scutellar length ^ 2.12 (1.96- 2.36), ? 2.57 (2.36-2.76); scutellar width ^ 2.32(2.20-2.52), "i 2.81 (2.60- 3.00); length antennal segments I S .75 (.64-. 82), 1 .83 (.78-. 88), II ^ 2.88 (2.54-3.22), 2 3.23 (2.98-3.40), III S 2.61 (2.25-2.97), $ 2.98 (2.79-3.20), IV S 4.06 (3.64-4.39), S 4.43 (4.28- 4.73); length labial segments I S 2.05 (1.88-2.24), V 2.33 (2.24-2.40), II S 2.55 (2.10-3.28), T 2.92 (2.76-3.06), III ^ 3.23 (2.96-4.04), v 3.57 (3.32- 3.80), IV S 1.90 (1.70-2.28), 9 2.00 (1.86-2.16). Comments. — The bright blood red coloration and the development of the posterior pronotal projections separate this species from others in Australia. These two characters together with the red head, completely black first anten- nal segment, and lack of a tooth on each posterior pronotal projection sep- arate it from all members of the genus known to me. Thunbergia augur is restricted to the eastern coastal region of Australia, penetrating as far south as northern New South Wales (Fig. 23). It is associ- ated with rainforest. Specimens have been collected on Parsonsia straminia and Araujia hortorum. Thunbergia marcida, sp. nov. Type. — Holotype ?, "Cape York, Queensland." in SAM. Description. — Color: Pale orange, clavus and corium darker. The fol- lowing infuscated: anterior half of pronotum except collar; basal % of scutellum; gular area; posterior half of prosternum; mesosternum medially; metasternum; center of propleuron; meso- and metapleura except lateral, posterior and ventral margins, and os- tiolar peritreme; abdominal sternum II except middle and lateral margin; and a transverse macula on abdominal sterna III to VII posteriorly, not reach- ing lateral margin, reduced on VII. Labium and legs light orangey brown becoming darker apically. Eye brown. Ocellus red. Hemelytral membrane with white basal angle, otherwise brown basally shading to whitish hya- line apically. Dark markings pruinose. Structure: Hairs short. Callus sepa- rated from pronotal collar by two rows of fine punctures; punctures of pos- terior pronotal lobe shallow, coarse; lateral pronotal margin slightly expla- nate; posterior pronotal margin scarcely produced posteriorly lateral to scutellum, slightly convex laterally, slightly concave medially; crossbar of T-shaped scutellar carina only moder- ately swollen; scutellum acute apically; propleuron narrowly impunctate cen- trally; labial segment I slighdy surpass- ing anterior prosternal margin, II about reaching mesocoxa. III about reaching posterior margin of abdomi- The Lygaeinae of Australia 375 nal sternum III, IV about reaching posterior margin of abdominiil ster- num V; abdominal sterna III and IV distinctly sulcate medially; abdominal sternum V faintly sulcate medially at anterior margin. Measurements: Holotype rC . Length 10.5; abdominal width 2.6; head length 1.54; width across eyes 1.88; pronotal length 1.68; pronotal width 2.72; scutellar length 1.52; scutellar width 1.68; antenna missing; labial segments I 1.48, II 1.72, III 2.42, IV 1.36. Comments. — The type male of Thun- bergia marcida is the only specimen known to me. Both antennae, the left fore tarsus, both mid tarsi, the left hind leg and the pygophore are missing. The specimen appears somewhat ten- eral and I believe that the infuscated areas would be black in a fully mature specimen. The general body color might also become darker orange. The small size and long labium sepa- rate this species from other members of the genus. Thunbergia rufifemur, sp. nov. Figures 23; 24; 63c, f, h, 1, m; 64b, h-j; 65c; 66c, d. Types.— Ho\oiyY>e ^, "Queensland, Richmond, VIII-1965, A. Fenwick" in QM. Paratypes: 2S , 1$, same data as holotype, in UQ; IS, "Al- lora, Queensland, 29-1-1951, J. Letchford"; 1$, "Caloundra, Queensland, 4-1-1958, L. Clark"; \$, "Queensland, Knob Lagoon, 3 miles N.W. of Doomadgee Mission, 22-V-1972, G. B. & S. R. Mon- teith"; 1?, "Queensland, Lake Mondarra, 12 miles from Mt. Isa, 3- XI- 1967, E. M. Exley, on Eucalyptus sp."; 1^, "Toowoomba, 2-VI-1937, N. T. M. Yeats" all in UQ 1?, "Rockhampton" in AM; 1?, "Peak-Downs" and 1 c? , "Sidney" in BMNH. Description. — Color: Pale orange. The following black: callus; base of scutellum except lateral margin; pro- sternum except narrow anterior mar- gin; meso- and metasterna medially; propleuron except narrow anterior, lat- eral, and acetabular margins; meso- pleuron except narrow lateral and acetabular margins; metapleuron rufifemur Fig. 24. Thunbergia rufifemur, sp. nov., dorsal view. Scale = 1 mm. 376 The University of Kansas Science Bulletin except narrow lateral, posterior, and acetabular margins and ostiolar per- itreme; and a transversely ovate mac- ula on each side of abdominal sterna III to VI. The following orangey red: head; basal -A of antennal segment I, at least ventrally; labial segment I dor- sally; and ostiolar peritreme. The fol- lowing purplish-brown: antennal segment I apically antennal segments II to IV; tibiae; tarsi; and sometimes apices of femora. Coxae, trochanters, and femora except sometimes apices red. Abdomen pale orange basally gradually deepening in color to become orangey red at apex. Hairs pale. Hem- elytral membrane with white basal an- gle, otherwise dark brown with bluish irridescence basally becoming hyaline at apex. Structure: Hairs fairly short; callus separated from pronotal collar by dou- ble row of punctures laterally, single row medially; few small punctures me- dially on callus; pronotum only slightly explanate laterally; posterior pronotal margin scarcely produced posteriorly, convex laterally, concave medially; T- shaped scutellar carina with crossbar only moderately swollen; only very center of propleuron impunctate; labial segment I reaching anterior prosternal margin, II reaching middle of meso- sternum. III reaching posterior margin of metacoxa, IV reaching abdominal sternum IV. Pygophore as in Figs. 63c, 1. Clasper as in Figs. 64h-j. Aedeagus similar to T. augur (Figs. 64c, d); phal- lothccal process as in Figs. 63h, 64b; ejaculaiory reservoir as in Figs. 631, m. Ovipositor as in Figs. 66c, d. Sper- matheca as in Fig. 65c. Measurements: AS, 7i. Length S 10.9 (10.6-11.3), V 12.6 (12.3-13.5); abdominal width S 3.2 (3.1-3.3), $ 3.6 (3.2-3.9); head length ' 1.22 (1.18- 1.30), . 1.48(1.36-1.60); width across eyes / 1.88(1.88-1.90), i 2.04(1.92- 2.14); pronotal length S 1.76 (1.72- 1.82), V 2.03 (1.78-2.20); pronotal width i 3.21 (3.10-3.38), 'i 3.70(3.28- 3.92); scutellar length S 1.66 (1.60- 1.72), $ 1.89 (1.64-2.14); scutellar width ? 1.77(1.70-1.86), 2 2.06(1.72- 2.20); length antennal segment I ? .57 (.53-. 62), 5 .61 (.55-. 68), II ^ 1.94 (1.90-1.99), 5 2.08 (1.93-2.27), III S 1.84 (1.76-1.90), i 1.97 (1.81-2.18), IV S 2.48 (2.37-2.56), i 2.48 (2.08- 2.77); length labial segments I ^ 1.34 (1.30-1.36), V 1.45 (1.32-1.46), II S 1.62 (1.52-1.72), $ 1.79 (1.56-1.92), III ? 2.02 (1.92-2.12), V 2.22 (2.08- 2.40), IV S 1.37 (1.32-1.40), ? 1.40 (1.28-1.48). Measurements holotype S . Length 10.6; abdominal width 3.1; head length 1.22; width across eyes 1.88; pronotal length 1.72; pronotal width 3.10; scutellar length 1.60; scutellar width 1.70; length antennal segments I .55, II 1.90, III 1.80, IV 2.37; length labial segments I 1.36, II 1.60, III 2.00, IV 1.40. Comments. — Thunbergia rufifemur is named for the distinctively colored femora. The species is easily recog- nized by the pale orange dorsum, or- angey red head, and bright red femora. A single female lacks the black callar marking and the black marking of the scutellum except two small basal spots. The dark markings on the thoracic venter of that specimen are represented by infuscated areas. The few specimens in collections in- dicate that the range is broad within Queensland, possibly extending into New South Wales (Fig. 23). It is appar- ently an inhabitant of drier areas than the other Australian members of the genus. Thunbergia torrida Distant 1918 Figures 23; 63b, e, i, n, o; 64a, e-g; 65g; 66e, f. The Lygaeinae of Australia 377 Caenocoris torridus Distant, 1918: 176- 177; Slater, 1964a:44. Thunbergw torrida Slater, 1978:854, 857. Types. — Lectotype i, "Townsville, Qld., F. P. Dodd." "Queensland. F. P. Dodd 1904-284." in BMNH. Paralectotypes: 1^, same data as lectotype, in BMNH; Iv, "Cook- town, Philip de la Garde, 1906-89; 2$, 3$, "Queensland, F. P. Dodd, 1904-284" all in BMNH. Redescription. — Color: Dark purplish- red. The following black: anterior V2 pronotum except very narrow anterior margin; scutellum except apex; broad gular area; prosternum except narrow anterior and coxal margins; meso- and metasterna medially; propleuron ex- cept narrow anterior margin and pro- duced posterolateral angle; meso- and metapleura except dorsal, posterior, and acetabular margins and ostiolar peritreme; and transverse fascia not reaching lateral margin near posterior margins of abdominal sterna III to VI. Abdomen except black fasciae greenish yellow, suffused with orange dorsally. Antenna, labium, femora, tibiae, and tarsi dark purplish-brown. Eye brown. Ocellus red. Hemelytral membrane with white basal angle, otherwise black with bronzy irridescence basally be- coming pale hyaline apically. Hairs pale yellow. Structure: Hairs fairly short; callus separated from pronotal collar by irreg- ular row of punctures; pronotum scarcely impressed behind callus, slightly explanate; callus sparsely punc- tate; posterior pronotal margin scarcely produced posteriorly lateral of scutel- lum, convex laterally, concave medi- ally; T-shaped scutellar carina with crossbar moderately swollen; propleu- ron completely punctate; labial seg- ment I reaching anterior prosternal margin, II reaching about anterior Vi of mesosternum, III reaching about middle of mesocoxa, IV almost reach- ing posterior margin of abdominal ster- num III. Pygophore as in Figs. 63b, e. Clasper as in Figs. 64e-g. Aedeagus similar to T. augur {Y\gs. 64d, c); phal- lotheca as in Figs. 63i, 64a; ejaculatory reservoir as in Figs. 63n, o. Ovipositor as in Figs. 66e, f. Spermatheca as in Fig. 65g. Measurements: 4c?, 5?. Length S 11.5 (10.8-12.0), ? 13.9 (12.7-^15.3); abdominal width ' 3.0(2.8-3.3), ; 3.9 (3.5-4.3); head length ' 1.32 (1.20- 1.44), 2 1.52 (1.34-1.68); width across eyes S 1.88(1.78-1.94), 5 2.08(1.96- 2.22); pronotal length ^ 1.78 (1.60- 1.88), V 2.10 (1.74-2.48); pronotal width f 3.02(2.80-3.16), $ 3.72(3.34- 4.12); scutellar length f 1.70 (1.62- 1.84), . 2.03 (1.88-2.16); scutellar width ' 1.72(1.58-1.88), X 2.15(1.92- 2.40); length antennal segments I S .58 (.55-. 59), ? .64 (.58-. 69), II ? 1.99 (1.92-2.14), ? 2.12 (1.96-2.38), III $ 1.91 (1.85-2.06), ? 2.13 (1.95-2.29), IV .f 3.14 (2.90-3.36), ^ 3.30 (2.85- 3.56); length labial segments I ' 1.35 (1.32-1.36), i 1.51 (1.44-1.68), II $ 1.62 (1.54-1.68), V 1.84 (1.78-1.90), III ^ 1.98 (1.78-2.20), . 2.26 (2.10- 2.42), IV i 1.30 (1.26-1.40), ? 1.48 (1.36-1.58). Comments. — The dark purplish-red dorsal coloration and greenish-yellow abdomen separate T. torrida from other Australian members of the genus. It is very similar structurally to T. rufifemur and in structure and color to a possibly undescribed species from Timor but differs slightly in antennal and labial segment measurements. Thunhergia torrida appears to be re- stricted to northeastern Queensland (Fig. 23). I have seen only one speci- men in addition to the type scries. 378 The University of Kansas Science Bulletin Thunbergia tumida, sp. nov. Figures 23; 65a; 66g, h. Type. — Holotype $, "Kuranda, Queensland, F. P. Dodd" in SAM. Description. — Color: Orange, clavus and cerium darkest. The following in- fuscated: pronotum medially immedi- ately behind callar area; basal % of scutellum; apex of clavus; gular area; posterior half of prosternum; mesoster- num medially; metasternum; propleu- ron centrally; mesopleuron posteriorly and on acetabulae; metapleuron ante- riorly and on acetabulae; and a medial, transverse fascia, extending about half- way to lateral margin near posterior margins of abdominal sterna III and IV. Abdomen except fasciae orange. Antenna except basal part of segment I, labium, and legs purplish-brown. Eye brown. Ocellus red. Hemelytral membrane with whitish basal angle, otherwise dark brown with greenish irridescence basally becoming pale hy- aline apically. Dark markings of body pruinose and so appearing lighter than they would otherwise. Structure: Ocellar protuberance very well developed; callar area and pronotal collar connected medially by remnant of median pronotal carina; pronotal punctation fine; posterior pronotal lobe with very pronounced slightly transverse swelling at hu- merus; posterior pronotal margin almost straight; T-shaped scutellar ca- rina with greatly swollen crossbar; hemelytra not completely covering ab- domen apically, scarcely reaching apex of abdomen; labial segment I surpass- ing anterior prosternal margin by about v., its length, II reaching about anterior Vi of mesosternum, III reach- ing about anterior '/( of metacoxa, IV reaching about middle of abdominal sternum III; abdominal sternum III distincdy sulcate medially, IV with faint indication of median sulcus. Ovi- positor as in Figs. 66g, h. Spermatheca as in Fig. 65a. Measurements: Holotype $. Length 16.0; abdominal width 4.7; head length 1.60; width across eyes 2.62; pronotal length 2.60; pronotal width 4.69; scutellar length 2.60; scutellar width 2.80; length antennal segments I .74, II 2.76, III 2.76, IV missing; length labial segments I 2.04, II 2.20, III 2.70, IV 1.76. Comments. — The type female, the only specimen known to me, is unfor- tunately in poor condition. It lacks the right antenna, segment IV of the left antenna, the right fore and hind legs beyond the coxae, the left hind and mid tarsi, and most of the dorsal hairs. It seems likely that the infuscated areas of the dorsum and of the thoracic venter are normally darker and more exten- sive. In particular the entire anterior half of the pronotum except the collar is probably normally blackish as this area is pruinose and in other species of the genus such pruinosity is confined to dark areas. There is no indication that the specimen is teneral. Thunbergia tumida and a closely re- lated species from New Guinea are the only members of the genus with pro- nounced humeral swellings. The New Guinea species is much darker except the legs which are much lighter, espe- cially the coxae and trochanters. It is also somewhat slimmer, more heavily punctate and has a noticeably longer labium. Woodwardiastes, gen. nov. Type-species Woodwardiastes exilis, sp. nov. Diagnosis. — Eyes stylate; head mod- erately declivent; buccula angulate anteriorly; antennal segment I very elongate, surpassing anteclypeus by ''/y The Lygaeinae of Australia 379 its length; metasternum swollen; dor- solateral lobe slightly beyond middle on each side of conjunctiva; ejaculatory reservoir with greatly expanded and flattened Y-shaped basal projection; vesica with lateral projections and hook-shaped pigmented lobe distal of ring sclerite; and spermatheca with elongate apical bulb and with sperm pump indistinctly differentiated from basal tube. Woodwardiastes is somewhat similar to Scopiastes and Scopiastella in general appearance. The elongate first anten- nal segment, angulate bucculae, and the genitalia all serve to separate it from these and other genera with styl- ate eyes. Description. — Color: The following dark brown to black: pronotum; heme- lytra; thoracic sterna except narrow anterior and posterior prosternal mar- gins; legs beyond trochanters except femora ventrally; antennal segments I to III; and extreme base of antennal segment IV. The following yellow: head; trochanters; apices of coxae; femora ventrally; and antennal seg- ment IV except extreme base. Abdo- men bright scarlet. Structure: Elongate, depressed, nearly parallel-sided, venter finely granulate, dull. Hairs very short, de- cumbent, a few slightly longer and upright. Head moderately declivent, slightly impressed anterior of ocellus; eye stylate; ocellus small, about level with vertex, as far from eye as from other ocellus; buccula concave, moder- ately prominent, sharply angulate anteriorly, tapered posteriorly, not reaching prosternum. Antennal seg- ment I surpassing anteclypeus by about ^7 its length; II longest. Labial segment I thickest, not reaching prosternum; segments II to IV progressively thin- ner. Pronotum shiny, subquadrate, narrower anteriorly than posteriorly, slightly constricted and impressed be- hind callus; posterior lobe slightly higher than anterior lobe; anterior margin straight; posterior margin slightly convex; lateral margin slightly sinuate; anterior lobe impunctate; pos- terior lobe very densely, moderately coarsely punctate; callus slightly swol- len; callar impression oblique, slightly sinuate, unbranched, slightly thick- ened near medial end; a shallow im- pression at posterior margin lateral of scutellum. Scutellum subacute api- cally; T-shaped median carina well de- veloped, crossbar strongly swollen; lateral fovea shallow, slightly punctate. Clavus and corium granulate, sub- shining, not covering outer lateroter- gites; costal corial margin slightly sinu- ate. Hemelytral membrane reaching apex of abdomen. Thoracic sterna im- punctate; posterior margins raised; metasternum swollen, sometimes strongly. Thoracic pleura finely, shal- lowly, moderately densely punctate; posterior metapleural margin slightly concave, directed either dorsovenirally or slightly posteroventrally; poste- rolateral metapleural angle slightly produced; ostiolar peritreme well de- veloped. Abdomen without dorsolateral gland openings; sternum II not cov- ered laterally by metapleuron; female sternum VII cleft to base, slightly con- cave laterally; male sternum VTI slightly concave. Pygophore rotund; median projection well developed, nar- row, subacute apically; tergum IX poorly differentiated from membrane of dorsal foramen. Clasper with short caudolateral lobe with acute, sinuate apical process; blade narrow, rounded in cross section, strongly curved, acute apically; shank shallowly excavated, a thin overhanging dorsal projection. Aedeagus with phallotheca strongly sclerotized ventrally, obliquely im- 380 The University of Kansas Science Bulletin pressed on either side ventrally near base, a membranous lateral swelling on each side near base; phallothecal proc- ess short, convex, dorsally directed; conjunctiva tubular, narrowed apically, a subconical dorsolateral lobe on each side slightly distal of middle, low dorsal lobe over ejaculatory reservoir; ejac- ulatory reservoir complete, Y-shaped basal projection with arms greatly ex- panded and flattened; vesica basally robust, subtubular, twisted apically; ring sclerite complete, broad basally, narrowed apically; vesica apically flat- tened, twisted, expanded into two lat- eral projections near ring sclerite, pigmented hook-shaped lobe near mid- dle, coiled and tubular near apex, cov- ered visibly by membrane to apex, twisted subapically; secondary gono- pore flared, without apical process. Ovipositor with narrow first valvula almost reaching apex of connecting membrane, bent inwards subapically; connecting membrane of first valve notched apically, two distinct subme- dian sclerites; second valvula thin, wid- est apically; connecting membrane of second valve strongly pigmented or lightly sclerotized submedially, cleft about half distance to base. Sperma- theca short; apical bulb elongate, pig- mented; subapical flange pigmented; basal tube and sperm pump unpig- mented, indistinctly separated, moder- ately thick. Key to the Species of Woodwardiastes 1. Thoracic pleura orange; abdo- men without longitudinal vitta on each side exilis, sp. nov. r. Thoracic pleura dark brown to black with narrow pale yellow margins; a broad dark brown to black longitudinal vitta on each side of abdomen monteilhi, sp. nov. Fig. 25. Woodwardiastes exilis, sp. nov., dorsal view. Scale = 1 mm. Woodwardiastes exilis, sp. nov. Figures 12b; 25; 67a-i. Type. — Holotype S, "Parada, May, ' 1969, I. E. C, ex light trap." in QM. Description. — Color: The following yellow: narrow anterior and posterior prosternal margins; narrow posterior and acetabular margins of thoracic pleura; and about basal half of meta- femur dorsally. Extreme apex of ante- clypeus and eye brown. Thoracic pleura except narrow posterior and acetabular margins but including ostio- lar peritreme pale orange. Ocellus red- dish orange. The following dark brown to black: abdominal sternum VII ex- cept broad lateral margin; genital seg- ments; a small rounded macula on each side of abdominal sterna V and VI. Structure: Buccula rather broadly angulate anteriorly; only lateral mar- gins of outer laterotergites visible dorsally; metasternum moderately swollen; posterior metapleural margin directed slightly posteroventrally. Scu- The Lygaeinae of Australia 381 tellum acute apically. Labial segment I extending about 'A distance to proster- num, II almost reaching procoxa, III almost reaching posterior margin of procoxa; IV reaching middle of meso- sternum. Pygophore as in Figs. 67h, j. Clasper as in Figs. 67e-g. Aedeagus as in Figs. 67a, b; ejaculatory reservoir as in Figs. 67c, d. Measurements: Holotype S. Length 7.7; abdominal width 2.1; head length .83; width across eyes 2.34; pronotal length 1.48; pronotal width 2.18; scu- tellar length 1.04; scutellar width 1.15; length antennal segments I .87, II 2.06, III 1.60, IV 1.86; length labial segments I .63, II .56, III .49, IV .59. Comments. — Woodwardiastes exilis and M. monteithi are very similar in general appearance. It is possible that the great disparity in size, distinctive ventral col- oration, and structural differences are due to sexual dimorphism. As other genera of Lygaeinae do not show such dimorphism I believe it best to consider them separate species. The name exilis refers to the slender general aspect of the species. The holotype male is the only specimen known to me. Woodwardiastes monteithi, sp. nov. Figures 12b; 68a-d. Type. — Holotype 'i , "Mount Lewis via Julatten, 3,500', 4-V-1970, G. B. Monteith" in QM. Description. — Color: The following pale yellow: narrow anterior and poste- rior prosternal margins; narrow pro- pleural margins; narrow lateral, posterior, and acetabular meso- and metapleural margins; ostiolar peri- treme; and extreme base of antennal segment I. The following dark brown to black: propleuron except narrow margins; mesopleuron except narrow lateral, posterior, and acetabular mar- gins; metapleuron except narrow lat- eral, posterior, and acetabular margins and ostiolar peritreme; abdominal ster- num VII except broad lateral margin; genital segments; and a very broad longitudinal vitta on each side of ab- dominal venter. Trochanters lightly in- fuscated ventrally. Structure: Buccula very acutely an- gulate anteriorly; outer laterotergites almost completely visible; metaster- num strongly swollen; posterior met- apleural margin directed almost dorsoventrally. Scutellum subacute apically. Labial segment I reaching or almost reaching prosternum, II reach- ing procoxa. III slightly surpassing procoxa, IV surpassing about anterior -'/3 of mesosternum. Ovipositor as in Figs. 68a, c, d. Spermatheca as in Fig. 68b. Measurements: Holotype 2. Length 10.4; abdominal width 3.3; head length .95; width across eyes 2.65; pronotal length 1.94; pronotal width 2.91; scutellar length 1.39; scutellar width 1.60; length antennal segments I 1.04, II 2.40, III 1.92, IV 2.08; length labial segments I .76, II .73, III .63, IV .71. Comments. — Woodwardiastes monteithi is very similar to W. exilis in structure and in dorsal coloration. The colora- tion of the venter is quite distinctive. In addition to the great disparity in size it differs from exilis in having the head slightly more declivent, the bucculae more strongly produced and more acutely angulate anteriorly, the first labial segment extending farther pos- teriorly, and the metasternum more strongly swollen. This species is named for the collector of the type specimen. Aspilocoryphus Stal Aspilocoryphus Stal, 1874:117; Slater, 1964a:30, 1964^:47; Hamid and Meher, 1973a:35-36. 382 The University of Kansas Science Bulletin Type-species Lygaeus fasciativentns Stal, 1858 (fixed by Distant, 1904). Diagnosis. — Base of head completely black; ostiolar peritreme black; pos- terolateral angle of metapleuron pro- duced, nearly covering abdominal ster- num II; body with short dense decum- bent hairs; paratergites separated from mediotergites II to VI; phallotheca with lateral spur. The genus Graptostethus, very similar io Aspilocoryphus in general appearance, usually has at least a pale spot at the base of the vertex, has the medio- and paratergites fused, and lacks the phal- lothecal spur. Four of the eight species presently assigned to Aspilocoryphus occur in the oriental region, three are known only from Africa, and one occurs only in Australia. Due to the external similar- ity between Aspilocoryphus and Grap- tostethus, and the lack of recent work on either genus, there is a good chance that some species are at present incor- rectly placed. Slater and Sperry (1973: 131-133) described the nymphs and biology of the African species A. fas- ciativentris ( S t al ) . The generic descriptions in this pa- per are based on characteristics held in common by all their Australian mem- bers. Since there is only one Australian Aspilocoryphus species the species de- scription fulfills that function. I have therefore omitted a generic redescrip- tion for this genus. Aspilocoryphus australicus Stal 1874 Figures 26; 27; 69a-l. Aspilocoryphus australicus Stal, 1874:117; Slater, 1964a:30; Slater, 1978:855. Types. — Lectotype V, "Austral, bo- real." in Stockholm. Paralectotypes, 2?, same data as holotype, in Stock- holm. australicus Fig. 26. Aspilocoryphus australicus Stal, dorsal view. Scale = 1 mm. Redescription. — Color: Pinkish or- ange. The following dark brown to black: head except two narrowly sepa- rated dorsolateral maculae on either side extending from laterad of ante- clypeus to base of eye between ocellus and antenniferous tubercle; large, rec- tangular, longitudinal macula on side of pronotum; scutellum except stem of T-shaped median carina and apex; hemclytral membrane except narrow white border becoming wider at apex, and large white discal macula narrowly connected to border; prosternum pos- teriorly; meso- and metasterna; The Lygaeinae of Australia 383 propleuron centrally; meso- and mcta- pleura anteriorly including ostiolar peritreme; irregular macula at postero- lateral angle of each pleuron; ante- rolateral angles of abdominal sterna II to VII; narrow posterior border of sterna IV to VI; sternum VII except posterolateral margin; antenna; legs; and labium. Dark dorsal markings of head, base of pronotal macula, scutel- lar fovea, membrane, and postero- lateral propleural macula darker than rest. Clavus and corium lightly infus- cated except posterior claval and ante- rior % of costal corial margin; veins more heavily infuscated. Buccula pal- lid. Eye brown. Ocellus red. Structure: Body subovate; finely granulate, dull; legs and antenna more coarsely granulate or finely rugose; la- bium smooth, shiny. Body above and below with dense decumbent pubes- cence; on head and pronotum in- terspersed with a few stout, longer, upright hairs; on abdominal venter with long upright hairs. Hairs nearly absent on base of pronotal macula, scutellar fovea, and posterolateral pro- pleural macula. Head fairly strongly declivent; vertex nearly flat; ocelli much closer to eyes than to each other; eye close to anterior pronotal angle; buccula prominent, gently tapering posteriorly, not reaching anterior pro- sternal margin. Antennal segment I barely surpassing tip of anteclypeus. Labial segment I thickest, scarcely sur- passing anterior prosternal margin, II thinner, reaching procoxa. III yet thin- ner, reaching mesocoxa, IV reaching metacoxa; II and III thickened distally, III and IV similar in thickness. Pro- notum subquadrate; nearly flat; mod- erately punctate anterior to and slightly posterior to callus; lateral margin and callus slightly swollen; anterior margin slightly concave; posterior margin slightly convex; lateral margin almost straight; median carina ob.solete; cailar impression slightly oblique, a short posterior branch at medial end, lateral end bifid. Scutellum with low T-shaped carina; fovea in angle of T shallow, with transverse striations; lateral mar- gin concave apically; apex acute, al- most spinelike. Clavus slightly convex basally, nearly flat apically. Corium with convex costal margin, veins slightly raised. Membrane almost reaching to slightly surpassing tip of abdomen. Prosternum coarsely punc- tate posteriorly. Propleuron coarsely punctate, less strongly centrally, very strongly and rather elongately at poste- rior margin of central dark marking. Mesopleuron with deep, elongate punctations slightly posterior to mid- line, a few punctations in area of poste- rolateral macula. Metapleuron with a few indistinct punctations in area of posterolateral macula; ostiolar peri- treme well developed, with evaporative area anteriorly; posterolateral angle produced; posterior margin nearly straight or slightly concave, directed anteroventrally. Abdominal sternum II almost completely covered by meta- pleuron; III to VII with dorsolateral gland openings; VI slightly shorter me- dially than laterally; VII of female cleft to base, posterior margin slightly con- cave laterally, curving gently to median cleft; VII of male concave posteriorly. Abdominal terga with inner and outer laterotergites II to VI well defined, VII fused and poorly separated from medi- otergite; distinct apodeme present at center of inner margin of each inner laterotergite II to \T; paratergites sep- arated from mediotergites 11 to VI, larger and more distinct posteriorly; mediotergites with apodemes; medi- otergite VII convex in male, concave in female. Pygophore as in Figs. 69e, f; median projection well de\eloped, bluntly rounded apicalK ; well de\cl- 384 The University of Kansas Science Bulletin oped ridge on each side of median projection separated from projection by a deep narrow channel; sHghtly bifid projection at middle of lateral walls of dorsal opening; tergum IX reduced to narrow elongate sclerite. Clasper as in Figs. 69h-j; blade slightly curved, sub- triangular in basal cross section, some- what flattened and hemispherical in apical cross section; shank shallowly excavated anterolaterally, without dor- sal overhang to excavation. Aedeagus as in Figs. 69c, g; phallotheca moder- ately sclerotized, phallothecal process with lateral shelflike projection, a flat- tened lateroventral spur basad of proc- ess; conjunctiva robust, short, with small, dorsolateral, elongate lobes dis- tally and low lobe over ejaculatory reservoir; ejaculatory reservoir as in Figs. 69a, b, complete, with lightly sclerotized basal projection fused with membrane; vesica basally short, single large conical lobe before middle; ring sclerite thick, heavily sclerotized, nar- rowly incomplete basally, produced apically, lightly pigmented subapical lobe and smaller though similar apical lobe; vesica apically robust, irregularly coiled, heavily pigmented on inner margin centrally; gonoporal process short, slightly coiled, covered visibly by membrane nearly to apex; secondary gonopore flaired, with short, stout, heavily pigmented triangular process. Ovipositor as in Figs. 69k, 1; first val- vula almost reaching apex of connect- ing membrane. Spermatheca as in Fig. 69d. Measurements: Length $ 6.6 (6.1- 7.0), i 7.2 (6.7-8.2); abdominal width $ 2.4 (2.1-2.5), $ 2.8 (2.6-3.0); head length $ .78 (.64-. 88), v .78 (.68- 1.04); width across eyes r^ 1.52 (1.44- 1.56), ■- 1.65 (1.54-1.78); pronotal length $ 1.42 (1.28-1.50), v 1.48 (1.36-1.62); pronotal width ^ 2.24 (2.12-2.40), ? 2.53 (2.32-2.72); scutel- lar length ^ .98 (.84-1.08), ? 1.09 (1.00-1.20); scutellar width ^ 1.26 (1.16-1.40), J 1.44 (1.28-1.60); length antennal segments I $ .45 (.40-. 48), i .50 (.47-. 56), II $ 1.05 (1.00-1.12), ? 1.11 (1.03-1.22), III ^ .83 (.78-. 90), ? .86 (.81-. 96), IV $ 1.08 (1.04-1.16), $ 1.09 (1.02-1.20); length labial seg- ments I i .75 (.68-. 80), i .83 (.72- .90), II ^ .64 (.56-. 70), 5 .70 (.60-. 80), III i .59 (.52-. 68), ? .65 (.56-. 72), IV $ .58 (.54-. 64), 2 .60 (.54-. 68). Comments. — This species is easily separated from other Australian Ly- gaeinae by the combination of short, decumbent body hairs, abdominal ster- num II nearly covered by the produced posterolateral angle of the metapleuron and the white discal macula of the hemelytral membrane. There is some variation in the extent of the black markings and in the hue of the lighter parts. In particular, the light markings on the head may be nearly completely obscured and in a few speci- mens all light markings are testaceous or nearly so. Aspilocoryphus mendicus (Fabricius) from the Oriental region is similar to A. australicus in both color and structure but differs in that the lateral light markings of the head occupy a larger area and are not divided into two mac- ulae on either side, the black longitudi- nal maculae of the pronotum are often divided behind the calli, thus forming four subquadrate maculae, and are equally dark anteriorly and posteriorly, the white discal macula of each hemely- tral membrane is less rounded, and the dark borders of abdominal sterna IV to VI occupy greater areas and are pro- duced medially to form an interrupted vitta. The shelflike projections of the phal- lothecal process of mendicus are slightly more produced than those of australicus and have the distal corners more The Lygaeinae of Australia 385 sharply squared off, the gonoporal process is sHghtly thicker, the "wings" of the ejaculatory reservoir are rela- tively smaller, the medial projections of the lateral wall of the dorsal opening of the pygophore are more distinctly bi- fid, the blade of the clasper is slightly shorter and less curved and the poste- rolateral lobe is slightly more globular when viewed laterally. These charac- teristics were consistent in the speci- mens examined. Nothing is known of the biology or food preferences of Aspilocoryphus aus- tralicus. Its distribution (Fig. 27) sug- gests that it prefers warm and dry conditions. Graptostethus Stal Lygaeus s.g. Graptostethus Stal, 1868: 73-74. Graptostethus Stal, 1872:42; Slater, 1964a:49, 1964Z»:51; Hamid and Meher, 1973^:37. Pyrrhobaphus (Graptostethus) Breddin, 1907:43 (part). Type-species Cimex servus Fabricius, 1787 (fixed by Distant, 1904) Diagnosis. — Posterolateral metapleu- ral angle produced, nearly covering abdominal sternum II laterally; base of vertex not black, at least small pale spot present; callar impression unbranched; mesopleuron lightly punctured ante- FiG. 27. Distribution oi Aspilocoryphus australicus Stal. 386 The University of Kansas Science Bulletin riorly, impunctate posteriorly; fused medio- and paratergites; and labium scarcely extending beyond metacoxa. The genus Aspilocoryphus is very simi- lar in general appearance but has dis- tinct para- and mediotergites II to VI and the head completely black basally. Graptostethus contains about 30 de- scribed species, all from the Eastern Hemisphere. It is primarily tropical. A complex of closely related species is present in the Oriental, Australian, and Pacific regions. Revisional work will be necessary before these species can be reliably separated. Of the four Australian species two are endemic, one also occurs in New Guinea, and the other is widespread throughout Af- rica, the Mediterranean region, and the Orient. Redescription. — Color: The Austral- ian species are for the most part quite similar in color pattern. The following description is an attempt to give a basic pattern or template from which the specific color patterns may be derived by reduction, enlargement, or fusion of the dark markings. This does not imply that it is an ancestral color pattern for the genus. Graptostethus cardinalis differs considerably from the pattern. Reddish orange. The following dark brown to black: base of head behind ocellus; tip of anteclypeus; transverse pronotal bar on callus not reaching lateral margin; round macula imme- diately behind callus; transverse bar immediately anterior to posterior pro- notal margin, interrupted medially; scutcllum except apex; claval and corial margins at claval suture; small discal corial spot; short longitudinal corial vitta at base of claval suture; subrec- tangular macula from midpoint to near apex of costal corial margin; posterior half of prosternum; meso- and meta- sterna except posterior margins; rounded macula at posterolateral pleu- ral angles; middle of propleuron; nar- row fascia from middle of propleuron to prosternum just anterior of procoxa; mesopleuron except acetabula and posterior and lateral margins; meta- pleuron except ostiolar peritreme, ace- tabula, and posterior and lateral margins; abdominal venter except lat- eral margin; genital segments; and abdominal terga I and II. Buccula, posterior margins of meso- and meta- sterna, spot at base of vertex, and apical corial margin pale yellow. Hem- elytral membrane dark brown, basal angle and apical margin white. Eye brown. Ocellus red. Basal angle of scutcllum, base of head ventrally, tho- racic sterna, and thoracic pleura except posterolateral pleural maculae and roundish anteromedial propleural area pruinose. Maculae thus appear darker than remainder of pleura. Structure: Somewhat ovate, broad; head, thorax, and hemelytra finely granulate, dull; abdomen, antenna, la- bium, and legs shiny. Head moderately to strongly declivent, moderately con- vex; buccula prominent, sloping grad- ually caudad, not reaching anterior prosternal margin; ocelli much closer to eyes than to each other; eye adjacent to anterior pronotal angle. Antennal segment I surpassing anteclypeus by V3 to V2 its length; II or IV longest. Labial segment I thickest; II slightly thinner than I; III and IV about equally thick, thinner than II; I or III longest. Pro- notum punctate anterior to and just behind callus; anterior margin concave (except in G. cardinalis); posterior mar- gin subsinuate, slightly convex medi- ally; lateral margin nearly straight, slightly convex and swollen posterior of callus, slightly concave anteriorly; callus slightly swollen, transverse; callar impression oblique, sinuate, un- branched; shallow transverse impres- sion behind callus interrupted laterally The Lvgaeinae of Australia 387 by low ridge; shallow impression at posterior margin laterad ol scutellum. Scutellum acuminate; T-shaped me- dian carina well developed; lateral fo- vea in angle of T shallow, transversely striate. Hemelytron incompletely cov- ering outer laterotergites III to VI; veins moderately raised; clavus widest subapically; claval commissure from V2 to subequal in length to scutellum; costal corial margin nearly straight ba- sally, slightly convex apically; apical corial margin straight or slightly con- vex. Prosternum punctate; anterior margin slightly raised. Meso- and met- asterna impunctate; posterior margins raised. Propleuron heavily punctate anteriorly, lightly punctate centrally; anterior margin slightly raised. Meso- pleuron sparsely, often indistinctly, punctate. Metapleuron at most very sparsely and indistinctly punctured; os- tiolar peritreme well developed; poste- rolateral angle produced posteriorly; posterior margin convex and angled anteroventrally in dorsal -Vi, convex and directed dorsoventrally in ventral Vi. Abdominal sternum II nearly cov- ered by metapleuron laterally (except in G. cardinalis); VI of female shorter medially than laterally; VII of female cleft to base, posterior margin slightly concave lateral to cleft; VII of male concave posteriorly; sterna without dorsolateral gland openings. Abdomi- nal terga with fused medio- and para- tergites; inner and outer laterotergites II to VI distinct, VII fused and indis- tinctly separated from median tergite; posterior margin of male tergum VII convex medially; posterior margin of female tergum VII concave medially. Pygophore slightly concave dorsally; median projection well developed; dor- sal opening laterally with ventral pro- jection just posterior to terga, sometimes forming depressed shelf; mesally directed variously sculptured projection just anterior to insertion of clasper. Clasper with moderately to well developed caudolateral lobe; blade curved, triangular in cross section ba- sally, flattened apically; shank deeply excavated anterolaterally, overhang de- veloped into ventrally directed hook. Aedeagus with moderately sclerotized phallotheca; phallothecal process free distally, slightly projecting laterally; conjunctiva moderately elongate, large low lobe on each side at base, long thin lobe on each side dorsolaterally at apex; ejaculatory reservoir complete, without basal projection, covered by low membranous lobe; vesica basally robust, twisted, narrowed at ring scle- rite, a large lateral lobe near ejacula- tory reservoir; ring sclerite complete, wider and "cup like" apically with two elongate "corners"; vesica apically moderately coiled, slightly pigmented apically; gonoporal process moderately long, covered visibly by membrane al- most to apex, less heavily pigmented distally; secondary gonopore with com- plex unpigmented process. Ovipositor with first valvula not reaching apex of connecting membrane; membrane connecting first valve cleft medially to level with first valvula, medial sclerite around cleft; second valvula subcylin- drical apically; connecting membrane of second valve lightly pigmented, cleft almost to base. Spcrmatheca long, ir- regularly coiled, tubular, heavily pig- mented except at base; sperm pump near base of pigmented area, unpig- mented; no flanges. Key to the Australian Species o/Graptostethus 1. Pronotum anterior of calli, in- cluding anterior margin, black; numerous long upright hairs on hemelytra cardinalis Stal r. Pronotum never black anterior of calli; hemelytral hairs at most 388 The University of Kansas Science Bulletin varipictus Fig. 28. Graplostethus spp. a-d. Dorsal view of named .species. These arc diagramatic representations — shape and relative size arc correct but contrast between different colors has been exaggerated. All species show considerable variation in color pattern. Scale = 1 mm. The Lygaeinae of Australia 389 moderately long, scmidecum- bent 2 2. Numerous long upright hairs on pronotum and scutellum; when present, dark markings of pro- notum usually not united, form- ing three separate maculae . . . servus (Fabricius) 2'. At most a few upright hairs of moderate length on pronotum and scutellum; when present, dark markings of pronotum usu- ally united to form single com- plexly shaped macula 3 3. Dorsal hairs very short, fine, and decumbent, usually covering postcallar maculae; dark hemely- tral markings always present; discal spot of corium usually not fused with other markings, if so then with costal macula pubescens, sp. no v. 3'. Dorsal hairs longer, coarser, and only semidecumbent, usu- ally absent on round postcallar maculae; dark hemelytral mark- ings sometimes absent; discal spot of corium usually fused with dark markings of claval suture . varipictus, sp. nov. Graptostethus cardinalis (St'al 1867) Figures 28b; 29; 70a, b, k; 71J-1, p; 72c. Lygaeus cardinalis Stal, 1867:163. Graptostethus (Graptostethus) cardinalis Stal, 1874:117. Graptostethus cardinalis. Slater, 1964^:50- 51. 7>/?^.— Type $, "Austral, boreal." "Thorey." in Stockholm. Redescnptwn. — Color: This species differs so markedly from the gener- alized generic description that a com- plete description is given below. Orangey red. The following dark brown to black: anteclypeus, base of head behind eye and ocellus; pronotum anterior of callus, including anterior margin, not reaching lateral margin; triangular macula behind callus, reach- ing lateral margin at humerus, almost reaching posterior margin and callus; scutellum except basal angle and apex; claval suture; narrow longitudinal vitta from near base of claval suture to mid- dle of corium; broad, short vitta paral- lel to apical corial margin from just beyond middle of costal margin to mid- dle of corium, connected with longitu- dinal vitta; hemelytral membrane except white basal angle and basal and apical margins; large rounded macula at posterolateral angle of metapleuron; posterior margins of abdominal sterna IV to VI, narrowly medially, broadly forming interrupted vitta laterally, not reaching lateral margin; abdominal sternum VII except lateral margin; genital segments; antennal segments II to IV and apex of I; labial segments II to IV and I apically and ventrally; apex of femora; tibiae; and tarsi. Eye dark brown. Ocellus red. Thoracic sterna infuscated except pallid posterior mar- gins. Thoracic pleura infuscated except dorsal (lateral) margins. Ostiolar per- itreme pallid basally, bright reddish orange apically. Structure: Narrow for genus. Body clothed moderately densely with long, fine, upright hairs; body except scutel- lum and disc of posterior pronotal lobe each side of middle densely covered with moderately long semidecumbent hairs, sparse on head, pronotum ante- rior of callus, and region of longitudi- nal corial vitta, shorter on abdominal venter. Head strongly declivent, slightly swollen immediately anterior of eye; buccula reaching or almost reach- ing anterior prosternal margin; eye slightly removed from anterior prono- taf angle. Antennal segment I surpass- 390 The University of Kansas Science Bulletin ing apex of anteclypeus by about half its length. Labial segment I reaching anterior prosternal margin, II reaching procoxa, III reaching about middle of mesosternum, IV reaching metaster- num. Pronotum very strongly punc- tured anterior to and immediately behind callus, slightly constricted and impressed behind callus; anterior mar- gin straight; lateral margin slightly convex anteriorly and posteriorly; cal- lus moderately swollen. Hemelytral membrane almost reaching abdominal apex. Propleuron coarsely punctate an- teriorly and posteriorly, impunctate medially. Mesopleuron with few very indistinct punctures. Metapleuron im- punctate. Pygophore as in Fig. 70k; ventral projection of side of dorsal opening invisible from above; lateral projection broadly bifid, dorsal channel absent. Clasper as in Figs. 71j-l; poste- rolateral lobe short; lateral hook well developed, very acute apically. Aedea- gus as in generic description; ejacu- latory reservoir as in Figs. 70a, b. Ovipositor as in Fig. 71p; first valvula surpassing first ramus. Spermatheca as in Fig. 72c; fairly short; sclerotized part basad of sperm pump moderately long. Measurements: 6S , 3?. Length S 6.8(6.2-7.0), . 7.3 (7.1-7.5); abdomi- nal width ' 2.2 (2.0-2.4), $ 2.7 (2.7- 2.8); head length S .72 (.64-. 78), ? .67 (.58-. 76); width across eyes ^ 1.67 (1.56-1.72), ? 1.79(1.76-1.82); prono- tal length ^ 1.58 (1.44-1.70), . 1.61 (1.56-1.64); pronotal width S 2.23 (2.04-2.32), S 2.57 (2.48-2.62); scutel- lar length ' 1.01 (.90-1.08), . 1.24 (1.20-1.30); scutellar width ' 1.17 (1.04-1.28), ? 1.37 (1.36-1.40); length antennal segments I S .46 (.43-. 51), $ .50 (.49-. 50), II ' 1.04 (.94-1.10),' $ 1.10(1.09-1.12), III T .85 (.76-. 92), $ .90 (.87-. 92), IV ^ 1.28(1.16-1.36), $ 1.27 (1.20-1.31); length labial seg- ments I S .72 (.64-. 84), ? .78 (.76- .80), II S .61 (.58-. 66), $ .69 (.68-. 70), III S .64 (.60-. 68), ? .73 (.70-. 76), IV S .66 (.64-. 70), i .70 (.66-. 72). Comments. — The black anterior pro- notal margin, constricted pronotum, and the long upright hairs on the heme- lytra separate this from all other species in the genus. The only other Grap- tostethus with dark anterior pronotal markings are large, with the typical generic pronotal shape, and only short semidecumbent hairs on the hemely- tra. As mentioned above, G. cardinalis differs markedly from the usual exter- nal appearance of the genus. The geni- talia are, however, typical of the genus and there can be no doubt that it is correctly placed. The dark markings are often reduced in size and in inten- sity but the anterior pronotal area, subapical corial vitta, and black pos- terolateral pleural maculae were pres- ent in all specimens examined. Graptostethus cardinalis is uncommon in collections but appears to be widely distributed in the northern half of Aus- tralia (Fig. 29). Nothing is known of its food plant or biology. Graptostethus pubescens, sp. nov. Figures 28c; 29; 70e, f, j; 71a-c, m; 72b, g, h. 7>/?^^.— Holotype 2, "1 km. N. of Cahill's Crossing (East Alligator R.), 12.25 S, 132.58 E, 31-X-1972. M Upton." in ANIC. Paratypes: 4{?, 6$, same data as holotype, in ANIC. Also: Northern Territory — 2$, 1$, Adelaide River, 13.15S, 131. 06E, 17.x. 1972, Upton and Bar- rett; 5^, 4?, 1 km N of Cahill's Crossing, 12.25S, 132. 58E, 8.xi. 1972, Upton and Feehan; 2^, 4$, 1 km N of Cahill's Xing, (E. Alligator R.), 12.25S, 132. 58E, 8-9. xi. 1972, The Lygaeinae of Australia 391 • G, cardinalis o G, pubescens ± I Fig. 29. Distribution ol Graploslcthus cardinalis Stal and C. pubescens. sp. no\'. J. C. Cardale; 1$, 1 km S of Cahill's Xing, 12.26S, 132. 58E, 3.xi.l972, Upton and Barrett; 2$, 5 km NNW of Cahill's Crossing, East Alligator River, 12.23S, 132. 57E, 8.vi.l973, R. L. Kitching; 2S , 5 km NNW of Cahill's Crossing, East Alligator River, 12.23S, 132. 57E, 8.vi.l973, R. L. Kitching, at light; I'', 1', Daly River Mission, 15. vi. 1974, J. F. Hutchinson; IS, Darwin, Night- cliff, 20.viii.l956, L. D. Crawford; 1*, Howard Springs, 4.iii.l967, M. S. Upton; 2$, Humpty Doo, 15. vi. 1961, C. S. Li, cat #16; 1$, 14 mi W Humpty Doo, 14. vi. 1964, K. R. Norris; 2S , 2?, 7 km SE of Litch- field Hmstd, 13.iii.l974, J. F. Hutchinson; 1 c^ , Magela Ck., 9 km SSE of Mudginberri H. S., 12.40S, 132. 54E, 6.xi.l972, M. Upton; IS, Magela Ck. , 2 km N of Mudginberri H. S., 12.35S, 132. 52E, 14. xi. 1972, M. Upton; 2-^, 3v, Nourbangie Ck., 8 km N of Mt. Cahill, 12.48S, 132. 42E, 21. V. 1973, Upton and Mclnnes; 1.^, 6 km SW by S of Oenpelli, 12.22S, 133. OlE, 30. v. 1973, Upton and Mclnnes (ANIC); 2?, Daly Riv. Mission, 22.i.l972, J. F. Hutchinson; Iv, 30 mi E. Dar- win, 12.xii.l923, G. F. Hill: U, 392 The University of Kansas Science Bulletin Oenpelli, xii.1918, D. P. Cahill (NMV); Ix, Humpty Doo, 21.vi. 1956, L. O. Crawford (PIBNTA); 2c?, 3$, Daly River, 30. vi. 1972, B. K. Head, on low bushes by river; 25, Darwin; 1 = , Koolpinyah, Octo- ber, G. F. Hill; IV, Roper R. (SAM); Queensland — ix, Archer R., 27.vi.196?, C. N. Smithers (AM); 2r, Weipa, 3-5. ii. 1976, G. B. Monteith, open forest (UQ). Description. — Color: Orange, darkest on head and abdomen, brighter on anterior pronotal lobe and thoracic venter, tawny on posterior pronotal lobe and coriaceous parts of hemely- tron. Differs from general generic col- oration in that: gular area infuscated; posterior transverse pronotal bar pro- duced forward to meet round postcallar macula, forming triangular macula on each side of pronotum; longitudinally ovate humeral macula connected with lateral end of triangular macula; round posterolateral pleural maculae poorly defined, no darker than other dark markings; narrow dark orange poste- rior margins on all abdominal sterna; white mark at basal angle of hemelytral membrane small, obscure; apical mar- gin of corium usually tawny orange. Antenna dark brown except segment I and sometimes II orangey brown. La- bium brown becoming darker apically. Coxae, trochanters, and femora or- angey brown. Tibiae and tarsi dark brown. All dark areas appear more brown than black. Structure: Moderately robust; densely clothed with very short, fine, decumbent hairs, slightly longer and more upright posteriorly on meso- and metapleura, interspersed with a few very short upright hairs dorsally on head and pronotum, interspersed with long upright hairs medially on abdomi- nal venter. Head moderately declivent. Antennal segment I surpassing ante- clypeus by about V3 its length. Labial segment I slightly thicker than II; II slightly thicker than III and IV; I surpassing anterior prosternal margin by about 'A its length; II reaching just beyond middle procoxa; III almost reaching posterior margin of meso- coxa; IV reaching posterior metacoxal margin. Hemelytral membrane greatly surpassing tip of abdomen. Proster- ; num shallowly punctured anteriorly, impunctate posteriorly. Propleuron shallowly punctured anteriorly; indis- tinctly punctate centrally; impunctate posteriorly. Mesopleuron very indis- tinctly punctate. Pygophore as in Fig. 70j; ventral projection of side of dorsal opening scarcely visible from above; lateral projection without anterior point, dorsal channel shallow. Clasper as in Figs. 71a-c; posterolateral lobe short; hook short, rather blunt. Aedea- gus as in generic description; ejacu- latory reservoir as in Figs. 70e, f. Ovipositor as in Figs. 71m, 72g, h; first valvula surpassed distally by first ra- mus. Spermatheca as in Fig. 72b; thin; sclerotized part basal to sperm pump relatively long. Measurements: Length ^ 8.4 (7.8- 9.1) ¥ 8.8 (8.5-9.2); abdominal width S 2.8 (2.7-3.1), $ 3.1 (2.9-3.4); head length 3 .77 (.66-. 85), i .80 (.68-. 89); width across eyes ^ 1.76(1.65-1.83), ? 1.80 (1.75-1.88); pronotal length S 1.62 (1.51-1.81), V 1.69 (1.58-1.78); pronotal width S 2.80 (2.58-3.05), 2 2.94 (2.84-3.11); scutellar length S 1.22 (1.11-1.30), V 1.29 (1.10-1.38); scutellar width S 1.52 (1.35-1.70), ? 1.60 (1.48-1.73); length antennal seg- ments I f .59 (.56-. 61), ? .58 (.56- .61), II S 1.37 (1.30-1.45), $ 1.36 (1.31-1.45), III S 1.00 (.95-1.07), $ .98 (.95-1.04), IV S 1.30 (1.25-1.36), ? 1.26 (1.19-1.31); length labial seg- ments I S 1.04 (1.01-1.06), ? 1.08 The Lygaeinae of Australia 393 (1.03-1.14), II S .97 (.95-1.00), v 1.01 (.96-1.04), III S 1.06 (1.00-1.10), ? 1.11 (1.06-1.18), IV S .81 (.79-. 85), ? .86 (.83-. 90). Measurements holotype $ . Length 8.8; abdominal width 3.1; head length .74; width across eyes 1.78; pronotal length 1.69; pronotal width 2.96; scu- tellar length 1.31; scutellar width 1.63; length antcnnal segments I .59, II 1.45, III 1.04, IV 1.29; length labial segments I 1.13, II 1.04, III 1.13, IV .90. Comments. — Graptostethus pubescens may be separated from other Austral- ian members of the genus by the very short decumbent hairs with no inter- spersed long upright hairs. It resembles Australian specimens of G. servus in the generally faded appearance of the color pattern but is more orange than red- dish in color. Graptostethus varipictus, sp. nov. , has much more distinct dark markings and is more brightly colored in general. Some specimens have greatly re- duced dark markings both in intensity and in area covered. The head, pro- notum, and thoracic venter are espe- cially affected. The dark discal spot on the corium is seldom fused with other dark areas but if fused then fused with the costal macula not the dark margin of the claval suture. Graptostethus pubescens is nearly re- stricted to the northern part of the Northern Territory (Fig. 29) with a few specimens taken on the western coastal part of the Cape York Peninsula. Noth- ing is known of its food plant or biol- ogy- Graptostethus servus (Fabricius 1787) Figures 28a; 30; 70c, d, 1, m; 71d-f, o; 72a, e, f, i, j. Cimex servus Vsbvxcixxs, 1787:300. Lygaeus servus Fabricius, 1794:156. Lygaeus manillensis wStal, 1859:240-241; Slater, 1978:855. Lygaeus ornatus Uhler, 1860:227. Lygaeus (Graptostethus) servus Stal, 1868: 74-75. Graptostethus (Graptostethus) servus Stal, 1874:117. Graptostethus servus Slater, 1964a: 57-59. Type. — Not seen. Type locality given by Fabricius as China. Redescription. — Color: Pale orangey red, dark areas appearing heavily in- fuscated rather than black. Differs from general generic coloration in that: head completely reddish except dark median longitudinal vitta from base to apex of anteclypeus, broader to sur- round ocelli basally; round postcallar macula very light or absent, indicated by hairless area causing it to appear darker than surrounding areas; poste- rior pronotal bar very light or absent; dark margin of claval suture frequently expanded to include short basal corial vitta and discal corial spot, some- times fused with costal macula to form oblique vitta across corium; proster- num completely reddish; major dark areas of thoracic pleura scarcely infus- cated; posterolateral maculae of all tho- racic pleura and anterior propleural macula indicated only by absence of pruinosity; posterior margins of ab- dominal sterna IV to VII, sternum II completely, and genital segments pale orange; apical margin of corium with broad pallid border; basal angle of hemelytral membrane and broad apical margin white. Antenna and labium brown. Legs orangey brown, becoming darker apically. Structure: Body moderately robust; densely clothed with fairly short semi- decumbent hairs interspersed (except on hemelytra) with long nearly upright hairs (these densest on head, callus, lateral pronotal margin, scutellum, and 394 The University of Kansas Science Bulletin medially on abdominal venter); short decumbent hairs lacking on round postcallar macula, thoracic sterna, meso- and metapleura posteriorly; hairs somewhat more upright on hem- elytra. Head moderately declivent; buccula almost reaching anterior pro- sternal margin. Antennal segment I surpassing anteclypeus by about V? its length. Labial segment I slightly thicker than II, II thicker than III and IV; I surpassing anterior prosternal margin by about Vs its length, II reach- ing middle of procoxa, III reaching middle of mesocoxa, IV reaching mid- dle of metacoxa. Hemelytral mem- brane slightly surpassing tip of abdomen. Prosternum moderatelv coarsely punctate anteriorly, impunc- tate posteriorly. Propleuron moder- ately coarsely punctate anteriorly, finely but distinctly punctate centrally, indistinctly punctate or impunctate posteriorly. Mesopleuron sparsely, finely, but distinctly punctate. Pygophore as in Figs. 77a, b; ventral projection of sides of dorsal opening not or scarcely visible from above; lateral projection slightly bifid, dorsal channel distinct. Clasper as in Figs. 71d-f; posterolateral lobe well devel- oped, directed somewhat dorsad; shank very deeply excavated, hook very strongly developed, blunt apically. Aedeagus as in Fig. 72e, f; ejaculatory reservoir as in Figs. 70c, d. Ovipositor as in Figs. 71o, 72i, j; first valvula surpassing first ramus distally. Sper- matheca as in Fig. 72a; sclerotized part basad of sperm pump relatively short. Measurements: Length '" 8.6 (7.9- 9.2), i 9.0 (7.9-9.9); abdominal width i 2.8 (2.7-3.0), ? 3.2 (2.7-3.5); head length ^ .78 (.61-. 88), 5 .80 (.64-. 99); width across eyes ? 1.81 (1.78-1.88), ? 1.90 (1.76-2.00); pronotal length ^ 1.80 (1.63-1.98), X 1.82 (1.58-2.01); pronotal width ' 2.86 (2.74-3.04), ? 3.05 (2.64-3.39); scutellar length $ 1.20 (1.11-1.40), $ L35 (1.15-1.55); scutellar width S 1.49 (L39-1.70), ? 1.69 (1.45-1.90); length antennal seg- ments I $ .58 (.54-. 62), V .59 (.56- .64), II ^ 1.37 (1.28-1.47), 5 1.36 (1.26-1.43), III $ LOO (.93-1.06), ? .99 (.93-1.09), IV ^ 1.39(1.28-1.51), ? L36 (1.26-1.43); length labial seg- ments I $ .98 (.95-1.05), 2 1.01 (.94- 1.06), II ^ .94 (.86-1.03), 4 .96 (.88- 1.01), III i .98 (.88-1.08), 'i L02 (.94-1.08), IV $ .82 (.78-. 88), 2 .85 (.79-. 90). Comments. — The dark median longi- tudinal vitta on the head separates G. servus from all other Australian inem- bers of the genus. The numerous long upright hairs, which give the pronotum a shaggy appearance from above, sepa- rate it from all except G. cardinalis which has the pronotum constricted behind the calli and a quite different color pattern. Australian specimens differ from typical G. servus in their generally paler coloration. Occasional individuals approach the coloring typi- cal of servus in India and the Orient. Specimens from Saibai Island, York Island, and Moa Island in the Torres Strait have the darker typical colora- tion as do occasional specimens from the Northern Territory. These darker individuals may indicate gene flow from populations of the species in New Guinea and Timor, respectively. Graptostethus servus is widely distrib- uted throughout Africa, the Medi- terranean region and the Orient. In Australia it is mainly northern in dis- tribution (Fig. 30). Nothing is known of its biology or food plant preferences in Australia. Slater (1964/^:54) has summarized host plant data for the entire range. Hoffman (1934) recorded the life history oi "" Graptostethus sp. near servus F. ", which may be this species, in China. The Lygaeinae of Australia 395 Fig. 30. Distribution of Grapiosteihus servus (Fabricius) in Australia. Graptostethus varipictus, sp. nov. Figures 28d; 31; 70g-i; 71g-i, n; 72d. Types. — Holotype 5, "Queensland, Brisbane, 23-III-1964. R. Kumar." in ANIC. Paratypes: \i, same data as holotype, in ANIC. Also: Vic- toria—1 S , Woodonga (NMV); New South Wales — 3i, Brunswick Heads, 9.ii.l962, D. E. Haverstein (ANIC); IV, Broadwater, iii.1924, E. G. Baber; 15, Broadwater, iii. 1924, D. S. North; 15, Broadwater, iv.l924, E. G. Baber (AM); North- ern Territory — IS , Jim Jim Water- hole, 12.57S, 137. 34E, 28.viii.1970, J. V. Peters (AM); 15, Darwin, Nightcliff, 20.viii.l956, L. D. Craw- ford; 1 ^' , Nourlangie Ck. , 8 km N of Mt. Cahill, 12.48S, 132. 42E, 21. v. 1973, Upton and Mclnnes; 1^, 6 km SW of S of OenpelH, 12.22S, 133. OlE, 30.V.1973, Upton and Mclnnes (ANIC); 15, (no locality), vii-viii.l2, from Prof. Spencer (NMV); 1', Groote Eylandt, Con- nexion I., N. B. Tindale; 25, Roper R., N. B. Tindale (SAM); 1.', Pine Creek, 22. xi. 1974, E. M. Exley and R. I. Storey, on Eucalyptus leclifica (UQ); Queensland— 25, Brisbane, 23. iii. 1964, R. Kumar; 1^. Bun- daberg, 2.xi.l971, H. Frauca; l^ Bundaberg, 16.x. 1974, H. Frauca; 396 The University of Kansas Science Bulletin IS, 19, Bundaberg, 17.x. 1974, H. Frauca; 1 ', Moreton I., Big Sand Hill, 3.iv.l966, T. G. Campbell (ANIC); 1 ', Bcntinck I., 27. v. 1960, P. Aitken, N. Tiridale, at light; IS, Cairns, A. M. Lea (SAM); 2?, nr. Red Cliff Pt. N of Cairns, 21.vi. 1971, E. C. Dahms(QM);2(?,Mac- kay. Far Beach, 5.vi.l971, E. C. Dahms; H, Pallarenda Pt., nr. Townsville, 26. i. 1965, E. C. Dahms, open forest, nets (QM); 1?, Bamaga, Cape York, 3-6. vi. 1969, G. B. Monteith; 1:^, Brisbane, 14. vii.1932; 1', Brisbane, 17.iii.l970, M. B. Malipatil; 1;, Brisbane, R. Kumar; IS, Brisbane, Long Pocket, 8.viii.l976, A. Slater; IS, Calam- vale, Johnston's Road, 23. xi. 1975, R. L Storey; 1 i , Captain Billy Ck., Cape York Pen., 142. 50E, 11.40S, 9-13. vii. 1975, G. B. Monteith; IS, Dunk L, 25.viii.1927; 3S, 59, Ma- reeba, 28. i. 1976, R. 1. Storey, on rose flowers; 1 • , Mission Beach, 21. V.1976, R. I. Storey; 3S, 2$, Dun- wich, Stradbroke l', 21-22. iii. 1970, M. B. Malipatil (UQ). Description. — Color: Bright reddish orange, apex of scutellum and cor- iaceous parts of hemelytra pales. Dark areas black. Differing from general ge- neric description in that: the entire head dark ventrally except buccula; posterior transverse pronotal bar ex- tended anteriorly, forming subtrian- gular macula by fusion with round postcallar macula, produced slightly anteriorly at humerus; round postcallar macula appearing darker than sur- roundings due to lack of hairs; apical '/:j-'/2 of raised lateral scutellar margin and apex orange; discal spot on corium lused with dark margin of claval su- ture; metapleural markings include an- terior half of acetabular area. White mark at basal angle of hemelytral membrane small, indistinct. Poste- rolateral maculae of thoracic pleura distinct, fused with dark central mark- ings, about equal in size. Antenna, labium, and legs dark brown. Apical margin of corium often with narrow pale border. Structure: Body robust; clothed densely with shortish, rather coarse, semidecumbent hairs, these usually ab- sent on postcallar macula; a few up- right or nearly upright hairs of similar length on pronotum and scutellum, most noticeable at lateral margin of pronotum at callus; meso- and meta- pleural hairs becoming longer and more upright posteriorly; scattered long nearly upright hairs, densest me- dially, on abdominal venter. Head moderately declivent. Antennal seg- ment I surpassing anteclypeus by about V:5 its length. Labial segment I thickest, II slightly thinner, III and IV slightly thinner than II; I surpassing anterior prosternal margin by about 'A its length, II reaching posterior margin of procoxa. III extending slightly beyond middle of mesocoxa, IV reaching or slightly surpassing posterior margin of metacoxa. Hemelytral membrane reaching or slightly surpassing tip of abdomen. Prosternum coarsely punc- tate anteriorly, indistinctly punctate or impunctate posteriorly. Propleuron coarsely punctate anteriorly, finely but distinctly punctate centrally, indis- tinctly punctate or impunctate posteri- orly. Mesopleuron moderately finely, distinctly punctate. Metapleuron with posterior margin slightly convex dor- sally and ventrally. Pygophore as in Fig. 70i; ventral projection of lateral margin of dorsal opening visible from above, forming depressed shelf; lateral projection deeply bifid, anterior point curving posterad to form slight apical lip, dorsal channel distinct. Clasper as in Figs. 71g-i; posterolateral lobe with The Lygaeinae of Australia 397 slight lump dorsally, not directed dor- sail v; hook short, broad. Aedcagus similar to G. servus (Fig. 72e); ejacula- tory reservoir as in Figs. 70g, h. Ovi- positor as in Fig. 7 In; first valvula surpassed by first ramus. Spermatheca as in Fig. 72d; thick; slightly thickened subapically; pigmented part basal to sperm pump of medium length. Measurements: Length ^ 8.8 (8.4- 9.4), $ 10.1 (9.2-10.9); abdominal width ' 3.1 (3.0-3.3), $ 3.6(3.2-3.8); head length ^ .82 (.68-. 96), ? .92 (.83-1.10); width across eyes S 1.86 (1.79-1.93), ^ 2.05(1.88-2.14); prono- tal length ' 1.75 (1.54-1.94), v 2.00 (1.78-2.18); pronotal width ' 3.01 (2.93-3.21), * 3.53(3.15-3.73); scutel- lar length f 1.26(1.21-1.33), . 1.48 (1.38-1.53); scutellar width ' 1.63 (1.55-1.74), * 1.97 (1.76-2.13); length antennal segments I S .65 (.63-. 69), ? .69 (.65-. 72), II S 1.55 (1.40-1.66), $ 1.67 (1.53-1.88), III ' 1.15 (1.08- 1.23), X 1.22 (1.09-1.38), IV S 1.62 (1.58-1.66), X 1.65 (1.56-1.74); length labial segments I ^ 1.14(1.08-1.20), ? 1.28(1.18-1.34), II ' 1.11 (1.04-1.19), $ 1.22 (1.08-1.30), III f 1.26 (1.16- 1.39), X 1.39 (1.25-1.53), IV S .98 (.90-1.04), ? 1.06 (.98-1.13). Measurements holotype i . Length 10.9; abdominal width 3.8; head length .96; width across eyes 2.14; pronotal length 2.15; pronotal width 3.66; scutellar length 1.63; scutellar width 1.95; length antennal segments I .72, II 1.71, III 1.23, IV 1.68; length labial segments I 1.28, II 1.24, III 1.53, IV 1.03. Comments. — Graptostethus varipictus can be distinguished from other Aus- tralian members of the genus by the large size, labium extending to or be- yond the hind coxae, and the short hairs interspersed with slightly longer upright hairs on pronotum and scutel- lum. The species is named for the variability shown in its color pattern. The color pattern of this species is extremely variable. In very dark speci- mens nearly the entire posterior lobe of the pronotum and entire corium are dark. The pale spot at the base of the vertex, often used in the past as a generic character, is sometimes absent. In lighter specimens the head may be almost completely orange, the black areas of the pronotum are present only as infuscations much reduced in area, and the scutellum is black only at the basal angles. A distinctive color form lacks any dark markings on the clavus and corium and has the posterior dark pronotal markings reduced to a narrow longitudinal vitta from the round post- callar macula nearly to the posterior margin and an indistinct spot at the humerus on each side. Graptostethus varipictus is restricted, in Australia, to the northern and eastern coastal regions, the range extending south to about the Queensland-New South Wales border (Fig. 31). I have also seen specimens from Papua New Guinea. Malipatil (1979) provides brief life history and food plant notes. Adults were taken on rose flowers in northern Queensland by Mr. R. I. Storey. Three Graptostethus specimens from New Caledonia (in the collection of the Bishop Museum) matching the original description of G. ocellatus (Montrou- zier) are very similar to the form of G. varipictus with unmarked hemelytra. They differ in their smaller size, the presence of pallid areas rather than orange markings ventrally on the tho- racic pleura, a comparatively slightly longer labium, slight differences in the shape of the claspers (especially the posterolateral lobe which is directed somewhat dorsad) and the less distinct channel of the lateral projection of the dorsal pygophore opening. The differ- 398 The University of Kansas Science Bulletin ences except in size are slight and it is possible that study of a larger series from New Caledonia would show the two species to be one. Some specimens of G. mgnceps Stal, G. diffusus (Walker), and G. mamllensts (Usinger nee Stal) also lack claval and corial markings. The first is smaller, lacks upright hairs on the pronotum and scutellum, and has a relatively much longer labium. The second has longer and more numerous upright hairs on the pronotum and scutellum (approaching G. servus in this respect) and pale yellow rather than orange markings on the thoracic pleura. The last has nearly obsolete dark markings on the thoracic pleura except for the very distinct posterolateral maculae and a similar macula anteriorly on each propleuron; the abdominal venter is nearly uniform brownish orange. Darker specimens of G. nigriceps usually have the corium except for very narrow margins infuscated. Dark specimens of G. diffusus and G. manillensis (Usinger nee Stal) have the discal macula and abbreviated longitudinal vitta of the corium fused with a very broad dark margin of the claval suture, thus ap- pearing to have two large clearly de- fined maculae on each hemelytron. Records of G. manillensis and G. nigri- ceps from Australia almost certainly refer to G. varipietus. Pyrrhobaphus Stal Lygaeus s.g. Pyrrhobaphus Stal, 1868: 73. Pyrrhobaphus ^va\, 1874:99, 116; Slater, 1964fl:187. Type-species Cimex leucurus Fabricius, 1787, monobasic. Stal (1868) erected this genus and the genus Graptostethus as subgenera of the omnibus genus Lygaeus Fabricius. His descriptions of the two subgenera are nearly identical, Pyrrhobaphus being separated from Graptostethus by the pos- session of relatively longer antenna and labium. Breddin (1905:121) pointed out that the discovery of species un- known to Stal, with the general appear- ance of P. leucurus (the type species of Pyrrhobaphus) but possessing characters intermediate between that species and members of Graptostethus, invalidated Stal's division between the two groups. He treated Graptostethus as a junior synonym o{ Pyrrhobaphus on the basis of line priority. No author since Breddin has followed his synonymy. Of the species placed in Pyrrhobaphus by Bred- din those species closely resembling P. leucurus in color or not readily placed in Graptostethus on the basis of their origi- nal description have been treated as members of Pyrrhobaphus. The large number of species, some of them wide- spread, corresponding more closely to Stal's description of Graptostethus than to his description of Pyrrhobaphus, have been described and discussed under the generic name Graptostethus. I have closely examined specimens of P. leucurus, including the male and fe- male genitalia, and can find no consist- ent generic differences between it and members of Graptostethus. As it would create confusion to revert to Breddin's synonymy at this time, I am treating the two genera as separate pending an appeal to the International Commis- sion on Zoological Nomenclature for the conservation of Graptostethus as the generic name for those species now contained in Graptostethus and Pyrrhoba- phus. The generic description given for Graptostethus applies to P. leucurus, the single Australian member of Pyrrhoba- phus, except that in P. leucurus the head is only slightly declivent, the labium extends well beyond the metacoxae, and the color pattern is quite different. The Lygaeinae of Ai'straiia 399 • G. varipictus o P. leucurus 1. Fig. Jil. Distribution of Graptustethus vanputus. sp. nov. and Pyrrhobaphus leucurus (Fabricius) in Australia. Pyrrhobaphus leucurus (Fabricius 1787) Figures 31; 32; 73a-i; 74a-d. Cimex leucurus Fabricius, 1787:299. Lygaeus leucurus Fabricius, 1794:154. Lygaeus (Pyrrhobaphus) leucurus Stal, 1868:73.' Pyrrhobaphus leucurus Stal, 1874:116; ' Slater, 1964a: 188; Slater, 1978:856- 857. Types. — Lectotype ?, "Is. Amstcr- ' dam" in BMNH (Banks Collection). Paralectotype ' , same data as lec- totype, in BMNH (Banks Collec- tion). Redescription. — Color: Dark reddish orange. The following black: base of head; tip of anteclypeus; gular area; pronotum except broad lateral mar- gin behind callus; scutellum; thoracic sterna and pleura except posterolateral angles of pro- and metapleura and narrow dorsal margin of mesopleuron; abdominal sterna except broad lateral margins; genital segments; fused me- dio- and paratergites; antenna except basal 7i of segment I; labivini; and legs. Basal 7i of antennal segment 1 light brown. Eye brown. Ocellus orange. Hemelytral membrane dark brown to black except white basal angle and narrow whitish margin widest apically. 400 The University of Kansas Science Bulletin eucurus Fig. 32. Pyrrhobaphus leucurus (Fabricius), dorsal view. Scale = 1 mm. Structure: elongate oval. Body with moderately dense, moderately long up- right hairs, longest on base of scu- tellum and medially on abdominal venter. The following with short, semi- decumbent hairs: head below and at base dorsally; callus; scutellar base; prosternum; propleuron centrally; meso- and metapleura anteriorly; and all margins of pronotal sterna. Head scarcely declivent, somewhat conical in appearance when viewed from above; buccula low, tapering gently poste- riorly, reaching or nearly reaching prosternum. Antennal segment I sur- passing anteclypeus by slightly less than half its length. Labial segment I surpassing prosternal margin by about half its length, extending about % dis- tance from anterior prosternal margin to proco.xa, II reaching middle of mes- ocoxa. III reaching abdominal sternum IV, IV reaching or almost reaching abdominal sternum V. Pronotum coarsely punctate before and slightly behind callus; callus and lateral margin moderately swollen. Scutellum with lateral margin basally almost same height as T-shaped carina, lower api- cally. Claval commissure about same length as scutellum. Prosternum and propleura coarsely punctate, deepest in front of and just behind center of pleu- ron, posterolateral area of pleuron nearly impunctate. Mesopleuron with a few very small shallow punctures. Metapleuron impunctate. Pygophore as in Figs. 73d, e; ventral projection of sides of dorsal opening slightly visible from above; lateral projection of dorsal opening slightly bifid, dorsal channel shallow. Clasper as in Figs. 73a-c; cau- dolateral lobe subconical; hook blunt but well developed. Aedeagus as in Fig. 73f, g; ejaculatory reservoir as in Figs. 73h, i. Ovipositor as in Figs. 74b- d; first valvula just reaching apex of first ramus. Spermatheca as in Fig. 74a; pigmented section basal of sperm pump long. Measurements: 6S , 2^. Length S 13.5(13.2-14.0), $ 13.7, 14.6; abdom- inal width S 4.2 (4.0-4.4), ? 4.5, 4.8; head length S 1.63(1.36-2.23), $ 1.52, 2.35; width across eyes S 2.08 (1.55- 2.28), . 1.70, 2.16; pronotal length S 2.27 (2.12-2.40), ? 2.16, 2.44; prono- tal width S 3.83 (3.74-4.00), i 4.00, 4.02; scutellar length ^ 1.62 (1.56- 1.73), ; 1.68, 1.75; scutellar width S 1.99 (1.80-2.12), $ 2.28, 2.33; length antennal segments IS 1.01 (.94-1.04), $ 1.08, II ^2.71 (2.64-2.78), i 2.71, III S 1.99 (1.90-2.04), $ 1.95, IV S 2.46 (2.18-2.57); length labial seg- ments I S 2.24 (2.lf-2.32), $ 2.29, 2.48, II ^ 2.36 (2.28-2.48), v 2.66, 2.68, III S 2.82 (2.72-2.90), 'i 3.10, 3.44, IV S 1.65 (1.56-1.74), v 1.74, 1.78. Comments. — The color pattern of P. leucurus is very distinctive. Some in- The Lygaeinae of Australia 401 dividuals of Oncopeltus spp. have a somewhat similar color pattern but arc easily separated from this species by their tumid scutellum and posteriorly directed [)ronotal projections. Pyrrhobaphus leucurus is a widespread species occurring from India to the Pacific region. The only Australian specimens known to me are a series of five males and one female collected near Airlie, Queensland, on October 10th, 1957, and presently in the Uni- \ersity of Queensland collection. I have taken nymphs of this species on the seed capsules oilpomoea sp. in the New Hebrides. Melanerythrus Stal Lygaeus s.g. melanerythrus Stal, 1868:75- 76. Melanerythrus China, 1930:115; Slater, 1964^:118. Brachy lygaeus Gross, 1965:24-26. syn. nov. Type- species Lygaeus mactans Stal, 1867, monobasic. Diagnosis. — Scutellum swollen; no lateral scutellar foveae; posterior pro- notal margin straight or slightly con- cave; ostiolar peritreme tumid but not greatly produced; spermatheca long, irregularly coiled, tubular, pigmented and wider apically. Gross (1965) erected the genus Brachylygaeus to contain Lygaeus mutilatus Erichson. While noting the similarity to Melanerythrus he felt that the reduc- tion in length of the hemelytra of that species, its lack of black medial mark- ings on the corium, and its lack of a white area at the base of the hemelytral membrane were sufficiently different to warrant the erection of a new genus. In the course of this study I have seen specimens of Melanerythrus bigut- tatus lacking black corial markings and specimens of Melanerythrus mactans with these markings reduced. The subma- cropterous specimen of B. mutilatus re- ferred to by Gross (1965) does have a small white mark at the basal angle of the membrane (this almost obscured by grease). Though to date no microp- terous specimens of either M. biguttatus or M. mactans have been discovered, the former shows considerable variation in development of the forewing. Some specimens showing more reduction than the submacropterous specimen of B. mutilatus. Based on these observa- tions and on the similarity of both male and female genitalia of the three species I have no hesitation in treating them as congeneric. As presently understood Melaneryth- rus contains five species, all with an entirely or primarily Australian dis- tribution. It may easily be separated from all other Australian Lygaeinae by being micropterous, or if macropterous by the combination of tumid scutellum and straight posterior pronotal margin. {Oncopeltus, the only other Australian genus with a tumid scutellum, has the posterior margin of the pronotum strik- ingly produced posteriorly to either side of the scutellum.) The shape of the head when viewed from above is also very distinctive due to a slight swelling anterior to the eyes. This gives it a rather elongate, parallel-sided appear- ance and a "pop-eyed" look due to the outline of the eyes not blending into that of the rest of the head. On the basis of the genitalia Melane- rythrus appears more closely related to Spilostethus than to other Australian Ly- gaeine genera. The general appearance of both male and female external geni- talia of the two genera being quite similar. Redescription. — Color: Orange or red and black in varying combinations. Structure: Size medium to large, 402 The University of Kansas Science Bulletin body length 6.2 to 10.7; macropterous, submacropterous, or micropterous; en- tire body granulate, dull; antenna, la- bium, and legs shiny. Head moderately to strongly declivent; wider than long; convex; swollen anterior to eye, ap- pearing parallel-sided; ocelli much closer to eyes than to each other; buc- cula prominent, almost reaching an- terior prosternal margin, convex, tapering gently posteriorly; eye slightly pedunculate, adjacent to anterior pro- notal angle. Antennal segment I sur- passing anteclypeus by slightly less than half its length, II longest, IV slightly shorter than II. Labium not surpassing metacoxa; segment I slightly surpassing anterior prosternal margin. Pronotum rounded laterally, slightly convex; punctations present or not; median carina indicated behind callus; callus swollen, sometimes strongly, reaching lateral margin; cal- lar impression slightly oblique, either unbranched or with short branch near lateral end; shallow depression behind callus reaching or almost reaching lat- eral margin; anterior margin concave, most strongly in micropter; posterior margin straight in macropter and sub- macropter, concave in micropter; lat- eral margin of micropter subparallel, strongly constricted behind callus, of macropter and submacropter subparal- lel anteriorly, angled strongly mesad from base to callus; shallow impression at posterior margin either side of scu- tellum. Scutellum tumid, slightly fo- veate at anterolateral angle; either extremely convex (micropterous spe- cies) or nearly flat; apex acute. Heme- lytra ot micropters with clavus and coriuin undifferentiated; membrane reduced to short flap, not or slightly extending posterior to scutcllar apex; macropters and submacroptcrs with costal corial margin slightly convex, veins slightly raised. Prosternum im- punctate; sometimes coarsely rugose; anterior margin slightly raised. Meso- and metasterna impuncate, coxal mar- gins raised. Propleuron impunctate centrally, punctate or coarsely rugose anteriorly and posteriorly. Mesopleu- ron impunctate except sparsely, shal- lowly near posterior margin. Meta- pleuron impunctate; ostiolar peri- treme well developed; posterior margin nearly straight, directed posteroven- trally, subparallel to posterior mes- opleural margin; convex bump near anterolateral angle; posterolateral an- gle rounded. Legs unarmed. Abdom- inal sternum II almost completely visible laterally; III to VII with ante- rolateral scent gland openings; VII of male concave posteriorly; VII of female cleft almost to base, sloping gently from lateral margin to cleft. Abdom- inal dorsum with medio- and par- atergites fused; inner laterotergites indistinctly separated from medio- and paratergites II to VI; all tergites fused on VII. Pygophore with median pro- jection well developed, ridge on each side separated from projection by nar- row channel; dorsal opening with de- pressed shelf laterally opposite terga. Clasper with large, thuinb-like cau- dolateral lobe; blade subtriangular in cross section basally, flattened apically; shank excavated mesally. Aedeagus with raised phallothecal process curved and with undercut projection; phal- lothecal surface with dorsal median ca- rina, depressed between carina and phallothecal process; conjunctiva ro- bust, with subapical dorsal lobe on each side; low lobe over ejaculatory reservoir; vesica robust, lobeless; ring sclerite almost complete, lightly scle- rotized; ejaculatory reservoir well developed, lacking basal projection; gonoporal process visibly covered by membrane almost to apex, secondary gonopore with short, acute process. The Lygaeinae of Australia 403 Ovipositor with connecting membrane of first valve cleft and with a well defined median sclerite; first valvula not reaching apex of connecting mem- brane; connecting membrane of second valve deeply cleft medially, moderately pigmented; second valvula rounded apically. Spermatheca moderately long, irregularly coiled, swollen api- cally, about apical 7s pigmented; sperm pump little differentiated from basal tube, basal to pigmented area. Key to the Species of Melanerythrus Stal 1. Calli red or orange; microp- terous; pronotum coarsely ru- gose except calli, nearly as wide across calli as across base 2 1'. Calli black; micropterous or not; pronotum punctate or nearly smooth, never coarsely rugose, much wider across base than across calli 3 2. Boldly marked with orange and black uptoni, sp. nov. 2'. Nearly completely dull red . . Ignitus, sp. nov. 3. Pronotum black and orange; macropterous mactans Stal 3'. Pronotum black; macropterous or not 4 4. Pronotum coarsely punctate; never micropterous biguttatus (Fabricius) 4'. Pronotum at most indistinctly punctate; usually micropterous. mutilatus (Erichson) Melanerythrus biguttatus (Fabricius 1775) Figures 33; 35; 75e, f; 76i-l; 77c, d, m; 78c, i; 79c, d. Cimex biguttatus Fabricius, 1775:720. Cimex australis Gmelin, 1790:2173. Lygaeus biguttatus Fabricius, 1794:158. Lygaeus divisus Walker, 1872:62-63. Melanerythrus biguttatus Distant, 1901a: 536; Slater, 1964a: 1 18-1 19. biguttatus Fig. 33. Melanerythrus biguttatus, (Fabricius), dorsal vifw. Scale = 1 mm. Type.— Ty^a ^, no label, in BMNH (Banks collection). The type locality given by Fabricius as "Nova Hollan- dia". Redescription. — Color: Orange. The following black: head laterally anterior of antenniferous tubercle; head ven- trally; head basally posterior of eye and ocellus; apical half anteclypeus; pro- notum; scutellum; rounded discal co- rial macula; thoracic sterna; thoracic pleura; anterolateral angles of abdomi- nal sterna III to VII; sterna VI and VTI medially; genital segments; and ter- gum VII medially. Fhe following dark brown to black; antenna; labium; and legs. Eye brown, Ocellus red. Heme- lytral membrane brown except white basal angle, discal spot, and apical margin. Structure: Head sharply declixent, hairs dorsally and laterally short, decli- 404 The University of Kansas Science Bulletin vent, densest laterally, hairs ventrally longer, upright, sparse. Labial segment I slightly surpassing anterior prosternal margin, II reaching procoxa. III reach- ing mesocoxa, IV slightly surpass- ing metasternum. Pronotum coarsely punctate except callus; callus slightly oblique, callar impression subsinuate, unbranched; median carina obsolete anterior to callus and at posterior mar- gin, well defined medially; hairs mod- erately long, semidecumbent, sparse, densest on callus. Scutellum slighdy convex, median carina well defined, shallow transverse striations laterally; hairs moderately long, semiupright, moderately dense. Clavus parallel sided; hairs semiupright, moderately long and dense. Corium with slightly convex costal margin; apex usually reaching tergum VI, occasionally reaching only tergum V; hairs semi- upright, moderately long and dense, absent on and immediately surround- ing discal macula. Hemelytral mem- brane of macropter reaching or nearly reaching tip of" abdomen; of subma- cropter reaching tergum VII or oc- casionally VI; basal cell absent in extreme submacropter. Prosternum with very shallow punctures; hairs very thin, moderately long, semiupright, sparse. Meso- and metasterna impunc- tate; hairs very thin, moderately long and dense, semiupright. Propleuron sparsely, coarsely punctate anteriorly and posteriorly; hairs moderately long and dense, semiupright, very thin. Mcsopleuron with a few coarse punc- tures posteriorly; hairs thin, moder- ately long and dense, semidecumbent anteriorly, semiupright posteriorly. Metapleuron with low anterolateral bump; hairs as on mesopleuron. Ab- dominal sterna with long semidecum- bent hairs, dense medially, gradually shorter and sparser laterally. Pygo- phore as in Figs. 75e, f. Clasper as in Figs. 76J-1; blade less flattened distally than other Me lanerythr us species. Aedea- gus generally as in M. mactans (Fig. 77i); phallotheca as in Figs. 76i, 77m, dorsum less sculptured than in other members of the genus; ejaculatory res- ervoir as in Figs. 77c, d. Ovipositor as in Figs. 78c, 79c, d. Spermatheca as in Fig. 78i. Measurements: Length S 6.9 (6.4- 7.4), $ 7.6 (6.2-8.2); abdominal width S 2.1 (1.9-2.3), 2 2.4 (2.0-2.6); head length ' .91 (.73-1.04), i 1.11 (.98- 1.25); width across eyes S 1.56 (1.48- 1.61), ? 1.66 (1.54-1.80); pronotal length S 1.40 (1.29-1.50), v 1.50 (1.34-1.63); pronotal width .^ 2.04 (1.93-2.19), $ 2.32 (1.98-2.60); scutel- lar length ' .91 (.78-1.03), $ .98 (.79- 1.08); scutellar width $ 1.10 (.98- 1.23), X 1.20 (.95-1.38); length anten- nal segments I _' .57 (.53-. 61), * .58 (.54-. 61), II ' 1.33 (1.21-1.46), ? 1.34 (1.24-1.46), III S 1.10 (1.02-1.21), ? 1.11 (1.03-1.20), IV ' 1.26 (1.14- 1.40), $ 1.22 (1.10-1.38); length labial segments I $ .74 (.66-. 83), ? .81 (.75- .86), II ' .80 (.73-. 88), 5 .89 (.84-. 95), III S .92 (.85-. 99), * L02 (.95-1.05), IV $ .92 (.89-. 99), 2 .97 (.93-1.05). Comments. — The combination of a completely black, punctate pronotum, and orange corium with a small black discal macula and a large white discal macula on the hemelytral membrane distinguish M. higuttatus from all other Australian Lygaeinae. In some speci- mens from Western Australia and Ti- mor the black corial macula is lacking or much reduced and the hemelytral membrane appears mottled with white due to irregular expansion of the white areas. In Australia M. biguttatus is appar- ently restricted to tropical and sub- tropical coastal regions (Fig. 35). I have also examined specimens from Timor and Papua New Guinea. Adults The Lygaeinae of Australia 405 have been collected on mangroves, maize seedlings, "Graminae", and Oe- nothera drummondii. I believe that only the latter two should be considered host records. Melanerythrus ignitus, sp. nov. Figures 35; 75a, b; 76e-h; 77e, f, i, 1; 78e, h; 79a, b. Types.— HoXoiyT^e $, "Herald Bight, Shark Bay, W.A., 5-VII-1943, C. P. Whitley" in AM. Paratypes: 9c?, 3$, "Shark Bay, W.A., 28- VII- 1964, L. E. Koch"; 1?, "Waroona Station, W.A., 28-VII-1974, L E. Koch"; li, "Marillana Station, 8- VIII-1958, A. M. Douglas" all in WAM. Description. — Color: Dull red. The following sometimes black: tergal scent gland scars; anterolateral angles of ab- nitus Fig. 34. Melanerythrus ignitus, sp. nov.. dorsal view. Scale = 1 mm. dominal sterna and outer laterotergites III to VII; abdominal terga VI and VII medially; abdominal sternum VII; and genital segments. Antenna and tarsi often apically infuscated. Labium red to reddish brown basally darkening to nearly black apically. Eye brown. Ocel- lus red. Hemelytral membrane brown at base, whitish apically. One specimen with black spot at anterior margin of propleuron. Structure: Micropterous. Head strongly declivent; hairs short, sparse on vertex, long upright hairs inter- spersed dorsally, hairs semidecumbent ventrally. Labial segment I slightly sur- passing anterior pronotal margin, II reaching procoxa. III reaching meso- coxa, IV reaching posterior metaster- nal margin. Pronotum subquadrate; coarsely rugose before and behind cal- lus; median carina well developed pos- teriorly, obsolete anteriorly, callus transverse, strongly swollen, occupies about half pronotal length laterally; callar impression oblique, sinuate, un- branched; sparse, interspersed short decumbent and long upright hairs. Scutellum strongly convex; lateral fo- vea well developed giving carinate appearance apically; lateral margin convex apically; apex bluntly angulate; interspersed moderately long, dense decumbent and long, sparse upright hairs. Clavus and corium fused, sub- triangular; posterior margin rounded apically; apical margin straight; hairs as on scutellum, very sparse on disc. Hemelytral membrane reduced, sub- triangular, rounded apically, reaching or slightly surpassing scutellar apex, not or barely reaching abdominal ter- gum II. Prosternum very coarsely punctate. Propleuron very coarsely punctate except center, nearly as long as combined meso- and metapleura; hairs moderately long, semidecum- bent, sparse. Meso- and metasterna 406 The University of Kansas Science Bulletin with dense, moderately long, upright hairs. Mesopleuron coarsely punctate posteriorly; hairs moderately long and dense, semiupright. Metapleuron with well developed anterolateral bump; hairs as on mesopleuron but denser. Abdominal sterna with dense, decum- bent, moderately long hairs sparsely interspersed with long semiupright hairs, both types densest medially and posteriorly. Abdominal terga with sparsely but evenly distributed long semiupright hairs interspersed medi- ally and on outer laterotergites with dense shorter decumbent hairs. Pygo- phore as in Figs. 75a, b. Clasper as in Figs. 76f-h. Aedeagus similar to M. mactans (Fig. 77i); ejaculatory reservoir as in Figs. 77e, f; phallothcca as in Figs. 76e, 771. Ovipositor as in Figs. 78e, 79a, b. Spermatheca as in Fig. 78h. Measurements: 9^, 6?. Length $ 6.2 (5.9-6.5). 2 7.0 (6.6-7.4); abdomi- nal width S 2.3 (2.0-2.5), $ 2.8 (2.5- 3.1); head length ^ 1.03 (.90-1.13), $ 1.04 (.85-1.19); width across eyes $ 1.46 (1.43-1.51), . 1.56 (1.50-1.60); pronotal length ' 1.07 (1.04-1.15), $ 1.19 (1.09-1.29); pronotal width $ 1.52 (1.50-1.56), 2 1.76 (1.63-1.85); scutellar length ^ .75 (.70-. 83), 5 .88 (.74-1.08); .scutellar width ^ .96 (.85- 1.03), $ 1.17 (1.05-1.20); length an- tennal segments I $ .55 (.53-. 60), $ .59 (.58-. 60), II S 1.22 (1.18-1.27), ? 1.39(1.33-1.44), III ' .98 (.93-1.03), $ 1.08 (1.03-1.14), IV ' 1.10 (1.07- 1.15), ^ 1.13 (1.08-1.15); length labial segments I ^ .85 (.83-. 89), 9 .92 (.89- .98), II ' .84 (.81-. 85), . .90 (.88-. 93), III $ .74 (.69-. 78), V .79 (.78-. 81), IV S .69 (.65-. 75), 9.74 (.70-. 78). Measurements holotypc . Length 6.8; abdominal width 2.6; head length 1.03; width across eyes 1.58; pronotal length 1.14; pronotal width 1.63; scu- tellar length .75; scutellar width 1.05; length antennal segments I .58, II 1.41, III 1.09, IV 1.15; length labial segments I .93, II-IV obscured by card mount. Comments. — Melanerythrus ignitus is named for both its color and the cli- mate of its type locality. The species appears to be restricted to western Aus- tralia (Fig. 35). The combination of microptery and virtual lack of black markings separates this species from all other Australian Lygaeinae. It is somewhat similar in appearance to members of the African genus Apterola Mulsant and Rey but the head and pronotum appear rather longer than in that genus and the calli are much more swollen. I have not seen the genitalia of Apterola. Melanerythrus ignitus and M. uptom show very pronounced structural dif- ferences from the rest of the genus, even more so than do micropterous specimens of M. mutilatus. However these differences are consistent with reduction in wing length. The male and female genitalia are typical of the genus. Melanerythrus mactans (Stkl 1867) Figures 35; 75g, h; 76m-p; 77g-j; 78b; 79g, h. Lygaeus mactans Stal, 1867:162. Lygaeus (Melanerythrus) mactans Stal, ' 1868:76. Lygaeus ruficeps Walker, 1872:63. Stalagmostethus mactans Kirkaldy, 1908: 355. Stalagmostethus melanesicus Kirkaldy, 1909:31. Spilostethus mactans Bergroth, 1916:33. Melanerythrus mactans China, 1930:115; Slater, 1964a: 119; Slater, 1978:857. Types. — Lectotype 9, "Austral occid." "Thorey. " in Stockholm. Paralec- totypes: 19, "Austral, boreal."; 19, "Ins. Fidschi." in Stockholin. Thp: Lycjaeinae of Australia 407 Redescription. — Color: Bright orange. The following black: head around eye and ocellus; sometimes tip of anteclyp- eus; buccula; pronotum except area of humeral angle; scutellum; clavus dis- tally; transverse medial corial fascia widened costally to include subapical third of costal margin; thoracic pleura and sterna; anterolateral angles of ab- dominal sterna III to VI; sternum VII; genital segments; tergum VII; and sometimes sterna IV to VI medially. The following dark brown to black: antennal segments I distally, II, III, usually IV; labial segments; and legs. Eye brown. Ocellus red. Hemelytral membrane brown except white basal angle, discal spot, and margins. Structure: Head moderately decli- vent; hairs short, sparse, decumbent dorsally, denser laterally and ventrally. Labial segment I surpassing anterior prosternal margin by about 'A its length, II reaching posterior margin of procoxa. III reaching posterior margin of mesocoxa, IV reaching posterior margin of metasternum. Pronotum shallowly punctate anterior to and im- mediately behind callus; median carina obsolete except immediately behind callus; callus almost transverse, im- pression often with short branch near lateral end; hairs short, decumbent, sparse posteriorly, denser anteriorly. Scutellum slightly convex; without me- dian carina; hairs short, decumbent, sparse except anterolaterally. Clavus parallel-sided; hairs short, dense, de- cumbent. Corium with moderately convex costal margin; hairs short, de- cumbent, sparse except narrow costal border slightly widened at black fascia. Hemelytral membrane reaching or sur- passing apex of abdomen. Prosternum impunctate; with shallow, poorly de- fined, transverse striations; hairs short, sparse, decumbent. Meso- and meta- sterna with dense, moderately long, upright hairs. Propleuron shallowly striate; a few coarse, elongate punc- tures; hairs short, decumbent, denser posteriorly. Mesopleuron with hairs short, decumbent, denser posteriorly. Metapleuron with poorly developed anterolateral bump; hairs short, de- cumbent, denser anteriorly. Abdomi- nal venter with dense, thicker, semide- cumbent hairs interspersed with longer more upright hairs. Pygophore as in Figs. 75g, h. Clasper as in Figs. 76n-p. Aedeagus as in Fig. 77i; phallotheca as in Figs. 76m, 77j; ejaculatory reservoir as in Figs. 77g, h. Ovipositor as in Figs. 78b, 79g, h. Spermatheca as in Fig. 78g. Measurements: Length $ ^.\ (7.3- 9.6), 5 9.6 (8.3-10.7); abdominal width f 2.9(2.8-3.1), i 3.1 (2.7-3.4); head length ' .97 (.85-1.20), ^ 1.00 (.80-1.20); width across eyes ^ 1.89 (1.73-2.03), + 1.92(1.78-2.08); prono- tal length $ 1.73 (1.40-1.95), I 1.82 (1.63-2.06); pronotal width ^ 2.57 (2.04-2.78), V 1.46 (1.33-1.75); length antennal segments I •$ .65 (.54-. 69), $ .64 (.58-. 68), II i 1.58 (1.41-1.68), $ 1.53 (1.45-1.61), III S 1.25 (1.00- 1.36), 2 1.17 (1.07-1.28), IV :' 1.39 (1.18-1.51), . 1.35 (1.23-1.53); length labial segments I S 1.09 (.95-1.16), 2 1.14(1.04-1.28), II i 1.03 (.98-1.10), $ 1.08(1.05-1.13), III $ .83 (.75-. 90), ? .86 (.83-. 90), IV S .76 (.66-. 83), i .77 (.73-. 81). Comments. — Melanerythrus mactans is easily recognized by its black pronotum with broad orange lateral margins of the posterior lobe, black corial fascia, and large white macula of the hemely- tral membrane. Some species of On- copeltus approach this appearance but their dark pronotal areas are at most piceous and they have pronotal projec- tions extending posteriorly on either side of the scutellum. 408 The University of Kansas Science Bulletin On clean, degreased, specimens the anterior lobe of the pronotum, the an- terolateral scutellar angles, and most of the thoracic venter are pruinose. Two females examined were slightly submacropterous, the hemelytral membrane reaching only the posterior margin of abdominal tergum VII. An apparently fully mature female from the Northern Territory, in the collec- tion of the University of Queensland, has yellow, rather than orange, and brown, rather than black, markings. Melanerythrus mactans has the widest distribution of any member of the ge- nus, occurring almost throughout Aus- tralia. It has been reported from Fiji and I have seen specimens from Timor. The Australian distribution (Fig. 35) indicates that the species prefers arid conditions. Adults and nymphs were collected in Western Australia under Solarium sp. and under Solanum lucani by Mr. R. I. Storey. Additional records which probably do not constitute true food plants are strawberries, pecan nut trees, and "in soil around cherry trees". Melanerythrus mutilatus (Enchson 1842) Figures 35; 75c, d; 76a-d; 77a, b, k; 78a, f; 79e, f. ® M. mutilatus * M. Ignitus ® M.uptoni L ± ± Fig. 35. Distribution of A/Wa«fry/Aruj marfanj (Stal), A/, biguttatus {fabncms). M. mutilatus (E.v'ich^.on) . M. ignttus. sp. nov. . and M. uptoni, sp. nov. in Australia. The Ly(;akinae of Australia 409 Lygaeus mutilatus Erichson, 1842:279; ' Slater, 1964a: 116. Brachylygaeus mutilatus Gross, 1965:24- 26'. " '^ Type. — Type lost (Woodward, 1962). Erichson lists Tasmania as the type locality. Redescription. — Color: Orangey red. The following black: base of head, widest laterally to include base of eye and ocellus; head laterally and ventrallv anterior to antenniferous tu- bercle; anteclypeus except base; pro- notum; scutellum; posterior and apical hemelytral margins; thoracic sterna and pleura; broad longitudinal vitta laterally on abdominal sterna IV to VI; abdominal sternum VII; genital seg- ments; abdominal terga I, VI, and VII; abdominal terga II to V medially; antenna; labium; and legs. Eye brown. Ocellus red. Hemelytral membrane of micropter brown basally, white api- cally; of submacropter brown with white mark at basal angle, subdiscal macula, and narrow apical border. Structure: Head moderately decli- vent; hairs moderately long, sparse, upright dorsally, semidecumbent ven- trally. Labial segment I reaching an- terior prosternal margin, II reaching procoxa, III reaching mesocoxa, IV reaching middle of metasternum. Pro- notum usually impunctate, occasion- ally with faint punctures anterior to callus; median carina well defined on posterior lobe, faintly indicated on an- terior lobe; hairs short, dense, decum- bent, interspersed with some longer, upright hairs. Scutellum quite convex; lateral fovea well developed distally giving appearance of broad median carina; faint transverse striations later- ally; hairs as on pronotum. Clavus and corium of micropter fused, truncate apically, reaching posterior margin of abdominal tergum I, veins absent, hairs as on pronotum, denser on mar- gins; of submacropter distinct, clavus slightly narrowed distally, costal corial margin slightly convex, veins slightly raised, hairs of clavus and of corial margins short, decumbent, moderately dense, on disc and interspersed on clavus and margins of corium moder- ately long, upright hairs. Hemelytral membrane of micropter a small convex flap not surpassing apex of scutellum; of submacropter reaching posterior margin of abdominal tergum VII, ve- nation reduced on one wing forming five cells on the other. Prosternum very shallowly punctate; anterior margin raised; hairs short, sparse, semiup- right. Meso- and metasterna impunc- tate; hairs moderately long, dense, semiupright. Propleuron with shallow punctures anteriorly and posteriorly, impunctate centrally; hairs short, semi- decumbent, sparsest anteriorly and posteriorly. Mesopleuron with scat- tered shallow punctures posteriorly, otherwise impunctate; hairs short, semidecumbent, moderately dense. Metapleuron with anterolateral bump moderately developed; hairs semide- cumbent, dense anteriorly, sparse and slightly longer posteriorly. Abdominal sterna with interspersed short, decum- bent and sparse longer upright hairs, denser medially and posteriorly on sternum VII. Abdominal terga with short sparse semiupright hairs, densest medially. Pygophore as in Figs. 75c, d. Clasper as in Figs. 76b-d. Aedeagus similar to M. maclans (Fig. 77i); phal- lotheca as in Figs. 76a, 77k; ejaculatory reservoir as in Figs. 77a, b. Ovipositor as in Figs. 78a, 79e, f. Spermatheca as in Fig. 78f. Measurements: 8 '^ , 10?. Length ' 8.3 (7.7-8.5), . 8.5 (8.0-9.3); abdomi- nal width ^ 2.7 (2.5-2.8), ? 3.0 (2.5- 3.4); head length ' 1.08 (.88-1.25), $ 1.06 (.95-1.20); width across eyes c? 410 The University of Kansas Science Bulletin 1.75 (1.68-1.80), 2 1.78 (1.71-1.88); pronotal length i 1.60 (1.45-1.68), ? 1.58 (1.40-1.93); pronotal width S 2.14 (1.93-2.28), 2 2.25 (1.85-2.73); scutellar length S 1.14(1.06-1.23), 2 1.22 (.95-1.48); scutellar width S 1.25 (1.13-1.34), 2 1.34(1.18-1.50); length antennal segments I S .70 (.64-. 74), $ .69 (.64-. 74), II ^ 1.55 (1.41-1.68), $ 1.54 (1.39-1.68), III S 1.15 (1.06- 1.23), 2 1.16 (1.08-1.23), IV ' 1.36 (1.25-1.50), 2 1.34 (1.28-1.44); length labial segments I S 1.09(1.00-1.18), 2 1.14 (1.05-1.20), II S 1.04 (.98-1.13), $ 1.09(1.03-1.13), III ' .86 (.83-. 90), $ .93 (.85-. 98), IV S .78 (.75-. 80), 2 .79 (.78-. 83). Comments. — The completely black pronotum combined with microptery, or in the case of submacropters with a black apical corial margin, separate Melanerythnis mutilatus from all other Australian Lygaeinae. Gross (1965) provides figures of the dorsum. Originally described from Tasma- nia, the locality data of known speci- mens indicate that this is an alpine or subalpine species. It is apparently re- stricted to southeastern Australia (Fig. 35). During the course of this study several males and females were taken on or under Craspedia macrocephala among snow gums near the summit of Mt. Gininni, ACT. Melancrythrus uptoni, sp. nov. Figures 35; 36; 78d, j. Type. — Holotype i, "Standley Chasm, 43 km. W. by S. of Alice Springs, N.T., ll-X-1972, M. Upton." in ANIC. Description. — Color: Orange. The following dark brown to black: head around eye and ocellus; tip of ante- clypeus; buccula; anterior and pos- terior pronotal margins; scutellum; abdominal terga I and VII completely, uptoni Fig. 36. Melanerythrus uptoni. sp. nov., dorsal view. Scale = 1 mm. II, V, and VI medially. III antero- medially, and IV antero- and poste- romedially; outer laterotergites II to VI anteriorly and narrowly at poste- rior margin; thoracic sterna; thoracic pleura except propleuron lateromedi- ally; abdominal sternum II anteriorly, VII completely, III to VI at ante- rolateral angle, and broad diffuse lon- gitudinal vitta medially from posterior part of II to VI, narrowed anteriorly; and legs. Eye, antenna, and labium brown. Ocellus orange. Hemelytral membrane brown with faint whitish border. Structure: Very similar to M. ignitus, sp. nov., differing in: pronotum ante- rior to callus less differentiated from callus; scutellum extending posterad of abdominal tergum I, less convex later- ally with apex more acutely angulate; hemelytral membrane less rounded. The Lygaeinae of Australia 411 not approaching scutellar apex or pos- terior margin of tergum I; punctures of pronotum, thoracic pleura, and tho- racic sterna shallower and slightly bet- ter defined; apex of second valvula (Fig. 78d) shorter, less flattened and with less concave ventral margin; sper- matheca as in Fig. 78j; decumbent hairs shorter and much less dense ex- cept on abdomen; long hairs on pro- notum, scutellum, and hemelytra relatively longer. Measurements: Holotype $. Length 6.4; abdominal width 2.4; head length .83; width across eyes 1.50; pronotal length 1.10; pronotal width 1.63; scu- tellar length .85; scutellar width 1.08; length antennal segments I .51, II 1.21, III .96, IV 1.10; length labial segments I .88, II .83, III .78, IV .71. Comments. — Melanerythrus uptom is named for the collector of the only known specimen, M. E. Upton. The micropterous form and orange pro- notum with blackish anterior and pos- terior margins distinguishes this species from all other Australian Lygaeinae. As mentioned above it is very similar structurally to M. Ignitus, sp. nov. It is possible that study of a series of speci- mens will show that the structural fea- tures are variable enough and/or that the male genitalia are similar enough to justify synonymizing the two species. Melanotelus Reuter Melanotelus Reuter, 1885:200, 202; Scudder, 1968:156; Slater and Sperry, 1973:156-157. Lygaeosoma Bergroth, 1918:57-58 (part); Slater, 1964a:64-65 (part), 1964^:49 (part). Type-species Melanotelus argillaceus Reu- ter, 1885 (fixed by Distant, 1904). Diagnosis. — Base of head black; pro- notum tumid, calli scarcely protrud- ing; posterolateral metapleural angle slightly produced posteriorly; para- and mediotergites III to VI separate; tergal apodemes on paratergites III and IV, on mediotergites V and VI; and very long, apically curved, upright hairs over whole dorsum. The genus Melanotelus was erected by Reuter (1885) for a group of medium sized lygaeines previously included in the omnibus genus Lygaeus Fabricius. Bergroth (1918) synonymized it with Lygaeosoma Spinola on the ground that the character separating the two genera (angulate vs. truncate posterior meta- pleural margins) was too variable to warrant a generic division. Scudder (1968) removed Melanotelus from syn- onymy, stating only that he found it "... a distinct genus." Slater and Sperry (1973) concurred, adding "... the species we have examined have a habitus very distinct from 'typical' Ly- gaeosoma species." Examination of the male and female genitalia and of the abdominal dorsum has provided excel- lent evidence of their separate generic status. In Lygaeosoma there is a lateral sclerite on either side of the conjunc- tiva, the gonoporal process is short and thick, the spermatheca is short with a round pigmented apical lobe, and the tergal apodemes of abdominal seg- ments III to VI are all on the para- tergites. In Aielanotelus there are no conjunctival sclerites, the gonoporal process is very long and thin, the sper- matheca is elongate and slightly swol- len apically, and the tergal apodemes of abdominal segments V and VI are on the mediotergites. Little is known of the biology of members of the genus. Slater and Sperry (1973) surmise that they feed on a variety of mature seeds and state that M. villosulus (Stal) was easily reared on sunflower seeds in the laboratory. Scudder's (1968) report of two female M. bipunctatus (Dallas) taken during 412 The University of Kansas Science Bulletin aerial trapping over the ocean about three kilometers off the coast of India indicates that species may have good powers of dispersal. The only biolog- ical information available for Austral- ian species is a note on a specimen collected at Kambah, ACT, by G. F. Hill stating that it was found in an Iridomyrmex (Formicidae) nest. Redescription. — Color: Head dorsally black or dark brown; black macula at humeral angle of pronotum; black macula at posterolateral angle of meta- pleuron (sometimes also of pro- and mesopleura); scent gland orifice pallid, sometimes infuscated basally; antenna, labium, and legs dark brown to black; hemelytral membrane black except white basal and apical margins; ab- dominal segment VII and genital seg- ments dark brown to black. Structure: Subovoid, widest across middle of abdomen; head, thorax, and hemelytra granulate, dull; abdominal venter, antenna, labium, and legs shiny; pronotum, scutellum, clavus, corium, and dorsum of head sparsely covered with long, upright hairs. Head declivent, convex; ocelli much closer to eyes than to each other; buccula promi- nent; lateral and ventral hairs short, decumbent. Labium not extending • M.rubiginosus o M geriae Fig. 37. Distribution ot Melanotelus mbiginosus (Walker) and M. genae, sp. nov. I'nK Lygaeinae of Australia 413 caudad of metacoxa (except in M. ar- gillaceus Reuter where from the descrip- tion it reaches abdominal segment III); segment I thickest, II shghtly thinner, III and IV thinner than II and about equally thick. Pronotum moderately tumid, subquadrate, widest basally, coarsely punctate; callus slightly raised; callar impression slightly ob- lique, sinuate, unbranched; shallow in- terrupted impression at posterior margin near humeral angle; median carina obsolete, barely indicated im- mediately behind callus; anterior mar- gin concave; lateral margin straight or slightly sinuate; posterior margin sub- sinuate, slightly convex. Scutellum wider than long; median carina T- shaped; lateral fovea in angle of T very deep. Corium with slightly convex cos- tal margin; veins slightly raised. Prosternum with anterior margin ex- tending anterad of posterior margin of eye; anterior margin raised; punctate; hairs sparse, short, and upright. Meso- and metasterna impunctate; hairs dense, short, and upright. Mesopleu- ron raised anteriorly; coarsely punc- tate; hairs longer and denser than on propleuron, densest posteriorly. Meta- pleuron raised anteriorly; coarsely punctate at least in part; hairs upright, longer than on mesopleuron, densest anteriorly; ostiolar peritreme well de- veloped; posterior margin concave, di- rected anteroventrally; posterolateral angle produced, somewhat rounded. Abdominal sternum II almost covered by metapleuron; female sternum VI slightly shorter medially than laterally; posterior margin of male sternum VII truncate; female sternum VII cleft al- most to base; sterna without dor- solateral gland openings; all segments hirsute. Abdominal dorsum with para- tergites III to VI separate, apparently fused with laterotergites on female VII, with mediotergite on male VII; poste- rior margin of female VII truncate, of male VII convex. Pygophore subcircu- lar in cross section; median hypandrial projection well developed; projection of lateral wall of dorsal opening just ante- rior to insertion of clasper. Clasper with well developed caudolateral lobe; blade broad, flattened; shank deeply excavated, well developed overhanging projection. Aedeagus with phallotheca lightly pigmented; phallothecal process curved, projecting slightly laterad, sculptured apically; conjunctiva short, robust, and subapical dorsolateral lobe on each side; ejaculatory reservoir complete, without basal projection, covered by low membranous lobe; ve- sica basally robust, twisted, lobed; ring sclerite complete, heavily pigmented, short, narrowed distally; vesica api- cally short, coiled, striate and lightly pigmented towards apex; gonoporal process long, coiled, visibly covered by membrane only at extreme base; sec- ondary gonopore simple, without ap- ical process. Ovipositor with first valvula not reaching apex of connect- ing membrane; connecting membrane of first valve deeply cleft medially, a median sclerite surrounding cleft. Spermatheca elongate, tubular, loosely coiled, pigmented distally, slightly swollen subapically; sperm pump at base of pigmented part, elongate, swol- len. Key to the Australian Species of Melanotelus 1. Clavus and corium with inter- spersed short, fine, decumbent and long, stout, upright hairs; base color brown to brownish red rubiginosus (Walker) r. Clavus and corium with only long, stout, upright hairs; base color brilliant scarlet geriae, sp. nov. 414 The University of Kansas Science Bulletin Fig. 38. Melanolelus geriae, sp. nov. , dorsal view. Scale = 1 inm. Melanotelus geriae, sp. nov. Figures 37; 38; 8()d-g, j, k, o; 81c-e. Types. — Holotypc '" , "Northern Terri- tory, 39 km SW of Wave Hill Home- stead, 17.39 S, 130.52 E, 13-VIII- 1969, M. Mendum." in ANIC. Par- atypes: 1 c? , same data as holotype, in ANIC; 1*, 7 km S of Inverway Homestead, 17.55 S, 129.38 E, 25- VI-1969, M. Mendum" in ANIC; Ic?, "Western Australia, Port Head- land (airport), 19-11-1973, E. M. Exley" in UQ; 4. . 1865. Hemiptera nova vel minus cognita. Ann. Soc. Entomol. Fr. ser. 4, 5:163-188. . 1867. Analecta Hemipterologica. Berliner Entomol. Z. 10:151-172, 381-394. .1868. Hemiptera Fabriciana. T. K. Svenska VetenskAkad. Handl. 7:1-148. 1870. Hemiptera insularuni Philippinarum- bidrag till Philippinska oarnes Hemipteren-fauna. Ofvers. VetenskAkad. Forh. 27:607-776. . 1872. Genera Lygaeidarum Europae dis- posuit. Ofvers. VetenskAkad. Forh. 29:37-62. — . 1874. Enumeratio Hemipterorum. Pt. 4. K. Svenska VetenskAkad. Handl. 12:1-186. Thunberg, C. p. 1822. Dissertatio entomologica de Hemipteris rostratis Capensibus. Part 4. Acad. Typogr. , Upsaliae. 6 p. Tillyard, p. 1926. The Insects of Australia and New Zealand. Angus & Robert.son, Ltd., Sydney. 560 p. Tryon, H. 1892. Zoology of British New Guinea. 11. Hemiptera. Ann. Queensland Mus. 2:13-24. TuRTON, W. 1800. A General System of Nature Through the Three Grand Kingdoms etc., Vol. 2. Voss & Morris, Swansea. 717 p. Uhler, p. R. 1860. Hemiptera of the North Pacific Exploring Expedition under Com'rs. Rodgers and Ringgold. Proc. Acad. Nat. Sci. Philadelphia. 12: 221-231. Walker, F. 1872. Catalogue of the Specimens of Hemiptera Heteroptera in the Collection of the British Museum. Part 5. British Museum (Natural History), London. 202 p. White, F. B. 1878. List of the Hemiptera of New Zealand. Entomol. Mon. Mag. 15:31-34, 73-76. Wolff, J. F. 1802. Iconcs Cimicum dcscripiionibus illustratae. Part 3. Joann. Jacobum Palm, Er- iangae. 40 p. Woodward, T. E. 1962. Lectotype selection for three of Erichson's species of Australian Lygaeidae (He- miptera: Heteroptera). J. Entomol. Soc. Queens- land 1:50-66. ZiMSEN, E. 1964. The Type Material of J. C. Fabri- cius. Munksgaard, Copenhagen. 656 p. 440 The University of Kansas Science Bulletin montanus Fig. 46. Arocatus spp. a-f. Abdominal dorsum of named species: female left half, male right half, g, h. A. montanus, sp. nov. , pygophore: g, dorsal view; h, lateral \'iew. The Lygaeinae of Australia 441 m 'J iM chiasmus fastosus Fig. 47. Arocatus spp. a-h. Pygophore of named species: a, c, e, g, dorsal view; b, d, t, h. laierai view. 442 The University of Kansas Science Bulletin rusticus continctus 0.5mm chiasmus 0.125mm montanus 0 5 mm continctus aenescens Fin. 48. Arocalus spp. a-1. Phallotheca of named species: a, c, e, g, i, k, dorsal view ot lett apex showing phallothecal process; b, d, f, h, j, 1, lateral view, m, n. A. continctus Distant, pygophore: m, dorsal view; n, lateral view. The Lygaeinae of Australia 443 Fig. 49, Arocatus spp. a-d, i-i, k-n. Ejaculaiory reservoir oi named species: a, c, f, h, k. m, lateral view; b, d. s, i, 1, n, dorsal view. e. A. rusticus (Stal), left lateral view of partially inflated aedeagus. j. A. contmctus Distant, left lateral view of inflated aedeagus. 444 The University of Kansas Science Bulletin montanus Fu;. 30. Arocalus spp. a-r. Left claspcr of named species: a, d, g, j, m, p, dorsal view; b, e, h, k, n, q, posterior vi c, f, i, 1, o, r, interolateral view. The Lygaeinae of Australia 445 continctus Fig. 51. Arocatus spp. a-f. Spermatheca of named species. 446 The University of Kansas Science Bulletin Fig. 52. Arocatus spp. a-f. Ovipositor of named species: a, c, e, first valve; b, d, f, second valve. The Lygaeinae of Australia 447 025mm fastosus continctus Fig. 53. Arocatus spp. a-1. 0\ipositor of named species: a, c, e, first valve; b, d, f, second valve; g-1, apex of second valvula, lateral view. 448 The University of Kansas Science Bulletin doddi 0.125mm 0.25mm Fig. 54. Astacops spp. a^ b, f, g. Ejaculatory reservoir of named species: a, f, dorsal view; b, g, lateral view, c-e, h-j. Left claspcr of named species: c, h, dorsal view; d, i, posterior view; e, j, interolateral view. The Lygaeinae of Australia 449 0.25 mm 0.5 mm Fig. 55. Astacops viridiventns Stal. a. Phallothcca, dorsal view of left apex showing phallothccal process, b, c. Pygophore: b, lateral view; c, dorsal view. d. Apex of left second valvula, lateral view, e, f. Inflated aedcagus: e, right lateral view; f, ventral view, g, h. Ovipositor: g, second v£dve; h, first valve, i. Spermatheca. 450 The University of Kansas Science Bulletin walkeri papuas Fig. 56. Scopiastella spp. a-e. Apex of pygophorc of named species, lateral view. f-t. Left clasper of named species: f, 1, 1, o, r, dorsal view; g, j, m, p, s, posterior view; h, k, n, q, t, interolateral view. The Lygaeinae of Australia 451 elegans woodwardi nigricoronatus ^^^^^,^^ costalis nigricoronatus militaris Fig. 57. Scopiaslella spp. a-i. Apex of left second valvula of named species, lateral view, j, I. S. nigricoronatus. sp. nov. ovipositor: j, first valve; 1, second valve, k, m. Spermatheca of named species. The University of Kansas Science Bulletin C I k/'il^"^^-^.. nigncoronatus laticeps Fig. 58. a, b, e, g, h. Scopiastella spp. a, b. S. militaris (Distant), pygophore: a, dorsal view; b, lateral view. e. S. nigncoronatus, sp. nov. , partially inflated aedcagus, left lateral view, g, h. 5. militaris (Distant), ejaculatory reservoir: g, lateral view; h, dorsal view, c, d, f, i, j, k. Scopiastes spp. c, d. 5. bicolor Distant, pygophore: c, dorsal view; d, lateral view. f. 5. degeni (Slal), partially inflated aedeagus, left lateral view. i-k. 5. laticeps Breddin. i, j. Ejaculatory rcscr\'oir: i, dorsal view; j, lateral view. k. Phallotheca, dorsal view of left apex showing phallothecal process. The Lygaeinae of Australia 453 laticeps perplexus yorkensis obscurus impeditus degeer taroomi hackeri scudderi kurandensis obliquus eylandtensis 0 25mm obliquus eylandtensis Fig. 59. Scopiastes spp. a-1. Apex of pygophore of named species, lateral view. m-r. Left clasper of named species: m, p, dorsal view; n, q, posterior view; o, r, interolateral view. 454 The University of Kansas Science Bulletin Fig. 60. Scopiasles spp. a-x. left clasper of named species: a, d, g, j, m, p, s. v, dorsal view; b, e, h, k, n, q, t, w, posterior view; c, f, i, 1, o, r, u, x, interolatcral view. The Lygaeinae of Australia 455 kurandensis bicolor obdquus degeeri facetus 0.25mm Fig. 61. Scopiastes spp. a-i. Left clasper of named species: a, d, g, dorsal view; h, e, h. posterior \iew; c, f, i, intcrolaicral view. j-o. Spcrmatheca of named species. 456 The University of Kansas Science Bulletin bicolor laticeps perplexus impeditus hackeri kurandensis degeen yorkensis obliquus 10 mm Fig. 62. Scopiastes spp. a-1. Apex of left second \al\ ula ot named species, lateral view. in-o. Ovipositor of named species: m, o, second valve; n, first valve. TnK Lygaeinaf, of Australia 457 torrida Fk;. 63. Thunbergia spp. a-f. Pygophore of named species: a-c, dorsal view; d-f, lateral view. g-i. Phallotheca of named species, dorsal view of left apex showing phallothecal process, l-o. Ejaculatory reservoir of named species: 1, n, lateral view; m, o, dorsal view. 458 The University of Kansas Science Bulletin ruf ifemur Fig. 64. Thunbngia spp. a, b. Phallotheca of named species, lateral view, c, d. /'. augur (S{k\). intlated aedeagus: c, ventral view; d, left lateral view. e-j. Left clasper of named species: e. h. posterior view; f, i, dorsal \iew; g, j, interolateral view. The Lygaeinae of Australia 459 torrida Fig. 65. Thunbergm spp. a-c, g. Spermatheca of named species, cl-f. T. augur (Stal), left rlasper: d. posterior view; e, dorsal view: f, interolateral view. 460 The University of Kansas Science Bulletin rufifemur torrida I.Omtn Fig. 66. Thunbergia spp. a-h. Ovipositor of named species: a, c, e, g, first valve; b, d. f, h. second valve. The Lygaeinae of Australia 461 exilis 0.25mm 0.5 mm 0.25mm 0 5mm Fig. 67. WoodwardiaUei exilis, sp. nov. a. PhaJlothcca, dorsal view of left apex showing phallothecal process, b. Partially inflated aedeagus, left lateral view, c, d. Ejaculatory reservoir: c, lateral view; d, dorsal view. c-g. Left clasper: e, dorsal view; f, posterior view; g, interolateral view, h, i. Pygophore: h, dorsal view; i. lateral view. 462 The University of Kansas Science Bulletin Fig. 68. WoodwardiasUs monleilhi, sp. nov. a, c, d. Ovipositor: a, apex of left second valvula, lateral view; c, second valve; d, first valve, b. Spermatheca. The Lygaeinae of Australia 463 Fig. 69. Aspilocoryphus australicus Stal. a, b. Ejaculatory reserxoir: a, lateral view; b, dorsal view. c. Pariially mllatcd aedeagus, left lateral view. d. Spermatheca. e, f. Pygophore: e, dorsal view; f, lateral view. g. Phallotheca, dorsal view of left apex showing phallothecal process, h-j. Left clasper: h, dorsal view; i, posterior view; j. inierolatcral view, k, 1. Ovipositor: k, first valve; 1, lateral view. 464 The University of Kansas Science Bulletin cardinalis 0.5 mm Fig. 70. Graptoslelhus spp. a-h. Ejaculatory reservoir of named species: a, c, e, g, lateral view; b, d, f, h, dorsal view, i-m. Pygophore of named species: i-1, dorsal view; m, lateral view. The Lygaeinae of Australia 465 pubescens varipictus Fig. 71 . Graptostethus spp. a-1. Left clasper of named species: a, d, g. j, dorsal view; b, e, h, k, posterior view; c, f. i. 1. intciolatcral view. m-p. Apex of left second valvula of named species, lateral view. 466 The University of Kansas Science Bulletin Fig. 72. Graptostelhus spp. a-d. Spermathcca of named species, i, j. G. servus (Fabricius): i, left lateral view of partially inflated aedeagus; j, phallotheca, dorsal view of left apex showing phallothecal process, g-j. Ovipositor of named species: g, i, .second valve; h, j, first valve. The Lygaeinae of Australia 467 eucurus 0.26mm 0 5mm 0.5 mm F.G. 73. Pyrrhobaphus leucurus (Fabricius). a-c. Left clasper: a, dorsal view; b, posterior v.ew; c. '"'"o ^ "^' ^ "^^ ^j e. Pygophore: d, dorsal view; e, lateral view. f. Partially inflated aedeagus, nght lateral v.ew. g. Phalio.hcca. dorsal view ofleft ape.x showing phallothecal process, h, i. Ejaculatory reservoir; h. lateral v.ew; ., dorsal view. 468 The University of Kansas Science Bulletin leucurus Fig. 74. Pyrrhobaphus leucurus (Fabricius). a. Spermatheca. b. Apex of left second valvula, lateral view, c, d. Ovipositor: c, first valve; d, second valve. The Lygaeinae of Australia 469 Fig. 75. Melanerythms spp. a-h. Pygophore of named species: a, c, e, g, lateral view; b. ci, 1, h. dorsal view. 470 The University of Kansas Science Bulletin Fig. 76. Melanerythrus spp. a, e, i, m. Phallotheca of named species, donsal view oi left apex sfiowing phallotheca] process, b-d, f-h, j-1, n-p. Left clasper of named species: b, f, j, n, dorsal view; c, g, k, o, posterior view; d, h, 1, p, interolateral view. The Lygaeinae of Australia 471 mactans mutilatus ignitus biguttatus 0 5 mm Fig. 77. Melanerythrus spp. a-h. Ejaculatory reservoir of named species: a, c, e, g, lateral view; b, d, f, h, dorsal view. i. M. mactans (Stal): inflated aedeagus, right lateral view. j-m. Phallotheca of named species, left lateral view. 472 The University of Kansas Science Bulletin mutilatus biguttatus biguttatus mactans uptoni ignitus mactans uptoni 0.25 nnm Fig. 78. Afelanerythrus spp. a-e. Apex of left second valvula of named .species, lateral view. t-h. Spermatheca oi named species. The Lygaeinae of Australia 473 ignitus Fig. 79. Melanerylhrus spp. a-h. Ovipositor of named species: a, c, c, g, first valve; b, d. 1, h. second valve. 474 The University of Kansas Science Bulletin rubiginosus rubiginosus rubiginosus Fig. 80. Melanolelus spp. a-f. Left clasper of named species: a, d, dorsal view; b, c. posterior view; c, f, interolateral view. g. M geriae. sp. nov. , left lateral view of inflated acdeagus. h, i. Phallotheca of named species, dorsal view of left apex showing phallotheca! process, j-m. Ejaculatory reservoir of named species: j, 1, lateral view; k, m, dorsal view, g, h. Median projection of left margin of dorsal pygophore opening of named species, p, q. M. rubiginosus (Walker), pygophore: p, lateral view; q, dorsal view. The Lygaeinae of Australia 475 1.0 mm 0.25mm rubigmosus Fig. Q\ . Melanotelus spp. a-d. Ovipositor of named species: a, c, first valve; b, d, second valve, e. f. Spcrmalheca ol named species. 476 The University of Kansas Science Bulletin pacificus pacificus \ 0.25mm Fig. 82. Spilostelhus spp. a-f. Pygophore of named species: a, c, e, lateral view; b, d, f, dorsal view. g-1. Ejaculatory reservoir of named species: g, i, k, lateral view; h, j, 1, dorsal view. m-o. Phallotheca of named species, dorsal view of left apex showing phallothecal process. The Lygaeinae of Australia 477 IiG. 83. Spilostethus spp. a-i. Left claspcr of named species: a, d, g, dorsal view; b, c, h, posterior view; c, f. i. interolateral view, j, k. Phallotheca of named species, left lateral view. 1. S. hospes (Fabricius), left lateral view of inflated aedeagus. 478 The University of Kansas Science Bulletin 0.5 mm Fic;. 84. SptlosMhus spp. a-c. Spcrmathcta ol named species, d-i. Ovipositor oi named species: g, i, second valve. 0.5mm , h, first valve; e, The Lygaeinae of Australia 479 purpurascens microps Fig. 85. Oncopeltm spp. a-1. Ejaculatorv reservoir of named species: a, c, c, g, i, k, lateral view, b, ci, f. h. j. 1. dorsal view. m-o. 0. sordidus (Dallas), m. Inflated aedeagus, right lateral view, n, <.. PvK<,phnr.-: n, laieral view; o. dorsal view. 480 The University of Kansas Science Bulletin purpurascens / /' sordidus quadngutatus 0 25 mm Fig. 86. Oncopellus spp. a-1. Left clasper of named species: a, d, g, j, dorsal view; b, e, h, k, posterior view; c, f, i, 1, interolateral view. The I.ygaeinae of Australia 481 1.0 mm Fig. 87. Oncopellm spp. a-c. 0. alromjm (Gudrin), a, b. Ovipositor: a, first valve; b, second valve, c. Spermatheca. d- i. Left clasper of named species: d, g, dorsal view; e, h, posterior view; f, i, interoiateral view. 2566^^050 UK If II II I III n II I III illi.iiil'll .llil!|::lLlilil...l. li;l ill 2044 093 362 jb; Date Due