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X^GAAHs/ H* 2 xnxixsNi z nvinoshxiws^s 3 1 aVaanzuB rar i es^smithsonian institution Noixnxix: co 5 co ^ ^ 5 5 „ * CO X X X X X X X X* W W X X X X X X 0) CO o s s u, c§ _>> o 3 D o t- "C 3 c cd 3 c O o *2 O T3 lx s 3 o D +s C/2 X (fl 00 Vh ex o 'cx cd C/2 0) Vh cx ao t-H cd Uh o X CO C/3 (continued on next page) Structure/ Character Ancestral State Derived State Basis for classfn Times derived Times lost Distribution of derived state by lineage (Fig. 186) 52 Lafontaine T3 C 1 'o' 3 c > ;3 > O oo Os — 1 ' — 1 (continued on next page) Structure/ Character Ancestral State Derived State Basis for classfn Times derived Times lost Distribution of derived state by lineage (Fig. 186) The Euxoa detersa group (Lepidoptera: Noctuidae) 53 OO IT) ro G c G HH 1-1 1—1 X X X X X W P-1 PJ PQ UJ cd H CD G O X cd G cd X aJ ’> > M O 0* O g m «cr >/-> 2 ’> _ <3-> •r £ cs S a Quaest. Ent., 1981, 17(1,2) Structure/ Character Ancestral State Derived State Basis for classf n Times derived Times lost Distribution of derived state by lineage (Fig. 187) 56 Lafontaine 3 13 X) C/3 cd cd X x> C/2 cd cd s T3 *3 > O > 3 > 3 O .Sr ’> O 1 3 Ovipositor - conical absent present setae (continued on next page) Structure/ Character Ancestral State Derived State Basis for classfn Times derived Times lost Distribution of derived state by lineage (Fig. 187) The Euxoa detersa group (Lepidoptera: Noctuidae) 57 o o oo =i x> 1/3 o c3 'c« o C/3 ’c« o C/3 3 d> C/3 S3 a ’> O O C/3 cd X) .CX "> O t-i C o =s Q Vh > e 2 of oo Ov OT VO oo o O CO O > OV OV of co 00 vd 03 - 1 stub = a terminal filament of 5 segments or less (Morihara and McCafferty 1979) 2 + = present, - = absent, ? — unknown. 3 (.n) = ratio of length of terminal filament to that of the lateral filaments. Caudal filaments in Mayfly nymphs 143 !| ii and salinity all increase after the confluence of the two streams. jj Caudal filaments and vulnerability to predation. There was no mortality or loss of nymphs in control cells of the in situ predator-prey experiment conducted on the Highwood River. Fewer B. bicaudatus and B. tricaudatus nymphs were found in cells containing H. pacifica nymphs. Contents of predator foreguts indicated that nymphs of both mayfly species were eaten without preference. We tentatively conclude from this short-term study that neither two-tailed nor three-tailed Baetis nymphs had an advantage in avoiding predators. Morphological Studies Morphological studies showed one pair of subapical setae on each B. tricaudatus claw and as many as three pairs on the claws of B. bicaudatus (Platte 1). Published records (Table 5) show that all species of Baetis nymphs with subapical setae on claws possessed a terminal filament which was greatly reduced (in some species to a stub) and also inhabited fast-flowing waters. In North America, subapical setae on claws has been reported in two species groups, B. lapponicus and R. hageni (Morihara and McCafferty 1979). In the western Palearctic, these setae are present in four groups: Baetis, B. lapponicus , Baetis and B. rhodani Mtiller-Liebenau 1969, 1971, 1974a). Species within these groups live in running waters at high altitudes and latitudes or on islands (McDunnough 1936, Miiller-Liebenau 1971, Edmunds et al., 1976, Cobb and Flannagan 1980). Nymphs of the one northern species, Baetis macani bundyae, that lack subapical setae on claws inhabit ponds (Lehmkuhl 1973). Miiller-Liebenau (1974b) found subapical setae on nymphal claws of all three species of Heterocloeon McDunnough ( = Rheobaetis)\ these nymphs also possess a reduced (one segment) terminal filament and inhabit swift-flowing rivers. We know of no other mayfly species in which nymphs possess subapical setae on claws. DISCUSSION Our laboratory study showed that stonefly predators ate more B. tricaudatus than R. hageni nymphs. Evidently, the flattened, clinging, R. hageni nymphs were more difficult for the stonefly nymphs to capture than the rounded, swimming, B. tricaudatus nymphs. This tends to support the hypothesis of Elynes (1970) that flattened nymphs adhering to substrate surfaces may offer protection against predation. Fewer Baetis nymphs with caudal filaments were eaten by predators during light periods. This was a minor advantage since predators fed significantly more at night. In the field, there was no significant difference between two- and three-tailed nymphs. (B. bicaudatus and B. tricaudatus) in avoiding the stonefly nymph, H. pacifica. Based on these studies, we discard the idea that the major function of the caudal filaments of mayfly nymphs is to deter predators. We also concur with Schmidt’s (1951) view that caudal filaments do not assist in the molting process. Apparently, movement of caudal filaments does not significantly aid nymphs during molt as suggested by Drenkelfort (1910). Species replacement along a stream channel is a common phenomenon (Ide 1935, Allan 1975, Hawkes 1975, Ward and Berner 1980). In headwater areas of mountain streams, Dodds Quaest. Ent., 1981, 17 (1,2) 144 Corkum and Clifford and Hisaw (1924) and Hora (1930) suggested that species of Baetis with progressively shorter terminal filaments live in habitats with increasing current velocities. Although both B. bicaudatus and B. tricaudatus nymphs occupied fast-water areas, we found no relationship betwen current velocity and stream site inhabited by these two- and three-tailed species of Baetis. The difference between no current and some current may be of major significance to an organism, whereas large differences among current velocities may be of minor importance (Philipson 1954). Edmunds et al., (1976) and Ward and Berner (1980) have indicated that B. bicaudatus nymphs live at high altitudes. Ward and Berner (1980) report the upstream limit of B. tricaudatus nymphs on St. Vrain Creek Colorado ca. 2400 m, several degrees of latitude south of our site. On the Highwood River, the upstream limit of B. tricaudatus at 1500 m ('$0o23’ N) corresponds to the entrance of Etherton Creek, In our study, no single environmental factor influenced the distribution of the two mayfly species along a stream section. Rather, a combination of factors were correlated with the upstream limit of the three-tailed, B. tricaudatus nymphs. Typically, fast-flowing water is well aerated and carries food to aquatic organisms. The organisms, however, must either avoid the force of the current or be able to maintain their position. Baetis nymphs frequently occupy the upper substrate surfaces and face into the current (Dodds and Hisaw 1924). Hora (1930) suggests that nymphs facing into torrential water must hold the caudal filaments together to aid water flow over their body. He conceives that the terminal filament may have been “pressed out of existence, metaphorically speaking, by the pressure of the lateral [filaments].’’ Perhaps caudal filaments act to reduce drag on nymphs exposed to current flow. From morphological studies of B. bicaudatus and B. tricaudatus nymphs, we observed subapical setae on claws of both of these fast-water species. All species of Baetis with subapical setae on claws also have a reduced terminal filament. We suggest that these structures are mechanoreceptors. If so, they should be able to detect water-borne vibrations (Mclver 1975). Although elliptical shape of the socket may restrict the directional movement of setae, it may be possible that these structures are used by nymphs to detect subtle changes in flow within the boundary layer ( sensu Ambiihl 1959). The setae may also assist in orienting nymphs into the current. These structures would be of particular importance to nymphs inhabiting upper substrate surfaces exposed to flowing water. Popovici-Baznosanu (1906) suggested that caudal filaments may function as a respiratory organ. Since Baetis nymphs with a reduced terminal filament appear to be found in specific habitats (high altitude and latitude), gradual reduction in an appendage may be a response to lower respiratory demands in these cooler, fast-flowing waters. ACKNOWLEDGEMENTS We thank J. J. H. Ciborowski, D. A. Craig, V. Gotceitas, B. K. Mitchell and I. Muller-Liebenau for their comments on this study. R. L. Baker provided helpful information on the study site and J. J. H. Ciborowski assisted with field work. We are grateful to R. W. Mandryk who prepared mayfly specimens from which SEM photographs were taken by G. Braybrook. Funding was provided by grants from the Boreal Institute for Northern Studies (Alberta Government Grant-In-Aid program) to L.D.C. and N.S.E.R.C. to H.F.C. Caudal filaments in Mayfly nymphs 145 LITERATURE CITED ALLAN, J. D. 1975. The distributional ecology and diversity of benthic insects in Cement Creek, Colorado. Ecology 56: 1040-1053. AMBUHL, H. 1959. Die Bedeutung der Stromung als okologischer Faktor. Schweizerische Zietschrift fur Hydrologie 21: 133-264. AMERICAN PUBLIC HEALTH ASSOCIATION. 1971. Standard methods for the examination of water and wastewater. 13th ed. A.P.H.A., New York. CLIFFORD, H. F. 1970. Variability of linear measurements throughout the life cycle of the mayfly Leptophlebia cupida (Say) (Ephemeroptera: Leptophlebiidae). The Pan-Pacific Entomologist 46: 98-106. COBB, D. G., and J. F. FLANNAGAN. 1980. The distribution of Ephemeroptera Biology in Northern Canada, pi 55-166. In Advances in Ephemeroptera. Eds. Flannagan, J. F., and K. E. Marshall. Plenum Publ. New York. CORKUM, L. D., and H. F. CLIFFORD. 1980. The importance of species associations and substrate types to behavioural drift, p. 331-341. In Advances in Ephmeroptera Biology. Eds. Flannagan, J. F., and K. E. Marshall. Plenum Publ. New York. DRENKELFORT, H. 1910. Neue Beitrage zur Kenntnis der Biologie und Anatomie von Siphlonurus lacustris Eaton. Zoologische Jahrbucher (Anatomie) 29: 527-617. DODDS, G. S., and F. L. HISAW. 1924. Ecological studies of aquatic insects. 1. Adaptations of mayfly nymphs to swift streams. Ecology 5: 137-148. EDMUNDS, G. E., Jr., S. L. JENSEN, and L. BERNER. 1976. The mayflies of North and Central America. University of Minnesota Press, Minneapolis. HAWKES, H. A. 1975. River zonation and classification, p. 312-374. In River Ecology. Ed. B. A. Whitton. Blackwell Sci. Publ., Oxford. HORA, S. L. 1930. Ecology, bionomics and evolution of the torrential fauna, with special reference to the organs of attachment. Philosophical Transactions of the Royal Society. B218: 171-282. HUGHES, D. A. 1966. On the dorsal light response in a mayfly nymph. Animal Behaviour. 14: 13-16. HYNES, H. B. N. 1970. The Ecology of Running Waters. University of Toronto Press, Toronto. IDE, F. P. 1935. The effect of temperature on the distribution of the mayfly fauna of a stream. Publications of Ontario Fisheries Research Laboratory 50: 1-76. KEFFERMULLER, M. 1980. Un Changement dans la Taxonomie de la Famille Baetide. p. 115-121. In Advances in Ephemeroptera Biology, eds. Flannagan, J. F,., and K. E. Marshall. Plenum Publ. New York. LEHMKUHL, D. M. 1973. A new species of Baetis (Ephemeroptera) from ponds in the Canadian arctic, with biological notes. The Canadian Entomologist 105: 343-346. McDUNNOUGH, J. 1936. A new arctic baetid (Ephemeroptera). The Canadian Entomologist 68: 33-35. McIVER, S. B. 1975. Structure of cuticular mechanoreceptors of arthropods. Annual Review of Entomology 20: 381-397. MORIHARA, D. K., and W. P. McCAFFERTY. 1979. The Baetis larvae of North America (Ephemeroptera: Baetidae). Transactions of the American Entomological Society 105: Quaest. Ent., 1981, 17 146 Corkum and Clifford 139-221. MULLER-LIEBENAU, I. 1969. Revision der Europaischen Arten der Gattung Baetis Leach, 1815 (Insecta, Ephemeroptera). Gewasser und Abwasser, 48/49. 214 S., Max-Planck-Gesellschaft. Dokumentationsstelle, Gottingen. MULLER-LIEBENAU, I. 1971. Ephemeroptera (Insecta) von den Kanarischen Inseln. Gewasser and Abwasser, 50/5 1 : 7-40. MULLER-LIEBENAU, I. 1974a. Baetidae aus Sudfrankreich, Spanien und Portugal (Insecta, Ephemeroptera). Gewasser und Abwasser, 53/54: 7-42. MULLER-LIEBENAU, I. 1974b. Rheobaetis : a new genus from Georgia (Ephemeroptera: Baetidae). Annals of the Entomological Society of America 67: 555-567. PERCIVAL, E. and H. WHITEHEAD. 1929. A quantitative study of the fauna of some types of stream-bed. Journal of Ecology. 17: 282-314. PHILIPSON, G. N. 1954. The effect of water flow and oxygen concentration on six species of caddis fly (Trichoptera) larvae. Proceedings of the Zoological Society of London. A. 124: 547-564. POPOVICI-BAZNOSANU, A. 1906. Contributions a l’etude de Forganisation des larves des Ephemerines. Archives de zoologie experimental et generate 5: 66-78. SCHMIDT, H. 1951. Amputation und Regeneration von Schwanzfaden und Abdominlsegmenten bei Larven der Ephemeridenart Cloeon dipterum L. und ihr Einfluss auf die Hautungsintervalle. Zool. (Phys) 62: 395-428. SNEATH, P. H. A. and R. R. SOKAL. 1973. Numerical Taxonomy. W. H. Freeman and Co., San Francisco. VENTER, G. E. 1961. A new ephemeropteran record form Africa. Hydrobiologia 18: 327-331. WARD, J. V. and L. Berner. 1980. Abundance and altitudinal distribution of Ephemeroptera in a Rocky Mountain stream, p. 169-177. In Advances in Ephemeroptera Biology. Eds. Flannagan, J. F., and K. E. Marshall. Plenum Publ. New York. THE DISTRIBUTION OF MOSQUITO (DIPTERA, CULICIDAE) LARVAE IN SOUTHERN ALBERTA, 1976-1978 P.J. Scholefield' G. Pritchard 2 M.A. Enfield3 Department of Biology University of Calgary Calgary, Alberta T2N1N4 Quaestiones Entomologicae 17:147-168 1981 ABSTRACT Twenty-seven species of Aedes, four species of Culiseta , two species of Culex and one species of Anopheles were collected during five surveys of water bodies in Alberta south of 52° N during 1976, 1977 and 1978. Identifications were made on 4th stage larvae and confirmed after rearing to adult. The species of Aedes showed four distribution patterns: Group 1 was largely restricted to the mountains and foothills; Group 2 was found in all parts of the study area; Group 3 occurred everywhere but in the prairie zone; Group 4 was largely restricted to the parkland and prairie. Because of low numbers, the distribution of seven species of Aedes could not be determined. Of the species in other genera, only Culiseta inornata (a Group 2 species) was common enough to determine distribution, although Cs. alaskaensis appears to belong to Group 1. Apart from these broad patterns, the habitat preferences of individual species could not be determined. Of the 20 commonest species, 18 were associated with 13-20 other species at one time or another, and two had nine associated species. The number of pools that formed by snow-melt or rain-fall, the dates of appearance of larvae, and the relative species abundance all changed markedly from year to year. RESUME Vingt-sept especes d’ Aedes, quatre especes de Culiseta, deux especes de Culex et une espece d’ Anopheles ont ete collectionnees au cours de cinq inventaires des plans d’eau situes au sud du 52ie me parallele Nord en Alberta, en 1976, 1977 et 1978. Les especes ont ete identifiees a partir des larves du quatrieme stade, et les identifications ont ete verifiees •Present address: Pesticide Chemicals Branch, Alberta Environment, Calgary 2Author to whom correspondence should be addressed 3Present address: A.D.A.S. Sub Centre, Wye, Kent, U.K. 148 Scholefield, Pritchard and Enfield grace a Velevage de ces memes larvaes jusqu’au stade adulte. La distribution des especes d’ Aedes formait quatre patrons: le groupe I etait principalement restreint aux montagnes et aux piedmonts; le groupe 2 se retrouvait dans toutes les regions inventoriees; le groupe 3 se retrouvait partout, sauf dans la prairie; le groupe 4 etait en grande partie restreint au “parkland” et a la prairie. La distribution de sept especes d’ Aedes n pu etre etablie a cause des faibles captures. Parmi les especes appartenant aux genres autres qu’ Aedes, seule Culiseta inornata etait assez commune pour pouvoir etablir sa repartition (el le se classe dans le groupe 2); Cs. alaskaensis semble appartenir au groupe I. Mis a part ces patrons de distribution generaux, Vhabitat prefere de chaque espece n’a pu etre determine. Parmi les vingt especes les plus communes, dix-huit etaient associees avec de treize a vingt autres especes a une occasion ou I’autre, et deux especes se retrouvaient chacune en association avec neuf especes. Le nombre d’etangs formes par la fonte des neige ou la pluie, les dates d’emergence des larvaes, et I’abondance relative des especes ont tous varies d’une annee a I’autre. TABLE OF CONTENTS Introduction 148 Methods . 149 Results 154 Group 1 - Mountains and Foothills Species 154 Group 2 - Ubiquitous Species . 156 Group 3 - Non-prairie Species . 157 Group 4 - Parkland and Prairie Species . 157 Species of Uncertain Distribution . 159 Discussion . 161 Acknowledgements . 166 References 166 INTRODUCTION Proper identification of species is basic to any biological study. Furthermore, studies of mosquito-carried diseases and mosquito abatement in Alberta cannot make efficient use of data already accumulated and analysed unless the species, both in the current programs and in work already done, are properly known. The problem of adult migration into an area in which a larviciding program has been carried out, can hardly begin to be solved without thorough knowledge of the species involved. In the report of the “Symposium on Biting Fly Control and Environmental Quality” held in Edmonton in 1972, the summary of the section on Behaviour and Ecology of Populations (Mclver, 1973) contained the following passage: “ When considering a control program, four superficially simple questions need to be asked about the pest species: 1 . What is it? 2. Where is it? 2. What is it doing? 4. How does it accomplish its observed behaviour?” Mosquito (Diptera, Culicidae) larvae in Southern Alberta 149 This report attempts to answer the first two questions for mosquitoes in Southern Alberta. Such a study was necessary because, although several reports about mosquitoes of Alberta have included species lists (e.g. Strickland, 1938; Happold, 1965; Pucat, 1965; Wada, 1965; Tawfik and Gooding, 1970), these studies represented a limited approach to species identification and all contained some misidentifications through use of incomplete or inaccurate keys. For example, Enfield (1977) reported that two species, Aedes euedes Howard, Dyar and Knab 1917 and Aedes mercurator Dyar 1920 have long remained unrecognized. Larvae of Aedes euedes key to Aedes riparius Dyar and Knab in Rempel (1950) and Carpenter and LaCasse (1955), while adult males have genitalia that resemble those of Aedes excrucians (Walker) . Aedes barri Rueger 1958 and Aedes beklemishevi Denisova 1955 are synonyms for Aedes euedes (Wood, 1977). Aedes mercurator larvae key to one of four other species in conventional keys. Until this species was recognized in Calgary in 1975 (Enfield, 1977), it was recorded only from Dawson, Yukon Territory (Dyar, 1920), and from the U.S.S.R. (Danilov, 1974). Many adults labelled Aedes stimulans (Walker) in the Canadian National Insect Collection are in fact Aedes mercurator , and its range is now known to extend across much of Northern Canada (Wood, 1977). These problems stimulated a survey of the mosquitoes in Southern Alberta. We felt that the area of Alberta approximately south of latitude 52°N, within which Calgary is conveniently central, was the maximum that could be covered in a study of this kind. Further, we thought that only by collecting extensively in all ecological zones and by both identifying larvae and rearing them to obtain associated adults, could familiarity with the whole group be gained and problems such as those involving Aedes mercurator and Aedes euedes be exposed. The data, which are summarized in this paper, may be found in a more extensive form, together with a key to 4th stage Aedes larvae occurring in southern Alberta in an unpublished report (Scholefield, Enfield and Pritchard, 1979). METHODS The study area was surveyed five times. Samples of larvae were collected along planned routes so that the major ecological zones and different types of water bodies were included in each trip. Distribution of sites that yielded larvae is shown in Figures 1 & 2. In the Spring (April-May) of 1976, mosquito larvae were collected from 115 sites, from 45 sites in the Spring of 1977, from 61 sites in the Summer (June- July) of 1977, from 124 sites in the Spring of 1978, and from 41 sites in the Summer of 1978. Most collections were obtained by driving along set routes and sampling water bodies which were seen from the road. However, spring sampling in the Southeast of the province in 1977 and 1978 was done during a helicopter tour by Mr. J. McIntosh (Alberta Environment) to the various abatement areas under his jurisdiction; one slough in or near each town was selected from the air, then sampled. In 1977 and 1978, a record sheet containing information on each site was filled out and a photograph of the site was attached. Larvae were captured by means of a standard pint dipper, and, wherever possible, dipping was continued at each site until about 50 larvae had been caught. The larvae were concentrated Quaest. Ent., 1981, 17 150 Scholefield, Pritchard and Enfield Fig. 1 . Sites which yielded mosquito larvae during spring (April-May) sampling. The 1 976 survey did not adequately cover the mountain zone and northern foothills and this was corrected in later years. In 1978 the whole region was surveyed but only those areas not visited in previous years are indicated. Also shown are the 4 ecological zones (heavy lines), the Columbia Ice-fields (Cl) and the Cypress Hills (CH), the major rivers (thin lines), and some towns (B = Banff, BR = Brooks, C = Calgary, CY = Cayley, CO = Coleman, D = Drumheller, F = Fort McLeod, J = Jasper, L = Lethbridge, M = Medicine Hat, N = Nordegg, R = Rocky Mountain House Mosquito (Diptera, Culicidae) larvae in Southern Alberta 1 :| Fig. 2. Sites which yielded mosquito larvae during summer (June-July sampling Quaest. Ent., 1981, 17 152 Scholefield, Pritchard and Enfield in a clear plastic cup by means of a cylindrical filter which was fitted into the top of the cup. As water containing larvae was poured in, the larvae were retained in the cup while excess water spilled out. Usually no more than about 25 larvae were kept in each cup, which was sealed with a plastic lid, and returned to the laboratory in a cooler. When samples were being transported, a stop of five minutes duration was made every 30 minutes to allow larvae to rise to the water surface and breathe. When 4th stage larvae were collected about one-third of the sample was killed in KAiAD 1 and preserved in 95% ethanol. These subsamples were potential replacements in case a whole sample should die during transportation, since mosquito larvae decay very rapidly after death, making identification impossible. Some pupae collected in the field represented species of which there were no larvae in the same sample. Thus collection of pupae for subsequent rearing to the adult stage was important, even though pupae are more difficult to keep alive during transportation. In the laboratory, larvae were kept in these same plastic cups at 20°C, in tap water which had been conditioned with dead slough grass for one week. The water was aerated to prevent formation of surface scum, and larvae fed on the micro-organisms on the surface of the decaying grass. Larvae were identified when in stage IV and some were returned to the cups to complete development; the remainder were killed in KAiAD and preserved in 95% ethanol. Because examination of preserved specimens in Edmonton and Calgary showed that many Aedes adults and larvae had earlier been misidentified, as many identified larvae as possible were reared to adults and then reidentified to reduce the chance of further misidentification. Pupae were placed individually or in groups of about 20 in cups, and the emerging adults were trapped in inverted plastic vials on top of a funnel made of plastic fly screen. Several hours after emergence, vials containing adults were placed in a freezer to kill the mosquitoes and within a few days the specimens were mounted on a pin with shellac and returned to the freezer for a few weeks to complete drying. A draft of a key now published in Wood et al., (1979) was the main basis for identification of larvae. Keys of Gjullin and Eddy (1972) and Carpenter and LaCasse (1955) (modified to include Aedes euedes and Ae. mercurator) were used for identification of adults and for identification of some larvae. Voucher specimens of stage IV larvae and adults of all species are stored in the Department of Biology, University of Calgary. 1 KAAD= 1 part kerosene, 10 parts 95% ethanol, 2 parts glacial acetic acid and 1 part dioxane. KAi AD = equal parts KAAD and 95% ethanol. Mosquito (Diptera, Culicidae) larvae in Southern Alberta 153 Fig. 3. Examples of the 4 types of distributions Group 1 - Mountains and Foothills: Ae. communis : Group 2 - Ubiquitous Ae. euedes : Group 3 - Non-Prairie: Ae. mercurator ; Group 4 - Prairie and Parkland: Ae. flavescens. Quaest. Ent., 1981, 17 154 Scholefield, Pritchard and Enfield The Atlas of Alberta (Government of Alberta, 1969) was used to define ecological zones in preference to the one used by Strickland (1938), because there appeared to be little evidence that the prairie/parkland zone was divisible into as many different areas for Aedes species as Strickland showed. Four ecological zones (Mountains, Foothills, Parkland, Prairie) were included within the study area (Figs. 1, 2 & 3). In 1977 and 1978, in addition to recording location, we attempted to categorize habitat type for each species. Four habitat characteristics were noted at each site: 1). whether the pond was open or shaded by trees; 2). whether the surface was shaded by slough grass or clear of vegetation; 3). whether or not there were willows, conifers or poplars around the margin; and 4). whether the water body was stagnant or was continually fed by seepage or continuous surface run-off. In 1978 water analysis was also made at each site containing larvae. The pH, conductivity, temperature, and dissolved oxygen content at the time of sampling were recorded with a view to further characterization of habitats of individual Aedes species. RESULTS Thirty-four species of mosquitoes in four genera were collected in this survey. Twenty-seven of these species belong to Aedes , most larvae of which inhabit temporary bodies of water formed by melting snow or rainfall. Although not important vectors of disease in Alberta, some species develop huge populations with a high nuisance rating. The species of Aedes show four distribution patterns (Fig. 3). Group 1 mosquitoes are largely restricted to the mountains and foothills; Group 2 mosquitoes occur in all 4 zones; Group 3 mosquitoes occur in 3 of the zones, but are missing from the prairie; and Group 4 mosquitoes are largely restricted to the parkland and prairie. Some overlap at the foothills-parkland border is not uncommon in species in Groups 1 and 4. Because of rarity, the distribution of seven species could not be ascertained. Group 1 - Mountain and Foothills species Aedes (Ochlerotatus) canadensis (Theobald) (15 collections) This species is included in the mountains-foothills group although specimens were taken at two sites west of Calgary at the western edge of the parkland zone. Most larvae were found in spring, but were also collected in summer in the mountains in 1977. Most were found in unshaded locations in forested areas, and in non-stagnant pools. The species was found on only three occasions by itself; at other times it was associated with one or more of nine species of Aedes. Aedes ( Ochlerotatus ) communis (de Geer) (36 collections) Classification of this species is discussed by Ellis and Brust (1973). It was one of the commonest species in the mountains and foothills, where larvae were found only in spring in snow-melt pools. Larvae were in forested areas, but in relatively open pools. The water in most pools was clear and cool and laboratory rearing was much more successful when the water was kept clear and well aerated. The species was found on 10 occasions by itself; at other times it Mosquito (Diptera, Culicidae) larvae in Southern Alberta 155 was associated with one or more of 20 other species. Aedes (Ochlerotatus) hexodontus Dyar (34 collections) Larvae of this species are also common inhabitants of open pools in the mountains, often found with larvae of Ae. communis and Ae. punctor . Predominantly a spring species, larvae were found at one location in the mountains in the summer of 1977. According to Carpenter and LaCasse (1955), some hatching may be delayed until long after the pool has been filled by snow-melt. It was rarely found by itself (four collections), occurring at one time or another with 19 other species. Aedes ( Ochlerotatus ) pionips Dyar (11 collec Not a common species, found predominantly in spring in open locations in forested areas of the foothills, although larvae were collected during the summer on one occasion in the mountains in 1977. It was found once by itself, on other occasions being associated with one or more of 17 other species. Larvae were difficult to rear, even in clear, well aerated water. Aedes ( Ochlerotatus ) pullatus (Coquillett) (47 collections) One of the most common mountain species, larvae were in open pools in both spring and summer. In 1977 it was found alone in a pool near the Columbia Icefields on 10th May (water temperature 14°C) and again on 5th July (water temperature 20°C). On both occasions 4th stage larvae and pupae were present, and the indication is therefore that eggs hatched on two occasions in 1977. The species was not present in this pool in the spring of 1978; Ae. communis was there instead. Aedes pullatus was found on 10 occasions by itself, and on other occasions with 20 other species. Aedes ( Ochlerotatus ) punctor (Kirby) (33 collections) Another common mountain - foothills species, collected mainly in spring but also taken in a few summer samples in 1977 and 1978. Larvae apparently prefer grass-covered, willow-fringed, open pools. Found on three occasions by itself, it occurred at other times with 23 other species of Aedes. Aedes (Ochlerotatus) schizopinax Dyar ) (1 1 collections) Like Aedes canadensis this species has been taken from the western edge of the parkland, but is otherwise restricted to the mountains and foothills. It is rather local, being found at only one location in 1978, but it did occur in fairly high numbers in certain locations in the southern part of the study area during the summer of 1977, when its larvae were among the most common in the Coleman area. Almost all samples were in open, grass-covered pools. Nielsen (1961) reports that larvae prefer small to moderately sized pools containing considerable vegetation and organic matter, often covered with scum. He also reports that adult females are not known to take blood meals. Aedes schizopinax was found only once by itself; at other times it occurred with one or more of 16 other species. During spring, larvae appeared later than those of the other species, but developed quickly both in the field and laboratory. Quaest. Ent., 1981, 17 156 Scholefield, Pritchard and Enfield Group 2 - Ubiquitous Species Aedes (Ochlerotatus) cataphylla Dyar (90 collections) A common species in all zones, apparently having a wide tolerance of habitat conditions. It is a spring species, and was the most frequently encountered species in the spring of 1976. Eggs apparently hatch on the first flooding, whether warm or cold, and this could account for the two records of larvae in the summer sampling in 1977. Larvae developed quickly and Carpenter and LaCasse (1955) report that it is one of the first Aedes species to appear on the wing. Fourth larval and pupal stages suffered heavy mortality when reared in the laboratory. Aedes cataphylla larvae were found alone on 14 occasions, and with 22 other species at other times, especially Ae. campestris, Ae.fitchii, and Ae. mercurator . Aedes ( Ochlerotatus ) euedes Howard, Dyar and Knab (45 collections) This species was fairly common in the 1976 survey, but was not recorded as frequently in 1977 or 1978. It is primarily a snow-melt pool species, but small numbers of larvae have been found in rain-water pools during the summer in the mountains. Larvae were found in many types of pools in all four ecological zones. It occurred only six times alone, at other times being associated with 17 other Aedes species. This was the most difficult of the commoner species to rear in the laboratory, partly because the larvae were very aggressive and apparently unable to withstand even slight crowding. Rate of development was about average, but overall development time was long because individuals grew to a larger size than most other species. In the spring of 1975, this species required six to seven weeks to complete development from egg to adult (Enfield, unpubl. obs.). Enfield (1977) and Wood (1977) discuss nomenclatural problems surrounding this species. Setae vary between western and eastern Canadian populations. In Alberta abdominal seta 7-II is multiple (more than four hairs), whereas eastern specimens have seta 7-II with three or fewer hairs (Wood, pers. comm.). Aedes ( Ochlerotatus ) fitchii (Felt and Young) (65 collections) This common species in 1976 was less so in 1977 and 1978, but was widely distributed in open habitats. As might be expected of a widely distributed species, it was rarely found alone (three collections), but was found at one time or another with 22 other species, frequently with Ae. cataphylla. It was found primarily in the spring, but also in the summer on two occasions in 1977. The larvae were among the slowest developers, many being in the second stage when other species in the same water body were in the fourth. Larvae were easy to rear. Aedes ( Ochlerotatus ) implicatus Vockeroth (44 collections) Another early species which was, however, found in a few locations in the summer of 1977. It has a wide tolerance of habitat conditions and is clearly not restricted to montane, forested regions as stated by Carpenter and LaCasse (1955). Found on seven occasions alone, it occurred at other times with 23 other species. The larvae were easy to rear in the laboratory. Aedes ( Ochlerotatus ) increpitus Dyar (17 collections) Mosquito (Diptera, Culicidae) larvae in Southern Alberta 157 Another species which was much commoner in 1976 than in the later years. Larvae prefer open ponds in the foothills in the southern half of the region, and were found in enormous numbers at Cayley in 1976. Although rare in mountains and prairie, this species is included in the ubiquitous category rather than creating a separate category for it. All records were from spring sampling except for a sole mountain record which was taken in the summer of 1977. It was rarely (two collections) found in the absence of other species. The larvae, like those of Ae. mercurator to which they are rather similar, are distinctly yellowish in life, unlike those of most black-legged species. Aedes ( Aedimorphus ) vexans (Meigen) (40 collections) Aedes vexans is normally one of the commonest species in this area. However in the dry years of this survey it was found on only two occasions in 1976 and 2 occasions in 1978. Normally thought of as a typical summer species, all of the 1976 and 1978 records were from spring sampling, and all of the records from the northern part of the region in 1977 were also taken in the spring. Aedes vexans normally requires water temperatures above 8°C and a period of prior conditioning for successful hatching (Horsfall, 1956). However in 1977 eggs were hatching at much lower water temperatures. Our records show that larvae are found usually in open, grass-covered ponds throughout the region. The larvae were found associated with 21 other species, and on only seven occasions were they the only Aedes larvae taken at a site. The larvae are quite easy to rear in the laboratory. Group 3 - Non-prairie Species Aedes (Ochlerotatus) excrucians (Walker) (40 collections) This species was widely distributed through the mountains, foothills and parkland region. Larvae were primarily in open, grass-covered ponds in forested areas at higher elevations. Individuals develop slowly, like those of Ae. fitchii, to which they are very similar in many respects. On only five occasions was Ae. excrucians the only species taken at a site; on other occasions it was associated with one or more of 20 other species. It was found only during spring, except for the single 1977 record. Aedes ( Ochlerotatus ) mercurator Dyar (40 collections) Except for two mountain samples in 1978 this species was taken during spring. Open, snow-melt pools in non-forested areas are preferred. Like Ae. euedes, Ae. mercurator larvae survive well in very cold water (Enfield and Pritchard, 1977). The confusion with Ae. stimulans in Alberta is discussed by Enfield (1977). Many larvae were associated with those of Ae. cataphylla. Group 4 - Parkland and Prairie Species Aedes ( Ochlerotatus ) campestris Dyar and Knab (76 collections) Quaest. Ent., 1981, 17 158 Scholefield, Pritchard and Enfield One of the commonest species in the study area, Ae. campestris was found in all 4 zones but was characteristic of open country and much commoner in the parkland and prairie than in the mountains and foothills. Most larvae were in open pools, without marginal vegetation. Rempel (1950) reports larvae from pools rich in organic matter with a pH on the alkaline side, but since all of the locations from which larvae were taken in this study were alkaline the latter criterion is not particularly helpful. Larvae were found during both spring and summer sampling. The species was found alone on six occasions, but at other times with 17 other species, frequently including Ae. dorsalis and Ae. cataphylla . The larvae were notably paler than those of other species and were very easy to rear in the laboratory, being able to withstand considerable crowding and fouling of the water. Development was rapid, except under crowded conditions when the rate slowed, although larvae survived well for long periods. Aedes (Ochlerotatus) dorsalis (Meigen) (66 collections) This species was one of the commonest in the survey and was found in all four zones, although it was taken at only one location in the mountains and at only a few in the foothills. Most larvae were in open pools without marginal vegetation. It is normally thought of as a summer species and was indeed uncommon in the spring of 1976 and was commoner during summer of 1977 than in the spring of that year. However, in 1978 it was much commoner during spring than in the summer. This was the only Aedes species present at 10 sampling sites, and was found with one or more of 16 species on other occasions. Associated species were Ae. vexans, Ae. cataphylla, and Ae. spencerii. Aedes ( Ochlerotatus ) flavescens (Muller) (73 collections) This was one of the most common species encountered in the survey. It was widely distributed over the parkland and prairie regions and was also taken in the foothills zone, although rarely. Like so many other spring species, larvae were taken during summer of 1977, presumably reflecting lack of snow-melt pools in that year. Larvae were generally at an earlier stage than those of other species in the same water body and this slow rate of development could be a disadvantage in years when pools dry up quickly. Because of its geographical distribution most larvae were in open habitats without marginal vegetation. Only twice found in the absence of other species, Ae. flavescens often occurred with Ae. campestris. According to Carpenter and LaCasse (1955), adult females may survive until late in the season, and in July 1977 females were biting during the day in the southern part of the region. Aedes ( Ochlerotatus ) melanimon Dyar (13 collections) This species was only found in the southern part of the parkland and prairie zones during the summer of 1977. Most larvae were in open, grass-covered ponds, without marginal vegetation and frequently fed by water from more permanent sources. First recorded in Canada from Brooks, Alberta by Burgess (1957), this species was taken in only 13 sites in this survey. It occurred with 9 other species, including Ae. vexans on 12 occasions and Ae. dorsalis on 11 occasions. Aedes ( Ochlerotatus ) spencerii (Theobald) (71 collections) Like Aedes flavescens, this species occurs occasionally in the foothills zone, but it is so widespread over the parkland and prairie that it is best considered in this latter group. It was one of the commoner species in this survey, even though it was only taken three times in 1977. Mosquito (Diptera, Culicidae) larvae in Southern Alberta 159 In that year, it presumably suffered, as did many other spring species, from the lack of suitable snow-melt pools in the parkland and prairie zones. Aedes spencerii was found alone more than any other species (27 times), and had a relatively low number of associated species (13) for such a common mosquito. In addition, there were usually few larvae of any species in the pools in which it was found. Species of Uncertain Distribution Aedes (Aedes) cinereus Meigen (11 collections) The few records for this species are from the mountains, foothills and western edge of the parkland, where larvae were found in pools fringed by poplars and willows. During this survey, it was found only during spring, but during a population study in 1974 eggs hatched both in the spring and later in the year along >yith Ae. vexans (Enfield and Pritchard, 1977). Aedes (Ochlerotatus) diantaeus Howard, Dyar and Knab (4 collections) This species is, according to Rempel (1950), rare and local and found only in forested regions of Canada. Most larvae were collected from open, grass-covered pools with marginal evergreens and willow, in water that was not completely stagnant. Carpenter and LaCasse (1955) report the larvae from pools of melting snow, and both collections in 1978 were made during spring. In 1977, however, larvae were taken in rainwater pools during the summer. These are the first records for southern Alberta outside of the National Parks (recently reported from Banff National Park; Wood, pers. comm.). Aedes diantaeus was reported by Happold (1965) from Flatbush, 150 km north of Edmonton. Aedes ( Ochlerotatus ) impiger (Walker) (4 collections) Larvae were found only three times: twice in the mountains in 1977 and once in the foothills in 1978. Open, grass-covered, non-stagnant pools with marginal evergreens were the typical habitat. A single adult female was taken in the parkland south of Lethbridge. Aedes (Ochlerotatus) intrudens Dyar (3 collections) A rare species whose larvae are found only in shaded, snow-melt pools with marginal willows and poplars in the foothills and mountains. Aedes (Ochlerotatus) nigromaculis (Ludlow) (1 collection) The rarest species of Aedes encountered in the survey, Ae. nigromaculis was found only once in an open, grass-covered pool formed from irrigation seepage into a grassy ditch, north-west of Lethbridge during the summer collecting period in 1977. According to Rempel (1950) it is a species of the open prairies and has been recorded previously from Lethbridge and Medicine Hat. Perhaps the record for Beaverlodge (55° 13’N, 119° 26’W) (Strickland, 1938) is in error. Aedes ( Ochlerotatus ) sticticus (Meigen) (6 collections) Quaest. Em.. 1981, 17 160 Scholefield, Pritchard and Enfield Enfield (1977), having seen larvae of both Ae. implicatus and Ae. mercurator misidentified as Ae. sticticus , suggested that the record by Pucat (1965) needed to be verified. Five samples taken in 1977 and one in 1978, in late May or June, certified its presence, although it is apparently not common. Larvae were taken from open or shaded grass-covered pools, generally with marginal willows. The species was not collected from the mountain zone. Aedes (Ochlerotatus) trichurus (Dyar) (3 collections) This species was not common, with larvae being found only in a few willow-fringed pools in the foothills and mountains. The larvae developed quickly, but were not easy to rear. They are large, and like those of species which have banded-legged adults, have a distinctly yellowish appearance in life. Genera other than Aedes The following species in the genera Anopheles , Culiseta and Culex overwinter as adults. In the spring they are the first mosquitoes to be seen on the wing, especially the genus Culiseta , whose adults are quite large. Adults appear as early as April if the weather is warm, but the larvae tend to appear later in the season (late springearly summer) after the appearance of the larvae of aedine species. The females lay their eggs on the water surface and there is no conditioning of the eggs, which hatch within a few days. Anopheles ( Anopheles ) ear lei Vargas (2 collections) There are no previous records of this species occurring in southern Alberta. Specimens were taken from a site near Twin Butte in 1977 and another site in Waterton National Park on the Red Rock Canyon Road in 1978. The abundance of this species is usually quite low and local and no serious outbreaks have been recorded. Culex (Culex) tarsalis Coquillett (10 collections) This species is represented across the entire southern half of the province, and is especially prevalent in the irrigated areas. Western equine encephalitis is associated with this species (Shemanchuk 1969), and if weather and irrigation conditions are correct, it could prove to be a problem. In the southern part of the study area it was often found in shallow pools in pasture fields flooded by irrigation water as well as roadside ditches. Culex (Neoculex) territans Walker (4 collections) This is the first record of Culex territans south of Edmonton. We found it locally only in the foothills. Most larvae were associated with bodies of water that contained a lot of vegetation. Culiseta ( Culiseta ) alaskaensis (Ludlow) (17 collections) Strickland (1938) reports this species from Banff National Park, the Nordegg area, and the Calgary/Ft. Macleod area. It would seem to prefer higher elevations, being found in our survey only in the mountains and foothills except for two records west of Calgary in the parkland. This species was on the wing in early May in Jasper National Park in 1978, and was a persistent biter. Mosquito (Diptera, Culicidae) larvae in Southern Alberta 161 Culiseta ( Culiseta ) incidens (Thomson) (2 collections) This species was very rare in the study area having been found only once at Rocky Mountain House in 1977 and once northeast of Banff in 1978. Culiseta ( Culiseta ) inornata (Williston) (47 collections) This is the most widespread Culiseta in southern Alberta and one of the most troublesome species in the Lethbridge area during the summer, when, regardless of weather conditions, irrigation produces ideal breeding grounds. Culiseta inornata is a troublesome biter of both man and livestock as well as wild ungulates, and has been associated with western equine encephalitis (Shemanchuk 1969). Culiseta (Culicella) morsitans (Theobald) (1 collection) Only one specimen of this species was taken during the study period, being taken south of the Jasper townsite during the spring of 1977. According to Rempel (1950) it is very rare in Western Canada, but has been previously reported from Alberta (Dyar, 1928; Wada, unpubl.). DISCUSSION During the three years of this study a number of changes have been made to published lists (e.g. Pucat, 1965) of mosquitoes in southern Alberta. Enfield (1977) revealed the presence of one species not previously recorded from Canada namely Aedes schizopinax . This species in fact, has a wide distribution in the southern part of the mountains and foothills. It had previously been found in Calgary by Robins (1972, unpubl.) and Scholefield (1973, unpubl.), but not officially recorded. Aedes euedes has a very wide distribution across the entire survey area (Fig. 3). Previously identified as Ae. barri or Ae. riparius, its naming as Ae. euedes has now been properly established (Wood, 1977). Aedes stimulans, which, according to records, had a wide distribution in Alberta, apparently does not occur in the province. All specimens identified as such, appear to be Aedes mercurator (Enfield, 1977; Wood, 1977), which is widely distributed in the mountains, foothills, and parkland of the study area (Fig. 3). Aedes increpitus is now recorded from southern Alberta, being first found in the Calgary area by Scholefield (1973, unpubl.). It was not found north of Calgary, however. Enfield (1977) indicated that the presence of this species in northern Alberta is suspect due to the fact that all the records he had seen from that area were actually Ae. mercurator. Another new record for southern Alberta is Aedes implicatus , being first recorded by Enfield (1977). It ranges across the entire southern half of Alberta, although it is local in the prairie. Northern records for Aedes hexodontus have been published by Pucat (1965), Graham (1969), and Tawfik and Gooding (1970). It too was first recorded in southern Alberta during the first year of this present study (Enfield, 1977). Its distribution is restricted to the mountains and foothills where it is quite common. The ranges of six other species ( Ae. canadensis, Ae. pionips, Ae. punctor, Ae. communis, Ae. trichurus and Ae. pullatus) have been somewhat extended. Previous reports in the literature listed Banff as the only site in southern Alberta (Strickland, 1938; Rempel, 1950). All are more or less restricted to the mountains and foothills, areas which have not been Quaest. Ent., 1981, 17 162 Scholefield, Pritchard and Enfield previously extensively collected. impiger diantaeus ci nereus i ntrudens melanimon f lavescens spenceri i campestri s dorsalis mercurator excrucians i ncrepi tus vexans impl icatus fi tchi i euedes cataphylla canadensi s pionips schizopinax punctor pullatus hexodontus communi s "i — r 8 pH 10 fig. 4. The range of pH of the sites from which larvae of 24 species of Aedes were collected. Mosquito (Diptera, Culicidae) larvae in Southern Alberta 163 1 1 1 1 r 1 1 r — i 1 0 50 100 MICROMHOS X10 Fig. 5. The specific conductance (mean ± 95% confidence limits) of the sites from which larvae of 15 species of Aedes were collected. Quaest. Ent., 1981, 17 164 Scholefield, Pritchard and Enfield Factors affecting mosquito distribution in southern Alberta are largely unknown. Most species encountered in this study have wide distributions, although 15 species were collected only from the mountain - foothills region and six were more or less restricted to the parkland - prairie. On a finer scale, habitat distinctions between species are impossible to make on the basis of information collected in this survey. Of the 20 species which were collected on at least 10 occasions, all but two were associated with 13-20 other species at one time or another, and the other two species had nine associated species. Clearly, with such overlap there is no such thing as a unique habitat for each species. This problem was further exposed when characteristics of the individual sites were considered. There were few obvious departures from the typical set of habitat conditions for the area, nor were there any obvious differences in pH (Fig. 4) or specific conductance (Fig. 5) of the water bodies inhabited by the different species. Water in the mountains and foothills has lower conductivity than water in the plains, and species restricted to these areas are therefore found in water with lower conductance values. This does not, of course, imply that conductance of the water is the reason for restriction of these species to those zones. Similarly, water bodies in the mountains and foothills are more likely to be shaded than are those in the parkland and prairies. Maire and co-workers (e.g. Maire and Aubin, 1976; Maire et al., 1978; Mailhot and Maire, 1978) have related the distribution of mosquito larvae to vegetation types in Quebec. However, the analysis is rarely clear-cut, even with only one half of the number of Aedes species that are present in southern Alberta. For example, Maire (1977) proposes four associations based on abundance of four species of Aedes and two of Culiseta. However, except for Ae. decticus, which is found only in open sites, all species occur in all habitats, and three species are abundant in more than one habitat type. A further difficulty in judging habitats as suitable or not for particular species is that relative species abundance can change quite markedly from year to year. There was a marked decline in number of sites at which the 18 most common species were taken from 1976 (355 sites) to 1977 (226 sites), even though there was no summer sample nor were the mountains well sampled in 1976. There was only slight recovery in 1978 (273 sites). Particular species worth noting are Ae. campestris, Ae. cataphylla, Ae. euedes , Ae. fitchii , Ae. flavescens, and Ae. mercurator which were all common in 1976, but much reduced in numbers in later years. Aedes euedes and Aedes mercurator are especially noticeable since together they were so abundant at a pond just northwest of Calgary in 1975 that they provided excellent material for a population study (Enfield and Pritchard, 1977). Also noticeable are Ae. vexans , normally thought of as the commonest species in this area, but which only achieved any level of relative abundance in 1977, and Ae. spencerii , which was uncommon in 1977, but the commonest species in 1978. Records were not kept of individual pools from year to year, except for a pool near the Columbia Ice Fields which had two hatches of Ae. pullatus in 1977, but none in 1978. In the latter year, Ae. communis was present instead. Overall changes in abundance of mosquitoes are clearly related to number of pools formed by snow-melt or rainfall. Availability of snow-melt pools depends upon amount of snow-fall, pattern of Chinook winds during winter, and speed at which snow disappears during spring. During this survey, there appeared to be fewer snow-melt pools in 1977, especially in the parkland and prairie, than in other years. Rain-filled pools depend on the pattern of precipitation during spring and summer, and on the level of the water table. There were fewer such water bodies in 1978 than in 1977. Overall, the three years of this study coincided with a period of relative drought which was inimical to development of mosquito populations. Our Mosquito (Diptera, Culicidae) larvae in Southern Alberta 165 experience with Aedes vexans exemplifies this trend. In 1972 (Robins, unpubl.) and 1973 (Scholefield, unpubl.), Ae. vexans was abundant in the Calgary area. In August of 1974 at an experimental site outside Calgary, larvae of Ae. vexans were abundant, but the pond dried up before larvae had completed development (Enfield and Pritchard, 1977). In 1975, the pond flooded at snow-melt, was dry by early June and remained so for the rest of the year and no Ae. vexans eggs hatched. At our experimental sites in N.W. Calgary it was necessary to artificially flood pond beds in 1976 and 1977 in order to have populations with which to work. In 1978 these ponds flooded naturally in April, but dried up permanently for the year at the beginning of July. No Ae. vexans larvae appeared. These trends applied generally over the whole study area. Related to the pattern of precipitation is the distinction between “spring” and “summer” species, a concept which is by no means as tidy as many hold it to be. Spring species are those whose eggs generally hatch on the first flooding of the year, this normally being the result of snow-melt, and an egg diapause restricts the number of generations per year to one. Exceptions to this latter statement may be Ae. pullatus and Ae. cinereus (see above), and perhaps other species (see Magnarelli, 1977). Montane and foothills species generally belong to this group, as might be expected since there will be an abundance of snow-melt pools in these regions. So-called “spring species” apparently hatch in the first flooding whether warm or cold and this accounts for their appearance in summer samples, especially in 1977 when there was a dearth of snow-melt pools. However temperature requirements for hatching in these species appear not to have been well studied. Hatching of eggs of summer species such as Ae. vexans and Ae. dorsalis is complicated (Horsfall, 1956). The basic pattern is that eggs that have completed embryogeny hatch only after a period of drying, flooding with water above 8°C and a lowering of oxygen concentration. Hovyever, there is variability between populations in response to these factors and even given these conditions many eggs do not hatch. This erratic hatching has been viewed by some authors as an inherent safeguard against unpredictable water conditions, but Gillett et al., (1977) have suggested that in Ae. aegypti this erratic hatching is due to the grazing of oxygen-consuming bacteria from egg surfaces by the first larvae that hatch. Egg diapause in Ae. vexans is also complicated (Taylor, pers. comm.). Again, our experience with Ae. vexans is illustrative of the complexity of the problem. In our population studies in 1974 (Enfield and Pritchard, 1977), eggs of Ae. vexans behaved according to the above formula; i.e. no eggs hatched when flooded with water at 6°C and four subsequent floodings at 20°C were required in order to hatch all the eggs. Few, if any eggs, were laid in our experimental field sites in 1974 or 1975 and so there were few eggs available in the ground in 1976 or 1977. When they were hatched artificially by flooding with water above the threshold for hatching in the laboratory in these years, the hatch was complete with one flooding, perhaps because of the low density and supporting the hypothesis of Gillett et al., (1977). However, more interestingly, in the survey, Ae. vexans larvae were generally taken in spring sampling rather than summer sampling and the evidence suggests that eggs were hatching in much colder water than the accepted threshold. Eggs of Ae. vexans collected from soil cores in 1977 were in fact hatched in the laboratory at temperatures below 8°C. The same remarks apply to Ae. dorsalis in 1978. In view of the confused literature on this subject (e.g. Horsfall et al., 1973) and in light of the above observations, a renewed study of the hatching requirements of Aedes eggs would seem to be in order. Such a study might go a long way towards explaining the distribution and Quaest. Ent., 1981, 17 166 Scholefield, Pritchard and Enfield abundance of different species from year to year. ACKNOWLEDGEMENTS It is with much gratitude that we acknowledge the encouragement, advice and assistance given to us by Dr. D.M. Wood of the Biosystematics Research Institute, Agriculture Canada, Ottawa. Dr. R.H. Gooding, Mr. Will Johnson, and members of the Pesticide Chemicals Branch, Pollution Control of Alberta Environment who assisted in the collection of larvae at various times. We are especially grateful to Mr. Jock McIntosh of Alberta Environment in this last regard. We also thank Dr. R.A. Ellis of the City of Winnipeg for his comments about a draft of this report. Funds for this project were generously provided by the Pesticide Chemicals Branch, Pollution Control Division Alberta Environment. REFERENCES Burgess, L. 1957. Note on Aedes melanimon Dyar, a mosquito new to Canada. Canadian Entomologist. 89:352. Carpenter, S.J. and W.J. LaCasse. 1955. Mosquitoes of North America (North of Mexico). University of California Press. Danilov, V.N. 1974. Restoration of the name Aedes (O.) mercurator Dyar to the mosquito known in the U.S.S.R. as Aedes riparius ater Gutsevich (Diptera: Culicidae) (in Russian). Parazitologiya 8(4):322-328. Dyar, H.G. 1920. The mosquitoes of British Columbia and Yukon Territory, Canada (Diptera, Culicidae). Insecutor Inscitiae Menstruus 8:1-27. Dyar, H.G. 1928. The mosquitoes of the Americas. Carnegie Institute, Washington, Publication No. 387:1-616. Ellis, R.A. and R.A. Brust. 1973. Sibling species delimitation in the Aedes communis (DeGeer) aggregate (Diptera: Culicidae). Canadian Journal of Zoology. 51:915-959. Enfield, M.A. 1977. Additions and corrections to the records of Aedes mosquitoes in Alberta. Mosquito News 37:82-85. Enfield, M.A. and G. Pritchard. 1977. Estimates of population size and survival of immature stages of four species of Aedes (Diptera: Culicidae) in a temporary pond. Canadian Entomologist 109:1425-1434. Gillett, J.D., E.A. Roman and V. Phillips. 1977. Erratic hatching in Aedes eggs: a new interpretation. Proceedings of the Royal Society of London B 196:223-232. Gjullin, C.M. and G.H. Eddy. 1972. The mosquitoes of the Northwestern United States. U.S. Department of Agriculture Technical Bulletin No. 1447. Ill pp. Government of Alberta and the University of Alberta. 1969. Atlas of Alberta. University of Alberta Press. Graham, P. 1969. Observations on the biology of the adult female mosquitoes (Diptera: Culicidae) at George Lake, Alberta, Canada. Quaestiones Entomologicae 5:309-339. Happold, D.C.D. 1965. Mosquito ecology in central Alberta. I. The environment, the species, and studies of the larvae. Canadian Journal of Zoology 43:795-819. Horsfall, W.R. 1956. Eggs of floodwater mosquitoes (Diptera: Culicidae). III. Conditioning Mosquito (Diptera, Culicidae) larvae in Southern Alberta 167 and hatching of Aedes vexans . Annals of the Entomological Society of America 49:66-71. Horsfall, W.R., H.W. Fowler Jr., L.J. Moretti and J.R. Larsen. 1973. Bionomics and Embryology of the inland floodwater mosquito Aedes vexans . University of Illinois Press. 211 pp. Magnarelli, L.A. 1977. Seasonal occurrence and parity of Aedes canadensis (Diptera: Culicidae) in New York State, U.S.A. Journal of Medical Entomology 13:741-745. Mailhot, Y. and A. Maire. 1978. Caracterisation ecologique des milieux humides a larves de moustiques (Culicides) de la region subarctique continental de’Opinaca (territoire de la Baie de James. Quebec). Canadian Journal of Zoology 56:2377-2387. Maire, A. 1977. Identification des biotopes a larves de moustiques des tourbieres de la Basse-Mauricie (Quebec Meridional). Naturaliste Canadien 104:429-440. Maire, A. and A. Aubin. 1976. Inventaire et classification ecologiques des biotopes a larves de moustiques (Culicides) de la region de Radison (territoire de la Baie de James, Quebec). Canadian Journal of Zoology 54:1979-1991. Maire, A., C. Tessier and L.Picard. 1978. Analyse ecologique des populations larvaires de moustiques (Diptera: Culicidae) des zones Riveraines du Fleuve SaintLaurent, Quebec. Naturaliste Canadien 105:225-241. Mclver, S. 1973. Summary - Behaviour and Ecology of populations. In : Hudson, A. (Editor). Biting Fly Control and Environmental Quality. Defence Research Board, Ottawa. DR 217. pp. 150-152. Nielsen, L.T. 1961. Aedes schizopinax Dyar in the Western United States. California Mosquito Control Association 29:21-24. Pucat, A. 1965. List of mosquito records from Alberta. Mosquito News 25:300-302. Rempel, J.G. 1950. A guide to the mosquito larvae of Western Canada. Canadian Journal of Research D 28:207-247. Scholefield, P.J., M.A. Enfield, and G. Pritchard. 1979. Identification and distribution of the aedine mosquitoes of southern Alberta. Alberta Environment Report. 1 14 pp. Shemanchuk, J.A. 1969. Epidemiology of Western encephalitis in Alberta: response of natural populations of mosquitoes to avian hosts. Journal of Medical Entomology 6:269-275. Strickland, E.H. 1938. An annotated list of the Diptera (flies) of Alberta. Canadian Journal of Research D 16:175-219. Tawfik, M.S. and R.H. Gooding. 1970. Observations on mosquitoes during 1969 control operations at Edmonton, Alberta. Quaestiones Entomologicae 6:307-310. Wada, Y. 1965. Population studies on Edmonton mosquitoes. Quaestiones Entomologicae 1:187-222. Wood, D.M. 1977. Notes on the identities of some common nearctic Aedes mosquitoes. Mosquito News 37:71-81. Wood, D.M., P.T. Dang, and R.A. Ellis. 1979. The Insects and Arachnids of Canada. Part 6. The Mosquitoes of Canada. Agriculture Canada. 390 pp. Quaest. Ent., 1981, 17 Patasson luna (Hymenoptera: Mymaridae) NOTE RECOVERY OF PATASSON LUNA (HYMENOPTERA: MYMARIDAE) A PARASITE OF THE ALFALFA WEEVIL, HYPERA POSTICA (COLEOPTERA: CURCULIONIDAE), IN ALBERTA The alfalfa weevil, Hypera postica (Gyll.), was introduced into the United States near Salt Lake City, Utah in 1904 and by 1907 had become a serious pest in Salt Lake County (Titus, 1910). In southeastern Alberta, this pest was first located in alfalfa fields in the valley of the Milk River in June 1954. No egg parasites were recorded in studies of the biology and distrubution of H. postica from 1955 to 1958 (Hobbs et al., 1959). However, in 1978, during studies of the weevil in the Brooks area of Alberta, several specimens of a parasite were reared from alfalfa weevil eggs. Adults sent to the Biosystematics Research Institute, Agriculture Canada, Ottawa, were tentatively identified by Dr. Carl M. Yoshimoto and later confirmed by Mr. M. Schauff of the University of Maryland, College Park, Maryland, to be Patasson luna (Girault). The characters used to separate this species from a very closely related species Anaphes pratensis Foerster, can only be seen under a microscope and only in the females. The two species can be easily mistaken for each other. This is apparently what happened in several releases. During the years 1911-1913 and 1925-1928, P. luna that had been collected in Italy were released in Utah in an attempt to establish a biological control agent for control of the alfalfa weevil (Chamberlain, 1924a; Clausen, 1956). Patasson luna was probably confused with A. pratensis in these releases, because, in 1926, A. pratensis was recovered from several fields near Salt Lake City, Utah (Hamlin et al., 1949), but P. luna was not recovered at this time. Patasson luna from France was introduced into alfalfa fields in California around 1933 (Essig and Michelbacher, 1933) but has not been recovered since its release (K. S. Hagen, per. comm.). However, P. luna was collected in Idaho in 1921 (M. Schauff, per. comm. 1980; due to their condition, only a tentative identification of P. luna has been given to these specimens), in Utah by D.W. Davies (Dysart and Day, 1976) in 1973, and in Alberta in 1978. Patasson luna has been reared from alfalfa weevil eggs in the eastern United States from Delaware, Indiana, Illinois, Maryland, Massachusetts, New Jersey, New York, Ohio, Pennsylvania, and West Virginia. In eastern Canada, it has been collected in Ontario and Quebec. The exact method of establishment in eastern North America has never been documented. It seems probable that P. luna was introduced into this area with a shipment of A. pratensis sent to Indiana for control of Hypera nigrirostris (F.) in 1928 (Christie, 1928, p. 43) and that it subsequently spread throughout most of the eastern United States. At Brooks, Alberta, most seed alfalfa is grown on irrigated land surrounded by mixed grains to the north and west, and by mixed grain or grasslands to the south and east. No alfalfa hay is imported into these irrigated areas. Apparently, P. luna entered by migrating over mountains from Idaho, or possibly from the southeast from Montana along the Missouri and Milk River systems and then across the grainland and grassland. In southern France, A. pratensis and Patasson spp. infest eggs of Sitona sp. and Hypera sp. in alfalfa crops (Aechlimann, 1977). Chamberlain (1924 b) and Hamlin et al., (1949) Quaest. Ent., 1981, 17 170 Schaber showed that P. luna and A. pratensis parasitize eggs of the clover leaf weevil, Hypera punctata (Fabr.) in the United States. When clover leaf weevils become scarce, both parasites will use eggs of H. postica for oviposition. The egg of H. punctata is much larger than H. postica and, according to Chamberlain (1924 b), P. luna places two eggs in each egg of H. postica. Patasson luna may also use a wide range of Sitona and other Hypera spp. as their primary host and as these hosts become scarce they migrate and infest H. postica eggs as they do in Europe. The overwintering method or stage of this parasite is not known in Alberta, but, since a few eggs of the clover leaf weevil persist through the winter (Clausen, 1956), it seems possible that it is in this host that P. luna survives the winter. During the spring of 1979, no specimens of P. luna were recovered from eggs of the alfalfa weevil. Therefore, the species is considered at present to be of limited importance in the control of the alfalfa weevil in Alberta. REFERENCES Aeschlimann, J.P. 1977. Notes on Patasson lameerei (Hym: Mymaridae), an egg parasite of Sitona spp. (Col: Curculionidae) in the Mediterranean Region. Entomophaga 22:111-114. Chamberlain, T.R. 1924a. Introduction of parasites of the alfalfa weevil into the United States. United States Department of Agriculture Circular. No. 301. 9 pp. Chamberlain, T.R. 1924b. Studies of the parasites of the alfalfa weevil in Europe. Journal of Economic Entomology 17: 623-632. Christie, G.I. 1928. Forty-first annual report of the Purdue University Agricultural Experiment Station. Purdue University, Lafayette, Indiana. Clausen, C.P. 1956. Biological control of insect pests in the continental United States. United States Department of Agriculture Technical Bulletin. No. 1139: 116. Dysart, R.J., and W.H. Day. 1976. Release and recovery of introduced parasites of the alfalfa weevil in eastern North America. United States Department of Agriculture Research Service, Production Research Report. No. 167: 2. Essig, E.O., and A.E. Michelbacher. 1933. The alfalfa weevil. California Agriculture Experiment Station Bulletin. No. 567. 99 pp. Hamlin, J.C., F.V. Lieberman, R.W. Bunn, W.C. McDuffe, R.C. Newton, and L.J. Jones. 1949. Field studies of the alfalfa weevil and its enviironment. United States Department of Agriculture Technical Bulletin. No. 975. 84 pp. Hobbs, G.A., W.O. Nummi, and J. Virostek. 1959. History of the alfalfa weevil, Hypera postica (Gyll.) (Coleoptera: Curculionidae), in Alberta. The Canadian Entomologist 91: 562-565. Titus, E.G. 1910. The alfalfa weevil. Utah Agriculture Experiment Station Bulletin. No. 110: 19-72. Burton D. Schaber Research Station, Agriculture Canada Lethbridge, Alberta T1J 4B1 Book reviews 171 BOOK REVIEWS Larsson, Sven Gisle. 1978. Baltic Amber - a Palaeobiological Study. Entomograph, vol. 1, 192 pp., 62 text-figures, 12 monochrome plates. Scandinavian Science Press Ltd., Klampenborg, Denmark. Price: 120 Danish Kroner, $25.32 Canadian. One of the reasons that man is so attracted by amber is because it is the earth’s most stable and therefore oldest natural biological product. As ambers go, European or Baltic amber, the subject of Larsson’s study, is relatively young (35-40 m. yrs. as opposed to 75-80 m. yrs. for Canadian amber). Nevertheless it has been lying around the shores of the present Baltic Sea at least 15 times longer than the fossils of the earliest known man in Kenya, and it is still very much the same as it was when it was first produced by the extinct pine, Pinites succinitera Conwentz. Larsson’s attractive book is the first English-language synthesis of biological information on Baltic amber. It is also the first volume in the Scandinavian Science Press new series, Entomograph, designed especially for large important entomological papers such as this. This study builds upon, and in some respects supersedes, Kjell Ander’s (1942), Die Insekten Fauna des Baltischen Bernsteins nebst damit Verkniipten Zoogeographischen Problemen, and Andolph Bachofen-Echt’s (1949), Der Bernstein und sein Einschlusse. The author’s theme (p. 7) is the rich content of fossils of terrestrial arthropods and plants that are contained in Baltic amber, and the excellent opportunities these fossils provide for learning the early history of plant and animal life. Although not stated, the main purpose of the book appears to be to outline the problems involved in interpreting the amber biota. It is also apparent throughout the book that a secondary purpose is to publicize the relatively large, newly amassed collection of Baltic .amber (8000 specimens), derived almost exclusively from Danish coasts, in the Zoological Museum, Copenhagen. The book brings to fruition Larsson’s long standing interests in Baltic amber, and culminates his efforts since 1949 to develop a permanent working collection of scientific specimens from Denmark. Being a Dane, he is thoroughly familiar with the geography of the parts of Europe where the amber was produced and deposited, and he brings first-hand knowledge to bear on those problems. He is a professional entomologist with close contacts with many other entomologicial specialists in the Zoological Museum, and this is another essential qualification for fulfillment of his purposes. In developing the geological and chemical framework provided for his presentation he was guided by Danish experts in these fields. So the book has a solid foundation. Larsson treated the subject in five main sections. Section 1 deals with the general characteristics of amber and resin, its stability and production and its chemical and physical properties. Section 2 is a treatise on the origins of the various ambers from the Baltic region. Section 3 treats the original flora of the amber territory. These three sections occupy 59 pages. Section 4 deals with the original fauna of the amber territory. This, of course, makes up most of the book (116 pp.), and because of the great predominance of insects in the Baltic amber inclusions, the book is primarily an entomological work. The section is divided along ecological lines into seven chapters entitled Plant Sucking Insects, Leaf-and Seed-Consumers, Gall Producers, Nectar Quaest. Ent., 1981, 17 172 Seekers, Insects Trapped While Resting, The Fauna of Moss and Bark, and The Hidden Fauna of Tree Trunks. The final section (Section 5) is a review describing the nature and conditions of the original amber forest as envisaged by the author. It also includes a list of the amber material in the Zoological Museum, Copenhagen. Each section has its own separate list of references (645 in all). An index restricted to the scientific names of genera and higher taxonomic categories is provided. The book contains 12 plates of original halftone photgraphs of inclusions (mostly insects) in amber and 62 line drawings, almost all of which are copied from the literature. The author succeds to some degree in restating most of the age-old problems involved in the study of the flora and fauna of Balltic amber, but few new facts or ideas are introduced. The book has real value as a reference text and it is very useful as a starting point for finding basic information on many aspects of amber and its inclusions. On the other hand, it has some weaknesses and faults, and it should not be accepted as ‘the last word’ on the subject. Coverage of the literature is incomplete. Different ideas and interpretations are frequently presented in such a way that the reader is left uncertain as to which he favors. For example, I am still not very confident about the biological status of Pinites succinifera or the extent of its role in the production of Baltic amber. Much of the discussion on the age, geographical origin and host origin of Baltic amber from diffetent parts of Europe is in the realm of speculation and the reader ends up with a feeling of uncertainty and confusion on many points. What is said in one place is sometimes contradicted in another. For example, it is stated on p. 103 that the only calyptrate fly known from amber belongs to Fabricius’ old and well known species, Fannia scalaris, and on p. 108 attention is drawn to the fact that one of the amber fossils of Cicindellidae is identical with the recent species Tetracha Carolina (L.). However, the final sentence of the Review reads “But the original Baltic amber fauna appears to be totally extinct.” The decision to divide the main section on the fauna of Baltic amber into seven chapters based on ecological considerations rather than on systematic grounds was unfortunate. This has led to a considerable amount of overlap and repetition, and it has created difficulty in locating certain data. For example, anyone seeking information on specific groups such as Coleoptera or Diptera must search through seven chapters. It would be better if all the arthropods were treated in a standard systematic sequence, and the ecological correlation were presented in tabular form. Similarly, the decision to break the bibliography into four sections was not a good one. In such a small book it would be preferable to consolidate all the references in one composite list. As it is, the reader must scan four lists if he wishes to locate all the papers cited for any given author, and of course there is a considerable amount of duplication in the different lists. The literary quality would have been improved by a good editing, but the book itself is very well produced. The paper, the printing, and the binding are excellent. In spite of its shortcomings it is the best single synthesis of paleobiological information available on Baltic amber, and anyone interested in the subject should have a copy. J.F. McAlpine Biosystematics Research Institute Agriculture Canada OTTAWA, Ontario, Canada K1A 0C6 Book reviews 173 GILLOTT, CEDRIC. 1980. Entomology. Plenum Press, New York and London, xviii + 729 pp. Hard cover $49.50; soft cover $22.50. Of the several books on general entomology published in recent years, this is one of the best. Its coverage of basic entomology is extensive, making it an excellent reference text. The author intends for the book to be used as a “text for senior undergraduates taking their first course in entomology”, and, indeed, it would be good if such students were given time to master the meterial in this text in one course. However, it covers too much ground for a single semester but could be appropriate for a two semester course. Section I (Chapters 1-10) on evolution and diversity , is perhaps a bit long (300 pages) for an introductory text. For example, Chapter 2 (Insect Diversity) contains 9 page discussion (a good one) on the origin and evolution of insect wings and a discussion of the Heslop-Harrison theory of pupal origin which concludes with the criticism that the theory lacks supporting evidence. The lengthy discussion of phylogenetic relationships of pterygotes is also excellent, but would be difficult reading for those unfamiliar with the terminology and having little experience with extant orders. Insect Evolution may have been a better title for Chapter 2 than Insect Diversity. Chapter 1 (Arthropod Evolution) discusses the three major theories of Arthropod evolution and comes down firmly in favor of the polyphyletic school. In spite of this, the major lineages suggested by this theory are not given equal coverage in the rest of the chapter. Some comments on the relative abundance of arachnids would have been helpful, in fact, arachnids and crustaceans could have been given equal space. Most of the drawings in this chapter are unlabelled. When compared with other chapters in the book, this one is somewhat sketchy, giving an unfortunate first impression. Chapter 3 (External Structure) presents a well balanced overview of its topic, and illustrated with relabelled drawings taken largely from Snodgrass’ works. Chapter 4 (Classification and Identification) wisely begins with clear definitions of terms such as systematics, classification, identification and taxonomy and comments on disagreements regarding these definitions. A short discussion of natural and artificial classification schemes follows. The chapter also includes an essay on the history of insect classification and a key to insect orders. All are well done. The remainder of Section I (Chapters 5-10) consists of a review of the insect orders, each covered in the following manner: order name, synonyms, common name and a short description. These are arranged to stand out clearly from the rest of the text, and are set in smaller type. Following each order’s description, are short sections on general structure, life history, phylogeny and classification, and literature (including several references for each order). Illustrations are appropriate and well done, and the phylogenetic trees given for most major orders are a helpful addition. These 5 chapters embody approximately one-third of the book. Section II (Chapters 11-18) covers aspects of anatomy and physiology in 150 pages. Chapter 11 is an excellent introduction to the insect integument, with highlights for this reviewer being the discussions of cuticular structure and color. Chapter 12 (Sensory Systems) is good as far as it goes. Some illustrations or photographs and a discussion of sensillar ultrastructure would have enhanced the presentation, as this area is by now well enough understood to make some generalizations. The physiology of chemoreception is also better known than this chapter suggests. Two helpful reviews in this regard, Kaissling (1971) and Hansen Quaest. Ent., 1981, 17 174 (1978) are not mentioned. Chapter 13 (Nervous and Chemical Integration) presents a concise and up to date overview of the endocrine system, but falls somewhat short in the part on nervous integration. The author attempts to summarize the physiology of neural integration, including a discussion of membrane physiology, in four pages with a single diagram showing a simple reflex circuit. The result is probably too general to be very useful. Extensive work has been done on insect walking which could have been used to illustrate current concepts of nervous integration in insects. Pearson et al. (1973) and Bowerman (1977) have written a reviews of this area. The discussion on pheromones could have been improved by including something on the importance of ‘minor’ pheromone components. The review by Seabrook (1978) could have helped here. Chapter 14 (Muscles and Locomotion), is generally well done, though a near classic work on insect flight by Nachtigall (1974) seems to have been overlooked. Chapter 15 presents an excellent overview of respiration in both terrestrial and aquatic insects. Chapter 16 (Food Uptake and Utilization) includes gut morphology, gut physiology and a discussion on metabolism, all are well done. Food selection and feeding, however, are treated quite superficially superficially. Insect host-plant relationships have been given considerable attention by ecologists, behaviourists and physiologists for several decades, and recent advances in these areas would form an excellent basis for a discussion of feeding behaviour. More serious is the lack of reference to Dethier (1976). In this book, “The Hungry Fly” extensive studies on feeding behaviour and related physiology covering more than 20 years are clearly summarized. Circulation, excretion and water balance are given good coverage in Chapters 17 and 18, while Chapters 19-21 present a thorough treatment of reproduction and development. The discussion of environmental and endocrine factors affecting female maturation and the figures comparing endocrine relationships in 4 insect species are excellent. Two chapters (22 and 23) on the abiotic and biotic environment give a good introduction to classical insect ecology. The final chapter, entitled Insects and Man, considers beneficial and harmful insects and stresses the importance of integrated pest management. Major problems with chemical control, particularly resistance to insecticides, are highlighted, and a good discussion of biological control with a table of suitable examples is presented. REFERENCES Bowerman, R.F. (1977) Control of arthropod walking. Comp. Biochem. Physiol. 56A: 231-247. Dethier, V.G. (1976) The Hungry Fly. Harvard University Press, Cambridge, London. Hansen. K. (1978) Insect chemoreception, In: Receptors and Recognition, Ser. B. Vol. 5. Taxis and Behaviour. G.L. Hazelbauer (ed.). Methuen. Kaissling, K.E. (1971) Insect olfaction. In: Handbook of Sensory Physiology. IV, Chemical Senses, Part 1, Olfaction. L.m. Beidler (ed.). Springer, Berlin, Hidelberg, New York. Nachtigall, W. (1974) Insects in Flight. George Allen and Unwin Ltd., London. Pearson, K.G., Fourtner, C.R. and Wong, R.K. (1973) Nervous control of walking in the Book reviews 175 cockroach. In: Control of Posture and Locomotion. R.B. Stein, K.G. Pearson, R.S. Smith, J.B. Redford (eds.). Plenum, New York. Seabrook, W.D. (1978) Neurobiological contributions to understanding insect pheromone systems. Ann. Rev. Entomol. 23: 471-485. B.K. Mitchell Department of Entomology The University of Alberta Edmonton, Alberta, Canada T6G 2E3 Quaest. Ent., 1981, 17 NOTICE PIFON, A NEW PERMANENT INTERNATIONAL FILE OF NATURALISTS In 1980 a new institution, the Oxycopis Pond Research Station was established in a wooded area in upstate New York. Among other projects the Institute compiles and maintains a coded information file of naturalists of the world, PIFON (acronym for Permanent International File of Naturalists). This international register includes well over 10,000 contemporary naturalists, and in addition a file of all persons listed in The Naturalist’s Directory ( International ) since the first edition in 1877. Thus PIFON is the world’s most complete listing of naturalists, past and present. This wealth of data is available for the use of any person registered in PIFON. The 44th edition of The Naturalists’ Directory ( International ) is being compiled from data in PIFON. All 10,000 persons were contacted in 1980 and all institutions in 1981 to urge registration in PIFON. Now the six 'parts of the new 44th edition (to be called volume 44) are being published. The six parts are: Part I: The Naturalists’ Directory of Insect Collectors and Identifiers ( International ); Part II: The Naturalists’ Directory of Plant Collectors and Identifiers ( International ); Part III: The Naturalists’ Directory of Bird Watchers ( International '); Part IV: The Naturalists’ Directory of Field Biologists ( International '); Part V: The Naturalists’ Directory of Rock, Mineral, and Fossil Collectors ( International '); Part VI: The Naturalists’ Directory ( International '): Subject, Geographical, and Name Index. Each part is complete in itself, but the last part is a general index and cross-reference for the other five parts for individuals and libraries using two or more parts. All persons registered in PIFON will receive a registration card showing their unique registration number. Among the other advantages of being registered is access to the unpublished data in the PIFON file. Registration is free. A charge of $1.00 to cover cost of photocopying and mailing is required of those using the file. Requests may be made at anytime by writing or by phone. To be registered in PIFON send the following information (please use block letters or type): 1) name (last or family name underlined; include Dr., Prof., Mr., Mrs., Ms., etc.); 2) mailing address (shortest form); 3) phone number (include area code); 4) group of main interest, area of specialization (e.g., family group name, etc.) and kind of interest (e.g., collecting, exchanging, systematics, ecology, etc.); 5) geographical area of greatest interest; 6) short statement giving interest details (not over 25 words); 7) if a taxonomist, list groups willing to identify for others; 8) signature and date. These data will be coded and entered into the file. Those already listed should send changes as they occur. No person is listed unless these data are supplied hy that person The right to edit all listings published in the Directory is reserved. The deadline for submitting entries for each edition is three months before publication. Parts will be issued every four months, resulting in a two year cycle for the entire volume. Regardless of the deadline dates, however, persons are urged to register immediately upon reading this note, thus making available their data to others using the PIFON file. 178 Notice The parts of the volume are sold by the publisher, Flora and Fauna Publications, by subscription only, either as single parts, or multiple parts, with a discount for subscriptions to the entire volume. Send registration and/or requests for further information to the compiler and ediitor: Dr. Ross H. Arnett, Jr., Oxycopis Pond Research Station, 90 Wallace Road, Kinderhook, NY 12106 USA. (Phone: (518) 758-7219).— R.H.A. Ouaest. Ent. 1981. 17 Publication of Quaestiones Entomologicae was started in 1965 as part of a memorial project for Professor E. H. Strickland, the founder of the Department of Entomology at the University of Alberta in Edmonton in 1922. It is intended to provide prompt relatively low-cost publication for comprehensive accounts of entomological research of greater than average length. However, shorter papers about insects in the Prairie Provinces of Canada are acceptable. Page charges are normally levied, the rate determined by printer’s charges. For information about current page charges, consult the Editor. Copy for all types of papers should conform to the Style Manual for Biological Journals, published by the American Institute of Biological Sciences, Second Edition, 1964, except that titles of periodicals should be given in full. For style of taxonomic papers, the Editor should be consulted. Two copies of a manuscript are requested. All manuscripts will be reviewed by referees. Abstracts are required: one in English, and one in another language, preferably French. Tables, including titles and footnotes, must not be more than 7 3/4 X 4 3/4 inches (19.7 X 12.1 cm). Copy for illustrations must accompany the manuscript, and be of such character as to give satisfactory reproduction at page size (less 1 /2 inch, or 1.2 cm on plates of full page size [7 3/4 X 5 inches, or 19.7 X 13.2 cm]). Reprints must be ordered when proofs are returned, and will be supplied at cost. Subscription rates are the same for institutions, libraries and individuals, $15.00 per volume of four issues, normally appearing at quarterly intervals; single issues $5.00. Back volumes and issues are available at the same cost. These prices supersede those previously indicated, and are subject to change as required by inflationary pressure on the value of money. Communications regarding subscriptions and exchanges should be addressed to the Subscription Manager, and regarding manuscripts to: The Editor, Quaestiones Entomologicae Department of Entomology University of Alberta Edmonton, Alberta, Canada T6G 2E3 Second Class Mail Registration Number 5222 He i QI3 Ba^ Quaest lones Entomolog icae A periodical record of entomological investigations# published at the Department of Entomology, University of Alberta, Edmonton, Canada. VOLUME 17 NUMBER 3-4 JULY-OCTOBER 1981 CONTENTS Lafontaine-Classification and phylogeny of the Euxoa detersa group (Lepidoptera: Noctuidae) 1 Oliver-Redescription and systematic placement of Oreadomyia albertae Kevan and Cutten-Ali-Khan (Diptera: Chironomidae 121 Corkum and Clifford-Function of caudal filaments and correlated structures in mayfly nymphs, with special reference to Baetis (Ephemeroptera) 129 Scholefield, Pritchard and Enfield-The distribution of mosquito (Diptera, Culicidae) larvae in Southern Alberta, 1976-1978 147 Note-Schaber-Recovery of Patasson luna (Hymenoptera: Mymaridae) a parasite of the alfalfa weevil, Hypera postica (Coleoptera: Curculionidae), in Alberta 169 Book Review-Larsson, Sven Gisle. 1978. Baltic Amber - a Palaeobiological Study. Entomograph, vol. 1 171 Book Review-Gillott, Cedric. 1980. Entomology 173 Hudson, J.E.-Studies on flight activity and control of mosquitoes in the Edmonton area, 1971-73 179 Fredeen, F.J.H.-Keys to the black flies (Simuliidae) of the Saskatchewan River in Saskatchewan 189 Doane, J.F.-Seasonal captures and diversity of ground beetles (Carabidae: Coleoptera) in a wheat field and its grassy borders in central Saskatchewan 211 Carter, A.-Aspects of the comparative ecology of populations of four Patrobus species (Coleoptera: Carabidae: Patrobini) at George Lake, Alberta 235 Roughley and Pengelly-Classification, phylogeny, and zoogeography of Hydaticus leach (Coleoptera: Dytiscidae) of North America 249 Book Review-Tschirnhaus, M von. 1981. Die Halm- und Minierfliegen im Grenzbereich Land-Meer der Nordsee 315 Editor’s Acknowledgements 317 Index 319 STUDIES ON FLIGHT ACTIVITY AND CONTROL OF MOSQUITOES IN THE EDMONTON AREA, 1971-73 J.E. Hudson Department of Entomology University of Alberta Edmonton, Alberta, Canada T6G 2E3 Quaestiones Entomologicae 17:179-188 1981 ABSTRACT The effects on mosquito activity of barrier strips of dichlorvos and propoxur sprayed on the vegetation in wooded ravines, were assessed by means of insect wind traps and by landing counts on human bait. Dichlorvos at 1.12kg a.i./ha reduced trap catches by 70% for one day after treatment; bite count data were inconclusive. Propoxur at 0.34kg a.i./ha reduced trap catches by 44% and bite counts by 100% for one day after treatment. From these results a continuation of the barrier strip method could not be recommended. Many other insects ( chiefly other Diptera) were caught in the traps, but there were no consistent reductions in catches in the treated strips. Catches of mosquitoes were consistently higher on the down-valley sides of the traps than on the up-valley sides, and catches were generally higher in traps in the ravines than in traps on the plain above. The effectiveness of malathion sprayed by helicopter was measured by kills of caged, wild-caught mosquitoes and by bite counts. Boom and nozzle application at 0.7kg a.i./ha killed only 27 % of the mosquitoes (mostly Aedes vexansj caged in the woods at Whitemud Creek. Ultra-Low-Volume application at 0.3kg a.i./ha killed only 8% of the caged mosquitoes (mostly Aedes fitchiij at Whitemud Creek, but 94 % of the caged mosquitoes at Victoria Park golf course, a more open area. RESUME Le dichlorvos et le propoxur furent appliques sur des bandes de vegetation en ravin boise, et leurs effets sur iactivite des maringouins furent etudies a I’aide de pieges eoliens, et en denombrant les individus atterrissant sur un appat humain. Le dichlorvos, applique a raison de 1.12 kg i.a./ha, reduisit les captures de 70% pendant une journee suivant le traitement; les resultats du denombrement des piqures ne furent pas concluants. Le propoxur en concentration de 0.34 kg i.a./ha reduisit les captures de 44%, et le nombre de piqures de 100%, pendant une journee suivant I’application. Ces resultats ne permettent cependant pas de recommander I’arrosage de bandes de vegetation pour controler les moustiques. De nombreux autres insectes (surtout des Dipteres) furent captures dans les pieges, mais il n’y eut pas de diminution uniforme des captures dans les bandes traitees. Les captures de moustiques furent toujours plus elevees pres du fond que pres du rebord des ravins, et furent generalement plus abondantes dans les pieges places dans les ravins que dans ceux situes sur la plaine environnante. L’efficacite du malathion repandu par helicoptere fut evaluee h partir de la mortalite de moustiques sauvages encages, et du denombrement des piqures. L’epandage a Vatomiseur d’une solution concentree h 0.7 kg i.a/ha ne tua que 27%, des maringouins (principalement Aedes vexansj encages dans un bois a Whitemud Creek. L’epandage a tres faible volume d’une solution a 0.3 kg i.a/ha ne tua que 8%> des maringouins ( surtout Aedes fitchiij encages a Whitemud Creek, mais tua 94% de ceux encages au terrain de golf de Victoria Park, qui est un endroit plus decouvert. 180 Hudson INTRODUCTION According to Klassen and Hocking (1964), the North Saskatchewan River valley and its tributary ravines are a major route of entry of adult mosquitoes into the city of Edmonton, because local air currents and the mosquitoes’ optomotor responses bring them down from the plains into the valleys, and the prevailing winds at night are down-valley. To stop the mosquito invasions, Klassen and Hocking applied lindane by aircraft, at 0.3kg/ha, to vegetation on the sides of the river valley, in three 90m wide barrier strips at the eastern and three at the western city limits. From bite counts and observations on male swarms, it was concluded that the barrier strips were effective, (Klassen and Hocking, 1963), and the method was used, without further evaluation, by Edmonton Parks and Recreation from 1959 to 1969, when lindane was banned. This report describes attempts in 1971 and 1972 to reassess the barrier strip method of control, and to test two substitutes for lindane, using traps designed to measure insect flight activity and direction, (Hocking, 1970; Hocking and Hudson, 1974). In the meantime, the City of Edmonton began using aerial sprays of Malathion against adult mosquitoes, and this report also describes attempts to measure the effectiveness of these sprays. MATERIALS AND METHODS The 1971 barrier strip experiment was performed in a wooded ravine which meets the North Saskatchewan River Valley on the south side at a point 18 km southwest of the City of Edmonton and 3 km east of Devon. The 1972 experiments were performed in the Whitemud Creek Valley, which runs North to Edmonton, at 0.8 and 1.6 km south of the city limits. The ravines at the two test sites were about 300m wide, and thickly wooded with aspen, poplar and spruce, with small streams running through. Two insecticides, dichlorvos (Vapona 20% e.c.,1 2,2 - dichlorovinyl dimethyl phosphate), and propoxur (Baygon 70% w.d.p.,2 ortho-isopropoxyphenyl methyl carbamate) were chosen for the tests because both are quick acting and have a fumigant action, properties they share with lindane. For the dichlorvos trial, a rectangular plot 150 x 70m was marked out on the west slope of the ravine near Devon, and on the 24 July 1971 all the vegetation in the outermost 10m of this plot was treated with the dichlorvos emulsion from hand compression sprayers, at a dosage of 1.11kg a.i.3 in 200 liter/ha. An insect trap was located in an unsprayed area 130 x 50m within the plot. A similar plot on the eastern slope was sprayed with the insecticide solvent only in water. For the propoxur trial, a 200m wide strip across the Whitemud Creek ravine, 0.8km south of the city limits, was sprayed by a helicopter with boom-and-nozzle equipment, at a dosage of 0.3kg a.i. in 39 liters of water per hectare, at 22:30 hrs on the 28 May 1972. A third trial in June 1972 at Whitemud Creek to re-evaluate lindane was abandoned because of a mistake by the helicopter pilot. A fourth trial in July at the same site was to test a new variant of the trap, not an insecticide. 'Emusifiable Concentrate 2Wettable powder 3a.i. = active ingredient Mosquitoes in the Edmonton area 181 Three variations of insect wind traps (Hocking, 1970) were used: the large (10m2) and medium-sized (2m2) ones were set up with one side facing up-valley and the other side down-valley, while the small ones (0.5m2, pivoted and wind-vaned) were set up in small clearings in the forest so they could turn freely. For the dichlorvos trial six traps were used, one large and two small on the west (treated) and the same numbers on the east (untreated) slope of the ravine. For the propoxur trial, five traps were used: one small and two large traps on the ravine bottom (one large inside the treated strip); and two small traps on the plain above, one each side of the ravine. Traps were emptied once or twice daily, and mosquito biting counts made near the traps when they were emptied. Mosquitoes from the traps were identified with the aid of Carpenter and LaCasse (1955). The effectiveness of the malathion sprays was assessed by putting wild-caught Aedes females into cylindrical cages made from 1.4mm mesh galvanized mosquito screen and 10cm plastic petri dishes and the cages were hung from trees and bushes in the areas to be sprayed. One hour after spraying, the mosquitoes were transferred to clean plastic cups and held in the laboratory for another 24h before the mortality count was made. Spray droplets were sampled by white glossy cards (Kromekote Bristol 29 1M), pinned to leaves near the cages, and a few cards were put into empty bioassay cages to measure the droplet penetration rate. The helicopters were fitted in 1972 with boom-and-nozzle equipment to spray malathion emulsion in drops of 500 microns at 0.7kg a.i. in 46 1 of water per hectare, in 1973 with Beecomist ultralow volume (ULV) rotary atomizers to spray undiluted technical malathion in drops of 40 microns at 0.3kg/ha. Reductions in trap catches and bite counts were calculated according to the formula: % reduction = 100(1 - (Tn Co / To Cn)) where Tn and Cn are the numbers caught in the treated and untreated areas, respectively, on day n after treatment, and To and Co are the corresponding values for the day before treatment. Mortalities of caged mosquitoes in the malathion-sprayed areas (T) were corrected for mortalities of mosquitoes caged in the unsprayed areas (C) by Abbott’s formula, as follows: Corrected mortality = 100(T - C/100 - C) RESULTS The wind traps caught many insects (Table I), and the mean catch per m2 at Devon was the highest obtained anywhere in the world, except Tahiti (Hocking, 1970). Of the catch at Devon, 72.4% of the total were female mosquitoes, 97.4% of them Aedes vexans (Meigen), the worst pest mosquito in western Canada. In the trials at Whitemud Creek, female mosquitoes, which never formed more than 15% of the total catch, were mostly Aedes cataphylla in May, Aedes fitchii in June and Aedes vexans again in July. In the dichlorvos experiment there was a 69.8% reduction in trap catch in the treated plot only on the day after treatment (Table II). There was no reduction in catch in the traps up-valley and down-valley from the treated plot. The bite counts were too low to draw any conclusions about the effectiveness of the treatment. In the propoxur experiment there was a 43.8% and 100% reduction in the trap catches and in the biting rates, respectively, in the treated strip the day after treatment (Table III). On the second day after treatment, both flight and biting activity within the treated strip returned to pre-treatment levels. Quaest. Ent. 1981, 17 (3,4) 182 Hudson The wind traps have the advantage that they can also detect changes in the activity for non-target insects, but the results obtained in the propoxur trial were inconsistent. On the day after treatment, there was a reduction in the catch of other Diptera in the treated strip, but no reduction in the catch of Hymenoptera, and the catch of Lepidoptera actually increased. According to Klassen and Hocking (1964) bigger catches could be expected in the traps in the ravines than on the plain, and bigger catches on the up-valley sides of the traps (i.e. mosquitoes flying down-valley) than on the down-valley sides. The 10m2 and 2m2 traps gave consistently higher catches in the ravine than on the plain, but the 0.5m2 traps gave higher catches on the plain, (Table IV). In three of four trials, catches were greater in the sides of the traps facing down-valley, i.e. the direction of flight was predominantly up the valley, and against the prevailing night winds, (Table V). At Whitemud Creek in July 1972, malathion applied by boom-and-nozzle killed only 27.4% of the caged mosquitoes, 84% of which were Aedes vexans, (Table VI), and four hours after spraying the biting rate in the treated area (4.5/min.) was almost as high as in an untreated area nearby (5.5/min). In the 1973 tests, 97% of the caged mosquitoes used were Aedes fitchii. At Whitemud Creek, the average corrected mortality in treated area was only 8% (average uncorrected 26%, average for untreated area 25%), and 0.09 droplets/cm2 were found on the test cards. At Victoria Park, the average mortality of the caged mosquitoes was 94% and 0.26 droplets/cm2 were found on the cards. From the numbers of the droplets found on the cards inside cages, it was estimated that the penetration rate was only 35.4%. In both 1973 tests, mosquitoes bit us in the treated area less than one hour after treatment, and 23% of 48 mosquitoes caught in the treated area died after 24hr holding in the laboratory, compared with 2% mortality in 41 mosquiitoes from an untreated area. DISCUSSION A barrier strip of insecticide should kill all the mosquitoes attempting to cross it, for several days after treatment, but the trap catches indicated reductions in mosquito activity for only one day after treatment, and no measurable reduction in mosquito activity up-valley or down-valley from the strips. The reasons for these poor results are uncertain and require further investigation, but on this basis we could not recommend the resumption of barrier strip spraying with dichlorvos and propoxur. Both dichlorvos and propoxur have very short residual life. Lindane was more effective because of residual properties. Dichlorvos is unstable in the presence of light and water. Propoxur is more stable but the volume of spray, 39 liters/ha, was insufficient for adequate leaf coverage. Brown (1951) states that 50 gallons/acre ( = 5621itres/ha) are required for adequate leaf coverage in woodland. Another important factor in the Edmonton region is the vulnerability of the spray deposits to the summer rains. Two attempts were made in 1972 to bioassay the residual toxicity of propoxur and lindane deposits on leaves, but on both occasions the toxicity of the deposits disappeared within 48h, after heavy rains. The insect wind traps were chosen for the trials because of their ability to sample a wide selection of flying insects, and to measure both flight density and direction of flight. However, in practice the traps proved to be less suitable for these studies than they had seemed. The ratio of mosquitoes to other insects was low in 3 of the 4 trials. The pivoted 0.5m2 traps did not keep facing properly into the light winds down in the ravines, though later a lighter, more efficient wind vane was developed, (Hocking and Hudson, 1974). In studies elsewhere, Hocking found that sometimes insects which had been flying upwind were caught in the upwind, not the Mosquitoes in the Edmonton area 183 downwind, sides of the traps, (Hocking and Hudson, 1974). The three variants of the trap gave different results in the same place for flight densities of mosquitoes, (see Table V), the 2m2 j fixed, pyramidal, traps giving the highest values per m2-hr. This was probably due to differences in area distribution with height. For future studies of mosquito dispersal it would be better to have a larger number of smaller traps, more specific for mosquitoes. The boom-and-nozzle application of malathion was ineffective, probably because of the very large droplets produced. The low temperature at the time of spraying, 10°C, was also unfavourable. The ultralow volume equipment produced drops of around 40 microns in diameter, which is much closer to the optimum of 25 microns for adult mosquitoes, (Weidhaas [ et al. 1970). Malathion ULV sprayed at 0.3 kg/ha. gave satisfactory kills of caged mosquitoes at Victoria Park golf course, a fairly open area, but not in the thickly wooded ravine at Whitemud Creek. In both areas, mosquitoes were taken biting less than one hour after treatment. These mosquitoes may have been protected by resting deep in the undergrowth, below the level of the caged mosquitoes (about 1.5m). The ULV application of malathion was i adopted as routine by Edmonton Parks and Recreation, with a higher dosage (0.5kg/ha) for thickly wooded areas. ACKNOWLEDGEMENTS These studies were supported by grants from the Alberta Agricultural Research Trust and City of Edmonton to the late Dr. Brian Hocking. Assistance of Hugh G. Philip, Brian M. Rolseth, Bernard Boisvert and Jacqueline Marr is acknowledged. The propoxur wettable powder was kindly donated by Chemagro. REFERENCES Brown, A.W.A.. 1951. Insect control by chemicals. New York, John Wiley. 817 pp. Carpenter, S.J. and W.J. LaCasse. 1955. Mosquitoes of North America (North of Mexico). Berkeley, University of California Press. 360 pp. Hocking, B. 1970. Insect flight and entomologists’ inheritance. Ent. News 81:269-278. Hocking, B., and J.E. Hudson. 1974. Insect wind traps: improvements and problems. Quaestiones Entomologicae 10:275-284. Klassen, W., and B. Hocking. 1963. Control of Aedes dispersing along a deep river valley. Mosquito News 23:23-26. Klassen, W., and B. Hocking. 1964. The influence of a deep river valley system on the dispersal of Aedes mosquitoes. Bulletin of Entomological Research 55:289-304. Weidhaas, D.E., M.C. Bowman, G.A. Mount, C.S. Lofgren and H.R. Ford. 1970. Relation of minimum lethal dose to the optimum size of droplets of insecticides for use in mosquito control. Mosquito News 30:195-200. Quaest. Ent. 1981, 17 (3,4) 184 Hudson Table I. Catches of mosquitoes and other arthropods in wind traps at Devon, 1971, and at Whitemud Creek, 1972. Locality Devon (1971) Whitemud Creek (1972) Dates 19-30 July 25-31 May 24-30 June 12- 17 July Trial Dichlorvos Propoxur Lindane1 New trap Number of traps 6 5 8 4 Total area (m2): 22 21.5 26 24 m2-hours: 5515 2750 4014 3295 Catch/m2-hr: 5.24 2.84 2.60 1.32 Catch composition: No. % No. % No. % No. % Female mosquitoes: Aedes vexans 20446 97.4 0 0.0 43 11.2 231 58.9 Aedes fitchii gp2 86 0.9 400 36.1 203 52.9 74 18.9 Aedes communis gp3 4 0.03 474 42.8 48 12.5 7 1.8 Other species 355 1.7 233 20.9 90 23.4 80 20.4 Sub-total 20891 72.4 1107 14.2 384 3.7 392 9.1 Male mosquitoes 640 2.2 1011 13.0 159 1.5 203 4.7 Other Diptera 5788 20.1 3837 49.3 6321 60.7 2583 60.1 Lepidoptera: 734 2.5 610 7.8 1989 19.1 468 10.9 Hymenoptera 706 2.4 863 11.0 738 7.1 405 9.4 Other arthropods: 121 0.4 368 4.7 826 7.9 244 5.7 Total 28880 7796 10417 4295 !Trial stopped because of pilot error 2In May 1972 mostly Ae. cataphylla * Aedes fitchii plus Ae. mercurator Mosquitoes in the Edmonton area 185 Table II. Catches of both sexes of mosquitioes in wind traps, and landing counts of females, in a dichlorvos treated and an untreated plot near Devon, July 1971 Before treatment 1 day 1 day After treatment 2 days 3 days Catches in traps/m2-hr Treated side 1.50 0.18 1.04 5.37 Untreated side 1.71 0.68 1.11 7.09 Landings/person/min. Treated side _i 3 3 4 Untreated side - 1 11 3 % reduction on treated side Trap catches - 69.8 0 0 !Sample not taken Quaest. Ent. 1981,17(3,4) 186 Hudson m Table III. Catches of mosquitoes (both sexes) in wind traps and bite counts before and after propoxur spraying, Whitemud Creek, May 1972. Before treatment 1 day lday After treatment 2 days 3 days Catches in traps/m2-hr Inside treated strip 0.68 0.57 2.26 0.70 Outside treated strip 0.67 1.00 1.32 0.74 On plain (untreated) 1.25 0.89 0.67 1.75 Bites/persons /min. Inside treated strip 5.5 0.0 1.3 1.3 Outside treated strip 8.2 2.6 2.0 3.0 On plain (untreated) 5.0 1.3 1.8 3.6 % reduction in treated strip vs. outside Trap catches _i 43.8 0 6.8 Bite counts - 100.0 3.0 35.4 Sample not taken. Mosquitoes in the Edmonton area 187 Table IV. Catches of mosquitoes of both sexes, per m2-hr, in insect wind traps in the ravine and on the plain, Whitemud Creek 1972. Trap variant: 0.5m2 2.0m2 10.0m2 Location: Ravine Plain Ravine Plain Ravine Plain May 0.30 0.50 _i - 0.75 - June 0.11 0.12 0.34 0.12 0.11 - July - - 0.36 0.12 0.26 0.05 Mean 0.20 0.31 0.35 0.12 0.37 - ‘Sample not taken. Table V. Catches of mosquitoes of both sexes, per m2-hr, in the large insect wind traps (10m2 variant) sited in the ravines on the up-valley and down-valley sides of the traps Locality: Devon -Whitemud Creek- Date: July 1971 May 1972 June 1972 July 1972 Down-valley side 4.89 0.94 0.04 0.18 Up-valley side 3.58 0.40 0.06 0.08 Quaest. Ent. 1981, 17(3,4) 188 Hudson Table VI. Assessment of aerial sprays of malathion against mosquitoes in the Edmonton area, 1972-1973. Location: Whitemud Cr. Whitemud Cr. Victoria Park Date: 1 6 /vii/72 22/vi/73 23 /vi/73 Time: 06.30 22.00 04.30 Temperature (°C): 10 21 16 Spray equipment: Boom + nozzle ULV ULV Dosage, kg a.i./ha: 0.7 0.3 0.3 No. of caged 307 210 424 mosquitoes: Mean mortality, %l: 21 A 8.0 94.0 Droplets on cards/cm2: - 0.09 0.26 Corrected for control mortality by Abbott’s formula KEYS TO THE BLACK FLIES (SIMULIIDAE) OF THE SASKATCHEWAN RIVER IN SASKATCHEWAN1 F.J.H. Fredeen Canada Agriculture Research Station 107 Science Crescent Saskatoon, Saskatchewan, Canada Quaestiones Entomologicae S7N0X2 17:189-210 1981 ABSTRACT Illustrated keys to the larvae, pupae and adults of 15 species of black flies known to occur in the North, South and Main Saskatchewan Rivers in Saskatchewan are presented. Keys are based mainly upon external characteristics. Life histories, trends in abundance and economic importance are discussed. Species treated include Ectemnia taeniatifrons (Enderlein), Metacnephia saskatchewana Shewell and Fredeen, Simulium euryadminiculum Davies, S. duplex Shewell and Fredeen, S. meridionale Riley, S. rugglesi Nicholson and Mickel, S. bivittatum Malloch, S. griseum Coquillett, S. vittatum Zetterstedt, S. arcticum Malloch, S. luggeri Nicholson and Mickel, S. decorum Walker, S. tuberosum (Lundstroem), S. venustum Say, and S. verecundum Stone and Jamnback. RESUME Clefs illustrees pour les larves, les pupaes, et les adultes de chacune des quinze especes des mouches noires connues d’exister dans les trois Rivieres de la Saskatchewan: la Nord, la Sud, et la Principale, sont presentees. Les clefs se basent principalement sur des charateristiques externelles. Des histoires naturelles, des tendanes d’abondance et de la signification economique sont discutees. Les especes sont Ectemnia taeniatifrons (Enderlein), Metacnephia saskatchewana Shewell et Fredeen, Simulium euryadminiculum Davies, S. duplex Shewell et Fredeen, S. meridionale Riley, S. rugglesi Nicholson et Mickel, S. bivittatum Malloch, S. griseum Coquillett, S. vittatum Zetterstedt, S. arcticum Malloch, S. luggeri Nicholson et Mickel, S. decorum Walker, S. tuberosum ( Lundstroem ), S. venustum Say, et S. verecundum Stone et Jamnback. INTRODUCTION For some 30 years the Agriculture Canada Research Station at Saskatoon has been involved in research on black flies inhabiting the Saskatchewan River (Map 1), particularly those species breeding in numbers sufficient to create economic problems. The river has changed considerably during this period, partly because of construction of hydropower dams and has trended toward relatively shallow, weedy conditions. Parallel with these changes, numbers of black fly species have increased, and dominance has shifted from Simulium arcticum Malloch to Simulium luggeri Nicholson and Mickel. This shift created new problems for residents in a large area of Saskatchewan because Simulium luggeri females attack humans as well as other animals and remain abundant all summer unlike Simulium arcticum. Direction of future changes will depend upon climatic conditions and upon artificial manipulations of the river, but 'Contribution No. 803, of the Research Station, Saskatoon 190 Fredeen Map 1. The North, South, and main Saskatchewan Rivers, in Saskatchewan. Black flies (Simuliidae) of the Saskatchewan River 191 because of its large size the Saskatchewan River has potential for creating large black fly outbreaks. Thus, surveillance, research and abatement may be required for many more years. These keys to larvae, pupae, and adults of the 15 species known to inhabit the river were prepared to aid future studies. External features are emphasized to facilitate sorting large samples. Taxonomic concepts and nomenclature of structures follow those used by Davies et al. (1962), Wood et al. (1963), Stone et al. (1965), Davies (1968), Stone and Snoddy (1969) and McAlpine et al. (1981). The keys are based upon specimens stored at the Agriculture Canada Research Station, Saskatoon, augmented by published descriptiions. CHECKLIST OF SPECIES OF SIMULIIDAE FROM THE SASKATCHEWAN RIVER SYSTEM, SASKATCHEWAN, CANADA. 1 . Ectemnia taeniatifrons (Enderlein) 2. Metacnephia saskatchewana Shewed and Fredeen 3. Simulium (Eusimulium) euryadminiculum Davies 4. Simulium (Eusimulium) duplex Shewed and Fredeen 5. Simulium (Byssodon) meridionale Riley 6. Simulium ( Byssodon ) rugglesi Nicholson and Mickel 7. Simulium (Psilopelmia) bivittatum Malloch 8. Simulium (Psilopelmia) griseum Coquillett 9. Simulium (Psilozia) vittatum Zetterstedt 10. Simulium (Gnus) arcticum Malloch 1 1 . Simulium (Phosterodoros) luggeri Nicholson and Mickel 12. Simulium (sensu stricto) decorum Walker 13. Simulium (sensu stricto) tuberosum (Lundstroem) 14. Simulium (sensu stricto) venustum Say 15. Simulium (sensu stricto) verecundum Stone and Jamnback KEYS TO GENERA AND SPECIES ADULTS, Males 1 Entire vein R of wing clothed with hairs (setae) dorsally (Fig. 1) 2 I' Basal section of vein R (Rb, Fig. 1) bare 5 2 (1) Ratio of length of basal section of radius (Rb, Fig. 1) to length of remainder of wing 1:2.5-1:3.0; basal cell present (Fig. 1); calcipala on first hind tarsomere narrow, pointed (Fig. 2); pedisulcus on second hind tarsomere insignificant 3 2' Ratio of length of basal section of radius to length of remainder of wing 1:2.9 - 1:3.9; basal cell absent; calcipala on first hind tarsomere broadly rounded (Fig. 3); pedisulcus deep 4 3 (2) Size smaller, wing 2.0-3.0 mm long; Sc setose ventrally; gonostylus with one minute terminal tooth (Fig. 4) Metacnephia saskatchewana Shewed and Fredeen 3' Size larger, wing 4.0 - 4.5 mm long; Sc bare ventrally; gonostylus with two terminal teeth Ectemnia taeniatifrons (Enderlein) Quaest. Ent., 1981, 17 (3,4) 192 Fredeen 4 (2') Mesepimeral tuft (Fig. 5) pale; ventral plate conical and about as wide as base of gonostylus, markedly convex and with ventral surface clothed in long pale bristles Simulium duplex Shewed and Fredeen 4' Mesepimeral tuft dark; ventral plate broadly rectangular, more than two times wider than base of gonostylus, ventral surface bare, shining, concave Simulium euryadminiculum Davies 5 (F) Gonostylus with three or more teeth and almost square; scutum with two silvery pollinose vittae Simulium vittatum Zetterstedt 5' Gonostylus with one or no teeth and longer than wide in most species; scutum generally without distinct vittae 6 6 (50 Scutellum yellow; lateral and anterior margins of scutum yellow; gonostylus flat, quadrangular, shorter than gonocoxite; ventral plate broad and membranous 7 6' Scutellum and margins of scutum not yellow; gonostylus more or less cylindrical, longer than wide and as long as, or longer than gonocoxite; ventral plate variously shaped 8 7 (6) Integument predominantly black; central area of scutum black with two anterior, lance-shaped, silvery-pollinose spots or short vittae Simulium bivittatum Malloch T Integument predominantly greyish-yellow; scutum greenish-grey with two anterior, rectangular, indistinctly pollinose spots Simulium griseum Coquillett 8 (6') Ventral plate relatively broad with basal arms extended laterally, slightly beyond margins of plate 9 8' Ventral plate spike or tooth-shaped with basal arms extended laterally well beyond margins of plate 11 9 (8) Scutum velvety black suffused with ashen pollinosity especially on anterior half (best seen with horizontal illumination); scutum without prominent shining, white bars or spots, but with three indistinct dark vittae; scutum densely covered with golden hairs; gonostylus without prominent medio-basal lobe Simulium meridionale Riley 9' Scutum velvety black with prominent shining white bar near each anterior corner; scutum sparsely clothed with golden or brown hairs; gonostylus with prominent medio-basal lobe 10 10 (9') Medio-basal lobe on gonostylus covered with fine hairs only Simulium rugglesi Nicholson and Mickel 10' Medio-basal lobe on gonostylus covered with short stout spines Simulium tuberosum (Lundstroem) 1 1 (8') Ventral plate markedly compressed laterally, “Y”-shaped when seen in ventral view 12 IF Ventral plate tooth-shaped in ventral view 13 12 (11) Ventral plate not prominently hairy; gonostylus about three times longer than wide; scutum with single pair of silvery-pollinose spots extended from anterior corners toward center of scutum Simulium arcticum Malloch Ventral plate prominently hairy; gonostylus about two times longer than wide; 12' Black flies (Simuliidae) of the Saskatchewan River 193 scutum with two pairs silvery pollinose spots, each occupying more than one third of disc Simulium decorum Walker 13 (IT) Posterior quarter of scutum with sparse, fine hairs; arms of ventral plate each with postero-medial projection Simulium luggeri Nicholson and Mickel 13' Posterior quarter of scutum with strong, erect hairs; arms of ventral plate without prominent lateral projections 14 14 (130 Ventral plate relatively broad with approximate outline of molar tooth; notched margins of keel of ventral plate flared outward in “V”-shape Simulium venustum Say 14" Ventral plate laterally compressed as narrow keel; notched margins of keel turned inward and parallel with one another Simulium verecundum Stone and Jamnback ADULTS, Females 1 Entire vein R of wing clothed with hairs (setae) dorsally (Fig. 1) 2 Y Basal section of vein R bare 5 2 (1) Basal cell present (Fig. 1); ratio of length of basal section of radius (Rb, Fig. 1) to length of remainder of wing 1:2.5-1:3.0; calcipala on first hind tarsomere narrow and pointed; pedisulcus on second hind tarsomere shallow and indistinct (Fig. 2) 3 2' Basal cell absent; ratio of length of basal section of radius to length of remainder of wing 1:2.9— 1:3.9; calcipala on first hind tarsomere prominent except in S. vittatum and rounded; pedisulcus on second hind tarsomere deep (Fig. 3) 4 3 (2) Frons with nearly parallel sides, narrowest near middle (Fig. 6); size larger (wing 4.0-4. 5 mm long); R dorsally with setae, but lacking black spinules .... Ectemnia taeniatifrons (Enderlein) 3' Frons wedge-shaped, narrowest anterio-ventrally (Fig. 7); size smaller (wing 2.0-3.0 mm long); R dorsally with setae plus black spinules Metacnephia saskatchewana Shewed and Fredeen 4 (2') Medial and two lateral vittae on scutum narrow, black Simulium duplex Shewed and Fredeen 4' Medial and two lateral vittae on scutum narrow, white Simulium euryadminiculum Davies 5 (F) Claw with prominent, thumb-like basal projection (Fig. 8) 6 5' Claw simple or with minute sub-basal tooth (Figs. 9, 10, 1 1) 7 6 (5) Foreleg entirely dark; width of frons at narrowest point about one-quarter as long as frons; frons and terminal abdominal terga grey-pollinose; scutum densely grey, pollinose with three narrow, brown vittae Simulium meridionale Riley 6' Fore coxa and about three-quarters of tibia pale yellow; width of frons at narrowest point about one-half as long as frons; frons and terminal abdominal terga shining dark brown; scutum thinly grey pollinose, subshining and without three narrow brown vittae Simulium rugglesi Nicholson and Mickel 7 (57) Small yellowish flies with postnotum contrastingly dark and with dark dorsal spots on abdomen and ends of some leg articles 8 Quaest. Ent., 1981,17(3,4) 194 Fredeen Fig. 1. Wing of Ectemnia taeniatifrons (Enderlein), showing: basal cell (b); stem vein (Sv); basal section of radius (Rb); subcosta (Sc); costa (C); radial sector (Rs); radius (R); media (M,, M2); submedial false vein (sf); cubitus, anterior branches (Cu,, Cu2), and Anals (An,, An2). Fig. 2. Tarsomeres of E. taeniatifrons , showing: pointed calcipala (c), on first tarsomere; and shallow pedisulcus (p), on second tarsomere. Fig. 3. Tarsomeres of Simulium duplex Shewed & Fredeen, showing: rounded calcipala (c), and deep pedisulcus (p). Fig. 4. Terminalia of male Metacnephia saskatchewana Shewed & Fredeen: parameral spines (p); ventral plate (v); ventral plate arm (a); gonocoxite (gc); gonostylus (gs); and terminal spine (s). Black flies (Simuliidae) of the Saskatchewan River 195 Quaest. Ent., 1981, 17(3,4) 196 Fredeen V Fig. 5. Male of Simulium duplex Shewell & Fredeen, left lateral aspect: mesepimeral tuft (t); scutum (s); scutellum (1); postnotum (p); gonostylus of terminalia (d). Figs. 6 and 7. Head, female, dorsal aspect, showing: antenna (a); eye (e); and frons (f), of : 6, Ectemnia taeniatifrons\ and 7, Metacnephia saskatchewana Shewell & Fredeen. Figs. 8-1 1. Third leg of female Simulium : claw (cl); pedisulcus on second tarsomere (p); calcipala (ca); first tarsomere (t); tibia (ti); femur (f); trochanter (tr); and coxa (c), of : 8. S. meridional Riley; 9, S. vittatum Zetterstedt; 10, S. luggeri Nicholson & Mickel; and 1 1, S', arcticum Malloch. Figs. 12 and 13. Terminalia of female Simulium , ventral view: cercus (c); anal lobe (a); hypogynial valve (h); and genital fork (g), of : 12, S. luggeri ; and 13, S. arcticum. Black flies (Simuliidae) of the Saskatchewan River 197 ! Quaest. Ent., 1981, 17 (3,4) 198 Fredeen 7 8 (7) 8' 9 (70 9' 10 (90 10' 11 (ioo rr 12 (110 12' 13 (120 13' 14 (13') 14' Greyish or brownish flies without contrastingly dark postnotum 9 Scutum with seven alternating stripes of yellowish-grey and brownish orange . Simulium bivittatum Malloch Scutum yellowish-grey without distinct striped pattern Simulium griseum Coquillett Scutum distinctly striped; color patterns of third leg segments as in Fig. 9 ... Simulium vittatum Zetterstedt Scutum not distinctly striped (indistinct vittae may be seen at certain angles of view) 10 Subcosta without setae (rarely one or two) on ventral surface; terminalia with hypogynial valves widely separated and not obscuring genital fork (Fig. 12); color pattern of third leg as in Fig. 10 Simulium luggeri Nicholson and Mickel Subcosta with row of setae on ventral surface (sockets visible if setae lost); terminalia with hypogynial valves almost meeting medially, thus obscuring genital fork (Fig. 13) 11 Hind tibia with distal half dark and proximal half pale (Fig. 11); claw with minute subbasal tooth Simulium arcticum Malloch Hind tibia dark except slanted “knee patch” (Fig. 10); claw without minute subbasal tooth 12 Frons and terminal abdominal terga thinly grey pollinose; anal lobe twice as long as cercus, almost square Simulium decorum Walker Frons and terminal abdominal terga shiny black or brown; anal lobe scarcely longer than cercus, wider than long 13 Fore coxa, trochanter and entire femora as dark as first tarsomere; setae on stem vein and base of costa dark; entire antenna dark; a small dark species (wing length often under 2.0 mm) Simulium tuberosum (Lundstroem) Fore coxa, trochanter and proximal portion of femur paler than first tarsomere; setae on stem vein and base of costa pale; pedicel of antenna relatively pale; size various; abdomen pale ventrally 14 Medial margins of hypogynial valves straight and slightly divergent distally; anterior margin of anal lobe not noticeably more sclerotized than rest of lobe; width and length of dorsal sclerites on third and fifth abdominal segments approximately equal Simulium venustum Say Medial margins of hypogynial valves concave with oval space between them; anterior margin of anal lobe noticeably more sclerotized than rest of lobe; dorsal sclerites on third and fifth abdominal segments approximately two times wider than long Simulium verecundum Stone and Jamnback PUPAE 1 Cocoon loosely woven throughout 2 1' Cocoon tightly woven, at least in posterior portion 3 2 (1) Cocoon sock-shaped, attached or not to short stalk; ten filaments in respiratory organ (Fig. 14) Ectemnia taeniatifrons (Enderlein) 2' Cocoon boot-shaped (elavated anterior collar), not attached to stalk; 17 to 19 Black flies (Simuliidae) of the Saskatchewan River 199 3 (10 3' 4 (3) 4' 5 (30 5' 6 (5) 6' 1 (60 r 8 (50 8' 9 (8) 9' 10 (9') 10' 11 (80 11' 12 (11) 12' 13 (110 13' filaments in respiratory tuft, irregularly branched (Fig. 15) Metacnephia saskatchewana Shewed and Fredeen Pupal respiratory organ with four or fewer filaments 4 Pupal respiratory organ with more than four filaments 5 Pupal respiratory organ of two (rarely three or four) stout filaments (Fig. 16) Simulium duplex Shewed and Fredeen Pupal respiratory organ of four slender filaments (Fig. 17) Simulium euryadminiculum Davies Pupal respiratory organ of six filaments . 6 Pupal respiratory organ of more than six filaments 8 Pupa and cocoon relatively small (about 3.0 mm long) and dark; respiratory filaments less than half as long as cocoon (Fig. 18) Simulium tuberosum (Lundstroem) Pupa and cocoon 4.0 mm or longer, pale; respiratory filaments more than half as long as cocoon 7 Respiratory filaments relatively fine, spread about 90 degrees at base (Fig. 19) Simulium venustum Say Respiratory filaments relatively coarse, spread up to 180 degrees at base (Fig. 20) Simulium verecundum Stone and Jamnback Respiratory organ with eight filaments 9 Respiratory organ with more than eight filaments 11 Respiratory filaments grouped as pairs on four long petioles (Fig. 21) Simulium rugglesi Nicholson and Mickel Respiratory filaments not paired 10 Cocoon more than four mm long, loosely woven (Fig. 22) . Simulium decorum Walker Cocoon about three mm long, tightly woven (Fig. 23) Simulium bivittatum Malloch and Simulium griseum Coquillett1 Pupal respiratory organ with 12 filaments 12 Pupal respiratory organ with more thar 12 filaments 13 Filaments in respiratory organ paired; cocoon boot-shaped (with an elevated anterior collar) (Fig. 24) Simulium arcticum Malloch Filaments in respiratory organ in groups of three; cocoon generally slipper-shaped (without anterior collar) (Fig. 25) Simulium luggeri Nicholson and Mickel Pupal respiratory organ with 16 filaments, in pairs; cocoon slipper-shaped (Fig. 26) Simulium vittatum Zetterstedt Pupal respiratory organ with 22 to 26 irregularly branched filaments; many cocoons boot-shaped (Fig. 27) Simulium meridionale Riley lSimulium bivittatum Malloch and S. griseum Coquillett may be reliably separated as adults. Quaest. Ent., 1981, 17 (3,4) 200 Fredeen Figs. 14-20. Cocoon, and one set of respiratory filaments of pupa, of: 14, Ectemnia taeniatifrons (Enderlein); 15, Metacnephia saskatchewana Shewed & Fredeen; 16, Simulium duplex Shewed & Fredeen; 17, S. euryadminiculum Davies; 18, S. Tuberosum (Lundstroem); 19, S. venustum Say; 20, S. verecundum Stone & Jamnback. Black flies (Simuliidae) of the Saskatchewan River 201 Figs. 21-27. Cocoon and one set of respiratory filaments of pupa, of: 21, Simulium rugglesi Nicholson & Mickel; 22, S. decorum Walker; 23, S. griseum Coquillett, and S. bivittatum Malloch; 24, S. arcticum Malloch; 25, S. luggeri Nicholson & Mickel; 26, S. vittatum Zetterstedt; and 27, S. meridionale Riley. Quaest. Ent., 1981, 17 (3,4) 202 Fredeen LARVAE 1 F 2 (1) 2' 3 (2') 3' 4 (30 4' 5 (20 5' 6 (5) 6' 7 (6) T 8 (60 8' 9 (80 9' 10 (50 10' 11 (10) IF Three unbranched anal gills (Fig. 28) 2 Three compound anal gills 5 Lateral margins of abdominal segments 5 to 8 extended ventrally to about twice depth of segments 1 to 4 (Fig. 28); prominent pair of ventral tubercles (Fig. 28); ventral view of head capsule (Fig. 29) Ectemnia taeniatifrons (Enderlein) Lateral margins of abdominal segments 5 to 8 not extended ventrally; anal sclerites darkly sclerotized (Fig. 30); ventral tubercles insignificant 3 Arms of anal sclerite as in Fig. 31; postgenal cleft extended to base of hypostomium (Fig. 32) Metacnephia saskatchewana Shewed and Fredeen Arms of anal sclerite as in Fig. 33; postgenal cleft extended less than half distance to base of hypostomium (Fig. 34) 4 Suboesophageal ganglion dark (Fig. 34); larva relatively large ( > 7 mm long when mature) and with prominent dark, greenish transverse bands Simulium vittatum Zetterstedt Suboesophageal ganglion almost invisible (Fig. 35); larva relatively small (< 5 mm long when mature) and without prominent dark transverse bands Simulium bivittatum Malloch and Simulium griseum Coquillett2 Eighth abdominal segment with two prominent ventral tubercles; abdominal integument sparsely clothed in fine short bristles 6 Eighth abdominal segment without prominent ventral tubercles; abdominal integument essentially bare 10 Postgenal cleft extended to base of hypostomium (Figs. 36, 37) 7 Postgenal cl€ft not extended to base of hypostomium (Fig. 38) 8 Prominent row of bristly tubercles in transverse reddish band encircling each of segments 1 to 5, smaller tubercles on remaining segments; suboesophageal gland colorless Simulium meridionale Riley Abdomen without bristly tubercles; suboesophageal gland dark Simulium rugglesi Nicholson and Mickel Ventral tubercles broadly rounded; postgenal cleft broadly rounded, extended about three quarters of distance to base of hypostomium (Fig. 38) Simulium luggeri Nicholson and Mickel Ventral tubercles distinctly conical; postgenal cleft extended no more than half distance to base of hypostomium 9 Postgenal cleft extended about one-third of distance to base of hypostomium, bilobed (Fig. 39) Simulium euryadminiculum Davies Postgenal cleft extended about one-half of distance to base of hypostomium, not bilobed (Fig. 40) Simulium duplex Shewed and Fredeen Medial spots on dorsal surface of head capsule dark 11 Medial spots on dorsal surface of head capsule pale 12 Suboesophageal ganglion dark; postgenal cleft extended about half way to base of hypostomium (Fig. 41) Simulium tuberosum (Lundstroem) Suboesophageal ganglion almost colorless; postgenal cleft extended about three-quarters of distance to base of hypostomium (Fig. 42) S. bivittatum and S. griseum may be reliably separated as adults only. Black flies (Simuliidae) of the Saskatchewan River 203 Fig. 28. Lateral view of posterior portion of abdomen of Ectemnia taeniatifrons (Enderlein) larva: anal gills; g); posterior ring of hooklets (r); ventral tubercle (t). Not drawn to scale. Fig. 29. Ventral view of head Capsule of E. taeniatifrons larva: hypostomium (h); suboesophageal ganglion (g); margin of postgenal cleft (p). Fig. 30. Lateral view of Metacnephia saskatchewana Shewed & Fredeen larva: anal gills (g); portion of posterior ring of hooklets (r). Not drawn to scale. Fig. 31. Dorsal view of anal sclerites of M. saskatchewana larva: anal sclerite (s); portion of posterior ring of hooklets (r). Not drawn to scale. Fig. 32. Ventral view of head capsule of M. saskatchewana larva: postgenal cleft (p). Fig. 33. Dorsal view of anal sclerites of Simulium vittatum Zetterstedt larva: anal sclerite (s); portion of posterior ring of hooklets (r). Not drawn to scale. Figs. 34-45. Ventral view of head capsule of Simulium larvae: suboesophageal ganglion (g); margin of postgenal cleft (p), of: 34, S. vittatum Zetterstedt; 35, S. griseum Coquillett and S. bivittatum Malloch; 36, S. meridionale Riley; 37, 5. rugglesi Nicholson & Mickel; 38, S. luggeri Nicholson & Mickel; 39, S. euryadminiculum Davies; 40, 5. duplex Shewed & Fredeen; 41, S. tuberosum (Lundstroem); 42, S. articum Malloch; 43, S. decorum Walker; 44, S', venustum Say; 45, S. verecundum Stone & Jamnback. (Scale applies to head capsules, only). Quaest. Ent., 1981, 17 (3,4) 204 Fredeen Simulium arcticum Malloch 12 (100 Mature larvae large (many 7 mm long); antenna extended as far as distal end of stalk of cephalic fan; postgenal cleft slightly longer than wide (Fig. 43) Simulium decorum Walker 12' Mature larvae smaller (many 6 to 6.5 mm long); tip of antenna extended beyond distal end of stalk of cephalic fan; postgenal cleft about as long as wide (Figs. 44,45) . . Simulium venustum Say and Simulium verecundum Stone and Jamnback3 NOTES ABOUT SPECIES 1. Ectemnia taeniatifrons (Enderlein) (Figs. 1, 2, 6, 14, 28, 29) Thirty years ago larvae and pupae of this species were widely distributed throughout the north, south, and main branches of the Saskatchewan River in Saskatchewan; none were found in tributaries. In recent years fewer specimens have been collected and those mainly near the confluence of the north and south branches near Prince Albert, the Red Deer River near Estuary and the South Saskatchewan River near Pike Lake. Ectemnia taeniatifrons is univoltine. Eggs laid in April and May hatch in late summer. At fall freeze-up, some larvae are almost half grown. Larvae are notable in that many are found attached to the tips of slender stalks up to 15 mm long, built from threads of salivary secretions and debris and attached with broad bases to substrates. If undisturbed, a larva attaches its cocoon on or near the tip of its stalk. Larvae continue growth under the ice and adults commence emerging three to five weeks after ice break-up. Adults commonly feed on sap seeping from the bruised bark of birch trees. Mating occurs in flight in thinly dispersed swarms in clearings and thereafter, the females seek blood from humans and other animals. Because of its rarity the species was not considered to be economically important. 2. Metacnephia saskat chew ana Shewed and Fredeen (Figs. 4, 7, 15, 30-32) Larvae and pupae occur regularly on water weeds collected in the spring from the Shell and Battle Rivers in Saskatchewan and occasionally from the North Saskatchewan River as far as about 50 km downstream from the outlets of those small tributaries. I believe that the species is not native to the Saskatchewan River, but that larvae appear there only after accidentally drifting in from tributaries. The cocoons possess the same shape as those of S. arcticum , but are loosely constructed and the pupa possesses 17 to 19 respiratory filaments instead of 12. Only one generation per year is known. Adults emerge four to eight weeks after ice break-up suggesting that the species overwinters as larvae under the ice of permanently flowing rivers. Feeding habits of adults are not known. Type specimens for this species were collected from the Shell River (Shewed and Fredeen, 1958) and since then it has been shown to be widely distributed in western Canada. *S. venustum and S. verecundum may be reliably separated in the pupal and adult stages only. Black flies (Simuliidae) of the Saskatchewan River 205 3. Simulium (Eusimulium) euryadminiculum Davies (Figs. 17 and 39) This species breeds abundantly in the Battle and Shell Rivers. Larvae occasionally drift into the North Saskatchewan River and have been collected as far as about 60 km downstream from those tributaries. There, adults are sometimes produced judging by empty pupal cases, but there is no evidence of permanent establishment in the Saskatchewan River. S. euryadminiculum is univoltine with adults emerging in May about three to five weeks after ice break-up. Blood-fed females have been taken from a dead common loon (Davies et al., 1962) and have driven chickens indoors in Saskatchewan, without causing losses. The most severe attack on chickens occurred May 16 to 18, 1981, near Prongua, Saskatchewan. 4. Simulium ( Eusimulium ) duplex Shewed and Fredeen (Figs. 3, 5, 16 and 40). Larvae and pupae of this species occur regularly in the Battle and Shell Rivers, the source of type specimens (Shewed and Fredeen, 1958) and in small numbers in the Saskatchewan River below their outlets. They were collected once from the Red Deer River immediately above its confluence with the South Saskatchewan River in southwestern Saskatchewan. S. duplex is univoltine, with pupation usually occurring in late May. Feeding habits of adults are not known. 5. Simulium (Byssodon) meridionale Riley (Figs. 8, 27 and 36). S. meridionale is widely distributed throughout many rivers and streams in Saskatchewan. Recently, larvae have become relatively abundant in the North Saskatchewan River, especially in the vicinities of the Wingard and Cecil ferries. They also occur in the Red Deer and South Saskatchewan Rivers in southwest Saskatchewan and occasionally downstream from Tobin Lake on the main Saskatchewan. Eggs overwinter in river bed sand and commence hatching within a month after ice break-up in the spring. The species is multivoltine. Larval populations often peak in May and June in the South Saskatchewan and in July in the north branch. It is sometimes abundant enough to be a pest of humans and other animals. In some localities such as Melville, Saskatchewan, it has been a pest of poultry and implicated indirectly in fatal proven Leucocytozoon infections in flocks of domestic poultry. Those outbreaks originated in small local streams not connected with the Saskatchewan River. Possibly sibling species of S. meridionale possessing different host preferences are involved. Surprisingly, summer-long emergences of relatively large numbers of S. meridionale in recent years from the North Saskatchewan River did not result in poultry losses until early July 1981, when six farmers in a 100,000 ha area near Nipawin, Saskatchewan reported losses. Veterinarians considered those fatalities due to exsanguination rather than disease. S. meridionale females were abundant inside chicken house windows. Deaths occurred too rapidly to allow collection of blood smears. Until more information is available poultry producers should avoid establishing new flocks out-of-doors near the North Saskatchewan River. Swarms of S. meridionale have been bothersome as far as 15 km from that river in recent years. Poultry producers already established should report immediately any massive illnesses in their flocks to the Western College of Veterinary Medicine at Saskatoon. Blood is required from living birds for diagnosis. Quaest. Ent., 1981, 17 (3,4) 206 Fredeen 6. Simulium (Byssodon) rugglesi Nicholson and Mickel (Figs. 21 and 37) Eggs, larvae and pupae of this species occur regularly and sometimes abundantly in small clear rivers such as, the Battle, Shell, and Torch, and in small numbers in the North Saskatchewan River below the outlets of those tributaries. Eggs overwinter in river bed sand and commence hatching about one month after ice break-up in spring. Larvae attain maximum abundance about mid-June, but may be found until September, suggesting more than one generation per summer. Shewed (1955) identified S. rugglesi as the species believed responsible for transmitting fatal Leucocytozoon infections to ducklings and goslings in eastern Canada. However, there have not been any reports of this black fly attacking poultry in Saskatchewan. 7. Simulium (Psilopelmiaj bivittatum Malloch and 8. S. (Psilopelmiaj griseum Coquillett (Figs. 23 and 35) I was not able to reliably separate immature stages of these two species. Adults reared from pupae collected throughout the ice-free season in 1976 showed that the two species occurred together that year in both the South Saskatchewan and Red Deer Rivers in southwest Saskatchewan. In earlier years larvae and pupae of both species were found every summer in the South Saskatchewan River. They were abundant in upper reaches, but rare in lower reaches of that river and absent from the North Saskatchewan River. Populations in the lower South Saskatchewan apparently were renewed annually by downstream drift of eggs and larvae from breeding sites much further upstream because specimens have not been collected since completion of a hydroelectric dam near Macrorie in 1968. At Estuary, newly-hatched larvae appeared in May, a few weeks after ice break-up. Development was rapid and by late May pupation commenced. Both species were multivoltine and attained maximum abundance in July and August. Occasional severe outbreaks until about 1965 used to force people indoors at Saskatoon and Medicine Hat (Alberta). Adults of these species also attacked horses, cattle, sheep and hogs, but apparently not poultry. 9. Simulium (Psilozia) vittatum Zetterstedt (Figs. 9, 26, 33, and 34). Although S. vittatum is the most widely distributed black fly species in Saskatchewan it apparently did not become established in the Saskatchewan River until the 1960’s, coincidental with appearance of several species of water weeds. Now immature stages occur throughout this river system and are particularly abundant in beds of weeds that provide attachment sites for larvae and pupae, and substrates suitable for egg masses on emergent leaves. S. vittatum overwinters as eggs in the river bed, and occasionally as larvae. It is univoltine and attains greatest abundance in late July and early August. As a pest it is not considered too bothersome, even when adults are abundant. These are relatively unobstrusive in attacks, quietly entering the ears of animals where they feed undisturbed. 10. Simulium (Gnus) arcticum Malloch (Figs. 11, 13, 24, and 42) S. arcticum was considered to be the only black fly species capable of killing livestock in Saskatchewan until recently when S. luggeri became the dominant species in the Saskatchewan River. Larvae and pupae used to occur throughout the entire Saskatchewan River in Saskatchewan, but were particularly abundant in rocky rapids in the final 1 50 km or so of each of the north and south branches and in the entire main river below their confluence. The most destructive outbreaks occurred from 1944 to 1947, inclusive, when more than 1,100 farm Black flies (Simuliidae) of the Saskatchewan River 207 animals, mainly cattle, were killed, including many expensive herd sires. Up to 3 million ha of farm lands in Saskatchewan were affected at irregular intervals by these outbreaks until advent of chemical larviciding in 1948 (Fredeen, 1958). Deaths were caused by direct toxemia. Native born mature mammals were relatively immune to the toxins, but outbreaks still caused indirect losses by interrupting grazing and breeding. Most outbreaks occurred in late May or early June, rarely in August. Since the early 1960’s numbers of larvae in these rivers gradually declined until this species no longer threatens livestock. The last outbreak in which fatalities occurred was in 1972 near Wingard. This decline seems not to have been caused by larviciding, as that affected only 150 km or shorter sections of these rivers, reinfested annually from untreated sections upstream. Rather, the decline coincided with a combination of several ecological changes including increases in amounts of dissolved plant nutrients, shallowing and clearing of water due to summertime impoundment behind new hydroelectric dams, and development of massive beds of aquatic plants. Reduced numbers of larvae of S. arcticum continue to inhabit rapids, however, and the species might return in bothersome numbers if eutrophication declines. S. arcticum overwinters as eggs in submerged river sand. First-instar larvae begin to appear 10 to 14 days after ice break-up in the spring and pupation commences four or more weeks later. A second, smaller generation often occurs in July or August. Downstream drift of larvae undoubtedly played an important part in development of outbreaks. When river levels declined as they usually did in May and early June, larvae released and drifted downriver to more suitable sites in rocky rapids. It was from such accumulation points that many destructive outbreaks originated. Accumulation of unfed females awaiting weather conditions suitable for flight also was a factor. There was evidence, however, that some females could lay the first batch of eggs autogenously, that is, from nutrients stored during larval life (Fredeen, 1963). Mammals within about 30 km of rapids were perhaps in greatest danger of attack. In 1947, mammals were killed near Simpson, Saskatchewan, some 200 km south of the nearest point of origin of black fly populations. Three morphologically different forms of S. arcticum have been collected in Saskatchewan. The pest species from the Saskatchewan River was identified as S. arcticum by Stone (1959) after comparing it with Malloch’s types. A second relatively small, pale variety is widespread in northern Saskatchewan and a third large, dark variety is in the Cypress Hills in southwest Saskatchewan. 11. Simulium ( Phosterodoros ) luggeri Nicholson and Mickel (Figs. 10, 12, 25, and 38). S. luggeri is the most northerly distributed species of Phosterodoros , and is the only one known to occur in western Canada. Until the late 1960’s breeding of S. luggeri Nicholson and Mickel in Saskatchewan was restricted to medium-sized clear rivers such as the Battle, Shell and Torch Rivers where there was vegetation for attachment by larvae. From these rivers larvae drifted in small numbers out into the North Saskatchewan River. However, they failed to colonize either branch of the Saskatchewan until perhaps 1968 when larvae were detected for the first time in the south branch near Fenton. By August, 1971, it was breeding so abundantly in that river that emerging swarms of adults stampeded cattle in a pasture near St. Louis. Also, for the first time in 1971 there was a distinct increase in numbers of larvae found attached to artificial substrates anchored in the north branch near Prince Albert, indicating that a breeding population had become established there. In 1976, larvae exceeded those of S. Quaest. Ent., 1981, 17 (3,4) 208 Fredeen arcticum in abundance and for the first time outbreaks of economic proportions originated repeatedly from that river. Since then, outbreaks have continued to severely affect people and livestock in some 18,000 km2 of farmlands and recreational areas in east central Saskatchewan. Even though S. luggeri does not poison animals as severely as does S. arcticum , the habit of swarming densely around heads of cattle causes stampeding. Grazing and breeding are repeatedly interrupted throughout the summer and fences are broken down. Young animals suffer broken bones and increased incidence of stress-related diseases such as pneumonia and diarrhoea. Summer-long outbreaks reached a peak in 1978, causing losses including unrealized weight gains that were estimated to have exceeded $1,400,000.00 Fredeen (in preparation). Relatively intensive larviciding prevented re-occurrences of these outbreaks in 1979. S. luggeri is capable of about five generations per summer in this region. Like S. arcticum it overwinters as eggs in submerged sand. Hatching commences about two weeks after ice break-up in the spring and adults emerge in late May or early June. Attacks on people and livestock continue during warm days until late in the fall. Egg masses are attached to emergent vegetation and also, single eggs, similar to those of S. arcticum may be dropped onto the water surface . 12. Simulium (sensu stricto ) decorum Walker (Figs. 22, and 43). This species is distributed sparsely throughout the province. Larvae occur most frequently in the outfalls from small stream impoundments. Single larvae sometimes collected from the North Saskatchewan River presumably originated in small streams. Not enough is known about S. decorum to predict whether it could commence breeding in fast flowing sections of the Saskatchewan River. It overwinters as eggs in wet river sand and apparently is capable of several generations per summer. The females attack a wide vareity of animals. In northern Saskatchewan adults are sometimes abundant enough to create severe local problems for' humans and other animals. 13. Simulium (sensu stricto) tuberosum (Lundstroem) (Figs. 18 and 41). Landau (1962) showed that S. tuberosum (Lundstroem) in southern Ontario consists of a complex of at least four cytogenetically distinct species. Several morphological forms occur in Saskatchewan indicating a complex here, also. Larvae occur sparsely but regularly in the North Saskatchewan River, especially downstream from tributaries that are normal habitats. Larvae were reliably reported from South Saskatchewan River for the first time in 1979 and again in 1980, when a few were collected from three sites, 20, 40 and 120 km upstream from the confluence with the north branch. This widespread distribution and lack of stream tributaries which might have contributed larvae, suggests that the species is now breeding in that river. S. tuberosum (Lundstroem) may be multivoltine and adults have been collected along with those of S. venustum Say from swarms attacking humans and other animals. However, nowhere in the southern half of the province has it been abundant enough to create problems by itself. 14. Simulium (sensu stricto) venustum Say (Figs. 19, and 44). S. venustum Say is one of the most widely distributed black flies in Saskatchewan and breeds in a wide variety of relatively unpolluted streams and rivers. Occasionally faster flowing sections of small rivers such as the Assioniboine, Big and Beaver have produced brief outbreaks Black flies (Simuliidae) of the Saskatchewan River 209 that have created problems for humans and other animals. In the early 1970’s S. venustum Say commenced breeding sparsely in the last 50 km or more of both branches of the Saskatchewan River. Although still uncommon there it should be watched because of the large productive capacity of the Saskatchewan River. S. venustum Say overwinters as eggs and is multivoltine, attaining greatest abundance in June. Rothfels (1975) considers this to be a species complex. 15. Simulium (sensu stricto ) verecundum Stone and Jamnback (Figs. 20, and 45). Larvae and pupae of this species occasionally have been colleted from the last 50 km or so of each branch of the Saskatchewan River. Although it may become more abundant it probably will not become economically important because it is not known to attack for blood. PRACTICAL CONSIDERATIONS The black fly fauna of the Saskatchewan River in Saskatchewan has changed considerably during the past 15 years coinciding with changes in river conditions. Originally S. arcticum was the dominant species. Sporadic outbreaks that killed livestock belonging to the earliest settlers in 1885 were attributed to this species. As recently as 1972 an outbreak of S. arcticum originating from an untreated portion of the North Saskatchewan River killed 18 farm animals near Wingard. Since then, however, S. luggeri has gradually replaced S. arcticum as the dominant species and the conversion is believed due to changes in river conditions that allowed growth of large beds of aquatic plants on previously barren river beds. Losses estimated to have exceeded $1,400,000.00 resulted from summer-long outbreaks of S. luggeri in 1978 (Fredeen, in preparation). Tendencies towards continuing changes in black fly populations were indicated by appearances of larvae of S. tuberosum (Lundstroem) and S. venustum Say in several sites in 1979 and 1980. Troublesome outbreaks of those two blood sucking species have been reported from several small prairie rivers. Thus their populations should be monitored annually along with those of blood sucking species in the relatively large Saskatchewan River. At present, chemical larviciding offers the only means of preventing outbreaks (Fredeen, 1977). Continued research may eventually provide other methods. Residents in areas recently affected by outbreaks of S. arcticum Malloch and S. luggeri Nicholson and Mickel require assurance that protection will be provided when required. It is hoped that this key to species inhabiting the Saskatchewan River will prove useful for monitoring and research. ACKNOWLEDGEMENTS I am indebted to Dr. L. Burgess, Canada Agriculture Research Station, Saskatoon, Saskatchewan, for reviewing this paper and Ralph Underwood for advice during preparation of the figures. REFERENCES Davies, L. 1968. A key to the British species of Simuliidae (Diptera) in the larval, pupal and adult stages. Freshwater Biological Association, Westmorland, England, scientific publication number 24. 126 pp. Davies, D.M., B.V. Peterson and D.M. Wood. 1962. The black flies (Diptera: Simuliidae) of Quaest. Ent., 1981, 17(3,4) 210 Fredeen Ontario. Part I. Adult identification and distribution with descriptions of six new species. Proceedings of the Entomological Society of Ontario (1961). 92: 71-154. Fredeen, F.J.H. 1958. Black flies (Diptera: Simuliidae) of the agricultural areas of Manitoba, Saskatchewan and Alberta. Proceedings, Tenth International Congress of Entomology 3: 819-823. Fredeen, F.J.H. 1963. Oviposition in relation to the accumulation of blood thirsty black flies (Simulium (Gnus) arcticum Mall (Diptera)) prior to a damaging outbreak. Nature (London) 200: 4910. p. 1024. Fredeen, F.J.H. 1977. Black fly control and environmental quality with reference to chemical larviciding in western Canada. Quaestiones Entomologicae 13: 321-325. Fredeen, F. J.H. In preparation. Outbreaks of Simulium luggeri in Saskatchewan. Landau, R. 1962. Four forms of Simulium tuberosum (Lundstr.) in southern Ontario: a salivary gland chromosome study. Canadian Journal of Zoology 40: 921-939. McAlpine, J.F., B.V. Peterson, G.E. Shewed, H.J. Teskey, J.R. Vockeroth and D.M. Wood. 1981. Manual of Nearctic Diptera. Volume 1, Monograph No. 27, Research Branch, Agriculture Canada, Ottawa, Canada, 674 pp. Rothfels, K.H. 1975. Personal communication. Department of Botany, University of Toronto, Canada. Shewed, G.E. 1955. Identity of the black fly that attacks ducklings and goslings in Canada (Diptera: Simuliidae). The Canadian Entomologist 87: 345-349. Shewed, G.E. and F.J.H. Fredeen. 1958. Two new black flies from Saskatchewan (Diptera: Simuliidae). The Canadian Entomologist 90: 733-738. Stone, A. 1959. Personal communication. United States Department of Agriculture, Entomology Research Division, Washington, D.C. Stone, A., C.W. Sabrosky, W.W. Worth, R.H. Foote and J.R. Coulson. 1965. A catalogue of the Diptera of America north of Mexico, Agruculture Handbook No. 276, Agricultural Research Service, United States of America, Washington, D.C. 1696 pp. Stone, A. and E.L. Snoddy. 1969. The black flies of Alabama (Diptera: Simuliidae). Bulletin 390. Agricultural Experiment Station, Auburn University, Alabama. 93 pp. Wood, D.M., B.V. Peterson, D.M. Davies and Helen Gyorkos. 1963. The black flies (Diptera: Simuliidae) of Ontario. Part II. Larval identification, with descriptions and illustrations. Proceedings of the Entomological Society of Ontario (1982) 19: 99-129. SEASONAL CAPTURES AND DIVERSITY OF GROUND BEETLES (COLEOPTERA: CARABIDAE) IN A WHEAT FIELD AND ITS GRASSY BORDERS IN CENTRAL SASKATCHEWAN4 J.F. Doane Research Station, Research Branch Agriculture Canada 107 Science Crescent Saskatoon, Saskatchewan, S7N 0X2 Canada Quaestiones Entomologicae 17:211-233 1981 ABSTRACT Pitfall trapping, at weekly intervals from spring to autumn during a 2-year period in a wheat field and its grassy borders near Clavet, Saskatchewan, captured 87 species of carabids in 25 genera. Seven percent of these species were captured only in the field, 30% only in the field border and 63 % in both habitats. Bembidion obscurellum Mtsch., Bembidion quadrimaculatum oppositum Say, Bembidion nitidum Kby., Amara lacustris Lee., Amara carinata Lee., Amara farcta Lee., Harpalus herbivagus Say and Pterostichus lucublandus Say were dominant in both habitats, while Bembidion timidum Lee. was dominant only in the field, and Amara quenseli Schnh. and Amara obesa Say, were dominant only in the border. The overall sex ratio in the four collections was about 1:1. However, data for some species differed considerably from this ratio. Species diversity and evenness were higher for the field border collections than for collections from the field. An average of about 15 species per week was trapped in the border compared to an average of 11 species per week in the field. RESUME Le Piegeage em fosses-collectionneuses dans un champs de ble sis pre de Clavet, Saskatchewan, et dans les bordures herbeuses de de celui-ci, aux ecartements d’une semaine a partir du printemps jusqu’ a partir du printemps jusqu’ a /’ automne pendant deux ans, a pris 87 especes de carabides. reparties en 25 genres. En proportion, les especes prises ont ete comme suit: seulement dans le champs, 7 p.c.; seulement dans les bordures, 30 p.c.; et a la fois dans le champs et dans les bordures, 63 p.c. Bembidion obscurellum Mtsch., Bembidion quadrimaculatum oppositum Say, Bembidion nitidum Kby., Amara lacustris Lee., Amara carinata Lee., Amara farcta Lee., Harpalus herbivagus Say et Pterostichus lucublandus Say etaient predominates parmi les especes qui se sont trouvees dans les deux habitats, tandis que Bembidion timidum Lee. a predomine parmi celles qui se sont rencontrees seulement dans le champs et Amara quenseli Schnh. et Amara obesa Say ont predomine parmi celles qui ont demeurees dans les bordures. La proportion sexuelle globale de quatre collections a ete approximativement en rapport de 1:1 , cependant, cette proportion a souvent varie considerablement de ce niveau dans le cas des especes particulieres. La diversite et iuniformite des especes ont ete plus elevees dans les collections prises des bordures que dans celles des champs. Une moyenne de 15 especes par semaine a ete captee dans les bordures a comparer dll especes par semaine en moyenne dans le champs. Contribution No. 793, from the Research Station, Saskatoon. 212 Doane INTRODUCTION Investigations of the way of life of carabids associated with agricultural lands have usually been conducted to determine the potential importance of dominant species as predators and their possible role in the population dynamics of crop pests (Rivard 1964; Frank 1971a; Kirk 1971a; Esau and Peters 1975; Finlayson and Campbell 1 976% Holliday and Hagley 1978). It is also important to determine species composition, population structure and diversity of carabid populations associated with various habitats, including arable lands. This information is potentially useful for assessing impact of agricultural or other practices on stability of carabid populations. Temporal changes or spatial differences in diversity could be a valuable biological index for assessing effect of agronomic practices on carabid populations (Dritschilo and Wanner 1980). Carabids are known to attack wireworm larvae (Fox and MacLellan 1956) and are potential predators of all stages of wireworms. This paper reports on pitfall trapping for carabids in a wireworm infested field in Central Saskatchewan to determine: (1) dominant species that might be important as predators of wireworms; (2) the species composition and diversity of collections. For comparison, similar collections were made in the uncultivated field border which also harboured a wireworm population. Observations on the sex ratio and seasonal occurrence of some species are also included. METHODS The general methods, plot arrangement, plot size, and vegetation surrounding the study field (52° 02' N., 106° 24' W.) near Clavet, Saskatchewan, were as described previously (Doane and Dondale 1979). The field study, cultivated and seeded to spring wheat in June in 1975 and 1976, was harvested each year in late August and early September. A pitfall trap was placed at the centre of each of 16 field plots and 15 field border plots. Carabids were trapped in these two habitats for a 48-hr period each week, usually from Monday until Wednesday, from April or May until mid-October, giving a total of 44 and 56 trap-days in 1975 and 1976 respectively. During the remainder of the week, traps remained in place, but carabids captured were released in order to avoid possible over trapping of some species. Sex of all carabids trapped was determined, and for species that comprised one per cent or more of the total annual collection from each habitat, chi-square analysis was used to test conformity to a 1:1 male-female ratio. Abundance of different species within collections was expressed as a percentage of the total collection for each year and habitat. To allow better comparison between years and habitats, in which different numbers of trap-days and traps were used respectively, total seasonal catch per species was also expressed as captures per metre of trap circumference per trap-day, since trap circumference is one factor influencing number of captures (Turner 1962; Thomas and Sleeper 1977). The inside rim of the pitfall trap used in the present studies, described previously (Doane 1961), had a circumference of 51.496 cm giving a total trapping circumference of 8.24 m (16 traps) and 7.72 m (15 traps) in the field and field border respectively. Diversity and evenness of carabid collections made for the two years and sites was determined using the Brillouin diversity index (H) and Simpson’s index of diversity (D) (Pielou 1969). Choice of the Brillouin index has been discussed previously for similar spider collections (Doane and Dondale 1979). The Brillouin index was calculated using a table compiled by Lloyd et al. (1968), while maximum diversity (Hmax) and evenness was determined by using Ground beetles (Coleoptera: Carabidae) in Central Saskatchewan 213 equations given by Pielou (1969). Simpson’s measure of diversity (D) (Pielou 1969) was used for comparison since it is worthwhile to have more than one measure of diversity. RESULTS AND DISCUSSION Abundance and seasonal diversity Numbers of genera and species captured in each habitat for each year, were quite consistent (Table 1). More individuals were captured in 1976 in both the field and its border, but on the basis of numbers trapped per metre of trap circumference per trap-day the greatest numbers were trapped in 1975 in the field (Table 2). Eighty-seven species were found in the four collections (Table 2); of these 7% were collected only from the field, 30% only from the field border and, 63% from both habitats. Dominant species, those that comprised 3% or more of the total individuals in at least one of the collections (Table 2), were the following: Bembidion obscurellum Mtsch., Bembidion quadrimaculatum oppositum Say, Bembidion nitidum Kby., Bembidion timidum Lee., Amara lacustris Lee., Amara carinata Lee., Amara fracta Lee., Amara quenseli Schnh., Amara obesa Say, Harpalus herbivagus Say and Pterostichus lucublandus Say. Of these, B. timidum was dominant only in the field and A. quenseli and A. obesa were dominant only in the border; the remainder were dominant in both habitats. For convenience, species abundance and seasonal occurrence are discussed under generic groupings. Seasonal activity of dominant species that were trapped at Clavet, Saskatchewan, agreed closely with that observed by Frank (1971a) for the same or congeneric species occurring in Alberta. Bembidion Eighteen species were captured in the field and field border during the 2-year study period, eight species in the field and 17 in the field border. B. obscurellum was by far the most abundant species collected in the field in 1975, comprising about 40% of the total individuals in the collection (Table 2), and about 85% of the individuals in that genus. Collection rate of 3.15/m trap circumference/trap-day was the greatest for any species during either year. In 1976, captures of B. q. oppositum were slightly in excess of B. obscurellum in the field. More rare species of Bembidion were trapped in the field border than in the field. Frank (1971a) captured 11 Bembidion species in a cultivated field at Calahoo, Alberta (about 32 km northwest of Edmonton). There, B. q. oppositum was the most abundant species followed by B. obscurellum , B. rupicola, B. nitidum and B. bimaculatum. Bembidion rupicola was rarely trapped in Saskatchewan at Clavet. At Clavet, Bembidion species were mainly active during mid-summer until about the first week in August, when activity, as determined by pitfall captures, ceased abruptly. A small amount of activity resumed in the late fall. Bembidion bimaculatum , collected in small numbers at Clavet, was an exception in that it was active mainly from mid-summer on. The main period of mating for the most abundant species B. obscurellum appeared to be in June, assuming that the observed increased male activity during that period reflected mating activity. Chi-square analysis showed that males were in excess of females (P<0.05) on June 4, June 18 and on July 2; on the latter date, 208 males and 1 13 females were captured. For the other 10 weekly collections so analyzed, the male-female ratio for B. obscurellum did not differ from a Quaest. Ent. 1981,17(3,4) 214 Doane 1:1 expectation. Bembidion quadrimaculatum oppositum was the second most abundant Bembidion species in the field collection in 1975, and in 1976 exceeded B. obscurellum in numbers. Amur a The genus Amara was represented by 18 species in the two habitats; 15 were collected in the field and 17 in the border. Amara lacustris and A. carinata were the most abundant species in the field (Table 2). A. latior and A. fracta were slightly more abundant in the field than the border while, A. quenseli and A. obesa were captured more often in the border. Species such as A. apricaria and A. littoralis were captured regularly, but in low numbers, while others were extremely rare (Table 2). In Alberta, Frank (1971a) found many of the same species, but the most abundant in Saskatchewan, A. lacustris , was rare at Calahoo, and A. carinata very abundant at Clavet, was not found at the Alberta site. Amara torrida, common in Alberta, was extremely rare at Clavet. The seasonal activity peaks of most of the Amara species occurred in late summer and fall, the same general pattern observed by Frank (1971a). However, there was some variation from this predominant activity period exhibited particularly by A. fracta. In 1975, A. fracta was active mainly from September to the close of the season, but in 1976, peak captures occurred in April and again in midsummer, with relatively few captures in the fall. Harpalus Nine species of Harpalus were trapped, but of these, only H. herbivagus was abundant in collections; it was taken regularly in both the field and border habitats. Lindroth (1968) considered this species rare on the prairies where he considered Harpalus pleuriticus Kby.. more common. Frank (1971a) found H. pleuriticus second in abundance to H. amputatus but did not capture H. herbivagus. Harpalus herbivagus had a long period of activity during the season with several pronounced peaks that varied somewhat in the two seasons. In 1975, this carabid was caught throughout the early part of the season in low numbers, and after July, captures per week steadily increased to a peak about the* third week in September. In 1976, however, it was abundant in field traps from April to mid-July and was then captured in low numbers to the end of the season. In the border, the same general pattern was observed, except that fewer beetles were trapped, and the peak captures were observed about mid-June. Teneral H. herbivagus adults were observed from mid-September until the end of the season. Pterostichus Pterostichus lucublandus was the most abundant of the five species trapped (Table 2). This is an extremely widespread species described as transamerican (Lindroth 1966) and extending south and east at least to North Carolina (Kirk 1971b). At Clavet it was found from early May to October in both the field and field border. Peak activity occurred from about the end of May until the first of July and then again from about the first week in August until late September. In general, the peaks of activity agreed very closely with the pattern of activity described by Kirk (1971b). Frank (1971a) concluded that this species was only active from May to July, but noted a second peak one year. However, Kirk’s observations and those made at Clavet confirm the existence of two activity peaks. The second peak is at least partially composed of young adults, many of which were teneral in September at Clavet. Adults trapped in the fall may be a mixture of newly emerged beetles and those of the previous generation (Kirk 1971b). Ground beetles (Coleoptera: Carabidae) in Central Saskatchewan 215 Agonunt Although five species of Agonum were trapped, few individuals per species were taken. Frank (1971a) found Agonum cupreum Dej. and Agonum placidum Say, two of the same species trapped at Clavet, to be quite common in Alberta. Sex ratios The male:female sex ratio for the 8741 carabids in the four collections, i.e., field and field border 1975, and 1976, were 48:52, 48:52, 53:47 and 50:50% respectively. Rivard (1966), for a collection of 10,000 carabids made over a 3-year period, observed a male:female ratio of 57:43 per cent. Although overall sex ratio of beetles trapped at Clavet was about 1:1, several species did not conform to this ratio (Table 3). Usually males were in excess of females; exceptions were A. carinata for which females were in excess of males in all four collections and P. lucublandus for which females outnumbered males in three of the four collections (Table 3). Disproportionate numbers of A. carinata males and females (overall about 3.5 times fewer males than females) points to unknown characteristics in the life-cycle or activity that appear to differ considerably from most of the species collected. Diversity Diversity and evenness were higher for the seasonal collections from the field in 1976 than in 1975 and, higher in the border collections in 1975 than in 1976 (Table 4). Both years, diversity of the border collections was greater than that of the field collections. Although, total numbers of species in the two habitats ranged from 45 to 66 (Table 1) during the two years, considerably fewer species were captured in any one week (Table 5). More species per week were captured in the border, but more individuals per week were trapped in the field. Higher diversity of carabid species in the field border than in the field appears to be consistent with the higher variety of plant and insect life in the uncultivated habitat. Lewis (1969) observed that insect diversity was highest in hedgerows and that it declined in adjacent crops as the distance from the hedgerow increased. Uncultivated lands therefore appear to be important to the ecology of adjacent crops by providing niches and shelter both for beneficial and pest species (Lewis 1965). Such areas may also provide alternate prey for predators when pest species are not available as prey in the field (van Emden 1965). At Clavet, 63 per cent of all carabid species were captured in both the field and its borders. Therefore, considerable movement likely occurs between the two habitats. This suggests that the uncultivated border may serve as shelter and a source of alternate prey for some species if the field becomes an unsuitable habitat, as for example, during periods of summerfallowing. Numbers of genera and species trapped each year in the field and its border (Table 1) and the average numbers of species trapped in these habitats per week (Table 5) were remarkably constant even for the field. Numbers of species in a habitat tend to reach an equilibrium depending upon the habitat structure (MacArthur 1965). Thus, an approximately constant number of species occur in the field and a few more in the field border as dictated by the available resources and structure of the habitat. Diversity indices of collections from the two habitats provide a convenient measure of carabid species numbers and the relationship between the numbers of species and individuals within species. Although these measures of diversity apply only to collections from the study Quaest. Ent. 1981, 17 (3,4) 216 Doane field and its borders, they will provide baseline information for studies in other fields and areas of Saskatchewan when a similar collection method is used. Potential of carabids as predators Many species of carabids are primarily phytophagous (Johnson and Cameron 1969; Thiele 1977), so that food preferences of dominant species and their response to potential prey require detailed study before the value of dominant species as predators can be assessed. Best and Beegle (1977) investigated the feeding of five species of carabids on a variety of plant and animal food in the laboratory and concluded that dead black cutworm larvae ( Agrotis ipsilon (Hiifnagel)) were preferred by all the species tested. Frank (1971b) investigated the potential of carabids from a cultivated field in Alberta as predators of eggs of the red-backed cutworm ( Euxoa ochrogaster Guenee) and reported that 21 species consumed eggs. These included species of Bembidion (8), Amara (7), Agonum (2), Harpalus (2), P. lucublandus and Trichocellus cognatus Gyll.. Serological tests showed that seven species fed on cutworm larvae or pupae in the field. Natural predation by carabids on wireworm larvae in pastures has also been demonstrated by the precipitin test (Fox and MacLellan 1956). Predatory species included members of the genera Agonum , Amara , Harpalus and Pterostichus. Perhaps the greatest chance for carabid predation in cultivated fields in Saskatchewan would be upon wireworm eggs and newly hatched larvae. The main period of oviposition in the field of the prairie grain wireworm, Ctenicera destructor (Brown), occurs from about the third week in May until mid- June (Doane 1963). These beetles lay eggs under clods of soil in summerfallow fields and in soil cracks (Doane 1967). Such eggs would be exposed to predation when at least four species of Bembidion , i.e., B. obscurellum, B. q. oppositum , B. nitidum and B. timidum are at peak populations and activity levels in the field. Bembidion species fed on eggs of the red-backed cutworm (Frank 1971b) while, in Ontario, B. q. oppositum and B: nitidum were shown to be predators of eggs of Hylemya brassicae (Bouche), the cabbage maggot (Wishart, Doane and Maybee 1956). The role of Bembidion species as potential predators of the egg and early larval stages of wireworms, therefore, is worthy of investigation. ACKNOWLEDGEMENTS Appreciation is expressed to George E. Ball, University of Alberta, for his kind assistance in identification and to Drs. Ball and L. Burgess for helpful suggestions for improving the manuscript. Thanks are extended also to R. K. Allan for valuable technical assistance, and to summer students, Margaret McCrimmon and Rozanne Tennent, for much of the sorting and pinning of collections. REFERENCES Best, L.R. and C.C. Beegle. 1977. Food preferences of five species of carabids commonly found in Iowa cornfields. Environmental Entomology 6: 9-12. Doane, J.F. 1961. Movement on the soil surface, of adult Ctenicera aeripennis destructor (Brown) and Hypolithus bicolor Esch. (Coleoptera: Elateridae), as indicated by funnel pitfall traps, with notes on captures of other arthropods. The Canadian Entomologist 93: 636-644. Doane, J.F. 1963. Studies on ovipostion and fecundity of Ctenicera destructor (Brown) (Coleoptera: Elateridae). The Canadian Entomologist 95: 1145-1 153. Ground beetles (Coleoptera: Carabidae) in Central Saskatchewan 217 Doane, J.F. 1967. The influence of soil moisture and some soil physical factors on the ovipositional behavior of the prairie grain wireworm, Ctenicera destructor. Entomologia experimentalis et applicata 10: 275-286. Doane, J.F. and C.D. Dondale. 1977. Seasonal captures of spiders (Araneae) in a wheat field and its grassy borders in central Saskatchewan. The Canadian Entomologist 111: 439-445. Dritschilo, W. and D. Wanner. 1980. Ground beetle abundance in organic and conventional cornfields. Environmental Entomologist 9: 629-631. van Emden, H.F. 1965. The role of uncultivated land in the biology of crop pests and beneficial insects. Scientific Horticulture 17: 121-136. Esau, K.L. and D.C. Peters. 1975. Carabidae collected in pitfall traps in Iowa cornfields, fencerows, and prairies. Environmental Entomologist 4: 509-513. Finlayson, D.G. and C.J. Campbell. 1976. Carabid and staphylinid beetles from agricultural land in the lower Frazer Valley, British Columbia. Journal of the Entomological Society of British Columbia 73: 10-20. Fox, C.J.S. and C.R. MacLellan. 1956. Some Carabidae and Staphylinidae shown to feed on a wireworm, Agriotes sputator (L.), by the precipitin test. The Canadian Entomologist 88: 228-231. Frank, J.H. 1971a. Carabidae (Coleoptera) of an arable field in central Alberta. Quaestiones Entomologicae 7: 237-252. Frank, J.H. 1971b. Carabidae (Coleoptera) as predators of the redbacked cutworm (Lepidoptera: Noctuidae) in central Alberta. The Canadian Entomologist 103: 1039-1044. Holliday, N.J. and E.A.C. Haglev. 1978. Occurrence and activity of ground beetles (Coleoptera: Carabidae) in a pest management apple orchard. The Canadian Entomologist 110: 113-119. Johnson, N.E. and R.S. Cameron. 1969. Phytophagous ground beetles. Annals of the Entomological Society of America 62: 909-914. Kirk, V.M. 1971a. Ground beetles in cropland in South Dakota. Annals of the Entomological Society of America 64: 238-241. Kirk, V. M. 1971b. Biological studies of a ground beetle Pterostichus lucublandus. Annals of the Entomological Society of America 64: 540-544. Lewis, T. 1965. The effect of shelter on the distribution of insect pests. Scientific Horticulture 17:74-84. Lewis, T. 1969. The diversity of the insect fauna in a hedgerow and neighbouring fields. Journal of Applied Ecology 6: 453-458. Lindroth, C.H. 1966. The ground beetles of Canada and Alaska, 4. Opuscula Entomologica, Supplementum 29, 240 pp. Lindroth, C.H. 1968. The ground beetles of Canada and Alaska, 5. Opuscula Entomologica, Supplementum 33, 295 pp. Lloyd, M., J. H. Zar, and J.R. Karr. 1968. On the calculation of information theoretical measures of diversity. American Midland Naturalist 79: 257-272. MacArthur, R.H. 1965. Patterns of species diversity. Biological Review 40: 510-533. Pielou, E.C. 1969. An introduction to mathematical ecology. Wiley-Interscience, New York. Rivard, I. 1964. Carabid beetles (Coleoptera: Carabidae) from agricultural lands near Belleville, Ontario. The Canadian Entomologist 96: 517-520. Quaest. Ent. 1981, 17(3,4) 218 Doane Rivard, I. 1966. Ground beetles (Coleoptera: Carabidae) in relation to agricultural crops. The Canadian Entomologist 98: 189-195. Thiele, H.U. 1977. Carabid beetles in their environments. Springer-Verlag, Berlin, New York. Thomas, D.B. and E.L. Sleeper. 1977. The use of pitfall traps for estimating the abundance of arthropods, with special reference to the Tenebrionidae (Coleoptera). Annals of the Entomological Society of America 70: 242-248. Turner, F.B. 1962. Some sampling characteristics of plants and arthropods of the Arizona desert. Ecology 43: 567-571. Wishart, G., J.F. Doane, and G.E. Maybee. 1956. Notes on beetles as predators of eggs of Hylemya brassicae (Bouche) (Diptera: Anthomyiidae). The Canadian Entomologist 88: 634-639. Ground beetles (Coleoptera: Carabidae) in Central Saskatchewan 219 Table 1. Summary of individuals, species and genera of carabids captured in pitfall traps in a wheat field and its borders near Clavet, Sask. from spring to autumn, 1975-76 Site No. of individuals3 Mean per Mean per trapb No. of genera No. of species 1325 Field 2810 175.8 16 45 Field border 1215 81.0 23 66 1916 Field 2957 184.8 16 51 Field border 1757 117.1 22 66 aTotal captured in 44 and 56 days of trapping in 1975 and 1976 respectively. b16 traps in the field and 15 traps in the field border. Quaest. Ent. 1981, 17 (3,4) Table 2. Species of carabids captured in pitfall traps expressed as number/meter trap circumference/trap day, and per cent of total captures, for collections made in a wheat field and its borders in 1975 and 1976 at Clavet, Saskatchewan1 220 Doane £ £ 6 g ^ .S 03 Xh O T3 S3 i /3 o 2 X B o o ko c3 U _ • O ■ a cs aJ *0 £ £ £ .3 Cd O TD to CO C/3 r- •*— » o X> I 2 O q O 5 o "5 3 § § § 3 73 2 -S 2, ^ o^OQ 3< c> Qp c «3 ^ oq ^ «3 goq £ a X ■S § 8 ■§ ’C § 5 S CQ D 00 cS Cu >< ^•S 3 § $ £ Field-1975 Field Border-1975 Field-1976 Field border- 1976 No./m trap % of total No./m trap % of total No./m trap % of total No./m trap % of total circum./trap captures circum./trap captures circum./trap captures circum./trap captures Ground beetles (Coleoptera: Carabidae) in Central Saskatchewan 221 O VO VO Tt o o d do VO > 03 C/3 u s X > JO G O 3 1 1 2 s 3 O § 03 3 ~3 3 O § X 3 3 3 O 3 o G c 03 -2 03 3 V. 8 "1 « r- 3 s a QQ "§ 5 oq ~o Vj 5 oq 1 V. S c Qj 3 CQ Vj >-N o o 3 OJ Field-1975 Field Border-1975 Field-1976 Field border- 1976 Species No./m trap % of total No./m trap % of total No./m trap % of total No./m trap % of total circum./trap captures circum./trap captures circum./trap captures circum./trap captures 222 Doane cd X5 Os (N o o o o © o »—i r-~ cd *33 in cd -O cd TD "§ Cu C a & £ a cj a 2 <3 S 8 x -I 03 v. C3 <3 C CU a w O C3 <5 O <3 <3 <3 <3 <3 43 V. S*. a C Q « a s d 5 43 o 5 d CD 43 S m I I- cd cd (continued on next page) Field-1975 Field Border-1975 Field-1976 Field border- 1976 No./m trap % of total No. /m trap % of total No./m trap % of total No. /m trap % of total circum./trap captures circum./trap captures circum./trap captures circum./trap captures Ground beetles (Coleoptera: Carabidae) in Central Saskatchewan 223 1 — ( so CO r-H oo r-H SO r-H >o os so 04 *— < © o d d o d d d d d T—f Os oo no to 04 to 04 to 04 r- 1—1 04 o i— i o O O O o o O o O © o o d d d o d o d d d o o CO r-~ o , — i 04 CO o SO d d d d d d CO o 04 H VO as i — i 04 O r-H o CO o O o o o o d d d d d d vo to 04 04 04 d d o 04 «o VO Os Ov t-H O O r-~ O o o o d d d o o d "3- o o 03 T3 oo o o d oo o o o 224 Doane VO r- o ov ^ “ Oh a O, cd S 2 i a a 3 <-> >» .iS cd O T3 O Oh InS> ^ 6^ O Oh Oh cd cd *-* a =3 £ .fa cd CJ TD Oh Oh Cd cd £ a a a >, .i3 cd O T3 O Oh IsP ^ O Oh Cl- cd cd *-< ^ | d' § >> Z u ^ o U Oh m s v a o a . 5 ‘5? X O "§ CN O O tO O % cd w 3 v. "3 OO OV O 'd- d d r- r- o o d d 0 O a 3 §■ 3 a Jo "3 m a 3 | £ § ^ §« b' a, X Q ^ 3 X ^ (continued on next page) Field-1975 Field Border-1975 Field-1976 Field border- 1976 No./m trap % of total No./m trap % of total No./m trap % of total No./m trap % of total circum./trap captures circum./trap captures circum./trap captures circum./trap captures 226 Doane 'si' co d co 04 o d jo T3 ^j- © d OV r-H O 04 o o d d jo cd T3 co co O co O o r- o d 04 O o d o p d vo d co vo co co i-h co odd •O) 04 o co co d vo o o d 04 O o vo o o d 04 O o ov o d 04 O d Jo cd T3 a a S> d X -1 o -s; l~ii &o S oa ^3 .>s § "5 ~o "5 c e> s: W ~ r a '37 | O fc «c> CU r'O V U a O ft, O b Cymindis planipennis Lee. 0.002 0.03 0.007 0.17 (continued on next page) Field-1975 Field Border-1975 Field-1976 Field border- 1976 No. /m trap % of total No. /m trap % of total No. /m trap % of total No. /m trap % of total circum./trap captures circum./trap captures circum./trap captures circum./trap captures Ground beetles (Coleoptera: Carabidae) in Central Saskatchewan 227 uo co r— vo VO < O — ^ d d d VO o m d d vo o o r-~ OV r- (N (N v CS T3 O d SO VO v cd T3 (N >, Si 03 J3 JO So £ B s 8 3 -- £ g ,5 -5 U C/3 U P-i cx, vr (3, ^ S' ^ S' O c-q 3 tq U too s >5 3 -3 Patrobus lecontei Chd. 0.003 0.04 0.009 0.25 0.002 0.03 0.005 (continued on next page) Field-1975 Field Border- 197 5 Field-1976 Field border- 1976 No./m trap % of total No./m trap % of total No. /m trap % of total No. /m trap % of total circum./trap captures circum./trap captures circum./trap captures circum./trap captures Ground beetles (Coleoptera: Carabidae) in Central Saskatchewan 229 cd -a Cu cd cd CN Tf O — < o o o r- C/5 S 2 c c cd TD cd X3 ~ O VO ON o o o o d d -o u Oh a, cd ° oq 0, ^H cd o — <3 3 1 2 I CO C $ -s cd W) rxj u. 0 03 X) 17 >— < 3 a o a Q o -3 f § 03 03 3 3 •Si C 3 -fC 3 § ~ 03 3 03 03 I 03 3 ■3 3 -3 co co O NO o r-~ d 3 3 tj 3 3 Cl, Q J E*5 in OQ 5 H T3 VO c t-~ cd On 1 -S «*H \0 K <3 - £ Table 3. Total seasonal captures of male and female carabids and chi-square analysis testing the hypothesis of a 1:1 male:female ratio for species that comprised one percent or more of the collections made in a wheat field and its borders near Clavet, Sask. 230 Doane VO r~~ On & u* « Id To c £ £ o o •*> Ph 2 o £ M "Id VO C/5 On D T Id 2 £ E NT) r- on 7 o3 ja o T3 "d O £ co u I cd 2 s a> 1) £ Ph 'o t— OO O ON O OO i/d co oo *— i icd co — co d" O co co wn oo oo d VO VO NO OO r- vo VO ON VO VO o o —• r"; CO O CO © VO 3* o QQ QQ 3 3 S 5 3 ‘c: QQ 3 3 S « a .5 5 QQ QCj oq CN v/d vo On CN O CN CN «— i CN d- O CN OO ON ON VO O oo CN —I — * CN VO VO oo d- CN O co d- o CO CO ON — 1 3 3 ^ w S | .3 -3 •S <3 3 .2 a ^ o 3^ No O oo d- OO rH OO d- O >— < On M I — d" <— < CN VO cn d- &5 50 5 a a a “3 a 3 H « a -o 3 No 3 .50 ^ £ 3 •S' §■ ■£ ^3-3 „ No 3 b •S' 5 3 Vo GX 3 3 3 No &o 5 § 3 g 3 O 3, T3 O X> Ph 2 13 2 1/3 2 2 vo 7 10 2 fx , VO r- ON 2 2 2 B , 3 2 rjj 2 2 8 i i £ *x t-~ ON t/j 2 7 2 2 £ 0 ■ ■ k k 1 1 a s « m vo vo r- oo o ,^ oo r- JQ oo so S3 2 O o oo 13 ~o x oo 40 .5 3 *•3 Uh 00 X 00 > '> 00 cfl T3 .C -5 3 *c _c o o co Oh '3 S 'x a c3 8.5. •ts to jO in oq Doane 0wa o 03 00 I I § c - x> X O 2 3 3 'G CX O ^1^8 .a § * *8 S & o, g> < £ £ 3 c/3 H X) 3 D X Q < nJ < C* PU Z (_i i >» 3 D Z* 3 %-» o 3 s 3 *—> ob ■ o> ob ■ ob s 03 X3 >> 03 03 c 3 3 -a G o 8- ’> 3 X < 1 03 ^3 < ~3 1—3 < i £ 3 cx a,) v X) 03 03 <— > XJ 4) £* C3 3 • <6 c =3 i 03 i >> cd £ i 03 c 3 1) w >> i >> cd •—3 -o O a!> G — ( ob 3 C 3 03 o •—3 u ’> £ £ o l mJ *— 3 < |— 3 o 3 03 0 1 5 -—5 T3 < o 3 3 cl ra £ XJ e XJ £ X cd o -3 £ XJ £ cl >3 a So 03 G c c _o #o #o *+3 X2 ■-S "35 35 C/3 c G G cd 3 3 t-4 t- Wh H H H 3 03 03 03 03 c G C 3 3 3 *“ 3 ^3 *— 3 0) 03 >> 03 jd 3 ^3 ,«2 C/3 GO 32 3 3 X> X5 3 3 00 c "C Oh C/3 O 8 8 to summer Ecology of four Patrobus species 245 Table 3. Numbers of adullts of Patrobus stygicus and various habitats in mid-summer P. lecontei pitfall trapped 1969 in P. stygicus Salix Shallow marsh Deep marsh July 18-25 25 21 21 July 25 - Aug. 3 12 4 48 37 25 69 P. lecontei July 18-25 52 15 0 July 25 - Aug. 3 27 21 79 36 0 Quaest. Ent. 1981, 17 (3,4) 246 Carter Figure 1. George Lake field station showing wet habitats and study areas. The station numbers of study areas were designated by the grid reference of the S.W. corner of the quadrant(s) in which each occurred. The habitats in which the study areas occurred and their station numbers were: adjoining the lake, main marsh, Salix and pools in sedge meadows (30, 31, 45 and 46); forest margin (29S), grassy forest clearings (83 W and 84W); small marsh (22N) and sedge meadows (22, 41N). Ecology of four Patrobus species 247 Quaest. Ent. 1981, 17 (3,4) Figure 2. Occurrence of larvae and adults of Patrobus species in wet habitats at George Lake. 248 Carter DECIDUOUS FORESTS, NEAR RIVERS Figure 3. Diagram of the possible phylogeny of the North American Patrobus (Neopatrobus) species, after Darlington (1938). A summary of the geographical distributions and habitat preferences of these species is included. Quaest. Ent. 1981, 17 (3,4) CLASSIFICATION, PHYLOGENY, AND ZOOGEOGRAPHY OF HYDATJCUS LEACH (COLEOPTERA: DYTISCIDAE) OF NORTH AMERICA1 RE. Roughley Department of Entomology University of Alberta Edmonton, Alberta, Canada T6G 2E3 Quaestiones Entomologicae 17:249-309 1981 D.H. Pengelly Department of Environmental Biology University of Guelph Guelph, Ontario, Canada N1G2W1 ABSTRACT The five Nearctic species of Hydaticus Leach are assigned to the subgenera Guignotites Brinck and Hydaticus sensu stricto. These are H. (H.) modestus Sharp, 1882 (=H. rugosus Poppius, 1905 new synonomy; type area- Ust Aldan, U.S.S.R.), H. (H.) piceus Leconte, 1863, and H. (H.) cinctipennis Aube, 1838 which is recognized as a valid taxon. The Nearctic species of H. (Guignotites) are H. (G.) rimosus Aube, 1838 and H. (G.) bimarginatus (Say, 1831). Because of the similarity between H. rimosus (type not located ) and H. bimarginatus a neotype is designated for H. bimarginatus from the LeConte collection. All of these taxa are keyed, diagnosed and described in terms of morphological features of adults, geographical distribution and habitat. Hydaticus riehli Wehncke, 1876 ( type area— Cuba) is considered to be a species of uncertain placement. A reconstructed phylogeny of genera and subgenera of Hydaticini indicated that this tribe is comprised of two genera: Hydaticus Leach and Prodaticus Sharp; and that Hydaticus includes the subgenera: Hydaticus (s. str.), Guignotites Brinck, Hydaticinus Guignot, and Pleurodytes Regimbart, new status. The reconstructed phylogeny of Hydaticini and the species group of Hydaticus s. str. which includes Nearctic species is used in conjunction with other features of these taxa to construct an hypothesis accounting for the differentiation and distribution of these taxa. The hypothesis is that the ancestral stock of Hydaticini is of gondwanian origin, and that present-day Africa has been a zoogeographic centre from which taxon pulses have originated. The Nearctic Hydaticus fauna is comprised of three separate colonizations of North America: via a North Atlantic land bridge (ancestral stock of H. piceus-H. cinctipennis) via Central America after a closure of the Panamanian portal (ancestral stock of H. modestus). Subsequent vicariance of the two ancestral stocks has produced the present pattern. ‘Part of an M.Sc. thesis submitted to the Faculty of Graduate Studies of the University of Guelph by the first author 250 Roughley and Pengelly RESUME Les cinq especes ^’Hydaticus nearctiques sont assignees aux sous-genres Guignotites Brinck et Hydaticus sensu stricto. Ce sont: H. (H.) modestus Sharp, 1882 (= H. rugosus Poppius, 1905, synonyme nouveau; region typique- list Aldan, U.R.S.S.), H. (H.) piceus LeConte, 1863, et H. (H.) cinctipennis Aube, 1838, cette derniere espece etant reconnue comme valide. Les especes nearctiques d’ H. (Guignotites) sont H. (G.) rimosus Aube, 1838 et H. (G.) bimarginatus (Say, 1831). Etant donne ressemblance entre H. rimosus (type non localise ) et H. bimarginatus, un neotype a ete designe pour H. bimarginatus dans la collection LeConte. Une cle d' identification de tous les taxons est presentee, ainsi qu’un diagnostic et une description des caracteres morphologiques des adultes, la repartition geographique et ihabitat de chaque taxon. La position taxonomique r/’Hydaticus riehli Wehncke, 1876 (region typique: Cuba) est consideree comme incertaine. Une analyse phylogenetique des genres et sous-genres d’Hydaticini indique que cette tribu comprend deux genres, Hydaticus et Prodaticus Sharp, et qu 'Hydaticus inclut les sous-genres Hydaticus (s. str.), Guignotites Brinck, Hydaticinus Guignot et Pleurodytes Regimbart, statut nouveau. L’arbre phylogenetique des Hydaticini et des especes ^’Hydaticus s. str. nearctiques est utilise en combinaison avec d’autres caracteristiques de ces taxons pour edifier une hypothese expliquant leur differentiation et leur repartition geographique. L’hypothese est la suivante: le groupe ancestral d’Hydaticini este d’origine gondwandienne, et I’Afrique actuelle a ete un centre evolutif ou des “vagues” successives de taxons ont origine. La faune nearctique ^’Hydaticus est composee de trois “ vagues " distinctes de colonisation de VAmerique du Nord: une qui entra par la connection Nord-Atlantique (ancetres de la lignee H. piceus-H. cinctipennisj, une qui arriva par I’Amerique centrale apres la fermeture de I’isthme de Panama (ancetre de la lignee H. bimarginatus-rimosusj, et une qui penetra par la connection beringienne (H. modestus). Ulterieurement, la vicariance des deux groupes ancestraux produisit le patron de repartition actuel. TABLE OF CONTENTS Introduction . 250 Materials and Methods . 251 Natural History . 254 Taxonomic Characters and Terms . 255 Classification . 257 Genus Hydaticus Leach . 257 Key to the species of Hydaticus of North America . 259 Species of Uncertain Placement . 277 Phylogeny . 277 Zoogeography . 285 Acknowledgements . 287 References . 288 Figures . 294 Index . 313 INTRODUCTION The genus Hydaticus Leach, 1817, as it is now known, occurs on all continents of the world except Antarctica. Zimmermann( 1920) recorded 104 species in his world catalogue, only four of which were recorded from North America. Later authors (Leng, 1920; Blackwelder, 1939; Blackwelder and Blackwelder, 1948) concerned with North American taxa, modified the list Hydaticus Leach (Coleoptera: Dytiscidae) of North America 251 and increased the total to five taxa. The only comprehensive treatments of adults of the world fauna of Hydaticus are those of Aube (1838) and Sharp (1882). The last treatment of Hydaticus in North America, by Crotch (1873), consisted of a literature review and summation. Regional works treating various taxa of Hydaticus were for the Pacific Northwest (Hatch, 1953), California (Leech and Chandler, 1956), Baja California (Leech, 1948), Alberta (Larson, 1975), Utah (Anderson, 1962), North Dakota (Gordon and Post, 1965), Indiana (Blatchley, 1910), Florida (Leng and Mutchler, 1918; Blatchley, 1919; Young, 1954), Maine (Malcolm, 1971), Virginia (Matta and Michael, 1977), parts of the West Indies (Chevrolat, 1963; Young, 1953), and Canada (Wickham, 1895; Beaulne, 1917). Original descriptions, check lists, nomenclatural notes, and natural history notes comprise most of the remaining literature on North American taxa of Hydaticus. Although only a few taxa of this genus are involved, there has been some confusion concerning their proper identities and limits (Wallis, 1939; Leech, 1948; Young, 1954; Larson, 1975). During this work adult specimens of the genus Hydaticus were examined in an attempt to understand specific limits, variation, and distribution. A phylogeny is reconstructed for the genera of the tribe Hydaticini, for the subgenera of Hydaticus and for the species groups which possess Nearctic representatives. MATERIALS AND METHODS Specimens of Hydaticus used in this study were borrowed from a number of institutions and private collections in North America and Europe. In addition, field collecting of specimens was carried out by the senior author in Ontario in 1975 and 1976; in 1978 specimens were examined during an extensive trip to various entomological museums in the U.S.A. and Canada. Some 2014 adult specimens of North American Hydaticus were examined. The collections from which material was borrowed and curators of these collections are listed below in association with abbreviations used in the text. BMNH British Museum (Natural History, London, England; M.E. Bacchus; BMUW University of Washington, Burke Museum, Seattle, Washington 98195; S.A. Rohwer; BPBM Bernice P. Bishop Museum, Honolulu, Hawaii 96818; G.A. Samuelson; CASC California Academy of Sciences, San Francisco, California 94118; D.H. Kavanaugh; CISC University of California, Berkeley, California 94720; J.A. Chemsak; CMNH Field Museum of Natural History, Chicago, Illinois 60605; E.M. Smith; CNIC Canadian National Collection, Agriculture Canada, Ottawa, Ontario, K1A 0C6; A. Smetana; CUIC Cornell University, Ithaca, New York 14853; L.L. Pechuman; DBUM University of Montreal, Montreal, Quebec, H3C 3J7; M. Coulloudon; DEFW University of Minnesota, St. Paul, Minnesota 55101; P.J. Clausen; ELSC California State College, Long Beach, California 90804; E.M. Fisher; EMUS Utah State University, Logan, Utah 84322; W.J. Hanson; FNYC F.N. Young, Indiana University, Bloomington, Indiana 47401; and University of Michigan, Museum of Zoology, Ann Arbor, Michigan 48109; I.J. Cantrall; GWWC G.W. Wolfe, University of Tennessee, Knoxville, Tennessee 37916; ICCM Carnegie Museum of Natural History, Pittsburgh, Pennsylvania 15213; G.E. Wallace; Quaest. Ent., 1981, 17 (3,4) 252 Roughley and Pengelly INHS Illinois State Natural History Survey, Urbana, Illinois 61801; D.W. Webb, including W.U. Brigham collection; ISUI Iowa State University, Ames, Iowa 50010; R.E. Lewis; JCAC J.C. Aube, 283 des Franciscains, Quebec, Quebec; JFBC J.F. Brimley (now part of CNIC); JFMC J.F. Matta, Old Dominion University, Norfolk, Virginia 23508; KSUC Kansas State University, Manhattan, Kansas 66502; H.D. Blocker; LACM Los Angeles County Museum of Natural History, Los Angeles, California 90007; C.L. Hogue; MCZC Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138; J.C. Scott and M. Thayer; MNHP Museum of Natural History, Prague, Czechoslovakia; J. Jelinek; MSUC Michigan State University, East Lansing, Michigan 48824; R.L. Fisher; MZHF Museum of Zoology, University of Helsinki, Helsinki, Finland; H. Silfverburg; NMDC N.M. Downie, 505 Lingle Terrace, Lafayette, Indiana 47901; NMSU New Mexico State University, Las Cruces, New Mexico 88001; J.R. Zimmerman; OSUC Ohio State University, Columbus, Ohio 43210; C.A. Triplehorn; OSUO Oregon State University, Corvallis, Oregon 97331; J. Lattin; PMNH Peabody Museum of Natural History, Yale University, New Haven, Connecticut 06520; C.L. Remington; ROMC Royal Ontario Museum, Toronto, Ontario, M5S 2C6; G.B. Wiggins; SCSC St. Cloud State University, St. Cloud, Minnesota 56301; R.D. Gunderson; SDSU South Dakota State University, Brookings, South Dakota 57006; E.U. Balsbaugh; SEMC Snow Museum, University of Kansas, Lawrence, Kansas 66045; G.W. Byers; SPMC Provincial Museum of Natural History, Wascana Park, Regina, Saskatchewan, S4P 3V7; R.R. Hooper; TAMU Texas A & M University, College Station, Texas 77843; H.R. Burke; UADE University of Arkansas, Fayetteville, Arkansas 72701 ; E.P. Rouse; UASM University of Alberta, Strickland Museum, Edmonton, Alberta, T6G 2E3; G.E. Ball; UBCZ University of British Columbia, Spencer Museum, Vancouver, British Columbia, V6T 1 W5; G.G.E. Scudder; UCDC University of California, Davis, California 95616; R.O. Schuster; UCSE University of Connecticut, Storrs, Connecticut 06268; R.E. Dubos; UDCC University of Delaware, Newark, Delaware 19711; P.P. Burbutis; UGIC University of Guelph, Guelph, Ontario, NIG 2W 1 ; D.H. Pengelly; UICU University of Illinois, Urbana, Illinois 61801; J. Sternberg; UMDE University of Maine, Orono, Maine 04473; E.A. Osgood, Jr.; UMRM University of Missouri, c/o S.O. Swadener, INHS; UCRC University of California, Riverside, California 92502; S I. Frommer; USNM United States National Museum, Washington, D.C. 20560; P.J. Spangler; UVCC University of Vermont, Burlington, Vermont 05401 ; R.T. Bell; UWOC University of Western Ontario, London, Ontario, N6A 3K7; W.W. Judd; WEHC W.E. Hilsenhoff, University of Wisconsin, Madison, Wisconsin 53706. Hydaticus Leach (Coleoptera: Dytiscidae) of North America 253 Standardized techniques were used in the dissection of the copulatory apparatus of adult males. The technique used for preparation of the aedoeagus was that of Smetana (1971, pp. 10-11). Illustrations of male copulatory apparatus were drawn to scale using a Bausch and Lomb microprojector. Figures of the dorsal view of the left half of the body, and hind leg were drawn using a Leitz Wetzlar binocular, stereoscopic microscope equipped with a 20 x 20 square, ocular reticule. All illustrations are oriented with the anterior end of the specimen toward the top of the figure. Scanning electron micrographs of various body parts were taken with an “ETEC autoscan” after plating the structures with gold. Measurements of body proportions were taken using a stereoscopic microscope fitted with an ocular reticule. Measurements were taken only when both end points of the structure were contained within the grid and were in sharp focus. Only mature specimens, in which elytra were joined completely along the suture and in which the head was drawn tightly into the pronotum, were used for measurements. The largest and smallest specimens of each species were chosen for the range of measurements. Measurements were made in terms of whole or parts of squares of the reticule and converted to millimeters. In addition, length of the median lobe was taken for adult males of each species. Samples from various localities were selected for measurement on the basis of available material. In general, 10 males and 10 females were measured for each locality; however, for two species this was not possible, and most of the appropriate specimens were measured. Two measurements were made on each specimen of the sample; viz. total length of body (TL) and greatest width of body (GW). A detailed description of the genus lists characteristics common to all species considered in this study. The following information is provided for each species treated: citation of the original description and references to other published taxonomic treatments, discussion of type material, summation of diagnostic characteristics, short descriptions of adults that accentuate the distinction among various species, notes on variation, notes about natural history based on either published records or field observations, distribution, chorological relationships, phylogenetic relationships, and material examined. All locality records and the general Nearctic distribution given for each species were based on specimens examined. Some literature records could be based on misidentifications but they have been recorded for those areas outside the geographic limits of this study. In the locality lists of specimens examined, the following information is provided: Country, State or Province, County or Regional Municipality or District, locality, date, collector, collector’s remarks, acronym of the collection from which the specimens were received, and the number of specimens received from that collection. Data were recorded as they appeared on the label except that the first two digits of the year were deleted and the months were converted to Roman numberals (i.e., 21 May, 1967 became 21.V.67). In a series of specimens from a single locality, the label with the most complete data for that locality was the one recorded. Complete locality records for such common species as H. modestus, H. piceus , and H. bimarginatus are omitted. Copies of these records are stored at the Department of Environmental Biology, University of Guelph and at the Strickland Museum, University of Alberta. Persons desiring copies of these lists should contact the authors or these Departments. The greatest problem facing any taxonomist is determining the best manner in which to recognize a biological species from series of museum specimens. Various techniques are used for inferring species status by means of clues supplied by the museum specimens. Our Quaest. Ent., 1981, 17 (3,4) 254 Roughley and Pengelly procedure involved grouping specimens by similarity of the male aedoeagus, especially the median lobe. The underlying assumption is that dissimilar median lobes of various phena, especially if correlated with consistent differences in other characters, represent reproductive isolating mechanisms necessary for fulfilment of the biological species definition (Mayr, 1963, pp. 91, 663) (for a more detailed discussion, especially of exceptions, see Erwin, 1970; Noonan, 1973; Whitehead, 1972). The next step involved associating females with males of that same taxon. Each species of Hydaticus of North America is sympatric with at least one other species in some part of its range. This area of sympatry was considered a test of species status versus subspecific status (Mayr, 1969; Ross, 1974). If no intermediate specimens were observed, species status was considered to be confirmed. The areas of sympatry are indicated under the treatment of each taxon in the section on chorological relationships. In practice, identification of North American specimens of Hydaticus is not difficult once the specific limits of variation are understood. Each taxon has distinctive features which should allow ready assignment of specimens. The subgenera of Hydaticus proposed by Guignot (1950) are used herein because they are monophyletic and represent natural evolutionary units. There is the probability of increased information from such a taxonomic division, and subgeneric characters are used within the key to species. The characters separating two subgenera were recently clarified, expanded, and strengthened by Franciscolo (1968). NATURAL HISTORY Very little is known about natural history of North American species of Hydaticus , but European authors have contributed more to the knowledge of natural history of this genus. Galewski (1971), in his work on Dytiscidae of Poland, recorded the preference of Hydaticus for ditches, pools, and ponds with abundant vegetation, a deep layer of plant debris or detritus and low acid content. Adults frequent temporary pools in early spring to take advantage of available food, expecially the larvae of snow-melt mosquito populations, but they spend the summer in more permanent waters. The breeding season of the European species is from late spring to early summer, mainly in May and June. This breeding season often coincides with development of preferred oviposition sites, which are aquatic, vascular plants, belonging to genera such as Alisma, Iris, and Typha. Adult females of Hydaticus possess genital valves which are long, narrow, and knife-like, but without serrations (Boving 1912; Galewski (1971)). This piercing type of ovipositor is well adapted for endophytic, egg deposition (Balduf, 1935). Larvae of Hydaticus are good swimmers, when compared to most other larvae of Dytiscidae, because of the dense fringe of swimming hairs on all legs and on the last two abdominal segments. The three larval stages are passed within the littoral zone of the breeding areas, although some individuals venture into open water. The large tracheae of larvae are filled with air; consequently larvae float to the surface when not swimming. Vegetation is used as resting and feeding sites, but the larvae are awkward when crawling. The prey of larvae are small naiads of Odonata and Ephemeroptera, and larvae of Trichoptera and Diptera (Galewski, 1971). Pupation occurs on land near the larval habitat in the typical dytiscid manner. In England, Balfour-Browne (1950) recorded emergence of adults from pupae during August and September. Many newly emerged adults invade temporary pools in late summer and autumn (Galewski, 1971). Adults overwinter under leaf litter and moss on land, often a considerable Hydaticus Leach (Coleoptera: Dytiscidae) of North America 255 distance from the nearest body of water (Galewski, 1964, 1971). There are few known natural enemies of Hydaticus. The gregarine, Bothriopsis histrio was recorded from individuals by Balduf (1935). Pujatti (1953) recorded the metacercariae of the trematode genus Lecithodendrium. The chalcidoid Mesotocharis syclospila Forst. (Hymenoptera: Eulophidae) parasitized eggs of Hydaticus within the stems of Alisma when that part of the plant was above water (Balduf, 1935). TAXONOMIC CHARACTERS AND TERMS Sexual dimorphism. - Adults of Hydaticus possess five main characters which are sexually dimorphic, other than those of the male and female genitalia. 1) In the North American taxa studied, female specimens are, on average, shorter and narrower and have a general outline which is more symmetrically oval than that of males. 2) Male specimens are smooth except for two rows of widely spaced, shallow punctures on each elytron. Most females have sculpturing on the prothorax and/or elytra. The sculpturing or rugosity varies from shallow, widely spaced, longitudinal depressions to deep, closely spaced, irregular folds or furrows. Rugosity varies markedly within and among populations; general trends are noted under each species description and discussion. 3) Protarsal claws of males are longer and are bent more sharply at the base than are their female counterparts. Mesotarsal claws are of different lengths and differ in amount of curvature between male and female specimens. Unlike other genera of Dytiscidae, claws of males differ only slightly among taxa. 4) The three basal tarsomeres of the forelegs of males are expanded into circular acetabula (Fig. 2), and posses sucker-like setae which adhere to the pronotum of the female during copulation. Four large suckers are located on the basal segment and these are supplemented by 16 to 18 smaller suckers. 5) The three basal mesotarsomeres of adult males are expanded into elongate, oval acetabula. Mesotarsal suckers are of about equal size inter se. Eight suckers are on the anterior side of the tarsal midline and seven on the posterior side (Fig. 3). The number of suckers on each acetabulum is variable and is difficult to ascertain on many specimens as the suckers are easily broken. Students of Dytiscidae (Wickham, 1893; Chatanay, 1910; Balfour-Browne, 1940, 1950; Larson and Pritchard, 1974) have studied adaptations within the family. Their works indicated that the form of the front tarsi of adult males and the arrangement and number of sucker-like setae were diagnostic for higher taxa. Within the North American males of Hydaticus this character was uniform yet varied. The same basic arrangement is found throughout; however, arrangement and number of modified setae varied between legs of an individual and within and among species. These were without diagnostic value in the taxa studied; however, trends in the number of suckers and their size and pattern are used as a phylogenetic character. Head. - In Nearctic Hydaticus , colour pattern and the presence or absence of large spots or maculae on the frons is an important diagnostic character. The frons varied in colour from a black band against a yellow ground colour to reddish-brown throughout, with infuscated areas. Pronotum. - Colour of pronotum varied from yellow to black. In most species there is a basal, black band of variable width. This band is useful in recognition of some specimens of H. rimosus because it extends to the anterior margin in the form of a wide band. H. piceus and H. cinctipennis specimens have a more or less unicolourous pronotum except for discal infuscations. Curvature of the posterior margin of the pronotum is useful for separation of subgenera. In adults of subgenus Guignotites, the posterior margin is sinuate. The portion along the midline is extended posteriorly, and the portions lateral to the midline are shortened, but the postero-lateral corners are recurved into an acute angle (Figs. 37, 43, 48, 49). In adults of Quaest. Ent., 1981, 17 (3,4) 256 Roughley and Pengelly the subgenus Hydaticus, the portions lateral to the midline are not as conspicuously shortened in length and the postero-lateral corners of the pronotum are truncate (Figs. 7, 8, 9, 22, 30). Metatarsal claws. - Shape of the claws of the hind legs is an aid in distinguishing between some males of the two species of the subgenus Guignotites. In H. bimarginatus , the anterior (shorter, inferior) claw is deflected downward at the apex (Fig. 44), whereas in H. rimosus it is not (Fig. 55). This character was considered unreliable because of the possible effect of abrasion on shape of the apex of the claws. Some males of H. bimarginatus lacked the apical deflections. The relative length and shape of the metatarsal claws are given in the description of each taxon. Metatibial chaetotaxy. - Balfour-Browne (1950) emphasized the use of arrangement, number, and size of spines on the legs for separating the higher taxa of Dytiscidae. Guignot (1950) and Franciscolo (1968) used chaetotaxy of metatibiae for separating the subgenera of Hydaticus. In North American specimens, the row of spines on the posterior (upper, nearest the venter) side of the metatibia is either in a straight line and parallel to the outer margin of the tibia (subgenus Hydaticus , Figs. 10, 23, 31); or the row is curved inward basally, and thus does not form a parallel line with the outer margin (subgenus Guignotites, Figs. 38, 50). Within the subgenus Guignotites, the number of large spines on the anterior (lower) side of the metatibiae is important in separating the two species. These spines were broken off in some individuals and if so, the large pits where the spines originated were counted. The most reliable way to determine number of spines per tibia is to average the number of spines on both metatibiae of the same specimen. The number of spines varies between specimens and between metatibiae of the same specimen, but the specific limits do not overlap and they are in accord with other taxonomic characters. In H. bimarginatus there are a number of smaller spines basally which were not counted. Under each species in which the lower, metatibial spines were used for diagnosis, a table is provided of the average number of spines per metatibia from the specimens used for the descriptive measurements. Metasternum. - The metasternum of dytiscid adults is displaced by the anterior extension of the hind coxae, resulting in the formation of a relatively narrow, transverse sclerite which is indented on each side of the median portion. The area lateral to the median indentation is termed the “metasternal wing” (Balfour-Browne, 1950). The straight anterior margin of the metasternal wing is a diagnostic characteristic of members of the genus Hydaticus. Dorsal colour pattern. - Franciscolo (1968) suggested that colour patterns in the subgenus Guignotites were useful in the recognition of some of the species only. The dorsal colour pattern, however, is useful in the identification of North American material due, in part, to the small number of taxa concerned. Specimens of H. piceus and H. cinctipennis are yellowish to reddish-brown, whereas those of the other taxa are black. Adults of all taxa have the general colour disrupted by various amounts of yellow which varies from an indistinct lateral border in H. piceus, to lateral and/or sub-lateral stripes among other taxa, to a highly developed pattern in some females of H. modestus (Fig. 9). Male genitalic appendages. - The terms used for the genitalic sclerites were those of Edwards (1950), except for the specialized structures of Hydaticus, as discussed by Guignot (1933) whose terms were retained. The male genital capsule, in situ, is rotated 90° to the left( i.e., left paramere is next to sternum 7). This sinistral twist of the genitalic armature is common to all Adephaga, except Gyrinidae (Edwards, 1950; Balfour-Browne, 1950). Components of the genitalic capsule are illustrated in Figs. 4 and 5, and consist of: 1) sternum 8 which is joined to a semi-circular Hydaticus Leach (Coleoptera: Dytiscidae) of North America 257 anterior arch; 2) the basal portion of sternum 9, pleuron 9, and tergum 9 which are fused into a dorso-ventral circular ring surrounding the more posterior structures in repose; 3) an elongate rod or spicule lying on the membrane between the parameres which is possibly a posterior section of sternum 9 separated from an anterior portion; 4) the preputial covering which is strengthened but not sclerotized at the apical portion of a membrane which joins the parameres ventrally; 5) the epipenite, on the dorsal side of the preputial covering, next to the median lobe; 6) symmetrical, hairless parameres which are parallel and of equal length; 7) the median lobe which extends slightly beyond the parameres and articulates with them basally. All of these structures are joined by an ensheathing membrane, except for the dorsal side of the parameres and the median lobe. The aedoeagus is composed of median lobe, parameres, and epipenite. The position and form of epipenite are important for subgeneric diagnoses. Males of the subgenus Hydaticus have the epipenite on the preputial covering (i.e., on the opposite side of the preputial covering to the median lobe, Figs. 4 and 5). In males of the subgenus Guignotites the epipenite is internal, on the preputial covering, next to the median lobe. The transition from external to internal position of the epipenite on the preputial covering reverses the orientation of the median and lateral arms of the epipenite (Figs. 4 and 41). The epipenite in males of the subgenus Guignotites is generally not as heavily sclerotized as that of males of subgenus Hydaticus. In the subgenus Hydaticinus, represented only by the South American H. rectus , the epipenite is still less sclerotized and is inserted into the apex of the preputial covering. The shape of the epipenite and of the apex of the median lobe are important in species diagnosis. The epipenite is a trilobed structure consisting of a median arm and a pair of lateral arms. Length of the lateral arms, when compared to length of the median arm, and shape of the three arms are important in specific assignment. In side view, the apex of the median lobe varies from truncate to acutely angled. The median lobe possesses lateral flanges (Figs. 4 and 5) of a tough membranous substance. The degree of posterior extension of these flanges in relation to the apex of the median lobe is a useful character. CLASSIFICATION Genus Hydaticus Leach Hydaticus Leach, 1817: 69, 72 [nec Hydaticus Schoenherr, 1825 (Coleoptera: Curculionidae)]. Nearctic references only - Aube, 1838: 155 (ex parte). - Crotch, 1873: 403. - Sharp, 1882: 907, 908. - LeConte and Horn, 1883: 67. - Horn, 1883: 380. - Wickham, 1895: 149. - Blatchley, 1910; 232. - Beauline, 1917: 124. - Blatchley, 1919: 314. - Leech, 1948: 419. - Young, 1953: 6. - Hatch, 1953: 235. - Young, 1954: 1 13. - Leech and Chandler, 1956: 332. - Anderson, 1963; 56. - Gordon and Post, 1965: 23. - Malcolm, 1972: 30. - Larson, 1975: 405. - Matta and Michael, 1977: 48. Type-specimens. - Dytiscus transversalis Pontoppidian, 1763; designated by Curtis, 1825: 95 (attributing the species to Fabricius), and by Crotch, 1873: 403. Hope, 1939: 131 cited Dytiscus hybneri Fabricius, 1787( = Dytiscus seminiger Degeer, 1774) as generitype Duponchel, 1845 (6): 728 designated Dytiscus fasciatus Fabricius, 1775 [=Sandracottus fasciatus (Fabricius, 1775)] as generitype but this species was not listed by Leach, 1817 (from Leech, 1948: 419). Diagnostic combination. - Dytiscinae with the following combination of characters: outer margin of metasternal wings straight; outer (shorter) spurs at apex of metatibiae acute; apex of prosternal process rounded; and posterior margins of four basal tarsomeres of hind legs with Quaest. Ent., 1981, 17 (3,4) 258 Roughley and Pengelly dense fringe of flat, golden hairs. Description. — Adults. Size medium (North American specimens, TL from 10.9 to 15.4 mm, GW from 5.9 to 8.0), body form ovate (ratio GW:TL 0.47 to 0.58), outline continuous, widest just behind middle. Colour various, from yellowish-brown to black; surface shining or dulled by rugosity. Microsculpture singular, isodiametric, very fine (difficult to see except at high magnification), small pores scattered throughout. Sculpture of elytron consisting of two linear rows of shallow punctures, not apparent on some females because of deep, irregular wrinkles on elytron and/or pronotum. Head. - Form as in subfamily, except inner anterior margin of eyes slightly emarginate, but not as in Colymbetinae (Balfour-Browne, 1950). Labrum excised at middle, yellowish to reddish-brown, with brush of fine, short hairs. Clypeus indistinct, fused with frons, yellowish to reddish-brown. Frons of most specimens darkened basally, reddish-brown to black, many specimens with yellowish maculae (Figs. 6,47). Occiput short, reddish-brown to black. Palps and antennae yellowish to reddish-brown, segments often infuscated apically. Structure of mouthparts not examined for specific or subgeneric differences. Thorax. - Pronotum with lateral margins acute but not margined; wider at base than at head, with lateral outline rounded; antero-lateral margins produced anteriorly as acute projections; base continuous with bases of elytra, posterior margin more or less sinuate, projected posteriorly opposite scutellum, curved forward lateral to scutellum. Scutellum visible, about 1.5 times as long as wide. Prosternal process with ventral surface convex basally to flat apically, apex broadly rounded, marginal bead complete except apically. Metasternum with short, deep notch for reception of prosternal process; metasternal wings with anterior margin straight, not extended to epipleura, posterior margin sinuate. Metacoxal plates as wide or wider than long (ratio from 1.0 to 0.77); metacoxal processes margined, rounded, separated by posterior incision; metacoxal lines convergent on metacoxal processes and divergent on metacoxal plates, effaced anteriorly. Legs. - Foretarsi of males with tarsomeres 1, 2, and 3 expanded as a broad circular acetabulum (Fig. 1). Tarsomere 5 longest, others subequal in length. Four rows of sucker-like, circular, modified setae or cupules on acetabulum; ventral surface of tarsomere 1 with two rows of four and five suckers each, tarsomere 2 with one row of seven suckers, tarsomere 3 with one row of six suckers. Tarsomere 4 with brush of short, golden setae along antero-ventral margin, tarsomere 5 with two long parallel rows of similar setae ventrally. Anterior claws slightly longer and less robust than posterior claws. Male protibiae and profemora hollowed out ventrally for reception of acetabula of protarsi and protibiae respectively (Fig. 1). Antennal comb and marginal setae of protibiae numerous and longer than those of mesotibiae. Protibiae with pegs along posterior margin near apex, opposed to pits along dorsal surface of tarsomeres 1 and 2 (Fig. 1) (Larson and Pritchard, 1974). Mesotarsi of males with tarsomeres 1, 2, and 3 moderately expanded as an elongate oval acetabulum (Figs. 2 and 3); tarsomere 5 longest, and others subequal, tarsomere 4 of some specimens longer than 2 or 3. Ventral surface of tarsomere 1 with seven suckers, 2 and 3 with four suckers. Acetabulum with long lateral setae (Fig. 2). Mesotibiae and mesofemora not modified for reception of mesotarsi or mesotibiae, respectively. Tarsomere 4 with brush of short golden setae along antero-ventral margin, tarsomere 5 with two long parallel rows ventrally. Claws of middle legs longer than those of forelegs, more evenly arcuate, subequal in length, and of similar shape. Protarsi of females not expanded; tarsomere 5 longest, others subequal. Protibiae and profemora not hollowed out ventrally. Claws subequal in length and similar in shape. Mesotarsi of female not expanded; tarsomere 5 longer than 1, 1 longer than remaining ones which are subequal. Mesotarsal claws longer than, or subequal to, protarsal claws. Otherwise, legs of females similar to those of males. Elytra. - Continuous with outer margin of pronotum at base, widest just behind middle, apices rounded or sinuate (Fig. 18). Shining or dulled by rugosity baso-laterally (Fig. 17). Epipleuron extended to about middle of last visible (6th) abdominal segment. Lateral margins with irregular row of long, fine setae extending from basal third almost to apex. Rows of translucent, rectangular spots beginning marginally in basal third, curving inward posteriorly and in most specimens confluent with division between lateral and sub-lateral stripes if the latter are present. Colour yellowish-brown to black, many specimens with yellow marginal stripes, and/or sub-lateral stripes, latter recurved suturally at base in some taxa, but in adults of most taxa curved away from margin posteriorly. Some specimens with transverse, yellow fasciae sub-basally, with longitudinal vittae from fasciae ending sub-apically (Fig. 9). Taxonomic placement. - Most authors placed Hydaticus in the tribe Hydaticini of the subfamily Dytiscinae, except Balfour-Browne (1950), who included it in the subtribe Hydaticina of the tribe Dytiscini. The only other genus in the tribe Hydaticini is the monobasic Prodaticus with P. pictus Sharp, 1882, known from Persia, Baluchistan, India and Arabia (Zimmermann, 1920). The dibasic Pleurodytes Regimbart with P. dineutoides (Sharp, 1882) known from Java and Borneo (Zimmermann, 1920; Vazirani, 1969) and P. epipleuricus (Regimbart, 1891) known from Tennasserim, Malewan (Vazirani, 1969) is also a member of Hydaticini. However, it does not deserve generic status and Pleurodytes is treated here as a subgenus (NEW STATUS) of Hydaticus s. lat. (see PHYLOGENY). The monobasic Notaticus with N. fasciatus as type species was described as belonging to the tribe Hydaticini. Hydaticus Leach (Coleoptera: Dytiscidae) of North America 259 Spangler (1973) placed N. fasciatus as a junior synonym of Aubehydrus speciosissimus Guignot 1942 and retained it within the subfamily Aubehydrinae. A key to the world genera of Hydaticini was presented by Zimmermann, 1919: 225, and by Zimmermann and Gschwendtner, 1937: 4. KEY TO THE SPECIES OF HYDATICUS OF NORTH AMERICA 1 (0) Pronotum with postero-lateral corners sharply and distinctly acute (best seen in side view), posterior pronotal margin markedly sinuate and recurved laterally (Figs. 37, 43, 48, and 49); metatibia with row of spines on disc of posterior (upper) surface curving inward basally, not parallel to outer tibial margins (Figs. 38 and 50); and epipenite at apex of preputial covering. (Subgenus Guignotites Brinck) 4 1" Pronotum with postero-lateral corners truncate, posterior pronotal margin moderately sinuate but not revurved laterally (Figs. 7, 8, 9, 22, and 30); metatibia with row of spines on disc of posterior surface straight, parallel to outer tibial margin (Figs. 10, 23, and 31). Epipenite anterior to apex of preputial covering (Figs. 4 and 5). (Subgenus Hydaticus ) 2 2 (L) Elytra with broad, lateral, yellow stripes, without sub-lateral stripes but some females vittate (Figs. 7, 8 and 9); colour black; frons black with two, narrow, transverse, yellow maculae (Fig. 6). Epipenite of aedoeagus W-shaped, lateral arms as long as or longer than median arm (Fig. 13); median lobe with lateral flanges extending almost to apex (Fig. 12)1 H. modestus Sharp 7! Elytra with broad, lateral, yellow stripes absent, narrow, indistinct, marginal area present or absent; colour reddish-brown; frons not black, without narrow, transverse, yellow maculae. Epipenite W-shaped, but length of lateral arms 0.66 or less than that of median arm 3 3 (20 Elytra with sub-lateral, longitudinal, yellow stripes, with or without basal fasciae (Fig. 22); form elongate oval, more parallel sided. Epipenite of aedoeagus with lateral arms less than 0.50 length of median arm (Fig. 26); apex of median lobe acute in side view (Fig. 24) H. cinctipennis Aube 3' Elytron without sub-lateral, longitudinal stripes, yellow marginal area present or not, basal fasciae absent (Fig. 30); form oval, sides more arcuate. Epipenite of aedoeagus with lateral arms about 0.66 length of median arm (Fig. 34); apex of median lobe with projection in side view (Fig. 32) H. piceus LeConte 4 (1) Median lobe of aedoeagus angulate at apex in side view (Fig. 39). Anterior (lower) disc of metatibiae with an average2 of more than 10 large spines (x = 1 1.8; max. = 14; min. = 9) (Fig. 45); basal black band of pronotum restricted in most specimens to basal 0.33, extended in few specimens to anterior margin as broad band; elytron with sub-lateral stripes without inward extensions of yellow, transverse fasciae absent (Fig. 37) H. bimarginatus Say 4' Median lobe of aedoeagus truncate at apex in side view (Fig. 51). Anterior disc of metatibiae with average of less than 10 large spines (x = 6.2; max. — 10; 1 While this manuscript was in press, a paper by Nilsson (1981) appeared which suggests that the valid name of this taxon is H. aruspex Clark, 1864. 2 one-half the total number of spines on both metatibiae. Quaest. Ent., 1981, 17 (3,4) 260 Roughley and Pengelly min. = 2) (Fig. 56); basal black band of pronotum often extended to or almost to anterior margin; elytra with sub-lateral stripes, sometimes with inward extensions of yellow, transverse fasciae present in some specimens (Figs. 48, 49) H. rimosus Aube Subgenus Hydaticus Guignot, 1950: 104. - Franciscolo, 1968: 48. Diagnostic combination. - Frons black or infuscated basally, reddish-brown. Frons, when reddish-brown, lacking distinct, contrasting maculae; maculae represented by variable infuscations (Figs. 21 and 29). Frons, when black, with distinct, yellow maculae (Fig. 6). Pronotum with posterolateral corners truncate, pronotal margin moderately sinuate posteriorally but not recurved laterally (Figs. 7, 8, 9, 22 and 30). Row of spines on disc of posterior surface of metatibiae straight (Figs. 10 and 23) or only slightly curved inward basally (Fig. 31). Elytra black to yellowish-brown; stripes absent, or only marginal, or only sub-lateral present; transverse basal fascia present or absent. Elytra, when black, with yellow colouration in form of transverse basal fasciae, and longitudinal vittae in some female specimens (Fig. 9). Epipenite of male genitalic capsule on opposite side of preputial covering to the median lobe (Figs. 4 and 5); heavily sclerotized. Hydaticus ( Hydaticus ) modestus Sharp3 Figs. 4-19. Distribution map, Fig. 20 Hydaticus modestus Sharp, 1882: 650. — Wallis, 1939: 126, 127. -- Hatch, 1953: 235. — Leech and Chandler, 1956: 332. — Anderson, 1962: 73. — Gordon and Post, 1965: — Larson, 1975: 405. Hydaticus americanus Sharp, 1882: 651. — Zimmermann, 1919: 225. - Wallis, 1939: 126, 127. Hydaticus stagnalis , Crotch, 1873: 404. — Horn, 1883: 280. — Wickham 1895: 150. — Blatchley 1910: 232, 233. — Beaulne, 1917: 124. — Zimmermann, 1919: 225, 226 {ex parte). — Zimmermann and Gschwendtner, 1937: 10 {ex parte). — F. Balfour-Browne, 1950: 300 {ex parte). — Zaitzev, 1953: 307 {ex parte). — Watts, 1970: 727. — nec Fabricius, 1787: 191. Hydaticus laevipennis , Sharp, 1882: 651. - Blatchley, 1910: 233. - Beaulne, 1917: 124. - nec Thomson, 1867: 88. Hydaticus rugosus Poppius, 1905: 23, 24. NEW SYNONOMY. Zaitzev, 1910: 44. — Zimmermann and Gschwendtner, 1937: 17.- Zaitzev, 1953:330. Hydaticus bimarginatus , Wickham, 1895: 150. — nec Say, 1931: 5. Notes on synonomy and type material. - Holotypes of H. modestus and H. americanus were examined by Larson (1975) and were not re-examined in this study. The holotype (9) of H. rugosus was borrowed from the Zoological Museum, University of Helsinki, Helsinki, Finland. It is labelled as follows: Ust Aldan; FI. Lena; B. Poppius: 877 (pink label); Mus. Zool. H:fors, Spec, type No. 1953, Hydaticus rugosus Popp. Nomenclature of Nearctic H. modestus and Palaearctic H. stagnalis (Fabricius, 1787) has long been confused. J. Balfour-Browne (1944, p. 355) proposed H. continentalis as a new name for H. stagnalis which was described originally as a member of Dytiscus and thus the latter name is itself a junior homonym of Dysticus stagnalis Fourcr., 1785, which is itself a junior synonym of Dytiscus semisulcatus O. Muller, 1776. Most subsequent European workers — e.g., F. Balfour-Browne (1950) and Guignot (1947), did not accept the name H. continentalis , 3While this manuscript was in press, a paper by Nilsson (1981) appeared which suggests that the valid name of this taxon is H. aruspex Clark, 1864. Table 1. Variation in the length and greatest width of body of selected samples of North American taxa of Hydaticus (Coleoptera: Dytiscidae). Measurements in mm. Hydaticus Leach (Coleoptera: Dytiscidae) of North America 261 oo -s: c IS cu o tp p p in p U r (*5 cd 4) ^ i- , f O E d, £ 6 o3 U £ c/5 00 TD C 23 c/5 £ s o3 c/5 ox) X O s X H H.rimosus Cuba;Bahamas,B-\V.I. <3 9 11.9 11.2-12.3 6.3 6. 2-6. 7 2 6 12.3 11.9-12.6 6.6 6.4-6.9 (continued on next page) Mexico; Guatemala: (Br. <5 15 12.4 11.9-12.7 6.7 6. 3-7.0 Hydaticus Leach (Coleoptera: Dytiscidae) of North America 263 Quaest. Ent., 1981, 17 (3,4) Table 2. The numbers of Hydaticus (H.) modestus Sharp with the development of yellow on the elytra, characterized as vittate (Vj.fasciate (F), and non-fasciate ( N ); and of elytral sculpture, characterized as rugose (R), and smooth (S). 264 Roughley and Pengelly cni D =3 a d Vh cd 0 £ c l-M M II Hydaticus Leach (Coleoptera: Dytiscidae) of North America 265 presumably because the North American H. modestus was, in their opinions, incompletely studied (see F. Balfour-Browne, 1950, pp. 299-300; and Guignot, 1947, p. 228, footnote #2) even though Wallis (1939) had presented evidence that the two were separate. The distinguishing features used by Wallis were that H. modestus is narrower and more parallel-sided than H. stagnalis and males of H. modestus are not vittate as are males of H. stagnalis. The anterior metatarsal claw of H. modestus is proportionately shorter and weaker than the posterior claw when compared to that of H. stagnalis. The elytral apices are sinuate in females of H. modestus (Figs. 9 and 18). All of these differences were confirmed in this study and, in addition, the aedoeagus of males differed consistently. The epipenite of H. continentalis (Fig. 15) has the middle arm much more expanded apically, and the lateral arms are shorter in relation to the median arm, when compared to H. modestus (Fig. 13). The median lobe of male H. continentalis (Fig. 16) is more rounded basally and is much longer (x = 2.9 mm, N = 5) than that of H. modestus males (x = 2.7 mm, N = 12). Thus, the correct name for the Palaearctic form is H. continentalis. Specimens of H. modestus run consistently to H. rugosus Poppius 1905 and not to H. continentalis (= H. stagnalis) in Zaitzev’s key (1953). The type of H. rugosus was examined and found to be within the range of variation of similar rugose specimens of H. modestus from Alaska, Alberta, and the North West Territories. Thus, H. rugosus is considered to be conspecific with H. modestus. This synonomy supports the distinctness of H. modestus from H. continentalis, because H. rugosus was described by Poppius who had an adequate knowledge of H. continentalis. The type of H. rugosus is from List Aldan which is in Yakutskaya region of the Central Siberian Uplands of the U.S.S.R. This specimen is illustrated in Figure 9. Wallis (1939) doubted the validity of H. cinctipennis Aube 1838 and suggested that it was possibly conspecific with H. modestus. However, he believed that there was insufficient evidence for placing the names in synonomy. It would appear that Wallis had not seen Aube’s rather detailed original description of H. cinctipennis as he cited only Sharp’s (1882) subsequent description and the taxonomic notes of Zimmermann (1919). In the present study, H. cinctipennis is considered as a valid and distinct taxon, separate from H. modestus. Blatchley (1910), as noted by Larson (1975), seemed to have assigned vittate females to H. stagnalis and the males and non-vittate females to H. laevipennis. Larson (1975), who examined the types of H. modestus and H. americanus , stated that the latter is a lightly sculptured female of H. modestus which is the valid name because of page priority. Diagnostic combination. - Adults are recognized by the narrow, transverse, yellow maculae on the frons (Fig. 4). They are the only North American members of the subgenus Hydaticus with a black ground colour, lateral stripes, and vittate females (Fig. 9). Median lobe and epipenite of aedoeagus are distinctive. Description. — Length from 12.8 to 15.4 mm, width from 6.8 to 8.0 mm, other measurements in Table 1. Form ovate, moderately convex. General colour black. Head black, except for clypeus and two narrow, yellowish to reddish-brown, transverse maculae on frons (Fig. 6). Pronotum yellowish to reddish-brown except for narrow, transverse, black band at base, band not extending to lateral margins and restricted to basal third (Fig. 7). Elytra piceous to black; lateral yellow stripes distinct, slightly recurved suturally at base, marginal, ending before elytral apex except in very few specimens, appearing divided medially because of rows of translucent rectangles (Fig. 7). Elytra, except for lateral stripes, piceous to black (Fig. 7), or with transverse, basal fasciae (Fig. 8), or some adult females with fasciae and two to 10 longitudinal vittae (Fig. 9), fasciae and vittae yellow to reddish-yellow. Ventrally, prosternum yellow to reddish-yellow; remainder of thorax black; abdominal sterna black, margins of many specimens piceous and with yellow to piceous maculations laterally. Profemora and mesofemora yellowish-red with infuscations, tibiae more infuscated, tarsi darkest. Hind legs piceous to black. Rugosity absent or present, in the form of a few, shallow depressions baso-laterally on elytra or deep, irregular wrinkles on pronotum and most of elytra except area along suture (Figs. 9, 17). Anterior metatarsal claw, bent downward at tip, less than 0.50 length of posterior claw. Median lobe of aedoeagus moderately long (x = 2.7 mm, N Quaest. Ent., 1981, 17(3,4) 266 Roughley and Pengelly = 12), truncate basally, apex notched in side view (Fig. 11). In ventral view, lateral flanges extending almost to tip, apex of median lobe not modified (Fig. 12). Epipenite with lateral arms equal or subequal in length to median arm, expanded medially, thickened and narrowed basally (Fig. 13). Parameres narrow, acute at apex, with narrow, translucent flange on apical third (Fig. 14). j Variation. - Adults of H. modestus vary in the amount of rugosity and yellow coloration on the elytra. Both of these characteristics show patterns of geographical variation. Data for selected samples of H. modestus are presented in Table 2 and shown schematically in Figure j 19. The pronotum and elytra of males are smooth. In females, however, rugosity varies from nil to almost all of pronotum and elytra being covered by deep, irregular wrinkles (Figs. 9, 17). Elytra of females from British Columbia, Washington, Oregon, and California are predominantly smooth, whereas those from Alaska and east of the Rocky Mountains to southeastern Manitoba are consistently rugose in sculpture. Specimens from scattered localities in northern Ontario and northern Quebec could indicate an eastward extension of this form. A few rugose females were seen in population samples from Wisconsin, Minnesota, and Michigan. Females from eastern Manitoba, eastern and mid-western U.S.A., southern Quebec, and southern Ontario are commonly smooth, whereas specimens from the southeastern part of the range are exclusively smooth (Table 2, Fig. 19). Elytral colouration is more complex than is rugosity: females show three states - vittate, fasciate, and non-fasciate; males show two states — fasciate and non-fasciate. The vittate condition is characterized by marginal stripes, transverse basal fasciae and from two, but commonly eight to ten, narrow longitudinal vittae (Fig. 9). The fasciate condition is characterized by stripes and basal fasciae only (Fig. 8). The non-fasciate condition is characterized by marginal stripes only. The degree of concentration of the three female conditions in each geographic area ranges from: vittate — 59% in New England, New York, and Pennsylvania to 80% in Manitoba and Saskatchewan; fasciate — 5% in northwestern U.S.A. and British Columbia to 36% in midwestern U.S.A. ; non-fasciate — 3% in midwestern U.S.A. to 21% in northwestern U.S.A. and British Columbia. The fasciate condition for males ranged from 91% in northwestern U.S.A. and British Columbia to 64% in midwestern U.S.A. (Table 2, Fig. 19). There is some correlation between the abundance of fasciate males and vittate females within areas, throughout the range. In specimens from northwestern U.S.A. and British Columbia 91% of males were fasciate and 74% of females were vittate. About half of males from Alberta were fasciate and about half of females were vittate. In specimens from Michigan, two-thirds of males were fasciate and two-thirds of females were vittate. Natural history notes. - Galewski (1964, 1971) presented evidence that adults of European Hydaticus overwintered out of water as they were found in forest debris or litter. Preliminary evidence for H. modestus indicates a similar pattern; for instance, adults from Framingham, Massachusetts were labelled “by sifting”, and “under stone, high dry hill” and were collected in January, March and April. Specimens from Arlington, Massachusetts were labelled “moss roots” and were taken in March and December. Fletcher and Gibson (1908) recorded specimens from moss but did not give time of year. A specimen was collected 13.ix.79 in leaf litter at George Lake, Alberta approximately 3 m from water line. Flight records are 13.iv.24 at Framingham, Massachusetts; 21.vi.08 at Algonquin, Illinois; 30.viii-3.ix.69 at Chaffey’s Locks, Leeds Co., and 14.ix.73 at Harrow, Ontario; 3.vi.75 at St. Cloud and l.ix. 70 at Itasca State Park, Minnesota, and 29.ix.56 at Creston, British Columbia. Although these records are few, the general indication is of two major periods of flight, spring and fall, perhaps indicating pre- and post-overwintering movements. However, it is also possible Hydaticus Leach (Coleoptera: Dytiscidae) of North America 267 that these beetles were intercepted during flight to temporary ponds or between ponds. A number of specimens were recorded by Hatch (1924) as occurring in beach drift in Charlevoix Co., Michigan. Larson (1975) noted that H. modestus was found most commonly in the forested regions of Alberta. Adults were taken in dense detritus, or emergent vegetation along margins of ponds. The senior author has taken them in similar situations in Ontario. In Mer Bleu marsh, near Ottawa, specimens were collected among emergent Carex, in the drainage ditch of a small beaver pond. Near Moffat, Halton R.M., Ontario, specimens were collected among dead Typha leaves of a permanent woodland pool on 24.iii.76. Andrews (1923) collected adults by dredging aquatic plants at Whitefish Point, Chippewa Co., Michigan. James (1970) included H. modestus in his key to the aquatic beetles of vernal woodland pools in Hastings Co., Ontario. Mature larvae of H. modestus were described by Watts (1970) (as H. stagnalis ) but the adequacy of this description was criticized by Galewski (1975). Records for teneral adults are: 24.vi.33 at Beach, Illinois; lO.viii at L’Anse, Michigan; 14.vii.27 at Long Beach, Long Island, New York; 12.vii.56 at Chatterton, 2.vii.l9 at Port Stanley and 4.vii.57 at Spanish, all in Ontario. Larson and Pritchard (1974) suggested that, for the males, there was a stridulatory function for the dorsal surface of the expanded protarsi and the accompanying pegs on the protibiae. Distribution. - Map, Fig. 20. In North America, this species is transcontinental and ranges as far south as California, Missouri, and the New England states. Zaitzev (1953) recorded H. rugosus from Ust’-Aldan, Yakutsk, Bulun, and Irkutsk, in Siberia, U.S.S.R. As well as the type of H. rugosus , the following specimens from the U.S.S.R. were examined: Sibir oc., Barnaul am Ob, v.20, leg. Babiy, (CUIC), 2; Barnaul am Ob, 25.iv.18, Babiy, (CUIC), 1. Chorological relationships. - H. modestus is sympatric with H. piceus in northeastern U.S.A. and from Alberta to Quebec in Canada. Possibly, H. modestus is sympatric with H. continentalis in Siberia, U.S.S.R. (Zaitzev, 1953). Phylogenetic relationships. - H. modestus is more closely related to a complex of Palaearctic Hydaticus , centering around H. continentalis, than it is to other Nearctic Hydaticus. Material examined. - The number of specimens examined was 378 66, and 500 99. Hydaticus ( Hydaticus ) piceus LeConte Figs. 1, 2, 21-27. Distribution map, Fig. 28. Hydaticus piceus LeConte, 1863: 23. - Crotch, 1873: 404. - Sharp, 1882: 653. -- Wickham, 1895: 150. - Beaulne, 1917: 125. Hydaticus piceous Gordon and Post, 1965: 23, misspelling. Notes on synonomy and type material. - LeConte, in the original description, gave a general description of the distribution as “Middle States and Canada”, but did not give a specific locality. The holotype is a male in the Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts. It is labelled as follows: 111.; Type 6086 (red label). Diagnostic combination. - The predominantly reddish-brown colour, absence of elytral stripes, shape of the median lobe and epipenite of the aedoeagus of males distinguish members of this species. Adults of H. piceus have been confused, in collections, with those of H. cinctipennis. The former are distinguished by smaller size, more oval shape, and lack of sub-lateral stripes and basal fasciae. Quaest. Ent., 1981, 17 (3,4) 268 Roughley and Pengelly Description. — Length 11.6 to 14.0 mm, width 6.6 to 7.9 mm, other measurements in Table 1. Form oval, moderately convex. General colour reddish-brown. Head yellowish to piceous posteriorly; infuscation of frons generally in form of inverted “Y” (Fig. 21), but in some specimens joined by indistinct, fuscous areas extended inward from eyes in the form of indistinct maculae. Pronotum yellowish to dark brown, lighter laterally, often with baso-central infuscation or blotches (Fig. 22). Elytra yellowish to dark reddish-brown, with small, isolated, obsolete irrorations; irrorations denser basally and laterally; lateral, yellow stripes absent, but lateral, yellowish margin present or absent, when present indistinctly delimited (Fig. 22). Ventrally, prosternum lightest in colour, yellow to yellowish-red; remainder of thorax piceous to black; abdominal sterna intermediate in colour, some specimens with yellow maculations laterally. Profemora and mesofemora of many specimens yellowish with infuscations, tibiae and tarsi usually darker. Hind legs corresponding to abdominal colour. Rugosity, baso-laterally on elytra and laterally on pronotal disc of females, shallow, and in some specimens reduced to isolated, elongated depressions. Anterior metatarsal claw abruptly bent before tip, posterior claw with tip deflexed. Median lobe of aedoeagus moderately long (x = 2.8 mm, N = 10), rounded basally; apex of median lobe, in side view, with projection (Fig. 24). In ventral view, lateral flanges narrow and extending almost to tip, apex of median lobe not modified (Fig. 25). Epipenite with lateral arms about two-thirds of the length of median arm (Fig. 26). Parameres narrow, acute at apex, with narrow, translucent flange on apical half (Fig. 27). Variation. - Colour of adults varies markedly. Many samples from single localities exhibit extremes of colour, although specimens from Ontario and Quebec are consistently darker. Presence or absence of marginal, yellowish area of the elytra is not related to geographic localities. This yellow area contrasts markedly in many adults with colour of the suture but colours evenly intergrade between the suture and the lateral margin. Natural history notes. - Needham and Williamson (1907) recorded adults of H. piceus from a permanent spring-fed pond at Lake Forest, Illinois where they were taken among Typha stalks in water as deep as 1 m. Sherman (1913) supported this record in his discussion of Dytiscidae of meadow ponds. Adults were recorded at large, coloured search-lights at Niagara Falls, Ontario by Stirret (1936). Only two other records of flight are available: St. Paul, 25.vi.21 and Albert Lea, 10.vii.23; both localities are in Minnesota. During the fall of 1975 and spring of 1976 adults were observed near Moffat, Halton R.M., Ontario. The habitat was a permanent woodland pool with deep, organic detritus and diverse flora of aquatic vascular plants. In September and October, 1975, adults of H. piceus, assumed to be newly emerged, were collected among emergent Carex. In April and May of 1976, specimens were collected consistently among Typha stalks. In the laboratory, females oviposited into the leaf petioles of Alisma plant ago-aquatica L. (Alismaceae). Eggs hatched after about two weeks and the larvae were fed on mayfly naiads and mosquito larvae collected from the pond. Larvae of H. piceus are agile swimmers and were observed to frequent open water as well as dense vegetation in the laboratory and in pond situations. Distribution. - Map, Fig. 28. The general range of this species extends from central Alberta and southern Manitoba southward to Missouri, eastward to the New England states and northward into Quebec. The Alberta locality (Sturgeon River at bridge 2mi. N.W. of Calahoo, l.vi.77, K.A. Shaw, in Myriophyllum sp. at river edge, [(UASM), 3<3<3] was unexpected since the most westerly previous record had been Winnipeg, Manitoba. This species was not found during the extensive survey by Larson (1975). Subsequently, specimens from Buchanan, Hudson Bay and Prieceville, Saskatchewan (SPMC) were examined, indicating a sporadic, localized distribution of this species along the southern edge of the boreal forest of western Canada. Chorological relationships. - H. piceus is sympatric with H. cinctipennis in the more southerly part of its range. It is also sympatric with H. modestus in northeastern United States, and in Alberta, Manitoba, Ontario, and Quebec in Canada. Phylogenetic relationships. - This species could be closely related to H. cinctipennis’, however, the shared derived characters uniting these two species are relatively weak. As well, the relationship of these two species to other Hydaticus is obscure. Hydaticus Leach (Coleoptera: Dytiscidae) of North America 269 Material examined. - The number of specimens examined was 169 92 and 206 66. Hydaticus ( Hydaticus ) cinctipennis Aube Hydaticus cinctipennis Aube, 1838: 191, 192. --Sharp, 1882: 651 . — Zimmermann, 1919: 225. -Wallis, 1939: 127. Notes on synonomy and type material. - The type-area specified by Aube was the United States and the Antilles. Mr. J.T. Huber kindly searched for the type of H. cinctipennis in various European museums, including the Aube and Dejean collections of the Paris Museum, the collections of the National Museum of Belgium, Brussels, and those of the British Museum of Natural History, London, but without success. H. cinctipennis has been a source of taxonomic confusion, possibly because of its rarity in collections. Wallis (1939) hesitated to synonomize this name with that of H. modestus because he had not seen specimens which agreed exactly with Sharp’s (1882) description and because Zimmermann (1919) considered H. cinctipennis to be valid. Aube’s original description does seem to apply very well to the specimens studied although the type series was not located. Diagnostic combination. - Large size, predominantly reddish-brown colour, short but distinct, yellow, sub-lateral stripes on the elytra and form of male aedoeagus distinguishes adults of this species from others of the genus in North America. Description. - Length 12.8 to 14.7 mm, width 6.9 to 7.6 mm, other measurements in Table 1. Form ovate, moderately convex. General colour reddish-brown. Frons yellowish-brown to dark-brown posteriorly, infuscation irregular but in form of indistinct maculae posteriorly in some specimens (Fig. 29). Pronotum of most specimens lighter laterally, some with baso-central infuscation (Fig. 30). Head and pronotum of some specimens unicolourous, yellowish-orange. Elytra reddish-brown to dark brown; sub-lateral, yellow stripes distinct, not recurved suturally at base, marginal basally but curved away from margin posteriorly, ending from two-thirds to three-quarters of elytral length; area between stripes and outer margins darker; transverse fasciae present (Fig. 30) or absent at base. Ventrally, prosternum lightest in colour, yellowish to yellowish-brown; remainder of thorax dark reddish-brown to black; abdominal sterna of many specimens darkest, some with yellow maculations laterally. Profemora and mesofemora of many specimens yellowish with infuscations, tibiae and tarsi of same colour or darker apically. Hind legs corresponding to abdominal colour. Rugosity absent, or on females: baso-laterally on elytra and laterally on disc of pronotum, shallow and, on some specimens, reduced to isolated, elongate depressions. Anterior metatarsal claw about 0.40 length of posterior claw, both slightly bent at tip, anterior claw more acutely bent in females than in males. Metatibiae ventrally with row of spines only slightly curved inward at base (Fig. 31), but not as noticeably as in adults of subgenus Guignotites. Median lobe of aedoeagus long x = 3.3 mm, N = 5), broadly rounded basally, apex acutely angled in side view (Fig. 32). In ventral view, lateral flanges ending well before tip, median lobe broadened apically (Fig. 33). Epipenite with lateral arms about same length as median arm but extending apically less than half the length of median arm (Fig. 34). Parameres broad, with wide, translucent, membranous flange on posterior half (Fig. 35). Variation. - Specimens from southern United States (Florida, Georgia, Mississippi) are smaller than more northern specimens (Table 1). Also, the head and pronotum of southern specimens are more uniformly orangish in marked contrast with darker, elytral colour. Elytra of most of the northern specimens are more uniformly coloured with respect to pronotal colour. Transverse fasciae at the base of the elytra were evident in 7 of 45 specimens examined. Fasciae are isolated, pale spots near the base, or full, distinct transverse bands as in Figure 30. Presence or absence of fasciae was not related to the sex of the specimen. Natural history notes. - Immature stages of H. cinctipennis are unknown. Habitat of adults is also unknown. Distribution. - Map, Fig. 28. The general distribution of H. cinctipennis is inferred from scattered localities as being the Atlantic coastal plain from New York south to Florida and west along the Gulf coastal plain to Mississippi, and north in the Mississippi River valley to Tennessee. Quaest. Ent., 1981, 17 (3,4) 270 Roughley and Pengelly Aube (1838) wrote that he had seen a specimen from the Antilles. This is an interesting record since no other North American species of Hydaticus ( s . str.) shows a tendency toward toleration of brackish-water situations which would be useful in colonizing and exploiting aquatic habitats within the Antilles. The only species of Hydaticus known with certainty from the Antilles are in the subgenus Guignotites, and these occur commonly in brackish water. Fleutiaux and Salle (1890, p. 352) provided a possible explanation for this anomaly. Felix Lherminier collected extensively in Guadeloupe and in South Carolina and, upon his death, his collections were sent to Chevrolat and Dupont. The collections, however, were mixed and certain species from the U.S.A. were labelled as if they had come from Guadeloupe. Aube (1838, p. viii-ix) acknowledged use of Chevrolat and Dupont collections. Therefore, it is possible that the specimen(s) Aube recorded from the Antilles actually came from South Carolina, U.S.A. which is well within the known range of H. cinctipennis. Chorological relationships. - This species is sympatric with H. piceus in the northerly part of its range and with H. bimarginatus in the southern and southeastern United States. Material examined. - The number of specimens examined was 23 66 and 22 $2 from the following localities: UNITED STATES OF AMERICA CONNECTICUT. Middlesex Co.: Cockaponsett St. Forest, 24.V.49, G.E. Pickford, fire pond, s. section, (PMNH), 1. DELAWARE. Sussex Co.: Glasgow, 29.vi.53, 30.viii.50, H.E. Milliron, electric light, (UDCC), 2; Rehoboth Beach, -.viii.34, (ICCM), 1. FLORIDA. Baker Co.: near Manning, xii. 30.47, F.N. Young, (FNYC), 1. Walton Co.: near Bruce on Fla. Hwy. 20, x.17.41, F.N. Young, hog wallow, (FNYC), 1. GEORGIA. Clinch Co.: 2.6 mil. W. Clinch Co.-Ware Co. line on Hwy #177, 8.vi.75, G.W. Wolfe, (GWWC), 1. Lowndes Co.: (county record only), v. 13.63, E. Hazard, (OSUC), 1. Pierce Co.: Blackshear, viii. 17.50, P.J. Spangler, (USNM), 1. MASSACHUSETTS, (state record only), (ICCM), 1. Bristol Co.: Dartmouth, 20.x. 06, (MCZC), 1; Fall River, iv. 19.18, N.S. Easton, (CNIC), 1; Freetown, viii. 16. 12, N.S. Easton, (CASC), 1; No. Attleboro, 6.ix.20, C.A. Frost, (MCZC), 1; Westport, iv. 1 8.03, (MCZC), 1. Middlesex Co.: Lexington, 20.iv.30, Darlington, (MCZC), 1. Norfolk Co.: Brookline, (MCZC), 1; Stoughton, -.vii.20, Blake, (USNM), 1. MISSISSIPPI. Jackson Co.: Ocean Springs, vi. 14.31, H. Dietrich, (CUIC), 1. NEW JERSEY. Atlantic Co.: Atlantic City, 7.4.01, (USNM), 1. Burlington Co.: Atsion, vi. 11.45, J.W. Green, (CASC), 1 . Morris Co.: Great Swamp at end of White Bridge Rd., 2.ix.75, G.W. Wolfe, (GWWC), 1 . NEW YORK. Richmond Co.: Staten Island, (USNM), 2, (MCZC), 1, -.vii.91, iv.16.05, -.viii.25, (USNM), 3, iv.16-05, E. Shoemaker, (USNM), 2; Bull’s Head, 1.4.21 (UASM), 1. Suffolk Co.: Long Island, M.L. Linell, (USNM), 1; Forrest Park, v.21.04, (USNM), 1; Orient, ix.3.49, R. Latham, (CUIC), 2; Riverhead, vi.5.36, vi.9.36, R. Latham, (CUIC), 2. PENNSYLVANIA. Somerset Co.: Windber, 8.xi.24, (ICCM), 1. RHODE ISLAND. Newport Co.: Tiverton, v.8.09, (MCZC), 1 . SOUTH CAROLINA. Charleston Co.: McClennanville, 2-3.vi.73, R. Turnbow, lite, (UMRM), 1. TENNESSEE. White Co.: Swamp along Rt. 42, 7 mi. N. Sparta, 19.vi.76, G.W. Wolfe, (GWWC), 2. VIRGINIA. Nansemond Co.: Dismal Swamp, iv.16-17.65, P.J. Spangler, (USNM), 2; Holland, vi.20.55, (JFMC), 1. Sub genus Guignotites Subgenus Guignotites Brinck, 1943: 141, new name for subgenus Isonotus Houlbert, 1934: 124, preoccupied by Lepeletier and Serville, 1828 (Coleoptera: Scarabaeidae). - Guignot, 1950: 104. - Franciscolo, 1968: 48. Diagnostic combination. - Frons piceous to black basally; yellowish anteriorly, with or without distinct maculae (Figs. 47, 36). Pronotum with postero-lateral corners sharply and distinctly acute, posterior pronotal margin strongly sinuate and recurved laterally (Figs. 37, 43, Hydaticus Leach (Coleoptera: Dytiscidae) of North America 271 48 and 49). Row of spines on disc of posterior surface of metatibiae curved inward basally, not parallel to outer tibial margins (Figs. 38, 50). Elytra black; lateral and sub-lateral stripes present in most specimens, fused in some specimens to form a single, wide lateral band. Elytra with increased yellow ornamentation in some specimens (Fig. 48). Epipenite of male genital capsule on same side of preputial cover as median lobe; lightly sclerotized. Hydaticus ( Guignotites ) bimarginatus (Say) Figs. 36-45. Distribution map. Fig. 46. Dytiscus bimarginatus Say, 1831: 5. — Say, 1834: 442. Hydaticus bimarginatus, LeConte, 1869: 556. — Crotch, 1873: 404. — Sharp, 1882: 654. — Blatchley, 1910: 233. — Leng and Mutchler, 1918:89. — Blatchley, 1919: 314. - Young, 1954: 113. — Matta and Michael, 1977: 48. Hydaticus fulvicollis Aube, 1838: 184. Hydaticus rimosusl Young, 1954: 113; nec Aube, 1838: 182. Notes on synonomy and type material. - Say’s type area is “Louisiana” but the types were lost (Le Conte, 1869a, p. VI). However, specimens in the LeConte collection, MCZC, are considered characteristic of Say’s species (Lindroth and Freitag, 1969). A male from the LeConte collection is hereby designated as NEOTYPE (MCZC type number 32443). It bears an orange, circular label (for southern States), and another label “ bimarginatus 4”. It is well within the range of variation described by Say. It lacks the left mesotibia and mesotarsus, and claws of the left metatarsus. A neotype is needed to clarify the concept of this taxon because of the similarity to H. rimosus and because the type of H. rimosus was not found. Aube (1838) recorded the type area of H. rimosus as the United States, but type material was not located. The most commonly cited original description of H. bimarginatus is that of Say (1834). This is possibly because the 1834 description is the one quoted by LeConte (1869b). However, Scudder (1899) discussed a paper written by Say in 1831 which contained the description of Dytiscus bimarginatus (see also Bequaert (1951) and Leech (1970, p. 241, footnote 1). In the original description, Aube suggested that H. fulvicollis could be identical to H. bimarginatus. Subsequent authors have considered the names to be synonymous. Because Aube gave a general description of the distribution as the United States and because H. bimarginatus is the only similar species in the type area, the two are considered synonymous herein, although the type of H. fulvicollis was not located. Specimens from Broward Co., Florida, which Young (1954) identified as MH. rimosus ? ” because of their colour pattern, are similar to a few other specimens from Texas, Louisiana, Maryland, and Virgina. However, shape of apex of median lobe of the aedoeagus and number of large spines on the metatibiae places them within the limits of H. bimarginatus as interpreted here. Young (1954) suggested that H. rimosus and H. bimarginatus could be subspecies. This could be so, for there is extensive overlap in colour pattern and a close similarity in other morphological characters. However, H. rimosus and H. bimarginatus are considered specifically distinct because of subtle, yet consistent, morphological differences. In addition, there was a small area of sympatry within the West Indies. Sharp (1882) noted differences in body shape of H. bimarginatus and H. rimosus. The pronotum of H bimarginatus is more abruptly curved outward at the anterior projections and the elytra are less distinctly widened post-medially. The pronotum of H. rimosus is more linear from the anterior projections to the base and the elytra are distinctly widened post-medially. In dorsal profile the sides of H. bimarginatus are more parallel than those of H. rimosus. These differences are noticeable in the sympatric specimens from Cuba; however, these differences Quaest. Ent., 1981,17(3,4) American taxa of Hydaticus ( Guignotites ) (Coleoptera: Dytiscidae ) 272 Roughley and Pengelly d s o3 £ C/3 6 23 E d U C/3 T3 C 23 2 00 a c ‘B a -5 ui Mexico; Guatemala; Br. Honduras; Honduras Hydaticus Leach (Coleoptera: Dytiscidae) of North America 273 are too subtle to be used as diagnostic characters. Diagnostic combination. - Adults of this species are recognized by combination of the following characters: small size, undeveloped sub-lateral, elytral stripes, average of more than ten large spines on lower disc of metatibiae, and angulate tip of median lobe of aedoeagus. Description. — Length 1 0.9 to 13.1 mm, width 5.8 to 7.2 mm, other measurements in Table 1. Form ovate, moderately convex. General colour black. Clypeus and frons yellowish to yellowish-red, except posterior, transverse black band and antero-lateral projections (Fig. 36), some specimens with maculae on frons enclosed by black areas. Pronotum yellowish to yellowish-red, except for transverse, black band at base; band not extended to lateral margins, band as wide as distance between inner limits of basal, recurved portions of elytral stripes and restricted to basal third in most specimens (Fig. 37), but in some specimens extended as a narrow band to anterior border. Elytra piceous to black; lateral and sub-lateral, yellow stripes distinct; lateral stripes marginal, separated from sub-laterals or joined to them in many specimens, in most specimens originating in basal third and ending sub-apically with yellow lobes (Fig. 37) and/or spots; sub-laterals recurved suturally at base, sub-marginal at base, but curved inward from margin posteriorly, ending from two-thirds the length of elytra to near apex of elytra, medial and post-medial, sutural extensions of yellow absent; yellow transverse, basal fasciae absent. Ventrally, prosternum lightest in colour, yellow to yellowish-red; remainder of thorax black; abdominal sterna piceous to black. Profemora and mesofemora yellow with varied amounts of infuscation, tibiae more infuscated, tarsi darkest. Hind legs piceous to black, corresponding to adominal colour. Rugosity of pronotum of females absent or, in most specimens, in form of isolated, circular areas not in contact with anterior or lateral borders of pronotum (Fig. 43); rugosity of elytra absent from most specimens but in some in form of elongate, shallow depressions baso-laterally. Anterior metatarsal claw about half the length of posterior claw, both claws bent downward at tip (Fig. 44), anterior more so in females than in males. Anterior disc of metatibiae with large spines (Fig. 45), (x = 12.2, max. = 16, min. = 8, N 1 150). Median lobe of aedoeagus short (x = 2.0 mm, N = 10), unevenly rounded basally, apex, in side view, angulate (Fig. 39). In ventral view, lateral flanges ending sub-apically, apex of median lobe not modified (Fig. 40). Epipenite with lateral arms extending about two-thirds the length of median arm, narrowed apically; median arm wide, rounded at tip, thickened medially (Fig. 41). Parameres narrow, acute at apex, with narrow, translucent flange on apical third (Fig. 42). Variation. - Adults are quite variable with respect to form of lateral, and sub-lateral elytral stripes and amount of darkening of pronotum. Most specimens have yellow marginal stripes beginning at about one-third of elytral length and extending to apex or ending sub-apically. Lateral stripes on some specimens have two to four apical extensions of yellow, although these are reduced to isolated spots in a few specimens. Sub-lateral stripes of most specimens have inner margins curvilinear and in some specimens these stripes are abruptly narrowed, usually at about two-thirds of elytral length. Inner edges of sub-lateral stripes are uneven in some specimens but distinct median and post-median sutural extensions of yellow are absent. In some specimens, lateral and sub-lateral stripes are united as a single, broad, lateral band (Fig. 37). Black colour at base of pronotum is relatively uniform in width but varies markedly in length. In most specimens, it is restricted to the basal third (Fig. 37), but in some it extends as much as three-quarters of pronotal length as a broad, semi-lunar area, or as a narrow, black band from wide, basal area to anterior margin. Black area does not extend the entire length of the pronotum as a wide, black band as in some specimens of H. rimosus (Fig. 49). Commonly, the frons is immaculate (Fig. 36), but inward, lateral and central, anterior infuscations enclose indistinctly defied maculae. These infuscations vary in darkness and in very few specimens approach the wide, black area seen in H. rimosus (Fig. 47). Number of large spines on disc of lower surface of metatibiae (Fig. 45) varies from nine to 18, with the average consistently greater than 10 in more than 150 specimens examined. In addition, males with an average of 10 to 12 spines were found to have the tip of median lobe of aedoeagus angulate in side view. Numbers of large spines on left and right metatibiae of specimens used for descriptive measurements are presented in Table 3. Natural history notes. - Young (1953, 1954) noted that most specimens were found in clear temporary fresh-water ponds, but also recorded occurrence of a few specimens in brackish-water habitats. Young (1954) regarded H. bimarginatus in Florida as an erratic Quaest. Ent., 1981, 17 (3,4) 274 Roughley and Pengelly occupant of the lowland and upland regions, but noted that it was more abundant in flatwoods situations. Matta (1973) found adults of H. bimarginatus in woodland pools and in thick vegetation of non-acidic, fresh-water ditches, also beetles were collected in a deep, permanent, sand-bottom pool with no vegetation and in a small bog characterized by large clumps of Typha latifolia L. and Juncus effusus L. in the Dismal Swamp of Virginia. Label data indicate a wide variety of aquatic habitats, from saline (brackish pools, brackish water, salt marsh), to permanent (woods pond, tupelo swamp, sink hole pond), to temporary or disturbed (canal, pool in canal, puddle in forest stream, temporary pond). Judging from the frequency of specimens labelled “electric light”, “black light”, and “black light trap”, ranging from February to November, adults of this species apparently come to light more readily than those of any other North American species of Hydaticus. Eggs, larvae, and pupae are undescribed. Geographical distribution. - Map, Fig. 46. The general range of this species includes the Atlantic and Gulf coastal plains of the U.S.A. (New York south to Florida and west to Texas) and parts of the West Indies. Chorological relationships. - H. bimarginatus is sympatric with H. cinctipennis over the whole range of the latter; and with H. piceus in the northern part of the Atlantic coastal plain. It is sympatric with H. rimosus in Cuba and the Bahama Islands, however, no zone of sympatry with H. rimosus was found on the continent. More intensive collecting along either side of the Mexico-U.S.A. border should provide valuable information. Phylogenetic relationships. - H. bimarginatus and H. rimosus are sister species that represent an invasion from the Neotropical realm. Material examined. - The number of specimens examined was 331 66, and 313 29. Hydaticus ( Guignotites ) rimosus Aube Figs. 3, 47-56. Distribution map, Fig. 46. Hydaticus rimosus Aube, 1838: 182, 183. — Chevrolat, 1863: 202, 203. — Sharp, 1882: 654. Notes on synonomy and type material. - Aube’s type of H. rimosus was not located. In his original description he stated that H. rimosus was found in Mexico and the Antilles. He included a possible record for Paraguay. As interpreted here, H. rimosus does not occur in continental U.S.A. H. stagnalis of Horn (1894) and H. bimarginatus of Horn (1896) from San Jose del Cabo, Territory of Baja California, Mexico are most likely variants of H. rimosus with less developed markings of the sub-lateal stripes (Leech, 1948). Diagnostic combination. - Adults are recognized by combination of: small size, often highly developed sub-lateral elytral stripes, average of less than ten large spines on lower disc of metatibiae, and truncate tip of median lobe of aedoeagus. Description. — Length 11.1 to 1 3.7 mm, width 6.2 to 7.0 mm, other measurements in Table 1. Form ovate, moderately convex. General colour black. Head black except clypeus and two, large, oval maculae on frons, yellowish to yellowish-red (Fig. 47). Pronotum yellowish to yellowish-red, except transverse, black band at base; band not extended to lateral margins, in most specimens as wide as distance between inner limits of basal, recurved portions of elytral stripes; in many specimens extending anteriorly as a broad, black band (fig. 49), and in some specimens enclosing a broad area of anterior margin. Elytra piceous to black; lateral and sub-lateral yellow stripes distinct, lateral stripes marginal, separated from sub-laterals in some specimens or both stripes joined, originating in basal third in most specimens and ending sub-apically with yellow lobes (Fig. 48) and/or spots; sub-lateral stripes sub-marginal at base, recurved suturally, and curving away from margin posteriorly (Fig. 49), ending from two-thirds the length of elytra to near apex, inner edges of sub-lateral stripes undeveloped (Fig. 49), or with sub-basal, inward extensions connected with basal, transverse fasciae, Hydaticus Leach (Coleoptera: Dytiscidae) of North America 275 and in a few specimens with medial and post-medial lobes of yellow present as only isolated spots, lobes as wide as stripe in most specimens (Fig. 48); yellow, transverse, basal fasciae present (Fig. 48) or absent (Fig. 49). Ventrally, prosternum lightest in colour, yellowish to yellowish-red; remainder of thorax black; abdominal sterna piceous to black. Profemora and mesofemora yellowish-red with variable amounts of infuscation, tibiae more infuscated, tarsi darkest. Hind legs piceous to black, corresponding to abdominal colour. Rugosity, in most specimens, restricted to pronotum of females; varying from small isolated areas laterally to broad bands from anterior to posterior margins, in most specimens not involving lateral margins. Anterior, metatarsal claw about half the length of posterior claw, bent downward at tip in females but not in males (Fig. 55). Anterior disc of metatibiae with average of less than ten large spines (Fig. 56), x = 6.7, max. = 10, min. = 2. N = 42). Median lobe of aedoeagus short x = 2.2 mm, N = 5), unevenly rounded basally, apex truncate in side view (Fig. 51). In ventral view, lateral flanges ending sub-apically, apex of median lobe not modified Fig. 52). Epipenite with lateral arms sub-equal to or slightly longer than median arm, rounded apically; median arm wide, rounded at apex, thickened medially (Fig. 53). Parameres narrow, acute at apex, with narrow, translucent flange on apical half (Fig. 54.) Variation. - No specimens of H. rimosus included in this study were without distinct maculae on the frons. Most had two anterolateral projections of black from the lower band of maculae (Fig. 47). Size of the black area at base of pronotum was usually greater in H. rimosus than in H. bimarginatus. In most specimens of H. rimosus , the length of the black area is greater than one-half of pronotal length, ending sub-apically in most specimens (Fig. 48), and in some specimens involving width of pronotum posterior to head. Lateral and sub-lateral yellow elytral stripes are extremely varied in size and extent (compare Figs. 48, 49). Specimens with the least developed stripes resemble those of H. bimarginatus. Basal recurved portions of sub-lateral stripes, in a few specimens, have short, posterior prolongations. The area between basal, recurved portions and sub-basal, inward extensions, in some specimens, is also yellow. Sub-basal inward extensions of yellow and basal transverse fasciae vary from complete and distinct, to isolated spots, to obscure yellowish-brown areas. Lateral and sub-lateral stripes, in a few specimens, are joined as wide, lateral stripes, much as in some specimens of H. bimarginatus , however, in H. rimosus the wide stripes are usually accompanied by yellow ornamentation. Apical portions of lateral stripes are dilated in most specimens (Fig. 48). As in H. bimarginatus , the number of large spines on disc of the lower surface of metatibiae (Fig. 56) varies appreciably. However, there are limits between the two species. Although some tibiae have only two, others have 10 spines, the average number of spines per tibia of the specimens observed was consistently less than 10 (Table 3). Natural history notes. - The immature stages of H. rimosus are undescribed and very little is known about the habitat of the adults. Specimens from Nayarit, Mexico were labelled “pool in drying stream bed” and “pool in stream”. Adults were collected at light in late July and early August in Sinaloa, Mexico. Geographical distribution. - Map, Fig. 46. This species is found in the Antilles, Mexico, and southward to at least Honduras. Aube’s record of H. rimosus from Paraguay is based on a single female specimen with a smooth pronotum. This could represent a distinct taxon as Aube (1838: 184) suggested because in this study no specimens of H. rimosus were received from south of Honduras, but of those received from other areas, the pronotum of at least a few females was smooth. Chorological relationships. - H. rimosus could be a southern sub-species of H. bimarginatus as suggested by Young (1954) because differences between adults of the two species are subtle. Intermediate specimens have not been discovered; however, too few specimens were seen from northern Mexico to indicate zones of contact. The two species appear to be sympatric in Cuba and the Bahama Islands. Phylogenetic relationships. - H. rimosus and H. bimarginatus are closely related species that, in aggregate, have their closest relatives within the Neotropical Region. Quaest. Ent., 1981, 17 (3,4) 276 Roughley and Pengelly Material examined. - The number of specimens examined was 37 68, and 35 $2 from the following localities: BAHAMAS, BRITISH WEST INDIES NEW PROVIDENCE: 4 mi. SW Nassau, iv.8.53, E.B. Hayden, (FNYC), 1. BRITISH HONDURAS Punta Gorda, vii.34, (CASC), 1. CUBA CAMAGUEY: Camaguey, xii.20.23, J. Acuna, (UGIC), 2. LAS VILLAS: Buenos Aires, Trinidad Mts., v.8-14,36, Darlington, 2500-3500’, (MCZC), 4. ORIENTE: Cuabitas, Stgo. de Cuba, v.51, P. Alayo, (UGIC), 1; Cauto El Cristo, (Cauto R.), viii.12.36, (UGIC), 1. Rangel Mts., P. de Rio, viii.24.36, Darlington, about 1500’, (MCZC), 1. Upper Ovando R., vii. 17-20.36, Darlington, 1000-2000’, (MCZC), 1. GUATEMALA El Salto, Esquintla, .34, F.X. Williams, (FNYC), 1. Naranjo, El Peten, iii.20.22, H.F. Loomis, (FNYC), 1. Peten Tikal, iv.9.56, T.H. Hubbell & I.J. Cantrall, at light at camp, (FNYC), 1. HONDURAS DEPT. MORAZAN: Zamorano, Esc. Agr. Pan., vii. 6. 48, T.H. Hubbell, 2600’, (carbonal), (FNYC), 1. MEXICO (Country record only), (MUSC), 1. BAJA CALIFORNIA: betw. San Jose del Cabo and Triunfo, (CNIC), 1; San Jose del Cabo, Fuchs, (CASC), 1. CAMPECHE: Hopelchen, 18 mi. E., xi.28.63, K.L. McWilliams, (NMSU), 1. CHIAPAS: Chuatemoc, viii.28.63, K.L. McWilliams, (NMSU), 1. COAHUILA: Matamoros, .v., (CASC), 1. COLIMA: 30 mi. NE Colima, xii.4.48, H.B. Leech, (CASC), 2, (FNYC), 1; Colima, 20 mi. W., vi.6.63, K.L. McWilliams, (NMSU), 2. JALISCO: La Huerta, 6 mi. N., x.25.66, A.H. Smith & J.R. Zimmerman, roadside puddle, (NMSU), 2; La Huerta, 6 mi. N. & 2 mi. E., iii.22.71, J.R. Zimmerman, (NMSU), 1. NAYARIT: 24 mi. N. Acaponeta, vii. 63, F.D. Parker & L.A. Stange, (UCDC), 1; 20.3 mi. W. Compostela, vi. 19.67, A.R. Hardy, (UCRC), 1; Sierra de Zapotan, xi.42, E. Paredes, pool in stream, (CASC), 3; 20 mi. SE Tepic, 23.ix.48, pool in drying steam bed, (CASC), 3. OAXACA: Oaxaca, 20.vii.37, Embury, 3000’, (CASC), 1. SAN LUIS POTOSI: N. Morelos, iii.21.59, (NMSU), 3; Paletla, xii.19.40, F.N. Young, Hydaticus Leach (Coleoptera: Dytiscidae) of North America 277 (FNYC), 2; Tamazunchale, Quinta Chilla Cts., 19.vi.71, A. Newton, Trop. sub-evergreen for., ca. 600’, blacklight trap, (INHS), 2. SINALOA: Coyatitan, 3 mi. E. on road to San Ignacio, iv.9.75, J.R. Zimmerman, (NMSU), 1; 5 mi. N. Mazatlan, vii.22.72, J. & M.A. Chemsak, A. & M.M. Michelbacher, (CISC), 1, vii.25.73, J. Chemsak, E.G. Linsley & A.E. Michelbacher, at lite, (CISC), 1, vii.26.73, J. Chemsak, at lite, (CISC), 1, vii.28.73, J. Chemsak, at lite, (CISC), 1, vii.29.66, J. Chemsak & J. Doyen, white lites, (CISC), 1, 30.vii.64, W.C. McGuffin, (CNIC), 2, viii.5.64, J.A. Chemsak & J. Powell, black and white lights, (CISC), 1, viii.5-7.64, H.F. Howden, (CNIC), 1. TABASCO: Villahermosa, 5 mi. S., viii.26.63, K.L. McWilliams, (NMSU), 3. TAMAULIPAS: ditch N. of Mante, vi.12.60, F.N. Young, (FNYC), 3. VERACRUZ: J.D. Covarrubia, 1 mi. N., viii. 26.62, J.R. Zimmerman, (NMSU), 1; Lake Catemaco, D.C. Robinson, (TAMU), 4; 13 km. WNW Potrero, vii. 16.48, H.B. Leech, (CASC), 1. YUCATAN: Chichen Itza, Xtolok Cenote, vi.28.32, E.P. Creaser, (MCZC), 1; Progreso, 3 mi. S., xi.24.63, K.L. McWilliams, (NMSU), 3. SPECIES OF UNCERTAIN PLACEMENT A female specimen of Hydaticus (G.) grammicus Germar, 1830, is labelled: FLA: Highlands Co.; Archbold Biol. Sta.; 13-X-1964; P.H. Arnaud, Jr., (CASC), Zaitzev (1953) reported this species from southern U.S.S.R., central and southern Europe, Iran, and Japan. The Florida specimen appears to be identical to specimens received from European collections. Most likely it is mislabelled but it could be an isolated occurrence of the species in North America. Leech (1970) discussed some intriguing additions to the California water beetle fauna which could have been introduced by means of aquaria supplies. The name Hydaticus riehli Wehncke, 1876, was not associated with any specimens studied. The type locality is Cuba but the collection of the Academia de Ciencias, Havana, Cuba contained no specimens assigned to this name. Sharp (1882, p. 782) was of the opinion that the species belonged to Thermonectus Dejean. PHYLOGENY Introduction Methods, principles, usefulness, and importance of cladistic analysis for extant faunas are presented by Hennig (1966), Brundin (1966), Ball and Erwin (1969), Erwin (1970), Whitehead (1972), Noonan (1973), and Kavanaugh (1972), 1978). Darlington (1970) and Ashlock (1980) discuss the limitations of cladistic techniques. Cladistic techniques were used to discover relationships but the formal classification is not cladistic. Characters and character states used in phylogenetic analysis are presented in Table 4 and cladograms are presented as Figs. 57 and 58. Each character is numbered and the derived or apotypic state is represented by a filled circle on the cladogram. The ancestral or plesiotypic state is represented by an open circle. Characters which have three states are considered to have both a derived and a highly derived state; the last is designated by a prime mark(')- An attempt to reconstruct the evolutionary history of a group, by cladistic techniques, employs analysis of transformation series (morphoclines) of two or more character states. Quaest. Ent., 1981, 17 (3,4) Hydaticus, and species of Hydaticus (s. str.) (Coleoptera: Dytiscidae 278 Roughley and Pengelly o o3 t-H o ’5L >» o CL. < O 13 >> >, c s- o o3 3 a" o £ c > CO c o o3 u c &- co 2 § fe O J oJ X X . o oo on — < ^ c C/3 X2 X) C/3 g s' C/3 X! s cd t- o c c3 73 cd 3 cd 73 X! N — ' o cd O a cd >1 > cd O > c/3 3 OX) cd a> c _o c 3 o < Quaest. Ent., 1981, 17 (3,4) Character Character state Basis for classification Weight Plesiotypic Apotypic 280 Roughley and Pengelly a cs JH 13 c O T3 -O U) O ~ ■> o3 C/2 i— « O/ U ^ X5 an — h • -h C g M *3 « .5 „ C3 O 0.05). Females specialized on a few common species occurring near dune margins. Surprisingly, dune dwellers are underrepresented among prey used. There is also heavy bias toward males of the dominant species of prey. Males of many species of bees and wasps frequent flowers or positions near flowers while seeking potential mates. Hunting females flew swiftly from the nesting area into vegetation surrounding the dune, where they were quickly lost from sight. In view of the prey captured, I think that two different methods of hunting are employed. The first is to attack and capture any apocritan seen, of appropriate size. This would produce a wide spectrum of prey and would not be biased toward a particular species or sex. A number of apocritans not commonly encountered in large number would be used. A second method of prey capture may be to attack and capture male bees or wasps that trail the female beewolf, as a potential mate. Male bees and wasps which are much larger or smaller than females of P. albopilosus may detect the error at greater distance; those whose females are of similar size and colour would likely continue pursuit and may possibly grapple with the beewolf. The three species of prey captured in the greatest number were males of Aphilanthops frigidus (F. Smith) Colletes sp. and Halictus rubicundus Christ.. Females of the three prey species approximate the size and appearance of female P. albopilosus. Neither the A. frigidus wasps nor the Colletes sp. and H. rubicundus bees were very abundant around the dunes. Prey selected by female P. albopilosus (Empress dunes) is consistent with data from a population examined by Evans (1975) at Albany, New York, but is inconsistent with data from the population examined by him at Roggen, Colorado. This may suggest a genetic basis, which is behaviourally manifested in either, 1) site of prey capture, or, 2) acceptability of potential prey after contact has been made: or, 3) prey selection is the result of opportunism. Quaest. Ent., 1982, 18 (1-4) 106 Hilchie Table 6. Prey records for wasps of Philanthus albopilosus Empress study site. 6 9 Ichneumonidae genus species 1 0 Eumenidae Stenodynerus anormis (Say) 0 1 Sphecidae Aphilanthops frigidus (F. Smith) 0 17 Crabro denningi R. Bohart 0 1 Diodontus sp. 1 3 Mellinus abdominalis Cresson 0 1 Microbembex Monodonta (Say) 1 1 Podalonia violaceipennis (Lep.) 0 1 Tachysphex tarsatus (Say) 1 0 Tachysphex exsectus W. Fox 0 1 10 species of wasps; 30 prey records Halictidae Agapostemon sp. 0 1 Halictus rubicundus Christ 0 23 Halictus sp. 1 0 Lasioglossum sp. 0 2 Sphecodes sp. 0 3 Colletidae Colletes sp. 0 19 Megachilidae Osmia sp. 1 0 7 species of bees; 50 prey records total prey records for each sex 6 74 Change in prey selected, may have occurred as a result of populations of P. albopilosus becoming isolated in different refugia during the Wisconsin glaciation. In a northern refugium (i.e. Nebraska Sand Hills, see below) changes in faunal composition coincident with changing climate may have led to a shift in prey selection. Decreasing abundance and number of suitable species of bees, may have selected populations of P. albopilosus that used increasing proportions of wasps. Selection of sphecids over other groups of wasps may be in part, due to prevalence of dune nesting species with similar environmental tolerances, relative abundance around dunes and a suitable body size. False burrows.. - A false burrow or accessory burrow is defined by Evans (1964b) as “any burrow started from the soil surface in close proximity to the true burrow and made by the same individual”. Evans suggests that these burrows serve to dupe nest parasites into ovipositing in an innapropriate place or to divert parasites into exploring empty holes where there are no hosts. Wasps which maintain or repair false burrows dig them early in the history Table 7. Some taxa of Hymenoptera collected in the Empress area which are available as potential prey* to the beewolf Philanthus albopilosus. Philanthus albopilosus Cresson 107 -a D CH cx CO c o c j— 3 o s- 3 X> c o o X) X) C D C o 3 C/3 C/3 «N c 3 3 * j**! T3 oo ■ — "O T3 o o X) CD a) c a) O D D D C e c c C 3 c 3 c C C C C O O 3 3 3 3 3 CX (D O 3 3 o O 3 3 3 XJ 03 C/3 T3 TD -a ~a T3 T3 T3 C/3 C/3 TD T3 T3 O C C c c 3 £ e C D D C C C < C o o o o O C/3 o o C C O o o ^ rl- (N I OS | — i > 03 GO R R X •S- 3 x> 3 03 CO CX CX CX m 03 -R -R CX CX c/3 H x> -R R X> -2 -R <3 O ft. 2- fft 03 03 >S ^ -R -R X) X) £ £ Quaest. Ent., 1982, 18 (1-4) (continued on next page) Table 7 (continued) 108 Hilchie T3 U CM- o o o c/3 c — — I — < — i | § o §, -3 2 i O Co U »3 ”3 3 3 .3 ■3 3 »3 3 O '‘-■4 §, .2 0« kS £ ** 3 X CL Cl Sr3 Sr3 "3 ”3 O O Vj c £ C C o C/5 o O .5 a S g o3 03 C/5 *— I 03 _Q o *3 CL OX) 03 s O 03 C/5 60 t- 03 cb . -H 03 t ■ 03 C/5 . 03 03 C/5 C/5 CL O C 2 H £ o 03 CL £ C/5 B 33 o X 60 33 C3 o3 < O c "C -a o o c 03 -a CL < >> 33 •d 03 C/5 T3 03 £ C , 03 o Ui C/5 o3 o3 2 o g c E | £ « 3 &-s « 60 ^ O C <» •r D oj b v- -i c u u •C T3 - H o c*h * X) o Quaest. Ent., 1982, 18 (1-4) Symbols used: a. abundant; c. common; nc. not common; # taxa used as prey but excluding prey records. Table 8. Occurrence of accessory burrows in the Sphecidae 110 Hilchie o 0) Cl H 3 CL JO t— £ ^ ^ r-~ no __ ON ON — — CO CO > CO I I 3 to .O l -C> a, a. rn no ON JJ ON > ON > ON > ON UJ uu UU UU S £ fe. S £ 3 ON g - _S 6 I J= — JO 4r O I3 a. CQ B. amoena Handlirsch 0-1 fu - pr - at fc Evans Philanthus albopilosus Cresson 111 of the nest, usually during or after initial closure. The habit is thought to have evolved from quarrying soil for a closure of the nest. Maintaining false burrows would simply entail use of one or a few of the holes as a source of soil for later closures. Maintenance of false burrows may provide continued protection from nest parasites. Females of P. albopilosus construct accessory burrows after completion of the nest. Associated with species which dig false burrows early, but do not maintain them, are various behavioural features which may reduce the effect of nest parasitism(Table 8). Females of Bembix provision their nests progressively. If parasites enter the nest the female may be able to supply enough prey, permitting survival of both larvae and parasites. This tactic may remove some of the selective advantage of maintaining false burrows. Females of one species, B. texana Cresson, construct false burrows during final closure. Debris is removed from the nest and parasites may be swept out during this process. False burrows prepared at this time may offer protection from invasion by parasites, at least until environmental effects obliterate all traces of the burrow. Construction and failure to maintain false burrows in a mass-provisioning species such as P. albopilosus is unexpected. The association of provisioning and nesting behaviours has changed, I propose that false burrows made by P. albopilosus serve as visual markers aiding in initial orientation to the nest in a habitat with an unstable substrate. A large premium would be placed on learning the correct location of the nest without advertising its true entrance during the first orientation flight. This would permit the female to take bearings on distant land marks and visually relate the location of her nest (van Iersel 1964). After orientation, false burrows would no longer have a function, and so they would not be maintained. A similar land mark function has been attributed to the mound building activities of Bembix littoralis Turner. Evans and Matthews (1975) reported B. littoralis females building a mound of soil at the entrance of their burrow, which may serve as a species specific marker identifying the nest location when other species of sand wasps were present. GEOGRAPHICAL DISTRIBUTION AND VARIATION Known Range and Locality Data, Philanthus albopilosus is almost transcontinental in distribution with the most eastern known locality at Albany, New York (Evans 1975) and the western most at Hatton, Washington. The most northern locality is 175 km north of Empress at Rutland, Saskatchewan. It is doubtful that the range extends much further north. The southern known extreme is on Padre Island, Texas, but the range probably extends into central Mexico. Localities of material studied are listed below. Numbers following locality names indicate the number of females examined followed by the number of males examined. Acronyms represent collections from which the material was borrowed. Material examined: CANADA. Alberta: Empress (11 km south), 31, 8 (GH); Medicine Hat, 6, 14, (CNC, UAS); Orion, 0, 2, (UAS). Manitoba: Aweme, 2, 8, (CNC, UCD); Harteny 0, 1, (SEM); Onah, 3, 0, (CNC). Saskatchewan: (see Empress, Alberta) Elbow, 5, 6, (CNC, UCD); Rutland, 3, 8, (CNC, UCD). MEXICO. Chihuahua: Samalyuca, 0, 1, (UCD). UNITED STATES. Arizona: Coconino County: Tuba City, 1, 0, (CUM); Navajo County: Hotenvilla, 1,1, (UCD); Indian Wells (16 miles south), 1, 0, (UCD); Judito Trading Post, 1, 2, (UCD); Joseph City, 0, 1, (UCB); Kayenta (19 miles southwest), 2, 0, (CAS); Colorado: Alamosa County: Great Sand Dunes National Monument, 0, 2, (CSU); Bent County: Caddoa (9 km east), 3, 16, (GH); Hasty, 1, 1, (CSU); Prowers County; Carlton, 1, 3, (CSU); Lamar (10 miles west), 0, 2, (CSU); Weld County: Roggen, 10, 1, (CSU,GH). Idaho (Strandtmann 1946). Iowa: Woodbury County: Sergeant Bluff, 1, 1, (USNM); Sioux City, 0, 1, (USNM). Illinois: Cook County: Chicago, 2, 5, (MCZ). Kansas: Kearny 112 Hilchie County: McKinney Lake, 0, 1, (MCZ); Stafford County: Salt Flats, 2, 4, (CUM, CNC, MCZ). Minnesota: Polk County: Muskoda, 0, 1, (CUM). Montana: 0, 2. Nebraska: Cumming County: West Point, 1 , 2, (USNM). New Mexico: Dona Ana County: Las Cruces, 0, 2, (CAS, SEM); Otero County: White Sands National Monument, 0, 5, (MCZ, UCD); San Juan County: Shiprock (near), 0, 1, (USNM); Socorro County: La Joya Wild Life Preserve, 8, 14, (CSU, MCZ,GH). New York: Albany County: Albany (Evans 1975); Colonie 3, 0, (CSU). North Dakota: Billings County: Medora, 0, 1, (MCZ); Ransom County: McLeod, 1, 1, (UCB), Sheldon, 2, 1, (CUM, MCZ); Richland County: Walcott (11 miles west), 1, 0, (UCD); Williams County: Williston, 1, 2, (MCZ). Ohio (Strandtmann 1946). South Dakota: Fall River County: Hotsprings, 1, 0, (MCZ). Texas: El Paso County: Fabens, 0, 2, (CAS, SEM); Hartley County: Romero, 2, 4, (SEM, UCD); Nueces County: Padre Island, 0, 1, (UCD); Ward County: Monahans State Park, 10, 22, (CSU, MCZ). Utah: Juab County: Eureka, 0, 1, (UCB); Little Sahara Recreation Area, 5, 19, (GH); Utah County: Utah Lake, 0, 1, (CUM): Millard County: Oak City (18 km north), 0, 1, (GH). Washington: Adams County: Hatton, 0, 1, (USNM). In total 234 specimens were examined, 100 females and 134 males. Positions of localities are mapped in Figure 3. Dotted lines define groups of populations which were united and treated as a single population to increase sample sizes for statistical purposes. Populations were grouped naturally using existing geographical barriers as boundaries. Most localities are in semi-arid to arid regions of the Great Plains with clusters in desert regions. A range extension appears to follow the arid interior into the Great Basin. Three eastern localities - Chicago, Illinois: Ohio and Albany, New York - are disjunctive. Beewolves are not found in regions where annual precipitation exceeds 91.5 cm (36 inches). Moisture may affect presence or absence of suitable habitats rather than having a direct influence on the species. Activities by man permit colonization of disturbed sites such as sand and gravel pits and road cuts. Geographical Variation in Colour Pattern Interpretation of species limits for many Philanthus wasps is based on maculation and punctuation patterns in adults. Several closely related species of Philanthus were once thought to be subspecies of P. politus. Recognition was in part based on coloration of tibiae, femora, and punctuation and shape of metanotal lamina. In P. zebratus nitens (Banks) several forms once recognized as distinct species have been united and given subspecies status. Diagnostic characters were maculations and punctures. In this paper specimens of P. albopilosus are placed in recognizable groups based on colour patterns. Results of analysis with a compound character index are shown in Figure 4 for females and in Figure 5 for males. To give some perspective to the index values used, Figure 6 a, b and d illustrate some aspects of a dark northern female. The total compound character index value for the specimen is zero. Figure 7 a, b and d illustrate a southern female which has a total compound character index value of 93. Figure 8 illustrates the typical frontal maculation of the head and an abdomen of a northern male. This male received a total index value of four. Figure 4 illustrates variation in maculation patterns in females. In the north (Areas 7, 8, 9, 10, 11 and 12) index values are concentrated toward the low range, indicating populations comprised of dark individuals. Most of the specimens received index values below 30. A few maculate specimens (Figure 7c) were present but index values did not exceed 70 units. Females from more southern localities (Areas 4,5 and 6) had extreme expansion of the yellow markings and few specimens were assigned index values less than 70. No populations are known to have intermediate index values. Females from mountain regions of the southern United States (Areas 1, 2 and 3) are intermediate in coloration, with index values between 30 and 70. These groups are the most geographically distant populations from those in the northern areas. Number 20 Philanthus albopilosus Cresson 113 Figure 4. Compound character index values for females of Philanthus albopilosus. Area numbers refer to those illustrated in Figure 3. Wasps 114 Hilchie Quaest. Ent., 1982, 18 (1-4) Figure 5. Compound character index values for males of Philanthus albopilosus. Area numbers refer to those illustrated in Figure 3. Philanthus albopilosus Cresson 115 Figure 6. Typical and maculated specimens of female Philanthus albopilosus (northern race). Figures 6a, 6b, and 6d, head, thorax and abdomen of a dark specimen from Empress, Alberta. Figure 6c, head of maculate specimen of the dark race at Medicine Hat, Alberta. Males of P. albopilosus also exhibit some trends in colouration (Figure 5) but these are not as distinctive. Individuals from northern populations (Areas 7 to 12) tended to be darker (Figure 8). Males from southern and western areas (Areas 1 to 6) tended to be pale. There is not a discrete break in index values but a trend to shift to darker forms in the northeast and lighter forms in the southwest. Index values for each area were averaged, female index values were plotted against male index values (Figure 9). The points form loose clusters: northern groups separate together, in contrast to southwestern populations and geographically intermediate populations. The points illustrate a break in variation, with the largest break occurring between the geographically intermediate group and the northern group. The phenotypically intermediate groups are the extreme southwestern populations which may grade into the geographically intermediate group. Figure 7. Typical specimens of females of the light race Philanthus albopilosus manuelito . Figures 7a, 7b, and 7d head, thorax and abdomen of specimen from Monahans, Texas. Figure 7c, head of specimen from Roggen, Colorado. Quaest. Ent., 1982, 18 (1-4) Philanthus albopilosus Cresson 117 Figure 8. Head and abdomen of male Philanthus albopilosus wasp from Empress, Alberta showing typical maculation pattern for most males. Taxonomic Interpretation The beewolf species P. albopilosus includes two major groups of populations. Populations of the dark form appear to form a cohesive geographic unit across the northern United States and southern Canada (Areas 7, 8, 9, 10, 11 and 12). Slight differences among northern populations may be due in part to “ founder effect ” (MacArthur and Wilson 1967), associated with invasion of new areas by few individuals. Physiographic barriers are not evident. However sand dune habitats are islands in a sea of grassland or forest, unsuitable for P. albopilosus. Therefore, these latter habitats serve as barriers. The southern light form is a slightly less coherent grouping. In Areas 2 and 3, an intermediate colour form is present, which is partly isolated by mountain divide systems from populations of the extreme yellow form found in Areas 4, 5 and 6. It appears that there has been a break in gene flow between northern populations and southern populations (Figure 9) for an extended period of time (probably since the Wisconsin glaciation). At present, gene flow is interrupted between some populations in the southern race by physiographic barriers. On the Great Plains gene flow appears restricted between northern areas (7 and 8) and southern areas (5 and 6). No populations of intermediate coloration were found between these areas, even though P. albopilosus is highly vagile and should have dispersed, forming intermediate populations. This pattern suggests character displacement. Character displacement occurs between ecologically similar sibling species, many pairs of which exhibit narrowly parapatric distribution. Lack of a physiographic boundary separating species indicates contact has recently occurred (Vuilleumier 1971). Taxonomists often 118 Hilchie Figure 9. Compound character index value comparisons of means for colour variation in wasps of Philanthus albopilosus. Numbers refer to areas illustrated in Figure 3. experience difficulty deciding between displacement and hybridization (Brown and Wilson 1956). Races of P. albopilosus might be specifically distinct, and thus cryptic species. The premise that the races have come into recent contact implies a barrier at some phase in the species history. Populations in Area 4 may be relatively recently derived from yellow populations further to the north which are in contact with the northern race. Origin from Areas 1, 2 or 3 is less likely because of mountain divide systems. A north-south movement on the Quaest. Ent., 1982, 18 (1-4) Philanthus albopilosus Cresson 119 Great Plains is more probable because of relative lack of geographic barriers. The major isolating event separating the northern from the southern race was probably the Wisconsinan glaciation. Isolation of the lineages may have ocurred late in the Sangamon interglacial stage. This would place the dichotomy at about 60,000+ years B.P. Whitehead (1972) suggested that speciation occurs at a rate of about one dichotomy per 3,000,000 years. On the basis of this hypothesis, races of P. albopilosus may not have been isolated long enough, to have evolved into separate species. Regardless of what actually happened in the past, populations of P. albopilosus may be grouped into definable races on the basis of phenotypic variation and these groups are restricted to geographic regions. For a few populations additional evidence is supplied by variation in aspects of life history and behaviour. Subspecies of Philanthus albopilsus In a species divided into various populations which are more or less isolated over time, when do the variants warrant taxonomic recognition? Many authors have considered the subspecies concept and each has a slightly different idea of what the limits should be; some are for recognition, others are against. I prefer the argument given by Willis (1976)“ ... if subspecies reflect to some degree the actual pattern of variation, as well as being convenient ‘handles’ for reference, their value seems sufficient to justify their recognition”. In philanthines, character differences in head shape, punctation, placement of ocellus and most importantly maculation patterns, provide useful tools for species recognition. In members of Philanthus albopilosus two major groups can be recognized. In certain regions, some specimens feature character states not characteristic of the population mean. Most female specimens (95%) and the majority of males can be assigned to one subspecies group or the other consistantly on the basis of maculation patterns. Philanthus albopilosus albopilosus Cresson Philanthus albopilosus , Proc, Ent. Soc. Phila., 5, 91, 1865 (5). P. simillimus Cresson, Proc, Ent. Soc. Phila., 5, 95, 1865 (2). For a complete synonomy see Strandtmann 1946. Female. - Length 10 to 13 mm. Black, marked with yellow as follows: clypeus 0 to 50% yellow, frons 0 to 40% yellow (Fig. 7a, c), vertex at most five yellow spots, antennae scape black above yellow below, pedicel black, flagellum black above infusciated below, thorax black to black with maculations (Fig. 6b), coxae black or black with apical spot, femora black apices yellow, tibiae yellow with black spot, tarsi pale to testaceous, sterna black may have lateral spots, tergum 1 with two lateral spots, terga 2 and 3 with biemarginate band, tergum 4 with broken band, tergum 5 with three spots, pygidium black. Male. - Length 7 to 1 1 mm. (Fig. 8). Yellow, frons yellow, vertex black, scape black above, pedicel black, flagellum with basal three yellow ventrally dorsally black infuscated toward tip, thorax black, pubesence white, apices of femora yellow, tibiae yellow, tarsi infuscated propodeum black, abdominal sterna black, terga 1,2, and 3 with yellow band, terga 4, 5, and 6 with elongate yellow spot; tergum 7 with or without elongate yellow spot. Type locality. - Illinois. Range. — CANADA: Alberta, Manitoba, Saskatchewan. UNITED STATES: Illinois, Iowa, Montana, Nebraska, New York, North Dakota, Ohio, South Dakota. 120 Hilchie Philanthus albopilosus manuelito new subspecies Female. - Length 10 to 13 mm. Yellow, marked with black, clypeus yellow, frons 10 to 20% black near vertex (Fig. 7a, c), vertex with five yellow spots or solid yellow band, antennae as in Philanthus albopilosus albopilosus , collar yellow, mesonotum with two longitudinal yellow bands, scutum yellow, scutellum yellow, proprodeum large lateral yellow spots (Fig. 7b), apices of coxae yellow, femora 30 to 50% yellow abdominal sterna with lateral spots, with or without medial spots, terga yellow with black along apical and basal margins (Fig. 7d). Male. - Length 8 to 1 1 mm. Similar to Philanthus albopilosus albopilosus , most specimens with additional maculations on scutellum, collar and tubercle of thorax and with or without additional spot on tergum 7. Type material. - Holotype; UNITED STATES: Texas, Ward County, Manahans State Park, 1-2 vi 1974, H. Evans and W. Rubink (female) [USNM]. Paratypes; 9 females, 22 males with same data. Range. — MEXICO: Chihuahua UNITED STATES: Arizona, Colorado, Idaho, Kansas, New Mexico , Texas, Utah. Derivation of specfic epithet. - Manuelito was a Navaho chief, who with the aid of other chiefs, persuaded the United States goverment in 1868 to let the Navaho return to their home lands. The bee wolf Philanthus albopilosus manuelito nests on reservation land. EVOLUTIONARY CONSIDERATIONS Evolution of the Geographic Pattern The occurence of geographical races of P. albopilosus Cresson indicates fragmentation of the ancestral range of the species, and hence, isolation of populations. At least two major refugia seem to be indicated, one in northeastern Nebraska, and the other in southwestern Arizona-New Mexico. Each region contained unique biotic and abiotic features. Divergence in structure and behaviour are related to historical events which might have been causative. Because of similarities between subspecies, they are of relatively recent origin. The Sangamon interglacial stage between Illioinian and Wisconsinan glaciations may serve as a reference point. Flint (1971) argued that conditions during the Sangamon probably ranged from arid in the southwest to cool and moist in the east. Thus climate and plant biomes may have been comparable with today’s. Dune habitats would have been present in many areas where they are currently found. During the early part of the Sangamon interglacial stage, populations of P. albopilosus Cresson were probably widespread (Figure 10a). Toward the end of the Sangamon interglacial (about 70,000 B.P.) the climate began to cool eventually giving rise to the Wisconsinan glaciation (from 55,000 to 10,000 B.P.). Cool wet conditions induced reforestation of much of the Great Plains, and of the northern Great Basin and Mohave Desert (Martin and Mehringer 1965). Increased moisture also caused reduction of sand dunes as ground cover increased, with consequent widespread extinction of and disruption of range of P. albopilosus Cresson. During this isolation, morphological and ecological differentation occurred. In large dune fields, however, porosity of the sand would locally induce aridity and thus blowouts may have developed. Thus populations of P. albopilosus Cresson (s.l.) in the large dune fields may have survived (Figure 10b). Toward the end of the Wisconsinan glacial stage, climate ameliorated, lifezones move north and up mountain slopes. Many stabilized dunes would have become active by loss of their plant Quaest. Ent., 1982, 18 (1-4) Philanthus albopilosus Cresson 121 Figure 10. Distribution patterns of Philanthus albopilosus through time, a) 70,000 B.P. hypothetical range of ancestral stock, b) Ca. 17,000 B.P. geographic isolation of ancestral stock at the zenith of the Wisconsin, c) Ca. 7,000 B.P. expansion of ranges from refugiom, d) recent distribution of (1) Philanthus albopilosus albopilosus (dark race) and P. a. manuelito (pale race). cover and being subject to wind erosion. Maximal rates of range extension may have occurred during the Hypsithermal due to increased dune and blowout development. In the southwestern United States, arid regions may have persisted during Wisconsinan glaciation, permitting dune fields to remain active. Sonoran Desert floras were derived relatively recently (Axelrod, 1979). The great diversity is due to a ‘vacuum’ effect, floristic elements from surrounding zones which adapted to increasing arididty and temperature, provide the bulk of taxa. A rise in rainfall of 250 to 300 mm would eliminate the Sonoran Desert as a regional lowland. This would allow an oak-conifer association to cover much of the lowland. Axelrod’s data suggest that taxa survived on the driest sites in bordering zones. A portion of the american southwest is proposed as a refugium for the southern subspecies, P. albopilosus manuelito . Very few sites suitable for a northern refugium are available east of the Mississippi River (Thorp et al., 1952). Most of eastern North America was cloaked in a continuous forest (Ross 1970, Martin 1958, Freitag 1973, and others). On the Great Plains, several sand dune areas were probably active during portions of the Wisconsinan glaciation. Smith (1965) demonstrated that there were several periods of dune building on the Nebraska Sand Hills, based on past movement of stabilized dunes. This large dune area may have served 122 Hilchie as a refugium for sand dwelling organisms which were able to adapt to a cooler climate. Willis (1967) proposed survival of at least five species of tiger beetles ( Cicindela : Cicindelidae) in the vicinity of the Nebraska Sand Hills. Following Wisconsin glaciation, many sand dunes developed and older ones became reactivated. Figure 10c illustrates proposed dispersal patterns for P. albopilosus Cresson from refugia. Expansion of Philanthus albopilosus manuelito onto the Great Plains should have allowed rapid colonization of most prairie sand dunes. Northward expansion may have been limited by presence of Philanthus albopilosus albopilosus acting as a cryptic species, genetically distinct, yet morphologically similar. Rare occurrence of Philanthus albopilosus manuelito in the Great basin may be the result of relatively recent post-Pleistocene dispersal. EVOLUTION OF COLOUR PATTERN AND BEHAVIOURAL FEATURES OF P. ALBOPILOSUS Colour pattern features of P. albopilosus Ancestral P. albopilosus were probably similar to P. a. manuelito . Extensive yellow contrasted with black, in typical warning maculation pattern, is similar to that seen on other sand dune sphecids (e.g. Microbembex, P. psyche). Adaptation to a cooler climate during Wisconsinan glaciation may have been the major evolutionary reason for differentiation by populations in the northern refugium. Females of northern, P. a. albopilosus are dark: this may be a functional adaptation for absorbing radiant heat, which may prolong the length of time per day in which they could hunt. Males may not require much darkening, as they could restrict their activities to the hot microenvironment of the sand dune surface. The very pale form of P. a. manuelito of the great plains is somewhat of a puzzle: Character displacement was suggested, but does not adequately explain populations of area 4, unless, there is considerable gene flow between populations on the plains. Mountain barriers would buffer populations in Areas 2 and 3 from influx of extreme pale forms. Present distribution of P. a. albopilosus is the result of post-Pleistocene dispersal into new dune habitats. Eastern and northwestern expansion of range has occurred. Southward expansion may have been blocked by P. a. manuelito . There may be enough genetic and behavioural differences reducing the tendency for the two groups to hybridize (cryptic species). A few northern populations contain well marked individuals. This may be the result of reversed selection toward a form with typical warning colouration pattern or they may represent remnants of the ancestral gene pool or limited introgression. The above discussion is primarily based on variation in the females; trends are present in males but are less marked. Behavioural features of P. albopilosus Evolutionary adaptations exhibited by P. albopilosus are similar to other digger wasps living in sand dune environments. Parallel evolution in development of false burrows originated from nest parasite pressures, which are intense in open environments. Orientation to the nest is difficult for females, as landmarks near the nest may change in an unstable dune environment. Use of distant landmarks is important, but fixation to these requires seeing the nest site while orienting and keeping nest parasites out. False burrows may serve both of these functions. Quaest. Ent., 1982, 18 (1-4) Philanthus albopilosus Cresson 123 Short occupancy time of the nest as compared to other species of Philanthus may be due to friable soil. Little time is spent digging in comparison to species which dig in hard soils. Few cells per nest would be a function of the ephemeral occupancy coupled with slow provisioning. A relatively long adult life span, nest location in areas devoid of vegetation and anti-parasite nest building behaviours may reduce selection pressures for the production of large numbers of offspring (“k” selection). Reduction in number of generations per year in northern populations is probably a result of shorter growing seasons either presently or during the species evolutionary history. Change in prey specificity from the ancestral limits may be a response to reduction in number and species of bees available during Wisconsinan time. Alternate prey species would be incorporated from other groups of Apocrita. These facets of life history serve to characterize P. albopilosus as a whole. This also serves in part, as a start in characterization of the northern fauna of Philanthus. ACKNOWLEDGEMENTS I wish to express my gratitude to the following people for assistance in this study; G. E. Ball, my thesis supervisor and sponsor who critically and constructively assisted and guided this manuscript to completion. W. G. Evans and A. L. Steiner for advice and assistance through the course of the study. I would like to thank R. M. Bohart for assistance in identification of eumenid and sphecid wasps. Thanks are due to the curators of museums from which material was borrowed. Thanks are extended to J. L. and B. F. Carr for my introduction to the wonders of sand dune communities; F. A. H. Sperling for assistance in drafting illustrations and checking the manuscript, and J. R. Spence for critically reviewing this paper. Financial assistance is acknowledged from The National Research Council of Canada grant A 1399 held by G. E. Ball and the Entomological Society of Canada Postgraduate Award. REFERENCES Alberta Environment, 1975. Climate of Alberta, tables of temperature, precipitation and sunshine. 90 pp. Alcock, J. 1974. 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Classification, phylogeny and zoogeography of Schizogenius putzeys (Coleoptera: Carabidae: Scaritini). Quaestiones Entomologicae 8: 131-348. Willis, H.J. 1967. Bionomics and zoogeography of tigerbeetles of saline habitats in central U.S.A. University of Kansas Science Bulletin 47: 143-313. Wright, H.E. jr. (Ed.). 1969. Quaternary geology and climate. National Academy of Sciences, Publication 1701. Wright, H.E. and D.G. Frey (Ed.). 1965. The Quaternary of the United States. Princeton University Press, Princeton, N.J. 922 pp. 2*References not seen Quaest. Ent., 1982, 18 (1-4) RHYSODINI OF THE WORLD PART III. REVISION OF OMOGLYMMIUS GANGLBAUER (COLEOPTERA: CARABIDAE OR RHYSODIDAE) AND SUBSTITUTIONS FOR PREOCCUPIED GENERIC NAMES Ross T. Bell Department of Zoology University of Vermont Burlington, VERMONT 05405 U.S.A. Quaestiones Entomologicae 18: 127-259 1982 Joyce R. Bell 24 East Terrace South Burlington, VERMONT 05401 U.S.A. ABSTRACT This paper is third of a series which will constitute a revision of the Rhysodini of the world. Two replacements are proposed for genera described in Part 1: Kupeus NEW NAME, for Kupea Bell and Bell 1978, preoccupied by Kupea Philpott 1930, and Tangarona NEW NAME for Tangaroa Bell and Bell 1978, preoccupied by Tangaroa Lehtinnen 1967. The remainder of the paper is a revision of Omoglymmius Ganglbauer. Pyxiglymmius Bell and Bell is reduced to the rank of subgenus. The subgeneric classification is extensively altered: eleven subgenera are recognized, compared with five in Part I. New subgenera are: Boreoglymmius NEW SUBGENUS, type- Omoglymmius americanus ( Castelnau ), three spp; North America, Japan; Laminoglymmius NEW SUBGENUS, type- Omoglymmius insularis (Grouvelle), seven spp. Malay Peninsula, Sumatra, Borneo, Nicobar Is., New Guinea; Caeconavitia NEW SUBGENUS, ^pe-Omoglymmius zimmermani Bell and Bell, one species, Fiji; Indoglymmius NEW SUBGENUS, type- Omoglymmius lineatus ( Grouvelle ), one species south India; Carinoglymmius NEW SUBGENUS, type- Omoglymmius carinatus ( Grouvelle ), three spp., Sumatra, Borneo, Andaman Is. The following new species are described ( type localities indicated): Omoglymmius (Hemiglymmius) hemipunctatus (SUMATRA (?), Serdang, Lau Raku); O. (H.) occultus (SUMATRA, only); O. (H.) rimatus (JAVA, Mt. Kawi); O. (H.) inermis (SUMATRA, only); O. (Pyxiglymmius) cristatus (PHILLIPPINE IS., Luzon, Mt. Makilung); O. (P.) hesperus (M ENT AWE I Is. Sipora, Sereina); O. (P.) krikkeni (SUMATRA: Alas Valley, Gumpang); O. (Laminoglymmius) inaequalis (NICOBAR IS.); O. (L.) actae (NEW GUINEA, Maffin Bay); O. (L.) trisinuatus (SUMATRA (?), Bangung); O. (L.) gorgo (SUMATRA, Siantar); O. (Navitia) stylatus (NEW HEBRIDES: Malekula); O. (Nitiglymmius) semioculatus (PHILIPPINE IS., Siargao, Dapa); O. (Orthoglymmius) microtis (RYUKYU IS.: Ishigaki, Takeda); O. (sensu stricto) imugani (PHILIPPINE IS., Luzon, Imugan; O. (s. str.) politus (PHILIPPINE IS., Luzon, Mt. Polis; O. (s. str.) crassicornis (PHILIPPINE IS., Negros, Horns of Negros); O. (s. str.) amplus (SUMATRA, Palembang); O. (s. str.) modiglianii 128 Bell and Bell ( MENTAWEI IS., Si Oban); O. (s. str.) evasus ( PHILIPPINE IS., Mindanao, E. Slope Mt. McKinley ); O. (s. str.) nemoralis (SARAWAK, Mt. Matang); O. (s. str.) fraudulentus ( SUMATRA , Palembang); O. (s. str.) coelebs ( PHILIPPINE IS., Palawan, Binaluan); O (s. str.) thoracicus (JAVA, Tangk. Prahoe); O. (s. str.) summissus (SUMATRA, Palembang); O. (s. str.) semperi (PHILIPPINE IS., only); O. (s. str.) data (PHILIPPINE IS., Luzon, Mt. Data); O. (s. str.) hiekei (PHILIPPINE IS., Luzon, Dalbalan); O. (s. str.) pectoralis (JAVA: Mt. Smetou); O. (s. str.) quadruplex (PHILIPPINE IS., Mindanao, Zamboanga, Kabasalan); O. (s. str.) duplex (PHILIPPINE IS., Luzon, Mt. Makiling); O. (s. str.) bouchardi (SUMATRA, Palembang); O. (s. str.) consors (SUMATRA, Palembang); O. (s. str.) repetitus (CELEBES, Tolitoli); O. (s. str.) opticus (LESSER SUNDA IS., Dammar Is.); O. (s. str.) viduus (KEI IS.); O. (s. str.) continuus (MOLUCCA IS., Sula Is., Mangole); O. (s. str.) wittmeri (MOLUCCA IS., Sula Is., Mangole); O. (s. str.) vadosus (MOLUCCA IS., Amboina); O. (s. str.) morditus (MOLUCCA IS., Morotai ); O. (s. str.) nasalis (MOLUCCA IS., Boeroe, Kajeli);0. (s. str.) bicarinatus (SCHOUTEN IS., Jobi); O. (s. str.) lindrothi (SOLOMON IS., Guadalcanal, Kukum); O. (s. str.) modicus (SOLOMON IS., Savo); O. (s. str.) rusticus (SOLOMON IS., Russell Is., Loani); O. (s. str.) manni (SOLOMON IS., Malaita, Auki); O. (s. str.) regius (SOLOMON IS., Isabel, Regi); O. (s. str.) gurneyi (SOLOMON IS., Bougainville); O. (s. str.) princeps (SOLOMON IS., Bougainville, Kokure, Crown Prince Ra.); O. (s. str.) renutus (SOLOMON IS., New Georgia, Mavovo); O. (s. str.) scopulinus (SANTA CRUZ IS., Reef I); O. (s. str.) mycteroides (SOLOMON IS., Kolombangara, Hunda); O. (s. str.) tabulatus (SOLOMON IS., Boku); O. (s. str.) classicus (ADMIRALTY IS., Los Negros); O. (s. str.) oroensis (NEW GUINEA, Oro Bay); O. (s. str.) fringillus (NEW GUINEA, Huon Peninsula, Finschhafen ); O. (s. str.) puncticornis (NEW GUINEA, Fly R., Kiunga); O. (s. str.) trepidus (NEW GUINEA, Wau, Morobe Dist .); O. (s. str.) patens (NEW GUINEA, Maffin Bay); O. (s. str.) cavea (NEW GUINEA, Ramoi Is.); O. (s. str.) sectatus (NEW GUINEA, Mt. Missim); O. (s. str.) ephemeris (NEW GUINEA, Nabire, s. Geelvink Bay); O. (s. str.) follis (NEW GUINEA, Wau, Morobe Dist.); O. (s. str.) iridescens (NEW GUINEA, Araboebivak); O. (s. str.) massa (NEW GUINEA, Eliptamin Val); O. (s. str.) biroi (NEW GUINEA, Sattelberg); O. (s. str.) denticulatus (NEW GUINEA, Wissel Lakes, Enarotadi); O. (s. str.) auratus (NEW GUINEA, Swart Val.); O. (s. str.) sus (NEW GUINEA, Katau); O. (s. str.) planiceps (NEW GUINEA, Wau, Morobe Dist.); O. (s. str.) lentus (NEW GUINEA, Sattelberg); O. (s. str.) asetatus (NEW GUINEA, Madang); O. (s. str.) sedlaceki (NEW GUINEA, Wau, Morobe Dist.). O. (Pyxiglymmius) aterrimus (Chevrolat) is synonymous with O. (P.) strabus (Newman). RESUME Cet article est la troisieme d'une serie qui constiteront une revue taxonomique des Rhysodini du monde. Deux nommes remplaces des autres car ils sont preoccupees: Kupeus NOUVEAU NOM pour Kupea Bell et Bell 1978, preoccupee de Kupea Philpott, 1930, et Tangarona NOUVEAU NOM pour Tangaroa Bell et Bell 1978, preoccupee de Tangaroa Lehtinnen 1967. Le reste de Particle est une revision taxonomique du genre Omoglymmius Ganglbauer. Pyxiglymmius Bell et Bell est redui au rang de sous-genre. Nous changeon beaucoup la classification des sous-genres: nous reconnaissons onze sous-genres en comparaison de cinque sous-genres en Partie I. Les sous-genres nouveaux sont: Boreoglymmius NOUVEAU SOUS-GENRE, type- Omoglymmius americanus (Castelnau), trois spp. Amerique du Nord, Japon; Laminoglymmius NOUVEAU SOUS-GENRE, type- Omoglymmius insularis (Grouvelle), sept, spp., Peninsule de Malaisie, Sumatra, Borneo, lies Nicobars, Nouvelle Guinee; Caeconavitia NOUVEAU SOUS-GENRE, type- Omoglymmius zimmermani Bell et Bell, une sp., Fiji: Indoglymmius NOUVEAU SOUS-GENRE, type- Omoglymmius lineatus ( Grouvelle ), une sp.. Inde du sud; Carinoglymmius NOUVEAU SOUS-GENRE, type- Omoglymmius carinatus ( Grouvelle ), trois spp., Sumatra, Borneo, lies Andamans. On decrit les especes nouvelles qui voice (en indiquant pour chacune la localite du specimen type): Omoglymmius (Hemiglymmius) hemipunctatus (SUMATRA (?), Serdang, Lau, Raku); O. (H.) occultus (SUMATRA sans localite Revision of Omoglymmius Ganglbauer 129 specifie); O. (H.) rimatus (JAVA, Mt. Kawi); O. (H.) inermis ( SUMATRA , sans localite specific); O. (Pyxiglymmius) cristatus (ILES PHILIPPINES, Luzon, Mt. Makiling); O. (P.) hesperus (ILES DE MENTAWEI, Sipora, Sereina ); O. (P.) krikkeni (SUMATRA, vallee d'Alas, Gumpang ); O. (Laminoglymmius) inaequalis (lies Nicobars ); O. (L.) actae (NOUVELLE GUINEE, Baie de Maffin); O. (L.) trisinuatus (SUMATRA (?), Bangung); O. (L.) gorgo (SUMATRA, Siantar ); O. (Navitia) stylatus (NOUVELLE HEBRIDES, Malekula); O. (Nitiglymmius) semioculatus (ILES PHILIPPINES, Siargao, Dapa); O. (Orthoglymmius) microtis (ILES DE RYUKYU: Ishigaki, Takeda); O. (sensu stricto) imugani (ILES PHILIPPINES, Luzon, Imugan); O. (s. str.) politus (ILES PHILIPPINES, Luzon, Mt. Polis); O. (s. str.) crassicornis (ILES PHILIPPINES, Negros, Horns of Negros); O. (s. str.) amplus (SUMATRA, Palembang ); O. (s. str.) modiglianii flLES DE MENTAWEI, Si Oban); O. (s. str.) evasus (ILES PHILIPPINES, Mindanao, versant est du Mont McKinley); O. (s. str.) nemoralis (SARAWAK, Mt. Matang); O. (s. str.) fraudulentus (SUMATRA, Palembang); O. (s. str.) coelebs (ILES PHILIPPINES, Palawan, Binaluan); O. (s. str.) thoracicus (JAVA, Tangk. Prahoe); O. (s. str.) summissus (SUMATRA, Palambang); O. (s. str.) semperi (ILES PHILIPPINES, san localite specifie); O. (s. str.) data (ILES PHILIPPINES, Luzon, Mt. Data); O. (s. str.) hiekei (ILES PHILIPPINES, Luzon, Dalbalan); O. (s. str.) pectoralis (JAVA, Mt. Smetou); O. (s. str.) quadruplex (ILES PHILIPPINES, Mindanao, Zamboanga, Kabasalan); O. (s. str.) duplex (ILES PHILIPPINES, Luzon, Mt. Makiling); O. (s. str.) bouchardi (SUMATRA, Palembang); O. (s. str.) consors (SUMATRA, Palembang); O. (s. str.) repetitus (CELEBES, Tolitoli); O. (s. str.) opticus (PETITES ILES DE SONDE, lies de Dammar); O. (s. str.) viduus (ILES DE KEI); O. (s. str.) continuus (ILES MOLUQUES, lies de Sula, Mangole); O. (s. str.) wittmeri (ILES MOLUQUES, lies de Sula, Mangole); O. (s. str.) vadosus (ILES MOLUQUES, Amboina); O. (s. str.) morditus (ILES MOLUQUES, Moratai); O. (s. str.) nasalis flLES MOLUQUES, Boeroe, Kajeli), O. (s. str.) bicarinatus (ILES DE SCHOUTEN, Jobi); O. (s. str.) lindrothi (ILES DE SOLOMON, Guadalcanal, Kukum); O. (s. str.) modicus (ILES DE SOLOMON, Savo); O. (s. str.) rusticus (ILES DE SOLOMON, lies de Russell, Loani); O. (s. str.) manni (ILES DE SOLOMON, Malaita, Auki); O. (s. str.) regius (ILES DE SOLOMON, Isabel, Regi); O. (s. str.) gurneyi (ILES DE SOLOMON; Bougainville); O. (s. str.) princeps (ILES DE SOLOMON , Bouganville, Kokure, Crown Prince Ra.); O. (s. str.) renutus (ILES DE SOLOMON , Nouveau Georgia, Mavovo); O. (s. str.) scopulinus (ILES SANTA CRUZ, lie de Reef); O. (s. str.) mycteroides (ILES DE SOLOMON, Kolombangara, Hunda); O. (s. str.) tabulatus (ILES SOLOMON, Boku); O. (S. str.) classicus flLES D’ ADMIRALTY, Los Negros); O. (s. str.) oroensis (NOUVELLE GUINEE, Baie d'Oro ); O. (s. str.) fringillus (NOUVELLE GUINEE, Huon Peninsula, Finschhafen); O. (s. str.) puncticornis (NOUVELLE GUINEE, Fly R., Kiunga); O. (s. str.) trepidus (NOUVELLE GUINEE, Wau, Morobe Dist .); O. (s. str.) patens (NOUVELLE GUINEE, Baie de Maffin ); O. (s. str.) cavea (NOUVELLE GUINEE, ‘lie Ramoi); O. (s. str.) sectatus (NOUVELLE GUINEE, Mt. Missim); O. (s. str.) ephemeris (NOUVELLE GUINEE, Nabire, sud de la Baie de Geelvink); O. (s. str.) follis (NOUVELLE GUINEE, Wau, Morobe Dist.); O. (s. str.) iridescens (NOUVELLE GUINEE, Araboebivak); O. (s. str.) massa (NOUVELLE GUINEE, Eliptamin Val.); O. (s. str.) biroi (NOUVELLE GUINEE, Sattelberg); O. (s. str.) denticulatus (NOUVELLE GUINEE, Lacs de Wissel, Enarotadi); O. (s. str.) auratus (NOUVELLE GUINEE, Swart Val.); O. (s. str.) sus (NOUVELLE GUINEE, Katau); O. (s. str.) planiceps (NOUVELLE GUINEE, Wau, Morobe Dist.); O. (s. str.) lentus (NOUVELLE GUINEE, Sattelberg); O. (s. str.) asetatus (NOUVELLE GUINEE, Madang); O. (s. str.) sedlaceki (NOUVELLE GUINEE, Wau, Morobe Dist.). Nous decouvrames que O. (Pyxiglymmius) aterrimus (Chevrolat) est un synonyme de O. (P.) strabus NEWMAN TABLE OF CONTENTS Introduction 1 30 Sources of Material 1 30 Genus Omoglymmius 131 Subgenus Hemiglymmius (Figs. 1-16) 132 New Subgenus Boreoglymmius (Figs. 17-26) 140 Subgenus Pyxiglymmius (Figs. 27-56) 146 New Subgenus Laminoglymmi us (Figs. 57-74) 1 56 Subgenus Navitia (Figs. 75,76,79,80) 164 New Subgenus Caeconavitia 166 New Subgenus Indoglymmius (Figs. 77,83) 1 67 Subgenus Nitiglymmius (Figs. 78,8 1 ,82) 168 Subgenus Orthoglymmius (Figs. 84-103) 169 New Subgenus Carinoglymmius (Figs. 104-1 12) 176 Subgenus Omoglymmius sensu stricto (Figs. 1 1 3-237) 180 Substitutions for Preoccupied Generic Names 253 Quaest. Ent., 1982, 18 (1-4) 130 Bell and Bell Aeknowledgements 254 Supplementary References 254 INTRODUCTION This paper, third of a series of five, comprises a revision of the large and difficult genus Omoglymmius Ganglbauer. Pyxiglymmius Bell and Bell, formerly regarded by us as a separate genus, is included as a subgenus. The discovery of many additional species has resulted in several changes in limits and definitions of subgenera. Two generic names proposed in Part I of this series are preoccupied, and substitutes are proposed. SOURCES OF MATERIAL The following abbreviations designate collections cited in this paper. Names of curators of respective institutions are in parentheses. AMS Instituut voor Taxonomische Zoologie, Amsterdam, Netherlands (J. Duffels) AP U.S. Dept, of Agriculture, Harrisburg, PA (K. Valley) ARK University of Arkansas, Fayetteville (E. P. Rouse) AU S. F. Austin State University, Nacogdoches, Texas (W. W. Gibson) BMNH British Museum, Natural History, London (R. Pope) BPBM Bernice P. Bishop Museum, Honolulu (G. Samuelson) BPM Barry P. Moore, Canberra City, Australia BSL Naturhistorisches Museum, Basel, Switzerland (W. Wittmer) CAG U.S. Dept, of Agriculture, Sacramento, CA (F. G. Andrews) CAS California Academy of Sciences, San Francisco (D. H. Kavanaugh) CMP Carnegie Museum of Natural History, Pittsburgh, PA (G. Wallace) CNHM Field Museum of Natural History, Chicago, IL (H. Dybas) CU Cornell University, Ithaca, NY (L. L. Pechuman) DM Dayton Museum, Ohio (A. J. Koestner) DY Daniel K. Young, E. Lansing, MI FLA U.S. Dept, of Agriculture, Gainesville, FL (R. Woodruff) GEN Museo Civico di Storia Naturale G. Doria , Genoa (R. Poggi) GLP Gary L. Peters, Corvallis, OR HL Harry J. Lee, Fairview Park, OH IO Iowa State University, Ames (R. Miller) ISNHS Illinois State Natural History Survey, Urbana (M. Sanderson) IU Indiana University, Bloomington KS Karl Stephan, Tucson, AZ KU Kagoshima University, Japan LA Los Angeles County Natural History Museum, CA (C. L. Hogue) LEI Rijksmuseum von Natuurlijke Historie, Leiden, Netherlands (J. Krikken) LEN Academy of Sciences, Leningrad (O. Kryzhanovskij) LS Louisiana State University, Baton Rouge (J. B. Chapin) LUN Zoological Institute, Lund, Sweden (R. Danielsson) MCZ Museum of Comparative Zoology, Harvard University, Cambridge, MA Revision of Omoglymmius Ganglbauer 131 (J. H. Lawrence) MNHB Museum fur Naturkunde der Humboldt-Universitat, Berlin, DDR (F. Hieke) MNHN Museum National d’Histoire Naturelle, Paris, France (A. Descarpentries) MN University of Minnesota, St. Paul (P. J. Clausen) MO University of Missouri, Columbia (W. R. Enns) MRAC Musee Royal de l’Afrique Centrale, Tervuren, Belgium (P. Basilewsky) MSU Michigan State University, E. Lansing NC North Carolina State University, Raleigh (D. A. Young) NMNH United States National Museum of Natural History, Washington, D. C. (T. L. Erwin) NMNZ National Museum of New Zealand, Wellington (R. G. Ordish) NMW Naturhistorisches Museum Wien, Austria (F. Janczyk) OK Oklahoma State University, Stillwater (W. A. Drew) OS Oregon State University, Corvallis (G. L. Peters) OSFS Oregon State Forest Sciences Collection, Corvallis OSU Ohio State University, Columbus (C. A. Triplehorn) PA Academy of Sciences, Philadelphia (D. C. Rentz) PU Purdue University, Lafayette, IN RCG R. C. Graves, Bowling Green, OH SATO Masataka Sato, Nagoya, Japan SDA U.S. Dept, of Agriculture, Brookings, SD (V. M. Kirk) SI Southern Illinois University, Carbondale (J. E. McPherson) TB Thomas Barr, University of Kentucky, Lexington UCD University of California, Davis UK University of Kansas, Lawrence (G. W. Byers) UL University of Louisville, KY (C. V. Covell) UM University of Michigan, Ann Arbor (I. J. Cantrall) UN University of Nebraska, Lincoln (B. C. Ratcliffe) UT Utah State University, Logan (W. J. Hanson) UW University of Wisconsin, Madison (J. R. Baker) VP Virginia Polytechnic Institute, Blacksburg (M. Kosztarab) WR William Rosenberg WRS Walter R. Suter, Carthage College, Kenosha, WI WS Washington State University, Pullman (W. J. Turner) Genus Omoglymmius Ganglbauer 1892 Type species. - Rhysodes germari Ganglbauer 1892 Description. — Paramedian grooves of pronotum deep, distinct; middle and hind tibiae each with a single terminal spur; antennal stylet, basal setae, midial angles of temporal lobes variously developed. The description has been altered from that in Part I, p. 72, to reflect incorporation of Pyxiglymmius as a subgenus. In the key to genera of Omoglymmiina, Part I, pp. 66-67, adults of Omoglymmius as presently defined, trace to 5'. The key to subgenera presented below supersedes that in Part I, p. 72. Quaest. Ent., 1982, 18 (1-4) 132 Bell and Bell KEY TO SUBGENERA 1 v 2 (1) 2' 3 (20 3' 4 (3) 4/ 5 (4) 5' 6 (50 6' 7 (40 7 8 (70 8' 9 (30 9' 10(90 10' Clypeal setae present Orthoglymmius Bell and Bell, p. 169 Clypeal setae absent Spur of middle tibia straight; female with deepest lateral pits on Sternum V; outer carina very narrow, 0.33 or less as wide as paramedian groove Carinoglymmius new subgenus, p. 176 Spur of middle tibia curved anteriorly; female with deepest pits on Sternum IV; outer carina in most species subequal in width to paramedian groove, or only slightly narrower (in O. ( s . str.) solitarius and O. ( Laminoglymmius ) inaequalis 0.33 as wide as paramedian groove) Apical stylet of antenna absent Apical stylet present Basal setae present on outer antennal segments Basal setae absent Medial margin of temporal lobe with broad emargination between two widely separated medial angles Pyxiglymmius Bell and Bell, p. 146 Medial margin of temporal lobe without emargination or with one or two shallow ones; each temporal lobe with one medial angle or with two or three close together Medial margin with translucent area (ill-defined in O. actae ); medial margin with distinct emarginations (except in O. inaequalis ) Laminoglymmius new subgenus, p. 1 56 Medial margin without translucent area and without emargination, medial angle single Boreoglymmius new subgenus, p. 140 Medial angles with translucent area Indoglymmius new subgenus, p. 167 Medial angles without translucent area Eye large, over 0.5 as deep as head; cornea distinctly facetted Omoglymmius sensu stricto Ganglbauer, p. 180 Eye reduced; less than 0.25 as deep as head; cornea not facetted Nitiglymmius Bell and Bell, p. 168 Outer antennal segments with basal setae. Hemiglymmius Bell and Bell, p. 132 Basal setae absent Eyes large, cornea facetted; frontal grooves obsolete Navitia Bell and Bell, p. 164 Eye reduced, less than 0.25 as deep as head; cornea not facetted; frontal grooves deep Caeconavitia new subgenus, p. 166 2 3 4 9 5 7 6 8 Subgenus Hemiglymmius Bell and Bell 1978 Type species. - Rhysodes africanus Grouvelle 1892. Description. — Basal setae on at least antennal Segments IX, X (in most species on more proximal segments as well); antennal stylet present; clypeal setae absent; eye large, normal, cornea facetted; frontal grooves deep, complete, dilated in many species; medial angle simple, no translucent area; postorbital tubercle present; marginal groove of pronotum deep; spur of middle tibia curved anteriorly; punctures of abdominal Sterna lll-V scattered; lateral pits of abdomen in female on Sternum IV, poorly developed. Revision of Omoglymmius Ganglbauer 133 This subgenus differs from others that have an antennal stylet and basal setae in having simple, unmodified medial angles on the temporal lobes. Limits of this subgenus have been altered by removal of those species which lack the antennal stylet ( O . americanus, O. hamatus, and O. lewisi ) to Boreoglymmius new subgenus. Restudy of types has shown that O. oberthueri belongs in Laminoglymmius new subgenus, while O. nicobarensis belongs in Orthoglymmius Bell and Bell. We also originally listed O. borneensis among the species of Hemiglymmius. We have not been able to locate the type for this species, but a detailed study of the original description suggests that the name is based on a species of Omoglymmius s. str. from Borneo, and not on the species formerly interpreted by us as being O. borneensis. Range of the subgenus Hemiglymmius as limited above includes Java, Borneo, and Sumatra, and a large part of Africa, from Liberia to eastern Zaire. O. germaini , described from Bolivia, is certainly a member of this subgenus. However, it is very similar to one of the two species forming Grouvelle’s concept of O. javanicus, so it seems certain that locality data associated with the types is erroneous. Phylogeny. - O. africanus, with its narrow, abbreviated median lobe, appears to stand apart from the Indonesian species. The latter constitute a closely related and poorly known complex whose interrelationships will not be easy to decipher until both sexes of all species have been collected. KEY TO SPECIES 1 Median lobe of head narrow, linear, short, ended opposite anterior margin of eye; gular tubercle absent; male without ventral tooth on anterior femur O. africanus (Grouvelle), p. 135 1 ' Median lobe narrow to broad, long, extended at least to level of middle of eye; gular tubercle present; male with ventral tooth on anterior femur 2 2 (L) Temporal lobe not extensively pollinose, its glabrous portion separated from antennal lobe only by pollinosity of antennal groove 3 2' Temporal lobe extensively pollinose anteriorly, its glabrous portion broadly separated from antennal lobe 6 3 (2) Inner carina of pronotum punctate 4 3' Inner carina impunctate O. hemipunctatus new species, p. 137 4 (3) Temporal lobe with 18-20 coarse punctures; Stria IV with five or six setae along its length O. occultus new species, p. 138 4' Temporal lobe with one to five punctures, or none; Stria IV with one or two setae near apex 5 5 (4') Pronotum widest anterior to middle; outer carina very broad anteriorly O. germaini (Grouvelle), p. 137 5' Pronotum broadest at middle; outer carina less broad anteriorly O. javanicus (Grouvelle), p. 136 6 (2') Median lobe glabrous; antennal lobe bounded posteriorly by prominent scarp O. rimatus new species, p. 139 6' Median lobe pollinose; antennal lobe not bounded posteriorly by scarp or with only a vestige of one 7 7 (6') Male with proximal tooth on anterior tibia; pronotum shorter, length/greatest Quaest. Ent., 1982, 18 (1-4) 134 Bell and Bell Plate 1. Figures 1-16, Subgenus Hemiglymmius. Figs. 1-8, Head and pronotum, dorsal aspect; Fig. 1, Omoglymmius (Hemiglymmius) africanus (Grouvelle); Fig. 2, O. (H.) javanicus (Grouvelle); Fig. 3, O. (H.) germaini (Grouvelle); Fig. 4, O. (H.) hemipunctatus new species; Fig. 5, O. (H.) occultus new species; Fig. 6, O. (H.) ineditus (Dajoz); Fig. 7, O. (H.) rimatus new species; Fig. 8, O. (H.)inermis new species; Figs. 9-16, Head, lateral aspect; Fig. 9, O. (H.) africanus (Grouvelle); Fig. 10, O. (H.) javanicus (Grouvelle); Fig. 11,0. (H.) germaini (Grouvelle); Fig. 12, O. (H.) hemipunctatus new species; Fig. 13, O. (H.) occultus new species; Fig. 14 O. (H.) ineditus (Dajoz); Fig. 15 O. (H.) rimatus new species; Fig. 16 O. (H.) inermis new species. Revision of Omoglymmius Ganglbauer 135 width about 1.26; calcars larger O. ineditus (Dajoz), p. 138 T Male without proximal tooth on anterior tibia; pronotum longer, length/greatest width about 1 / 36; calcars smaller O. inermis new species, p. 140 Omoglymmius ( Hemiglymmius ) africanus (Grouvelle 1892) Figs. 1, 9 Rhysodes africanus Grouvelle 1892: 299. Omoglymmius ( Hemiglymmius ) africanus (Grouvelle) Bell and Bell 1978 Type material. - LECTOTYPE sex not recorded, labelled: “Assinie,” no other data (MNHN). According to the original description there were two specimens in the type series, so the second specimen, if still extant, is a paralectotype. Assinie was a former French colony, centering about the town of the same name, now part of the Ivory Coast. A specimen labelled “Lac Albert (Albrecht), Afrique Allemande” (MNHN), is labelled as a paratype, but is probably not one in modern terms, it is not mentioned in the original description. The locality is ambiguous, as Lake Albert was not in German East Africa. Description. — Length 5. 8-6. 7 mm. Antennal stylet short, acute; basal setae well-developed on Segments IX, X, sparse on Segments V I- V III; absent from Segments IV, V; Segments I-XI coarsely punctate; median lobe narrow, with sides parallel, very short, tip even with anterior margin of eye, apex obtuse; median lobe varied from coarsely punctate to impunctate; temporal lobe with medial angles nearly rectangular, rather widely separated in most specimens (but nearly contiguous in some); posteriomedial margin oblique between medial and occipital angles; temporal lobe varied from finely punctate to impunctate; distinct postantennal depression between antennal base and eye, extended medially from lateral margin, but not to frontal groove; temporal setae absent; postorbit entirely finely pollinose; postorbital tubercle well-developed, in most specimens with well-developed pilosity conspicuously coarser and longer than pollinosity of postorbit. Pronotum relatively short, length/greatest width averaging 1.18; widest at middle; sides curved, convergent both basally and apically; margin not sinuate anterior to hind angle; median groove relatively broad, about 0.33 as broad as inner carina at middle; marginal groove dilated, as wide as outer Carina; both carinae narrow, inner slightly narrower than outer one; inner carina impunctate (in some specimens one or two); outer carina finely punctate. Elytra relatively long and narrow (though distinctly wider than pronotum); elytral striae broad, rather shallow; strial punctures large, deep, pilose; intervals convex, very narrow; each interval distinctly narrower than a strial puncture; pattern of setae varied, when fully developed more extensive than in any other member of subgenus; Stria I with one to three in apex; Stria II with three in apex; Stria IV with continuous series of six to seven setae; medial side of apical tubercle with single seta; apical stride with three or four setae; apex of Stria VII with seven to ten setae; some specimens without certain setae (most frequently, setae absent from middle portion of Stria IV); hind coxa without setae; abdominal Sternum VI with pair of setae in both sexes; female with shallow, indistinct lateral pits on Sterna IV and V; male without ventral tooth on anterior femur and without proximal tooth on anterior tibia; middle tibia without trace of calcar; calcar of hind tribia very small, triangular, acute. The short, narrow median lobe gives adults of this species a distinctive appearance, in most specimens enhanced by well-separated medial angles, which make the head appear almost “split”. Variation is extensive. A few specimens, especially those from the most eastern part of the range, have the medial angles almost contiguous. Amount of punctation of medial and temporal lobes is markedly variable, and there is also considerable variation in development of elytral setae. Neither of these two characters seem to vary geographically. Perhaps more than one species is represented among African Hemiglymmius , although we can see no clear-cut pattern in the material available to us. Range. - This is extensive for a rhysodine, from Liberia on the west to the eastern borders of Zaire in the east. In addition to the type series, we have seen specimens from the following localities;CAMEROONS: one male, two females, Lolodorf, coll. G. Schwab (MCZ); one male, “Neu-Kamerun” No. 3209-48, coll. Tessmann, S.G. (MNHB); EQUATORIAL GUINEA: one female, Nkolentangan, XI-07.-V-08, coll. G. Tessman, S.G. (MNHB); one male. Is. Fernando Poo, Musola, 500-800 m.s.m., I- 1 1 1- 1 902, coll. L. Fea (GEN); LIBERIA: four males, three females, Mt. Coffee, Feb-March 1895, O.F. Cook Colin. (NMNH); ZAIRE: Quaest. Ent., 1982, 18 (1-4) 136 Bell and Bell (all specimens in MRAC) one male, Haute-Uele: Moto, 1923, coll. L. Burgeon; one female, Kasai: Ngombe,-1921, coll. Dr. H. Schouteden; one female, Ituri: Yindi, 18-VI-1948, coll. A.E. Bertrand; one female, Lulua: Kapanga, X-1933, coll. G.F. Overlaet; one female, Mayidi, 1942, coll. Rev. P. Van Eyen; two males, Territoire de Sandoa, 18-XI-1948, coll. N. Leleup; one male, two females, Tschuapa, Lac Tumba, Mabali, 350 m., 29- 1 X- 1955, coll. N. Leleup, under bark of dead tree. Omoglymmius { Hemiglymmius ) javanicus (Grouvelle) Figs. 2, 10 Rhysodes javanicus Grouvelle 1903: 1 10. Omoglymmius (Hemiglymmius) javanicus (Grouvelle) Bell and Bell 1978. Type material. - LECTOTYPE (here designated) male labelled: “JAVA, Gounod Gedeh, Ledru 1898, TYPE” (MNHN); PARALECTOTYPES one male, two females, same labels as lectotype (MNHN); one female with same label as lectotype and additional label “Rhysodes javanicus ty. Grouv.” (MNHN); one female, four males with same label as lectotype and additional labels “R. javanicus ty. Grouv., 1952-Oberthiir Colin.”, “Syn. Rhysod. A Grouvelle 1903” (MNHN); one male, with same label as lectotype and additional labels “ex. coll. Oberthiir 1904-175, syn, Rhysod. A. Grouvelle, 1903, co-type, javanicus Grouv. ex typis” (BMNH). A female specimen, labelled: “Tougou, Java Occid., J.D. Pasteur 3-96, Type, Rhysodes javanicus Grouv.” (MNHN), is not conspecific with the series from Gounod Gedeh, but is probably O. ineditus (Dajoz) or O. rimatus n. sp. Description. — Length 6. 7-8.0 mm. Basal setae well developed on antennal Segments V-X; Segments I- 1 V each with pollinose band on dorsal surface; Segments V-X coarsely punctate; Segment XI distinctly longer than wide; corpus cylindrical, narrower than Segment X; stylet short, acute. Head in most specimens slightly longer than wide (average of 11 specimens width/length 0.963; range 0.909-1.02); median lobe impunctate, glabrous, elongate oval, its tip bluntly rounded, just anterior to level of posterior margin of eye; frontal space 1.5 wider than long; medial angles slightly obtuse, contiguous; temporal lobe convex, not at all pollinose; antennal groove pollinose, continous with pollinose orbital groove; temporal lobe with one to six rather fine punctures, or none; temporal setae none to four; temporal setae more frequent than in related species; antennal lobe glabrous, distinctly raised above level of temporal lobe; lateral margin of head shallowly sinuate between antennal lobe and eye; postorbit pollinose; postorbital tubercle small about 0.3 of length of eye. Pronotum relatively broad, length/greatest width 1/19; widest near middle; base, apex distinctly, nearly equally narrowed; lateral margins slightly curved except oblique near base and apex; margin shallowly sinuate just anterior to hind angle; paramedian grooves nearly parallel; outer carina widest just anterior to middle; both inner, outer carinae coarsely punctate. Elytral striae broad, rather shallow; strial punctures coarse throughout; intervals convex, narrow, each narrower than one strial puncture; Stria II with one or two setae near apex; Stria IV with one or two near apex; subapical striole with seta; Stria VI 1 with several setae near apex; abdominal sterna moderately densely punctate; 25-29 punctures in each half of Sternum V; male with ventral tooth on anterior femur, proximal tooth on anterior tibia; middle calcar small but distinct; hind calcar slightly larger than middle one. This species differs sharply from the sympatric O. ineditus in absence of pollinosity from the temporal lobe; longer, narrower head; and much shallower emargination of head margin anterior to the eye. It is more similar to O. hemipunctatus, O. occultus , and O. germaini. The first species lacks punctures on the inner carina of the pronotum, the second has many coarse punctures on the temporal lobe and a complete series of setae in Stria IV, and the third has the pronotum inflated anteriorly and the postorbital tubercle much larger. Range. - O. javanicus occupies the western 2/3 of Java, where it seems to be the commonest member of the subgenus. The easternmost locality is Kopeng, near Jogjakarta. In addition to type material we have seen the following specimens; one male, five females, Gunong Tengkoeban Prahoe, elev. 1000-1400 m., June 20, 1933, coll, van Doesburg (LEI); one male, two females, Kopeng, elev. 1610 m.. May 14, 1933, coll, van Doesburg (LEI); one female, Preanger, no date, coll. P.F.S. Sijthoff (LEI). Revision of Omoglymmius Ganglbauer 137 Omoglymmius ( Hemiglymmius ) germaini (Grouvelle) Figs. 3, 1 1 Rhysodes germaini Grouvelle 1903: 109. Omoglymmius ( Hemiglymmius ) germaini (Grouvelle) Bell and Bell 1978 Type material. - LECTOTYPE (here designated) female labelled: “Cochabamba, Bolivia” (MNHN) PARALECTOTYPE male, same data (MNHN); the paralectotype has a slightly deformed pronotum, with an irregular interruption of the lateral margin near the apex. This species is very close to O. javanicus , and might prove to be conspecific with it. It is very likely that the collecting data are erroneous, and that the true locality is in one of the Greater Sunda Islands. Vulcano and Pereira (1975b) illustrated a specimen under the name R. germaini Grouvelle labelled as coming from Rio Madeira, Brazil. Their specimen lacks postorbital tubercles and antennal stylets, and is not O. germaini nor a member of Subgenus Hemiglymmius. We have not studied this specimen, and cannot form a definite opinion about its specific identity nor its subgeneric placement. We hope that more specimens will be collected. If it is not a mislabelled specimen from some other part of the world, it represents the first authentic record of Genus Omoglymmius from South America. In the paper of Vulcano and Pereira two figures were transposed, so that illustrations of their Omoglymmius appear under the name Clinidium rojasi Chevrolat. Description. — Length 7 .0-7 .2 mm. Basal setae well-developed on antennal Segments V-X; Segments I- 1 V with pollinose band on dorsal surface. Segments V-X coarsely punctate; Segment XI longer than wide, corpus cylindrical, narrower than Segment X; stylet short, acute. Head slightly longer than wide; median lobe impunctate, glabrous, elongate oval, its tip bluntly rounded near level of posterior margin of eye; frontal space 1.5 wider than long; medial angles slightly obtuse, contiguous; temporal lobe convex, not at all pollinose; antennal groove pollinose, continuous with pollinose orbital groove; temporal lobe with one to six coarse punctures; temporal seta absent; antennal lobe glabrous, distinctly raised above level of temporal lobe; lateral margin of head shallowly sinuate between antennal lobe and eye; postorbit finely pollinose; postorbital tubercle 0.3-0. 5 of length of eye. Pronotum with length/greatest width 1.21-1.24, widest anterior to middle; base, apex equally narrowed; lateral margins more markedly curved than in O. javanicus ; margin shallowly sinuate just anterior to hing angle; paramedian grooves distinctly convergent anteriorly; outer carina broadest at anterior 0.25 of length; both inner, outer carinae coarsely punctate. Elytral striae broad, rather shallow; strial punctures coarse throughout; intervals convex, narrow, each narrower than one strial puncture; Stria II with two setae near apex; Stria IV with two setae near apex; subapical stride with one seta; Stria VII with several setae near apex; abdominal sterna densely punctate with about 33 punctures in each half of Sternum V; male with ventral tooth on anterior femur, proximal tooth on anterior tibia; middle calcar more prominent than in O. javanicus , longer than hind calcar, similar to that of O. javanicus. We regard this species as only doubtfully distinct from O. javanicus , distinguished by the pronotum more inflated anteriorly and perhaps by the larger middle calcar. Omoglymmius (Hemiglymmius) hemipunctatus new species Figs. 4, 12 Type material. - HOLOTYPE female, labelled: “Corporaal, Serdang, Lau Raku, 4-1910, cat. no. 6” (LEI). This locality is ambiguous, but probably refers to the former Province of Serdang in Sumatra, and not the town by the same name in northwestern Java. Description. — Length 7.0 mm approx, (specimen broken). Basal setae well-developed on antennal Segments V-X; Segments I- 1 V each with pollinose band on dorsal surface; Segments II-XI coarsely punctate; Segment XI distinctly longer than wide, the corpus nearly cylindrical, slightly narrower than Segment X; stylet acute, longer than in O. germaini. Quaest. Ent., 1982, 18 (1-4) 138 Bell and Bell Head as long as wide; median lobe glabrous, elongate-oval, its tip bluntly rounded just posterior to middle of eye; median lobe impunctate; frontal space 1.5 wider than long; medial angles slightly obtuse, narrowly separated; temporal lobe convex, not at all pollinose, but with pollinose strip in antennal groove and extended into orbital groove; temporal lobe with 16-18 coarse punctures, but without temporal setae; antennal lobe glabrous, distinctly raised above level of temporal lobe; lateral margin of head shallowly sinuate between antennal lobe and eye; postorbit pollinose; postorbital tubercle small, about 0.4 of width of eye. Pronotum slightly narrower than in O. germaini. length-greatest width 1.23;widest near middle, apex more narrowed than base; lateral margin distinctly sinuate anterior to hind angle; pronotal carinae convex, nearly equal; inner carina impunctate, outer carina coarsely punctate. Elytral striae broad, rather shallow; strial punctures coarse in anterior 80%, becoming distinctly finer near apex; Stria II with seta at apex; Stria IV with three setae near apex; apical stride with one seta; several setae near apex of Stria VII; abdominal sterna densely punctate, with about 33 punctures in left half of Sternum V. Male unknown. This species is close to O. javanicus, but differs in having the temporal lobes strongly punctate and the inner pronotal carina entirely impunctate. Omoglymmius (Hemiglymmius) occultus new species Figs. 5, 13 Type material. - HOLOTYPE male, labelled: “SUMATRA” (MNHN); PARATYPE female, same data (MNHN). Description. — Length 6. 4-7. 7 mm. Basal setae well-developed on antennal Segments V-X; Segments 1-IV each with pollinose band on dorsal surface; Segments IV-XI coarsely punctate; Segment XI distinctly longer than wide, corpus slightly narrowed near base, only slightly narrower than Segment X; stylet short, acute. Head as wide as long; median lobe impunctate, glabrous except for minute pollinosity on clypeus; median lobe elongate oval, its tip obtuse, even with middle of eye; median space slightly wider than long; medial angles obtuse, nearly contiguous; temporal lobes convex, not pollinose, but with pollinose strip in antennal groove and extended along orbital groove; temporal lobe with approximately 15 coarse punctures; temporal setae absent; antennal lobe glabrous, distinctly raised above level of temporal lobe; lateral margin of head shallowly sinuate between antennal lobe and eye; post-orbit pollinose; postorbital tubercle larger than in O. javanicus, its width 0.5 of width of eye. Pronotum relatively narrow, length/greatest width 1.30; widest near middle; apex more markedly narrowed than base; margin distinctly sinuate anterior to hind angle; pronotal carinae convex, nearly equal in width; inner carina with six to 12 scattered coarse punctures, most of them near base; outer carina densely, coarsely punctate. Elytral striae broad, rather shallow; strial punctures coarse throughout, not becoming finer near apex; Stria 11 with one to two setae near apex; Stria IV with five or six setae scattered along its length; apical striole with one seta; several setae near apex of Stria VII; abdominal sterna coarsely, densely punctate; male with ventral tooth present on anterior femur, with proximal tooth present on anterior tibia; middle calcar small but distinct; hind calcar equal to middle one. This species is unique among Indonesian Hemiglymmius in having a complete series of setae in Stria IV. It also differs from O. hemipunctatus in having punctures on the inner pronotal carina, and from O. javanicus and O. germaini in having more punctures on the temporal lobe and in having a larger postorbital tubercle. Omoglymmius ( Hemiglymmius ) ineditus (Dajoz) Figs. 6, 14 Rhysodes ineditus Dajoz 1975: 8-9. Omoglymmius ( Hemiglymmius ) ineditus (Dajoz) Bell and Bell 1978 Type material. - HOLOTYPE sex not listed, labelled: “JAVA:Tji Salimar, Province de Preang dans le Sud-Ouest de File, altitude 3,000 pieds, septembre 1897” (MNHN). We have not been able to study this type, which has apparently been misplaced. We provisionally conclude, on the basis of the description and figure, that it applies to the species described below. Revision of Omoglymmius Ganglbauer 139 Description. — Length 5.8-7. 1 mm. Basal setae well-developed on antennal Segments V-X; Segments 1-1 V each with pollinose dorsal band; Segments V-X coarsely punctate; Segment XI distinctly longer than wide, corpus nearly cylindrical, narrower than Segment X; stylet short, acute. Head slightly wider than long; greatest width/length 1.000-1.075; mean 1.050; median lobe pollinose, impunctate, narrow, margins nearly parallel, tip bluntly rounded at level of middle of eye; frontal space 1.5 wider than long; medial angles obtuse, contiguous; temporal lobe convex, anterior half pollinose, posterior half glabrous, shining, with one to four fine punctures or none; temporal seta absent; antennal lobe pollinose, continuous with anterior part of temporal lobe, without trace of scarp; lateral margin of head deeply sinuate between antennal lobe and eye, postantennal groove oblique; postorbit pollinose; postorbital tubercle very small, less distinct than in O. rimatus. Pronotum rather narrow; length/greatest width 1.26, widest near middle, apex strongly narrowed, base moderately narrowed; margin scarcely sinuate anterior to hind angle; pronotal carinae convex, outer and inner ones nearly equal in width at middle; outer carina with 14-18 punctures; inner carina with one to eight coarse punctures or none. Elytral striae broad, rather shallow, intervals convex, narrow; each interval narrower than one strial puncture; latter coarse, closely spaced; Stria 11 with one or two setae near apex; Stria IV with single seta at apex; subapical striole with one seta; Stria VI 1 with several setae near apex; abdominal sterna densely punctate; each half of Sternum V with 38-48 punctures; male with ventral tooth on anterior femur and proximal tooth on anterior tibia; middle calcar triangular, about 0.5 as long as spur; hind calcar triangular, slightly larger than middle one. This species is marked by the complete absence of posterior scarp of the antennal lobe. The latter connects to the narrow, convex anterior portion of the temporal lobe, forming an angular pollinose ridge. The median lobe is pollinose. O. rimatus is similar, but has a glabrous median lobe and a distinct scarp posterior to the antennal lobe. O. inermis is intermediate in having an indistinct scarp posterior to the antennal lobe. It has the median lobe pollinose, as in the present species, but differs from both O. ineditus and O. rimatus in the male lacking the proximal tooth on the anterior tibia. Range. - Western Borneo, western Java, and eastern Sumatra. We have studied specimens from the following localities: BORNEO; one female, Mowong, West Borneo, coll. F. Muir, Sept. 1907 (BMNH); JAVA: one male, Pangalengen, elev. 4000 feet, 1893, coll. H. Fruhstorfer (MNHN); one male, Java-Orient (MNHN); SARAWAK: one female, Mount Matang, West Sarawak, coll. G. K. Bryant, January, 1914 (BMNH); SUMATRA: ten males, four females, Palembang (MNHN); two males, one female, Palembang (BMNH); one female, Palembang (AMS). Variation. - The two specimens from Borneo lack punctures of the inner carina, and might not be conspecific with the others, which have four to eight such punctures. Omoglymmius (Hemiglymmius) rimatus new species Figs. 7, 15 Type material. - HOLOTYPE male, labelled: “JAVA, Mt. Kawi” (MNHN). PARATYPES two females (one mounted on same pin as holotype), same label as holotype (MNHN); one female labelled: “Mt. Kawi, Java, A. Grouvelle” (LEI); two females, labelled: “Mt. Kawi, Java, Juli 1934, v. Doesburg” (LEI). Description. — Length 6. 2-8. 8 mm. Basal setae of antenna well-developed on Segments IV-X; Segents I- 1 V each with dorsal pollinose band; Segments V-X coarsely punctate; Segment XI distinctly longer than wide, corpus nearly cylindrical, narrower than Segment X; stylet short, acute. Head as long as wide; median lobe glabrous, impunctate or with a few coarse punctures; median lobe narrow, margins nearly parallel, tip bluntly rounded at level of middle of eye; frontal space 1.5 wider than long; medial angles obtuse, contiguous or slightly separated; temporal lobe convex; its anterior 0.33 pollinose; posterior portion glabrous, shining with few punctures; temporal seta absent in most specimens, a few specimens with small temporal seta; antennal lobe glabrous, raised above level of temporal lobe and separated from it by distinct scarp; pollinosity of temporal lobe in most specimens ended posteriorly, opposite middle of eye; lateral margin of head deeply sinuate between antennal lobe and eye, postantennal groove oblique; postorbit pollinose; postorbital tubercle about 0.3 as long as eye. Pronotum with length/greatest width about 1.23; widest near middle; apex strongly narrowed, base moderately narrower; margin scarcely sinuate anterior to hind angle; pronotal carinae convex, outer and inner ones nearly equal in width at middle; outer carina with 20-25 punctures; inner carina with 11-14 punctures. Elytral striae broad, rather shallow; intervals convex, narrow, each narrower than one strial puncture; strial punctures coarse, close together; Stria 11 with single seta at apex; Stria IV with two setae at apex; subapical striole with single seta; Stria VII with several setae near apex; abdominal sterna moderately densely punctate: each half of Sternum V with 27-37 Quaest. Ent., 1982, 18 (1-4) 140 Bell and Bell punctures; male with ventral tooth on anterior femur and proximal tooth on anterior tibia; middle calcar acute, about O. 5 as long as spur; hind calcar triangular, its proximal margin concave, smaller than middle one. This species has a distinct postantennal groove and extensive pollinosity on the temporal lobe as in O. ineditus , but differs sharply in having the median lobe glabrous and the antennal lobe glabrous, strongly elevated, and separated from the temporal lobe by a distinct scarp. Range. - West Central Sumatra and eastern Java. This species appears to be allopatric to O. ineditus, and to occupy a markedly disjunct range. In addition to the type series, we have seen specimens from the following localities: JAVA; five males, five females, Gg. Moeria, Tjolo, elev. 700-1000 m., P. H. v. Doesburg (LEI); one male, two females, Mt. Moeria, 3-4000' (MNHN); three males, two females (on same pin), Malang (MNHN); SUMATRA: two males, one female, Pajakombo (MNHN). Variation. - The series of specimens from Gunong Moeria have the pollinosity of the temporal lobe less extensive than in specimens from other localities. Omoglymmius ( Hemiglymmius ) inermis new species Figs. 8,16 Type material. - HOLOTYPE male, labelled: “SUMATRA” (MNHN). PARATYPE male, labelled: “SUMATRA, don. Grouvelle 1901” (GEN). Description. — Length 7.0 mm. Basal setae well-developed on antennal Segments V-X; Segments I- 1 V each with pollinose dorsal band; Segments V-X coarsely punctate; Segment XI distinctly longer than wide, corpus nearly cylindrical, narrower than Segment X; stylet short, acute. Head slightly wider than long, slightly more transverse than in preceding species, median lobe pollinose, impunctate, narrow, its sides nearly parallel, its tip bluntly rounded at level of middle of eye; frontal space 1.5 wider than long; temporal lobe convex, its anterior half pollinose, a rather broad band of pollinosity extended posteriorly medial to eye; medial angles obtuse, contiguous; posterior half of temporal lobe otherwise glabrous, shining with few fine punctures; temporal seta absent; antennal lobe pollinose except for narrow area along lateral margin; antennal lobe continuous with temporal lobe, not marked off by a well-defined scarp, though fine trace of scarp is present; lateral margin of head distinctly sinuate between antennal lobe and eye; postorbit pollinose; postorbital tubercle small. Pronotum more elongate than in O. ineditus , length/greatest width 1.36; widest near middle, apex strongly narrowed, base more narrowed than in O. ineditus , lateral margin not sinuate anterior to hind angle; median groove narrow; pronotal carinae convex, nearly equal, punctate; inner carina more sparsely punctate than outer one. Elytral striae broad, rather shallow; intervals convex, narrow; each interval narrower than a strial puncture; strial punctures coarse, closely spaced; two setae in apex of Stria II; one in apex of Stria IV; one in apical stride; several in apex of Stria VII; abdominal sterna coarsely punctate; 41 coarse punctures in left half of Sternum V, additional very minute punctures present on sterna; male with ventral tooth on anterior femur, but without proximal tooth on anterior tibia; calcars. very small. This species is very similar to the two preceding ones, from which it differs in the absence of the proximal tooth on the anterior tibia of the male. It is also larger than the males of the sympatric O. ineditus, and in addition, has a more elongate pronotum and smaller calcars. The female is unknown. Subgenus Boreoglymmius new subgenus Type species. - Rhysodes americanus Castelnau 1836 Description. — Basal setae on antennal Segments V-X, antennal stylet absent; outer segments not at all punctate; Segment I finely pollinose, others glabrous; clypeal setae absent; eye large, normal, deeper than long; cornea facetted; temporal setae absent; frontal grooves deep, complete; medial angle simple, no translucent area; postorbital tubercle present or absent; marginal groove of pronotum deep; spur of middle tibia curved anteriorly; punctures of abdominal Sterna III-V scattered; lateral pits of female varying, either limited to abdominal Sternum IV and shallow, or else deep and extended over Sterna III and IV; anterior femur of male with ventral tooth; anterior tibia of male without proximal tooth; hind calcars of male larger than middle ones. This subgenus resembles Hemiglymmius in most respects, but differs sharply in lacking the antennal stylet. It is also close to Omoglymmius s. str. from which it differs in presence of basal Revision of Omoglymmius Ganglbauer 141 setae on antennal segments. Three allopatric species are known, two from North America, and one from Japan. Phytogeny. - O. hamatus is structurally as well as geographically intermediate between the two other species. It resembles O. americanus in the presence of a small, shallow lateral pit restricted to Sternum IV in the female, and in the absence of a postorbital tubercle. O. lewisi, in contrast, has a deep lateral pit extended over Sterna 111 and IV, and has a well-developed postorbital tubercle. On the other hand, O. hamatus and O. lewisi are very similar in the shape of the temporal lobe and of the hind calcar. Two possible phylogenies seem worthy of consideration: either O. lewisi and O. hamatus share an ancestor, and O. americanus is more distantly related, or else O. hamatus and O. americanus are more closely related than either is to O. lewisi. We believe that the latter phylogeny is the more likely one. The presence of a postorbital tubercle in O. lewisi is a similarity to Hemiglymmius. According to this hypothesis, the ancestral species had most of the characters of O. hamatus except that the postorbital tubercle was present. Perhaps this species evolved in eastern Asia. Later it invaded western North America, and still later it split into two daughter species after the range was interrupted at the Bering Strait. The population that remained in Asia developed enlarged lateral pits on the abdominal sterna of the female, and became O. lewisi. The population in western North America lost the postorbital tubercle. This species later split into two: O. hamatus , essentially like the ancestral North American species, and O. americanus , in eastern North America, which evolved a shorter, wider head, and which, in the male, had the hind calcars secondarily simplified. According to this view, the similarites between O. americanus and O. (s. str.j germari of Europe represent evolutionary convergence. KEY TO SPECIES 1 Postorbital tubercle present; female with deep lateral pit, extending over abdominal Sterna III and IV O. lewisi (Nakane), p. 141 Y Postorbital tubercle absent; lateral pit of female confined to Sternum IV, small and shallow 2 2 (L) Frontal space twice as wide as long; medial angles of temporal lobe narrow, obtuse, nearly contiguous; posteriomedial margin distinctly concave between medial and occipital angles O. americanus (Castelnau), p. 144 2' Frontal space at most slightly wider than long; medial angles broadly rounded; posteriomedial margin not emarginate O. hamatus (Leconte), p. 143 Omoglymmius (Boreoglymmius) lewisi (Nakane) NEW COMBINATION Figs. 17, 20,21,24 Rhysodes lewisi Nakane 1973: 5 (described in Japanese and designated by the author as a manuscript name.) Omoglymmius (Hemiglymmius) lewisi (Nakane) Bell and Bell 1978 Omoglymmius lewisi Nakane 1978: 129-130 (described in English) Type material. - HOLOTYPE male, Kirishima, Kyushu, Japan, 26- VII- 1 97 1 , T. Nakane lgt. ALLOTYPE female, Kurama, Kyoto, Honshu, Japan, 22-IV-1952, F. Takahashi lgt. Quaest. Ent., 1982, 18 (1-4) 142 Bell and Bell Plate 2. Figures 17-26, subgenus Boreoglymmius. Figs. 17-19, Head and pronotum, dorsal aspect; Fig. 17, Omoglymmius ( Boreoglymmius ) lewisi (Nakane); Fig. 18, O. (B.) hamatus (Leconte); Fig. 19, O. (B.) americanus (Castelnau); Fig. 20, Metasternum and abdomen, ventral aspect, O. (B.) lewisi (Nakane); Figs. 21-23, Head, lateral aspect; Fig. 21, O. (B.) lewisi (Nakane); Fig. 22, O. (B.) hamatus (Leconte); Fig. 23, O. (B.) americanus (Castelnau); Figs. 24-26, Hind tibia, male, apical portion; Fig. 24, O. (B.) lewisi (Nakane); Fig. 25, O. (B.) hamatus (Leconte); Fig. 26, O. (B.) americanus (Castelnau). PARATYPES one male, one female, Fujimidai, Shinano, Honshu, Japan, 1 1 -VI- 1 944, S. Osawa lgt.; one male, Kurama, Kyoto, Honshu, Japan, 1 1 -VII- 1956, T. Nakane lgt.; one male, Mt. Kooya, Wakayama, Honshu, Japan, 1 1 - VII- 1 948. M. Hayashi lgt. (all KU). We have not seen the holotype, but have studied excellent drawings kindly furnished to us by Dr. Nakane. Description. — Length 6. 5-8.0 mm. Antennal Segment I coarsely punctate; head distinctly longer than wide; clypeus with few punctures near margin, continuous with median lobe; latter rhomboidal, narrow, pointed posteriorly even with anterior third of eye; temporal lobe coarsely, very sparsely punctate, punctures very scarce in medial half; anteriomedial margin oblique, scarcely sinuate; medial angles obtusely rounded, contiguous; frontal space longer than wide; posteriomedial margin slightly convexly curved; occipital angle distinct; posteriolateral margin oblique between occipital angle and eye; orbital groove with pollinose strip narrowed posteriorly, extended about to middle of eye; postorbital tubercle with well-marked dorsal margin, but with ventral margin indistinct; postorbit with few coarse Revision of Omoglymmius Ganglbauer 143 punctures near eye, not pollinose. Pronotum moderately elongate, length/greatest width 1.2; widest near middle, narrowed both anteriorly and posteriorly, margin sinuate anterior to hind angle; median and paramedian grooves relatively narrow, punctate; carinae broad; marginal groove fine, linear; outer carina punctate throughout; inner carina with few punctures near base. Elytra long, narrow, intervals narrow, slightly convex; striae relatively shallow; punctures fine; Stria IV with one or two setae near apex; apical stride with two setae; several setae near apex of Stria VII; male with middle calcar acute; hind calcar large, almost twice as wide as tip of tibia, its tip acute, its dorsal side with pronounced “shoulder” with group of large setae (Fig. 24); lateral abdominal pit of female, deep, long, narrow, extended over Sterna Ill-IV (Fig. 20). This species differs from other members of the subgenus in presence of a postorbital tubercle and in form of the lateral pit of the female. Range. - Restricted to Japan. We have studied the following specimens: one male, one female, Mt. Kooya (Wakayama Prefecture), VII-1 1-1948, coll. Masao Hayasi (SATO); one female, Mie University Forest, Ichishi-Gun (Mie Prefecture), 24-VI-1956, coll. M. Sato (SATO). In addition, we have seen one male labelled: “JAPAN: G. Lewis, 1910-320, C. F. Baker Colin. 1927” (NMNH). This specimen is labelled as a “cotype” of Rhysodes crassiusculus Lewis.. This can be a true cotype if the date on the label represents the acquisition date and not the collection date. The description of R. crassiusculus Lewis seems to be a composite, as the description of the head seems to be based on the present species, while the secondary sexual characters are clearly those of the Japanese species of subgenus Pyxiglymmius. We follow Nakane in restricting the name R. crassiusculus to the latter species. Omoglymmius ( Boreoglymmius ) hamatus (Leconte) NEW COMBINATION Figs. 18,22, 25 Rhysodes aratus Chevrolat (nec Newman) 1873a: 209. Rhysodes hamatus Leconte 1875: 163. Type material. - HOLOTYPE, sex not recorded, labelled: “CALIFORNIA” (MCZ, 6830). Description. — Length 6. 2-8. 8 mm. Antennal Segment I coarsely punctate in most specimens (few specimens with reduced punctation), finely pollinose; head longer than wide; clypeus with few punctures near margin; clypeus continuous with median lobe; latter rhomboidal, narrow, pointed posteriorly, tip even with anterior third of eye; temporal lobe coarsely, rather sparsely punctate; anteriomedial margin oblique, only shallowly sinuate; medial angles obtusely rounded, contiguous or narrowly separated; frontal space about as wide as long; posteriomedial margin convexly curved; occipital angle obtusely rounded; posteriolateral margin oblique to eye; orbital groove short, narrow, its posterior end opposite middle of eye; postorbit convex, but not in form of distinct tubercle; postorbit finely pollinose. Pronotum longer and narrower than in related species, length/greatest width 1.33 or greater; both base and apex distinctly narrowed; lateral margin not sinuate anterior to hind angle, both inner and outer carinae sparsely, rather coarsely punctate. Elytra long, parallel-sided; striae rather deep, coarsely punctate; each puncture about as wide as an interval; Stria IV with one to three setae near apex; apical striole with one seta; Stria VII with several setae near apex; male with hind calcar much larger than middle calcar, with pronounced “shoulder” on dorsal surface, separated from shaft of tibia by deep emargination (Fig. 25); female with lateral pit of abdomen on Sternum IV, shallow, ill-defined. This species is distinguished from O. lewisi by absence of the postorbital tubercle, and form of the hind calcar of the male and of the lateral abdominal pit of the female. Compared to O. americanus , this species is much narrower and more elongate, and with better developed punctation. Available bionomic information was summarized by Bell (1970). Range. — Widespread in forested regions of California and Oregon. In California, represented by many specimens from the Sierra Nevada and in the mountains north of Los Angeles, as well as by a few records from the Coast Range north of San Fancisco. In Oregon, known from many localities in the Cascade Range, and from isolated mountain ranges in southcentral and northeastern parts of the state, but not recorded from the Coast Range. There are also records from several localities in the northern third of Idaho. There is a single record from southeastern Washington, and another, quite isolated one, from the Huachuca Mountains of southeastern Arizona. There are also records without specific localities from British Columbia, Nevada, and Texas. Quaest. Ent., 1982, 18 (1-4) 144 Bell and Bell We have seen specimens from the following localities: ARIZONA: Huachuca Mountains (Cochise County) (CNHM). CALIFORNIA: Ahwahnee (CNHM); Alhambra (CAS); Areata (LA); Arrowhead (LA, CAG); Bartlett Springs (CAS); Barton Flats (San Bernardino Co.) (UCD, OS, CAG); Bass Lake (LA, CAS); Bear Valley (Santa Barbara Co.)(CAS); Big Bear Lake (San Bernardino Co.), 6700 ft. (LA); Big Meadows (UCD); Big Tree Grove (MCZ); Big Trees (Calaveras Co.) (UK, CU, AP, CAS); Blodgett Forest (El Dorado Co.), 4400 ft. (LA); Buck’s Lake (Plumas Co.) (UCD); Butte Co. (CNHM); Calaveras Co. (CU); Calistoga (CAS); Camino (CAG); Camp Connell (Calaveras Co.) (CAG); Carrville (Trinity Co.), 2500 ft. (LA, CAS); Cascade Lake (El Dorado Co.) (BSR1); Castle Crag (CMP, MCZ); Cedar Pass (Modoc Co.) (LA, UCD); China Flats (El Dorado Co.) (CAS; Chester (CNHM); Chiquito Creek (Madera Co.), 4100 ft. (CU, UCD); Cisco (MNHB); Cole (CAS); Coleville (CNHM); Coffey Creek (Trinity Co.) (UCD); Dalton Creek (Fresno Co.), 4800 ft. (CAS, CU); Dardanelle (Tuolumne Co.) (LA); Dark Creek (San Jacinto Mts.) (LA); East Crag (CNHM); Desolation Valley, Echo Lake (El Dorado Co.), 7500 ft. (CU, BSRI); El Dorado Co. (CNHM); Facht (Lassen Co.) (CAS); Fallen Leaf Lake (El Dorado Co.) (CAS); Feather Falls (Butte Co.) (LA); Fish Camp (MCZ); Fresno Flats, 3500 ft. (ISNHS); Georgetown (CAG); Giant Forest, 6400 ft. (CU); Grass Valley (LA); Hackamore (OSFS); Happy Camp (MCZ); Hat Creek P.O. (Shasta Co.) (LA); Hassock Meadows (Tulare Co.) (CAS); Hayfork (CAS); Hobart Mills (CAG); Huntington Lake (UCD); Ice House (El Dorado Co.) (CAG); Kaweah (CAS); Kyburz (El Dorado Co.) (UCD); Lake Almanor (Plumas Co.) (CAG); Lake Tahoe, 6465 ft. (LA, PA); Leland Meadows (Tuolumne Co.) (UCD); Madera Co. (CAS); Manzanita Creek (Lassen Co.) (UCD); Mariposa Co. (CNHM); Mather (Tuolumne Co.) (OSU, UCD, WRS); Meadow Creek (Plumas Co.) (CAS); Mendocino Co. (CNHM): Miami (Madera Co.) (CU, AP, LA); Mineral (UCD); Mokelumne (CAS); Morgan Summit (Tehama Co.) (CAG); Mount Aukum (El Dorado Co.) (LA); Mount Home (Tulare Co.) (UCD); Northfork (CU); Old Station (Shasta Co.) (LA); Paradise Valley (King’s River), 7000 ft. (CAS); Pasadena (MO); Peanut (Trinity Co.) (LA); Pinecrest (Tuolumne Co.) (UT); Placer Co. (MO, CNHM, PA); Plumas Co. (CNHM); Pollock Pines (LA); Pyramid Ranger Station (OSFS); Rainbow Falls (Madera Co.) (UCD); Red’s Meadow (Madera Co.) (LA); Riverton (El Dorado Co.) (CAG); San Bernardino Mts. (MCZ); Sequoia National Park (CAS, CAG); Shasta Co. (CNHM); Shasta Retreat (Siskiyou Co.) (CAS); Sierraville (CAG); Sissons (Siskiyou Co.) (MO, CNHM, MCZ); Strawberry (El Dorado Co.) (CAS, LA); Sugar Pine (CAS); Tahquitz Valley (San Jacinto Mts.) (LA); Tallac (CAS, SD); Trinity Co. (CNHM); Truckee (CAS); Tulare Co. (CNHM, MCZ); Tuolumne Co. (CNHM); Uncle Tom’s (CAG); Viola (LA, CAS); Virner (CAG); Walker Mine (OSFS); Weaverville (CAS); West Fork (San Gabriel Canyon) (UT); Whitehall (El Dorado Co.) (LA); Yosemite (UK, LA, CAS); Yuba pass (Sierra Co.) (LA, CAG). CANADA: “B.C.” (CMP). IDAHO: Clearwater (Idaho Co.) (DY); Coeur d’ Alene (CAS); Harvard (Latah Co.) (WS); Moscow (WS); Moscow Mt. (WS). NEVADA: “Nevada” (MNHB). OREGON: Beaver-Sulfur (Jackson Co.) (OS); Brownsboro (OS); Butte Falls (Jackson Co.) (UCS, OS); Cave State Park, 5 mi. nw of Culver, 1775 ft. (UT); Colton (CAS); Deming Creek (near Bly) (MSU); Fish Lake (Jackson Co.) (UCD); Huckleberry Mt. (Jackson Co.), 5500 ft. (CNHM); Indian Fork Creek (near Sisters) (UT, OS); Josephine Co. (CNHM); Klamath Co. (CNHM, OS); Klamath Falls (MSU); Lake of the Woods (Klamath Co.), 5300 ft. (CNHM); Mahama (Marion Co.) (LA); McAllister Soda Springs (Jackson Co.) (CNHM); McDonald Forest (Benton Co.) (OS); Moon Prairie (OS); Portland (CAS); Ruch (Jackson Co.) (LA); Union (CNHM); Union Creek (Jackson Co.), 3100-3500 ft. (CNHM); Upper Klamath Lake (MUS); Wildwood Camp (Ochoco National Forest) (OS). TEXAS: “Tex.” (MN). WASHINGTON: Dayton(WS). Omoglymmius (Boreoglymmius) americanus (Castelnau) NEW COMBINATION Figs. 19, 23,26 Rhysodes americanus Castelnau 1836: 58. Rhysodes exaratus Serville 1825: 308 (nec Dalman 1823; nec Erichson 1848). Rhysodes aratus Newman 1838: 664. Omoglymmius americanus (Castelnau) Bell 1975: 351. Type material. - HOLOTYPE male, labelled: “America Sept., De Castelnau 2-43” (MNHN). Description. — Length 6. 0-7. 9 mm. Segment I of antenna pollinose, impunctate or with very few punctures; head as long as wide; clypeus impunctate, continuous with median lobe; latter relatively broad, rhomboid, tip pointed opposite anterior third of eye; temporal lobe very finely punctate, impunctate in appearance at low magnifiction; anteriomedial margin very deeply sinuate, anterior half longitudinal, posterior half oblique; medial angles narrow, obtusely pointed, contiguous; frontal space twice as wide as long; posteriomedial margin shallowly emarginate posterior to obtuse but Revision of Omoglymmius Ganglbauer 145 distinct occipital angle; lateral margin slightly curved between occipital angle and eye; orbital groove relatively well-developed, extended to posterior margin of eye; postorbit flat, pollinose, without trace of tubercle. Pronotum relatively short and broad, length/greatest width is 1.09; widest near middle, both base and apex narrowed; apex less narrowed than in related species; pronotal carinae with very fine punctures, impunctate in appearance at low magnification; elytra broader and less parallel-sided than in O. hamatus-, strial punctures less coarse than in O. hamatus , each puncture clearly narrower than an interval; Stria IV with one to four setae near apex; hind calcar of male acute, triangular, not “shouldered” dorsally (Fig. 26); lateral abdominal pits of female as in O. hamatus. This species differs from other members of the subgenus in having the hind calcar triangular. It is easily distinguished from O. hamatus by broad frontal space, deeply emarginate anteriomedial margin of the temporal lobe, emarginate posteriomedial margin, and much finer punctures of temporal lobes and pronotal carinae. Available bionomic information was summarized by Bell (1970). Range. - Eastern North America, north to central New York, southern Ontario, and the vicinity of Minneapolis, Minnesota; west to southeastern Nebraska and eastern Texas; south to the Gulf of Mexico and northern Florida. Two specimens labelled “Arizona” (ISNHS) are probably mislabelled, as is certain for a specimen labelled “Nederl. Indie, A. Roller, Colin. Drescher” (AMS). A specimen labelled “Castle Crag, California” (CMP) is also very likely mislabelled. Since Bell (1970) we have seen specimens from the following additional localities: ALABAMA: Auburn (BSRI); Chiat State Park (UW); Mobile (MO). ARKANSAS: Arkadelphia (CNHM); Bluff City (Ouachita Co.) (ARK); Rosston (Nevada Co.) (ARK). DELAWARE: “Del.” (MCZ). FLORIDA: Alachua Co. (UM); Florida Caverns State Park (Jackson Co.) (FLA); Gainesville (CNHM). ILLINOIS: Algonquin (ISNHS); Carbondale (SI); Crab Orchard Lake (Williamson Co.) (SI); East Dubuque (ISNHS); Evanston (CNHM); Funk’s Grove (ISNHS); Galena (ISNHS); Harrisburg (ISNHS); Lusk Creek (Pope Co.) (SI); Moline (MN); Murphysboro (SI); New Lennox (CNHM); Normal (ISNHS); Pine Hills Field Station (Union Co.) (BSRI): Pulaski (ISNHS, CNHM); Urbana (CNHM), 10); Utica (UVM); White Heath (UM). INDIANA: Bloomington (IU); Brown County State Park (IU); Crowthersville (PU, FLA); Dubois County (PU); Indianapolis (FLA); Jennings Co. (PU); Knox Co. (PU); Kosciusko Co. (CU); Lafayette (PU); Lagrange Co. (PU); Posey Co. (PU); Smith (CNHM); Smith Station (La Porte Co.) (CNHM); Vigo Co. (PU). IOWA: Ames (IO); Jefferson Co. (UVM). KANSAS: Douglas Co., 900 ft. (MCZ, UK, CU, MNHN); Riley Co. (CNHM). KENTUCKY: Anchorage (UL). LOUISIANA: Baton Rouge (LS); Chastine (Natchitoches Parish) (MO); Magnolia (LS); Natchitoches (MO); Opelousas (MNHN): Ruston (LS). MARYLAND: College Park (LA); Elk Neck State Park (UD); Sparrows Point (CNHM). MICHIGAN: Allegan Co. (MSU); Ann Arbor (UN); Clinton Co. (HL, UVM); Detroit (MO); Frazer (MSU); Grand Rapids (UN); Kalamazoo (MCZ); Lansing (MSU); Washtenaw Co. (UM). MINNESOTA: Hennepin Co. (MN); Ramsay Co. (MN). MISSISSIPPI: Avera (CU); Chocktaw Co. (RCG); Greer (UW); Lafayette Co. (BSRI); New Augusta (CU). MISSOURI: Ashland (MO); Columbia (MO); Louisiana (MO); McBaine (MO); Meramee Highland (St. Louis Co.) (MO); Saint Louis Co. (CMP, MO); Sikeston (MO); Warrensburg (CNHM). NEBRASKA: Ashland (UN); Lincoln (UN). NEW JERSEY: “N.J.” (MCZ). NEW YORK: Buffalo (CNHM); Lodus (MCZ); McLean (Tompkins Co.) (CU); McLean Bogs (Cortland Co.) (CU); Niagara (CNHM). NORTH CAROLINA: Southern Pines (UI, NMNH). OHIO: Bowling Green (RCG); Cedar Swamp (Champaign Co.) (FLA); Columbus (MSU); Dayton (DM); Delaware Co. (WS); Franklin Co. (NC); Montgomery Co. (DM); Oxford (NC); Wood Co. (RCG). OKLAHOMA: Bradem (OK). ONTARIO: Essex Co. (KS, RCG); Kingsville (CNHM); Port Alma (WR); Tilbury (UVM, KS). PENNSYLVANIA: Allegheny (CMP); Harrisburg (AP); Jeanette (CMP); New Cumberland (AP); Norwich (MNHB); Pittsburgh (CMP). SOUTH CAROLINA: Clemson (UVM); Florence (SDA); Sumter (UVM); Wedgefield (UVM). TENNESSEE: Cookeville (TB); Madison Co. (CU, MCZ); Shelby Co. (DY). TEXAS: Nacogdoches (AU). VIRGINIA: Blacksburg (VP, NMNH); Great Falls (NMNH); Mount Vernon (NMNH); Warrenton (UK). WISCONSIN: Broadhead (Green Co.) (UW). Quaest. Ent., 1982, 18 (1-4) 146 Bell and Bell Subgenus Pyxiglymmius Bell and Bell 1978 NEW STATUS Type species. - Rhysodes strabus Newman 1838. Description. — Eleventh antennal segment without true stylet, its apex in most species obtuse (apex acute and resembling a stylet in O. cristatus)\ basal setae on at least antennal Segments VIII-X (on VI-X in most species); clypeal setae absent; frontal grooves deep, complete; medial margin of temporal lobe with two medial angles well separated from one another, emargination between them deep except in O. lederi-, medial margin without distinct translucent area (in some specimens of O. strabus, slight suggestion of translucent area); eye large, deeper than long; cornea facetted; postorbital tubercle present; gular tubercle prominent except in O. crassiusculus ; punctures of abdominal sterna scattered; female with prominent lateral pit in Sternum IV; in most species pit complex, with anteriolateral ridge or brace, and with longitudinal striation; spur of middle tibia curved anteriorly at apex; male with proximal tooth on anterior tibia and ventral tooth on anterior femur; calcars large, conspicuous, that of middle tibia slender, nearly cylindrical, tibial spur apparently inserted in its base; hind calcar very large, compressed, triangular, separated from spur by distinct emargination; hind tibia of male bent near middle, swollen proximad to middle. In this subgenus there are two medial angles on each temporal lobe. This feature separates it from all other subgenera except for some species of Laminoglymmius. Members of the latter subgenus have the medial angles much closer together and, in addition, all species except O. actae have a conspicuous translucent area in the region of the medial angles. This taxon was ranked as a genus in Part I. Further study has convinced us that it should be reduced to subgeneric status. O. rugosus and O. insularis have been removed to Laminoglymmius New Subgenus. The range of Pyxiglymmius extends through most of the Oriental Region, from Japan and Viet Nam to Java, Sumatra, Borneo, and the Andaman and Mentawei Islands. It is also represented on Luzon in the Philippine Islands. Phylogeny. - A phylogenetic tree for the subgenus is presented in Diagram 1. We postulate that the common ancestor of the subgenus, Species 1, had the following characters: median lobe longer than wide, acutely pointed; medial margin of temporal lobe deeply emarginate; female with lateral groove on metasternum; lateral abdominal pits of female with small anteriolateral brace, but without striations. Six of the nine species clearly form a monophyletic group, descended from Species 3. In the latter species, the median lobe became markedly transverse, a specialization not seen in any of the other subgenera of Omoglymmius. In the six species of this group, the postorbit is flat, and is bounded ventrally by a distinct suborbital tubercle. The three remaining species, crassiusculus, subcaviceps , and lederi, agree in having the median lobe elongate (as is usual in Omoglymmius s. lat .), and in having the postorbit strongly convex and not bounded ventrally by a suborbital tubercle. They may have arisen from a common ancestor (Species 2), although none of the common characters is clearly a synapomorphy. It is also possible that they descended separately from Species 1. O. crassiusculus and O. lederi lack the lateral groove on the metasternum of the female. We believe this is a secondary loss. It might be a synapomorphy. O. subcaviceps has a well-developed lateral metasternal groove as in the remainder of the subgenus. Of the line descended from Species 3, O. cristatus of Luzon possibly stands apart from the others. The pointed apical antennal segment and reduced metasternal grooves are clearly specializations, while the absence of striations from the lateral abdominal pit may represent a character state more primitive than that in the remaining species. The remaining species were descended from Species 4, in which deep lateral grooves were retained on the metasternum of the female and the lateral abdominal pits had become longitudinally striate. We hypothesize two daughter species to Species 4. One of these. Species 5, greatly increased the number of setae on the elytra, giving rise to O. pilosus and O. krikkeni. Revision of Omoglymmius Ganglbauer 147 KRIKKENI PILOSUS HESPERUS ARMATUS STRABUS CRISTATUS LEDERI CRASSIUSCULUS SUBCAVICEPS Phylogenetic Diagram 1. Reconstructed Phylogeny of species of Pyxiglymmius. On the other hand, it is possible that supernumerary setae developed independently, correlated with large size (cf. O. ( Laminoglymmius ) rugosus ). In Species 6, the number of setae on the elytron did not increase. It gave rise to O. strabus, O. armatus, and the aberrant O. hesperus. O. strabus has a wide range, including Sumatra, Borneo, the Malaysian Peninsula and Java. The two remaing species appear to represent offshoots of O. strabus which have speciated on isolated islands (the Andamans and Nicobars in the case of O. armatus and the Mentawei Islands for O. hesperus). Quaest. Ent., 1982, 18 (1-4) 148 Bell and Bell Plate 3. Figures 27-40, Subgenus Pyxiglymmius. Figs. 27-32, Head and pronotum, dorsal aspect; Fig. 27, Omoglymmius (Pyxiglymmius) subcaviceps (Grouvelle); Fig. 28, O. (P.) crassiusculus (Lewis); Fig. 29, O. (P.) lederi (Lewis); Fig. 30, O. (P.) cristatus new species; Fig. 31,0. (P.) strabus (Newman); Fig. 32, O. (P.) armatus (Arrow); Figs. 33-38, Head, lateral aspect; Fig. 33, O. (P.) subcaviceps (Grouvelle); Fig. 34, O. (P.) crassiusculus (Lewis); Fig. 35, O. (P.) lederi (Lewis); Fig. 36, O. (P.) cristatus new species; Fig. 37, O. (P.) strabus (Newman); Fig. 38, O. (P.) armatus (Arrow); Fig. 39, Hind tibia, male; O. (P.) subcaviceps (Grouvelle); Fig. 40, Antennal Segment XI, O. (P.) cristatus, new species; Revision of Omoglymmius Ganglbauer 149 Plate 4. Figures 41-56, Subgenus Pyxiglymmius. Figs. 41-43, head and pronotum, dorsal aspect; Fig. 41, Omoglymmius (Pyxiglummius) hesperus new species; Fig. 42, O. (P.) pilosus (Grouvelle); Fig. 43, O. (P.) krikkeni new species; Figs. 44-46, Head, lateral aspect; Fig. 44, O. (P.) hesperus new species; Fig. 45, O. (P.) pilosus (Grouvelle); Fig. 46, O. (P.) krikkeni new species; Figs. 47-49, O. (P.) pilosus (Grouvelle), male; Fig. 47, Femur, tibia, anterior leg; Fig. 48, Middle tibia, apical portion; Fig. 49, Femur, tibia, hind leg; Fig. 50, Metasternum and abdomen, ventral aspect, O. (P.) pilosus (Grouvelle); Fig. 51 Metasternum and abdomen, right half, O. (P.) krikkeni new species; Fig. 52, Right elytron, dorsal aspect, O. (P.) krikkeni new species; Fig. 53, Elytron, metasternum, abdomen, left lateral aspect, O. (P.) krikkeni new species; Figs. 54-56, O. (P.) krikkeni new species, male; Fig. 54, Femur, tibia, anterior leg; Fig. 55, Middle tibia; Fig. 56, Hind tibia. Quaest. Ent., 1982, 18 (1-4) 150 Bell and Bell KEY TO SPECIES 1 Median lobe distinctly longer than wide, its apex opposite middle of eye; postorbit convex, in form of deep but short, ill-defined postorbital tubercle 2 Y Median lobe transverse, as wide as long, its tip opposite anterior margin of eye; postorbit flat, bounded ventrally by well-defined sub- or postorbital tubercle 4 2 (1) Medial emargination of temporal lobe relatively shallow, its depth about 0.25 of its length; basal setae of antenna on Segments VIII-X O. lederi (Lewis), p. 152 2' Medial emargination of temporal lobe deep, its depth 0.5 or more of its length; basal setae on Segments V or VI-X 3 3 (2') Temporal lobe broadly rounded posteriorly; postorbital tubercles scarcely visible in dorsal view; anteriomedial margin of temporal lobe strongly curved O. subcaviceps (Grouvelle), p. 150 3' Temporal lobe with distinct occipital angle, margin markedly oblique between occipital angle and eye; orbital tubercles prominent in dorsal view; anteriomedial margin of temporal lobe oblique . O. crassiusculus (Lewis), p. 151 4 (L) Apex of Segment XI of antenna pointed; metasternum of female without lateral grooves O. cristatus new species, p. 152 A' Apex of Segment XI obtuse; metasternum of female with deep lateral grooves ... 5 5 (4') Setae absent from Stria II; one or two setae present near apex of Stria IV 6 5' Five or more setae each in Striae II, IV 8 6 (5) Outer carina of pronotum distinctly narrower than paramedian groove; outer carina nearly straight, of even width O. armatus (Arrow), p. 154 6' Outer carina wider than paramedian groove, widest near middle, tapered both anteriorly and posteriorly 7 7 (6') Postorbital tubercles large, prominent in dorsal view O. hesperus new species, p. 1 54 7' Postorbital tubercles small, not prominent in dorsal view O. strabus(Nev/man), p. 153 8 (5') All elytral striae setose O. pilosus (Grouvelle), p. 155 8' Stria I and III devoid of setae O.krikkeni new species, p. 156 Omoglymmius (Pyxiglymmius) subcaviceps (Grouvelle) NEW COMBINATION Figs. 27, 33, 39 Rhysodes subcaviceps Grouvelle 1903: 122. Pyxiglymmius subcaviceps (Grouvelle) Bell and Bell 1978 Type material. - HOLOTYPE female, labelled: (VIET NAM) “Tonkin sept. Ha-giang, Frontiere de Chine, coll. A. Weiss, 1901” (MNHN). Description. — Length 7-9 mm. Antennal Segment XI 1.5 longer than wide, tip of cone obtuse; basal setae of antenna sparse on Segment V, well-developed on Segments VI-X; head about as long as wide; median lobe longer than wide, pointed posteriorly, its lateral angles very obtuse; anteriomedial margin strongly curved; frontal space thus U-shaped; first medial angles contiguous, separated from second medial angles by rather small but deep emargination, wich is nearly as deep as long; temporal lobes rounded posteriorly, occipital angles very indistinct; temporal lobe with few fine, irregularly scattered punctures; postorbit convex, in form of a deep but inconspicuous tubercle. Revision of Omoglymmius Ganglbauer 151 Pronotum longer than wide, its greatest width less than width across humeral tubercles of elytra; apex of pronotum strongly narrowed, base moderately narrowed, widest point at middle; lateral margin distinctly sinuate anterior to hind angle; paramedian groove narrow, about 0.33 as wide as inner carina at middle; paramedian groove distinctly but inconspicuously punctate; carinae of equal width, both inner and outer ones finely punctate; marginal groove narrow. Elytra narrow, elongate; apical tubercle very small; Striae III, IV not sinuate posteriorly; strial punctures large, close together, each about as wide as an interval; male with short lateral groove in posterior part of metasternum (in female, a much deeper, conspicuous pit); male with proximal tooth of anterior tibia prominent, near middle of length of tibia; male with acute ventral tooth on anterior femur; middle calcar acute, slightly shorter than spur; calcar of hind leg very large, its dorsal margin slightly convex, its apex forming a right angle (Fig. 39; female with deep lateral pit in Sternum IV; margin of elytron angulate opposite Sternum IV. This species resembles O. crassiusculus in having the postorbit broadly convex, and in having both pairs of medial angles contiguous. It differs from O. crassiusculus in having the temporal lobes broadly rounded posteriorly and in having the anteriomedial margin of the temporal lobe emarginate. The elongate median lobe and the small, contiguous anterior medial angles will separete it from O. strabus and its relatives. Range. - Northern Viet Nam. In addition to the holotype, we have seen: a second specimen from the type locality, collected by Bonifay, 1912 (MNHN) and one male, Hoa Bihn, Tonkin, coll. A. de Cooman, BM 1929-299 (BMNH); two females, N. Viet Nam hills, 50 km. N. Thai-Nguen, 9-1-1964 and 12-1-1964, coll. Kabakov (LEN); two males, same locality, 300 m. 15-12-1092 and 5-2-1963, coll. Kabakov (LEN); one male, S.W. Kui-Tchau, 300 m„ 19-2-1963, coll. Kabakov (LEN); one male, Mts. S.W. Dong-Hai, 23-XI-1961, coll. Kabakov (LEN). Omoglymmius (Pyxiglymmius) crassiusculus (Lewis) NEW COMBINATION Figs. 28, 34 Rhysodes crassiusculus Lewis 1888: 80. Pyxiglymmius crassiusculus (Lewis) Bell and Bell 1978 Type material. - LECTOTYPE (here designated) male, labelled: “(JAPAN) Kika, Chizenji, 17-19 April, 1880” (BMNH) PARALECTOTYPES one male, four females, Hakone (BMNH); one female, Miyanoshita (BMNH); one female, Nikko, Japan, coll. G. Lewis, 1910-320, C. F. Baker Colin. (NMNH); one male, Kiga, Japan, G. Lewis 1910-320; from C. F. Baker, July 1923 (Cotype No. 26386) (NMNH). Dates on NMNH paralectotypes seem to refer to the accession date of the Lewis collection, and thus do not preclude them being part of the original material of Lewis. One specimen labelled as a cotype of R. crassiusculus (NMNH) actually pertains to Omoglymmius (Boreoglymmius) lewisi Nakane. Description. — Length 6. 5-8.0 mm. Antennal Segment XI 1 .5 longer than wide, its tip obtuse; basal setae sparse on Segments V-VI1, numerous on Segments VIII-X; head slightly longer than wide; median lobe longer than wide, pointed posteriorly; anteriomedial margin of temporal lobe oblique, frontal space thus V-shaped; first medial angles contiguous, separated from second medial angles by deep emarginations and are longer than in O. subcaviceps; posteriomedial and posteriolateral margins of temporal lobe oblique, joined at obtuse but distinct occipital angle; temporal lobe with few irregularly scattered punctures; postorbit convex, in form of deep but inconspicuous postorbital tubercle. Pronotum narrow, elongate; length/greatest width 1.22; its width conspicuously less than width across humeral tubercles; apex markedly narrowed; base moderately narrowed; lateral margin distinctly sinuate anterior to hind angles; paramedian groove narrow, less than 0.25 width of inner carina at middle; paramedian groove distinctly but inconspicuously punctate; carinae of equal width, both inner and outer ones finely and irregularly punctate; marginal groove narrow. Elytra narrow, elongate; apical tubercle more distinct than in O. subcaviceps-. Striae IV, V united posteriorly in a shallow depression; strial punctures large, close together, each about as wide as an interval; one seta apex of Stria IV, one in stride, several in VII near apex; metasternum without lateral grooves in either sex; male with proximal tooth of anterior tibia prominent, near middle of length of tibia; male with low, obtuse tooth on ventral margin of anterior femur; middle calcar acute, slightly shorter than spur; hind calcar very large, its dorsal margin straight, its apex in form of acute angle; female with deep, distinct lateral furrow on abdominal Sterna I- 1 1; female with very large lateral pits on Sternum IV, separated medially by less than width of one of them; lateral pit with small, inconspicuous anteriolateral brace; pit smooth, without striae or punctures; ventral margin of elytron conspicuously angulate opposite Sternum IV in female, slightly angulate in male. Quaest. Ent., 1982, 18 (1-4) 152 Bell and Bell The distinct occipital angles give the head of this species a “diamond” shape, separating it from all other members of the subgenus. Range. - Japan. In addition to type material, we have seen one specimen from Shuzenji Izu (CAS). Omoglymmius (Pyxiglymmius) lederi (Lewis) NEW COMBINATION Figs. 29, 35 Rhys odes lederi Lewis 1888: 82. Pyxiglymmius lederi (Lewis) Bell and Bell 1978 Type material. - HOLOTYPE female, labelled: “Caucasus, coll. Saunders, G. Lewis, 1901-31” (BMNH). This distinctive species has not been collected subsequently in the Caucasus, and we suspect that the locality label is erroneous. The species probably really comes from one of the poorly collected countries of the Oriental Region. Description. — Length 8.9 mm. Antennal Segment XI slightly longer than wide; apex of cone obtuse; basal setae sparse on Segments Vlll-IV; numerous on Segment X; head as wide as long; median lobe elongate, narrow, pointed posteriorly; separated from clypeus by shallow transverse impression; anteriomedial margin of temporal lobe oblique; both first and second posterior medial angles narrowly separated; first medial angle separate from second medial angle by broad but relatively shallow emargination which is twice as long as deep; occipital angle indistinct, temporal lobe rounded posteriorly; temporal lobe with a few coarse, irregularly scattered punctures; postorbit convex, in form of deep but inconspicuous postorbital tubercle. Pronotum narrow, elongate; length/greatest width 1 /26; apex markedly narrowed; base moderately so; lateral margin distinctly sinuate anterior to hind angle; paramedian groove narrow, 0.33 as wide as inner carina at middle; carinae of equal width, both outer and inner carinae finely, irregularly punctate, more densely so than in O. crassiusculus\ marginal groove narrow. Elytra narrow, elongate; striae coarsely punctate; no setae in Stria IV, one in apical stride, several in Stria VII near apex; ventral margin of elytron conspicuously angulate opposite lateral abdominal pit in Sternum IV of female; metasternum without lateral groove in female; abdominal Sternum I with lateral pit in female; Sternum IV of female with moderately large lateral pits (pits separated by more than twice the width of one of them); pits of Sternum IV without anteriolateral braces, irregularly pitted, but without longitudinal stria. Male unknown. Omoglymmius ( Pyxiglymmius ) cristatus new species Figs. 30, 36, 40 Type material. - HOLOTYPE female, labelled: “PHILIPPINES, Mt. Makiling, Laguna (Province), 1-1 1931, F. C. Hadden Colin. ”(CAS). The type locality is on Luzon. Description. — Length 8.0 mm. Antennal Segment XI distinctly longer than wide, apex of cone acutely pointed; basal setae numerous on Segments V-X (Fig. 40); head slightly wider than long; antennal lobes more elevated than in other members of subgenus; clypeus coarsely punctate, separated from median lobe by shallow transverse impression; median lobe rhomboidal, wider than long; anteriomedial margin of temporal lobe oblique; first medial angles widely separated; second medial angles contiguous; temporal lobe rounded posteriorly; temporal lobe convex, with about 30 coarse punctures; postorbit flat dorsad to tubercle; latter located more dorsad than in O. strabus and related species; postorbital tubercle about 0.33 as deep and 0.33 times as wide as eye. Pronotum relatively short; length/greatest width 1.16; base of pronotum moderately narrowed; apex very strongly narrowed, lateral margin slightly sinuate anterior to hind angles; paramedian grooves punctate, rather broad, about 0.66 as wide as inner carinae at middle; carinae of equal width at middle, both inner and outer ones coarsely punctate; marginal groove rather fine, about 0.25 as wide as outer carina at middle. Elytra moderately elongate; apical tubercle small; strial punctures very large, each equal in width to an interval; one seta present in apex of Stria IV, one in apical stride; several near apex of Stria VII; female with shallow lateral groove in posterior fourth of lateral margin of metasternum; female with indistinct lateral impression on Sternum I and large, deep lateral pits on Sternum IV, separated medially by less than twice width of one of them; lateral pit of Sternum IV with prominent anteriolateral brace and with confused punctuation, but without longitudinal striation. Ventral margin of female elytron indistinctly angulate opposite Sternum IV. Revision of Omoglymmius Ganglbauer 153 Male unknown. This species is similar to O. strabus, from which it differs in having the apex of the antenna pointed, the head shorter and wider, the antennal lobes more elevated, and the temporal lobes more coarsely and densely punctate. Omoglymmius (Pyxiglymmius) strabus (Newman) NEW COMBINATION Figs. 31, 37 Rhysodes strabus Newman 1838: 663-664. Rhysodes aterrimus Chevrolat 1873: 209. NEW SYNONYMY Pyxiglymmius strabus (Newman) Bell and Bell 1978 Type material. - ( R . strabus) HOLOTYPE female, labelled: “Java, Ent. Club, 44-12” (labelled as type) (BMNH); (R. aterrimus) HOLOTYPE female, labelled: “Malacca, S. S”, “typus” (NMW). Description. — Length 7. 5-9.0 mm. Antennal Segment XI slightly wider than long, its tip obtusely rounded; basal setae numerous on Segments V-X; head slightly wider than long; clypeus with a few coarse punctures or impunctate, continuous with median lobe; median lobe rhomboidal, wider than long; anteriomedial margin of temporal lobe oblique; first medial angles widely separated; second medial angles contiguous or very narrowly separated; temporal lobe rounded posteriorly; temporal lobe convex, with one to 12 coarse punctures; postorbit flat dorsad to tubercle; latter located opposite ventral third of eye; postorbital tubercle about 0.33 as deep and 0.33 as long as eye. Pronotum rather short, length/greatest width ranging from 1.12-1.19; base of pronotum moderately narrowed; apex stongly narrowed; lateral margin slightly sinuate anterior to hind angles; paramedian grooves rather broad, punctate, about 0.66 as wide as inner carina at middle; inner carina slightly narrower than outer carina at middle; both pairs of carinae coarsely punctate, inner ones more sparsely so than outer ones; marginal groove fine, less than 0.25 as wide as outer carina at middle. Elytra moderately elongate; apical tubercle very small; strial punctures large, each as wide as an interval; one seta present in apex of Stria IV; one in apical striole, several near apex of Stria VI 1; female with broad, distinct groove along each lateral margin of metasternum; female with indistinct lateral impression on abdominal Sternum I, and large, deep lateral pits on Sternum IV, which are separated medially by more than twice the width of one of them; lateral pit of Sternum IV with prominent aneriolateral brace and with longitudinal striation; ventral margin of female elytron not angulate opposite Sternum IV; male with well-developed proximal tooth at basal third of anterior tibia; male with ventral tooth on anterior femur; calcar of middle tibia acute, about 0.5 as long as spur; calcar of hind tibia large, triangular, acute, with dorsal margin curved. This species may be separated from the sympatric O. pilosus and O. krikkeni by the small number of setae on the elytra. The type specimens of O. strabus and O. aterrimus at first glance look like separate species, that of O. strabus having the temporal lobes nearly impunctate, while that of O. aterrimus is distinctly punctate. However, numerous specimens from Java show that this is individual variation. O. armatus, O. hesperus , and O. cristatus are similar to O. strabus , and appear to have developed from populations of the latter species isolated respectively on the Andaman Islands, the Mentawei Islands, and Luzon. For differences between O. strabus and each of these species, see discussion of the latter. Range. - Borneo, Java, Sumatra, and the Malay Peninsula. In addition to the types, we have Studied the following specimens: BORNEO: one male, one female, R. Kapah, trib. of R. Tinjar, 3-X, 1932, on bark, felled timber, Oxford Univ. Exp., coll. B. M. Hobby & A. W. Moore, BM 1933-254 (BMNH); JAVA: 14 males, 5 females. Preanger (MNHN); one male, Mt. Tengger (MNHN); two females, without specific locality (LEI); MALAYA: one male, Perak, coll. Doherty, 58794, Fry. Colin. 1905-100 (BMNH); SUMATRA: one male, Alas Valley, Gumpang, foothills of Mt. Kemiri, 10-V1-1972, ca 780 m., coll. J. Krikken (LEI); one female, Palembang (MNHN). Records for O. aterrimus from the Andaman and Nicobar Islands refer to O. armatus. Quaest. Ent., 1982, 18 (1-4) 154 Bell and Bell Omoglymmius (Pyxiglymmius) armatus. (Arrow) NEW COMBINATION Figs. 32, 38 Rhysodes armatus Arrow 1901: 85. Pyxiglymmius armatus (Arrow) Bell and Bell 1978 Grouvelle (1903) synonymized the names R. armatus and R. aterrimus Chevrolat 1873. Bell and Bell (1978) showed this to be an error. Type material. - LECTOTYPE (here designated) male, labelled: “Andamans (Roepstorff)” (BMNH). PARALECTOTYPES one female, same data as lectotype (teneral specimen) (BMNH); one female, labelled: “Nicobars (Roepstorff)” (BMNH). Description. — Length 6. 1-8.0 mm. Antennal Segment Xi slightly longer than wide, its apex obtuse; basal setae sparse on Segment V, numerous on Segments VI-X; head slightly wider than long; clypeus impunctate; separated from median lobe by a more or less distinct transverse impression; median lobe short, strongly transverse, its posterior margin varying from nearly transverse to distinctly angled at middle (i.e., median lobe triangular to rhomboid); anteriomedial margin of temporal lobe oblique; first medial angles separated; second medial angles contiguous; temporal lobe rounded posteriorly; temporal lobe convex, with 10-15 scattered punctures; postorbit flattened dorsad to tubercle; latter more prominent than in O. strabus, about 0.5 as deep and 0.66 as long as eye. Pronotum more elongate than in O. strabus , length/greatest width 1.32, widest anterior to middle; in some specimens distinctly dilated anterior to middle, in others almost parallel-sided; margin various from scarcely to markedly sinuate anterior to hind angles, latter obtuse; paramedian grooves broad, punctate, nearly straight, as wide as or wider than inner carina at middle; both pairs of carinae very narrow; outer carina in some specimens slightly narrower than inner one, in others, opposite; both carinae sparsely, coarsely punctate; marginal groove very deep and dilated, as wide as outer carina or nearly so. Elytra moderately elongate; apical tubercle larger than in O. strabus ; strial punctures large, each equal in width to one interval; one seta in apex of Stria IV, one on apical striole, several near apex of Stria VII. Female with deep lateral grooves on metasternum; lateral pits of Sternum IV of female separated by 1.25 times width of one of them; lateral pit with anteriolateral brace and longitudinal striae. Proximal tooth of anterior tibia present, but various in size and position; ventral tooth on anterior femur present, also various in size and position; middle calcar approximately equal in length to spur; hind tibia various in degree of swelling in proximal third, and especially in size of calcar. The deep, dilated marginal grooves of pronotum are diagnostic, and form a striking contrast with the very fine grooves of O. strabus. There are considerable differences among specimens we assign to O. armatus , including: shape of pronotum, size of postorbital tubercles, depth of lateral metasternal grooves, size of lateral abdominal pits of females; and particularly in size of the hind calcars of males. It seems possible that there are subspecific or perhaps specific differences among the populations on the various islands, but available specimens are too few and too inexactly labelled to allow definite conclusions. Range. - Andaman and Nicobar Islands. In addition to type material, we have seen one male from the Andaman Islands (MNHN), and two males and four females from the Nicobar Islands (MNHN). Omoglymmius ( Pyxiglymmius ) hesperus new species Figs. 41, 44 Type material. - HOLOTYPE male, labelled: “Mentawei:Sipora, Sereinu V-VI-94, Modigliani, 51” (MNHN). This island, also spelled “Sipoera” or “Sipura,” is in the Indian Ocean, west of Sumatra. PARATYPES two females, three males, labelled: “Mentawei, Si Oban, IV-VIII, Modigliani 94” (GEN). Description. — Length 7. 9-9.0 mm. Antennal Segment XI slightly wider than long, its apex obtuse; basal setae absent from Segment V, sparse on VI, VII, numerous on VII-X; head slightly longer than wide, smaller relative to pronotum than in other Pyxiglymmius-, clypeus punctate, pollinose, continuous with median lobe; median lobe rhomboid, wider than long, its posterior angle obtuse; anteriomedial margin of temporal lobe oblique; first medial angles distinctly Revision of Omoglymmius Ganglbauer 155 though narrowly, separated; second medial angles contiguous; occipital angles very obtuse but distinct; posteriolateral margin of temporal lobe slightly oblique; each temporal lobe with seven to 10 coarse punctures; postorbit slightly concave dorsad to postorbital tubercle; latter large, 0.75 as long and 0.66 as deep as eye, prominent in dorsal aspect; width across postorbital tubercle greater than width across eyes. Pronotum short, length/greatest width 1.16; its greatest width greater than width of elytra across humeri; widest point posterior to middle; apex of pronotum very strongly narrowed; base moderately narrowed; lateral margin scarcely sinuate anterior to hind angle; paramedian grooves broad, punctate, shallower than in other members of subgenus; paramedian groove equal in width to inner carina at middle; distinctly narrower than outer one at middle; both carinae coarsely, densely punctate; marginal groove fine, about 0.20 as wide as outer carina at middle. Elytra relatively short, broad; strial punctures smaller than in related species, each puncture less than half width of one interval; one seta in apex of Stria IV, one in apical stride; several setae near tip of Stria VII. Male with proximal tooth of front tibia small, more proximad than in related species; ventral tooth of anterior femur of male small, at apical third; calcar of middle tibia appearing truncate, 0.5 as long as spur (but calcar in holotype may be broken or worn), hind tibia only slightly swollen proximad to middle; hind calcar very large, its tip slightly obtuse. Female with short, deep lateral groove in metasternum; lateral margin of latter not flared; lateral pits of Sternum IV rather small, separated by more than 5 times width of one of them; pit with very strong anteriolateral brace, without longitudinal striation. This species has quite different body proportions from other members of the subgenus, with a short, wide pronotum, a very small head, and short elytra. The paramedian grooves are shallower and more coarsely punctate than in related species, and the postorbital tubercles are larger than in other Pyxiglymmius, although they are small in comparison to those of Omoglymmius (Laminoglymmius) gorgo. Omoglymmius ( Pyxiglymmius ) pilosus (Grouvelle) NEW COMBINATION Figs. 42, 45, 47-50 Rhysodes pilosus Grouvelle 1903: 123-124. Pyxiglymmius pilosus (Grouvelle) Bell and Bell 1978 Type material. - LECTOTYPE (here designated) male, labelled: “Bengkalis-Maindron 1885” (SUMATRA) (MNHN); PARALECTOTYPES one female, same data as lectotype. These specimens are labelled as “Types” in the Grouvelle Collection (MNHN), but are not mentioned in the original description. Additional PARALECTOTYPES, SUMATRA; two males, four females, Palembang (MNHN); one male, Palembang, Fry Colin. 1905-100, labelled “co-type” (BMNH); one female, Palembang, 1901-267 (BMNH); BORNEO: two males, Occ. riv. Sambey, pres Ngabang, coll. J. B. Ledru, 1897, labelled “co-type” (BMN). Since these localities are mentioned in the original description, they must also represent paralectotypes. Grouvelle also mentions specimens from the rivers Pontianac and Mander and Guenong-Ampar in Borneo Occ. We have not located these specimens, but they may be among recently discovered material in the Oberthur Collection in Paris. Description. — Length 7.8-10.0 mm. Antennal Segment XI slightly longer than wide, its apex obtuse; basal setae numerous on Segments V-X; head slightly longer than wide; clypeus coarsely punctate; median lobe continuous with clypeus; median lobe rhomboidal, wider than long, its lateral and posterior angles acute; anteriomedial margin of temporal lobe oblique; first medial angles broadly separated; emargination between first and second medial angles very deep; second medial angles narrowly separated; nearly oblique between second medial angle and occipital angle; each temporal lobe with 4-7 very coarse punctures in lateral half, some of these with very short setae; postorbit flat dorsad to tubercle; latter prominent, 0.33 as deep and 0.67 as long as eye; width across postorbital tubercle slightly greater than width across temporal lobes. Pronotum rather short, length/greatest width 1.13, widest near middle; base moderately and apex strongly narrowed; lateral margin at most slightly sinuate anterior to hind angle; median and paramedian grooves broad, rather shallow, prominently punctate; inner carinae slightly narrower than outer carinae at middle; both pairs of carinae very coarsely, rather densely punctate; marginal groove shallow, rather broad, punctate. Elytra moderately elongate; apical tubercle very small; strial punctures very large, some of them slightly irregular in arrangement; all striae with numerous setae; female with deep, broad lateral groove on metasternum, lateral margin of Quaest. Ent., 1982, 18 (1-4) 156 Bell and Bell groove conspicuously flared, groove continuous with lateral pit on abdominal Sternum 1; female with very large lateral pits on Sternum IV, separated medially by less than width of one of them (Fig. 50) pit of Sternum IV with anteriolateral brace and longitudinal striae; medioposterior border of pit straight, oblique; ventral margin of female elytron angulate opposite abdominal Sternum I; male with shallow lateral pit on Sternum IV and faint suggestion of lateral furrow on metasternum. Male with proximal tooth on anterior tibia located slightly dorsad of middle of tibia; male with ventral tooth on anterior femur located distad to middle (Fig. 47); calcar of middle tibia about 0.67 as long as spur (Fig. 48); calcar of hind tibia very large, obtusely pointed (Fig. 49). Presence of setae on all elytral striae will separate this species from all other members of the subgenus. In this character it agrees with the sympatric Omoglymmius ( Laminoglymmius ) rugosus, which differs in having a setose pronotum and differently shaped temporal lobes. Range. - Sumatra and Borneo. In addition to type material, we have seen the following specimens (all from Sumatra): one male, Bedagei, int. Sumatra s O.K., 600 ft., 2 Sem. 89, coll. 1. Z. Kannegieter (MNHN); one male. Deli, coll. Dr. Busey (AMS); one male, Manna, coll. M. Knappert (LEI); two males, one female, Roagu, no other data (MNHN); two females, Tandjong, Morawa, Serdang, N. O. Sumatra, coll. Dr. B. Hagen (LEI); two females, Tebing-Tinggi, coll. Dr. Schultheiss (MNHN). Omoglymmius (Pyxiglymmius) krikkeni new species Figs. 43,46,51-56 Type material. - HOLOTYPE male, labelled: “N. SUMATRA, Alas Valley, Gumpang: Foothills Mt. Kemiri, 3° 47' N, 97° 27' E, 10-VI-1972, ca. 780 m., coll. J. Krikken, no. 10, from rotten trunk, lowland multistratal evergreen forest” (LEI). PARATYPES one male, eight females, same data as type (LEI). Description. — Length 8.3-10.3 mm. Antennal Segment XI slightly wider than long, its apex obtuse; basal setae numerous on Segments V-X; head slightly longer than wide; clypeus finely punctate; median lobe continuous with clypeus; median lobe rhomboidal, wider than long; anteriomedial margin of temporal lobe oblique, first medial angles broadly separated; emargination between first and second medial angles shallower than in O. pilosus\ second medial angles narrowly separated; margin distinctly rounded between second medial angle and occipital angle; each temporal lobe with three to seven coarse punctures; postorbit flat dorsad to tubercle, latter prominent 0.33 as deep and 0.67 as long as eye; width across postorbital tubercles slightly less than width across eyes. Pronotum rather short, length/greatest width 1.21; widest near middle, base strongly narrowed, apex moderately narrowed; lateral margin slightly sinuate anterior to hind angles; median and paramedian grooves broad, rather shallow, punctate; inner carina slightly narrower than outer carina at middle; both pairs of carinae with few, irregularly distributed punctures, nearly absent from anterior third; marginal groove fine, shallow. Elytra moderately elongate; Striae 111, IV slightly sinuate anterior to very small apical tubercle; strial punctures very coarse; elytron with row of setae in Interval III slightly laterad to Stria 11, and another row in Interval V slightly laterad to Stria IV; two or three setae in Interval VI slightly laterad to Stria V; row of setae in Stria VII; distinct depression in Intervals IV, V just anterior to apical tubercle (Fig. 52); female with broad, deep lateral groove on metasternum, lateral margin of groove conspicuously flared (Fig. 53); female with deep lateral pit on abdominal Sternum 1; female with very large lateral pits on Sternum IV, separated medially by slightly more than the width of one of them; pit of Sternum IV with anteriolateral brace and longitudinal striae; medioposterior margin of pit curved; ventral margin of elytron of female expanded, angulate, opposite hind coxa, and slightly angulate opposite lateral pit of Sternum IV (Fig. 51); male with small, shallow lateral groove on metasternum and slight, irregular lateral pits on Sterna III, IV, V. Male with proximal tooth of anterior tibia slightly dorsad to middle; male with ventral tooth of anterior femur rounded, nearer to apex than to middle (Fig. 54); calcar of middle tibia acute, longer than spur (Fig. 55); calcar of hind tibia very large, its dorsal margin straight, its tip acute (Fig. 56). This very large species is easily recognized by the pattern of setae on the elytra. We dedicate this species to its discoverer, Dr. J. Krikken, in appreciation of his help in our studies. Subgenus Laminoglymmius new subgenus Type species. - Rhysodes insularis Grouvelle 1903 Description. — Segment XI of antenna obtuse, without stylet; basal setae on distal antennal segments; clypeal setae absent; medial margin of temporal lobe with pale translucent area (conspicuous in all species except O. actae, in which it is Revision of Omoglymmius Ganglbauer 157 obsolete); some species with similar translucent area at tip of median lobe; medial margin of temporal lobe various from entire ( O . inaequalis ) to bilobed or trilobed, with one, two, or three medial angles; most species with postorbital tubercle conspicuous, on some species located nearly suborbitally; punctures of abdominal sterna scattered, numerous; female with small, simple lateral pits in Sternum IV, male with shallow lateral pits in Sternum IV; cleaning organ located entirely distad to base of anterior tarsus; spur of middle tibia curved; males, where known, with hind calcar small, at or near base of spur. Most members of this subgenus are recognized by the presence of a translucent medial margin, with one or two shallow emarginations on the temporal lobe separating two or three medial angles, which are close together. Two species are atypical: in O. actae , the translucent area is scarcely developed, though there is a shallow but distinct emargination. In O. inaequalis the medial margin is not emarginate, though the translucent area is well marked. Those species with the medial margin of the temporal lobe distinctly emarginate might be confused with members of the subgenus Pyxiglymmius. In the latter subgenus, the two medial angles are widely separated from one another, and the emargination between them is deep in all species except for O. lederi. O. inaequalis , which lacks the medial emargination, might be mistaken for Subgenus Indoglymmius. The latter has a well-marked translucent area on the median margin, but differs in lacking basal setae from the antennae. The subgenus is largely confined to a small part of the Oriental Region, where it is found in Sumatra, Borneo, the Malay Peninsula, and the Nicobar Islands. O. actae is found in New Guinea, far from the other species. It is aberrant in the poor development of the translucent areas, and is perhaps not really related to the other species. Laminoglymmius is especially well represented in Sumatra, where there are four sympatric species. Phytogeny. - A possible phylogeny for the subgenus is presented in Diagram 2. We postulate that: Species 1, the ancestor of the subgenus, had two medial angles on each temporal lobe, both included in a distinct translucent area; clypeus was continuous with the median lobe, punctures of pronotum coarse, moderately dense and relatively evenly distributed; and elytral setae were absent except for the tip of Stria VII and perhaps the apical striole. We suggest that O. rugosus is a highly modified descendant of Species 1, having developed the following features: a transverse groove between the clypeus and the median lobe; numerous setae on the head, pronotum, and elytra; a third opaque medial angle posterior to the second one and very dense, confluent pronotal punctures. All other species are probably descended from Species 2. This was similar to Species 1 except that the pronotal punctures became much sparser than is typical for most species of Omoglymmius. Species 2 was probably very similar to O. oberthueri, which may be regarded as an almost unmodified descendent. O. actae is a similar species, marked by the virtual disappearance of the translucent areas of the temporal lobe. These two species show no obvious evidence of close relationship with each other, or with the four remaining species. The latter appear to be descended from two daughter species of Species 2 (Species 3 and 4). Species 3 and 4 might have a common ancestor more recent than Species 2. The only evidence for this is the presence in all four daughter species of a translucent area at the tip of the median lobe. In Species 3, the reduction of punctation of the pronotum was extreme, O. insularis lacking punctures entirely and O. inaequalis having only a single puncture on the outer carina. In Species 4, the punctuation was less reduced, approximately six punctures being retained on each outer carina, but the pronotum became rounded in outline, the median lobe became narrow and elongate, with nearly parallel sides, and the postorbital tubercles were enlarged to the point of being visible in dorsal view. Species 3 gave rise to O. insularis and O. inaequalis. In O. insularis , the median lobe became concave, all punctures were lost from the outer carina, and the temporal lobe lost all its Quaest. Ent., 1982, 18 (1-4) 158 Bell and Bell Phylogenetic Diagram 2. Reconstructed Phylogeny of species of Laminoglymmius. punctures. O. inaequalis retained a single puncture at the middle of each outer carina, while the outer carina became very narrow, the postorbital tubercle was displaced ventrally to lie opposite the lower margin of the eye, and the anteriomedial margin of the temporal lobe became oblique, so there is only one medial angle. Species 4 gave rise to O. trisinuatus and O. gorgo. In O. trisinuatus, a third medial angle was developed, while in O. gorgo , the postorbital tubercles became greatly enlarged. KEY TO SPECIES 1 Median lobe separated from clypeus by transverse pollinose groove; temporal Revision of Omoglymmius Ganglbauer 159 Plate 5. Figures 57-74, subgenus Laminoglymmius. Figs. 57-63, Head and pronotum, dorsal aspect; Fig. 57, Omoglymmius ( Laminoglymmius ) rugosus (Grouvelle); Fig. 58, O. (L.) inaequalis new species; Fig. 59, 0. (L.) actae new species; Fig. 60, O. (L.) insularis (Grouvelle); Fig. 61, O. (L.) oberthueri (Grouvelle); Fig. 62, O. (L.) trisinuatus new species; Fig. 63, O. (L.) gorgo new species ; Figs. 64-66, O. (L.) rugosus (Grouvelle), male; Fig. 64, Femur, tibia, anterior leg; Fig. 65, Middle tibia; Fig. 66, Hind tibia; Fig. 67, Hind tibia, male, O. (L.) actae new species; Figs. 68-74, Head, lateral aspect; Fig. 68, O. (L.) rugosus (Grouvelle); Fig. 69, O. (L.) inaequalis new species; Fig. 70, O. (L.) actae new species; Fig. 71, O. (L.) insularis (Grouvelle); Fig. 72, O. (L.) oberthueri (Grouvelle); Fig. 73, O. (L.) trisinuatus new species; Fig. 74, O. (L.) gorgo new species. Quaest. Ent., 1982, 18 (1-4) 160 Bell and Bell lobes, genae, pronotum, and elytra with numerous setae O. rugosus (Grouvelle), p. 160 r Median lobe continuous with clypeus; head, pronotum without setae; setae of elytron limited to a few in apex of Stria VII and none or one on apical tubercle ... 2 2 (1) Outer carina distinctly narrower than inner carina 3 2' Outer carina not narrower than inner carina 4 3 (2) One medial angle on temproal lobe; translucent area present; outer carina 0.25 width of inner carina; outer carina with one puncture near middle, inner carina impunctate O. inaequalis new species, p. 161 3' Two medial angles on temporal lobe; latter without translucent area; outer carina 0.5 width of inner carina; outer carina with five punctures, inner carinae with one or two O. actae new species, p. 161 4 (2') Median lobe flat; outer carina punctuate 5 4' Median lobe concave; pronotum entirely impunctate O. insularis (Grouvelle), p. 162 5 (4) Inner carina of pronotum with three or four coarse punctures; median lobe of head rhomboidal O. oberthueri (Grouvelle), p. 162 5' Inner carina impunctate; median lobe of head narrow, its margins parallel or nearly so 6 6 (5') Temporal lobe with three medial angles; postorbital tubercle relatively small, its length about 0.5 of diameter of eye O. trisinuatus new species, p. 163 6' Temporal lobe with two medial angles; postorbital tubercle very large, its length equal to diameter of eye O. gorgo new species, p. 163 Omoglymmius (Laminoglymmius) rugosus (Grouvelle 1903) NEW COMBINATION Figs. 57, 64-66, 68 Rhysodes rugosus Grouvelle 1903: 122-123. Pyxiglymmius rugosus (Grouvelle) Bell and Bell 1978 Type material. - HOLOTYPE sex not recorded, labelled: “SINGAPORE, Raffray” (MNHN). Description. — Length 6. 9-8. 5 mm. Segment XI of antenna wider than long, rounded at apex; basal setae present on Segments V-XI; head relatively short; clypeus punctate, microsculptured, separated from median lobe by deep transverse pollinose groove; antennal rim very thick, elevated; frontal space minute; temporal lobe with three medial angles, first and second ones translucent, each narrowly separated from corresponding angle of opposite temporal lobe; third medial angles opaque, more broadly separated from each other; temporal lobe very convex, shining, with about 15 coarse punctures near lateral margin, most of punctures with stout but very short setae; temporal lobe rounded posteriorly; postorbital tubercle moderate in size, with deep concavity dorsad to it; postorbital tubercle densely coarsely punctate, irregular network of ridges between punctures; some punctures with short setae. Pronotum moderately long; length/greatest width 1.15; widest point distinctly anterior to middle, apex narrowed, lateral margins oblique, nearly straight posteriorly, in form of long but shallow sinuation anterior to rectangular hind angle, pronotal grooves broad, rather shallow, ill-defined, finely pollinose, coarsely punctate; carinae very coarsely, irregularly rugose, punctate; each puncture with seta; inner and outer carinae nearly equal at middle; marginal groove ill-defined medially, but with its lateral margin raised, in form of fine carina. Elytral striae very broad, intervals narrow, about 0.5 as wide as striae; intervals convex; strial punctures very irregular, in places in form of two parallel rows within stria; all striae with many setae; apical tubercle very small; female with distinct lateral pits on Sternum IV. Male with sharp ventral tooth at apical third of anterior femur (Fig. 64); female with sinuation in this posititon; proximal tooth at middle of anterior tibia in male; middle tibia of male with a small, acute calcar (Fig. 65); hind calcar small, obtuse, distinctly dorsad to level of spur (Fig. 66). Revision of Omoglymmius Ganglbauer 161 The numerous setae of the pronotal carinae, genae, and temporal lobe separate this large species from all others. Large size, completely setose striae and partly irregular strial punctation give a superficial similarity to the sympatric Omoglymmius (Pyxiglymmius) pilosus, but the form of the temporal lobes and the absence of pronotal setae eliminate the latter species. Range. - Singapore, Sumatra and Borneo. In addition to the type, we have studied the following specimens: BORNEO: one male, one female, Mt. Matang, West Sarawak, coll. G. E. Bryant, 31-1-14 (BMNH); SUMATRA: one male, one female, Palembang (MNHN). Omoglymmius (Laminoglymmius) inaequalis new species Figs. 58, 69 Type material. - HOLOTYPE female, labelled: “Nicobars” (MNHN). This specimen was identified by Grouvelle as Rhysodes nicobarensis Grouvelle (1895b). The latter species, as shown by the holotype, also in Paris, belongs to Subgenus Orthoglymmius , and, despite the name, was collected from the Andaman Islands. Our placement of R. nicobarensis in Subgenus Hemiglymmius (Bell and Bell 1978) was based on a study of O. inaequalis , and not the true R. nicobarensis. Description. — Length 6.5 mm. Antennal Segment XI as wide as long, its apex obtusely rounded; basal setae sparse on Segment V, numerous on Segments VI-X; clypeus impunctate, continuous with median lobe; latter also impunctate, broad, its tip pointed, translucent; anteriomedial margin of temporal lobe almost straight, small but distinct semicircular translucent area present anterior to medial angle; only one medial angle present; margin oblique between medial and occipital angles; temporal lobe convex, shining, with 10-12 coarse punctures present in lateral half; postorbital tubercle large, 0.5 as deep and 0.66 as long as eye, its dorsal margin only slightly above ventral margin of eye (so it could better be termed a “suborbital tubercle”); width across potorbital tubercles slightly less than width across temporal lobes. Pronotum relatively short, length/greatest width about 1.1; widest point slightly anterior to middle, sides strongly curved, convergent to apex, which is narrow; base relatively broad, lateral margin not sinuate anterior to hind angle, which is slightly obtuse; paramedian groove deep, broad, more than 0.33 as wide as inner carina at middle; outer carina very narrow, less than 0.25 as wide as inner carina at middle; outer carina with one minute puncture near middle, this puncture with minute seta; inner carina entirely impunctate; marginal groove dilated, over 0.5 width of outer carina at middle. Elytra relatively short, broad; striae very shallow; intervals flat, finely microsculptured, with faint sheen; strial punctures very small, each about 0.20 as wide as an interval; elytron with several setae in apex of Stria VII; female with large, deep lateral pit on Sternum IV; female with small ventral tooth on anterior femur; male unknown. This is the only member of the subgenus in which there is only one medial angle. The very narrow outer carina with a single minute setiferous puncture is also diagnostic. The combination of a single medial angle with a well-developed translucent area is suggestive of Subgenus Indoglymmius. Members of the latter subgenus differ in lacking basal setae on the antennae, and in having the inner and outer carinae of equal width. Omoglymmius (Laminoglymmius) actae new species Figs. 59, 67, 70 Type material. - HOLOTYPE male, labelled: “NEW GUINEA, Maffin Bay, IX- 1944, E. S. Ross” (CAS). Description. — Length 5.5 mm. Antennal Segment XI slightly wider than long, its apex rounded; basal setae numerous on Segments V-X; clypeus continuous with median lobe; latter rhomboidal, with a few coarse punctures; head distinctly longer than wide; anteriomedial margin of temporal lobe oblique; latter with two minute medial angles which are located close together; without obvious translucent area near medial angles (under strong light, translucent area is faintly suggested); temporal lobe convex, shining, with 12-15 coarse scattered punctures; postorbital tubercle small, 0.25 as deep and 0.33 as long as eye, located slightly above ventral margin of eye; width across postorbital tubercles less than width across eyes. Quaest. Ent., 1982, 18 (1-4) 162 Bell and Bell Pronotum moderately elongate, length/ greatest width 1.20; widest near middle, lateral margins curved; apex strongly narrowed; base moderately narrowed; lateral margin not sinuate anterior to hind angle; latter obtuse; paramedian grooves deep, rather narrow; outer carina distinctly narrower than inner one, about 0.33 as wide as inner one at middle; outer carina with four or five scattered coarse punctures, without setae; inner carina with one or two coarse punctures; marginal groove dilated, about 0.33 as wide as outer carina at middle. Elytra narrow, convex; elytral striae deep, coarsely punctate, each puncture about 0.6 as wide as in interval; intervals narrow, convex, shining; several setae present in apex of Stria VII; male with distinct proximal tooth on anterior tibia and ventral tooth at apical fourth of anterior femur; middle calcar small, triangular; hind calcar obtuse, triangular, located 0.33 of length of tibia dorsad to spur (Fig. 67); female unknown. This species is the only member of the subgenus which does not have well marked translucent areas near the medial angles of the temporal lobes. It is recognized as a member of Subgenus Laminoglymmius by the double medial angles. This character and the presence of basal setae on the antenna will separate it from superficially similar species of Omoglymmius s. str. from New Guinea. Omoglymmius (Laminoglymmius) insularis (Grouvelle) NEW COMBINATION Figs. 60,71 Rhysodes insularis Grouvelle 1903: 124-125. Pyxiglymmius insularis (Grouvelle) Bell and Bell 1978 Type material. - HOLOTYPE female, labelled: “Bengkalis, Maindron 1885” (MNHN). The type locality is a small coastal island near Sumatra. Description. — Length 7.0 mm. Segment XI of antenna as wide as long, its apex rounded; basal setae sparse on Segments V-VI, dense on Segments VII-X; head slightly longer than wide; clypeus continuous with median lobe; clypeus impunctate; median lobe elongate oval, deeply concave at center, its margins^ raised; tip of median lobe rounded, translucent; temporal lobe with two medial angles, separated by very shallow emargination; first medial angles separated from one another by approximately the width of one of them; second medial angles separated by about half the distance; both angles and adjacent margins in a translucent area; temporal lobe evenly rounded posteriorly, laterally; temporal lobe entirely smooth, impunctate scarcely overhanging postorbit, latter smooth, glabrous; postorbital tubercle small but well-defined; gular tubercle slightly prominent. Pronotum moderately elongate, length/greatest width 1.25; widest anterior to middle, lateral margins curved, apex strongly narrowed; base moderately narrowed, lateral margin distinctly sinuate anterior to hind angle, which is nearly rectangular; paramedian grooves deep, rather narrow, outer carina slightly narrower than inner one at middle; both carinae entirely impunctate, without setae; marginal groove relatively narrow. Eltyra moderately elongate, interval V slightly elevated anteriorly; striae relatively narrow, each puncture about 0.25 as wide as interval; several setae present in apex of Stria VII; female with deep, round, small lateral pit on abdominal Sternum IV and with small ventral tooth on anterior femur; male unknown, The deeply concave, oval median lobe separates this species from all other Rhysodini. Omoglymmius ( Laminoglymmius ) oberthueri (Grouvelle) NEW COMBINATION Figs. 61,72 Rhysodes oberthueri Grouvelle 1903: 1 18-119. Omoglymmius ( Hemiglymmius ) oberthueri (Grouvelle) Bell and Bell 1978 Type material. - HOLOTYPE female, labelled: “Poulo-Pinang, Raffray” (MNHN) Description. — Length 7.0 mm. Antennal Segment XI as wide as long, its apex rounded; basal setae numerous on Segments V-X; head slightly longer than wide; clypeus continuous with median lobe; latter rhomboidal, its tip pointed; clypeus and median lobe impunctate; anteriomedial margin of temporal lobe oblique; two medial angles, separated by shallow but rather long emargination; each medial angle narrowly separated from that on opposite temporal lobe; medial angles and emargination within well-marked translucent area; temporal lobe with nine or 10 coarse punctures in posteriolateral third; posteriolateral margin of temporal lobe evenly rounded; temporal lobe slightly overhanging postorbit; latter glabrous, impunctate; postorbital tubercle rather large, about 0.5 as long and 0.5 as deep as eye; its dorsal margin on level with middle of eye. Revision of Omoglymmius Ganglbauer 163 Pronotum relatively short, broad, length/greatest width 1.13; widest slightly anterior to middle; sides curved, apex strongly narrowed, base moderately narrowed; sides curved, sides oblique just anterior to obtuse hind angle, paramedian grooves deep, rather narrow; outer carina scarcely narrower than inner one; outer carina with seven to 1 2 coarse, scattered punctures; inner carina with three or four coarse punctures near middle of its length, otherwise impunctate. Elytra relatively short, broad; striae deep; Intervals III, V slightly broader, more elevated than others anteriorly; strial punctures coarse, each more than half as wide as interval; Stria IV with one seta in apex; apical stride with one seta, Stria VII with several setae near apex; female with very deep, large lateral pits on Sternum IV; each pit as long as Sternum IV; pits separated medially by less than twice the width of one of them; female with distinct ventral tooth on anterior femur; male unknown. The rhomboid median lobe and the presence of two medial angles make this species most like O.actae. It differs from the latter species in having a well-marked medial translucent area on each temporal lobe, as well as in having the medial angles of each lobe more separated from one another. Omoglymmius (Laminoglymmius) trisinuatus new species Figs. 62, 73 Type material. - HOLOTYPE male, labelled: “Bangung” (MNHN). This is possibly Bangoen (modern spelling “Bangun”). east of Pematangsiantar, in northern Sumatra. Description. — Length 6.9 mm. Antennal Segment XI wider than long, its apex bluntly rounded; basal setae numerous on Segments V-X; head longer than wide; clypeus continuous with median lobe; clypeus, median lobe impunctate; median lobe elongate, narrow, its tip translucent; anteriomedial margin of temporal lobe curved; temporal lobe with three very small medial angles, separated by very shallow sinuations; first pair well separated; second pair very narrowly separated; third pair almost contiguous; all angles and adjacent margin translucent; posteriomedial margin slightly oblique; temporal lobe convex, shining, with three coarse punctures near lateral margin; postorbit pollinose; postorbital tubercle 0.3 as deep and 0.3 as long as eye, its dorsal margin on level with lower third of eye; postorbital tubercles visible in dorsal view, width across them slightly less than width across eyes. Pronotum short, broad, length/greatest width 1.05; widest slightly anterior to middle, apex very strongly narrowed; base slightly narrowed; lateral margin scarcely sinuate anterior to obtuse hind angles; paramedian grooves deep, about 0.5 as wide as inner carina at middle; outer carina only slightly narrower than inner one at middle; inner carina impunctate; outer carina with five to seven punctures near its lateral margin; marginal groove narrow. Elytra relatively short, broad; strial punctures small, well separated from one another, less than 0.3 as wide as an interval; Interval V slightly elevated anteriorly; elytron with setae near apex of Stria VII, otherwise without setae; male with faint lateral pit in Sternum IV; female unknown. Male with small ventral tooth on anterior femur; middle calcar very small, obtuse; hind tibia with small obtuse calcar at level of spur. Three medial angles of the temporal lobe separate this species from all others except for O. rugosus, from which it differs in lacking setae on the temporal lobe and pronotum, as well as many other characters. Shape and punctation of the pronotum, narrow median lobe, and secondary sexual characters of the male indicate that its true relationships are with O. gorgo, which differs in having the postorbital lobes greatly enlarged. Omoglymmius ( Laminoglymmius ) gorgo new species Figs. 63, 74 Type material. - HOLOTYPE male, labelled: “SUMATRA: Siantar, NGSSI exp, 1937, Mann” (NMNH). PARATYPES one male, one female, labelled “Bedagei, int., Sumatra’s O.K. 600', 2de de Sem, '89, coll. I. Z. Kannegieter” (MNHN). Description. — Length 6. 0-8.0 mm. Antennal Segment XI as wide as long, its apex rounded; basal setae numerous on Segments V-X; head longer than wide (if postorbital tubercles are not considered); clypeus continuous with median lobe; latter very narrow, its tip narrowed, rounded, opaque; clypeus finely punctate; median lobe impunctate; anteriomedial margin of temporal lobe curved; temporal lobe with two small medial angles, separated by shallow emargination; first medial angles slightly separated; second medial angles contiguous; first medial angles translucent; second medial angles less distinctly so; temporal lobe convex, impunctate; postorbit pollinose; postorbital tubercle very large, longer than eye, 0.6 Quaest. Ent., 1982, 18 (1-4) 164 Bell and Bell as deep as eye, its upper and lower margins parallel, its apex subtruncate; width across postorbital tubercles much greater than that across eyes. Pronotum short, relatively broad, length/greatest width 1.04; widest near middle, apex very markedly narrowed; base moderately narrowed, sides markedly curved; lateral margin distinctly sinuate anterior to obtuse hind angle; paramedian grooves deep, about 0.5 as wide as inner carina at middle; outer carina only slightly narrower than inner one at middle; inner carina impunctate; outer carina with five to seven punctures near its lateral margin; marginal groove narrow. Elytra relatively short, broad; strial punctures small, well separated from one another, less than 0.3 as wide as one interval; Interval V slightly elevated anteriorly; several setae near apex of Stria VII, elytron otherwise without setae; female with small but distinct lateral pit on abdominal Sternum IV; male without lateral pit; male with small ventral tooth on anterior femur; female with very small tooth in this location; middle calcar scarcely evident; hind calcar very small but distinct (smaller than in O. trisinuatus). The grotesque enlargement of the postorbital tubercles easily separates this species from other Laminoglymmius . Subgenus Navitia Bell and Bell 1978 Type species. - Rhysodes intrusus Grouvelle 1903 Description. — Basal setae of antennae entirely absent; antennal stylet present, conical, acute; clypeal setae absent; eye large, normal, cornea facetted; frontal grooves very shallow; frontal space wide, crescentic; medial angles of temporal lobes simple; translucent areas absent from temporal lobes; one temporal seta on each lobe; postorbital tubercle absent; gular grooves each containing an enlarged pit. Pronotal carinae impunctate except for a few punctures at base of outer carina; inner and outer carinae equal in width; marginal groove of pronotum abbreviated posteriorly, replaced there by row of punctures; apex of elytral Stria VII not impressed; most posterior puncture of Stria III enlarged; Stria II with one seta at apex in most specimens (absent in some specimens); Stria IV with three or four setae, distributed along its entire length; subapical stride with 2 setae; Stria VII with three to five setae near apex; lateral pits present on abdominal Sternum IV in both sexes; spur of middle tibia curved anteriorly; punctures of abdominal Sterna III-V each aligned in single transverse row. This small and distinctive subgenus combines the characters of several other subgenera. An antennal stylet and the simple medial angles of the temporal lobes suggest Hemiglymmius. The latter subgenus differs in having well-developed basal setae on the antennae and deep frontal grooves. Alignment of abdominal punctures in a single row on each sternum is suggestive of Orthoglymmius , but in the latter subgenus, the middle tibial spur is straight, and clypeal setae are present. The habitus is similar to Omoglymmius s. str ., but the latter subgenus lacks the antennal stylet and has deeper frontal grooves. The frontal grooves are shallower than in any other members of Subtribe Omoglymmiina. In O. intrusus , they are glabrous and scarcely visible, giving a superficial resemblance to Dhysores and related genera. In Navitia , however, the frontal space is crescentic, while in Subtribe Dhysorina it is small and round. In O. stylatus, the shallowness of the frontal grooves is less conspicuous because they are pollinose. Navitia is known from Fiji and the New Hebrides. KEY TO SPECIES 1 Marginal groove confined to anterior 0.85 of pronotum, followed posteriorly by three to four very crowded punctures; pronotal epipleura with line of punctures ventrad to marginal groove; frontal grooves pollinose O. stylatus new species, p. 165 V Marginal groove confined to anterior 0.50-0.66 of pronotum, followed posteriorly by five widely separated punctures; pronotal epipleura without line of punctures; frontal grooves glabrous, scarcely evident O. intrusus (Grouvelle), p. 166 Revision of Omoglymmius Ganglbauer 165 Plate 6. Figures 75, 76, 79, 80, Subgenus Navitia-, Figures 77, 83 Subgenus Indoglymmius-, Figures 78, 81,82, Subgenus Nitiglymmius; Figs. 75-78, Head and pronotum, dorsal aspect; Fig. 75, Omoglymmius (Navitia) stylatus new species; Fig. 76, O. (N.) intrusus (Grouvelle); Fig. 77, Omoglymmius (Indoglymmius) lineatus (Grouvelle); Fig. 78, O. ( Nitiglymmius ) semioculatus new species; Figs. 79-80, Prothorax, lateral aspect; Fig. 79, O. ( Navitia ) intrusus (Grouvelle); Fig. 80, O. (TV.) stylatus new species; Fig. 81, Hind tibia, male, O. ( Nitiglymmius ) semioculatus ; new species; Figs. 82-83, Head, lateral aspect; Fig. 82, O. ( Nitiglymmius ) semioculatus new species; Fig. 83, O. (Indoglymmius) lineatus (Grouvelle); Omoglymmius ( Navitia ) stylatus new species Figs. 75, 80 Type material. - HOLOTYPE male, labelled: “NEW HEBRIBES.Malekula, XII-1929, coll. L. E. Cheesman, BM 1930-393 (1669)” (BMNH). PARATYPES two females, N. E. Malekula, 1-1930, coll. L. E. Cheesman, BM-1930-178 (BMNH); one female, Malekula: Ounua, IV-V-1929, coll. L. E. Cheesman, BM 1929-371 (531) (BMNH); one female, Aoba:Banka Harijitoa, above Dundy, elev. 1500-2000 ft., IX-8-1958, collector not specified (CNHM). Quaest. Ent., 1982, 18 (1-4) 166 Bell and Bell Description. — Length 5. 0-6. 2 mm. Head cordate, slightly longer than broad; temporal lobes distinctly flattened; frontal grooves slightly impressed, with fine pollinosity; temporal lobe with few coarse punctures in posteriolateral third, near temporal setae; orbital groove relatively well-developed, extending posteriorly to or beyond middle of temporal lobe. Pronotum markedly varied in proportions, widest anterior to middle; lateral margin with distinct sinuation anterior to hind angles in some specimens, in others sinuation slight or absent; marginal groove of pronotum entire except for basal 0.16, replaced there by very crowded row of three or four punctures; pronotal epipleuron with row of punctures ventrad to marginal groove (Fig. 80). Elytron with most posterior puncture of Stria III moderately enlarged; punctures of abdominal Sternum V not confluent laterally; male with ventral tooth on anterior femur; calcar of middle tibia obsolete; hind calcar small, triangular, its tip slightly obtuse, distinctly proximad to base of spur. This species is easily distinguished from O. intrusus by the more complete marginal groove of the pronotum, line of punctures on the pronotal epipleura, and deeper, pollinose frontal grooves. Grouvelle (1903) mentioned a specimen tentatively identified as R. intrusus from the New Hebrides. We have not located this specimen, but we suspect that it really represents O. stylatus. Omoglymmius (Navitia) intrusus (Grouvelle) Figs. 76, 79 Rhysodes intrusus Grouvelle 1903:1 12-113. Omoglymmius ( Navitia ) intrusus (Grouvelle) Bell and Bell 1978 Type material. - LECTOTYPE (here designated) female, labelled: FIJI “Viti,” no collector or date (MNHN). PARALECTOTYPE: Grouvelle (1903) mentioned a second specimen in the Oberthiir collection. It may still exist among Oberthiir material recently rediscovered in Paris. Description. — Length 5. 0-6.0 mm. Head broader, more convex than in O. stylatus, temporal lobes not flattened; frontal grooves scarcely impressed, shining, glabrous; temporal lobe entirely without coarse punctures, but with sparse minute ones; orbital grooves obsolete. Pronotum widest near middle; sinuation anterior to hind angle shallow but distinct; marginal groove distinct in anterior 0.50 to 0.67, replaced posteriorly by row of five widely spaced punctures; pronotal epipleura without row of punctures (Fig. 79). Elytron with most posterior puncture of Stria III greatly enlarged, so that Stria IV also seems to lead to it; punctures of Sternum V confluent near lateral margin; male unknown. The frontal grooves are virtually absent in this species. Glabrous frontal grooves, the more abbreviated marginal groove on the pronotum, absence of punctures from the pronotal epipleura, and confluent punctures of Sternum V are the most conspicuous differences from O. stylatus. Shape of the pronotum shows an unusual amount of variation in both species. O. intrusus is known only from the Fijian islands of Viti Levu and Ova Lau. In addition to the lectotype, we have examined the following specimens; one female, labelled “Viti Levu, Naivithuia, Taileru-Fiji, V 1 1 1- 1 6- 1 937, coll. J. M. Valentine” (BPM); one female, labelled “Ova Lau, Fiji, Draiba Trail, V 1 1-38, rotten log, coll. E. C. Zimmerman” (BPBM). Subgenus Caeconavitia new subgenus Type species. - Omoglymmius (Nitiglymmius) zimmermani Bell and Bell 1978:77. Description. — Basal setae of antenna entirely absent; antennal stylet present, conical, obtuse; clypeal setae absent; eye small, only slightly visible in dorsal aspect; depth of eye about 0.33 in depth of head; facetting of cornea much reduced, barely visible at high magnification; frontal grooves deep, narrow; medial angles of temporal lobes simple; translucent areas absent from temporal lobes; temporal setae absent; postorbital tubercle absent; each gular groove with enlarged pit. Marginal groove of pronotum entire; elytral striae distinctly impressed; abdominal Sterna III-V each with transverse row of punctures; Sternum IV with deep and Sternum V with shallow lateral pit in both sexes. This subgenus contains one species, from Fiji. We originally assigned it to Subgenus Nitiglymmius. Futher study of the genus has led us to attribute greater importance to presence Revision of Omoglymmius Ganglbauer 167 or absence of the antennal stylet, and has convinced us that O. zimmermani is more closely related to Navitia than to the remaining species of Nitiglymmius. Characters shared between Navitia and Caeconavitia include an antennal stylet, enlarged gular pits, and alignment of punctures of abdominal sterna into one transverse line per sternum. On the other hand, O. zimmermani cannot simply be regarded as a species of Navitia with reduced eyes, as it lacks two specializations typical of Navitia : obsolete frontal grooves and an incomplete marginal groove on the pronotum. We therefore consider it best to isolate this species in a subgenus of its own. Omoglymmius ( Caeconavitia ) zimmermani (Bell and Bell) NEW COMBINATION Omoglymmius ( Nitiglymmius ) zimmermani Bell and Bell, 1978:77. Type material. - Part 1:77. Description. - Part 1:77. Indoglymmius new subgenus Type species. - Rhysodes lineatus Grouvelle 1908 Description. — Segment XI of antenna obtuse, without stylet; basal setae of antenna absent; clypeal setae absent; medial margin of temporal lobe with pale translucent area; medial angles simple; frontal grooves straight and linear; temporal lobe with line of punctures along orbital groove, otherwise impunctate; temporal setae absent; frontal grooves linear; frontal space very small; postorbital tubercle absent; median groove of pronotum coarsely conspicuously punctate; punctures pollinose, groove otherwise glabrous; paramedian grooves coarsely punctate, but inconspicuously so, punctures partly hidden by pollinosity; pronotal carinae impunctate; borders between carinae and grooves somewhat irregular, with carinae encroached by punctures of grooves; elytral striae scarcely impressed; Stria IV with one seta at apex; apical stride with one seta; apex of Stria VII with several setae; abdominal sterna with scattered punctures; both sexes with deep lateral pits on abdominal Sternum IV, and with shallow ones on Sternum V; ventral tooth on anterior femora in both sexes. The only species in this subgenus resembles Laminoglymmius in having a conspicuous translucent area near the medial angle of the temporal lobe, but differs in lacking basal setae on the antennae. The simple medial angles separate it from all Laminoglymmius except for O. (L) inaequalis. The very narrow outer pronotal carinae separate the latter species from Indoglymmius. The translucent area is the most conspicuous character distinguishing Indoglymmius from Omoglymmius s. str. Omoglymmius ( Indoglymmius ) lineatus (Grouvelle) NEW COMBINATION Figs. 77, 83 Rhysodes lineatus Grouvelle 1908:319-320. Omoglymmius (s. str.) lineatus (Grouvelle) Bell and Bell 1978 Type material. - LECTOTYPE (here designated) male, labelled: “(INDIA), Chambaganor, Madura, 48” (MNHN) PARALECTOTYPES 17 specimens, sexes not recorded, same data as lectotype (MNHN; one male (labelled as “cotype”) labelled: “INDIA, Shembaganur, Madura, Andrews Bequest BM 1922-22” (BMNH) According to Grouvelle, the type series was collected by R. P. Dubreuil. Description. — Length 5. 6-6. 2 mm. Antennal Segment XI conspicuously narrower than Segment X; head slightly longer than wide; pronotum moderately elongate, length/greatest width 1.28; lateral margin distinctly sinuate anterior to nearly rectangular hind angle; middle calcar of male minute; hind calcar small, triangular, its apex obtuse. In addition to the type series, we have seen one male and one female, labelled “Key Inseln, coll. Reithoffer” (BSL). This refers to the Kei Islands, west of New Guinea. It is highly Quaest. Ent., 1982, 18 (1-4) 168 Bell and Bell unlikely that the species occurs both in the Kei Islands and southern India. Probably either the type series or the Basel specimens have incorrect locality data. Since there is no independent record from either place, the true provenance of this species must be listed as uncertain. Subgenus Nitiglymmius Bell and Bell 1978 Type species. - Omoglymmius ( Nitiglymmius ) fulgens Bell and Bell 1978 Description. — Antennal Segment XI without apical stylet; clypeal setae absent; frontal grooves deep but narrow to linear; eye reduced, in profile view, entirely dorsad to middle of head; ommatidia not distinct; cornea not facetted; cornea with pinkish tint; silvery-white reflecting disc medial to cornea and at angle to it; marginal grooves of pronotum fine, linear entire or partly or completely effaced; elytral stria not impressed, but represented by rows of punctures; spur of middle tibia with tip curved anteriorly; punctures of abdominal sterna scattered or in single transverse row on each segment; lateral pits present on Sternum IV in both sexes (where known); ventral surface with bluish opalescence (such opalescence on dorsal surface also in most species); hind wings vestigial. The description has been modified from that in Part I in accordance with removal of O. zimmermani to Subgenus Caeconavitia and addition of O. (N.) semioculatus new species from Philippines. The range of Nitiglymmius as here limited is New Guinea, the Solomon Islands and Siargao Island in the Philippines. KEY TO SPECIES (Supersedes that in Part I: 76-77) 1 Elytron with longitudinal scarp present at base of Stria IV; elytral striae slightly impressed, rather coarsely punctate; eye minute, less than 0.1 of depth of head. O. semioculatus new species, p. 169 T Elytron without scarp at base of Stria IV; elytral stria not at all impressed, very finely punctate; eye less minute, more than 0.1 of depth of head 2 2 (T) Pronotum relatively elongate, its margin evenly curved to apex, not constricted to form “collar.” O. greensladei Bell and Bell, (part I; 78) 2' Pronotum less elongate, its sides abruptly narrowed anteriorly, in form of “collar” at apex 3 3 (20 Postorbital tubercles distinct 4 3' Postorbital tubercles absent 5 4 (3) Pronotum distinctly narrowed at apex; lateral margin of pronotum sinuate just anterior to hind angles; posterior median pit large, as wide as basal impression . O. hornabrooki Bell and Bell (Part I: 81) 4' Pronotum only slightly narrowed anteriorly, lateral margin not sinuate, posterior median pit very small, much narrower than basal impression O. offafinus Bell and Bell (Part I; 81) 5 (30 Marginal groove of pronotum present 6 5' Marginal groove absent O. lustrans Bell and Bell (Part I: 80) 6 (5) Frontal grooves very fine, linear; median lobe long, acutely pointed, extending posterior to middle of temporal lobes O. fulgens Bell and Bell (Part 1:80) 6' Frontal grooves broader, not linear; median lobe short, obtusely pointed, not extended to middle of temporal lobes O. toxopei Bell and Bell (Part I: 82) Revision of Omoglymmius Ganglbauer 169 Omoglymmius ( Nitiglymmius ) semioculatus new species Figs. 78,81,82 Type material. - HOLOTYPE male, labelled: “PHILIPPINEN: Mindanao, Dapa, leg. Boettcher, 30-9-16” (NMHB). Despite the label, the locality is not on Mindanao, but on the small island of Siargao, northeast of it. Description. — Length 6.2 mm. Form elongate for genus; dorsal surface not opalescent; ventral surface scarcely opalescent; head elongate, its lateral margin markedly sinuate anterior to eye; median lobe elongate, its apex rather acute, opposite middle of temporal lobe; frontal grooves long, wider than in other members of subgenus; orbital groove absent; frontal space larger than in other members of subgenus; medial angles rectangular, nearly contiguous; margin posterior to medial angle sinuate; one temporal seta; temporal lobe with about six fine punctures, restricted to lateral region; eye minute, oval, visible only in lateral view; postorbital tubercle absent; mentum punctate; gular region extensively pollinose. Pronotum elongate; length/greatest width 1.35; anterior end in form of short, ill-defined “collar”; lateral margin curved into “collar” anteriorly, slightly sinuate just anterior to hind angle; outer carina slightly narrower than inner one, with about nine punctures scattered along its lateral margin; inner carina impunctate; median groove deep, anterior and posterior median pits equal, moderately large; lateral margin of paramedian groove distinct, but medial margin sloped gradually from inner carina; marginal grooves broader than in other members of subgenus, complete; epipleura impunctate. Elytra elongate, more parallel-sided than in other members of subgenus; elytron with short, longitudinal scarp at base of Stria IV; elytral striae shallowly impressed, rather coarsely punctate; Stria IV with one seta near apex; Stria VII with one seta near apex; elytron in profile with highest point at apical fifth, sloped rather abruptly posterior to this point; metasternum with entire surface densely, coarsely, shallowly punctate; abdominal sterna similarly punctate; anterior femur of male without ventral tooth; hind calcar of male large, its apex obtuse, raised above spur, margin between apex and spur sinuate; margin proximal to calcar distinctly emarginate (Fig. 81). This species differs from all other members of the subgenus in the scarp at the base of Stria IV very reduced eyes, extensive pollinosity in the gula impressed, coarsely punctate elytral striae, posteriorly displaced elytral “hump” and the coarsely punctate metasternum and abdominal sterna. Form of the hind calcar is also distinctive. Subgenus Orthoglymmius Bell and Bell 1978 Type species. - Rhysodes sulcicollis Lewis 1888 Description. — Antennal Segment XI without stylet; basal setae of antenna absent; clypeal setae present; frontal grooves deep, broad; margins of temporal lobe oblique both medial to and lateral to occipital angle, posterior temporal lobe is pointed; temporal setae absent; eye large, deeper than long; cornea facetted; ommatidia distinct; marginal groove of pronotum complete; elytral setae relatively well-developed, with complete row on Stria IV, and in some species, with complete rows in Striae II and VI as well; spur of middle tibia straight or curved; metasternal punctures limited to medial band and lateral margin; abdominal Sterna 1 1- V each with single transverse row of very coarse punctures, or with punctures coalesced to form transverse sulci; female with lateral pits better developed on Sternum V than on Sternum IV; hind coxa with seta; Sternum VI with pair of setae in both sexes; males, as far as known, with ventral tooth on anterior femur but without proximal tooth on anterior tibia; hind calcar, where known, bisinuate; hind wings and genitalia not investigated. The straight spur of the middle tibia and deepest lateral pits on the fifth, rather than the fourth, abdominal sternum, distinguish this subgenus from Omoglymmius s. str.. Only a few members of the latter subgenus have the punctures of each abdominal sternum in a single transverse row, as is consistently true in Orthoglymmius. In addition, Orthoglymmius specimens have clypeal setae and have the temporal lobes pointed posteriorly. The Subgenus Orthoglymmius is confined to a part of the Oriental Region, where it is known from the Himalaya, the Indochina Peninsula, the Ryukyu Islands, and southern Japan. It is apparently absent from southern India and from Indonesia. Phylogeny. - This subgenus is perhaps the most inadequately known major group of Rhysodini. It consists of a complex of very similar and closely related species. Of the eight species, both sexes are known in only two. Several of the species described by Grouvelle are Quaest. Ent., 1982, 18 (1-4) 170 Bell and Bell known only from type material, and are so similar to one another that it would be premature to reach final conclusions about their validity. The group is so uniform that it is almost a waste of time to speculate about interrelationships among the species. The group will need a futher revision when adequate series of specimens are available and the genitalia can be studied. KEY TO SPECIES 1 V 2 (1) 2' 3 (10 3' 4 (3) 4' 5 (40 5' 6 (50 6' 7 (60 7 Medial angles of temporal lobes well separated, obtuse; posterior margin of temporal lobe without pilosity 2 Medial angles contiguous or almost so; posterior margin of temporal lobe fringed with pilosity 3 Postorbital tubercle distinct, 0.33 as long and 0.33 as deep as eye, glabrous except for tuft of pilosity at tip; tip of median lobe obtuse O. sulcicollis (Lewis), p. 170 Postorbital tubercle reduced to minute vestige with small pilose tuft at its tip; median lobe acute at tip O. microtis new species, p. 172 Postorbital tubercle pilose 4 Postorbital tubercle glabrous O.feae (Grouvelle), p. 172 Median groove closed anteriorly O. alticola (Grouvelle), p. 173 Median groove open anteriorly 5 Lateral margin of pronotum obtusely angulate at middle O. longiceps (Grouvelle), p. 173 Lateral margin of pronotum evenly curved at middle 6 Pronotal margins nearly parallel; lateral margin scarcely sinuate anterior to hind angle O. cavifrons (Grouvelle), p. 174 Pronotal margins more curved; pronotal base more distinctly narrowed; lateral margin distinctly sinuate anterior to hind angle 7 Orbital groove relatively wide, extending posteriorly to level of middle of eye; posteriomedial and posteriolateral margins of temporal lobe both distinctly sinuate; occipital angle obtusely pointed; postorbital tubercle large, prominent in dorsal view O. crenatus (Grouvelle), p. 175 Orbital groove narrow, short, ending at anterior 0.33 of eye; posteriomedial and posteriolateral margins of temporal lobe oblique or very slightly sinuate; occipital angle obtuse; postorbital tubercle smaller, barely visible in dorsal view. O. coomani (Arrow), p. 1 76 Omoglymmius (Orthoglymmius) sulcicollis (Lewis) Figs. 84, 92 Rhysodes sulcicollis Lewis 1 888:8 1 . Omoglymmius (Orthoglymmius) sulcicollis (Lewis) Bell and Bell 1978 Type material. - HOLOTYPE female, labelled: “JAPAN: Oyayama, Higo (Island of Honshu), 1881, coll. G. Lewis 1910-320” (BMNH). Description. — Length 6.0 mm. Antennal Segment XI as long as wide, its cone well-developed; clypeal setae present; median lobe short, tapered posteriorly, its apex obtuse, located anterior to widest point of head; anteriomedial margin of temporal lobe oblique, scarcely sinuate; medial angles obtuse, widely separated from one another; posteriomedial Revision of Omoglymmius Ganglbauer 171 Plate 7. Figures 84-103, Subgenus Orthoglymmius. Figs. 84-91, Head and pronotum, dorsal aspect; Fig. 84, Omoglymmius (Orthoglymmius) sulcicollis (Lewis); Fig. 85, O. (O.) microtis new species; Fig. 86, O. (O.) feae (Grouvelle); Fig. 87, O. (O.) alticola (Grouvelle); Fig. 88, O. (O.) longiceps (Grouvelle); Fig. 89, O. (O.) cavifrons (Grouvelle); Fig. 90, O. (O.) crenatus (Grouvelle); Fig. 91, O. (O.) coomani (Arrow); Figs. 92-99, Head, lateral aspect; Fig. 92, O. (O.) sulcicollis (Lewis); Fig. 93, O. (O.) microtis new species; Fig. 94, O. (O.) feae (Grouvelle); Fig. 95, O. (O.) alticola (Grouvelle); Fig. 96, O. (O.) longiceps (Grouvelle); Fig. 97, O. (O.) cavifrons (Grouvelle); Fig. 98, O. (O.) crenatus (Grouvelle); Fig. 99, O. (O.) coomani (Arrow); Figs. 100-103, Hind tibia, male, apical portion; Fig. 100, O. (O.) longiceps (Grouvelle); Fig. 101, O. (O.) cavifrons (Grouvelle); Fig. 102, O. (O.) crenatus (Grouvelle); Fig. 103, O. (O.) coomani (Arrow); Quaest. Ent., 1982, 18 (1-4) 172 Bell and Bell and posteriolateral margins oblique, not margined by pilosity; occipital angle obtuse but distinct; orbital groove narrow, ended just anterior to middle of eye; postorbital tubercle small, 0.33 as deep and 0.33 as long as eye, with tuft of long pollinosity. Pronotum moderately elongate, length/greatest width 1.25; widest at middle, sides strongly curved, narrowed to apex, slightly narrowed to base; lateral margin with slight sinuosity anterior to hind angle; paramedian grooves narrower than carinae at level of middle of pronotum; carinae broad, impunctate; pronotal setae absent. Elytra rather short, broad for subgenus; striae subequal, intervals equally convex; base of Interval V not elevated; Stria I with two to four setae near apex; Stria 11 with two setae near apex (in one specimen these setae are actually slightly laterad to stria in medial part of Interval III); Stria IV with five or six setae in form of complete row; apical stride with three setae; Stria VI with two setae near humerus; Stria VII with about seven setae near apex. Female with shallow depression at middle of metasternum, abdominal Sterna II-V each with a transverse row of punctures; male unknown. Widely separated medial angles and absence of marginal pollinosity from the vicinity of the occipital angles separate this species from all other except O. microtis. The postorbital tubercle is much better developed than in the latter species. In addition to the holotype, we have studied one female, from the collection of Dr. Sato, labelled simply “Japan”. Omoglymmius (Orthoglymmius) microtis new species Figs. 85, 93 Type material. - HOLOTYPE female, labelled: “Loochoo, Is. Ishigaki, Takeda, 17-IV-1962, S. Tamai leg.” (SATO). Ishigaki is an island in the Sakashima Retto, the southernmost group in the Ryukyu Islands. Description. — Length 5.3 mm. Antennal Segment XI as long as wide, its cone distinct; clypeal setae present; median lobe more elongate, more pointed than in O. sulcicollis, its tip even with greatest width of head; anteriomedial margin of temporal lobe shallowly sinuate, nearly oblique; medial angles obtuse, widely separated; posteriomedial, posteriolateral margins oblique, not margined by pilosity; occipital angle obtuse; orbital groove narrow, ended just anterior to middle of eye; postorbital tubercle minute, less than 0.2 as long as eye, located immediately ventrad to tuft of short pilosity. Pronotum moderately elongate, length/greatest width 1.25; widest at middle, sides curved, narrowed to apex, slightly narrowed to base, side slightly sinuate anterior to hind angle; paramedian grooves narrower than carinae at level of middle of pronotum; carinae broad, impunctate; pronotal setae absent. Elytra longer and more cylindrical than in O. sulcicollis ; stria subequal, intervals equally convex; base of Interval V not at all elevated anteriorly; Stria I with three or four setae near apex; Stria II with one to two setae near apex; Stria IV with six setae forming complete row; apical stride with two setae; Stria VI with two setae near humerus; Stria VII with about six setae near apex. Female with shallow depression at middle of metasternum; abdominal Sterna II-V each with transverse row of punctures; male unknown. This species differs from all other members of the subgenus in the extreme reduction of the postorbital tubercle. It is otherwise very similar to O. sulcicollis of Japan. It contrasts sharply with O. cavifrons of the nearby island of Taiwan in shape of the temporal lobes. Omoglymmius ( Orthoglymmius ) feae (Grouvelle) Figs. 86, 94 Rhy socles feae Grouvelle 1895b:76 1-762. Omoglymmius ( Orthoglymmius ) feae (Grouvelle) Bell and Bell 1978 Type material. - HOLOTYPE female, labelled: “BURMA, Carin Cheba, 900-1100 m, XII-88, coll. L. Fea” (MNHN). This may be a unique type, although the original dscription implies the existence of a series of specimens. Revision of Omoglymmius Ganglbauer 173 Description. — Length 6.9 mm. Antennal Segment XI as long as wide, its cone distinct; clypeal setae present; median lobe tapered posteriorly, its tip obtuse; anteriomedial margin distinctly curved, emarginate; medial angles acute, contiguous; posteriomedial and posteriolateral margins oblique, slightly sinuate, margined by pilosity; occipital angle distinct; orbital groove narrow, ending anterior to middle of eye; postorbital tubercle large, 0.5 as long and 0.66 as deep as eye, glabrous. Pronotum moderately long, length/greatest width 1.22; widest near middle, strongly curved and narrowed to apex; slightly to base; lateral margin slightly sinuate anterior to hind angle; paramedian grooves narrower than carinae at level of middle of pronotum; carinae broad, impunctate; pronotal setae absent. Elytra moderately long; striae subequal, intervals equally convex; base of Interval V not at all elevated; Stria 1 with four setae near apex; Stria II with three setae near apex; Stria IV with seven setae in form of complete row; apical striole with one seta; Stria VI with two setae near humerus; Stria VII with six or seven setae near apex; female without impression in metasternum; abdominal Sterna Il-V each with transverse row of punctures. The single specimen assigned to this species differs from other members of the group in having the postorbital tubercle glabrous. Omoglymmius (Orthoglymmius) alticola (Grouvelle) Figs. 87, 95 Rhysodes alticola Grouvelle 1913: 99-100. Omoglymmius ( Orthoglymmius ) alticola (Grouvelle) Bell and Bell 1978 Type material. - HOLOTYPE male, labelled: “Kobo, 400 feet, Abor Exped, 30-XI-ll, Kemp” (NMHN). The Abor Hills are now in the State of Arunachal Pradesh, in northeastern India. Description. — Length 5.5 mm. Antennal Segment XI as long as wide, its cone distinct; clypeal setae present; median lobe tapered posteriorly, its tip obtuse; anteriomedial margin strongly curved and emarginate; medial angles acute, contiguous; posteriomedial and posteriolateral margins oblique; margined by pilosity; occipital angle obtuse; orbital groove narrow, ended anterior to middle of eye; postorbital tubercle large, 0.5 as long and 0.67 as deep as eye; postorbital tubercle and postorbit pilose. Pronotum moderately long, length/greatest width 1.25, widest near middle, strongly curved and narrowed to apex, moderately incurved and narrowed to base; lateral margin scarcely sinuate anterior to hind angle; paramedian grooves narrower than outer carinae near middle; inner carinae relatively narrow, about 0.67 as wide as outer ones at middle; median groove closed at anterior end; carinae impunctate; pronotal setae absent. Elytra relatively long, narrow; stria subequal, intervals equally convex; Interval V not at all elevated at base; Stria 1 with four setae near apex; Stria III with three setae near apex; Stria IV with complete row of six or seven setae; medial face of apical tubercle with one seta; apical striole with three setae; Stria VI with three setae near humerus; Stria VII with six setae near apex, male with shallow median impression on metasternum and ventral tooth on anterior femur; middle calcar small, acute; hind calcar bisinuate; female unknown. The distinctive marks of this species are that the median groove is closed anteriorly and that the inner carina is distinctly narrower than the outer one. The species most nearly sympatric is O. crenatus. The latter species differs in having the median groove open anteriorly, the inner carinae as broad as the outer ones, the orbital groove both wider and longer, and the occipital angles more pointed. Omoglymmius ( Orthoglymmius ) longiceps (Grouvelle) Figs. 88, 96, 100 Rhysodes longiceps Grouvelle 1910: 324-325. Omoglymmius (Orthoglymmius) longiceps (Grouvelle) Bell and Bell 1978 Type material. - HOLOTYPE female, labelled: “BURMA,” without definite locality or collector (MNHN). According to the original description, there was a type series of four specimens, but we have been unable to locate the other three. Quaest. Ent., 1982, 18 (1-4) 174 Bell and Bell Description. — Length 6-7 mm. Antennal Segment XI as long as wide, its cone distinct; clypeal setae present; head relatively elongate; median lobe tapered posteriorly, its tip obtuse; anteriomedial margin curved, emarginate; medial angles acute, nearly contiguous; posteriomedial and posteriolateral margins oblique, slightly sinuate, margined by pilosity; occipital angle distinct; orbital groove narrow, ended anterior to middle of eye; postorbital tubercle large, 0.67 as long and 0.67 as deep as eye, prominent in dorsal view; postorbital tubercle and postorbit pilose. Pronotum moderately long, length/greatest width 1.21; widest near middle, margin in form of obtuse angle at widest point; sides oblique and slightly curved both anterior and posterior to angle; apex strongly narrowed; base distinctly narrowed; margin not sinuate anterior to hind angle; paramedian grooves narrower than carinae at level of middle of pronotum; carinae broad, impunctate; pronotal setae absent. Elytra moderately long; striae subequal; intervals equally convex; base of Interval V not elevated; Stria I with five setae near apex; Stria II with three setae near apex; Stria IV with five or six setae in complete row; medial face of apical tubercle with one seta; apical striole with two setae; Stria VI with one or two setae near humerus; Stria VII with about five setae near apex; both sexes with lateral pit in abdominal Sternum V; male with shallow concavity at middle of metasternum; abdominal Sterna 1 1- V each with transverse row of punctures. Male with ventral tooth on anterior femur; middle calcar acute; hind calcar with obtuse triangular point and rectangular “shoulder” proximad to it, separated from point by shallow emargination (Fig. 100). In addition to the holotype, we have studied one male specimen, labelled “Carin Ghecu 1300-1400 m, L. Fea, II-III 88, coll. Jul. Moser” (MNHB). This specimen bears the label “/?. feae Grouv.” but resembles the holotype of R. longiceps in having the postorbital tubercles pollinose, and in the shape of the pronotum. In our opinion, it represents the male of R. longiceps. A female, labelled: “Carin-Ghecu 300-1400 m.; L. Fea, V-88” (GEN), and a male, labelled: “Tenasserim, Meetan, Fea, April, 1887” (GEN) also appear to be O. longiceps. O. coomani is closely related, but appears to differ consistently in the narrower pronotum without distinct lateral angles, the shorter, more rounded occipital angles, and in the somewhat more elongate median lobe. Omoglymmius (Orthoglymmius) cavifrons (Grouvelle) Figs. 89, 97, 101 Rhysodes cavifrons Grouvelle 1914: 33-34. Omoglymmius ( Orthoglymmius ) cavifrons (Grouvelle) Bell and Bell 1978 Type material. - LECTOTYPE (here designated) sex not recorded, labelled: “Kosempo. Formosa, coll. H. A. Sauter, 191 1” (MNHN). PARALECTOTYPES: The original description mentions a type series of four specimens. We have seen one specimen from the MNHB collection labelled “Kosempo, 11-08, Sauter, S.V.” which is a probable paralectotype. The remaining two specimens, if still in existence, would also be paralectotypes. Description. - Length 5-6 mm. Antennal Segment XI as long as wide, its cone distinct; clypeal setae present; median lobe short, its tip obtuse, anterior to middle of eye; anteriomedial margin distinctly curved, sinuate; medial angles contiguous or nearly so, acute; posteriomedial and posteriolateral margins oblique, margined with pilosity; occipital angle nearly rectangular; orbital groove rather broad, ended posterior to middle of eye; postorbital tubercle rather large, 0.67 as deep and 0.67 as long as eye; postorbital tubercle and postorbit pilose. Pronotum rather elongate, length/greatest width 1.26; widest near middle, lateral margins curved and convergent anteriorly; apex strongly narrowed; margins straight, almost parallel posteriorly, base only slightly narrowed; margin not sinuate anterior to hind angle; paramedian grooves narrower than carinae at level of middle of pronotum; carinae broad, impunctate, pronotal setae absent. Elytra rather long, narrow; striae subequal; intervals equally convex; base of Interval V not at all elevated; Stria I with three or four seta near apex; Stria II with two setae near apex; Stria IV with six setae in form of complete row; medial side of apical tubercle with one seta; apical striole with two or three setae; Stria VI with three setae near humerus; Stria VII with about five setae near apex. Abdominal Sterna II-V each with transverse row of punctures; both sexes with lateral pits in Sternum V; male with ventral tooth on anterior femur; middle calcar acute; hind calcar with two obtuse angles, one above the other, separated by shallow sinuation (Fig. 101). This species differs from O. coomani and other species from the mainland of Asia in having the pronotum longer and narrower, with the lateral margins straighter and more nearly Revision of Omoglymmius Ganglbauer 175 parallel. It differs sharply from O. microtis of the Ryukyu Islands in having the postorbital tubercles much larger and more strongly pilose, and in having the medial angles of the temporal lobes contiguous or nearly so. Range. - O. cavifrons is restricted to Taiwan. In addition to the lectotype, we have seen the following specimens: three males, two females, Chipen Spa, Taitung Hsien, 3-6, V-1972, M. Sakai leg. (SATO); one male, one female, Chipon, Formosa, 1 V- 1 8-32, coll. L. Gressitt (CAS); one male, one female, Taiwan: Hori (Puli), July, 1954, native collector (BPBM). Miwa (1931) cites specimens from Kosempo, Sokutsu and Chip-Chip. Omoglymmius (Orthoglymmius) crenatus (Grouvelle) Figs. 90,98, 102 Rhysodes crenatus Grouvelle 1903: 119. Omoglymmius crenatus (Grouvelle) Bell and Bell 1977 Omoglymmius ( Orthoglymmius ) crenatus (Grouvelle) Bell and Bell 1978 Type material. - HOLOTYPE: According to the original description, female, Bhoutan, coll. Maria Borsti, in collection of R. Oberthiir. We have not been able to locate the type specimen, but have described and illustrated a female specimen corresponding well to the original description, from Balu-Jhura, Bhutan (Bell 1977). Description. — Length 5.5 mm. Antennal Segment XI as long as wide, its cone distinct; head relatively elongate; clypeal setae present; median lobe tapered posteriorly, its tip acute, slightly anterior to widest point of head; anteriomedial margin distinctly curved, emarginate; medial angles acute, nearly contiguous; posteriomedial margin shallowly emarginate; posteriolateral margin oblique; both margined with pilosity; occipital angle distinct; orbital groove relatively broad, ended opposite middle of eye; postorbital tubercle 0.5 as deep and 0.5 as long as eye; postorbital tubercle and postorbit pilose. Pronotum moderately elongate, length/greatest width 1.23; widest slightly anterior to middle, markedly and very abruptly narrowed at apex; slightly narrowed to base; sides oblique, not sinuate anterior to hind angles; paramedian grooves narrower than carinae at level of middle of pronotum; carinae broad, impunctate; pronotal setae absent. Elytra relatively long, narrow; Striae subequal; intervals equally convex except for more elevated base of Interval V, latter with fine microsculpture opaque; Stria I with four setae near apex; Stria II with three setae near apex; Stria IV with complete row of four or five setae; one seta on medial face of apical tubercle; apical striole with four setae; Stria VI with two setae near humerus; Stria VII with four setae near apex; both sexes with shallow median impression on metasternum; abdominal Sterna 1 1 - V each with transverse row of punctures; both sexes with deep lateral pit in Sternum V; male with ventral tooth on anterior femur; middle calcar acute; hind calcar with two very obtuse lobes, one proximad to the other, separated by shallow sinuation (Fig. 102). In this species, the base of Interval V is more distinctly elevated than in any other member of the subgenus. O. coomani is similar but has the base of Interval V less elevated and without microsculpture. It also has the lobes of the hind calcar less rounded and the postorbital tubercles smaller. O. cavifrons resembles this species in having the orbital groove dilated, but differs in having the base of Interval V not at all elevated, and shape of the frontal space and pronotum different. O. alticola is possibly sympatric with O. crenatus in the eastern Himalaya. O. alticola differs in having the orbital grooves fine, base of Interval V not at all elevated, median groove of the pronotum closed anteriorly, and inner pronotal carinae conspicuously narrower than the outer ones. Range. - Foothills of the Himalayas in India, Bhutan and also in Laos. We have seen the following additional Specimens:BHUTAN; one female, Balu-Jhura, 200 m, 28-4-72, coll. Basel Natural History Museum Exped. (BSL); INDIA: one male, Haldwani Dist., Kumaon, coll. H. Gower Champion, British Museum, 1953-156 (BMNH); LAOS: two males, ULR farm, 3200', Xieng Khouang Prov., 2 mi. NW, Long Tieng, 18-Jan-1968 and 3-Mar-1968, coll. G. L. Peters (GLP). Quaest. Ent., 1982, 18 (1-4) 176 Bell and Bell Omoglymmius ( Orthoglymmius ) coomani (Arrow) Figs. 91, 99, 103 Rhysodes coomani Arrow 1942: 180. Omoglymmius ( Orthoglymmius ) coomani (Arrow) Bell and Bell 1978 Type material. - LECTOTYPE (here designated) male, labelled: “Tonkin: Hoabinh, coll. A. de Cooman, BM 1940-13” (BMNH). PARALECTOTYPES (all same collection data as lectotype), one male labelled “co-type”, BM 1929-299; one male, same data as lectotype; one male, BM 1929-299; one male, BM-1925-251 (all BMNH). Description. — Length 5. 8-6. 5 mm. Antennal Segment XI as long as wide, its cone distinct; clypeal setae present; median lobe tapered posteriorly; its tip obtusely pointed; anteriomedial margin of temporal lobe distinctly curved, emarginate; medial angles acute, contiguous, posteriomedial and posteriolateral margins oblique, margined by pilosity; occipital angles more obtuse than in O. crenatus, orbital groove narrow, short, ended anterior to middle of eye; postorbital tubercle rather small, 0.33 as long and 0.33 as deep as eye; postorbital tubercle and postorbit pilose. Pronotum moderately long, length/greatest width 1.23; widest near middle, curved to both base and apex; base more narrowed than in related species; side not sinuate anterior to hind angle; paramedian grooves narrower than carinae at level of middle of pronotum; carinae broad, impunctate; pronotal setae absent. Elytra rather narrow; striae subequal; intervals equally convex except that base of Interval V is slightly elevated (varied in degree of distinctness); Interval V not microsculptured near base; Stria I with four setae near apex; Stria II with two or three near apex; Stria IV with complete row of seven to nine setae; medial face of apical tubercle with one to three setae; apical striole with three to five setae; Stria VI with two to four setae near humerus; Stria VII with six to eight setae near apex. Both sexes with lateral pit in Sternum V; male with ventral tooth on anterior femur; middle calcar acute; hind calcar with two lobes separated by sinuation; these lobes less rounded than in O. crenatus (Fig. 103). This rather nondescript species varies in the degree to which the base of Interval V is elevated. Those specimens in which the elevation is rather distinct are most liable to be mistaken for O. crenatus. The latter species has a large postorbital tubercle, occipital angle more distinct and orbital groove longer and more dilated. Those specimens with the base of Interval V only slightly elevated are liable to be mistaken for O. longiceps or O. feae. In the former species, the median lobe is shorter and more obtuse, while the latter species has a glabrous postorbital tubercle. Range. - O. coomani is known from Vietnam, Thailand and Java. In addition to the type series, we have seen the following specimens: java: one female, Mt. Djampangs (MNHN); Thailand: two females, Khao-Yai Nat. Pk., 750 m, VI 1-26-62, coll. E. S. Ross, D. Q. Cavagnaro (CAS); two males, E. slope of Doi Sutep, 875-950 m, 1 5- V 1 1- 1 962, coll. E. S. Ross and D. Q. Cavagnaro (CAS); one male, “SIAM”, Mouhot, colln. G. Lewis, 1901-31 (BMNH). VIETNAM: one female, Hoabinh, Tonkin, labelled with the manuscript name “R. tonkinensis” in Grouvelle's handwriting (MNHN); one female, 30 km NW of Pleiku, elevation 300 m, 10-V-1960, coll. L. W. Quate (BPBM); one female, nr. Tam-Dao, 900 m (N. Vietnam), 30-X-1963. coll. Kabakov (LEN); one male, five females, Tam-Dao, Son-Zuong, 200-300 m, 31-1. 1-2, 20-11, 24-11, 1962, coll. Kabakov (LEN); two males, one female, NE Thai-Nguen, 1 9-X 1 1 - 1 962, coll. Kabakov (LEN); three males, three females, 50 km No. Thai-Nguen, 300-400 m, 2-1 1 1, 3-3, 5-2, 19-12, 1963, coll. Kabakov (LEN); one male, NE Thai-Nguen, 1 1-1-1964, coll. Kabakov (LEN); one female, Tong-Hoa Phu (MNHN). Subgenus Carinoglymmius new subgenus Type species. - Rhysodes carinatus Grouvelle 1903 Description. — Antennal Segment XI without stylet; basal setae of antenna present or absent; clypeal setae absent; frontal grooves deep, broad; margins of temporal lobe rounded posteriorly in one species, angulate posteriorly in other species; temporal setae absent; eye large, deeper than long; cornea facetted; ommatidia distinct; marginal groove of pronotum complete, dilated; outer carina of pronotum very narrow, linear; elytral setae relatively well-developed, with complete or partial rows in Striae 11, IV, VI; metasternal punctures evenly scattered; spur of middle tibia straight or nearly so; abdominal Sterna 1 1 - V each with single transverse row of very coarse punctures; female with lateral pits better developed on Sternum V than on Sternum IV; hind coxa with a seta; Sternum VI with pair of setae in both sexes; males, as far as known, with ventral tooth on anterior femur but without proximal tooth on anterior tibia; hind wings and genitalia Revision of Omoglymmius Ganglbauer 177 not investigated. The straight spur of the middle tibia and presence of deepest pits on the fifth, rather than fourth abdominal sternum separate this subgenus from all subgenera except Orthoglymmius. Absence of clypeal setae and very narrow outer carina separate Carinoglymmius from the latter subgenus. The Subgenus Carinoglymmius is restricted to a small part of the Oriental Region, where it is known from Borneo, Sumatra, Mentawei, and the Andaman Islands. Phylogeny. - O. nicobarensis is clearly less specialized and more isolated than the other two species, having the inner pronotal carina broad, pronotal setae absent, and the elytral stria equally deep. O. carinatus and O. hexagonus form a pair of highly specialized and closely related species, with the inner carinae linear, the outer carinae each with a row of punctures, pronotal setae present, and elytral Stria II and IV much deeper than the others. In these two species, both antennae and legs are much thickened in a manner reminiscent of some species of Rhyzodiastes. KEY TO SPECIES 1 Inner carina of pronotum more than twice width of outer carinae; outer carina impunctate; elytral striae of equal depth; intervals not carinate; pronotal setae absent O. nicobarensis (Grouvelle), p. 177 V Inner carina very narrow, subequal to outer carina; outer carina punctate; Stria II, IV much deeper than other striae; some elytral intervals carinate; pronotum with row of setae in marginal groove 2 2 (1) Posterior margin of temporal lobe obtusely pointed; pronotum clearly hexagonal O. hexagonus (Grouvelle), p. 178 2‘ ' Posterior margin of temporal lobe rounded; sides of pronotum more evenly curved O. carinatus (Grouvelle), p. 179 Omoglymmius ( Carinoglymmius ) nicobarensis (Grouvelle) NEW COMBINATION Figs. 104, 108 Rhysodes nicobarensis Grouvelle 1895b: 762. Type material. - HOLOTYPE female, labelled: “I. ANDAMAN” (MNHN). This specimen is labelled as the type, and the original description, despite the specific name, gives the locality as the Andaman Islands. A second specimen identified as this species by Grouvelle (MNHN), is described above as Omoglymmius (Laminoglymmius) inaequalis new species. It was collected in the Nicobar islands, and is responsible for the listing of R. nicobarensis from the latter islands by Hincks (1950). Our placement of R. nicobarensis in subgenus Hemiglymmius (1978) was based on this specimen and not the true R. nicobarensis. Description. — Length 5.5 mm. Antennal Segment XI longer than wide, with well-developed cone; basal setae absent; medial lobe of head pointed posteriorly; anteriomedial margin of temporal lobe shallowly sinuate; medial angles obtuse, distinctly separated; orbital groove abbreviated; postorbital lobe small. Pronotum slightly longer than in related species, length/greatest width 1.29; widest near middle, sides curved, apex strongly narrowed, base only slightly narrowed; lateral margin not sinuate anterior to hind angle;xparamedian grooves dilated, as broad as inner carinae, almost three times as broad as outer carina; marginal groove dilated; inner carina broad; outer carina linear, only 0.33 as broad as inner one, impunctate; pronotum; without setae. Elytra moderately elongate; striae of equal depth; Interval V slightly elevated at base; Stria I with one seta in apex; Stria II with four near apex; Stria IV with complete series of approximately seven setae; apical striole with two setae; Stria Quaest. Ent., 1982, 18 (1-4) 178 Bell and Bell Plate 8. Figures 104-112, Subgenus Carinoglymmius. Figs. 104-106, Head and pronotum, dorsal aspect; Fig. 104, Omoglymmius ( Carinoglymmius ) nicobarensis (Grouvelle); Fig. 105, O. (C.) carinatus (Grouvelle); Fig. 106, O. (C.) hexagonus (Grouvelle); Fig. 107, Left elytron, dorsal aspect, O. (C.) hexagonus (Grouvelle); Figs. 108-110, Head, lateral aspect; Fig. 108, O. (C.) nicobarensis (Grouvelle); Fig. 109, O. (C.) carinatus (Grouvelle); Fig. 110, O. (C.) hexagonus (Grouvelle); Fig. Ill, Hind tibia, male, apical portion, O. (C.) hexagonus (Grouvelle); Fig. 112, Antennal Segments IX-XI, O. (C.) hexagonus (Grouvelle); VI with complete series of about six; Stria VII with four near apex; female with rather deep lateral pit in abdominal Sternum V; male unknown. This species is easily recognized within the subgenus by the combination of broad inner and linear outer pronotal carinae. This feature is duplicated in other subgenera from the Andaman and Nicobar Islands: Omoglymmius (Laminoglymmius) inaequalis Bell and Bell, and Omoglymmius (s. str.) solitarius (Arrow), a striking and unexplained example of convergent evolution. Omoglymmius (Carinoglymmius) hexagonus (Grouvelle) NEW COMBINATION Figs. 106, 107, 110-112 Rhysodes hexagonus Grouvelle 1903: 120-121. Omoglymmius ( Orthoglymmius ) hexagonus (Grouvelle) Bell and Bell 1978 Type material. - LECTOTYPE (here designated) male, labelled: (SUMATRA) “Palembang”, collector and date unspecified (MNHN). PARALECTOTYPES one male, Revision of Omoglymmius Ganglbauer 179 three females, same data (MNHN). Description. — Length 5. 0-6. 2 mm. Antennae very short, thick; Segments IV-XI transverse, cylindrical (Fig, 1 12); Segment XI with cone very low, apex almost flat; Segments II-IV each with broad apical pollinose ring; Segments V-XI each with narrow basal pollinose ring; Segments V-X each with short basal setae without pollinose ring; head elongate; median lobe nearly parallel-sided, its apex broadly rounded; frontal grooves dilated; anteriomedial margin very deeply emarginate; medial angles obtuse, well separated; temporal and antennal lobes separated by broad pollinose space; occipital angle obtuse but distinct; postorbital tubercle distinct though small. Pronotum with length/greatest width 1.22, its outline hexagonal, sides nearly parallel in middle 0.33; oblique, moderately convergent in basal third; oblique, strongly convergent in apical third; paramedian groove dilated, almost three times as broad as either carina; marginal groove dilated, with approximately 10 marginal setae; setae clavate; inner and outer carinae subequal, both very narrow, sublinear; inner one straight, impunctate; outer one slightly curved, with row of punctures. Elytra elongate, cylindrical; Intervals III and V carinate, convex; Intervals I, II and IV very low and narrow; all striae coarsely punctate (Fig. 107); Stria II with seven to ten setae; Stria IV with eight to eleven setae (most posterior of these on medial face of carinate Interval V); apical stride with three or four setae; Stria VI with eight or nine setae; Stria VII with five to eight setae near apex; setae clavate; male with prominent ventral tooth on anterior femur; calcar of middle tibia small, acute; calcar of hind tibia rather long, its margins nearly parallel, its tip obtusely rounded (Fig. 111). This species and O. carinatus are very similar, having the inner pronotal carinae very narrow, the outer carinae punctate, the marginal groove setose, and Intervals III and V of the elytra elevated and carinate. In O. hexagonus the occipital angles are obtuse and distinct, while in the latter species they are completely rounded. Range. - Sumatra and Borneo. In addition to the type material, we have seen one specimen, labelled:“W.SARAWAK, Mt. Matang, coll. G. E. Bryant, Dec. 1913” (BMNH). Possibly also Mentawei. We have examined two specimens from Mentawei, labelled: “Mentawei:Sipora, Sereinu, V-VI-94, Modigliani” (GEN). One of these appears to us to be O. hexagonus and the other O. carinatus. Both are labelled “ Rhysodes carinatus Grouvelle.” The O. hexagonus has a “typus” label, and the R. carinatus a “syntype” label. It is possible that O. carinatus is not distinct from O. hexagonus , but more material is needed to decide the question. Omoglymmius (Carinoglymmius) carinatus (Grouvelle) NEW COMBINATION Figs. 105, 109 Rhysodes carinatus Grouvelle 1903: 121. Omoglymmius (Orthoglymmius) carinatus (Grouvelle) Bell and Bell 1978 Type material. - LECTOTYPE (here designated) female, labelled: “BORNEO: River Sambey pres Ngabang, 1897, coll. J. B. Ledru” (MNHN). PARALECTOTYPE male, labelled: “Mentawei.Sipora, Sereinu, V-VI-94, Modigliani” (GEN) As mentioned above a second male with the same label is O. hexagonus , if the latter is specifically distinct. Description. — Length 6 mm. Antennae very short, thick; Segments IV-XI transverse, cylindrical; Segment XI with cone very low, so apex is almost flat; Segments II-IV each with broad apical pollinose ring; Segments V-XI each with narrow basal pollinose ring; Segments V-X each with short basal setae within pollinose ring. Head broader than in O. hexagonus ; median lobe nearly parallel-sided, its apex rounded; temporal lobe separated from antennal lobe by broad pollinose space; anteriomedial margin of temporal lobe deeply emarginate; medial angles rounded, well separated; temporal lobe evenly rounded posteriorly, occipital angle indistinct; postorbital tubercle distinct though small. Pronotum with length/greatest width 1.21; widest near middle, narrowed to both base and apex; sides more evenly curved than in O. hexagonus ; outline not clearly hexagonal; paramedian groove dilated, almost three times as broad as either carina; marginal groove dilated, with seven to nine setae; inner and outer carinae subequal, both very narrow; inner ones straight, impunctate; outer one more curved than in O. hexagonus , with a row of punctures which extends almost to anterior end. Elytra elongate, cylindrical; Intervals III, V carinate, convex; Intervals 1, II and IV very low and narrow; all striae coarsely punctate; Stria II with seven or eight setae; Stria IV with 10-13 setae (the most posterior of these are located on the medial face of the carinate Interval V); apical stride with three or four setae; Stria VI with five or six setae; Stria VII with 6 setae in apex; female with moderately deep lateral pit in abdominal Sternum V; male with legs as in O. hexagonus. This species is close to O. hexagonus , but differs in having a wider head, temporal lobes rounded posteriorly, and the sides of the pronotum not distinctly hexagonal. Quaest. Ent., 1982, 18 (1-4) 180 Bell and Bell Subgenus Omoglymmius sensu stricto Ganglbauer 1892 Type species. - Rhysodes germari Ganglbauer 1892 Description. — Antennal Segment XI without stylet; basal setae of antennae absent; clypeal setae absent; frontal grooves deep, varied in width; margin of temporal lobe rounded lateral to occipital angle in all species except O. germari ; temporal setae one to four or none; eye deeper than long, in most species fully developed, more than 0.5 of depth of head; in few species, reduced in size, but not less than 0.33 of depth of head, and extended well below middle of side of head; cornea facetted, transparent, colorless; marginal groove of pronotum complete; setae of pronotum and elytra varied in development; spur of middle tibia curved anteriorly; punctures of abdominal sterna scattered in most species, forming single transverse row in a few species; lateral pits on abdominal Sternum IV in female, and at least suggested in male; ventral surface in most species without opalescent sheen; hind wings fully developed in most species (but probably reduced in those species with reduced eyes). This subgenus is most likely to be confused with Orthoglymmius and Nitiglymmius. Members of the former subgenus have clypeal setae, and have the lateral pits of the abdomen best developed on Sternum V. They also have the posteriolateral margin of the temporal lobe oblique, and the punctures of the abdomen arranged in a single transverse row on each sternum. Omoglymmius (s. str.) germari has the temporal lobes shaped somewhat like those of Orthoglymmius but has the sternal punctures scattered. A few species of Omoglymmius s. str. have the sternal punctures as in Orthoglymmius , but have the temporal lobes rounded laterally. A few species of Omoglymmius s. str. approach Nitiglymmius in having the eyes reduced, but the process has not gone as far, and the cornea is facetted and transparent, while the eye is 0.40 or more of the depth of the head and extends well below the middle in lateral view. The Subgenus Omoglymmius s. str. ranges from the Solomon Islands and the Carolines in the east to Europe in the west. There is only one species in Europe, two in India, and one in the Ryukyu Islands. There are possibly two species in northern Australia. Rhysodes ichthyocephalus Lea appears, on the basis of an excellent photograph of the type, to belong to this subgenus, and B. P. Moore has another, undescribed Australian species. As we have not studied these species, they have not been included in our keys. Phylogeny. - This subgenus contains more than 70 species which are very uniform in external appearance. The few characters of diagnostic value occur in many combinations, and some species widely separated localities are superficially very nearly alike. A few species, such as O. germari , are obviously somewhat isolated from the remainder. A few others, such as O. duplex and O. quadruplex, show evidence of close relationship with one another. Beyond these observations, attempts to construct a phylogeny would be highly speculative on the basis of present data. In the future, an analysis based on additional characters, such as male genitalia, would be more profitable. The key to species is of necessity very long and cumbersome. Therefore, we have added several regional keys. KEY TO SPECIES (WORLD) 1 Posteriolateral margin of temporal lobe oblique, meeting posteriomedial margin at prominent occipital angle; posteriomedial margin also oblique, posterior part of temporal lobe nearly rectangular O. germari (Ganglbauer), p. 194 V Posteriolateral margin of temporal lobe rounded; occipital angle not prominent; posteriomedial margin convex, oblique, or emarginate 2 2 (E) Antennal lobe, postantennal area densely microsculptured, not separated by antennal groove O. continuus new species, p. 226 Revision of Omoglymmius Ganglbauer 181 2' Antennal lobe glabrous, raised above level of postantennal area, or else separated from latter by distinct antennal groove 3 (2') Inner carina distinctly narrower than outer carina at middle; pronotal grooves coarsely, densely punctate O. malabaricus (Arrow), p. 207 3' Inner carina equal to or wider than outer carina at middle; pronotal grooves impunctate or sparsely punctate 4 (30 Temporal lobe nearly circular; frontal space very small, longer than wide; median lobe short, rhomboid; carinae of pronotum only moderately narrowed at base 4' Temporal lobe reniform or oblique; frontal space larger, in most species wider than long; median lobe either not rhomboid, or, if rhomboid, elongate; inner carina either pointed posteriorly or else constricted just anterior to base, with extreme base widened 5 (4) Medial angles of temporal lobes acute, contiguous; Stria IV with three to five setae along its length; total length 5. 1-5.2 O. sakuraii (Nakane), p. 207 5' Medial angles of temporal lobes obtuse, slightly separated; Stria IV with one or two setae near apex; total length 6. 1-7.1 O. laticeps Bell, p. 206 6 (4') Postorbital, suborbital tubercles absent 6' Either postorbital or suborbital tubercle present 7 (6') Fourth interval in form of sharp raised carina O. bicarinatus new species, p. 230 T Fourth interval flat or somewhat elevated, but not carinate 8 (70 Pronotum with two or three marginal setae, inserted in prominent punctures O. bucculatus (Arrow), p. 229 8' Pronotum without marginal setae 9 (80 Outer carina at middle equal to or only slightly narrower than inner carina, 0.66 or more of width of inner carina 9' Outer carina at middle 0.5 or less of width of inner carina 10 (9) Elytron with short pollinose longitudinal scarp at base of Stria IV, or with glabrous scarp ( O . summissus), or with pollinose spot but with scarp indistinct ( O . politus) 10' Elytron without longitudinal scarp or isolated pollinose spot at base of Stria IV 11 (10) Either or both pairs of pronotal carinae with six or more punctures (in most species with many more) IF Pronotal carinae without punctures, or with one or two punctures on inner carina or with one to five on outer carina 12(11) Inner carina with many punctures {very fine in O. vadosus) 12' Inner carina with none to two punctures 13 (12) Abdominal punctures indistinct, limited to lateral bands of pollinosity on Sterna III- V ; elytral punctures shallow, scarcely pollinose O. vadosus new species, p. 227 13' Punctures of abdominal sterna distinct, separated or barely coalescent laterally; elytral striae with distinct, pollinose punctures 14 (13') Median lobe of head narrow, its margins only slightly curved; outer carina of pronotum broadest at anterior margin, slightly flattened anteriorly; outer carina with lateral margin not sinuate anterior to hind angle O. summissus new species, p. 216 3 4 5 6 7 63 9 10 60 11 45 12 43 13 30 14 Quaest. Ent., 1982, 18 (1-4) 182 Bell and Bell 14 ' Median lobe relatively broad, its margins more strongly curved; outer carina widest near middle, in many specimens sinuate anterior to hind angle 15 15 (14") Orbital groove complete, continuously pollinose to base of temporal lobe 16 15' Pollinosity of orbital groove incomplete, ended posteriorly near posterior margin of eye, or else interrupted there 18 16 (15) Head relatively broad posteriorly, its lateral margin abruptly rounded near base; latter transverse; anteriomedial margin transverse; frontal space broadly U-shaped O. fringillus new species, p. 240 16' Head not broad posteriorly, lateral margin evenly rounded from eye to base; frontal space more narrow; anteriomedial margin oblique 17 17 (16') Outer carina markedly narrowed posteriorly, width at base less than 0.5 of greatest width; median lobe narrow, its junction with clypeus distinctly constricted; apex of median lobe obtuse; postantennal area convex, extensively pollinose O. wittmeri new species, p. 227 17' Outer carina scarcely narrowed posteriorly; width at base 0.9 of greatest width; median lobe relatively broad, its junction with clypeus scarcely constricted; apex of median lobe rounded; postantennal area less extensively pollinose O. gurney i new species, p. 235 18 (15') Tip of median lobe narrow, subacute; pronotum narrow, lateral margins strongly curved; eye somewhat reduced, with posterioventral margin oblique; antennal segments V-XI impunctate O. semperi new species, p. 217 18' Tip of median lobe broadly rounded or obtuse; pronotum quadrate or subquadrate, in most specimens with lateral margins less rounded; eye round, not reduced; antennal Segments VII-X, in some species with punctures indistinct 19 19 (18") Outer antennal segments, including X, with numerous coarse punctures 20 19' Outer antennal segments with punctures sparse, in some species extended only to Segment IX 22 20 (19) Postantennal region of temporal lobe in form of narrow, pollinose ridge; pronotum subquadrate O. oroensis new species, p. 240 20' Postantennal region glabrous posterior to antennal groove; pronotum distinctly narrowed anteriorly 21 21 (20') Medial angle of temporal lobe rounded; posteriomedial margin rounded; elytral punctures round, coarse O. puncticornis new species, p. 241 21' Medial angle obtusely pointed; posteriomedial margin slightly sinuate; strial punctures elliptical, fine, sparse O. viduus new species, p. 226 22 (19') Pronotum elongate, lateral margins only slightly curved; temporal lobe with 30-50 punctures; median lobe obtusely rounded; marginal groove deeper posteriorly O. scopulinus new species, p. 237 22' Pronotum shorter; temporal lobe with 20 or fewer punctures; medial angle produced; posteriomedial margin sinuate in most specimens; marginal groove not deeper posteriorly 23 23 (22') Median lobe with approximately 10 fine punctures O. vicinus (Grouvelle), p. 239 23' Median lobe impunctate 24 24 (23') Median lobe narrow, its tip acute 25 Revision of Omoglymmius Ganglbauer 183 24' Median lobe relatively broad, its tip obtuse 26 25 (24) Pronotum subquadrate, widest anterior to middle; outer carinae dilated, divergent near base O. classicus new species, p. 238 25' Pronotum subcircular, markedly narrowed at both base and apex; outer carinae narrow, parallel at bases O. princeps new species, p. 236 26 (24') Outer carinae narrowed to neither dilated nor divergent bases 27 26" Outer carinae scarcely narrowed to bases, latter either sinuate medially or else distinctly dilated 28 27 (26) Pronotum subquadrate, lateral margins nearly parallel; base of pronotum scarcely narrowed O. lindrothi new species, p. 232 21' Pronotum oval, lateral margins strongly curved; base of pronotum distinctly narrowed O. rusticus new species, p. 234 28 (26') Medial margin of temporal lobe distinctly angulate opposite tip of median lobe; frontal space more nearly U-shaped O. modicus new species, p. 233 28' Medial margin of temporal lobe scarcely angulate opposite tip of median lobe; frontal space more nearly V-shaped 29 29 (28') Medial angles of temporal lobe produced, blunt; posteriomedial margin emarginate; lateral margins of pronotum markedly curved; lateral pits of abdominal Sternum IV shallow in female O. manni new species, p. 234 29' Medial angles not produced; posteriomedial margin not emarginate; lateral margins of pronotum less curved, subparallel; lateral pits of Sternum IV deep, round in female O. regius new species, p. 235 30 (12') Median lobe very narrow, elongate, margins nearly parallel, tip acute; base of outer carina markedly narrowed O. crassicornis new species, p. 21 1 30' Median lobe moderate in width or broad, margins either parallel or not parallel; base of outer carina less narrowed 31 31 (30') Median lobe with sides parallel, tip obtuse; greatest width of outer carina at anterior 0.33; base of Interval V elevated laterad to longitudinal scarp of Stria IV; Antennal Segments V-XI punctate 32 3L Median lobe lance-shaped, margins not parallel; outer carina either of uniform width or else with greatest width at middle; base of Interval V not elevated; outer antennal segments with punctures indistinct or absent 33 32 (31) Lateral margin of outer carina deeply sinuate anterior to hind angle O. amplus new species, p. 21 1 32' Lateral margin scarcely sinuate anterior to hind angle O. modiglianii new species, p. 212 33 (31') Lateral margin of temporal lobe almost straight; anteriomedial margin of temporal lobe abruptly bent; frontal space very broad O. morditus new species, p. 228 33' Lateral margin of temporal lobe rounded; anteriomedial margin abruptly bent or rounded; frontal space narrower 34 34 (33') Head as broad as long or broader than long; frontal space broad, U-shaped 35 34' Head longer than broad; frontal space V- or U-shaped 38 35 (34) Outer carina slightly narrower than inner carina at middle O. caelatus Bell and Bell, p. 230 35' Outer carina equal in width at middle to inner carina 36 Quaest. Ent.. 1982, 18 (1-4) 184 Bell and Bell 36 (35') Antennal groove very narrow; temporal lobes very flat; length 5. 0-6. 8 mm . O. oceanicus Bell and Bell, p. 231 36' Antennal groove broader; temporal lobes convex; length 6. 5-7. 2 mm 37 37 (360 Pronotum nearly quadrate, scarcely narrowed posteriorly O. batchianus (Arrow), p. 223 37' Pronotum distinctly narrowed posteriorly, markedly narrowed anteriorly O. humeralis (Grouvelle), p. 224 38 (340 Antennal Segments V-VIII, and in some specimens also IX, X finely punctate 39 38' Antennal Segments V-XI impunctate 41 39 (38) Inner carina of pronotum constricted just anterior to base, latter broadened . .40 39' Inner carina not constricted just anterior to base; latter truncate O. re nut us new species, p. 236 40 (39) Preorbital pit extensively pollinose; median lobe broad O. trepidus new species, p. 242 40' Preorbital pit with pollinosity restricted; median lobe narrow O. cavea new species, p. 243 41 (38') Temporal lobe punctate over most of its surface; median lobe narrow, tip acute; sides of pronotum nearly parallel O. philippensis (Chevrolat), p. 209 41' Punctures of temporal lobe limited to lateral margin; median lobe slightly broader, tip more obtuse; pronotum widest at middle, sides curved 42 42 (41') Eye large, round; basal scarp of Stria IV distinct; male with ventral tooth on anterior femur O. imugani new species, p. 210 42' Eye slightly reduced; basal scarp of Stria IV very small, scarcely more than lateral margin of pollinose spot; male without ventral tooth on anterior femur. O. politus new species, p. 210 43 (11') Median lobe broad, margins rounded; abdominal Sterna IV, V with punctures not coalescent O. opticus new species, p. 225 43' Median lobe narrow, elongate; Sterna IV, V with punctures coalescent near lateral margins, forming shallow pits 44 44 (43') Anterior part of temporal lobe forming narrow, pollinose ridge; glabrous area of temporal lobe separated from antennal lobe by nearly length of antennal lobe; temporal setae one to three (in most specimens two); temporal lobe with up to three punctures near lateral margin or none O. duplex new species, p. 220 44' Anterior part of temporal lobe glabrous nearly to antennal lobe, separated from latter only by rather broad antennal groove; 1 temporal seta; temporal lobe with many punctures O. bouchardi new species, p. 221 45 (10') Inner, outer, or both pairs of pronotal carinae punctate 46 45' Pronotal carinae entirely impunctate 56 46 (45) Inner carina with three or more (in most species many) punctures 47 46' Inner carina with one or two punctures or none 49 47 (46) Head twice as long as wide; median lobe elongate, margins almost parallel O. nasalis new species, p. 228 47' Head slightly or not at all longer than wide; median lobe lance-shaped, margins not parallel 48 48 (47') Median lobe constricted at junction with clypeus; tip of median lobe subtruncate; Antennal Segments IV-IX impunctate; base of outer carina not Revision of Omoglymmius Ganglbauer 185 48 (47') Median lobe constricted at junction with clypeus; tip of median lobe subtruncate; Antennal Segments IV-IX impunctate; base of outer carina not sloped into basal impression; eye slightly reduced, about 0.33 of length of temporal lobe in profile view O. data new species, p. 217 48' Median lobe not constricted at junction with clypeus; tip of median lobe acute; all antennal segments impunctate; eye not reduced, about 0.5 of length of temporal lobe; base of outer carina sloped gradually into basal impression O. ephemeris new species, p. 244 49 (46') Intervals II, IV distinctly elevated anteriorly; head elongate anterior to eye; frontal grooves very narrow O. mycteroides new species, p. 237 49' Intervals II, IV not elevated; head not elongate; frontal grooves moderately broad 50 (49') Prosternum with precoxal carinae; temporal seta absent O. thoracicus new species, p. 216 50' Prosternum without precoxal carinae; one temporal seta 51 (50') Stria IV with five or six setae along its entire length 51' Stria IV with one or two setae near apex 52 (51) Outer carina scarcely narrowed at base O. coelebs new species, p. 215 52' Outer carina strongly narrowed to base; latter bluntly pointed 53 (52') Median lobe not constricted at junction with clypeus; tip of median lobe rounded; pronotum elongate O. malaicus (Arrow), p. 214 53' Median lobe constricted at junction with clypeus; tip of median lobe acute; pronotum more nearly quadrate 54 (53') Bases of outer carinae markedly divergent O. fraudulentus new species, p. 214 54' Bases of outer carinae scarcely divergent O. nemoralis new species, p. 213 55 (51') Antennal Segments I-XI with faint, irregular punctures; lateral pits of abdominal Sternum IV narrow, oblique, slit-like; temporal lobe, outer carina of pronotum finely punctate O. sectatus new species, p. 243 55' Antennal Segments I-IV punctate; Segments V-X impunctate; lateral pits of SternumlV shallow, round; temporal lobe, outer carina coarsely punctate O. evasus new species, p. 212 56 (45') Temporal lobe punctate; temporal seta one 56' Temporal lobe impunctate; temporal setae two to four 57 (56) Median lobe narrow, elongate; anterior femur of male with ventral tooth; hind calcar acute, angular 57' Median lobe broader, lance-shaped; anterior femur of male without ventral tooth; hind calcar prominent, apex rounded. O. gracilicornis (Grouvelle), p. 244 58 (57) Abdominal Sterna III-V with coarse punctures, these coalescent near lateral margin; Stria IV with one or two setae near apex. O. consors new species, p. 221 58' Abdominal Sterna III-V with finer, isolated punctures, not at all coalescent laterally; Stria IV with five setae along its length. . O. hiekei new species, p. 218 59 (56') Precoxal carinae absent; outer carina as broad as inner one at middle O. quadruplex new species, p. 219 59' Precoxal carina present; outer carina 0.66 as broad as inner carina at middle O. pectoralis new species, p. 218 50 51 52 55 53 54 57 59 58 Quaesi. Ent., 1982, 18 (1-4) 186 Bell and Bell 60 (9X) Outer carina of two planes meeting at sharp edge, one vertical, other sloped toward paramedian groove; pronotum hexagonal to subquadrate; Stria IV with five to seven setae along its length O. tabulatus new species, p. 238 60' Outer carina not formed of two planes; pronotum not hexagonal; Stria IV with one or two setae near apex 61 61 (600 Outer carina less than 0.4 as wide as inner carina at middle; Antennal Segments V-X punctate O. solitarius (Arrow), p. 208 61 ' Outer carina about 0.5 as wide as inner one at middle; antennal Segments V-X with punctures absent or indistinct 62 62 (6L) Pronotol carinae impunctate; temporal lobe with eight to 10 punctures near lateral margin O. impletus Bell and Bell, p. 231 62' Outer carina punctate; temporal lobe with 20-30 punctures O. patens new species, p. 242 63 (60 Suborbital tubercle present 64 63' Postorbital tubercle present 68 64 (63) Outer carina more than 0.6 as wide as inner carina at middle; marginal groove not dilated 65 64' Outer carina about 0.4 as wide as inner carina at middle; marginal groove dilated O. pulvinatus (Grouvelle), p. 252 65 (64) Outer carina broadest just anterior to base; base of inner carina strongly narrowed; inner carina impunctate O. sedlaceki new species, p. 252 65' Outer carina narrowed posteriorly; base of inner carina less narrowed; inner carina punctate 66 66 (650 Median lobe broad, tip subtruncate; pronotum nearly quadrate; metasternal punctures limited to midline, margins O. biroi new species, p. 251 66" Median lobe narrow, tip acute; pronotum not quadrate; metasternum entirely punctate 67 67 (660 Median lobe punctate; outer carina equal to or slightly narrower than inner carina at middle; one temporal seta O. cheesmanae (Arrow), p. 250 67' Median lobe impunctate; outer carina distinctly narrower than inner carina at middle; temporal seta absent O. asetatus new species, p. 250 68 (630 Postorbital tubercle visible only in lateral view 69 68' Postorbital tubercle visible in dorsal view 72 69 (68) Lateral margins of pronotum nearly parallel; median lobe broad, rounded; pollinosity of orbital groove extended to posterior margin of eye O. quadraticollis (Arrow), p. 223 69' Lateral margins more curved; pronotum not subquadrate; median lobe broad to narrow, obtuse to rounded; pollinosity of orbital groove less extensive 70 70 (690 Inner carina truncate at base; outer carina narrowed to base O. repetitus new species, p. 222 70' Inner carina pointed at base; outer carina dilated at base 71 71 (700 Median lobe broad, tip rounded; medial margin of base of outer carina sinuate; pollinose area of orbital groove broad; temporal lobe with scattered, very minute punctures; both pronotal carinae with very minute punctures O.follis new species, p. 245 7L Median lobe angulate at tip; medial margin of base of outer carina not sinuate; Revision of Omoglymmius Ganglbauer 187 pollinosity of orbital groove limited; temporal lobe with few minute punctures near lateral margin; pronotal carinae impunctate O. iridescens new species, p. 245 72 (680 Outer carina slightly narrower, more convex than inner one; carinae impunctate; anterior end of outer carina elevated, obliquely truncate O. planiceps new species, p. 248 72' Outer carina as wide and no more convex than inner one; anterior end of outer carina neither elevated nor obliquely truncate; carinae punctate or impunctate . 73 (720 Metasternum entirely punctate 73' Metasternum with punctures limited to midline, margins 74 (73) Inner, outer carinae with numerous fine punctures; outer carina narrowed to base; strial punctures round, pilose, coarse O. sus new species, p. 248 74' Outer carina punctate; inner carina impunctate; outer carina dilated at base; strial punctures fine, especially in Striae I-III 75 (740 Postorbital tubercles very large, divergent; outer carina with many fine punctures; medial angle of temporal lobe rounded O. capito (Grouvelle), p. 249 15' Postorbital tubercles smaller, scarcely divergent; outer carina with one or two fine punctures; medial angle obtuse O. lentus new species, p. 249 76 (730 Postorbital tubercle very large; apex of pronotum less narrowed; strial punctures elliptical, very fine O. auratus new species, p. 247 76' Postorbital tubercle small; pronotum more narrowed anteriorly; strial punctures round, moderately fine 77 (760 Median lobe broad, tip rounded; width of pronotum at middle subequal to width at base O. massa new species, p. 246 11' Median lobe narrower, tip obtusely rounded; width of pronotum at middle clearly less than width at base O. denticulatus new species, p. 247 73 74 76 75 77 KEY TO EURASIAN SPECIES (INCLUDES RYUKYU AND ANDAMAN ISLANDS) 1 Outer carina equal to or wider than inner carina at middle 2 1 ' Outer carina 0.25 as wide as inner carina at middle (Andaman Is.) O. solitarius (Arrow), p. 208 2 (1) Inner carina with anterior half distinctly narrower than posterior half; greatest width of inner carina distinctly less than that of outer carina (South India). . . O. malabaricus (Arrow), p. 207 2' Inner carina not distinctly narrower in anterior half; inner and outer carinae equal in width 3 3 (2') Median lobe constricted at base of clypeus; temporal seta absent 4 3' Median lobe not constricted at junction with clypeus; one temporal seta (Malay Peninsula) O. malaicus (Arrow), p. 214 4 (3) Orbital groove absent; prosternum with precoxal carinae; posteriolateral margin of temporal lobe rounded 5 4' Orbital groove represented by strip of pollinosity; prosternum without precoxal carinae; posteriolateral margin of temporal lobe sinuate (Europe, western Asia). O. germari (Ganglbauer), p. 194 Quaest. Ent., 1982, 18 (1-4) 188 Bell and Bell 5 (4) Medial angles of temporal lobes acute, contiguous; Stria IV with two to four setae along its length; total length 5. 1-5.2 mm (Ryukyu Is.) O. sakuraii (Nakane), p. 207 5' Medial angles of temporal lobe obtuse, slightly separated; Stria IV with 1-2 setae near apex; total length 6. 1-7.1 mm (Bhutan) O. laticeps Bell, p. 206 KEY TO SPECIES OF GREATER SUNDA AND PHILIPPINE ISLANDS 1 At least one pair of pronotal carinae punctate V Pronotal carinae entirely impunctate 2 (1) Inner carina with several to many punctures (only three or four in O. summissus) 2' Inner carina impunctate or with only one or two punctures on each carina 3 (2) Pronotum quadrate; outer carina widest near anterior margin, where flattened, angulate; outer carina only slightly narrower at base than at middle; eye large (Sumatra) O. summissus new species, p. 216 3' Pronotum not quadrate, its lateral margins markedly curved; outer carina distinctly narrowed to base, apex, widest at middle; eye reduced 4 (30 Base of Stria IV with small pollinose longitudinal scarp; pollinosity of orbital groove extended at least to middle of eye; depth of eye about 0.6 of depth of head; posterior margin of eye oblique, nearly straight (Philippines, island unknown) O. semperi new species, p. 217 4' Base of Stria IV without longitudinal scarp; orbital groove absent; depth of eye less than 0.5 of depth of head, its posterior margin curved (Luzon) O. data new species, p. 217 5 (2') Base of Stria IV with longitudinal pollinose scarp, or without scarp but with pollinose spot {O. politus ) 5' Base of Stria IV without longitudinal pollinose scarp and without pollinose spot 6 (50 Pronotum widest at middle or else with lateral margins almost parallel; median lobe of head narrow, its tip acute; base of Interval IV not elevated 6 ' Pronotum widest well anterior to middle; median lobe wide, margins almost parallel, tip rounded; base of Interval IV distinctly elevated 7 (6) Base of outer carina very markedly narrowed; median lobe very narrow, margins nearly parallel; punctures of abdominal Sterna IV and V each form transverse row (Negros) O. crassicornis new species, p. 21 1 7' Base of outer carina only moderately narrowed; median lobe broader; punctures of Sterna IV, V numerous, scattered 8 (7') Temporal lobe puncture over most of surface; median lobe narrow, tip acute; sides of pronotum nearly parallel (Luzon, Mindoro, Palawan, Siargao) O. philippensis (Chevrolat), p. 209 8" Temporal lobe punctate only near lateral margin; median lobe broader, tip more obtuse; pronotum widest at middle, sides curved 9 (8') Eye large, round; basal scarp of Stria IV distinct; male with ventral tooth on anterior femur (Luzon) O. imugani new species, p. 210 2 15 3 5 4 6 11 7 10 8 9 Revision of Omoglymmius Ganglbauer 189 9' Eye slightly reduced; basal scarp of Stria IV very small, scarcely more than slight elevation of lateral margin of pollinose spot; male without ventral tooth on anterior femur (Luzon) O. politus new species, p. 210 10 (6') Lateral margin of outer carina deeply sinuate anterior to hind angle (Sumatra) O. amplus new species, p. 21 1 10' Lateral margin scarcely sinuate anterior to hind angle (Mentawei) O. modiglianii new species, p. 212 1 1 (5") Prosternum with precoxal carinae (Java) O. thoracicus new species, p. 216 1 V Prosternum without precoxal carinae 12 12 (1 L) Outer carina not narrowed at base (Palawan). O. coelebs new species, p. 215 1 7! Outer carina narrowed at base 13 13 (120 Stria IV with one seta near apex, none or one at base (Mindanao) O. evasus new species, p. 212 1 3' Outer carina with four or five setae, forming continuous row 14 14 (130 Bases of outer carinae strongly divergent (Sumatra) O. fraudulentus new species, p. 214 14' Bases of outer carinae not divergent (Sarawak) O. nemoralis new species, p. 213 15 (10 Basal transverse scarp of elytron with continuous pollinose area medial to base of Stria IV 1 6 15' Basal transverse scarp without continuous pollinose area medial to base of Stria IV 17 16(15) One temporal seta; median lobe with sides parallel, tip truncate (Sumatra, Borneo) O. consors new species, p. 221 16' Two to four temporal setae; median lobe lance-shaped, tip acute (Mindanao) . O. quadruplex new species, p. 219 17 (15') Base of Stria IV with longitudinal pollinose scarp 18 17' Base of Stria IV without scarp 19 18 (17) Temporal lobe with many punctures; one temporal seta; temporal lobe glabrous anteriorly, separated from antennal lobe only by pollinose antennal groove (Sumatra) O. bouchardi new species, p. 221 18' Temporal lobe with one or two punctures near lateral margin; in most specimens, two temporal setae (in a few, one or three); temporal lobe connected to antennal lobe by pollinose longitudinal ridge (Luzon, Palawan, Sibuyan, Mindanao) O. duplex new species, p. 220 19 (17') Prosternum with precoxal carinae (Java) O. pectoralis new species, p. 218 19' Prosternum without precoxal carinae (Luzon) O. hiekei new species, p. 218 KEY TO SPECIES OF “WALLACEA” (CELEBES, MOLUCCAS, LESSER SUNDAS, KEI AND SCHOUTEN ISLANDS) 1 Fifth interval elevated, carinate (Yapan). O. bicarinatus new species, p. 230 V Fifth interval not elevated nor carinate 2 2 (L) Outer carina of pronotum much narrower than inner carina; outer carina with one to three setae inserted in conspicuous punctures (Sumbawa) Quaest. EnU, 1982, 18 (1-4) 190 Bell and Bell O. bucculatus (Arrow), p. 229 2 ' Pronotal carinae of nearly equal width at middle; outer carina without setae 3 3 (20 Head about twice as long as wide; median lobe elongate; its margins parallel (Buru) * O. nasalis new species, p.228 3' Head at most slightly longer than wide 4 4 (3') Postorbital tubercle present 5 4' Postorbital tubercle absent 6 5 (4) Pronotum quadrate, its margins nearly straight and parallel; median lobe broad, its apex obtuse; orbital groove extensively pollinose (Tanimbar) O. quadraticollis (Arrow), p. 223 5' Pronotum not quadrate, its margins rounded; median lobe narrower, its apex acute; orbital groove with pollinosity restricted to anterior end (Celebes) O. repetitus new species, p. 222 6 (40 Lateral margin of temporal lobe almost straight; anteriomedial margin of temporal lobe abruptly bent; frontal space very broad (Morotai) O. morditus new species, p. 228 6' Lateral margin of temporal lobe rounded; anteriomedial margin bent or rounded; frontal space narrower 7 7 (60 Abdominal sterna with extensive lateral pollinose area, but without distinct punctures (Amboina) O. vadosus new species, p. 227 1' Abdominal sterna with distinct punctures, in some specimens coalescent laterally 8 8 (70 Anterior portion of temporal lobe extensively pollinose, glabrous portion of temporal lobe broadly separated from antennal lobe 9 8' Anterior part of temporal lobe not pollinose; glabrous portion of temporal lobe separated from antennal lobe only by narrow antennal groove 10 9 (8) Pronotum subquadrate; one to three temporal setae; medial angle of temporal lobe rounded; Segment I of antenna with apex inflated laterally (Mangole) O. continuus new species, p. 226 9 ' Pronotum not subquadrate, its base distinctly narrowed; one temporal seta; medial angle of temporal lobe obtuse; Segment I of antenna with apex not inflated (Mangole) O. wittmeri new species, p. 227 10 (8') Pronotum with sides strongly curved; temporal lobe with 25-30 rather coarse punctures; outer carina with 35-40 punctures (Kei Is.) O. viduus new species, p. 226 10' Pronotum subquadrate, its margins only slightly curved; temporal lobe with 10-15 punctures; outer carina with 10 or fewer punctures 11 11 (100 Frontal space 0.33 of width of temporal lobe; eye relatively convex (Dammar) O. opticus new species, p. 225 1 Y Frontal space wider, about 0.43 of width of temporal lobe; eye less convex 12 12 (1 10 Pronotum nearly quadrate, scarcely narrowed posteriorly (Batjan) O. batchianus (Arrow), p. 223 12' Pronotum distinctly narrowed posteriorly, and rather strongly so anteriorly (Ternate, Halmahera) O. humeralis (Grouvelle), p. 224 Revision of Omoglymmius Ganglbauer 191 KEY TO SPECIES FROM SOLOMON ISLANDS 1 Outer carina much narrower than inner one, its dorsal surface flat, meeting lateral surface at sharp angle; Stria IV with four or five setae (Bougainville, New Georgia) O. tabulatus new species, p. 238 Y Outer carina subequal to inner carina at middle; outer carina more or less convex, its dorsal surface not in form of lateral margin with lateral face 2 2 (L) Head elongate, about twice as long as wide; median lobe elongate, narrow, its sides nearly parallel; elytron without longitudinal scarp at base of Stria IV (New Georgia, Guadalcanal) O. mycteroides new species, p. 237 2' Head not elongate, at most slightly longer than wide; median lobe broader, its sides not parallel; elytron with short longitudinal scarp at base of Stria IV 3 3 (2') Pronotum with sides strongly curved, conspicuously wider at middle than at base or apex; lateral margin sinuate anterior to hind angle 4 3' Pronotum with sides only slightly curved, nearly parallel (pronotum subquadrate); lateral margin not sinuate anterior to angle 5 4 (3) Inner carina punctate; pronotum more expanded at middle, more constricted to base and apex (Bougainville) O. princeps new species, p. 236 4' Inner carina impunctate; pronotum less expanded at middle (New Georgia) . O. renutus new species, p. 236 5 (3') Pollinosity of orbital groove complete to hind margin of eye, conspicuous; elytral striae with continuous pollinosity (Bougainville, Choiseul, Faro) O. gurney i new species, p. 235 5' Pollinosity of orbital groove incomplete; elytral striae with pollinosity confined to punctures 6 6 (50 Pronotum elongate, its sides nearly parallel; temporal lobe with 30-35 punctures; medial angle of temporal lobe broadly rounded; inner carina with 15-18 punctures; total length larger, about 7.3 mm (Santa Cruz) O. scopulinus new species, p. 237 6' Pronotum not elongate; temporal lobe with less than 30 punctures; medial angle of temporal lobe produced, oblique; inner carina with two to eight punctures 7 7 (6') Outer carina as broad as inner carina at middle; base of outer carina only slightly narrower than its greatest width, latter slightly greater than width of inner carina at base; medial margin of outer carina sinuate near base 8 7" Outer carina slightly narrower than inner carina at middle; base of outer carina more narrowed, its width equal to or slightly less than width of inner carina at base; medial margin of outer carina not sinuate near base 10 8 (7) Anteriomedial margin of temporal lobe curved gradually; frontal space more V-shaped 9 8' Medial margin of temporal lobe bent abruptly near middle, frontal space more U-shaped (Savo,Ngela) O. modicus new species, p. 233 9 (8) Posteriomedial margin of temporal lobe distinctly emarginate between occipital and medial angles; latter produced, closer together; margins of pronotum strongly curved; lateral pits of Sternum IV in female relatively shallow and indefinite (Malaita) O. manni new species, p. 234 9' Posteriomedial margin scarcely emarginate; medial angles not produced, more Quaest. Ent., 1982, 18 (1-4) 192 Bell and Bell separated from one another; margins of pronotum less curved: lateral pits of Sternum IV in female deeper, rounder (Isabella) . . O. regius new species, p. 235 10 (7') Pronotum subquadrate, lateral margins nearly parallel; base of pronotum scarcely narrowed (Guadalcanal) O. lindrothi new species, p. 232 10' Pronotum with lateral margins strongly curved; base of pronotum strongly narrowed (Russell Is., San Cristobal) O. rusticus new species, p. 234 KEY TO SPECIES FROM NEW GUINEA AND ADMIRALTY ISLANDS 1 Neither postorbital nor suborbital tubercles present 2 V Either postorbital or suborbital tubercle present 12 2 (1) Frontal space elongate, narrow, V-shaped or slightly U-shaped; antennal groove narrow 3 2' Frontal space broad, strongly U-shaped; antennal groove broad 8 3 (2) Base of Stria IV with longitudinal pollinose scarp 4 3' Base of Stria IV without longitudinal pollinose scarp 6 4 (3) Median lobe narrow, tip acute; pollinosity very limited at lateral end of antennal groove 5 4' Median lobe broader, tip obtuse; pollinosity expanded at lateral end of antennal groove O. trepidus new species, p. 242 5 (4) Outer carina 0.5 as wide as inner carina at middle; lateral pit of Sternum IV of female rounded O. patens new species, p. 242 5' Outer carina as wide as inner carina at middle; lateral pit of Sternum IV of female triangular, broad O. cavea new species, p. 243 6 (3') Pronotal carinae impunctate; Stria IV with two to four setae; male without ventral tooth on anterior femur; hind calcar of male large, obtuse; total length 4.5 mm O. gracilicornis (Grouvelle), p. 244 6' Pronotal carinae punctate; Stria IV with one seta near apex; male with ventral tooth on anterior femur; hind calcar acute; most specimens longer than 4.5 mm .7 7 (6') Apex of median lobe acute; median lobe with seven to 10 punctures; lateral pits of abdominal Sternum IV rounded; middle calcar of male elongate O. ephemeris new species, p. 244 T Apex of median lobe obtuse; median lobe with one puncture or none; lateral pits of Sternum IV narrow, oblique; middle calcar of male very small O. sectatus new species, p. 243 8 (27) Pronotum subquadrate, margins only slightly curved 9 8' Pronotum not subquadrate, margin strongly curved O. puncticornis new species, p. 241 9 (8) Outer carina narrowed to base; margin not sinuate anterior to hind angle; length 4. 6-6. 2 mm 10 9' Outer carina narrowed to subbasal constriction, posterior to which it is broadened; margin sinuate anterior to hind angle; length 6.0-7. 2 mm 11 10 (9) Median lobe with a few scattered punctures (New Guinea) O. vicinus (Grouvelle), p. 239 10' Median lobe impunctate (Admiralty Is.) O. classicus new species, p. 238 Revision of Omoglymmius Ganglbauer 193 1 1 (90 Frontal space broad-U-shaped; posterior margin of temporal lobe nearly transverse O. fr ingill us new species, p. 240 11' Frontal space more elongate, heart-shaped; posterior margin of temporal lobe more evenly rounded O. oroensis new species, p. 240 12 (F) Suborbital tubercle present 13 12' Postorbital tubercle present 17 13 (12) Outer carina more than 0.6 as wide as inner carina at middle; marginal groove not dilated 14 13' Outer carina about 0.4 as wide as inner carina at middle; marginal groove dilated O. pulvinatus (Grouvelle), p. 252 14(13) Outer carina broadest just anterior to base; base of inner carina strongly narrowed; inner carina impunctate O. sedlaceki new species, p.252 14' Outer carina narrowed posteriorly; base of inner carina less narrowed; inner carina punctate 1 5 15 (14x) Median lobe broad, tip subtruncate; pronotum subquadrate, lateral margins only slightly curved; posterior margin of temporal lobe nearly transverse; punctures of metasternum restricted to midline, lateral margins. O. biroi new species, p. 25 1 15' Median lobe narrow, tip acute to subacute; pronotum not subquadrate, lateral margins strongly curved; entire disc of metasternum punctate 16 16 (15') Median lobe punctate; outer carina equal to or slightly narrower than inner carina at middle; one temporal seta O. cheesmanae (Arrow), p. 250 16' Median lobe impunctate; outer carina distinctly narrower than inner carina at middle; temporal seta absent O. asetatus new species, p. 250 17 (12') Postorbital tubercle visible only in lateral view, hidden in dorsal view 18 17' Postorbital tubercle visible in dorsal as well as lateral view 19 18 (17) Median lobe broad, tip obtuse; medial margin of base of outer carina sinuate; antennal groove extended laterally into broad pollinose area; temporal lobe, pronotal carinae with scattered fine punctures . . O.follis new species, p. 245 18' Median lobe narrow, lance-shaped, tip acute; medial margin of outer carina not sinuate near base; antennal groove extended laterally into short, narrow pollinose area; temporal lobe with very few punctures; pronotal carinae impunctate O. iridescens new species, p. 245 19 (17') Metasternum with punctures limited to midline, margins 20 19' Metasternum with punctures scattered over entire disc 22 20 (19) Postorbital tubercle larger; anterior margin of outer carina abruptly truncate; punctures of elytral striae very fine, elliptical over most of disc, obsolete near base O. auratus new species, p. 247 20' Postorbital tubercle smaller, anterior margin of outer carina evenly sloped; strial punctures coarse, round on most of disc, becoming fine, elliptical near base 21 21 (20') Median lobe broad, tip rounded; postorbital tubercle of moderate size; medial angles of temporal lobes sharper, more produced O. massa new species, p. 246 21' Median lobe narrow, tip obtuse; postorbital tubercle very small; median angles obtuse, scarcely produced O. denticulatus new species, p. 247 22 (19') Outer carina more convex than, slightly narrower than inner one at middle Quaest. Ent., 1982, 18 (1-4) 194 Bell and Bell O. planiceps new species, p. 248 22' Outer carina equal in convexity, width to inner one at middle 23 23 (22') Both pairs of carinae punctate; strial punctures round, coarse, pollinose O. sus new species, p. 248 23' Inner carina impunctate; outer carina with few scattered punctures; strial punctures shallow, those of Striae I-III finer than those of outer striae 24 24 (230 Postorbital tubercles relatively small, not divergent posteriorly; medial angle of temporal lobe slightly produced; posteriomedial margin slightly emarginate between medial, occipital angles O. lentus new species, p.249 24' Postorbital tubercles large, divergent posteriorly; me angle of temporal lobe rounded; posteriomedial margin not emarginate O. capito (Grouvelle), p. 249 KEY TO SPECIES FROM THE CAROLINE ISLANDS 1 Outer carina of pronotum punctate; inner carina narrowest anterior to base, broadened from there to base 2 T Outer carina impunctate; inner carina tapered to point at base (Yap) O. impletus Bel! and Bell, p. 231 2 (1) Temporal lobe with margin sinuate between medial angle and occipital angle (Palau) O. caelatus Bell and Bell, p. 230 2' Temporal lobe with margin evenly convex between medial and occipital angles (Kusiae, Ponape, Palau) O. oceanicus Bell and Bell, p. 231 Omoglymmius (sens u stricto ) germari (Ganglbauer) Figs. 113, 121 Rhysodes exaratus Erichson 1847: 301 (nee Dalman 1823; nee Serville 1825) Rhysodes aratus Chevrolat 1873a: 209 (nee Newman 1838) Rhysodes americanus Reitter 1882: 140 (nee Castelnau 1836) Rhysodes germari Ganglbauer 1892: 534 Omoglymmius (sensu stricto ) german (Ganglbauer) Bell and Bell 1978 Type material. - None designated. NEOTYPE (here designated) female, labelled: “Majevica, Bosna, CNHM 1955, Karl Branczik colln., ex Eduard Knirsch” (in Yugoslavia)(CNHM). PARANEOTYPES three males, four females, labelled: “Majevica, Bosna, VI. Zoufal” (MNHB); one male, labelled: ’’Majevica, Bosna (Bosna = Bosnia), VI Zoufal, coll. G. Falkenstrom’’ (LUN). Description. — Length 6. 8-8.0 mm. Antennal segments impunctate; median lobe impunctate, rhomboid, its apex subacute; frontal space U-shaped, anteriomedial margin of temporal lobe abruptly curved; medial angles obtuse, slightly separated; posteriomedial margin oblique; occipital angle obtuse, distinct; posteriolateral margin oblique, parallel to posterior half of anteriomedial margin; posteriolateral margin slightly sinuate just posterior to eye; orbital groove pollinose, extended to level of hind margin of eye; temporal lobe with five to 12 coarse punctures, restricted to lateral portion; temporal seta absent; small postorbital tubercle present, or this tubercle merely suggested or entirely absent. Pronotum moderately elongate, length/greatest width 1.29; widest near middle, apex markedly narrowed; base nearly equal to greatest width; lateral margin with slight sinuation anterior to hind angle; outer carina slightly narrower than inner one at middle; medial margin of outer carina slightly sinuate anterior to base; base of outer carina narrowed; inner carinae narrowed at base; prpnotal carinae finely punctate; punctures more numerous on outer carina; inner carina with two to 10 punctures; pronotum without setae; prosternum without precoxal carinae. Revision of Omoglymmius Ganglbauer 195 Plate 9. Figures 1 13-129, Subgenus Omoglymmius sensu stricto. Figs. 1 13-120, Head and pronotum, dorsal aspect; Fig. 1 13, Omoglymmius (s. str.) germari (Ganglbauer); Fig. 114, O. (s. str.) laticeps Bell ; Fig. 1 15, O. (s. str.} sakuraii (Nakane); Fig. 116, O. (s. str.) malabaricus (Arrow); Fig. 117, O. (s. str.) solitarius (Arrow); Fig. 1 18, O. (s. str.) philippensis (Chevrolat); Fig. 1 19, O. (s. str.) imugani new species; Fig. 120, O. (s. str.) politus new species; Figs. 121-128, Head, lateral aspect; Fig. 121, O. (s. str.) germari (Ganglbauer); Fig. 122, O. (s. str.) laticeps Bell; Fig. 123, O. (s. str.) sakuraii (Nakane); Fig. 124, O. (s. str.) malabaricus (Arrow); Fig. 125, O. (s. str.) solitarius (Arrow); Fig. 126, O. (s. str.) philippensis (Chevrolat); Fig. 127, O. (s. str.) imugani new species; Fig. 128, O. (s. str.) politus new species; Fig. 129, Prothorax, ventrolateral aspect, O. (s. str.) sakuraii (Nakane). Quaest. Ent., 1982, 18 (1-4) 196 Bell and Bell Plate 10. Figures 130-139, Subgenus Omoglymmius sensu stricto. Figs. 130-138, Head and pronotum, dorsal aspect; Fig. 130, Omoglymmius (s. str.) crassicornis new species; Fig. 131, O. (s. str.) amplus new species; Fig. 132, O. (s. str.) modiglianii new species; Fig. 133, O. (s. str.) malaicus (Arrow); Fig. 134, O. (s. str.) evasus new species; Fig. 135, O. (s. str.) nemoralis new species; Fig. 136, O. (s. str.) fraudulentus new species; Fig. 137, O. (s. str.) coelebs new species; Fig. 138, O. (s. str.) thoracicus new species; Fig. 139, Prothorax, ventral aspect, O. (s. str.) thoracicus new species. Revision of Omoglymmius Ganglbauer 197 Plate 1 1. Figures 140-149, Subgenus Omoglymmius sensu stricto. Figs. 140-146, Head and pronotum, dorsal aspect; Fig. 140, Omoglymmius (s. str.) summissus new species; Fig. 141, O. (s. str.) semperi new species; Fig. 142, O. (s. str.) data new species; Fig. 143, O. (s. str.) hiekei new species; Fig. 144, O. (s. str.) pectoralis new species; Fig. 145, O. (s. str.) quadruplex new species; Fig. 146, O. (s. str.) duplex new species; Figs. 147-148, Head, lateral aspect; Fig, 147, O. (s. str.) semperi new species; Fig. 148, O. (s. str.) data new species; Fig. 149, Prothorax, ventral aspect, O. (s. str.) pectoralis new species; Quaest. Ent., 1982, 18 (1-4) 198 Bell and Bell Plate 12. Figures 150-158, Subgenus Omoglymmius sensu stricto. Figs. 150-151, Head, pronotum and anterior portion of; elytra, dorsal aspect; Fig. 150, Omoglymmius (s. str.) bouchardi new species; Fig. 151, O. ( s . str.) consors new species; Figs. 152-155, Head and pronotum, dorsal aspect; Fig. 152, O. (s. str.) quadraticollis (Arrow); Fig. 153, O. fs. str.) batchianus (Arrow); Fig. 154, O. (s. str.) humeralis (Grouvelle); Fig. 155, O. fs. str.) repetitus new species; Figs. 156-158, Head, lateral aspect; Fig. 156, O. (s. str.) repetitus new species; Fig. 157, O. f s . str.) quadraticollis (Arrow); Fig. 158, O. fs. str.) batchianus (Arrow). Revision of Omoglymmius Ganglbauer 199 Plate 13. Figures 159-171, Subgenus Omoglymmius sensu stricto. Figs. 159-165, Head and pronotum, dorsal aspect; Fig. 159, Omoglymmius (s. str.) opticus new species; Fig. 160, O. (s. str.) viduus new species; Fig. 161, O. (s. str.) continuus new species; Fig. 162, O. (s. str.) wittmeri new species; Fig. 163, O. (s. str.) vadosus new species; Fig. 164, O. (s. str.) nasalis new species; Fig. 165, O. (s. str.) morditus new species; Figs. 166-167, Head, lateral aspect; Fig. 166, O. (s. str.) continuus new species; Fig. 167, O. (s. str.) wittmeri new species; Figs. 168-169, Antennal Segments I, II; Fig. 168, O. (s. str.) wittmeri new species; Fig. 169, O. (s. str.) continuus new species; Figs. 170-171, Hind tibia, male, apical portion; Fig. 170, O. (s. str.) continuus new species; Fig. 171, O. (s. str.) wittmeri new species; Quaest. Ent., 1982, 18 (1-4) 200 Bell and Bell Plate 14. Figures 172-181, Subgenus Omoglymmius sensu stricto. Figs. 172-179, Head and pronotum, dorsal aspect; Fig. 172, Omoglymmius (s. str.) bucculatus (Arrow); Fig. 173, O. (s. str.) caelatus Bell & Bell; Fig. 174, O. (s. str.) impletus Bell & Bell; Fig. 175, O. (s. str.) oceanicus Bell & Bell; Fig. 176, O. (5. str.) lindrothi new species; Fig. 177, O. (s. str.) modicus new species; Fig. 178, O. (s. str.) rusticus new species; Fig. 179, O. (s. str.) manni new species; Fig. 180, Hind tibia, male, apical portion, O. (s. str.) manni new species; Fig. 181, Head, pronotum and anterior portion of elytra, O. (s. str.) bicarinatus new species. Revision of Omoglymmius Ganglbauer 201 Plate 15. Figures 182-191, Subgenus Omoglymmius sensu stricto. Figs. 182-188, Head and pronotum, dorsal aspect; Fig. 182, Omoglymmius (s. str.) regius new species; Fig. 183, O. (s. str.) gurneyi new species; Fig. 184, O. (s. str.) princeps new species; Fig. 185, O. (s. str.) renutus new species; Fig. 186, O. (s. str.) scopulinus new species; Fig. 187, O. (s. str.) mycteroides new species; Fig. 188, O. (s. str.) tabulatus new species; Fig. 189, Head, lateral aspect, O. (s. str.) gurneyi ; new species; Figs. 190-191, Hind tibia, male, apical portion; Fig. 190, O. (s. str.) regius new species; Fig. 191, O. (s. str.) tabulatus new species; Quaest. Ent., 1982, 18 (1-4) 202 Bell and Bell Plate 16. Figures 192-201, Subgenus Omoglymmius sensu stricto. Figs. 192-200, Flead and pronotum, dorsal aspect; Fig. 192, Omoglymmius (s. str.) classicus new species; Fig. 193, O. (s. str.) vicinus (Grouvelle); Fig. 194, O. (s. str.) oroensis new species; Fig. 195, O. (s. str.) fringillus new species; Fig. 196, O. (s. str.) puncticornis new species; Fig. 197, O. (s. str.) trepidus new species; Fig. 198, O. (s. str.) patens new species; Fig. 199, O. (s. str.) cavea new species; Fig. 200, O. (s. str.) sectatus new species; Fig. 201, Middle tibia, male, apical portion, O. (s. str.) sectatus new species. Revision of Omoglymmius Ganglbauer 203 Plate 17. Figures 202-215, Subgenus Omoglymmius sensu stricto. Figs. 202-208, Head and pronotum, dorsal aspect; Fig. 202, Omoglymmius (s. sir.) ephemeris new species; Fig. 203, O. (s. str.) gracilicornis (Grouvelle); Fig. 204, O. (s. str.) follis new species; Fig. 205, O. (s. str.) iridescens new species; Fig. 206, O. (s. str.) massa new species; Fig. 207, O. (s. str.) denticulatus new species; Fig. 208, O. (s. str.) auratus new species; Fig. 209, Middle tibia, male, apical portion; O. (s. str.) ephemeris new species; Fig. 210, Hind tibia, male, apical portion, O. (s. str.) gracilicornis (Grouvelle); Figs. 211-215, Head, lateral aspect; Fig. 211, O. (s. str.) iridescens new species; Fig. 212, O. (s. str.) follis new species; Fig. 213, O. (s. str.) massa new species; Fig. 214, O. (s. str.) denticulatus new species ; Fig. 215, O. (s. str.) auratus new species; Quaest. Ent., 1982, 18 (1-4) 204 Bell and Bell Plate 18. Figures 216-227, Subgenus Omoglymmius sensu stricto. Figs. 216-221, Head and pronotum, dorsal aspect; Fig. 216, Omoglymmius (s. str.) sus new species; Fig. 217, O. (s. str.) planiceps new species; Fig. 218, O. (s. str.) lentus new species; Fig. 219, O. (s. str.) capito (Grouvelle); Fig. 220, O. (s. str.) cheesmanae (Arrow); Fig. 221, O. (s. str.) asetatus new species; Figs. 222-227, Head, lateral aspect; Fig 222, O. (s. str.) sus new species; Fig. 223, O. (s. str.) planiceps new species; Fig. 224, O. (s. str.) lentus new species; Fig. 225, O. (s. str.) capito (Grouvelle); Fig. 226, O. (5. str.) cheesmanae (Arrow); Fig. 227 , O. (s. str.) asetatus new species; Revision of Omoglymmius Ganglbauer 205 Plate 19. Figures 228-237, Subgenus Omoglymmius sensu stricto. Figs. 228-233, Head and pronotum, dorsal aspect; Fig. 228, Omoglymmius (s. str.) biroi new species; Fig. 229, O. (s. str.) pulvinatus (Grouvelle); Figs. 230-233, O. (s. str.) sedlaceki new species; Fig. 230, Mt. Kaindi form A; Fig. 231, Kamira form C; Fig. 232, Kassem form B; Fig. 233, Sepalakambang form E; Figs. 234-236, Head, lateral aspect; Fig. 234, O. (s. str.) biroi new species; Fig. 235, O. (s. str.) pulvinatus (Grouvelle); Fig. 236, O. (s. str.) sedlaceki new species; Fig. 237, Head, dorsal aspect, O. (s. str.) sedlaeeki\ new species, Mt. Otto form D. Quaest. Ent., 1982, 18 (1-4) 206 Bell and Bell Elytron rather long, narrow for subgenus; elytral striae impressed, coarsely punctate; base of Stria IV with very small glabrous longitudinal scarp; Stria IV with one seta near apex; Stria VII with several near apex; metasternum with disc entirely coarsely punctate; abdominal sterna with punctures coarse, scattered; lateral pit of female deep, rounded; anterior femur of male with ventral tooth; middle calcar of male very small; hind calcar of male large, obtuse, its apex slightly lobate. The strongly oblique posteriolateral margin of the temporal lobe separates this species from all other members of the subgenus. The temporal lobes of O. (Boreoglymmius) lewisi (Nakane) are similarly oblique, but the latter species differs in presence of the basal setae on the antennae, a much larger postorbital tubercle, and in having the base of the pronotum distinctly narrowed. Early authors repeatedly confused O. germari with the North American O. (Boreoglymmius) americanus (Castelnau). The latter species differs in the presence of basal setae, in having the temporal lobes rounded, and in having the hind calcar of the male acute. Range. - Widely distributed in Europe, in and south of the Alps, and perhaps isolated, in the Caucasus and in the Kopet Dagh Mountains of northeastern Iran. Like Rhysodes sulcatus, an “Urwaldrelikte”, now restricted to small areas of primeval forest. Dajoz (1975) lists localities and gives a bibliography of published records. In addition to these records, we have examined specimens from the following localities: FRANCE: one male, Monguillon, Basses Alpes (SATO); IRAN: one female, Astrabad, Hauser coll. (MNHB); two males, one female, Kopet Dagh, coll. Hauser (CNHM); ITALY: two males, Lazio, Bassano di Sutri, coll. A. Fiori, P. Luigioni (MNHB); U.S.S.R.: (AZERBAIDZHAN S.S.R.) one male, Lenkoran, Tangyarud, coll. Kirichenko (LEN); one female, Hamarat, Caspian Sea area, coll. Leder (UVM); (GRUZIAN S.S.R.) one female, Batum, coll. G. Lewis (BMNH), two females, same locality (MNHN); one male, Lagodekhi Nature Reserve, coll. Kirekchuk (UVM), one male, one female, same locality (LEN); seven males, four females, Talyche (MNHB); CAUCASUS (unlocated); two females, Kadi-Keri (MNHB). The specimens listed for UVM were obtained by exchange from the Academy of Sciences, Leningrad. Variation. - This species is unusually varied in the development of the postorbital tubercles. This variation is in part geographical. Almost all specimens from west of the Black Sea have the tubercle relatively markedly developed. The single exception is the specimen from Monguillon, France. Most specimens from the Caucasus and eastern Iran have the tubercle very slightly developed or absent, though a few have it as well developed as in European specimens. It is possible that more than one species is involved, but further study is necessary. Bionomics. - Dajoz (1975) summarizes available information as follows (quoted, in translation): “found in Slovakia in the rotten wood of a beech ( Fagus silvatica L.) invaded by ants of the genus Camponotus, but this species is not a myrmecophile. In Bulgaria, in the red bark (?) of an old Oak ( Quercus sp.), larvae and soft imagos in September. In the Gargano Mountains, found in dead beech wood with its larva. In Greece, at Mount Ossa, at about 1400 meters, in July, we (Dajoz) have found them under the bark of dead beeches in the company of larvae of Cerambycidae, Buprestidae, and diverse Clavicornia such as Dechomus sulcicollis Germar and Uleiota planata Fab.” The specimen from Lagodekhi, U.S.S.R. is labelled as being from beech forest, in old dead trunk of beech. Omoglymmius (sensu stricto) laticeps (Bell) Figs, 114, 122 Omoglymmius laticeps Bell 1977: 157-158. Omoglymmius (sensu stricto) laticeps (Bell) Bell and Bell 1978. Type material. - HOLOTYPE female, labelled: “BHUTAN: Nobding, 41 km. E of Wangdi Phodrang, 2800 m. Basel Natl. Hist. Exped., 1972” (BSL). PARATYPES two females, labelled: “BHUTAN: Dorju La, Wangdo Phodrang, 1300 m.” (BSL); one female, Revision of Omoglymmius Ganglbauer 207 labelled: “BHUTAN: Tango, 22 km. N of Thimphu” (BSL). Description. — Length 6.1 -7.1 mm. Antennal segments impunctate; head relatively short, broad; median lobe short, broad, obtusely pointed at tip; frontal space short, narrow, nearly V-shaped, its margins only shallowly sinuate; medial angles rounded, nearly contiguous; posteriomedial margin rounded evenly into posteriolateral margin; orbital groove replaced by row of punctures, extended nearly to posterior margin; temporal lobe otherwise without punctures; temporal setae absent; postorbital tubercle minute; in profile view appearing as a slight convexity. Pronotum with length/greatest width 1.26; widest near middle, apex and base markedly narrowed, lateral margins markedly curved; lateral margin scarcely sinuate anterior to hind angle; carinae subequal at middle; medial margin of outer carina shallowly sinuate anterior to base; base of outer carina markedly narrowed; inner carina slightly narrowed to base; inner carina with two or three fine punctures near base; carinae otherwise impunctate except for extremely minute punctures; median groove coarsely, sparsely punctate; paramedian groove narrow, with two or three coarse punctures; pronotum without setae; prosternum with fine precoxal carinae. Elytron relatively long, narrow; striae impressed, coarsely punctate; base of Stria IV without scarp; Stria IV with one or two setae near apex; subapical stride with one seta; Stria VII with five to seven setae near apex; metasternum with very coarse punctures concentrated in midline and near lateral margins; female with lateral pit of abdominal sternum IV deep, round; female with minute ventral tooth on anterior femur; male unknown. This species, the only Omoglymmius s. str. known from the Himalaya, is very similar to O. sakuraii Nakane of the Ryukyu Islands. The two species are similar by combination of a precoxal carina, a reduced frontal space, very reduced punctation of the pronotal carinae, and a group of punctures replacing the orbital groove. O. sakuraii differs in having acute medial angles and in having more setae on Stria IV. Omoglymmius (sensu stricto) sakuraii (Nakane) Figs. 115, 123, 129 Rhysodes (Omoglymmius) sakuraii Nakane 1973: 5 (description in Japanese). Omoglymmius sakuraii Nakane 1978: 130-131 (redescription in English). Omoglymmius ( sensu stricto) sakuraii (Nakane) Bell and Bell 1978. Type material. - HOLOTYPE female, Hatsuno, Amami-Oshima, Japan, 30-III- 1 965, S. Sakurai lgt. (KU). We have not seen the type, but have studied two males from the type locality (SATO). Description. — Length 5. 1-5.2 mm. Antennal segments impunctate; head relatively short, broad; median lobe short, broad, its tip obtuse to subtruncate; frontal space short, narrow, nearly V-shaped, its margin only shallowly sinuate; medial angles acute, contiguous; posteriomedial angle rounded, in form of very obtuse angle with posteriolateral margin; orbital groove absent; one to four coarse punctures medial to eye; temporal setae absent, postorbital tubercle minute. Pronotum longer and narrower than in O. laticeps, length/greatest width 1.43 widest near middle; sides subparallel except near narrowed base and apex; lateral margin not sinuate anterior to hind angle; carinae subequal at middle; medial margin of outer carina sinuate anterior to base; base of outer carina narrowed; inner carina narrowed slightly to base; both pairs of carinae impunctate; median and paramedian grooves narrow, punctured as in O. laticeps ; pronotum without setae; prosternum with fine precoxal carinae (Fig. 129). Elytron relatively long, narrow; striae impressed, coarsely punctate; base of Stria IV without trace of scarp; Stria IV with two to four setae, most specimens with one seta anterior to middle; apical striole without seta; Stria VII with four setae near apex; metasternum with very coarse punctures concentrated in midline and near lateral margin; abdominal sterna with punctures very coarse, scattered; female with deep round lateral pit in Sternum IV; male with shallow pit in this position; male with ventral tooth on anterior femur (condition of anterior femur of female not mentioned in original description); middle calcar of male minute; hind calcar of male small, triangular, obtuse. This species is very similar to O. laticeps , but differs in having the pronotum narrower and more parallel-sided, and the medial angles acute and contiguous. Omoglymmius (sensu stricto) malabaricus (Arrow) Figs. 116, 124 Rhysodes malabaricus Arrow 1901: 86-87. Quaest. Ent., 1982, 18 (1-4) 208 Bell and Bell Omoglymmius (sensu stricto) malabaricus (Arrow) Bell and Bell 1978. Type material. - LECTOTYPE (here designated) male, labelled: “S. INDIA, Malabar, Pascoe collection” (BMNH). PARALECTOTYPE one specimen, sex not recorded, same data as lectotype (BMNH). Description. — Length 6. 5-7.0 mm. Antennal segments impunctate; head relatively short, broad; median lobe short, broad, obtusely pointed at apex; frontal space short, broad U-shaped, its lateral margin bent abruptly; temporal lobe continued anteriorly as oblique, glabrous carina; medial angles rounded, nearly contiguous; posteriomedial margin evenly rounded into posteriolateral margin; orbital groove fine, obsolescent; temporal lobe with eight to ten coarse punctures near posterior margin; temporal setae absent; postorbital tubercle minute. Pronotum with length/greatest width 1.38; widest near middle; apex and base strongly narrowed; lateral margins curved; lateral margin distinctly sinuate anterior to hind angle; outer carina broader than inner carina; latter distinctly broader behind than in front of middle; base of outer carina narrowed, narrowly truncate; inner carina narrowed near base; outer carina with three or four punctures near lateral margin; inner carina impunctate; both median and paramedian grooves coarsely punctate; pronotum without setae; prosternum without precoxal carinae. Elytron relatively elongate, narrow; striae impressed, coarsely punctate; base of Stria IV without any trace of scarp; Stria IV with three to six setae; apical stride without setae; Stria VII with several setae near apex; metepisternum with coarse punctures near lateral margins, but impunctate in midline; punctures of abdominal Sterna III-V coarse, in form of irregular transverse rows; lateral pits of Sternum IV of female deep, round; male anterior femur with ventral tooth. The punctate pronotal grooves and short, broad head make this isolated species superficially similar to the two preceding ones. However, O. malabaricus differs from them in lacking precoxal carinae; in having the inner carinae strongly narrowed anteriorly; and in having the anterior end of the temporal lobe in the form of an oblique ridge. It is the only Omoglymmius s. str. . in southern India. Range. - In addition to the type material, we have seen specimens from the following localities: four females, Anamalai Hills, Cinchona, 1050 m., VI- 1957 coll. P. S. Nathan (BPBM). Omoglymmius (sensu stricto ) solitarius (Arrow) Figs. 117, 125 Rhysodes solitarius Arrow 1942: 179. Omoglymmius (sensu stricto ) solitarius (Arrow) Bell and Bell 1978. Type material. - HOLOTYPE male, labelled: “ANDAMAN, 76.14” (BMNH). Description. — Length 6 mm. Antennal Segments V-X punctate; head slightly longer than broad; median lobe short, narrow, its sides nearly parallel, its apex broadly rounded; frontal space large, V-shaped; its lateral margins only slightly curved; medial angles produced, their apices obtuse, narrowly separated; posteriomedial margin emarginate; posteriolateral margin rounded; orbital groove short, ended opposite middle of eye; temporal lobe with nine or 10 shallow punctures, restricted to lateral third; one temporal seta; postorbital tubercle absent. Pronotum rather short, length/greatest width 1.24; broadest near middle, margins markedly, evenly curved; base and apex markedly narrowed; lateral margin not sinuate anterior to hind angle; outer carina only 0.3 as wide as inner carina at middle; outer carina of nearly even width, markedly curved; its medial margin not sinuate near base; inner carina widest at middle, tapered anteriorly; narrowed just anterior to base, broadened at extreme base; pronotal carinae entirely impunctate; pronotal setae absent; precoxal carina absent. Elytron relatively long, narrow; striae impressed, coarsely punctate; elytral Intervals III, V slightly elevated and dilated anteriorly; Stria II with three setae near apex; subapical stride with one seta; Stria VII with five or six setae near apex; metasternum coarsely punctate near lateral and anterior margins and in midline; abdominal Sterna III-V with scattered coarse punctures; male with shallow lateral pit on Sternum IV (female unknown); male with ventral tooth on anterior femur; calcar of middle tibia prominent, acute; hind calcar shorter than middle one, triangular, slightly obtuse; This species can be distinguished from all others in the subgenus by the very narrow, evenly curved outer carinae. It is the only member of the subgenus from the Andaman Islands, where there are members of several other subgenera which also have the outer carinae markedly narrowed. Revision of Omoglymmius Ganglbauer 209 Omoglymmius ( sensu stricto) philippensis (Chevrolat) Figs. 118, 126 Rhyzodes philippensis Chevrolat, 1875: 183 (note variant spelling of generic name). Rhysodes philippensis (Chevrolat) Grouvelle 1903. Rhysodes philippinensis (Chevrolat) Hincks 1950. Omoglymmius (sensu stricto) philippensis (Chevrolat) Bell and Bell 1978. Type material. - LECTOTYPE (here designated) male, labelled: “Ins. Philippine, Chevrolat, Chevl. Or. Typ., philippensis ” (NMW). According to the original description, the type was received from Henri Deyrolle. PARALECTOTYPES: It is uncertain whether additional specimens exist from the original type series. Description. — Length 5. 0-6. 7 mm. Antennal Segments I- IV coarsely punctate; V either punctate or impunctate; VI-X1 impunctate; head 1.5 longer than broad; median lobe short, narrow, lance-shaped, its apex pointed; frontal space large, V-shaped, its lateral margin curved; medial angles obtuse, slightly separated; posteriomedial margin oblique, shallowly emarginate; posteriolateral margin evenly curved; occipital angle distinct; orbital groove fine, becoming obsolete near middle of length of eye; temporal lobe with approximately 10 coarse punctures in posteriolateral portion; one temporal seta; postorbital and suborbital tubercles absent; eye large, round. Pronotum moderately elongate, length/greatest width 1.27; widest near middle, base and apex narrowed; lateral margins moderately curved; lateral margin not sinuate anterior to hind angle; carinae subequal at middle; medial margin of outer carina distinctly sinuate just anterior to base; outer carina widest near middle, distinctly narrowed to base; inner carina narrowed near base, but extreme base broadened, truncate; outer carina with 10-28 (most specimens 12-15) coarse punctures; inner carina impunctate or with only one or two punctures; pronotum without setae; prosternum without precoxal carinae. Elytron relatively long, narrow; striae impressed, coarsely punctate; base of Stria IV with very small longitudinal pollinose scarp; Stria IV with one to five setae or none (in most specimens with one at base and one or two near apex), in a few with unbroken row of four or five setae; apical striole with one seta or none; apex of Stria VII with several setae; metasternum coarsely punctate; punctures of Sterna I1I-V scattered, numerous; female with lateral pit of Sternum IV deep, round; male with ventral tooth on anterior femur; middle calcar acute, rather small about 0.5 of length of spur; hind calcar triangular, its apex slightly lobate, obtuse. This species is the most widespread of a large group of very similar forms, and is the standard for comparison for them. It is a species with a rather long head, impunctate inner carina and punctate outer carina, with a very small scarp at the base of Stria IV. Among species sympatric with it in Luzon, O. imugani is the most similar, but differs in shape of pronotum and in reduction of punctuation on outer carinae and temporal lobes. O. duplex differs in having more than one temporal seta and in shape of the head. O. hiekei differs in having the outer carinae impunctate and in lacking the longitudinal scarp on Stria IV. O. semperi and O. data have the inner carinae punctate and the eyes reduced. In Palawan, O. philippensis is sympatric with O. coelebs. The latter species has the outer carinae broad to the base, and the basal scarp of Stria IV entirely absent. Other Philippine species will be compared with O. philippensis when they are described below. Range. - Luzon, Mindoro, Palawan, and Siargao. We have seen the following specimens: LUZON: one female, Los Banos, coll. Baker (MCZ), one male, same locality, 27-Juli, 1914, leg. Boettcher (MNHB); four males, Mt. Makiling, Laguna, V-l-31, F. C. Hadden Colin. (CAS); four males, three females, Mt. Makiling, Baker (NMNH); one male, Pangil, Laguna, V-21-31, F. C. Hadden Colin. (CAS); one male, Quezon Park, Tayabas, alt. 1000 ft., VI-5-32, F. C. Hadden Colin. (BPBM); MINDORO; one male (MNHN); PALAWAN: one female, Binaluan, leg. Boettcher (MNHB); SIARGAO, two males, Dapa, leg. Boettcher (MNHB). Variation. - There appears to be considerable variation in development of setae of Stria IV; however, these are minute and it is difficult to tell whether or not they have been broken off in some specimens. The specimen from Palawan might be a distinct species; it has five setae in Stria IV (maximum of four in other O. philippensis ). Quaest. Ent., 1982, 18 (1-4) 210 Bell and Bell Omoglymmius { sensu stricto) imugani new species Figs. 119, 127 Type material. - HOLOTYPE female, labelled: “Imugin, N. Viscaya, Baker” (NMNH). PARATYPES one female, labelled: “Philippinen, Imugan, 6-6-1916, leg. Boettcher” (MNHB), one male, same data, 16-6-1917 (MNHB), one female, same data, 30-6-1917 (MNHB). Description. — Length 4. 8-6. 7 mm. Antennal Segments I- 1 1 1 coarsely punctate, Segment IV with few punctures or impunctate, Segments V-Xl impunctate; head about 1.5 longer than wide; median lobe short, lance-shaped, its apex obtusely pointed in some specimens, rounded in others; frontal space large, V-shaped; its lateral margin weakly curved; medial angles obtuse, slightly separated; posteriomedial margin oblique, weakly emarginate; posteriolateral margin evenly curved, occipital angle distinct; orbital groove fine, tending to be broken into series of punctures; temporal lobe impunctate except for those in orbital groove; one temporal seta; postorbital and suborbital tubercles absent; eye large, round. Pronotum moderately long, length/greatest width 1.26; widest near middle; apex more narrowed than base; lateral margins curved; lateral margin not sinuate anterior to hind angle or else slightly sinuate there; carinae subequal at middle; medial margin of outer carina slightly sinuate anterior to base; outer carina widest at middle, slightly narrowed to base; inner carina narrowed near base, at most slightly broadened at base; outer carina with six to nine coarse punctures; inner carina impunctate; pronotum without setae; prosternum without precoxal carinae. Elytron relatively long, narrow; striae impressed, coarsely punctate; base of Stria IV with longitudinal pollinose scarp; Stria IV with one seta or none at base and one or two at apex or none; subapical striole with one seta or none; apex of Stria VII with several setae; metasternum coarsely punctate along margins and in posterior half of midline; anterior half of midline punctate in some specimens, impunctate in others; disc of metasternum impunctate; punctures of abdominal Sterna III-V sparse, in form of transverse row on each sternum; female with lateral pit of Sternum IV deep, round; male with ventral tooth on anterior femur; middle calcar acute; hind calcar triangular, its apex lobate, obtuse. This species differs from O. philippensis in having the pronotum more narrowed anteriorly, and in having the punctation reduced, especially on the temporal lobes, outer pronotal carina, metasternum and abdomen, and in having a wider median lobe. In most respects, O. imugani is nearly identical to O. politus , but the two differ sharply in male secondary sexual characters; the former species has the ventral tooth of the anterior femur present and the middle calcar large, while in the latter species, the femoral tooth is absent and the middle calcar is minute. In O. imugani , the eye is large, while in O. politus , it is reduced. Omoglymmius (sensu stricto ) politus new species Figs. 120, 128 Type material. - HOLOTYPE male, labelled: “Philippinen, Luzon Prov., Lepanto, Mt. Polis, 2-1917, leg. Boettcher” (MNHB). PARATYPE female, same data as holotype (MNHB). Description. — Length 5. 7-6. 6 mm. Antennal Segments I-III coarsely punctate; Segments IV-XI impunctate; head 1 .5 longer than wide; median lobe short, lance-shaped, its apex obtusely pointed, its width greater than in O. imugani ; frontal space large, V-shaped, its lateral margins feebly curved; medial angles obtuse, slightly separated; posteriomedial margin oblique, slightly emarginate; posteriolateral margin evenly curved, occipital angle distinct; orbital groove in form of row of fine punctures; temporal lobe otherwise impunctate; one temporal seta; postorbital, suborbital tubercles absent; eye slightly reduced. Pronotum moderately long; length/greatest width 1.29; widest near middle; apex more narrowed than base; lateral margins curved; shallowly sinuate anterior to hind angle; carinae subequal at middle; medial margin of outer carina slightly sinuate anterior to base; outer carina widest near middle, narrowed to base; inner carina narrowed near base, but dilated at extreme base; outer carina with six to 10 coarse punctures; inner carina impunctate; pronotum without setae; prosternum without precoxal carinae. Elytron relatively long, narrow; striae impressed, coarsely punctate; base of Stria IV with pollinose scarp minute, indistinct; Stria IV with one seta at apex, none at base; subapical striole with one seta; apex of Stria VII with several setae; metasternum coarsely punctate near margins and in midline, its disc otherwise impunctate; punctures of abdominal Sterna III-V sparse, in form of one transverse row on each sternum; female with lateral pit on Sternum IV deep, round; male without ventral tooth on anterior femur; middle calcar minute; hind calcar as in O. imugani. Revision of Omoglymmius Ganglbauer 211 This species is very similar to O. imugani except in the secondary sexual characters of the male. The series of both species at hand are so small that it is impossible to be sure of the reliability of nonsexual characters. The slightly reduced eye, broader median lobe, more elongate pronotum with more distinct sinuation of the lateral margin, and broader base of the inner carina are all possible differences from O. imugani. Omoglymmius ( sensu stricto ) crassicornis new species Fig. 130 Type material. - HOLOTYPE female (Negros I.). PHILIPPINES, labelled: “Dumaguete, P.I., May 1925, John Chapman” (NMNH). PARATYPE female, labelled: “Dumaguete, P.I., Horns of Negros, J. W. Chapman Coll.” (MCZ). Description. — Length 6. 0-6. 5 mm. Outer segments of antennae stouter than in related species; Antennal Segments I-VI punctate; fine, sparse punctures on Segments VII, VIII in one specimen; IX-XI impunctate; head 1.5 longer than broad; median lobe narrow, lance-shaped, longer than in O. philippensis, its apex obtusely pointed; frontal space large, V-shaped, its lateral margin shallowly curved; medial angles obtuse, slightly separated; posteriomedial margin oblique, posteriolateral margin evenly curved; occipital angle distinct; orbital groove distinct, extended to posterior margin of eye; temporal lobe with four to nine coarse punctures in posteriolateral portion; one temporal seta present; postorbital and suborbital tubercles absent; eye large, round. Pronotum moderately elongate, length/greatest width 1.17 to 1.34; widest near middle, base, apex narrowed; lateral margins curved, subangulate in middle in one specimen; lateral margin not sinuate anterior to hind angle; carinae subequal at middle; medial margin of outer carina scarcely sinuate anterior to base; outer carina widest near middle, distinctly narrowed to base; inner carina narrowed near base, but extreme base broadened, its width nearly equal to width of inner carina at middle; outer carina with 12-17 coarse punctures; inner carina impunctate or with only one or two coarse punctures; pronotum without setae; prosternum without procoxal carinae. Elytron relatively long, narrow; striae impressed, coarsely punctate; base of Stria IV with very small longitudinal pollinose scarp; Stria I with 0-1 seta near apex; Stria II with one or two setae near apex; Stria IV with one or two setae near apex; subapical stride with one seta or none; Stria VII with several setae near apex; metasternum with coarse punctures near margin and in midline, but with disc otherwise impunctate; punctures of Sterna III-V coarse, in part in form of a single transverse row on each sternum; female with lateral pit on Sternum IV deep, round; male unknown. This species is the only Omoglymmius s. str. known from the island of Negros. It is similar to O. philippensis but differs in the following points: punctures of the abdominal sterna form transverse rows, punctures of the metasternum are restricted to the margins and midline; the median lobe of the head is longer; punctures of the antenna extend to more distal segments, and the outer antennal segments are stouter. Variation. - The two specimens differ markedly in shape of the pronotum. Nevertheless, they are markedly similar otherwise, and are probably conspecific. Probably one of the two is an extreme variant. More specimens will have to be collected to determine the limits of variation. Omoglymmius (sensu stricto ) amplus new species Fig. 131 Type material. - HOLOTYPE female, labelled: “SUMATRA, Palembg., Grouvelle” (LEI). The locality probably refers to the Province, rather than the city of Palembang. PARATYPE female labelled: “Sumatra, dono Grouvelle, 1901”, (GEN). Description. — Length 6.5 mm. All antennal segments coarsely punctate; head 1.5 longer than wide; median lobe short, rather broad, its tip broadly rounded; frontal space large, V-shaped, its lateral margin curved; medial angles obtuse, narrowly separated; posteriomedial margin oblique, nearly straight; posteriolateral margin evenly curved; occipital angle distinct; orbital groove fine, extended to posterior margin of eye; temporal lobe with five to seven coarse punctures near temporal seta; one temporal seta; postorbital, suborbital tubercles absent; eye large, round. Quaest. Ent., 1982, 18 (1-4) 212 Bell and Bell Pronotum moderately long, length/greatest width 1.20; widest slightly anterior to middle; apex rather suddenly narrowed; margin markedly oblique near apex; base markedly narrowed; lateral margin strongly sinuate anterior to hind angle; inner carina slightly narrower than outer one at middle; medial margin of outer carina shallowly sinuate just anterior to base; outer carina widest at anterior third, constricted opposite lateral sinuation, broadened at extreme base; base of outer carina directed obliquely laterad; inner carina narrowed just anterior to base; extreme base broadened; outer carina with 21 punctures; inner carina impunctate; pronotum without setae; prosternum without precoxal carina. Elytron relatively long, narrow; striae impressed, coarsely punctate; base of Stria IV with longitudinal pollinose scarp; Interval V with tubercle laterad to base of Stria IV; Stria II with one or two setae near apex; Stria IV with one seta near apex; subapical stride without seta; apex of Stria VII with several setae; metasternum with coarse, irregularly scattered punctures; punctures of abdominal Sterna III-V numerous, scattered; female with lateral pit of Sternum IV deep, round; male unknown. This species and the next resemble O. philippensis in most respects, but have the pronotum broadest anterior to the middle. Of the other species from Sumatra, O. summissus has a subquadrate pronotum, punctate inner carinae, and lacks a longitudinal basal scarp on Stria IV; O. fraudulentus likewise lacks the scarp, and has the temporal lobes very coarsely punctate; O. bouchardi has the pronotum nearly impunctate, and has the punctures of the abdominal sterna largely confluent. Omoglymmius (sensu stricto) modiglianii new species Fig. 132 Type material. - HOLOTYPE female, labelled: “MENTAWEI, Si Oban, IV-VIII, Modigliani 94” (GEN). PARATYPE female (teneral), same data as holotype (GEN). Description. — Length 5. 7-6.0 mm. All antennal segments punctate; punctures coarse on proximal segments, finer on distal segments; head 1.5 longer than wide; median lobe short, its tip broadly rounded; frontal space large, nearly U-shaped, its lateral margin more markedly curved than in O. antplus\ medial angles nearly rectangular, narrowly separated; posteriomedial margin oblique, slightly sinuate; posteriolateral margin evenly curved; occipital angle distinct; orbital groove fine, extended to posterior margin of eye; temporal lobe with 13-15 very coarse punctures in its lateral half; one temporal seta; postorbital, suborbital tubercles absent; eye large, rounded. Pronotum moderately elongate; length/greatest width 1.26, widest anterior to middle, apex rather suddenly narrowed; margin oblique near apex; base strongly narrowed; lateral margin sinuate anterior to hind angle; inner carina slightly narrower than outer one at middle; medial margin of outer carina shallowly sinuate just anterior to base; outer carina widest at anterior third, scarcely constricted opposite lateral sinuation; base of outer carina not turned obliquely laterally; inner carina narrowed just anterior to base, but extreme base broadened; outer carina with about 20 coarse punctures; inner carina impunctate; pronotum without setae; prosternum without precoxal carinae. Elytron relatively long, narrow; striae impressed, coarsely punctate; base of Stria IV with small longitudinal scarp; Interval V laterad to base of Stria IV with low tubercle; Stria I with one seta near apex or none; Stria II with up to two near apex or none; Stria IV with one seta near apex; subapical stride with one seta; apex of Stria VII with several setae; metasternum with coarse, irregularly distributed punctures; punctures of abdominal Sterna III-V numerous, scattered; female with lateral pit of Sternum IV deep, round; male unknown. This species is the only member of the subgenus known from the Mentawei Islands. It is close to O. ampins , but differs in having the pronotum more elongate, its lateral margin less sinuate, the base of the outer carina not turned outwards, and the temporal lobe with the punctures coarser and more numerous. Omoglymmius ( sensu stricto) evasus new species Fig. 134 Type material. - FIOLOTYPE male, labelled: “E. slope Mt. McKinley, Davao Prov., MINDANAO, elev. 3300 ft., IX-28-1946, F. G. Werner leg., lot #92, dead stump, CNHM Phil. Zool. Exp. (1946-47)” (CNHM). PARATYPES one female, same data as holotype (CNHM); one male, three females, same locality as holotype, IX- 1946, H. Hoogstrall leg. (CNHM). Revision of Omoglymmius Ganglbauer 213 Description. — Length 5. 0-7. 2 mm. Antennal Segments I - 1 1 1 coarsely punctate; Segment IV with a few, rather fine punctures; Segments V-XI impunctate; head 1.3 longer than broad; median lobe rather short, its apex broadly rounded; frontal space moderate-sized, V-shape, its lateral margins curved; medial angles obtuse, slightly separated; posteriomedial margin oblique; posteriolateral margin evenly curved; occipital angle distinct; orbital groove fine, extended to hind margin of eye; temporal lobe with three to nine coarse punctures in lateral portion; one temporal seta; postorbital, suborbital tubercles absent; eye large, round. Pronotum moderately elongate; length/greatest width 1.34, widest near middle, base and apex narrowed; apex much less narrowed than in O. philippensis\ lateral margins moderately curved; lateral margin shallowly sinuate anterior to hind angle; carinae subequal in middle; medial margin of outer carina sinuate just anterior to base; outer carina widest in anterior third, transversely truncate at apex; inner carina narrowed near base, but extreme base broadened, truncate; outer carina with 15-18 coarse punctures; inner carina impunctate; pronotum without setae; prosternum without precoxal carinae. Elytron relatively narrow, long; striae impressed, coarsely punctate; base of Stria IV without longitudinal scarp; Stria II with one seta near apex; Stria IV with one seta near apex; most specimens with one seta at base of Stria V near humerus; subapical stride without setae; Stria VII with several setae near apex; metasternum with coarse punctures near margin and scattered near midline; disc otherwise impunctate; punctures of abdominal Sterna III-V coarse, sparse, scattered; female with lateral pit of Sternum IV round, rather shallow; male with small, obtuse ventral tooth on anterior femur; middle calcar prominent, acute; hind calcar triangular, its apex obtuse. This species differs conspicuously from O. philippensis in the shape of the pronotum, particularly of the outer carinae, which are truncate anteriorly. Absence of the longitudinal scarp at the base of Stria IV, the smaller number of punctures on the temporal lobes, and metasternum are among numerous additional differences. The truncate apex of the outer carina will separate O. evasus from similar species in nearby regions, although O. nemoralis of Sarawak approaches it. The latter species has the setae of the elytra much more numerous, and the punctures of the outer carina fewer in number and limited to its posterior portion. Range. - Mindanao, Philippine Islands. In addition to the type material, we have seen the following specimens: two males, two females, Baracatan, 1500 m., June 27-29-1977, M. Sato leg. (SATO); two females, Mainit Hot Spring, 28- V 1 1- 1 970, M. Sato leg. (SATO); one male, four females, Todaya, July 30, 1970, M. Sato leg. (SATO). Omoglymmius (sensu stricto) nemoralis new species Fig. 135 Type material. - HOLOTYPE male, labelled: “Mt. Matang, W. Sarawak, 17-1-14, G. Bryant Coll. 1919-147” (BMNH). PARATYPES one male, same data as holotype (BMNH); one male, labelled: “Sarawak. Matang Riv., 3 1/2 mile, J.E.A. Lewis, 1910-116” (BMNH); one female, Semengoh For. Res., 15 mi. s. Kuching, 22-IX-1966, J.F.G. Clarke, Thelma M. Clarke (NMNH). Description. — Length 5. 6-6.0 mm. Antennal Segments I- V III coarsely punctate; Segments IX-XI impunctate; head 1.5 longer than broad, distinctly more elongate than in O. evasus\ median lobe rather narrow, its apex obtusely pointed; frontal space moderate-sized, V-shaped, its lateral margins curved; medial angles obtuse, slightly separate; posteriomedial margin oblique; posteriolateral margin evenly curved; occipital angle distinct; orbital groove fine, not extended to hind margin of eye, interrupted in some specimens; temporal lobe with seven to nine coarse punctures in lateral portion; one temporal seta; postorbital, suborbital tubercles absent; eye large, round. Pronotum moderately elongate; length/greatest width 1.26; widest near middle; base, apex narrowed; lateral margins moderately curved; lateral margin shallowly sinuate anterior to hind angle; carinae subequal in width at middle; medial margin of outer carina scarcely sinuate anterior to base; outer carina widest in anterior third, transversely truncate at apex; inner carina narrowed near base, but extreme base broadened, truncate; outer carina with nine to 16 coarse punctures, anterior 0.25-0.5 of outer carina impunctate; inner carina impunctate; pronotum without setae; prosternum without precoxal carinae. Elytron relatively narrow, long; striae impressed, coarsely punctate; base of Stria IV without longitudinal scarp; Stria II with two setae near apex; Stria IV with six or seven setae forming uninterruped row from base to apex; base of Stria V without seta; subapical striole with one seta or none; Stria VII with several setae near apex; metasternum coarsely punctate, with most punctures near midline or margins; punctures of Sterna III-V coarse, sparse, scattered; female with lateral pit of Sternum IV round, moderately deep; male with small, obtuse ventral tooth on anterior femur; middle calcar Quaest. Ent., 1982, 18 (1-4) 214 Bell and Bell of male acute, smaller than in O. evasus\ hind calcar triangular, apex obtuse. This species is allopatric with its closest ally, O. evasus of Mindanao. From the latter species it can be separated by the more numerous elytral setae and the absence of punctures from the anterior part of the outer carina. O. fraudulentus of Borneo and Sumatra and O. malaicus of Malay Peninsula are also very similar. In the former species, the outer carinae are markedly divergent at their bases, while in the latter, the pronotum is narrower, with its margins more nearly parallel, with apex of the median lobe rounded. The only other member of the subgenus known to occur on Borneo is O. consors. The latter species differs strikingly in completely impunctate outer carinae and the presence of a transverse band of pollinosity on the basal scarp of the elytron. This species is possibly O. borneensis (Grouvelle) 1903, p. 119-120). We have been unable to locate type material for O. borneensis (Type locality R. Sambey, coll. Ledru), indicated as being in the Oberthiir Collection. There are several points of disagreement between the original description and O. nemoralis, and it is quite possible that Grouvelle’s name applies to O. consors , O. fraudulentus, or to some other member of the subgenus, not yet found in Borneo. Omoglymmius (sensu stricto) fraudulentus new species Fig. 136 Type material. - HOLOTYPE male, labelled: “SUMATRA, Palembg” (MNHN); PARATYPES three males, eight females, labels identical to holotype (MNHN). Description. — Length 5. 5-6. 4 mm. Antennal Segments I-1V coarsely punctate; a few punctures on V, VI; Segments VII-XI impunctate; head 1.5 longer than broad; median lobe rather narrow, its apex obtusely pointed; frontal space moderate-size, V-shaped, its lateral margins curved; medial angles obtuse, slightly separated; posteriomedial margin less distinctly oblique than in the two preceding species, posteriolateral margin evenly curved; occipital angle indistinct; orbital groove fine, interrupted; temporal lobe with six to 10 coarse punctures in lateral portion; one temporal seta; postorbital, suborbital tubercles absent; eye large, round. Pronotum moderately elongate; length/greatest width 1.29; widest near middle; apex narrowed; base slightly narrower than greatest width; lateral margins moderately curved; lateral margin markedly sinuate anterior to hind angle; carinae subequal in width at middle; outer carinae abruptly bent near base, strongly divergent there; outer carina widest anterior to middle, narrowed, obtusely rounded at apex; outer carina with three or four (Borneo specimen) or 14-16 (Sumatra specimens) coarse punctures; inner carina impunctate; pronotum without setae; prosternum without precoxal carina. Elytron relatively narrow, long, striae impressed, coarsely punctate; base of Stria IV wihout longitudinal scarp; Stria II with two setae near apex; Stria IV with complete series of five setae; subapical stride with one seta; Stria VII with several setae near apex; metasternum coarsely punctate; punctures of abdominal Sterna II1-V coarse, rather dense; female with lateral pit on Sternum IV round, rather shallow; male with small, obtuse ventral tooth on anterior femur; calcars as in O. nemoralis. This species is distinguished from its closest relatives by the sharply divergent bases of the outer carinae. Among the other members of the subgenus on Sumatra, O. consors comes closest, but differs in having a transverse band of pollinosity on the basal scarp of the elytron, and in having the outer carina completely impunctate. In Borneo, the most similar species is O. nemoralis , which does not have the bases of the outer carinae divergent. Range. - Sumatra and Borneo. In addition to the type material, we have seen one female from Sumatra, labelled: “Tebbing Tinggi, VI-37 , Brues” (MCZ), and one male from Borneo, labelled: “Miri Riv. VIII, Mjoberg Colin., W. 1S/I. Funge Bequest-’ (CAS). The latter specimen differs from those of Sumatra having only three or four punctures on the outer carina. It does not seem to differ otherwise, and is provisionally assigned to this species. Omoglymmius (sensu stricto ) malaicus (Arrow) Fig. 133 Rhysodes malaicus Arrow 1901: 85-86. Revision of Omoglymmius Ganglbauer 215 Omoglymmius ( sensu stricto) malaicus (Arrow) Bell and Bell 1978. Type material. - LECTOTYPE (here designated) male, labelled: “Penang, Lamb, Pascoe Coll.” (BMNH). PARALECTOTYPE female, same label as lectotype (BMNH). Penang is a small offshore island near the Malay Peninsula. Description. — Length 6.0 mm. Antennal Segments I-V punctate, punctures of Segment V very small; Segments VI-XI impunctate; head 1.5 longer than broad; median lobe rather narrow, its sides parallel, its apex rounded; frontal space moderate-size, U-shaped, its lateral margins curved; medial angles obtuse, very narrowly separated; posteriomedial margin oblique; posteriolateral margin evenly curved; occipital angle distinct; orbital groove fine, interrupted; temporal lobe with seven to nine coarse punctures in lateral portion; one temporal seta; postorbital, suborbital tubercles absent; eye large, round. Pronotum elongate, length/greatest width 1.43; widest slightly anterior to middle, narrowed at apex, scarcely narrowed to base, lateral margins nearly parallel; lateral margin not sinuate anterior to hind angle; carinae subequal in width at middle; outer carinae not divergent at base; inner carina constricted anterior to base, latter broadened; outer Carina with 10-14 coarse punctures; inner carina impunctate; pronotum without setae; prosternum without precoxal carinae. Elytron relatively long, narrow; striae impressed, coarsely punctate; base of Stria IV without longitudinal scarp; Stria 1 1 with two setae near apex; Stria IV with complete series of four to seven setae; subapical striole with one seta; Stria VII with several setae near apex; metasternum coarsely punctate; punctures of abdominal Sterna 1 1 I-V coarse, rather dense; female with lateral pit round, rather deep; male with ventral tooth on anterior femur more prominent than in preceding species; middle calcar acute, prominent; hind calcar triangular, its apex obtuse. This is the only member of the subgenus known from the Malay Peninsula. It is closest to O. fraudulentus of Sumatra, but differs in shape of the pronotum, and lacks the divergent bases of the outer carinae which are so prominent in the latter species. Range. - Malay Peninsula and offshore islands. In addition to type material, we have seen the following specimens: one female, labelled: “Poulo Penang, Raffray, compared with type, G.J.A., correctly named, G.J.A.” (MNHN); one specimen, sex not recorded, labelled “Penang-Bryant” (BMNH); one specimen, sex not recorded, labelled “Perak, Doherty” (BMNH). Omoglymmius (sensu stricto) coelebs new species Fig. 137 Type material. - HOLOTYPE male, labelled: “N. Palawan, P. I. Binaluan, Nov.-Dez.-1913, leg. G. Boettcher, from H. P. Loding, 1935” (NMNH). Description. — Length 6.0 mm. Antennal Segments I-IV coarsely punctate; Segments V-VII with few punctures; Segments VIII-XI impunctate; head 1.5 longer than broad; median lobe with margins nearly parallel, apex rounded; frontal space moderate in size, U-shaped, its margins curved; medial angles obtuse, seperated; posteriomedial margin oblique; posteriolateral margin evenly curved; occipital angle indistinct; orbital groove fine, not attaining posterior margin of eye, interrupted; temporal lobe with eight or nine coarse punctures in lateral portion; one temporal seta; postorbital, suborbital tubercles absent; eye large, round. Pronotum moderately elongate, length/greatest with 1.26; base, apex narrowed; lateral margins moderately curved; lateral margin not sinuate anterior to hind angle; carinae subequal in width at middle; medial margin of outer carina scarcely sinuate anterior to base; outer carina of almost even width, transversely truncate at both apex and base; inner carina constricted just anterior to base; latter broad, truncate; outer carina with 15-19 coarse punctures; anterior 0.25 of outer carina impunctate; inner carina impunctate; pronotum without setae; prosternum without precoxal carinae. Elytron relatively narrow, long; striae impressed, coarsely punctate; base of Stria IV without longitudinal scarp; Stria I with one seta near apex; Stria II with one to three setae near apex; Stria IV with complete series of four or five setae; apical striole with one seta; Stria VII with several setae near apex; metasternum with coarse punctures near anterior and lateral margins, otherwise nearly impunctate; punctures of Sterna 1 1 I-V coarse, rather dense, forming broad transverse band on each sternum; male with small, obtuse ventral tooth on anterior femur; middle calcar acute, small; hind calcar triangular, its apex obtuse, slightly lobate; female unknown. This species is closest to O. evasus and O. nemoralis, from which it differs conspicuously in having the outer carinae not narrowed posteriorly. On Palawan it is sympatric with O. philippensis. The latter species is longer and narrower, and has the pronotum narrowed both at the base and apex. In addition, it has a small longitudinal scarp at the base of Stria IV. Quaest. Ent., 1982, 18 (1-4) 216 Bell and Bell Omoglymmius (sensu stricto) thoracicus new species Figs. 138, 139 Type material. - HOLOTYPE male (JAVA), labelled: “Tangk.-Prahoe, Juni- 1933, V. Doesburg” (LEI). Description. — Length 5.9 mm. All antennal segments coarsely punctate; head 1.5 longer than wide; median lobe oval, its apex broadly rounded; frontal space broad, U-shaped, its lateral margins abruptly bent; medial angles obtuse, slightly separated; posteriomedial margin oblique; posteriolateral margin evenly rounded; occipital angle distinct; orbital groove absent; temporal lobe with nine to 12 rather fine punctures in posteriolateral half; temporal seta absent; postorbital and suborbital tubercles absent; eye large, round. Pronotum moderately elongate; length/greatest width 1.24; widest near middle; base and apex narrowed; lateral margins strongly curved; lateral margin slightly sinuate anterior to hind angle; carinae subequal in width at middle; medial margin of outer carina distinctly sinuate just anterior to base; outer carina widest anterior to middle, narrowed to base and apex; inner carina narrowed just anterior to base, latter broad, truncate; outer carina with 13-16 rather fine punctures; inner carina impunctate; pronotum without setae; prosternum with sharp precoxal carina on each side, extended 0.75 of distance from coxa to anterior margin (Fig. 1 39). Elytron relatively long, narrow; striae impressed, coarsely punctate; base of Stria IV without longitudinal scarp; Striae I, II without setae; Stria IV with one seta near apex; subapical stride with one seta; apex of Stria VII with several setae; metasternum coarsely punctate; punctures of abdominal Sterna 1II-V scattered, numerous; male with small, obtuse, ventral tooth on anterior femur; middle calcar acute, subequal to spur; hind calcar small, triangular, obtuse, margin emarginate between calcar and tibial spur; female unknown. In dorsal aspect this species looks like O. philippensis and its relatives, except that the punctures of the temporal lobes and outer carinae are finer and the temporal seta is absent. In ventral view it is easily distinguished by well-developed precoxal carinae. It shares this character with another Javanese species, O. pectoralis. The latter species, however, entirely lacks punctures on the outer carinae, and has four temporal setae. Omoglymmius ( sensu stricto ) summissus new species Fig. 140 Type material. - HOLOTYPE female, labelled: “SUMATRA, Palembg.” (MNHN). (The type was part of a group of specimens of several species mounted on a single pin; consequently, the label on the holotype was copied by us.) Description. — Length 6.1 mm. Antennal Segments I-VI1 coarsely punctate; Segments VIII-XI impunctate; median lobe short, lance-shaped, its apex obtuse; frontal space large, V-shaped, its lateral margins only slightly curved; medial angles very obtuse, separated; posteriomedial margin slightly oblique; posteriolateral margin evenly rounded; occipital angle indistinct; trace of orbital groove present near anterior margin of eye, at junction with distinct preorbital pit; temporal lobe with 10-14 coarse punctures in posteriolateral half; one temporal seta; postorbital, suborbital tubercles absent; eye large, round. Pronotum moderately elongate; length/greatest width 1.31; widest near middle; base slightly narrowed; apex narrowed abruptly; lateral margins nearly straight, almost parallel; lateral margin not sinuate anterior to hind angle; inner carinae slightly wider than outer ones at middle; medial margin of outer carina not sinuate anterior to base; outer carina widest anterior to middle; its apex obliquely truncate; inner carina narrowed near base, broadened, truncate at base; outer carina with 12-18 coarse punctures; inner carina with four or five punctures in posterior half; pronotum without setae; prosternum without precoxal carinae. Elytron relatively long, narrow; striae impressed, coarsely punctate; base of Stria IV with slight suggestion of longitudinal scarp, but latter entirely glabrous; Stria II with two setae near apex; Stria IV with three setae in anterior third, one near apex; subapical stride with one seta; apex of Stria VII with several setae; metasternum entirely coarsely punctate; punctures of abdominal Sterna III-V rather fine, scattered, not at all confluent; female with deep, round lateral pit on Sternum IV; male unknown. This species differs from O. philippensis and similar species by punctures on the inner carina. Shape of the pronotum is closest to that of O. malaicus. The latter species has a longer, narrower pronotum with outer carinae more narrowed anteriorly, and without punctures on the inner carina. Among Sumatran species, O. consors and O. bouchardi have both inner and outer Revision of Omoglymmius Ganglbauer 217 carinae impunctate, while O. fraudulentus has only the outer carinae punctate. The latter species also differs from O. summissus in having the bases of the outer carinae divergent. Omoglymmius (sensu stricto ) semperi new species Fig. 141. 147 Type material. - HOLOTYPE male, labelled: “Phillip. Islands, Semper” (MNHN) PARATYPES two females, with labels identical to that of holotype and the additional label “Fry Colin. 1905-100” (BMNH). Description. — Length 5. 4-6. 4 mm. Antennal Segments Ml coarsely punctate; Segment III with a few fine punctures; Segments 1V-IX impunctate; head 1.5 longer than wide; median lobe short, lance-shaped, its apex acute; frontal space V-shaped, its margins scarcely curved; medial angles obtuse, well separated; posteriomedial margin oblique, slightly emarginate; posteriolateral margin evenly curved; occipital angle distinct; orbital groove narrow but distinct, extended posteriorly at least to middle of eye; temporal lobe with nine to 12 coarse punctures in posteriolateral portion; one temporal seta; postorbital, suborbital tubercles absent; eye reduced, its depth about 0.6 of depth of head, its posterior margin oblique, nearly straight. Pronotum elongate; length/greatest width 1.44; widest near middle, base, apex narrowed; lateral margin curved, scarcely sinuate anterior to hind angle; carinae subequal in width at middle; medial margin of outer carina scarcely sinuate anterior to base; outer carina widest near middle, its apex markedly narrowed; inner carina narrowed just anterior to base; latter broadly truncate; outer carina with 25-30 coarse punctures; inner carina with 10-14 coarse punctures concentrated in anterior and posterior thirds of carinae, middle third impunctate or nearly so; pronotum without setae; prosternum without precoxal carinae. Elytron relatively long, narrow; striae impressed, coarsely punctate; base of Stria IV with small, pollinose longitudinal scarp; Stria II without setae; Stria IV with one seta at apex; subapical stride with one seta; Stria VII with several setae near apex; metasternum entirely coarsely punctate; punctures of abdominal Sterna 1II-V coarse, rather dense, those of 111 in form of irregular transverse row; those of IV, V scattered; female with deep round lateral pit on Sternum IV; male with small obtuse ventral tooth on anterior femur; middle calcar acute, very small, less than 0.3 as long as spur; hind calcar triangular, its apex obtuse, slightly lobate, its upper margin angulate. This species and the next one differ from O. philippensis by punctures on the inner carina and by reduction of the eyes. In this species, the eyes are only slightly reduced in depth, but are sharply reduced in length, with the posterior margin forming an oblique, nearly straight line. This species is dedicated to the collector, Professor Carl Semper (1832-1893) of the University of Wurzburg, whose distinguished career included several years of pioneer collecting in the Philippines. It is not known on what island he collected this species. Its similarity to O. data suggests that it might be a localized montane form from northern Luzon. Omoglymmius ( sensu stricto ) data new species Figs. 142, 148 Type material. - HOLOTYPE male, labelled: “P.I., Mt. Data, Mt. Province, 5000 ft., Ill- 1 6- 193 1 , F.C. Hadden Colin.” (CAS). PARATYPE female, labelled: “Philippines, Mt. Prov., Abatan, Buguias, 60 km. S. of Bontoc, 1800-2000 m., 9-VI-1964, coll. H.M. Torrevillas” (BPBM). Description. — Length 6. 0-6. 6 mm. Antennal Segments 1, II coarsely punctate; Segments 1I1-XI impunctate; head 1.5 longer than wide; median lobe lance-shaped, notched at junction of clypeus; apex obtuse; frontal space V-shaped, its margins sharply curved; medial angles obtuse, contiguous; posteriomedial margin slightly curved; posteriolateral margin evenly curved; occipital angle very indistinct; orbital groove absent; temporal lobe with 25-30 rather fine, scattered punctures; one temporal seta; postorbital, suborbital tubercles absent; eye strongly reduced, its depth less than 0.5 of depth of head, its posterior margin curved. Pronotum moderately elongate; length/greatest width 1.21; widest near middle; base moderately narrowed; apex strongly narrowed; lateral margins curved; lateral margin not sinuate anterior to hind angle; carinae subequal in width at middle; medial margin of outer carina scarcely sinuate anterior to base; outer carina widest anterior to middle, its apex narrowed; inner carina only slightly constricted just anterior to base, latter broadly truncate; outer carina with 25-30 Quaest. Ent., 1982, 18 (1-4) 218 Bell and Bell rather fine punctures; inner carina with eight to 10 punctures; pronotum without setae; prosternum without precoxal carinae. Elytron relatively long, narrow; striae impressed, coarsely punctate; base of Stria IV without longitudinal carina; Stria II without setae; Stria IV with one apical seta; subapical striole without seta; apex of Stria VII with several setae; metasternum entirely coarsely punctate; punctures of abdominal Sterna as in O. semperi\ female with round lateral pit on Sternum IV; male with small obtuse ventral tooth on anterior femur; middle calcar minute; hind calcar triangular, its apex obtuse, its upper margin not angulate. This species is similar to O. semperi in most characters, bur differs in form of the eye, which is rounder and more reduced, in the shorter, broader pronotum, and in shape of the hind calcar of the male. Omoglymmius (sensu stricto ) hiekei new species Fig. 143 Type material. - HOLOTYPE male, labelled: “Philippinen, Luzon, Dalbalan (underside label), S. Boettcher, 3-1-17” (MNHB). PARATYPES one female, same data as holotype (MNHB; one male, same locality and collector as holotype (underside label), dated 1-1917 (MNHB); one female, labelled “Mt. Makiling, Luzon, Baker” (MCZ). Description. — Length 6. 0-6. 5 mm. Antennal Segments 1, II punctate; Segments III-XI impunctate; head about 1.5 longer than wide; median lobe narrow, lance-shaped, its apex obtusely pointed; frontal space large, U-shaped, its lateral margin curved; medial angles obtuse, slightly separated; posteriomedial margin oblique, slightly emarginate; posteriolateral margin evenly curved; occipital angles distinct though slight; temporal lobe with orbital groove extended as irregular row of punctures; one temporal seta; postorbital, suborbital tubercles absent; eye large, round. Pronotum long; length/greatest width 1.40; widest near middle; both ends narrowed; apex slightly more so than base; lateral margins curved; lateral margin scarcely sinuate anterior to hind angle; carinae subequal in width at middle; medial margin of outer carina slightly sinuate anterior to base; outer carina widest at middle, narrowed to base and apex; inner carina narrowed just anterior to base; latter broadly truncate; outer carina impunctate or with one or two minute punctures near base; inner carina impunctate; pronotum without setae; prosternum without precoxal carinae. Elytron relatively long, narrow; striae impressed, coarsely punctate; base of Stria IV without longitudinal scarp; Stria II with two setae near apex; Stria IV with five setae in continous series; subapical Stria with one seta; apex of stria VII with several setae; metasternum with coarse punctures near margins and midline, and few scattered on disc; punctures of abdominal Sterna III-V sparse, in form of irregular transverse row on each sternum; female with lateral pits of Sternum IV deep, round; male with very small, obtuse ventral tooth on anterior femur; middle calcar acute, prominent, nearly as long as spur; hind calcar triangular, its tip obtuse, slightly lobate. This species is characterized by the absence or near absence of punctures from the pronotal carinae. It is most similar to O. imugani and O. politus , but the latter two species have six to 10 conspicuous punctures on the outer carina. In addition, O. imugani has very few setae on the elytra, while O. politus has slightly reduced eyes and the male lacks the ventral tooth on the anterior femur. We dedicate this species to Dr. Fritz Hieke, of the Museum fur Naturkunde der Humboldt-Universitat, in token of his help in this work, in arranging for us to borrow the historically important rhysodid collection of his museum. Omoglymmius ( sensu stricto) pect oralis new species Figs. 144, 149 Type material. - HOLOTYPE female, labelled: “JAVA, Mt. Smetou” (label is handwritten, and spelling of place name may be inaccurate)(MNHN). Description. — Length 7.0 mm. Antennal Segments I- 1 1 1 coarsely punctate; Segments IV-XI impunctate; head 1 .25 longer than broad; median lobe short, oval, its apex obtusely pointed; frontal space rather narrow, its sides curved; medial angles nearly rectangular, contiguous; posteriomedial margin slightly curved; posteriolateral margin evenly curved; occipital angle indistinct; orbital groove absent; temporal lobe impunctate; four temporal setae; antennal lobe elevated above anterior end of temporal lobe: postantennal groove represented by broad pollinose scarp; postorbital, suborbital Revision of Omoglymmius Ganglbauer 219 tubercles absent; eye large, round. Pronotum rather short, length/greatest width 1.19; subquadrate, widest near middle; base, apex scarcely narrowed; lateral margins only slightly curved; lateral margin not sinuate anterior to hind angle; outer carina about 0.6 as wide as inner one at middle; inner carina only slightly narrowed anterior to base; latter broadly truncate; both pairs of carinae impunctate; marginal groove dilated; pronotum without setae; prosternum with sharp precoxal carina on each side, extending 0.75 of distance from coxa to anterior margin (Fig. 149). Elytron relatively long, narrow; striae impressed, coarsely punctate; base of Stria IV without longitudinal scarp; Stria IV with two setae near base and two near apex; subapical stride with 1 seta; Stria VII with several setae near apex; metasternum impunctate except for row along each lateral margin; abdominal Sterna 1II-V each with coarse punctures tending to form transverse row; female with deep, reniform lateral pit on Sternum IV; male unknown. This species resembles O. thoracicus, also of Java, in having precoxal carinae, but is otherwise very different. In dorsal aspect it resembles O. quadruplex of Mindanao, but the pronotum is more quadrate, and the anterior ridge of the temporal lobe less extensively pollinose. Omoglymmius (sensu stricto ) quadruplex new species Fig. 145 Type material. - HOLOTYPE male, labelled: “P.I., Kabasalan, Zamboanga, 20 ft., III-9-32, coll. H.C. Muzzell, F.C. Hadden Colin.” (BPBM). PARATYPES seven males, one female, same data as holotype (BPBM); two males, one female, same data as holotype, no collector cited (BPBM); one male, three females, same locality as holotype, V-4-1932, no elevation given (CAS); one male, same locality as holotype, F. C. Hadden Colin., no collector cited, VI- 1-32 (CAS). Description. — Length 5.2-7. 1 mm. Antennal Segments I-V sparsely, coarsely punctate; Segments VI-XI impunctate; head slightly longer than broad; median lobe short, oval, its tip obtusely pointed; frontal space rather narrow, its sides curved; medial angles rectangular, slightly separated; posteriomedial margin slightly emarginate; posteriolateral margin evenly curved; occipital angle indistinct; antennal ridge elevated above level of temporal lobe; latter forming narrow, pollinose ridge anteriorly; temporal lobe impunctate; orbital groove broad, extended beyond middle of eye; temporal setae four in most specimens (three present in a few specimens); two pairs of gular setae present; postorbital, suborbital tubercles absent; eye large, round. Pronotum rather broad, length/greatest width 1.23; widest near middle; base moderately narrowed, apex more narrowed; lateral margins curved; lateral margin not sinuate anterior to hind angle; outer carina slightly broader than inner one at middle; medial margin of outer carina sinuate anterior to base; inner carina constricted near base, latter broad, truncate; both pairs of carinae impunctate; marginal grooves not dilated; pronotum without setae; prosternum without precoxal carinae. Elytron relatively long, narrow; striae impressed, coarsely punctate; basal margin of elytron completely pollinose medial to base of Stria IV; base of latter without longitudinal scarp; Stria I with one to five setae near apex; Stria II with one seta near apex, Stria IV with one seta near base, one or two setae near apex; subapical stride with one seta; apex of Stria VII with several setae; metasternum coarsely, sparsely punctured near lateral margins and midline; disc otherwise impunctate; abdominal Sterna II I-V each with punctures in form of transverse row; lateral portions of Sterna 1 1 I-V I with punctures confluent, forming irregular transverse impressions; lateral pit of Sternum IV of female scarcely distinct; anterior femur of male with prominent, acute, ventral tooth; middle calcar acute, nearly as long as spur; hind calcar broadly rounded, raised well above spur. This species, O. pectoralis and O. duplex form a well-marked group with more than one temporal seta, the head broad, and the glabrous part of the temporal lobe separated from the antennal ridge by an elongate, pollinose ridge. From the first species, O. quadruplex is separated by the absence of precoxal carinae; from the second, by the greater number of temporal setae, the less extensive orbital groove, the extragular setae, several setae on the apex of Stria I, pollinosity on the basal margin medial to Stria IV. Range. - Mindanao in the Philippines. In addition to the type material we have seen the following specimens from eastern Mindanao: one male, Lawa, Davao Prov., IV-24-30. coll. C. F. Clagg, F. Psota Colin, “at light”(CNHM); two males, Madaum, Tagum, Davao Prov., sea level, F. G. Werner leg., CNHM Philip. Zool. Exp. 1946-47 (CNHM). Quaest. Ent., 1982, 18 (1-4) 220 Bell and Bell Omoglymmius (sensu stricto) duplex new species Fig. 146 Type material. - HOLOTYPE male, labelled: “Mt, Makiling, Laguna, P.I., 1-1-1931, F. C. Hadden Colin.” (CAS). PARATYPES three males, six females, same data as holotype (CAS); the following paratypes all have the same locality data as the holotype but with these dates: one female, IV- 1 - 193 1 ; one female, IV-24-1931; one male, VI-1-31; one female, VI-19-31; one male, one female, VI-29-31; one male, 3-3-32, 4000 ft., one male, II 1-5-32; two males, four females, IV- 1-32; two males, three females, IV- 1 -32, 400 ft.; one male, one female, V-5-32, 800 ft.; one male, VII- 1-32, 400 ft. (all CAS); one male, one female, same locality data as holotype, IV-6-32, 500 ft. (BPBM); one female, same locality data as holotype, VI-1-32, 500 ft. (BPBM); two females, Mt. Makiling, Luzon, coll. C. F. Baker (NMNH); one male, two females, same data as previous specimens, July 1923 (NMNH); one male, one female, Mt, Makiling, Luzon, P.I., coll. H. C. Muzzall, F. C. Hadden Colin. ,B. Malkin Colin., XII-6- 1931 (CNHM). Description. — Length 5. 7-7. 8 mm. Antennal Segments I- IV sparsely, coarsely punctate; Segments V-Xl impunctate; head slightly longer than broad; median lobe short, oval, its tip obtusely pointed; frontal space rather broad, U-shaped, its lateral margins strongly curved; median angles obtuse, contiguous; posteriomedial margin oblique; posteriolateral margin evenly curved; occipital angle relatively distinct; antennal ridge elevated above level of temporal lobe; anterior end of latter forming narrow, pollinose ridge; temporal lobe impunctate or with few minute punctures; orbital groove represented by broad, rather irregular band of pollinosity which reaches posterior margin of temporal lobe; temporal setae two in most specimens three or one in a few specimens; one pair gular setae present; postorbital, suborbital tubercles absent; eye large, round. Pronotum moderately elongate, length/greatest width 1.31; widest near middle; base moderately narrowed; apex more narrowed; lateral margins curved; lateral margin not sinuate anterior to hind angle; medial margin of outer carina sinuate anterior to base; carinae subequal at middle; inner carina constricted at base, latter broad, truncate; outer carinae impunctate in most specimens (four to six punctures in the specimen from Mindanao); marginal grooves not or but slightly dilated; pronotum without setae; prosternum without precoxal carinae. Elytron relatively long, narrow; striae impressed, coarsely punctate; basal scarp of elytron not pollinose; base of Stria IV with small longitudinal pollinose scarp; setae absent from apex of Stria I; Stria 1 1 with one seta near apex; Stria IV with one to three near apex, but none near base; subapical stride with one seta; apex of Stria VII with several setae; metasternum coarsely, sparsely punctate; abdominal Sterna III-V with coarse, sparse punctures; lateral portions of Sterna I1I-VI with punctures confluent, in form of irregular impressions; lateral pit of Sternum IV of female scarcely distinct; anterior femur of male with prominent, acute, ventral tooth; middle calcar acute, about 0.5 as long as spur; hind calcar triangular, very obtuse. This is the commonest and most widely distributed species in the Philippines. The broad head, impunctate temporal lobes, the pollinose ridge connecting the antennal rim with the temporal lobe, and, in most specimens, two temporal setae, will separate it from O. philippensis , the other widespread species in the islands. The most similar is O. quadruplex. The latter has more temporal setae, extragular setae, several setae on the apex of Stria I, a broader pronotum, and a reduced orbital groove. Range. - Philippines (Luzon, northern Palawan, Sibuyan Island and one specimen from northern Mindanao). This latter specimen is different from all others in that it has four to six punctures on the outer carina. In all other respects it seems to be O. duplex , though it is possibly not conspecific. In addition to the type locality of Mt. Makiling, we have seen Specimens from the following localities: Luzon: one female, Alabang, Rizal Prov., XI I- 1945, B. Malkin Colin. (NMNH); two females, S.E. Bataan, July-Aug. 1945, coll. Darlington (MCZ); one male, one female, Mt. Bulasan, Okt., 1917, leg Bottcher (MNHB); one male, two females, Dingalan Bay, June-Aug. 1945, coll. Darlington (MCZ); five females, Imugin, N. Viscaya, Baker (NMNH); ten females, Imugan, May, 1916, May & June 1917, leg. Bottcher (MNHB); two males, two females, Los banos, March, July, 1914, leg. Bottcher (MNHB); one female, Los Banos, 8-9, 2-914, Bottcher (MNNH); one male, Los Banos, Colin. Baker (NMNH); one male, Los Banos, Colin. Baker (MCZ); one female, Los Banos, May-June, 1947, 2nd growth, 100-1500 ft. F. G. Werner leg. (CNHM); two males, one female, Malinao, Tayabas, Colin. Baker, 5989 (NMNH); one male, Manila (MNHN); one female, Massisiat, Abra Prov., Revision of Omoglymmius Ganglbauer 221 V-1946, 3500 ft., CNHM Zool. Exp. 1946-47 (CNHM); one female, Montalvan, E. Simon (MNHN); one female, Ripang, 11-1918, Bottcher, Colin, Jul. Moser (MNHB); one male, three females, Zambales Prov., X 1 1- 1 9 1 7, Bottcher (MNHB); MINDANO: one female, Surigao, 4-2-1915, leg. Bottcher (MNHB); PALAWAN: one male, two females, Binaluan, 13-1-1914, leg. Bottcher (MNHB); SIBUYAN I.: one male, Colin. Baker (NMNH). Variation. - The specimens from Mindanao and Sibuyan are possibly not conspecific with those of Luzon. The Mindanao specimen has four to six punctures on the outer carina; that from Sibuyan differs from all others seen by us in having a ventral tooth on the hind femur of the male (in addition to the usual one on the front femur). Omoglymmius (sensu stricto) bouchardi new species Fig. 150 Type material. - HOLOTYPE female, labelled: “SUMATRA Palembang, ty-Grouv” (MNHN). PARATYPES two females, same locality data (but not cited as “ty”) (MNHN); one female, same locality data as holotype, labelled also “ty-Grouv” (LEI); one female, same locality data as holotype, labelled “Grouvelle, Fry Colin. 1905-100” (BMNH); one male; labelled “Sumatra” (MNHN). The “ty” labels probably were intended to designate type specimens. Grouvelle did not describe this species. Description. — Length 6. 0-6. 8 mm. Antennal Segments 1-IV sparsely, coarsely punctate; Segments V-XI impunctate; head 1.25 longer than broad, median lobe oval, its tip broadly rounded, its sides slightly parallel; frontal space rather broad, U-shaped, its lateral margins markedly curved; median angles obtuse, narrowly separated; posteriomedial margin curved; posteriolateral margin evenly curved; occipital angle indistinct; antennal lobe separated from temporal lobe by moderately wide antennal groove; temporal lobe with 15-20 rather fine punctures in posteriolateral portion; orbital groove broad, extending to posterior margin of eye; one temporal seta; postorbital, suborbital tubercles absent; eye large, round. Pronotum moderately broad, length/greatest width 1.21; widest near middle, base moderately narrowed; apex more strongly narrowed; lateral margin sinuate anterior to hind angle; medial margin of outer carina sinuate anterior to base; inner carinae slightly broader than outer carinae at middle; inner carina constricted near base; latter broadened, truncate; outer carina impunctate or with one or two punctures; marginal grooves slightly dilated; pronotum without setae; prosternum without precoxal carinae. Elytron relatively long, narrow; striae impressed, coarsely punctate; basal scarp of elytron glabrous except for longitudinal scarp at base of Stria IV and pollinose spot at base of Stria 11; Stria II with one seta near apex; Stria IV with one seta near apex; subapical stride with one seta; Stria VII with several setae near apex; metasternum coarsely punctate; abdominal Sterna II1-V with pollinose transverse sulci separated at midline; sternal punctures incorporated by sulci; lateral pit of Sternum IV in female shallow; front femur of male with prominent, acute ventral tooth; middle calcar acute, about 0.5 as long as spur; hind calcar triangular, its apex rounded, slightly lobate, its dorsal margin obtusely angulate. This species and O. consors are Sumatran species with pronotal carinae impunctate or nearly so, median lobe of head narrow and truncate, and abdominal sterna with the punctures confluent, in form of transverse sulci. O. consors differs in having the basal margin of the elytron entirely pollinose, and antennal lobes more markedly elevated above the temporal lobes. The remaining species from Sumatra have the outer carinae punctate. Grouvelle had intended to name this species in honor of M. Bouchard, who brought him the specimens, and whose observations on Rhysodidae in the dead limbs of freshly felled trees in the Sumatran forest were cited by Grouvelle (1903). We are pleased to carry out his intentions. Range. - Sumatra. In addition to the type material we have seen two males, labelled “Z.O. Serdang, Sumatra’s O.K. ± 1000', Dec. 89, 1.Z. Kannegieter” (MNHN). Omoglymmius (sensu stricto ) consors new species Fig. 151 Type material. - HOLOTYPE male, labelled: “SUMATRA, Palembang” (MNHN). PARATYPES three males, two females, same data as holotype (MNHN). Quaest. Ent., 1982, 18 (1-4) 222 Bell and Bell Description. — Length 5. 3-6. 5 mm. Antennal Segments I-Ill faintly punctate; Segments 1V-X1 impunctate; head 1 .5 longer than broad; median lobe parallel-sided, its tip broadly rounded; frontal space broad, U-shaped, its lateral margins markedly curved; medial angles obtuse to rounded, narrowly separated; posteriomedial margin curved; posteriolateral margin evenly curved; occipital angle indistinct; antennal lobe raised above level of temporal lobe, connected to latter by pollinose ridge; temporal lobe with three to six fine punctures; orbital groove complete though narrow; one temporal seta; postorbital, suborbital tubercles absent; eye large, round. Pronotum rather elongate, narrow; length/greatest width 1.36; widest near middle, base slightly narrowed, apex strongly narrowed; lateral margins nearly parallel; margin not sinuate anterior to hind angle; medial margin of outer carina markedly sinuate anterior to base; inner carina slightly broader than outer one at middle; inner carina constricted near base, latter truncate but only slightly broadened; both carinae impunctate; marginal grooves not dilated; pronotum without setae; prosternum without precoxal carinae. Elytron relatively long, narrow; striae impressed, coarsely punctate; basal margin of elytron entirely pollinose, base of Stria IV concealed, thus longitudinal scarp not evident; Stria II with one or two setae near apex; Stria IV with one or two setae near apex; subapical striole with one seta; Stria VII with several setae near apex; metasternum coarsely punctate; punctures of abdominal Sterna III-V in transverse sulci contiguous across midline or only narrowly interrupted there; lateral pit of Sternum IV in female round, deeper than in O. bouchardi\ front femur of male with prominent acute ventral tooth; middle calcar acute, about 0.5 as long as spur; hind calcar triangular, its apex obtuse, its dorsal margin not angulate. This species differs from all others in presence of a complete band of pollinosity on the basal transverse scarp of the elytron. Otherwise, it is rather similar to O. bouchardi, from which it also differs in having the pronotum more elongate and parallel-sided, and transverse sulci of the abdomen at most narrowly interrupted. In Borneo, completely impunctate outer carinae and the transverse sulci of the abdominal sterna separate it from O. fraudulentus and O. nemoralis. The name was chosen by Grouvelle, who died without publishing a description. Range. - Sumatra, Borneo. In addition to the type material we have seen one male, labelled: “Borneo Occ. Setipas pres Singawang, J. B. Ledru 1897” (MNHN). Omoglymmius { sensu stricto) repetitus new species Figs. 155, 156 Type material. - HOLOTYPE female, labelled: “Nord-Celebes, Toli-toli, Nov.-Dez. 1895, H. Fruhstorfer” (MNHN). Description. — Length 6.9 mm. Antennal Segments I-IV coarsely punctate; outer segments with punctures successively finer; those of X scarcely visible; Segment XI impunctate; head slightly longer than wide; median lobe broad, lance-shaped, its apex very obtuse; frontal space broad, U-shaped, its lateral margins markedly curved; medial angles rounded, well separated; posteriomedial margin curved into posteriolateral margin; latter curved most markedly opposite temporal seta; occipital angle absent; antennal lobes raised above level of temporal lobes, these and slope connected to temporal lobe pollinose; posterior margin of temporal lobe narrowly fringed by pollinosity; orbital groove absent; temporal lobe with about 12 very fine punctures near lateral margin; one temporal seta; small postorbital tubercle present; eye large, round. Pronotum rather short, length/greatest width 1.15; widest near middle; base slightly narrowed; apex more markedly narrowed; lateral margins slightly curved, markedly so near apex; margin shallowly sinuate anterior to hind angle; inner carina slightly broader than outer one at middle; medial margin of outer carina shallowly sinuate anterior to base; outer carina widest anterior to middle; narrowed to both base and apex; inner carina narrowed posteriorly, its base obliquely truncate; outer carina with seven or eight fine punctures; inner carina with about three fine punctures; pronotum without setae; prosternum without precoxal carinae. Elytron relatively long, narrow; striae impressed, coarsely punctate; base of Stria IV with very small, pollinose, longitudinal scarp; Stria IV with one seta near apex; subapical striole with one seta; Stria VII with several setae in apex; metasternum coarsely punctate; abdominal Sternum III with transverse row of punctures, IV, V with transverse bands of coarse punctures, confluent laterally on IV, V, VI; female with deep round lateral pit on Sternum IV; female with small, very obtuse ventral tubercle on anterior femur; male unknown. Pollinosity of antennal lobes is characteristic of this species. Presence of a small postorbital tubercle separates it from all species of Borneo and the Philippines, and from all other species from Wallacea except O. quadraticollis. The latter species has glabrous antennal lobes and a differently-shaped pronotum. Several species from New Guinea are superficially similar in having very small postorbital tubercles, but differ in having the antennal lobes glabrous, and Revision of Omoglymmius Ganglbauer 223 striae not impressed. Omoglymmius (sensu stricto) quadraticollis (Arrow) Figs. 152, 157 Rhysodes quadraticollis Arrow 1901: 87. Omoglymmius ( sensu stricto ) quadraticollis (Arrow) Bell and Bell 1978. Type material. - LECTOTYPE (here designated) male, labelled: “Tenimbar I., Doherty, 1900-150” (BMNH). PARALECTOTYPES two females, same data as lectotype (BMNH). The locality, usually spelled “Tanimbar”, is a group of islands between Timor and New Guinea. Description. — Length 5. 2-7.0 mm. Antennal Segments I-X punctate; Segment XI impunctate; head slightly longer than wide; median lobe broad, nearly circular, its apex broadly rounded; frontal space broad, U-shaped, its margins markedly curved; medial angles broadly rounded, nearly contiguous, posteriomedial margin curved into posteriolateral margin; latter more evenly curved than in O. repetitus\ occipital angle absent; antenal lobes glabrous, raised above level of temporal lobes, slope between antennal and temporal lobes broadly pollinose; posterior margin of temporal lobe narrowly fringed by pollinosity; orbital groove represented by broad pollinose band tapered to point near posterior margin of eye; temporal lobe with four to seven fine punctures mostly anterior to temporal setae; one temporal seta; small postorbital tubercle present, visible only in profile view; eye large, round. Pronotum rather short, length/greatest width 1.15; widest distinctly behind middle; base not narrowed; apex slightly narrowed; pronotum nearly rectangular, lateral margins nearly straight, slightly convergent anteriorly; margin not sinuate anterior to hind angle; carinae subequal in width at middle; medial margin of outer carina scarcely sinuate anterior to base; outer carina of nearly even width except at rounded apex; inner carina narrowed posteriorly, its base narrowly truncate; outer carina with 10-17 fine punctures; inner carina with about three fine punctures; pronotum without setae; prosternum without precoxal carinae. Elytron relatively long, narrow; striae impressed, coarsely punctate; transverse basal scarp of elytron pollinose; longitudinal basal scarp of Stria IV at most merely suggested; Stria IV with one seta near apex; subapical stride with one seta; Stria VIII with several setae near apex; metasternum finely, nearly uniformly punctate; abdominal Sterna II1-V with coarse punctures in form of irregular transverse row on each sternum; both sexes with lateral pit in Sternum IV, deeper in female; male with obtuse ventral tooth on anterior femur; middle calcar acute, very small, less than 0.25 of length of spur; hind calcar nearly triangular, its apex acute, only slightly above level of spur, its dorsal margin indistinctly angulate. This species is characterized by its almost rectangular pronotum, which is widest near the base. A small postorbital tubercle separates it from all other species of Wallacea except for O. repetitus of Celebes. The latter does not have a rectangular pronotum, and also has the median lobe pointed posteriorly. Omoglymmius (sens u stricto) batchianus (Arrow) Figs. 153, 158 Rhysodes batchianus Arrow 1901: 86. Omoglymmius ( sensu stricto) batchianus (Arrow) Bell and Bell 1978. Type material. - HOLOTYPE female, labelled: “Batchian, Pascoe Coll. 93-60” (BMNH). Description. — Length 7. 0-7.2 mm. Antennal Segments I-X punctate. Segment I coarsely so, punctures finer on more distal segments, those of Segment X extremely fine; Segment XI impunctate; head slightly longer than broad; median lobe broad, its tip obtusely pointed; frontal space broad, U-shaped, its margins markedly curved; medial angles rounded, well separated; posteriomedial margin curved into posteriolateral margin; latter evenly curved; occipital angle absent; antennal lobe raised above level of temporal lobe, slope between them broadly pollinose; orbital groove nearly absent, represented by narrow strip of pollinosity medial to anterior margin of eye; temporal lobe with three to five fine punctures between vestige of orbital groove and temporal seta; one temporal seta; postorbital, suborbital tubercles absent; eye large, round. Pronotum rather short, broad; length/greatest width 1.10; widest slightly behind middle; base very slightly narrowed; apex distinctly narrowed; lateral margins nearly parallel in basal half, more curved, convergent in apical half; margin not sinuate anterior to hind angle; inner carinae slightly wider than outer ones at middle; medial margin of outer carina not sinuate anterior to base; outer carina of nearly even width except at slightly narrowed extreme apex; inner carina widest at Quaest. Ent., 1982, 18 (1-4) 224 Bell and Bell basal third, narrowed anteriorly, posteriorly, base obtuse, no trace of constriction anterior to base; outer carina with 20-25 very fine punctures; inner carina with four to six fine punctures; pronotum without setae; prosternum without precoxal carinae. Elytron rather short, striae shallowly impressed, finely punctate; base of Stria IV with pollinose longitudinal scarp; Stria IV with one seta near apex; subapical stride with one seta; Stria VII with several setae near apex; metasternum coarsely punctate; abdominal Sterna III-V each with transverse band of coarse punctures; Sterna 111-V1 with punctures confluent laterally in form of irregular transverse sulci; Sternum IV in both sexes in form of deep, round lateral pit; male with obtuse ventral tooth on anterior femur; female with vestigial tooth or angle on anterior femur; middle calcar obtuse, very small, less than 0.25 as long as spur; hind calcar nearly triangular, its apex obtuse, only slightly above level of spur, its dorsal margin sinuate. This is a species with a broad, almost quadrate pronotum and shallow striae with fine punctures. It is close to O. humeralis of Ternate, but the latter species has the lateral margins of the pronotum more strongly curved. Shallow elytral striae, fine punctures, and tapered bases of inner carinae are points of resemblance to several species from New Guinea. Of these, the closest are O. fringillus and O. oroensis, both of which lack postorbital and suborbital tubercles. In these species, the glabrous part of the temporal lobe is much more broadly separated from the antennal lobe. Range. - Probably restricted to Batjan (also spelled “Batchian” and “Bacan”, a group of islands in the North Moluccas. Neither Arrow nor Grouvelle adequately compared this species with O. humeralis (Grouvelle) of Ternate and Halmahera. Differences in shape between pronota of the two appear to be equal to specific differences in other groups of Omoglymmius s. str ., but confirmation of their distinctness will have to await the collection of more material. In addition to the holotype, we have studied one male, labelled “Batjan, Bernstein” (LEI). Omoglymmius (sensu stricto) humeralis (Grouvelle) Fig. 154 Rhysodes humeralis Grouvelle 1895a: 157. Omoglymmius ( sensu stricto) humeralis (Grouvelle) Bell and Bell 1978. Type material. - LECTOTYPE (here designated) female, labelled: “MOLUQUES, Ternate, Raffray et Maindron 78” (MNHN). PARALECTOTYPE female, labelled: “MOLUQUES, Gilolo, Raffray & Maindron 78” (MNHN). The original description gave the locality as “Celebes”, but Grouvelle later (1903), listed it as “Ternate”. For this reason, we chose the Ternate specimen as lectotype. “Celebes” perhaps referred to a political unit, and not the island. “Gilolo” is an old name for Halmahera, the largest island in the North Moluccas. The specimens from the two islands are not identical, and are quite possibly specifically distinct, although this is difficult to judge on the basis of one specimen from each island. The description below is based entirely on the lectotype. Points of difference of the paralectotype are listed below, under the heading of “variation.” Description. — Length 6.5-7. 1 mm. Antennal Segments I- IX punctate; Segment I coarsely so; punctures finer distally, those of Segment IX very fine; Segments X, XI impunctate; head slightly longer than broad; median lobe broad, rather long, its tip obtusely pointed; frontal space very broad, more than twice as broad as long; U-shaped, its sides markedly curved; median angles obtuse, well separated; posteriomedial margin curved into posteriolateral margin; latter evenly curved; occipital angle absent; antennal lobe raised above level of temporal lobe, slope between them broadly pollinose; orbital groove nearly absent, represented by broad but very short strip of pollinosity opposite anterior margin of eye; temporal lobe with 12-14 very fine punctures in lateral half; one temporal seta; postorbital, suborbital tubercles absent; eye large, round. Pronotum moderately long, length/greatest width 1.37; widest near middle, base distinctly narrowed; apex more strongly so; lateral margins nearly evenly curved; margin not sinuate anterior to hind angle; inner carinae slightly wider at middle than outer carinae; medial margin of outer carina shallowly sinuate anterior to base; outer carina widest just anterior to middle; inner carina widest near middle, narrowed, obtuse anteriorly, narrowed to point posteriorly; no constriction near base of inner carina; outer carinae with 1 8-23 very fine punctures; inner carina with six or seven very fine Revision of Omoglymmius Ganglbauer 225 punctures; pronotum without setae; prosternum without precoxal carinae. Elytra rather short; stria shallowly impressed, finely punctate; base of Stria IV with pollinose longitudinal scarp; Stria IV with one seta near apex; subapical stride with one seta; Stria VII with several setae near apex; metasternum coarsely punctate; abdominal Sterna III-V each with sparse coarse punctures, in form of one irregular transverse row on each sternum; punctures of Sterna 1V-VI confluent laterally, in form of rudimentary transverse sulci (these less distinct than in O. batchianus)-, female with deep, round lateral pit on Sternum IV; female with minute, obtuse ventral tooth on anterior femur; male unknown. The most similar species to this one are O. batchianus, O. viduus and O. opticus. The first, from Batjan, has the pronotum with sides almost parallel; the second one, from the Kei Islands, has the pronotal margins more strongly curved, and the temporal lobe with more and coarser punctures, while the third, from Dammar, has the frontal space much narrower. Range. - In addition to the lectotype, we have studied the following specimens: two females, labelled: “Ins. Ternate, Doherty VII" (MCZ); two females, labelled “Ternate, Kannegieter (Museum Natura Artis Magistra)” (AMS). Variation. - As stated above, the paralectotype, from Halmahera, is rather different from the lectotype, and might represent another, unnamed species, possibly restricted to Halmahera. We do not wish to name it until more specimens from Halmahera are available. The length is 6.5mm. The principal points of difference from the specimens from Ternate are as follows: median lobe shorter, more broadly rounded posteriorly; frontal space smaller; medial angles more obtuse, more narrowly separated; orbital groove represented by a much narrower strip of pollinosity; temporal lobe with only about six minute punctures, these forming irregular row anterior to temporal seta; pronotum shorter, length/greatest width 1.18; inner carinae subtruncate, much less narrowed at base; fewer pronotal punctures (about 10 on outer carina; one to three on inner carina); basal scarp of Stria IV exceptionally long, its length almost twice distance from scarp to elytral humerus (in Ternate specimens it is shorter than distance to humerus). Omoglymmius (sensu stricto) opticus new species Fig. 159 Type material. - HOLOTYPE female, labelled: “Dammer Jnsel” (MNHB). PARATYPE female, same data as holotype. The label probably refers to the Damar Islands, which lie in the Banda Sea, northeast of Timor and northwest of Tanimbar. Description. — Length 6. 2-6. 7 mm. Antennal Segments I-X punctate, those of distal segments very fine; Segment XI impunctate; head slightly longer than wide; median lobe broad, lance-shaped, its apex very obtuse, nearly rounded; frontal space rather narrow, U-shaped, its lateral margins distinctly curved; medial angles very obtuse, slightly separated; posteriomedial margin evenly curved into posteriolateral margin; occipital angle absent; antennal lobe glabrous, separated from temporal lobe by broad band of pollinosity continued as very short orbital groove; latter ended opposite anterior 0.25 of eye; temporal lobe with 10-12 fine, scattered punctures; one temporal seta; postorbital, suborbital tubercles absent; eye large, round, more convex than in other Omoglymmius s. str., clearly visible in dorsal view. Pronotum rather short; length/greatest width 1.18; widest near middle; base, apex scarcely narrowed; pronotum nearly rectangular; lateral margins very feebly curved; margin not sinuate anterior to hind angle; inner carina slightly broader than outer carina at middle; medial margin of outer carina very shallowly sinuate anterior to base; outer carina widest anterior to middle; apex subtruncate; inner carina narrowed at base, slightly constricted anterior to base; outer carina with two to four fine punctures; inner carina with one puncture, or none; pronotum without setae; prosternum without precoxal carinae. Elytron relatively long, narrow; striae impressed, coarsely punctate; basal scarp pollinose; base of Stria IV with small pollinose longitudinal scarp; Stria IV with one seta at apex; subapical striole with one seta; Stria VII with several setae near apex; metasternum finely punctate; abdominal Sterna III-V with coarse, rather sparse punctures in form of broad band on each sternum; female with deep, round lateral pit on Sternum IV; male unknown. This species is closest in apperance to O. quadraticollis of the Tanimbar Islands. It resembles the latter in having the pronotum nearly rectangular but differs in lacking the postorbital tubercle, in having the pronotal carinae nearly impuctate: and in having the eyes very convex. Shape of the pronotum and very convex eyes separate it from O. batchianus and its relatives. Quaest. Ent., 1982, 18 (1-4) 226 Bell and Bell Omoglymmius ( sensu stricto) viduus new species Fig. 160 Type material - HOLOTYPE male, labelled: “Key Ins.” (MNHB). PARATYPES one male, one female, same data as holotype (MNHB); one female, labelled:“Key Inseln, coll. Plason, Coll. Reithoffer” (BSL). Description. — Length 6. 2-7. 2 mm. Antennal Segments I-X distinctly punctate; Segment XI impunctate; head slightly longer than wide; median lobe broad, lance-shaped, its apex obtuse; frontal space broad, U-shaped; its lateral margins markedly curved; medial angles acute, nearly contiguous; posteriomedial margin slightly sinuate; posteriolateral margin nearly evenly curved; occipital angle very obtuse; postantennal groove narrow; orbital groove abbreviated, scarcely extended to anterior margin of eye; temporal lobe with 20-25 rather fine punctures, scattered over its entire surface; one temporal seta; postorbital, suborbital tubercles absent; eye large, round. Prothorax rather long, length/greatest width 1.32; widest near middle; base slightly narrowed; apex more markedly narrowed; lateral margins curved; margin not or but slightly sinuate anterior to hind angle; inner carina slightly broader than outer one at middle; medial margin of outer carina not distinctly sinuate anterior to base; outer carina widest near middle, markedly narrowed to apex; inner carina with basal part narrowed parallel-sided; base truncate; outer carina with 21-27 fine punctures; inner carina with five to seven fine punctures; pronotum without setae; prosternum without precoxal carinae. Elytra relatively long, narrow; striae impressed, coarsely punctate; base of Stria 1 with longitudinal pollinose scarp; Stria IV with one seta near apex; subapical stride with one seta; Stria VII with several setae near apex; metasternum with coarse but very shallow punctures; abdominal Sterna 1I1-V with coarse, irregularly distributed punctures; these coalescent laterally to form indistinct transverse sulci; both sexes with deep, round lateral pit on Sternum IV; male with ventral tooth on anterior femur; middle calcar very small, indistinct; hind calcar triangular, its apex obtusely pointed; its dorsal margin slightly sinuate. This species differs from O. quadraticollis of the Tanimbar Islands in having the pronotum much more elongate and less quadrate. Numerous punctures of the temporal lobes and outer carina separate it from O. opticus of the Kei Islands. Deeper elytral striae and numerous punctures of the outer carinae separate it from O. batchianus and similar forms. Omoglymmius ( sensu stricto ) continuus new species Figs. 161, 166, 169, 170 Type material. - HOLOTYPE male, labelled: “Mangole (Sula Inseln), VII-XII, 1977, Wegener, DOY-K” (BSL). This is one of the islands east of Celebes, and lies southwest of Batjan and north of Buru. Description. — Length 6.9 mm. First segment of antenna with marked swelling around base of principal tactile setae (Fig. 169); Segments I-X punctate; punctures of distal segments very fine; Segment XI impunctate; head slightly longer than wide; median lobe oval, rather broad; its apex broadly rounded; frontal space broad, U-shaped; medial angles broadly rounded, nearly contiguous; posteriomedial margin evenly curved; posteriomedial margin broadly curved, margin more markedly curved at eye; occipital angle absent; antennal lobes pollinose, not sharply elevated above anterior end of temporal lobe; anterior end of latter extensively pollinose, boundary of pollinosity extending obliquely posteriomedially; pollinosity of orbital groove extending 0.75 of length of eye; temporal lobe with four to six very fine punctures; two or three very coarse punctures, probably each bear temporal seta (but only one present on either side in holotype, the other perhaps being broken off); postorbital, suborbital tubercles absent; eye large, round. Pronotum subquadrate; rather short, length/greatest width 1.17; widest near base; sides nearly parallel; apex slightly narrowed; lateral margin not at all sinuate anterior to hind angle; inner carina slightly wider than outer one; medial margin of outer carina not sinuate anterior to base; outer carina of nearly even width; inner carina constricted anterior to base; latter broad, truncate, outer carina with 11-12 very fine punctures; inner carina with nine to 1 1 very fine punctures; lateral margin of inner carina sloped gradually into paramedian groove; pronotum without seta; prosternum without precoxal carinae. Elytron moderately narrow; inner ( I- 1 1 1 ) striae finely punctate, very shallowly impressed; outer striae deeper, more coarsely punctate; base of Stria IV with very small longitudinal scarp; elytral setae absent except for several near apex of Stria VII; metasternum densly punctate; abdominal Sterna III-V densely punctate; male with deep, round lateral pit on Sternum IV; male with prominent ventral tooth on anterior femur; middle calcar acute, very small; hind calcar subtriangular, its apex truncate, raised above level of spur, its ventral margin emarginate; its dorsal margin angulate (Fig. 1 70); female unknown. Revision of Omoglymmius Ganglbauer 227 This species differs from other members of the subgenus in having the lateral margin of the inner carina sloped gradually into the paramedian groove, as in most species of Rhyzodiastes. The indistinct boundary between the antennal and temporal lobes is also unique. It differs sharply from O. wittmeri, the other species known from Mangole in these characters, and also in the subquadrate pronotum, in having the base of Antennal Segment I markedly swollen, in having temporal lobes scarcely punctate, and, apparently, in having more than one temporal seta. Omoglymmius ( sensu stricto ) wittmeri new species Figs. 162, 167, 168, 171 Type material. - HOLOTYPE male, labelled: “Mangole (Sula Inseln) VII-XII,1977, Wegener, DRP-K” (BSL). PARATYPES two females, same data as holotype but collected by V. & G. Wegener (BSL). Description. — Length 5. 3-6. 7 mm. First segment of antenna scarcely swollen at base of principal tactile seta (Fig. 168); Antennal Segments I-X punctate; punctures of distal segments very fine; Segments XI impunctate; head slightly longer than wide: median lobe lance-shaped, its apex very obtuse; frontal space broad, U-shaped; medial angles obtuse, narrowly separated; posteriomedial margin oblique; posteriolateral margin evenly curved; occipital angle indistinct; antennal lobe glabrous, raised above level of temporal lobe; anterior end of temporal lobe extensively pollinose, boundary of pollinosity extended obliquely posteriomedially; pollinosity of orbital groove extended posterior to eye, merged with narrow fringe of pollinosity along posterior margin of temporal lobe; temporal lobe with 12-18 rather fine punctures; one temporal seta; postorbital, suborbital tubercles absent; eye large, round. Pronotum rather short; length/greatest width 1.14; widest slightly anterior to middle; base moderately narrowed; apex more markedly so; lateral margins curved; lateral margin scarcely sinuate anterior to hind angle; carinae subequal at middle of length; medial margin of outer carina shallowly sinuate anterior to base; outer carina widest at anterior third; narrowed both at base and apex; inner carina constricted just anterior to base; latter truncate, scarcely broadened; outer carina with 18-25 punctures; inner carina with four or five fine punctures; medial margin of outer carina separated from paramedian groove by sharp, distinct scarp; pronotum without setae; prosternum without precoxal carinae. Elytron narrow, elongate; elytral striae impressed, coarsely punctate; base of Stria IV with longitudinal scarp; Stria IV with one seta near apex; subapical stride with one seta; Stria VII with several setae near apex; metasternum with dense, coarse but shallow punctures; abdominal Sterna III-V densely punctate; punctures near lateral margin of III, V, VI coalescent; both sexes with deep round lateral pit on Sternum IV; male with prominent ventral tooth on anterior femur; female with vestige of anterior femoral tooth; middle calcar small, triangular, about 0.3 as long as spur; hind calcar triangular, its apex slightly acute, at level of spur, its distal margin straight (Fig. 171). The glabrous, sharply defined antennal lobe, rounded pronotal margins, sharp outer boundary of the inner carina, lack of a swelling on Antennal Segment I, and different shape of the hind calcar differentiate this species from the sympatric O. wittmeri. The temporal lobe has a continous pollinose margin extended from the postantennal groove to the posterior margin of the temporal lobe. This feature is not found in other Moluccan species, but is duplicated in a few species from other regions, such as O. fringillus of New Guinea and O. gurneyi of the Solomon Islands. The former differs in the shape of the head, while the latter has a nearly quadrate pronotum. We dedicate this species to Dr. Wittmer of the Basel Natural History Museum in gratitude for his help in this study. Omoglymmius ( sensu stricto) vadosus new species Fig. 163 Type material. - HOLOTYPE female, labelled: “Amboina,X-XI-07, coll. E. Muir” (BPBM). Amboina, now Ambon, is in the South Molluccas, south of Ceram. Quaest. Ent., 1982, 18 (1-4) 228 Bell and Bell Description. — Length 7.0 mm. Antennal Segments I-X punctate; punctures of distal segments very fine; Segment XI impunctate; head slightly longer than wide; median lobe broad, lance-shaped, its apex very obtuse; frontal space broad, transverse, U-shaped; margin abruptly curved, medial angles obtuse, very narrowly separated; posteriomedial margin oblique, posteriolateral margin nearly evenly curved; occipital angle distinct; antennal lobe raised above level of temporal lobe, separated from latter by rather broad pollinose strip; latter continuous with short but broad orbital groove, ended opposite middle of eye; temporal lobe with six to eight very minute punctures; one temporal seta (holotype has two coarse punctures on right temporal lobe, and perhaps a second seta was broken off); postorbital, suborbital tubercles absent; eye large, round. Pronotum moderately long, length/greatest width 1.26; widest near middle; base, apex narrowed; lateral margins markedly curved; margin scarcely sinuate anterior to hind angle; inner carina slightly broader than outer carina at middle; medial margin of outer carina not sinuate near base; outer carina widest anterior to middle; inner carina tapered to base; outer carina with 12-14 extremely minute punctures, visible only under high magnification; inner carina with two or three minute punctures; pronotum without setae; prosternum without precoxal carinae. Elytron long, rather narrow; striae shallowly impressed; punctures of inner striae rather fine, those of outer striae coarser; base of Stria IV with pollinose longitudinal scarp; Stria IV with one seta near apex; subapical stride without seta; Stria VII with several setae near apex; metasternum sparsely, shallowly punctate; abdominal sterna with shallow, pollinose, lateral transverse sulci, with one or two isolated punctures between the sulci on each sternum; female with deep lateral pit on Sternum IV; male unknown. O. vadosus is characterized by the extremely fine punctures on the head, pronotum, elytra and metasternum. The propleuron is minutely pollinose as are the transverse sulci of the abdomen. Omoglymmius (sensu stricto) morditus new species Fig. 165 Type material. - HOLOTYPE male, labelled: “Morotai Is., Dutch E. I., Sept. 1944, Darlington” (MCZ). This island is in the North Moluccas, northeast of Halmahera. Description. — Length 5.8 mm. Antennal Segments I, II coarsely punctate; III, IV impunctate, V-IX very minutely punctate (appearing impunctate except under high magnification); Segments X, XI impunctate; head slightly longer than broad; median lobe short, broad, its tip broadly rounded; frontal space very broad, U-shaped; its margins bent abruptly; medial angles obtuse, well separated; posteriomedial margin oblique; occipital angle indistinct; posterior margin of head nearly transverse, in form of obtuse angle with nearly straight lateral margin of temporal lobe; antennal lobe raised above level of temporal lobe, slope between them broadly pollinose; orbital groove absent; posterior margin of temporal lobe narrowly fringed by pollinosity; temporal lobe with about 12 fine punctures in posteriolateral half; one temporal seta; postorbital, suborbital tubercles absent; eye large, round. Pronotum rather short, length/greatest width 1.17; subquadrate; widest near middle, slightly narrowed to base and apex; lateral margins very slightly curved; margins not sinuate anterior to hind angle; inner carina subequal to outer carina at middle; medial margin of outer carina not sinuate anterior to base; outer carina of nearly even width except at extreme base, apex; inner carina widest anterior to middle; tapered to point at base; outer carina with 13-15 punctures; inner carina with three or four very fine punctures; pronotum without setae; prosternum without precoxal carinae. Elytron rather long, narrow; striae deeply impressed, coarsely punctate; base of Stria IV with very small longitudinal scarp; transverse basal scarp of elytron pollinose; Stria IV with one seta near apex; subapical stride with one seta; Stria VII with several setae near apex; metasternum coarsely, shallowly punctate; abdominal Sterna III-V very coarsely, sparsely punctate; punctures of each sternum tending to form transverse row; male with ventral tooth on anterior femur; middle calcar very small, obtuse; hind calcar triangular, apex obtuse; upper margin slightly sinuate; female unknown. Among Moluccan species, this one is most similar to O. batchianus in the subquadrate form of the pronotum, but O. morditus differs from all other Moluccan species in the subquadrate form of the head, with a distinct angle between the lateral and posterior margins of the temporal lobe. Omoglymmius ( sensu stricto ) nasalis new species Fig. 164 Type material. - HOLOTYPE female, labelled: “Kajeli, Boeroe” (MNHN). The island, now spelled “Buru”. is in the South Moluccas, south of Mangole, and west of Ceram and Revision of Omoglymmius Ganglbauer 229 Amboina. Description. — Length 7.7 mm. Antennal Segments I-X coarsely punctate; Segment XI impunctate; head nearly twice as long as broad; preocular portion produced; median lobe elongate, its margins nearly parallel; its apex slightly narrowed, rounded; medial angles obtuse, slightly separated; frontal space very broad, U-shaped, its margins markedly curved; medial angles rounded; posteriomedial margin oblique; posteriolateral margin evenly curved; occipital angle indistinct; antennal lobe far from eye; pollinose oblique ridge between antennal lobe, frontal space; deep oblique groove posteriolateral to ridge; surface of temporal lobe between oblique groove and eye pollinose, representing short but broad orbital groove; temporal lobe with four to six punctures near temporal seta; one temporal seta; postorbital, suborbital tubercles absent; eye large, round. Pronotum rather short, length/greatest width 1.18; widest near middle; base slightly narrowed; apex more markedly narrowed; lateral margins curved; margin distinctly sinuate anterior to hind angle; inner carinae slightly broader than outer carinae at middle; medial margin of outer carina angulate but not sinuate near base; outer carina of nearly even width except at extreme base, apex; inner carina widest near middle; broadly truncate anteriorly, constricted just anterior to broadened, truncate base; outer carina with 25-28 coarse punctures; inner carina with 12-13 coarse punctures; pronotum without setae; prosternum without precoxal carinae. Elytron rather elongate, narrow; striae impressed; coarsely punctate, becoming finely so near apex; base of Stria IV without longitudinal scarp; Stria IV without setae; subapical striole with one seta; Stria VII with several setae near apex; metasternum densely, rather finely punctate; abdominal Sterna III-VI finely, densely punctate; female with large, deep, round lateral pit in Sternum IV; male unknown. The elongate snout separates this species from all others from Wallacea except for O. bucculatus. The latter has pronotal setae, several temporal setae, and has the outer carinae more narrowed, and lacks the diagonal groove anterior to the eye. Omoglymmius (sensu stricto) bucculatus (Arrow) Fig. 172 Rhysodes bucculatus Arrow 1901: 88-89. Omoglymmius ( sensu stricto ) bucculatus (Arrow) Bell and Bell 1978. Type material. - LECTOTYPE (Here designated) female, labelled: “SUMBAWA Id. (Doherty) 1900-150” (BMNH). PARALECTOTYPES three females, one male, same data as type (BMNH). Description. — Length 5. 9-7. 4 mm. Antennal Segments I-X punctate; Segment I coarsely punctate; punctures becoming finer on distal segments, nearly obsolete on Segment X; Segment XI impunctata; head nearly twice as long as wide, preocular portion produced; median lobe elongate, gradually tapered posteriorly; apex obtuse; frontal space narrow, V-shaped, its margins oblique or slightly curved; medial angles obtuse, slightly to distinctly separated; posteriomedial margin slightly emarginate; posteriolateral margin evenly curved; occipital angle distinct; antennal lobe separated from anterior end of temporal lobe by narrow postantennal groove; small pit present anterior to eye; orbital groove absent; temporal lobe with nine to 15 moderately coarse punctures; one to three temporal seta; postorbital, suborbital tubercles absent; eye large, round. Pronotum various in length, length/greatest width 1.23 in short specimens, up to 1.33 in longer ones; widest near middle; base, apex markedly narrowed; lateral margins curved; margin not sinuate anterior to hind angle; outer carina 0.35 to 0.5 as wide as inner one at middle; lateral margins markedly curved; margin not sinuate anterior to hind angle; marginal groove dilated; outer carina with medial margin sinuate near base; inner carina widest near middle, constricted just anterior to base, latter broadened, truncate; outer carina with one to eight coarse punctures or none in addition to two large setiferous punctures; inner carina with one to 1 3 coarse punctures or none; two or three prominent setae on each outer carina; precoxal carinae absent. Elytron relatively long, narrow; striae impressed, coarsely punctate; base of Stria IV without longitudinal scarp; Stria II with one to three setae near apex or none; Stria IV one seta near base and one to two near apex; subapical striole without seta; Stria VII with several setae near apex; metasternum with coarse punctures near margins and in midline, otherwise impunctate; punctures of abdominal Sterna III-V coarse, in form of transverse row on each sternum; both sexes with deep lateral pit in Sternum IV, this larger in female; male with small acute ventral tooth on anterior femur; middle calcar acute, as long as spur; hind calcar subtriangular, apex obtuse or truncate, proximal margin angulate. Pronotal setae are unique in the subgenus. The species is also characterized by the elongate head, and, in most specimens, by more than one temporal seta. Range. - Restricted to the island of Sumbawa, in the Lesser Sundas. In addition to type material, we have Studied the following specimens: one male, one female, labelled: “B. Aroe Hassa, Quaest. Ent., 1982, 18 (1-4) 230 Bell and Bell Sumbawa 2-5000’ Doherty IX, X” (year not indicated)(MCZ); two males, two females, labelled: “Sumbawa, W. Doherty” (MNHN); two females, labelled: “Sumbawa”, collector not indicated (MNHN). Variation. - We are provisionally treating this as one variable species, but it is possible that two or more species are involved. Most of the specimens can be assigned to one or other of two morphs. The first, and most common, is characterized by having the mentum densely punctate, the pronotum more elongate, and the hind calcar narrowly truncate at apex. The lectotype belongs to this form. The second form has the mentum impunctate except for a few punctures near its base, the pronotum less elongate and the hind calcar with apex obtuse. One specimen (BMNH), however, agrees with the first form in the shape of the pronotum, but agrees with the second one in the absence of punctures on the mentum and in the shape of the hind calcar. Omoglymmius ( sensu stricto) bicarinatus new species Fig. 181 Type material. - HOLOTYPE male, labelled: “Ins. Jobi, N. Guinea (Doherty)” (MNHN). This island, now called “Yapen”, formerly spelled “Japen”, is one of the Schouten Islands, north of Geelfink Bay on the northwest side of New Guinea. Description. — Length 4.8 mm. Antenna with Segments I- 1 1 1 coarsely punctate; Segments IV-X very minutely punctate; Segment XI impunctate; head about 1.5 longer than wide; median lobe lance-shaped, its apex obtusely pointed; frontal space rather narrow, its margins weakly curved; medial angles obtuse, widely separated; posteriomedial margin emarginate; posteriolateral margin oblique, in form of obtuse angle with posterior margin, which is transverse; occipital angle distinct; prominent preorbital pit; orbital groove absent; temporal lobe with about 16 fine punctures; one temporal seta; postorbital, suborbital tubercles absent; eye large, round. Pronotum relatively short, narrow; length/greatest width 1.14; shape irregular and probably somewhat deformed in holotype; widest anterior to middle; base, apex narrowed; margins curved, margin not sinuate anterior to hind angle; outer carina slightly narrower than inner carina at middle; medial margin shallowly sinuate anterior to base; outer carina widest anterior to middle, narrowed to both base, apex; inner carina constricted just anterior to base, latter slightly broadened; outer carina with 10-11 fine punctures; inner carina with six or seven fine punctures; pronotum without setae; prosternum without precoxal carinae. Elytron relatively short, narrow; striae impressed, coarsely punctate; Stria IV with longitudinal pollinose scarp at base; Interval V forming narrow, elevated carina extended from humerus to subapical tubercle; Interval V bounded on both sides by strip of pollinosity; Stria IV with one seta at base and one at apex; subapical stride with one seta; Stria VII with several setae near apex; metasternum coarsely, densely punctate; abdominal Sterna 1II-V densely punctate; male with deep, round lateral pit on Sternum IV; male without ventral tooth on anterior femur; middle calcar acute, longer than spur; hind calcar triangular, small, shorter than spur, its apex acute. The elevated cariniform fifth interval is characteristic of this species. Absence of a ventral tooth on the anterior femur of the male is highly unusual; it is shared only with O. gracilicornis of New Guinea, O. politus of Luzon, and two species from the Caroline Islands. Omoglymmius ( sensu stricto ) caelatus Bell and Bell NEW COMBINATION Fig. 173 Omoglymmius caelatus Bell and Bell 1981: 57-58. Type material. - HOLOTYPE male, labelled: “PELELIU I., Palau Islands, West coast, 2 Feb. 1948, H. S. Dybas” (BPBM). PARATYPES one male, two females, labelled: “PELELIU I. Palau Islands, East coast, 27 Jan. 1948, H. S. Dybas” (BPBM); one female, labelled: “Palau Islands, Koror I., 17 Jan. 1948. H. S. Dybas” (BPBM). See Bell and Bell, 1981 for complete data. Description. — Length 5.8-7. 1 mm. Outer antennal segments faintly punctate; head slightly longer than wide; frontal grooves very narrow; one temporal seta; posteriomedial margin emarginate; postorbital, suborbital tubercles absent. Pronotum moderately long; length/greatest width about 1.28; outer carina with about 25-30 coarse punctures; inner carina impunctate or with one or two minute punctures; inner carina constricted just anterior to base; latter broadened; Revision of Omoglymmius Ganglbauer 231 marginal groove dilated. Elytron with longitudinal scarp at base of Stria IV; striae shallow, scarcely impressed; punctures fine; male with minute vestige of ventral tooth on anterior femur; middle calcar very small, obtuse; hind calcar with apex obtuse, dorsal margin obtusely angulate. For more complete description, see Bell and Bell (1981). This species differs from O. impletus of Yap in having the outer carinae densely punctate and from O. oceanicus of Kusiae, Ponape, and Palau in having the posteriomedial margin of the temporal lobe emarginate, in having more punctures on the outer carina and in a greater constriction anterior to the base of the inner carina. It is a rather nondescript species, superficially similar to O. philippensis and its relatives in having a marked contrast between the coarsely, densely punctate outer carina and an impunctate inner one. O. philippensis differs in having the striae markedly impressed and coarsely punctate, and in having the ventral tooth of the anterior femur prominent. O. patens of New Guinea is similar, but lacks the basal scarp on Stria IV. Members of the O. lindrothi complex of the Solomon Islands are also similar, but have the inner carina punctate. Range. - Known only from the Palau Islands in the western Carolines. Omoglymmius (sensu stricto) impletus Bell and Bell NEW COMBINATION Fig. 174 Omoglymmius impletus Bell and Bell 1981: 58-60. Type material. - HOLOTYPE male, labelled: “Yap group, Tomil Dist. Jul-Au 50, R. J. Goss” (BPBM). PARATYPES two males, three females, same data as holotype (BPBM). See Bell and Bell (1981) for complete data. Description. — Length 5.2-6. 1 mm. Outer antennal segments impunctate; head slightly longer than wide; frontal grooves very narrow; frontal space longer than broad, its margins slightly curved; posteriomedial margin emarginate, narrowly fringed with pollinosity; one temporal seta; postorbital, suborbital tubercles absent. Pronotum relatively short, length/greatest width 1.18; marginal groove dilated; outer, inner carinae entirely impunctate; inner carina acutely pointed at base. Elytron with longitudinal scarp at base of Stria IV; striae impressed; punctures fine, shallow; male without ventral tooth on anterior femur; middle calcar very small, obtuse; hind calcar triangular, its apex obtusely pointed. For a more complete description, see Bell and Bell (1981). Among the species from the Caroline Islands, this species is easily recognized by the entirely impunctate pronotal carinae and the acutely pointed base of the inner carina. This combination of characters also occurs in several species from New Guinea, such as O. planiceps. Such species lack the basal scarp on Stria IV, and many of them have either postorbital or suborbital tubercles. Range. - Known only from the Islands of the Yap Group, in the western Caroline Islands. Omoglymmius (sensu stricto) oceanicus Bell and Bell NEW COMBINATION Fig. 175 Omoglymmius oceanicus Bell and Bell 1981: 60-61. Type material. - HOLOTYPE male, labelled: “Kusiae, Mutunlik, 22 m., 1-31-53, J.F.G. Clarke” (BPBM). PARATYPES one male, two females, same locality and collector as holotype (BPBM); one female, Kusaie, Malem, 1 9-XII- 1 937, Teiso Esaki (BPBM). See Bell and Bell (1981) for complete data. Description. — Length 5. 0-6. 8 mm. Outer antennal segments scarcely punctate; head slightly longer than wide; frontal grooves rather narrow; frontal space slightly broader than long, V-shaped, its margins slightly curved; Quaest. Ent., 1982, 18 (1-4) 232 Bell and Bell posteriomedial margin curved nearly evenly into posteriolateral margin; occipital angle absent; one temporal seta; postorbital, suborbital tubercles absent. Pronotum moderately elongate; length/greatest width about 1.28; marginal groove narrow; outer carina with about 10-14 very fine punctures; inner carina impunctate; base of inner carina more broadly dilated than in O. caelatus. Elytron with longitudinal scarp at base of Stria IV; striae impressed, coarsely punctate; male with minute ventral tooth on anterior femur; middle calcar acute, very small; hind calcar with apex truncate, dorsal margin sinuate. This species differs from O. caelatus in having the outer carina more finely and sparsely punctate, the inner carinae more broadly truncate at the base, and the posterior margin of the temporal lobe evenly curved. Range. - Widely distributed in the Caroline Islands. Known from Kusiae, Ponape, and Palau. For complete data, see Bell and Bell (1981). The lindrothi complex This group consists of the following nine species: Omoglymmius lindrothi, O. modicus, O. rusticus, O. manni, O. regius, O. princeps, O. gurneyi, O. renutus, and O. scopulinus. These very similar allopatric forms correspond to the pignoris complex of Kaveinga and could also be regarded as subspecies of a single species. Since Rhysodini from continental landmasses are rarely differentiated into subspecies, we prefer to regard the forms from the Solomon Islands as distinct species. It is, nevertheless, convenient to describe the O. lindrothi complex as a whole before describing the individual species. Description. — Length 4. 9-7. 5 mm. Antennal Segments I-X punctate; I- 111 coarsely punctate; distal segments with punctures very fine; Segment XI impunctate; head slightly longer than broad; orbital groove absent or present; temporal lobe punctate; one temporal seta; postorbital, suborbital tubercles absent; eye large, round. Pronotal carinae subequal at middie of length; both outer, inner carinae punctate; pronotum without setae; prosternum without precoxal carinae. Elytron relatively long, narrow; base of Stria IV with longitudinal scarp; in most specimens with one seta at apex of Stria IV, one in subapical stride, one or both of these setae absent in some specimens; apex of Stria VII with several setae. Shape of temporal lobes, pronotum, and calcars, punctuation of temporal lobes, metasternum and abdominal sterna, and depth of stria and lateral pits of the abdomen vary among the species of this complex. Among species from outside the Solomon Islands, the most similar species is O. classicus of the Admiralty Islands. It resembles O. princeps in having the tip of the median lobe pointed, but has a much more quadrate pronotum, with the base much less narrowed. Omoglymmius (sensu stricto) lindrothi new species Fig. 176 Type material. - HOLOTYPE male, labelled: “SOLOMON IS. Guadalcanal, Kukum, 25-11-1963, P. Greenslade, 11.007, B.M. 1966-477” (BMNH). PARATYPES two males, same data as holotype (BMNH); one female, Kukum, 19-4-1963, 5078; one female, Kukum, 8-12-1962, 3047; two males, one female (on same pin), Kukum, 15-11-1962, 2898; one male, Honiara, Kukum, 20-2-62, 830; two males, one female (on same pin), Honiara, Kukum, 6-2-62, 669; one female, Honiara, Kukum, 4-11-61, 68;one male, Guadalcanal, Manna, 23-7-1962, 2507; two males (on same pin), Guadalcanal, Mt. Austen, 19-9-672, 1297 (BM 1963-5) (all the preceding specimens collected by P. J. M. Greenslade, B. M. 1966-477)(all BMNH); one female, labelled: “Guadalcanal, Tambalias, 30 km. W. Honiara, 22-V-64, sweeping, R. Straatman” (BPBM). This species is apparently confined to Guadalcanal. Revision of Omoglymmius Ganglbauer 233 Description. — Length 5. 0-6. 2 mm. Median lobe broad, its tip broadly rounded; frontal space broad, U-shaped, its lateral margin curved; medial angles rounded, slightly separated; posteriomedial margin only slightly rounded, nearly oblique; posteriolateral margin evenly curved; occipital angle obsolete; orbital groove represented by short but broad band of pollinosity ended opposite middle of eye; temporal lobe with about 20 fine punctures. Pronotum moderately long, length/greatest width 1.27; widest near middle; base slightly narrowed; apex distinctly narrowed; lateral margins almost straight except near apex; lateral margin not sinuate anterior to hind angle; medial margin of outer carina straight, not sinuate anterior to base; outer carina broadest well anterior to middle, evenly narrowed posteriorly; basal part of inner carina narrow, its sides parallel; outer carina with 20-27 fine punctures; inner carina with four to six very fine punctures. Elytral striae moderately impressed, not pollinose except for pollinosity within punctures; latter moderately coarse; female with Sternum IV of abdomen with lateral pit deep, semicircular; male with ventral tooth on anterior femur; female with small ventral tooth or tooth entirely absent; hind calcar obtuse, subtruncate. The subquadrate pronotum, with outer carina not at all sinuate near base and inner carina with base very narrow, are the marks of this species. Of species from nearby islands, O. modicus of Savo and Nggela, is closest in form of pronotum. However, this latter species has the pronotum shorter, its lateral margins more distinctly curved, and its frontal space broader. The notation “sweeping” on the specimen from Tambalis, is probably a mistake, as there are no other records of Rhysodini from sweeping vegetation. We dedicate this species to the memory of the eminent carabid taxonomist, Carl H. Lindroth. Omoglymmius (sensu stricto) modicus new species' Fig. 177 Type material. - HOLOTYPE male, labelled: “SOLOMON IS., Savo, 5-4-62, P.J.M. Greenslade, 1327, B. M. 1966-477” (BMNH). PARATYPES two females (on same pin), one male, one female (on same pin), same data as holotype, labelled “1390” (BMNH); one male, same collector and locality as holotype, dated 6-4-62, “1455”(BMNH). Description. — Length 5. 0-6. 3 mm. Median lobe broad, its tip broadly rounded; frontal space very broad, U-shaped, its lateral margin deeply, evenly curved; medial angles obtuse, slightly separated; posteriomedial margin oblique; posteriolateral margin evenly curved; occipital angle indistinct; orbital groove represented by short band of pollinosity which scarcely extended past anterior margin of eye; temporal lobe with about 20 fine punctures. Pronotum short, broad; length/greatest width 1.17; widest near middle; base slightly narrowed; apex markedly narrowed; lateral margins curved; margin shallowly sinuate anterior to hind angle; medial margin of outer carina scarcely sinuate anterior to base; outer carina broadest near middle, slightly narrowed to base; narrowed almost to point at apex; basal part of inner carina less narrowed than in O. lindrothi, base narrowly truncate; outer carina with 22-25 fine punctures; inner carina with six to eight fine punctures. Elytral stria impressed, pollinosity limited to punctures; latter coarse, mostly elongate; female with deep, semicircular lateral pits on Sternum IV; male with prominent ventral tooth on anterior femur; female with or without small femoral tooth; hind calcar small, triangular, its apex obtuse. The pronotum in this species is shorter and less quadrate than in O. lindrothi and the medial margin of the temporal lobe is obtusely angulate opposite the median lobe. O. rusticus of nearby Russell Islands, has the margins of the pronotum much more markedly curved and the hind calcar is differently shaped. O. regius of Isabella has the outer carina more distinctly narrowed posteriorly and the frontal space more nearly V-shaped. Range. - Savo and Nggela Islands in the Solomons. In addition to the type series from Savo, we have seen the following specimens from Nggela: one female, labelled “Florida Is., Nggela L, Haleta, 0-50 m., 18-X-1964, R. Straatman, light trap” (BPBM); three females (on same pin), labelled “Nggela, Toa, 8-1-62, 524, P.J.M. Greenslade, B.M. 1966-477” (BMNH). 'see p. 254 for correction Quaest. Ent., 1982, 18 (1-4) 234 Bell and Bell Omoglymmius (sensu strict o) rusticus new species Fig. 178 Type material. - HOLOTYPE male, labelled: “SOLOMON IS., Russell Is., Loani, 31-5-1963, 6174, P. Greenslade, B.M. 1966-477” (BMNH). PARATYPES three females (on same pin), labelled: “Russell Is., Barika 30-5-1963, 6169”; one male, one female (on same pin), labelled: “Russell Is., Yandina, in logs, 22-24-11-1967, 13464” two males (on same pin). Russell Islands, Yandina, same data as preceding entry. All specimens collected by P.J.M. Greenslade, B.M. 1966-477 (all BMNH). Description. — Length 5. 0-6. 3 mm. Median lobe broad, its tip rounded; frontal space very broad, U-shaped, its margins deeply, evenly curved; medial angles narrow, slightly produced, obtuse; narrowly separated; posteriomedial margin slightly emarginate; posteriolateral margin evenly curved; occipital angle indefinite; orbital groove scarcely developed, but pollinose preorbital impression present; temporal lobe with about 20 very fine punctures. Pronotum moderately long; length/ greatest width 1.24; widest near middle, base, apex strongly narrowed; lateral margins markedly curved; margin not sinuate anterior to hind angle; medial margin of outer carina scarcely sinuate anterior to base; outer carina widest anterior to middle, both base and apex strongly narrowed; basal part of inner carina strongly narrowed, base narrowly truncate; outer carina with about 12 fine punctures; inner carina with six to eight fine punctures. Elytral striae impressed; punctures of Stria I, II fine, scarcely pollinose; those of outer striae coarser, distinctly pollinose; female with deep, semicircular lateral pits on Sternum IV; male with small ventral tooth on anterior femur; female with vestige of femoral tooth; hind calcar triangular, its apex obtuse, its upper margin curved. This species contrasts with the two preceding ones in the strongly curved lateral margins of the pronotum. The latter has the base almost as narrow as the apex. Form of the hind calcar also differs from that of O. modicus of the nearby islands of Nggela and Savo. Range. - Apparently identical forms are found in the Russell Islands and on San Cristobal. In addition to the type series, from the Russells, we have studied the following specimens from San Cristobal: one^male, one female (on the same pin), labelled; “Cristobal, Kinkia, 24-8-1962, P. Greenslade, 2193” (BMNH). Future study may show that the forms on the two islands owe their similarity to convergence, rather than close relationship. Omoglymmius ( sensu stricto) manni new species Figs. 179, 180 Type material. - HOLOTYPE male, labelled: “Auki, Sol. Isl., W. M. Mann” (MCZ). PARATYPE female, same data as holotype (MCZ). The type locality is on Malaita Is. Description. — Length 5. 7-6.0 mm. Median lobe broad, its tip broadly rounded; frontal space broad, its lateral margin more shallowly curved than in O. modicus; medial angles obtuse, slightly produced, nearly contiguous; posteriomedial margin oblique; posteriolateral margin evenly curved; occipital angle indistinct; orbital groove short but broad band of pollinosity, ended near middle of eye; temporal lobe 10-15 fine punctures. Pronotum moderately long, length/greatest width 1.19; widest near or slightly posterior to middle; base distinctly narrowed; apex markedly narrowed; lateral margins markedly curved; margin not sinuate anterior to hind angle; medial margin of outer carina shallowly sinuate anterior to base; outer carina curved, of nearly even width except for narrowed apex; basal part of inner carina constricted; outer carina with 22-25 fine punctures; inner carina with eight or nine very fine punctures. Basal scarp of elytron pollinose; elytral striae impressed; coarsely punctate; pollinosity limited to punctures; both sexes with shallow lateral pit on Sternum IV; male with prominent ventralTooth on anterior femur; female with minute one; hind calcar with proximal margin slightly concave, apex obtuse (Fig. 180). Shape of the pronotum in this species is close to that of O. rusticus , though the apex is more narrowed. However, the latter species has the hind calcar differently shaped and the lateral pit of Sternum IV of the female much deeper. We dedicate this species to the collector, the myrmecologist and former director of the National Zoological Garden, William M. Mann, who contributed many specimens of tropical Revision of Omoglymmius Ganglbauer 235 Rhysodini to American museums. Omoglymmius (sensu stricto) regius new species2 Figs. 182,190 Type material. - HOLOTYPE male, labelled: “SOLOMON IS., Isabel, Regi, 22-8-1963, P. Greenslade, 9554, B.M. 1966-477“ (BMNH). PARATYPES one female, labelled: “Isabel, Tatumba, 24-8-1963, P. Greenslade, 9587“ (BMNH); three males, one female (on same pin), labelled: “Isabel, Tatumba, 2-8-1962; 2432, P. Greenslade, B.M. 1966-477“ (BMNH). Description. — Length 5. 0-6. 6 mm. Median lobe broad, its tip rounded; frontal space moderately broad, less U-shaped than in O. manni, its lateral margins more oblique, less emarginate; medial angles obtuse, not produced, well separated; posteriomedial margin curved; posteriolateral margin curved; occipital angle absent; orbital groove pollinose, extended to or beyond middle of eye, slightly developed; temporal lobe with 12-15 very fine punctures. Pronotum moderately long; length/greatest width 1.19; widest slightly posterior to middle; base moderately narrowed (less so than in O. manni)', apex strongly narrowed; lateral margins moderately curved, less so than in O. manni', margin not sinuate anterior to hind angle; medial margin of outer carina shallowly sinuate anterior to base; outer carina oblique, of nearly even width except at apex, where narrowed; basal parts of inner carinae less narrowed than in O. manni', outer carina with about 20 fine punctures; inner carina with seven or eight fine punctures. Basal scarp of elytron pollinose; striae impressed, coarsely punctate; pollinosity limited to punctures; female with deep semi-circular lateral pits on Sternum IV; male pits fairly deep; male with prominent ventral tooth on anterior femur; female with minute one; hind calcar small, triangular, tip obtuse (Fig. 190). This species differs from O. manni in the more oblique, less curved margins of the pronotum and in the more separated medial angles of the temporal lobes. In general, shape is similar to O. gurneyi of Bougainville and Choiseul, but the latter has much more extensive pollinosity, including completely pollinose elytral striae. Omoglymmius (sensu stricto ) gurneyi new species Figs. 183, 189 Type material. - HOLOTYPE male labelled: “BOUGAINVILLE I., Nov. 1944, A.B. Gurney” (NMNH). PARATYPES one male, same locality and collector as type, dated 11- VI-44 (NMNH); one male, labelled: “Choiseul, Malangono, 25-8-1963, P. Greenslade, 9141, B.M. 1966-477“ (BMNH); one female, labelled: “Fauro I. Toumoa village, 30 m., 1 2- 1 V- 1 964, P. Shanihan, light trap” (BPBM). Description. — Length 6.2-7. 5 mm. Median lobe broad, its tip rounded; frontal space broad, its margin sharply bent at middle; medial angles obtuse, not produced, well separated; posteriomedial margin curved; posteriolateral margin evenly curved; occipital angle absent; orbital groove broadly pollinose, continued posteriorly as pollinose band along posterior margin of temporal lobe; temporal seta located within this band; temporal lobe with 15-25 fine punctures. Pronotum moderately long; length/greatest width 1.22; widest near middle; base slightly narrowed; apex markedly narrowed; lateral margins slightly curved except where moderately markedly curved near apex; margin scarcely sinuate anterior to hind angle; medial margin of outer carina sinuate anterior to base; outer carina slightly broadened at base, narrowed at apex, otherwise of nearly even width; base of inner carina gradually narrowed; outer carina with 30-35 fine punctures; inner carina with 12-17 fine punctures. Basal scarp of elytron pollinose; striae impressed, deep; coarsely punctate; elytral striae, except for Stria V, with punctures connected by continuous pollinosity; female with deep, semicircular lateral pit on Sternum IV; male with shallow one; male with prominent ventral tooth on anterior femur; female with small one; hind calcar triangular, its apical angle nearly rectangular, both proximal, distal margins straight. This species is separated from the others by continuous pollinosity of the striae, and extensive pollinosity along the posterior margin of the temporal lobe. It differs from all species except O. scopulinus of Santa Cruz in the high number of punctures in the pronotum and temporal lobes. O. princeps is sympatric with this species in Bougainville. It differs markedly in the shape of pronotum, as well as in having elytral pollinosity confined to the punctures. 2see p. 254 for correction Quaest. Ent., 1982, 18 (1-4) 236 Bell and Bell The specimen from Choiseul has the pollinosity of the elytral striae and temporal margins less developed, though still continuous. Otherwise, it seems identical to those from Bougainville. We dedicate this species to the distinguished orthopterist, Dr. Ashley B. Gurney, who collected the type specimen. Omoglymmius ( sensu stricto) princeps new species Fig. 184 Type material. - HOLOTYPE male, labelled: “SOLOMON Is., Bougainville (S). Kokure, nr. Crown Prince Range, 900 m., VI-8-1956, coll. E. J. Ford, Jr” (BPBM). Description. — Length 6.0 mm. Median lobe obtusely pointed posteriorly; frontal space broad, its margin moderately curved at middle; medial angles obtuse, separated; posteriomedial margin oblique; posteriolateral margin evenly curved; occipital angle distinct; orbital groove absent, but pollinose preorbital pit present; temporal lobe with about 20 fine punctures. Pronotum moderately long; length/greatest width 1.20; widest near middle; base, apex very markedly narrowed; lateral margins markedly curved; margin markedly sinuate anterior to hind angle; medial margin of outer carina sinuate well anterior to base; outer carina narrower than inner carina at middle and also at base; outer carina widest anterior to middle; inner carina constricted near base, latter broadened; outer carina with 10-11 fine punctures; inner carina with three or four fine punctures. Basal scarp of elytron not pollinose; striae impressed; strial punctures coarse, separated from one another by less than diameter of one puncture; male with prominent ventral tooth on anterior femur; hind calcar triangular, its apex obtuse; female unknown. The markedly rounded pronotal margins, absence of orbital groove and marginal pollinosity of the temporal lobe, and the lack of pollinosity between strial punctures separate this species easily from the sympatric O. gurneyi. The most similar species is O. renutus from Rendova and Mavovo. The latter species has a pronotum of similar shape, but has fewer punctures on the temporal lobes and pronotal carinae, while the hind calcar has a different shape. Omoglymmius (sensu stricto ) renutus new species Fig. 185 Type material. - HOLOTYPE male, labelled: “SOLOMON IS., New Georgia, Mavovo, 30-12-1964, P. Greenslade (cacao?) 16183, B.M. 1966-477” (BMNH). PARATYPE female, labelled: “SOLOMON IS., New Georgia, Rendova Is., 17-8-1963, P. Greenslade, B.M. 1966-477” (BMNH). Description. — Length 5. 0-6. 4 mm. Median lobe obtusely pointed posteriorly; frontal space broad, its margin moderately curved at middle; medial angles obtuse, nearly contiguous; posteriomedial margin oblique, slightly sinuate; posteriolateral margin evenly curved; occipital angle obsolete; orbital groove absent, but pollinose preorbital pit present; temporal lobe with 10-12 fine punctures. Pronotum long, length/greatest width 1.31; widest near middle; base, apex very markedly narrowed; lateral margins markedly curved; margin strongly sinuate anterior to hind angle; medial margin of outer carina sinuate anterior to base; outer carina narrower than inner one at middle and also at base; outer carina widest anterior to middle; inner carina narrowed anterior to base, latter rather broadly truncate; outer carina with about 10 very fine punctures; inner carina impunctate. Basal scarp of elytron not pollinose; striae impressed, rather finely punctate, punctures separated by more than diameter of one of them; male with prominent ventral tooth off anterior femur; female with smaller one; hind calcar triangular, its tip slightly lobate, truncate; female with deep semicircular lateral pit on Sternum IV; male with shallower one. The markedly rounded pronotal margins separate this species from all others except for O. princeps of Bougainville. The latter species differs in having the inner carina punctate, the temporal lobe with more punctures, and the hind calcar not truncate. Revision of Omoglymmius Ganglbauer 237 Omoglymmius (sensu stricto) scopulinus new species Fig. 186 Type material. - HOLOTYPE female, labelled: “SOLOMON IS., Santa Cruz, Reef Is., 30-6-1962, P. Greenslade 2131, B.M. 1966-477” (BMNH). Description. — Length 7.3 mm. Median lobe obtusely pointed posteriorly; frontal space moderately wide, its margins moderately curved at middle; medial angles rounded, well separated; posteriomedial margin evenly curved; posteriolateral margin evenly curved; occipital angle absent; orbital groove obsolete, ended slightly behind anterior margin of eye; temporal lobe with 30-35 fine punctures. Pronotum very long; length/greatest width 1.34, widest near middle; base, apex slightly narrowed; lateral margins slightly curved; margin slightly sinuate anterior to hind angle; medial margin of outer carina not distinctly sinuate anterior to base; outer carina distinctly narrower than inner one at middle; outer carina with base slightly narrower than that of inner carina; inner carina constricted just anterior to base; latter slightly broadened; rather narrowly truncate; outer carina with 18-21 fine punctures; inner carina with about 14 fine punctures. Elytral striae impressed; strial punctures coarse; Intervals III, V more convex than others, in form of inconspicuous raised carinae near base; female without ventral tooth on anterior femur; female with deep semicircular lateral pit on Sternum IV; male unknown. The long pronotum, the shape of the temporal lobe and its numerous punctures differentiate this species from all others. Omoglymmius ( sensu stricto ) mycteroides new species Fig. 187 Type material. - HOLOTYPE male, labelled: “SOLOMON Is., New Georgia gp. Kolombangara, hunda 2018, 1963, P. Greenslade, 8906, B.M. 1966-477” (BMNH). PARATYPES two males (on same pin) labelled: “SOLOMON IS., Guadalcanal, Mt. Tonapan, 3,500', 5-5-1963, P. Greenslade, 18120, B.M. 1966-477” (BMNH). Description. — Length 6. 0-6. 7 mm. Antennal Segments 1, II coarsely punctate; Segments Ill-XI impunctate; head nearly twice as long as wide preocular portion produced; median lobe elongate, its margins convergent in posterior half; its tip obtusely pointed; frontal space narrow; medial angles obtuse, contiguous, posteriomedial margin slightly emarginate; posteriolateral margin evenly curved; occipital angle distinct; antennal lobe far from eye; temporal lobe with anterior portion in form of oblique, pollinose ridge; margin of head in form of deep preorbital impression between eye and antennal lobe; orbital groove represented by short, broad pollinose impression ending opposite middle of eye; temporal lobe with eight to 16 fine punctures, mostly near temporal seta; one temporal seta; postorbital, suborbital tubercles absent; eye large, round. Pronotum moderately long, length/greatest width 1.20; widest near middle; base slightly narrowed; apex markedly narrowed; lateral margins curved; margin not sinuate anterior to hind angle; inner carina slightly broader than outer carina at middle; medial margin of outer carina angulate, sinuate anterior to base; outer carina curved, of nearly even width except at extreme apex and base; inner carina widest near middle, narrowed to base, latter truncate; outer carina with 23-25 coarse punctures; inner carina with four or five very fine punctures; pronotum without setae; prosternum without precoxal carinae. Elytron rather elongate, narrow; striae impressed, coarsely punctate; base of Stria IV without longitudinal pollinose scarp; Intervals III, V raised above others, subcarinate in anterior third; Stria IV with one seta near apex; subapical stride with one seta; Stria VII with several setae near apex; metasternum coarsely punctate; abdominal Sterna 1I1-V coarsely, sparsely punctate, punctures in form of irregular transverse row, especially near midline; lateral pit of Sternum IV barely discernible in either sex; male with ventral tooth on anterior femur; middle calcar acute, nearly as long as spur; hind calcar small, its apex an obtuse angle; female unknown. This isolated species differs strikingly from other species of the Solomon Islands in having a long snout. In this respect, it is superficially similar to O. bucculatus of Sumbawa and O. nasalis of Buru. In head structure, it is intermediate; the anterior part of the temporal lobe forms an oblique, pollinose ridge, as in O. nasalis , but there is no oblique groove latered to the ridge, as in the latter species. It differs from both these species in the subcarinate bases of Intervals III and V, a character shared with O. scopulinus , of the lindrothi group. Quaest. Ent., 1982, 18 (1-4) 238 Bell and Bell One might expect the populations on Guadalcanal and Kolombangara to represent distinct species, as is so in other groups of Rhysodini in the area. There are minor differences in shape of body and form of calcars, but these could be within the range of variation of a single population. Pending collection of more specimens, we consider the forms on the two islands to be conspecific. Omoglymmius (sensu stricto) tabulatus new species Figs. 188, 191 Type material. - HOLOTYPE male, labelled: “SOLOMON Is., Bougainville (S.) Boku-50 m., VI-5-56, coll. J.L. Gressitt” (BPBM). PARATYPE female, labelled: “SOLOMON IS., New Georgia gp., Kolombangara, nr. Kusi in log, 2-9-1965, leg. P. Naturaga, pres. P.J.M. Greenslade, B.M. 1966-477” (BMNH). Description. — Length 4. 3-4. 7 mm. Antennal Segments I, II coarsely punctate; Segments III-XI impunctate; head slightly longer than wide; median lobe rather short, its apex obtusely rounded; frontal space small, narrow, U-shaped, longer than wide; medial angle obtuse, slightly produced, contiguous; posteriomedial margin shallowly sinuate; posteriolateral margins evenly curved; occipital angles distinct, rather far apart; orbital groove abbreviated, extended to middle of eye; temporal lobe with 14-15 very coarse punctures; one temporal seta; postorbital, suborbital tubercles absent; eye large, round. Pronotum small, short, narrow; length/greatest width 1.31; lateral margins parallel except near base, apex; pronotum thus hexagonal; outer carina formed of two planes meeting at a straight line, medial plane is sloped toward paramedian groove, outer plane is nearly vertical; outer carina in dorsal view about 0.67 as broad as inner carina; outer carina with about 16 coarse punctures; inner carina with three to seven coarse punctures; pronotum without setae; prosternum without precoxal carinae. Elytron relatively long, narrow; striae impressed, coarsely punctured; base of Stria IV without longitudinal scarp; Stria II with one to four setae; Stria IV with four or five setae; subapical stride with one seta; apex of Stria VII with several setae; metasternum coarsely punctate; abdominal Sterna III-V with scattered coarse punctures; Sternum IV with deep, semicircular lateral pit in both sexes; anterior femur with small, obtuse tooth in male, without tooth in female; middle calcar obtuse, about 0.5 as long as spur; hind calcar triangular, longer than deep, its apex obtuse, its proximal margin convexly curved (Fig. 191). Form of the outer carina in this species is unique, with separate dorsal and lateral surfaces, separated by a sharp edge. Among species from the Solomon Islands it is unique in its small size and in having numerous elytral setae. The specimen from Kolombangara is only provisionally associated with the holotype. It differs in having fewer punctures on temporal lobe, these limited to margin; inner carina with fewer punctures, and the second stria with four rather than one seta. These might be specific differences, but more specimens are needed to confirm it. Omoglymmius (sensu stricto) classicus new species Fig. 192 Type material. - HOLOTYPE female, labelled: “ADMIRALTY IS., Los Negros, XI-45, W. Wagner jr., D. Greether” (BPBM). Description. — Length 6.0 mm. Antennal Segments I- 1 X punctate, distal segments very finely so; Segments X-XI impunctate; median lobe short, lance-shaped, its apex obtusely pointed; frontal space large, almost twice as wide as long, U-shaped, its margins markedly curved; medial angles obtuse, well separated; posteriomedial margin oblique, slightly sinuate; posteriolateral margin evenly curved; occipital angles very obtuse; trace of orbital groove present, near anterior margin of eye, posterior to pronounced preorbital pit; temporal lobe with about 23 fine punctures; one temporal seta; postorbital, suborbital tubercles absent; eye large, round. Pronotum moderately long; length/greatest width 1.24; widest near middle; base slightly narrowed; apex more markedly narrowed; lateral margins weakly curved; margin markedly sinuate anterior to hind angle; inner carinae slightly wider than outer ones at middle; medial margin of inner carina markedly sinuate anterior to base; outer carina widest anterior to middle, markedly narrowed anteriorly; inner carina narrowed just anterior to base; base broadened, truncate, Revision of Omoglymmius Ganglbauer 239 very finely pollinose; outer carina with about 22 fine punctures; inner carina with eight to 10 fine punctures; pronotum without setae; prosternum without precoxal carina. Elytron relatively long, narrow; striae impressed, coarsely punctate; transverse basal scarp very finely pollinose; base of Stria IV with pollinose longitudinal scarp; Stria IV with one seta at apex; subapical stride with one seta; Stria VII with several setae near apex; metasternum with most punctures near margins or midline, disc otherwise nearly impunctate; abdominal Sterna II1-V with punctures scattered, well separated in medial portion, becoming coalescent laterally; female with deep, round lateral pit in Sternum IV: female without ventral tooth on anterior femur; male unknown. This species resembles members of the lindrothi complex. The sinuation of the lateral margin of the pronotum suggests O. princeps and O. renutus, but the lateral margins are much less curved than in the latter two species. Another similar species is O. vicinus of New Guinea. The latter species differs in the shape of the temporal lobe, and in having the median lobe punctate. Omoglymmius ( sensu stricto) vicinus (Grouvelle) Fig. 193 Rhysodes vicinus Grouvelle 1895a: 157. Omoglymmius ( sensu stricto) vicinus (Grouvelle) Bell and Bell 1978. Type material. - HOLOTYPE female, labelled: “NOUV. GUINEE, Dorey, Baie de Geelvink, Raffray & Maindron 78” (MNHN). Additional specimens listed in Grouvelle (1903) are not authentic types, but merely additional localities. Description. — Length 4. 6-6. 2 mm. Antennal Segments 1-IV coarsely punctate; Segments V-1X very finely punctate; Segments X-Xl impunctate; median lobe short, oval, its apex obtusely rounded; median lobe with three to six punctures; frontal space twice as broad as long, U-shaped, its margins markedly, abruptly curved; median angles obtuse, well separated, lobate; posteriomedial margin oblique, emarginate; posteriolateral margin evenly curved; occipital angle obtuse; orbital groove represented by a narrow line of pollinosity of varied length; antennal lobe separated from temporal lobe by broader pollinose area than in O. classicus\ temporal lobe with 18-22 fine punctures; one temporal seta; postorbital, suborbital tubercles absent; eye large round. Pronotum moderately long, length/greatest width 1.22; widest near middle; base slightly narrowed; apex markedly narrowed; lateral margins slightly curved; margin sinuate anterior to hind angle; inner carina slightly wider than outer carina at middle; medial margin of outer carina slightly sinuate anterior to base; outer carina widest anterior to middle, markedly narrowed to apex; inner carina narrowed just anterior to base; latter narrowly truncate; outer carina with 20-30 very fine punctures; inner carina with six to 12 very fine punctures; pronotum without setae; prosternum without precoxal carinae. Elytron relatively long, narrow; striae impressed, coarsely punctate; base of Stria IV with longitudinal pollinose scarp; Stria IV with one seta at apex; subapical stride with one seta; Stria VII with several setae near apex; metasternum entirely punctate; punctures of abdominal Sterna III-V very coarse, scattered, confluent near lateral margin; in some specimens with very fine punctures medial to and posterior to the coarse principal punctures; female with deep semicircular lateral pit on Sternum IV; male with shallow one; both sexes with ventral tooth on anterior femur; middle calcar minute, obtuse; hind calcar triangular, its apex acute, its dorsal margin straight. Among species from New Guinea, this one is recognized by the distinct sinuation anterior to the hind angles and the U-shaped frontal space. No other species has both of these characters. In form it is rather isolated among New Guinean species, and is really closer to O. classicus of the Admiralty Islands, from which it differs in presence of punctures on the median lobe and in shape of the temporal lobe. Range. - North Coast of New Guinea from Geelvink Bay eastward, and on Goodenough Island, in the D’Entrecasteaux Islands near the eastern end of New Guinea. In addition to the type material, we have seen the following specimens: two males, Friedrich-Wilh.-hafen, Biro, 96 (MNHN); one male, one female, Dutch New Guinea, Maffin Bay, IX-1944, coll. E. S. Ross (CAS); one female, Stephansort, Astrolabe Bay, Biro 1900 (MNHN); two females, Goodenough Is., Gennais 1890, L. Loria (GEN). Quaest. Ent., 1982, 18 (1-4) 240 Bell and Bell Omoglymmius(sensu stricto ) oroensis new species Fig. 194 Type material. - HOLOTYPE female, labelled: “PAPUA, N. G., Oro Bay, Dec. 43-Jan. 44, Darlington” (MCZ). Description. — Length 7.0 mm. Antennal Segments I- IV coarsely punctate; Segments V-X more finely punctate; Segment XI impunctate; head distinctly longer than wide; median lobe short, oval, its tip rounded; median lobe with a few punctures; frontal space longer than broad, narrow, nearly V-shaped, its margins shallowly curved; medial angles obtuse, markedly separated; posteriomedial margin oblique, short; posteriolateral margin bent near temporal puncture, its posterior portion oblique; occipital angle very obtuse; orbital groove distinct, ended posterior to middle of eye; anterior portion of temporal lobe a convex, pollinose ridge; temporal lobe with 14-16 rather fine punctures; one temporal seta; postorbital, suborbital tubercles absent; eye large, round. Pronotum rather short, subquadrate; length/greatest width 1.14; base slightly narrowed; apex markedly narrowed; lateral margins slightly curved; margin scarcely sinuate anterior to hind angle; outer carina approximately 0.67 as wide as inner carinae at middle; medial margin of outer carina sinuate just anterior to base; outer carina widest anterior to middle, strongly narrowed at apex; inner carina narrowed to base; latter narrowly truncate; outer carina with 10-20 fine punctures; inner carina with three or four fine punctures; pronotum without setae; prosternum without precoxal carina. Elytron relatively long, narrow; striae impressed, coarsely punctate; transverse basal scarp pollinose; base of Stria IV with longitudinal pollinose scarp; Stria IV with one seta near apex; subapical stride with one seta; Stria VII with several setae near apex; metasternum entirely punctate; abdominal Sterna III-V coarsely punctate; punctures confleunt near lateral margin; female with lateral pit on Sternum IV shallow; female with minute ventral tubercle on anterior femur. This species is somewhat similar to O. vicinus except for shape of the pronotum and the more extensive pollinosity between antennal and temporal lobes. Variation. - We provisionally include two other specimens with the holotype, although they differ from it in some respects. One is a male, length 6.0 mm. labelled: “PAPUA, N. G., Dobodura, Mar. -July 1944, Darlington” (MCZ), It has a wider median lobe than the holotype. It has a ventral tooth on the anterior femur. The middle calcar is very small and obtuse, while the hind calcar is triangular, with an obtuse apex, and with its distal margin distinctly emarginate. The second specimen is a female, length 7.2 mm., labelled: “PAPUA-Kokoda, 200 ft. VI-1933, L. E. Cheesman, B. M. 1933-456” (BMNH). It differs from the holotype in having the temporal lobe more evenly rounded posteriorly, and in having two temporal setae on each side. More material will be necessary to ensure that these specimens are really variants of O. oroensis. Omoglymmius (sensu stricto ) fringillus new species Fig. 195 Type material. - HOLOTYPE male, labelled: “NEW GUINEA (NE) Huon Peninsula, Finschhafen, 150 m., 1 4-1 V- 1 963, Sedlacek” (BPBM). PARATYPES one male, same data as holotype (BPBM); one female, labelled; “NEW GUINEA (NE), Eliptamin Valley, 1200-1350 m., July 1-15, 1959, W. W. Brandt” (BPBM); one male, labelled: “Lae, Aug. 1944, F. E. Skinner” (BPBM); one female, labelled: “Lae 24-8-65”(NMNZ). Description. — Length 6. 2-7.0 mm. Antennal Segments I-IV coarsely punctate; Segments V-X finely punctate; Segment XI impunctate; head as broad as long; median lobe short, oval, its apex obtusely rounded; median lobe impunctate; frontal space broader than long, strongly U-shaped, margins parallel anteriorly, abruptly curved, nearly transverse posteriorly; medial angles obtuse, nearly rounded, separated, posteriomedial margin curved into posterior margin; posteriolateral margin curved into posterior margin; occipital angle absent; antennal lobe separated from temporal lobe by broad pollinose area; orbital groove distinct, extended to posterior margin of eye; temporal lobe with 15-17 fine ! punctures; posterior margin of temporal lobe with narrow fringe of pollinosity; one temporal seta; postorbital, suborbital tubercles absent; eye large, round. Pronotum short, quadrate; length/greatest width 1.16; widest near middle; base scarcely narrowed; apex markedly j narrowed; lateral margins nearly parallel except near apex, where curved, markedly convergent; margin not sinuate anterior to hind angle; outer carina distinctly wider than inner one at middle; medial margin of outer carina sinuate Revision of Omoglymmius Ganglbauer 241 anterior to base; outer carina widest anterior to middle; apex markedly narrowed; inner carina narrowed to base, latter very narrowly truncate; outer carina with 22-27 fine punctures; inner carina with five to seven fine punctures; pronotum without setae; prosternum without precoxal carinae. Elytra relatively long, narrow; striae impressed, coarsely punctate; transverse basal scarp pollinose; base of Stria IV with longitudinal pollinose scarp; Stria IV with one seta near apex; subapical striole with one seta; Stria VII with several setae near apex; metasternum coarsely punctate; abdominal Sterna III-V very coarsely punctate, punctures confluent near lateral margin on Sterna III-VI; both sexes with shallow lateral pits on Sternum IV; male with prominent ventral tooth on anterior femur; female with small tooth; middle calcar small, obtuse; hind calcar triangular, its apex sharp; both distal and proximal margins straight. This is a species with deep, coarsely punctate striae, a broad head, and a quadrate pronotum. Among New Guinean species it is closest to O. oroensis, but differs from the latter in having a shorter, broader head, a more nearly quadrate pronotum and an impunctate median lobe. Like the two preceding species, it inhabits the lowlands of the north coast. Omoglymmius (sensu stricto ) puncticornis new species Fig. 196 Type material. - HOLOTYPE male, labelled: “NEW GUINEA:PAPUA, Kiunga, Fly River, VII- 1 5-2 1 , Wm. W. Brandt” (BPBM). PARATYPES one male, labelled: “Nuova Guinea, Fly River, 1876-77, L. M. D'Albertis” (MNHN), one female same label as paratype (GEN). Description. — Length 5. 8-6. 5 mm. Antennal Segments I-X coarsely punctate; Segment XI impunctate; head distinctly longer than wide; median lobe short, its tip obtuse; frontal space broader than long, U-shaped, its lateral margins curved; medial angles obtuse, narrowly separated; posteriomedial margin curved into posteriolateral margin; occipital angle absent; preorbital pollinose impression distinct, but orbital groove absent; antennal lobe separated from temporal lobe by rather broad pollinose area; temporal lobe with 22-27 punctures; one temporal seta; postorbital, suborbital tubercles absent; eye large, round. Pronotum rather short, length/greatest width 1.13; widest near middle; base moderately narrowed; apex markedly so; lateral margins curved; margin not sinuate anterior to hind angle; inner carina slightly wider than outer carina at middle; medial margin of outer carina scarcely sinuate anterior to base; outer carina widest anterior to middle, markedly narrowed to apex; inner carina constricted anterior to base, latter pollinose; outer carina with about 23 rather coarse punctures; inner carina with seven to 10 finer punctures; pronotum without setae; prosternum without precoxal carinae. Elytron relatively long, narrow; striae impressed, coarsely punctate; transverse basal scarp pollinose; base of Stria IV with pollinose longitudinal scarp; Stria IV with one seta near apex; subapical striole with one seta; Stria VII with several setae near apex; metasternum entirely punctate; abdominal Sterna III-V with coarse, scattered punctures; both sexes with deep semicircular lateral pits on Sternum IV; male with prominent ventral tooth on anterior femur; middle calcar very small, obtuse; hind calcar triangular, its apex obtuse, its proximal margin straight. The very coarse punctures on the outer antennal segments are distinctive. This species differs from other similar species from New Guinea in having the lateral margins of the pronotum curved, but without a sinuation anterior to the hind angle. Curved margins eliminate O. fringillus and O. oroensis, while the absence of a sinuation eliminates O. vicinus , as do the coarse punctures of the outer antennal segments. O. viduus of the Kei Islands, is the most similar extralimital species. It differs in having the posteriomedial margin of the temporal lobe sinuate, and elytral punctures much finer. Variation. - A pair (male GEN, female MNHN) labelled “Ighibirei, New Guinea, VII-VIII 90, Loria”, resemble O. puncticornis in most respects, so we provisionally include them in the species. However, the inner carinae of the pronotum are entirely impunctate; and the abdominal punctures tend to form a single transverse row on each sternum. We have been unable to locate Ighibirei. These specimens may represent another species, but final judgment must await the collection of more specimens. Quaest. Ent., 1982, 18 (1-4) 242 Bell and Bell Omoglymmius (sensu stricto) trepidus new species Fig. 197 Type material. - HOLOTYPE male, labelled: “NEW GUINEA (NE), Wau, Morobe Dist., 1200 m„ 1 5-30-IX- 1 962, J. Sedlacek” (BPBM). Description. — Length 6.0 mm. Antennal Segments 1- 1 V coarsely punctate; Segments V-XI very finely punctate; head distinctly longer than wide; median lobe short, impunctate, its tip very obtuse, almost rounded; frontal space slightly broader than long, nearly V-shaped, its margins only shallowly curved; medial angles obtuse, slightly lobate; posteriomedial margin emarginate; posteriolateral margin evenly curved; occipital angles very obtuse; preorbital pollinose impression very prominent, continued posteriorly as very short orbital groove; antennal lobe separated from temporal lobe by short, narrow antennal groove; temporal lobe with 21-24 punctures; one temporal seta; postorbital, suborbital tubercles absent; eye large, round. Pronotum rather short, length/greatest width 1.17; widest near middle; base moderately narrowed; apex markedly narrowed; lateral margins markedly curved; margin not sinuate anterior to hind angle; inner carina slightly wider than outer carina at middle; medial margin of outer carina shallowly sinuate anterior to base; outer carina anterior to middle, very markedly narrowed to apex and to base; latter rather narrowly truncate; inner carina constricted near base, latter broadly truncate; outer carina with 16-18 very fine punctures; inner carina impunctate; pronotum without setae; prosternum without precoxal carinae. Elytron rather short, broad; striae shallowly impressed, strial punctures rather fine; especially in inner striae; transverse basal scarp iridescent, not pollinose; base of Stria IV with very short longitudinal pollinose scarp; Stria IV without seta near apex; subapical stride with one seta; Stria VII with several setae near apex; metasternum largely finely punctate, but with impunctate area at either side of midline; abdominal Sterna III-V with scattered punctures; with deep, semicircular lateral pit on Sternum IV; with ventral tooth on anterior femur; middle calcar scarcely evident; hind calcar triangular, its apex rounded; female unknown. The large preorbital pit and the rounded median lobe separate this species from O. patens and O. cavea , while the presence of a small basal carina on Stria IV and rounded, rather than oblique lateral pits of Sternum IV, separate it from O. sectatus. Omoglymmius ( sensu stricto ) patens new species Fig. 198 Type material. - HOLOTYPE female, labelled: “Maffin Bay, Dutch New Guinea, IX- 1944, E.S. Ross” (CAS). Description. — Length 5.2 mm. Antennal Segments I-IV coarsely punctate; Segments V-X very finely punctate; Segment XI impunctate; head distinctly longer than wide; median lobe short, with six scattered punctures; median lobe lance-shaped, its tip obtusely pointed; frontal space broader than long, nearly V-shaped, its lateral margins shallowly curved; medial angles obtuse, well separated; posteriomedial margin oblique; anteriomedial margin evenly curved; occipital angle indistinct; preorbital pollinose impression small; orbital groove absent; antennal groove short, narrow; temporal lobe with 23-27 fine punctures; one temporal seta; postorbital, suborbital tubercles absent; eye large, round. Pronotum moderately long, length/greatest width 1.22; widest near middle, base moderately narrowed; apex more markedly narrowed; lateral margins markedly curved; margin not sinuate anterior to hind angle; marginal groove dilated, much broader than in O. trepidus ; outer carina narrow, about 0.5 as wide as inner carina at middle; medial margin of outer carina scarcely sinuate anterior to base; outer carina widest anterior to middle; slightly narrowed at base; markedly narrowed at apex; inner carina constricted anterior to base; latter broad, truncate; outer carina with 18-22 fine punctures; inner carina impunctate; pronotum without setae; prosternum without precoxal carinae. Elytron rather long, narrow; striae impressed; strial punctures coarse; transverse basal scarp glabrous; base of Stria IV with pollinose longitudinal scarp rather long, its length equal to width of two elytral intervals; Stria IV without seta near apex; subapical stride with one seta; Stria VII with several setae near apex; metasternum coarsely punctate near margins, otherwise impunctate; abdominal Sterna III-V with punctures coarse, sparse, in form of irregular transverse row on each sternum; not confluent near lateral margins; female with deep, semicircular lateral pits on Sternum IV; female with prominent ventral tooth on anterior femur; male unknown. Outer carinae of the pronotum are narrower than in any other member of the subgenus in New Guinea. This easily separates this species from the two most similar species, O. trepidus and O. cavea. Revision of Omoglymmius Ganglbauer 243 Omoglymmius ( sensu stricto) cavea new species Fig. 199 Type material. - HOLOTYPE female, labelled: “NEW GUINEA, Ramoi II., Beccari, 1875” (GEN). Description. — Length 5.9 mm. Antennal Segments I- 1 1 coarsely punctate; Segments 1II-XI very finely punctate; head distinctly longer than broad; median lobe short, impunctate, lance-shaped, its tip obtusely pointed; frontal space broader than long, nearly V-shaped, its lateral margins shallowly curved; medial angles obtuse, well separated; posteriomedial margin oblique, shallowly emarginate; posteriolateral margin evenly curved; occipital angle obtuse; preorbital pollinose impression small; orbital groove absent; antennal groove short, narrow; temporal lobe with 17-18 fine punctures; one temporal seta; postorbital, suborbital tubercles absent; eye large, round. Pronotum moderately long, length/greatest width 1.30; widest near middle; base moderately narrowed; apex markedly narrowed; lateral margins markedly curved; margin shallowly sinuate anterior to hind angle; marginal groove not dilated; outer carina slightly narrower than inner one at middle; medial margin of outer carina scarcely sinuate anterior to base; outer carina widest anterior to middle, slightly narrowed at base, markedly narrowed at apex; inner carina constricted anterior to base; latter broad, truncate; outer carina with 16-18 rather coarse punctures; inner carina impunctate; pronotum without setae; prosternum without precoxal carinae. Elytron rather long, narrow; striae impressed; strial punctures moderate, finer than in O. patens ; transverse basal scarp glabrous; base of Stria IV with rather short pollinose longitudinal scarp; Stria IV with seta near apex; subapical stride with one seta; Stria VII with several setae near apex; metasternum entirely punctate; abdominal Sterna III-V rather densely, finely punctate, punctures scattered; female with deep triangular lateral pits on Sternum IV, these strongly narrowed posteriorly; with ventral tooth on anterior femur; male unknown. This species resembles O. trepidus and O. patens in having inner carinae impunctate, base of Stria IV carinate, and anteriomedial margin of the temporal lobe shallowly sinuate. It differs from the former in having the tip of the median lobe pointed and preorbital pollinose impression very small, and from the latter species in having the outer carinae broad, marginal groove not dilated and lateral pits of Sternum IV triangular. Shape of lateral pits is intermediate between the semicircular one of most Omoglymmius and the narrow oblique ones of O. sectatus. Omoglymmius (sensu stricto ) sectatus new species Figs. 200, 201 Type material. - HOLOTYPE male, labelled: “NEW GUINEA:(NE) Mt. Missim, 950-1000 m., 10-VIII-64, J. Sedlacek” (BPBM). PARATYPE female, labelled: “NEW GUINEA (NE), Wau. Morobe Dist., 1200 m., 2-VI-62, J. & M. Sedlacek” (BPBM). Description. — Length 5.8 mm. Antennal Segments I-IV coarsely punctate; Segments V-XI finely, irregularly punctate; head distinctly longer than broad; median lobe short, impunctate, lance-shaped, its tip narrowly rounded; frontal space longer than broad, V-shaped, its margin shallowly curved; medial angles obtuse, well separated; posteriomedial margin oblique; posteriolateral margin evenly curved, occipital angle scarcely evident; antennal groove narrow, its lateral end scarcely broadened; orbital groove absent; temporal lobe with 15-21 fine punctures; one or two temporal setae; postorbital, suborbital tubercles absent; eye large, round. Pronotum moderately long, length/greatest width 1.25; widest slightly anterior to middle; base moderately narrowed; apex markedly narrowed; lateral margins markedly curved; margin not sinuate anterior to hind angle; outer carina subequal to inner carina at middle; medial margin of outer carina scarcely sinuate anterior to base; outer carina widest at anterior third of length, moderately narrowed, subtruncate at apex; inner carina constricted anterior to base; latter broad, truncate; outer carina with 10-12 very fine, irregularly scattered punctures; inner carina with two or three very fine punctures; pronotum without setae; prosternum without precoxal carinae. Elytron rather narrow, elongate; striae impressed; strial punctures coarse; base of Stria IV without longitudinal scarp; Stria IV with one seta near apex; subapical stride with one seta; Stria VII with several setae near apex; metasternum coarsely punctate in midline and near margins; disc otherwise impunctate; abdominal Sterna III-V with coarse, scattered punctures; both sexes with narrow, oblique, slit-like lateral pit on Sternum IV; male with ventral tooth on anterior femur; female without such tooth; middle calcar very small, acute (Fig. 201) hind calcar triangular, its apex acute, its proximal margin straight, distal margin emarginate. This small, narrow species is easily identified by the oblique, slit-like lateral pits on Sternum IV. Quaest. Ent., 1982, 18 (1-4) 244 Bell and Bell Omoglymmius (sensu stricto) ephemeris new species Figs. 202, 209 Type material. - HOLOTYPE male, labelled: “NEW GUINEA (NW), Nabire, S. Geelvink Bay, 0-30 m., 2-9-VII-1962, J. L. Gressitt coll,” (BPBM). PARATYPE female, labelled: “NEW GUINEA: Papua, Kiunga, Fly River, VII-23-25- 1 957. coll. Wm. W. Brandt” (BPBM). Description. — Length 5.0 mm. Antennal Segments I-IV coarsely punctate; Segments V-XI finely punctate; head 1.3 longer than broad; median lobe short, coarsely punctate, lance-shaped, its tip obtusely pointed; frontal space broader than long, V-shaped, lateral margins shallowly curved; medial angles obtuse, well separated; posteriomedial margin oblique; posteriolateral margin evenly curved; occipital angle distinct; antennal groove narrow, its lateral end not broadened; orbital groove absent; temporal lobe with 25-30 rather coarse punctures; one temporal seta; postorbital, suborbital tubercles absent; eye large, round. Pronotum moderately long, length/greatest width 1.21; widest near middle; base moderately narrowed; apex strongly so; lateral margins markedly curved; margin not sinuate anterior to hind angle; outer carina subequal to inner carina at middle; medial margin of outer carina not sinuate anterior to base; outer carina widest anterior to middle, moderately narrowed, rounded anteriorly; median margin of outer carina indistinct opposite basal impression; inner carina constricted anterior to base, latter very broadly truncate, nearly closing paramedian groove posteriorly; outer carina with 13-15 punctures; inner carina with five or six rather fine punctures; pronotum without setae; prosternum without precoxal carinae. Elytra narrow, elongate; striae impressed, coarsely punctate; base of Stria IV without longitudinal scarp; Stria IV with one seta near apex; subapical stride with one seta; Stria VII with one seta near apex; metasternum coarsely punctate; abdominal Sterna III-V with coarse scattered punctures; both sexes with deep lateral pits on Sternum IV; lateral pits nearly semicircular, but anteriomedial margin oblique; male with distinct ventral tooth on anterior femur; female with very small, obtuse one; middle calcar acute, nearly as long as spur (Fig. 209); hind calcar triangular, its apex acute, slightly lobate, its proximal margin shallowly sinuate; its distal margin nearly straight. This is a small, narrow species which lacks the basal scarp on Stria IV. It differs from O. sectatus having the lateral pits of the abdomen nearly round and the hind calcar much longer and more acute. Although the two specimens come from distant localities, they are markedly similar, suggesting that this is a widely distributed lowland species. Omoglymmius (sensu stricto) gracilicornis (Grouvelle) Figs. 203,210 Rhysodes gracilicornis Grouvelle 1895a: 157. Omoglymmius ( sensu stricto ) gracilicornis (Grouvelle) Bell and Bell 1978 Type material. - HOLOTYPE male, labelled: “Baie du Geelvink, Raffray et Maindron 78” (MNHN). Description. — Length 4.5 mm. Antennal punctures not recorded; head slightly longer than broad; head nearly as broad as pronotum; median lobe short, impunctate, its tip obtuse; frontal space as long as broad, V-shaped, its lateral margins only slightly curved; medial angles nearly rectangular, well separated; posteriomedial margin oblique; posteriolateral margin evenly curved; occipital angle distinct; antennal groove narrow, its lateral end not broadened; orbital groove absent; temporal lobe with 10-12 punctures restricted to lateral half; one temporal seta; postorbital, suborbital tubercles absent; eye round, rather small. Pronotum moderately elongate; length/greatest width 1.30; widest near middle; base slightly narrowed; apex moderately narrowed; lateral margins slightly curved; margin not sinuate anterior to hind angle; outer carina equal in width to inner carina at middle; medial margin of outer carina not sinuate anterior to base; outer carina widest anterior to middle; subtruncate apex; inner carina constricted anterior to base; latter narrowly truncate; inner and outer carina entirely without punctures; pronotum without setae; prosternum without precoxal carinae. Elytra rather elongate, narrow; striae impressed, coarsely punctate; base of Stria IV without longitudinal scarp; Stria II with one seta near apex; Stria IV with two to four setae; subapical striole without seta; Stria VII with several setae near apex; metasternum with coarse punctures near lateral margins, a few coarse punctures in midline, otherwise impunctate; abdominal Sterna III-V with coarse punctures in form of transverse row on each sternum; with deep semicircular lateral pit on Sternum IV: without ventral tooth on anterior femur; middle calcar very small, obtuse; hind calcar prominent, its apex broadly rounded; female unknown. Revision of Omoglymmius Ganglbauer 245 This small, narrow species has more elytral setae than any similar species from New Guinea, and is also recognized by impunctate pronotal carinae and broadly rounded hind calcar (Fig. 210). A second specimen (MNHN), from Astrolabe Bay, labelled as R. gracilicornis by Grouvelle, is a different species, O. vicinus (Grouvelle). Omoglymmius (sensu stricto) follis new species Figs. 204,212 Type material. - HOLOTYPE female, labelled: “NEW GUINEA (NE), Wau, Morobe Distr., 1700-1800 m., 7-X-1962, J. & M. Sedlacek” (BPBM). PARATYPES one male, one female, labelled: “N. GUINEA. Okapa, Putosa, 2-11-1964, R. Hornabrook” (NMNZ). Description. — Length 7. 8-8. 8 mm. Antennal Segments I, 11 coarsely punctate; Segments 1I1-X very finely, sparsely punctate; Segment XI impunctate; head slightly longer than broad; median lobe short, broad, nearly quadrangular; its tip broadly truncate, with narrow pollinose posterior border; frontal space broader than long, its anteriomedial margin curved at middle; medial angles obtuse, well separated; posteriomedial margin oblique; posterior margin transverse, bordered with pollinosity, meeting posteriolateral margin at obtuse angle near temporal seta; antennal groove broad, dilated at lateral end in form of pollinose preorbital impression; latter extended posteriorly medial to eye for short distance, in form of vestige of orbital groove; temporal lobe with seven or eight extremely minute punctures, visible only under high magnification; one temporal seta; postorbital tubercle small, about 0.25 as long as eye, visible only in lateral view; eye large, round. Pronotum short, length/greatest width 1.04, widest near middle; base slightly narrowed, apex very markedly narrowed; lateral margins slightly curved in basal half, becoming markedly curved near apex; margin shallowly sinuate anterior to hind angle; outer carina slightly narrower than inner carina at middle; outer carina broadest at base, slightly constricted anterior to base, slightly broader to anterior third, tapered to point at apex; inner carinae long oval, tapered to point at base; outer carina with 12-15 very minute punctures, visible only at high magnification; inner carina with about 12 equally minute punctures; pronotum without setae; prosternum without precoxal carinae. Elytra rather narrow, elongate; striae not impressed; strial punctures coarse; base of Stria IV with longitudinal scarp; transverse basal scarp of elytron pollinose from base of Stria IV to suture; Stria IV without seta at apex; subapical stride with one or no setae; Stria VII with few setae near apex; metasternum punctate near lateral and posterior margins, its disc impunctate; abdominal Sterna III-V with punctures scattered, fine near midline, coarse, coalescent laterally; both sexes with deep, semicircular lateral pit on Sternum IV; male with ventral tooth on anterior femur; female without this tooth; middle calcar minute, obtuse; hind calcar triangular, its apex obtuse, distance from tibial spur to proximal end of calcar more than twice distance from tip of calcar to its base. This large species has the striae coarsely punctate but not impressed, and has a small postorbital tubercle, visible only in lateral view. The most similar species is O. iridescens. The latter species has the median lobe sharply pointed posteriorly, rather than truncate, outer carinae not suddenly dilated at base, and medial angles of temporal lobe contiguous. Omoglymmius (sensu stricto ) iridescens new species Figs. 205,211 Type material. - HOLOTYPE male (left outer pronotal carina malformed), labelled: “NIEUW GUINEA Exp., KNAG 1939, Araboebivak, 19-X-1939” (LEI). PARATYPE female, labelled: “Nieuw Gunea Exp., KNAG 1939, Koteboe Vallei ann Paniai meer, 24- VIII- 1939” (LEI). Description. — Length 6. 6-6. 9 mm. Antennal Segments I- 1 1 coarsely punctate; Segments III-X very finely punctate; Segment XI impunctate; head longer than broad; median lobe short, broad, rhomboid; posteriolateral margins oblique, tip angulate; frontal space slightly broader than long, V-shaped, its lateral margins oblique, not sinuate; medial angles nearly rectangular, contiguous; posteriomedial margin slightly sinuate; posteriolateral margin evenly curved; occipital angle absent; posterior margin of temporal lobe narrowly bordered with pollinosity; antennal groove narrow, not dilated laterally; orbital groove absent; temporal lobe with five or six punctures near eye; one temporal seta; postorbital tubercle small, about 0.20 as long as eye, visible only in lateral view; eye large, round. Pronotum short; length/greatest width 1.05, widest at base, lateral margins oblique, slightly convergent except near curved apex, markedly convergent; apex markedly narrowed; outer carina slightly narrower than inner carina at middle; Quaest. Ent., 1982, 18 (1-4) 246 Bell and Bell outer carina broadest at base, gradually evenly narrowed from there nearly to more abruptly narrowed apex; inner carina long, oval, tapered to obtuse point at base; pronotal carinae impunctate; pronotum without setae; prosternum without precoxal carinae. Elytra rather narrow, elongate; striae not impressed; strial punctures fine, separated by several times the diameter of one puncture, especially fine in lateral striae and at bases of all striae; elytral intervals flat, iridescent; base of Stria IV with longitudinal scarp; transverse basal scarp of elytron pollinose from base of Stria IV to suture; Stria IV with one seta near apex; subapical stride with one seta; apex of Stria VII with several setae; metasternum with blue reflections, impunctate except for one row of punctures along each lateral margin, cluster of punctures in posterior part of midline; abdominal Sterna III-V with scattered punctures which are not coalescent laterally; both sexes with moderately deep semicircular lateral pit on Sternum IV; male with ventral tooth on anterior femur; female without this tooth; middle calcar acute, about 0.33 as long as spur; hind calcar triangular, its apex angulate, obtuse; distance from tribal spur to proximal end of calcar about 1.5 longer than distance from tip of calcar to its base. A large species with a broad pronotum, resembling O. follis in having a small postorbital tubercle, visible only in lateral view, but differing in having the pronotum broadest at base, the median lobe pointed posteriorly, the striae finely punctate, and in many other details. Omoglymmius ( sensu stricto ) massa new species Figs. 206, 213 Type material - HOLOTYPE male, labelled: “NEW GUINEA (NE), Eliptamin Valley, 1200-1350 m., July 1-15, 1959, W. W. Brandt coll.” (BPBM). PARATYPES one female, same locality and collector as type but dated Aug. 16-30, 1959 (BPBM); one female, Wau, Morobe Dist., 1200 m., 14-IX-1961, Sedlacek & native (BPBM); one female, Koibuga, 1500 m., 3- VII- 1 963, coll. H. W. Clissold (BPBM); one female, 13 km. SE Okapa, 1650-1870 m., 26-VIII-1964, J. & M. Sedlacek(BPBM). Description. — Length 6. 5-8. 2 mm. Antennal Segments I, II coarsely punctate; Segments III-X very finely punctate; Segment XI impunctate; head slightly longer than wide; median lobe short, oval, its apex broadly rounded; frontal space broader than long, V-shaped, its margins oblique, very shallowly sinuate; medial angles obtuse, very narrowly separated; posteriomedial margin oblique; posterior margin transverse, bordered with pollinosity, meeting posteriolateral margin at obtuse angle near temporal seta; antennal groove rather broad, connected to very short orbital groove which ends just posterior to anterior margin of eye; temporal lobe with six to eight very fine punctures near medial margin of eye; one temporal seta or no temporal seta, in some specimens present unilaterally; postorbital tubercle about 0.4 as long as eye, clearly visible in dorsal view; eye large, round. Pronotum, length/greatest width 1.10; widest near middle, base slightly narrowed; apex markedly narrowed; lateral margins slightly curved except near markedly curved apex, convergent; lateral margin scarcely sinuate anterior to hind angle; outer carina slightly narrower than inner carina at middle; outer carina broadest at base, narrowed just anterior to base, then nearly parallel-sided to anterior 0.3, tapered to apex; inner carina long, oval, tapered to point at base; both pairs of carinae impunctate; pronotum without setae; prosternum without precoxal carinae. Elytra rather narrow, elongate; stria not impressed; strial punctures moderately coarse, round; base of Stria IV with longitudinal pollinose scarp; transverse basal scarp of elytron pollinose from base of Stria IV to base of Stria II; Stria IV without seta near apex; subapical striole with one seta; apex of Stria VII with several setae; metasternum with row of punctures along each lateral margin; disc of metasternum impunctate, with bluish opalescence; abdominal Sterna III-V with coarse punctures in form of irregular transverse row on each sternum, punctures coalescent near lateral margin; female with deep oval lateral pit on Sternum IV; male with shallow, ill-defined lateral pit on Sternum IV; male with ventral tooth on anterior femur; female without this tooth; middle calcar very minute, obtuse; hind calcar small, triangular, its apex obtusely angled, its proximal margin longer than distal margin. This species and O. denticulatus have the postorbital tubercle of moderate size, larger than that of the two preceding species, and clearly visible from above, but smaller than in O. auratus. This species differs from O. denticulatus in having the median lobe broadly oval, a slightly larger postorbital tubercle and more produced medial angles on the temporal lobe. Range. - Mountains of northeastern New Guinea. In addition to the type material we provisionally include the following specimens: two males, four females, labelled “Keefu, Okapa, Eastern Highlands, New Guinea, 17-4-1965, coll. R. Hornabrook” (NMNZ); one female, labelled “Morae, 6000, Eastern Highlands, Kuku Kuku, 1-3-64, R. Hornabrook” (NMNZ). Revision of Omoglymmius Ganglbauer 247 Omoglymmius ( sensu stricto) denticulatus new species Figs. 207,214 Type material. - HOLOTYPE female, labelled: “NEW GUINEA (NW), Wissel Lakes, Enarotadi, 1900-2000 m., 2-1 l-VII-62, coll. J. Sedlacek” (BPBM). PARATYPE female, labelled: “N. Guinea, Hatam VII, Beccari, 1875, Rhysodes pulvinatus ” (GEN). It is not conspecific with male lectotype of R. pulvinatus. Description. — Length 7. 7-8. 5 mm. Antennal Segments I, II coarsely punctate; Segments III-X very finely, sparsely punctate; Segment XI impunctate; head slightly longer than wide; median lobe moderately narrow, its tip subtruncate; frontal space longer than wide, V-shaped; lateral margins oblique; medial angles rather sharp, well separated; posteriomedial margin curved, extended to slightly oblique posterior margin; both margins bordered by pollinosity; posteriolateral margin in form of obtuse angle with posterior margin; antennal groove rather broad, slightly dilated at lateral end in form of preorbital impression; orbital groove absent; temporal lobe with six or seven fine punctures in lateral half; one temporal seta; postorbital tubercle about 0.2 as long as eye, clearly visible in dorsal view; eye large, round. Pronotum rather short, length/greatest width 1.14; broadest at base; apex strongly narrowed; lateral margins oblique in posterior half, curved, markedly convergent in anterior half; lateral margin not sinuate anterior to hind angle; outer carina slightly narrower than inner carina at middle; outer carina markedly sinuate just anterior to base, broadest at base, narrowed to point at apex; inner carina long oval, tapered to point at base; both pairs of carinae entirely impunctate; pronotum without setae; prosternum without precoxal carinae. Elytra rather narrow, elongate; striae not impressed; strial punctures moderately coarse, round; base of Stria IV with longitudinal pollinose scarp; transverse basal scarp of elytron pollinose from base of Stria IV to base of Stria I; Stria IV without seta near apex; subapical stride with one seta; Stria VII with several setae near apex; metasternum with row of punctures along each lateral margin; disc of metasternum impunctate, with bluish opalescence; abdominal Sterna III-V with coarse, scattered punctures; punctures coalescent near lateral margin; female with deep, oval lateral pit on Sternum IV; male unknown. This species is most like O. massa of the eastern part of New Guinea, but differs in the size of the postorbital tubercle and the shape of the medial angle of the temporal lobe. It is also similar to O. iridescens, which is sympatric or nearly sympatric with it. This species differs from O. iridescens in having a larger postorbital tubercle. O. auratus , also of West Irian, is also similar, but has the pronotum narrowed at the base and has much larger postorbital tubercles. Omoglymmius (sensu stricto) auratus new species Figs. 208,215 Type material. - HOLOTYPE male, labelled: “NEW GUINEA:Neth., Swart Val: W. ridge 1800-2000 m., Nov. 19, 1958, J. L. Gressitt” (BPBM). Description. — Length 7.0 mm. Antennal Segment I punctate, pollinose; Segments II-IV impunctate; Segments V-X finely punctate; Segment XI impunctate; head, including postorbital tubercles, wider than long; median lobe short, broadly rounded; frontal space V-shaped, longer than wide, margins oblique, scarcely sinuate; medial angles obtuse, separated; posteriomedial margin curved evenly into posteriolateral margin; posterior margin bordered with pollinosity; antennal groove rattier broad, slightly dilated at lateral end; orbital groove absent; temporal lobe with nine or 10 very fine punctures in lateral half; one temporal seta; postorbital tubercle as deep as eye, about 0.9 as long as eye; postorbital tubercles prominent in dorsal view, head width across them greater than width across eyes; eye large, round. Pronotum moderately short, length/greatest width 1.13; widest near middle; base slightly narrowed; apex more markedly narrowed; lateral margins curved; margin not sinuate anterior to hind angle; outer carina slightly narrower than inner one at middle; outer carina broadest near middle, very slightly narrowed at base, strongly narrowed anteriorly, apex obliquely truncate; medial margin of outer carina not sinuate anterior to base; inner carina long-oval, tapered to point at base; both pairs of carinae impunctate; pronotum without setae; prosternum without precoxal carinae. Elytra rather narrow, elongate; striae not impressed; strial punctures very fine, elliptical; base of Stria IV with short pollinose longitudinal scarp; transverse basal scarp of elytron pollinose from base of Stria IV to base of Stria II; Stria IV with one seta at apex; subapical striole with one seta; Stria VII with several setae near apex; metasternum nearly impunctate, with a few fine punctures near lateral margins, microsculptured but not opalescent; abdominal Sterna III-V with one transverse row of coarse punctures on each; those of III, IV widely interrupted at midline; that of V not interrupted; with shallow lateral pit on Sternum IV; with very obtuse ventral tooth on anterior femur; middle calcar small, obtuse; hind calcar small triangular, its apex obtuse, slightly lobate; female unknown. Quaest. Ent., 1982, 18 (1-4) 248 Bell and Bell This species is easily recognized by the large postorbital tubercles. Omoglymmius (sensu strict o) sus new species Figs. 216, 222 Type material. - HOLOTYPE male, labelled: “NUOVA GUINEA, Katau, L.M. D’Albertis” (GEN). PARATYPES one male, two females, same label as holotype (GEN); one female, labelled: “Nuova Guinea, Fly River, L.M. D’Albertis” (GEN); one damaged male with this label is probably conspecific, but we think it best not to designate it as a paratype (MNHN). Description. — Length 6. 8-8.0 mm. Antennal Segments I- VI 1 1 coarsely punctate; Segments IX, X more finely punctate; Segment XI impunctate; head distinctly longer than wide; median lobe lance-shaped, its tip very obtuse; frontal space nearly as long as wide, V-shaped, its margins only shallowly curved; medial angles obtuse, well separated; posteriomedial margin emarginate; posteriolateral margin evenly curved, occipital angle obtuse; antennal groove rather broad, slightly dilated at lateral end; orbital groove absent; temporal with 17-18 fine punctures; one temporal seta; orbital tubercle about 0.3 as long as eye, visible in dorsal view; eye large, round. Pronotum elongate; length/greatest width 1.30; widest near middle; base slightly narrowed; apex very markedly narrowed; lateral margins slightly curved posteriorly, markedly curved anteriorly; margin sinuate anterior to hind angle; outer carina slightly narrower than inner one at middle; medial margin of outer carina angulate, not sinuate just anterior to base; outer carina of nearly even width in posterior half, markedly narrowed near apex; inner carina narrowly subtruncate at base; outer carina with 24-29 fine punctures; inner carina with 19-25 fine punctures; pronotum without setae; prosternum without precoxal carinae. Elytron rather narrow, elongate; striae impressed; strial punctures coarse; base of Stria IV with short longitudinal pollinose scarp; basal transverse scarp pollinose from Stria IV to suture; one seta near apex of Stria IV, one seta in subapical stride; about four setae near apex of marginal stria; metasternum entirely punctate; abdominal Sterna III-V with coarse, scattered punctures; female with deep, narrowly oval, oblique lateral pit on Sternum IV; male with shallow, ill-defined pit on Sternum IV; male with ventral tooth on anterior femur; female with obtuse angle in this position; middle calcar very small, almost absent; hind calcar of moderate size, broadly rounded. Punctures on both inner and outer carinae and the elongate pronotum separate this species from others with a medium-sized postorbital tubercle. Omoglymmius ( sensu stricto) planiceps new species Figs. 217, 223 Type material. - HOLOTYPE female, labelled: “NEW GUINEA, Morobe District, Wau, 2-3-X-1969, James E. Tobler” (CAS). Description. — Length 7.0 mm. Antennal Segments I-X coarsely punctate; Segment XI with few punctures; head distinctly longer than wide; median lobe short, broad, its apex rounded; frontal space slightly wider than long; its margins curved; medial angles nearly rectangular, but blunt; well separated; posteriomedial margin oblique; posteriolateral margin evenly curved; occipital angle distinct; antennal groove short, connected laterally to shallow preorbital impression; very narrow orbital groove traceable almost to posterior margin of eye; temporal lobe with about 12 punctures, most in form of row medial to eye; one temporal seta; postorbital tubercle very short but deep, visible though inconspicuous in dorsal view; temporal lobe markedly flattened in lateral view; eye large, round. Pronotum short, length/greatest width 1.12; widest posterior to middle; base very slightly narrowed; apex strongly narrowed; lateral margins nearly straight posteriorly, curved, convergent anteriorly; margin slightly sinuate anterior to hind angle; marginal groove dilated; in dorsal view, outer carina appearing 0.6 as wide as inner carina at middle; outer carina convex, directed dorsolaterad, apparently narrower in dorsal view than in dorsolateral view; medial margin of outer carina shallowly sinuate just anterior to base; outer carina widest at base; inner carina obtusely pointed posteriorly; both pairs of carinae entirely impunctate; pronotum without setae; prosternum without precoxal carinae. Elytra moderately long; striae not impressed, represented only by rows of fine, shallow, elongate punctures; base of Stria IV with longitudinal pollinose scarp; Stria IV without seta; subapical striole with one seta; Stria VII without setae (or perhaps broken off); metasternum entirely punctate; metasternum, prosternum with faint bluish opalescence; abdominal Sterna III-V with fine, scattered punctures; female with moderately deep semicircular lateral pit on Sternum IV; female without ventral tooth on anterior femur; male unknown. Revision of Omoglymmius Ganglbauer 249 Flattened temporal lobes are distinctive in this species. The dilated marginal groove and narrow, convex outer carina also separate it from any other species from New Guinea. If the very short postorbital tubercle were overlooked, this species would trace to O. trepidus in the key to species from New Guinea. It differs from the latter species in having the temporal lobe with fewer punctures, the outer carina impunctate and narrow and the pronotum not narrowed at the base. Omoglymmius (sensu stricto ) lent us new species Figs. 218, 224 Type material. - HOLOTYPE male, labelled: “D. N. Guinea, Sattelberg” (MNHB). Description. — Length 7.2 mm. Antennal Segments 1-X coarsely punctate; Segment XI with one or two coarse punctures; head 1.3 longer than wide; median lobe broad, its apex obtuse; frontal space much broader than long, nearly U-shaped, its margins abruptly curved; medial angles obtuse, separated; posteriomedial margin oblique, curved gradually into posterior margin; latter nearly transverse laterally, bent abruptly into lateral margin; occipital angle absent; width across temporal lobes greatest just posterior to eyes, margins convergent anteriorly; antennal groove moderately broad, its lateral end slightly dilated; orbital groove absent; temporal lobe with seven or eight punctures near eye; temporal seta apparently absent, but possibly rubbed off in the holotype; postorbital tubercles about 0.33 as long as eye, visible in dorsal view, width across them slightly greater than that across eyes; latter large, round, more protuberant than usual in subgenus. Pronotum short, length/greatest width 1.07; widest near middle; base moderately narrowed; apex very markedly narrowed; lateral margins markedly curved; margin sinuate anterior to hind angle; outer carina 0.6 as wide as inner carina at middle; medial margin of outer carina sinuate just anterior to base; outer carina widest at base, abruptly narrowed anterior to base, from there nearly even width, abruptly narrowed near apex; inner carina with base narrowed, rounded; outer carina with two or three fine punctures; inner carina impunctate; median groove markedly narrowed at middle; pronotum without setae; prosternum without precoxal carinae. Elytron with bluish opalescence; striae not impressed, represented by rows of fine, shallow, elongate punctures; base of Stria IV with oblique pollinose scarp; transverse scarp not pollinose; Stria IV without seta near apex; subapical stride with one seta; Stria VII with several setae near apex; metasternum with bluish opalescence, entirely punctate; abdominal Sterna III-V with fine, scattered punctures; male with shallow lateral pits on Sternum IV; male with ventral tooth on anterior femur; middle calcar acute, small, about 0,25 as long as spur; hind calcar triangular, its apex obtuse, its proxijnaJ margin straight; female unknown. The species closest in appearance to this one is O. massa. In addition to the absence of punctures on the metasternal disc, the latter species differs in the shape of the temporal lobes, the shorter, more rounded median lobe, and the very broad base on the pronotum. The broad based pronotum and flattened temporal lobes distinguish O. planiceps from O. lentus. Omoglymmius (sensu stricto ) capito (Grouvelle) Figs. 219, 225 Rhysodes capito Grouvelle 1895a: 157-158. Omoglymmius (sensu stricto ) capito (Grouvelle) Bell and Bell 1978 Type material. - HOLOTYPE male, labelled: “Nouv. Guinee, Dorey, Baie du Geelvink, Raffray & Maindron 78” (MNHN). Description. — Length 8.3 mm. Antennal Segments V-X coarsely punctate; Segment XI missing in holotype; head as broad as long; median lobe lance-shaped, tip obtuse; frontal space broader than long; nearly V-shaped, margins oblique, shallowly sinuate; medial angles rounded, well separated; posteriomedial margin curved into posteriolateral margin; occipital angle absent; antennal groove moderately broad, not expanded at lateral end; orbital groove absent; temporal lobe with three or four coarse punctures between eye and temporal seta, also about 10 fine punctures more anteriorly, medially; one temporal seta; postorbital tubercles about 0.8 as long as eye, prominent in dorsal view; width across them much greater than across eyes; postorbital tubercles narrower, more divergent than in O. auratus ; eye large, round. Pronotum short, length/greatest width 1.09, widest near middle; base slightly narrowed; apex more markedly narrowed; lateral margin curved, moderately so in basal half, markedly so near apex; margin not sinuate anterior to hind angle; outer carina slightly narrower than inner one at middle; medial margin of outer carina sinuate anterior to base; Quaest. Ent., 1982, 18 (1-4) 250 Bell and Bell outer carina broadest just anterior to base, abruptly narrowed just anterior to it, then slightly broadened to anterior 0.33 of length; apex narrowed; inner carina with base narrowed, rounded; outer carina with 20-30 fine punctures; inner carina impunctate; pronotum without setae; prosternum without precoxal carinae. Elytron rather broad, surface not opalescent, but opaque, faintly microsculptured; striae not impressed, represented by rows of fine, elongate punctures; base of Stria IV with longitudinal pollinose scarp; Stria IV with one seta at apex; subapical stride without seta; Stria VII with several setae near apex; metasternum completely finely, shallowly punctate; abdominal Sterna 11I-V with scattered punctures; male with shallow lateral pit on Sternum IV; male with acute prominent ventral tooth on anterior femur; middle calcar acute, about 0.2 as long as spur; hind calcar triangular, apex nearly rectangular, proximal margin straight; female unknown. The prominent postorbital tubercles of this species are comparable to those of O. auratus. The latter species has postorbital tubercles broader, outer carinae impunctate, the paramedian grooves much narrower, and disc of the metasternum impunctate. Range. - Known only from the type locality, on the Vogelkop Peninsula at the west end of New Guinea. Grouvelle (1903) later cited additional localities. We have not located these specimens, and do not know whether they really belong to this species. Omoglymmius (sensu stricto) cheesmanae (Arrow) Figs. 220, 226 Rhysodes cheesmanae Arrow 1942: 180-181 Omoglymmius (sensu stricto ) cheesmanae (Arrow) Bell and Bell 1978 Type material. - HOLOTYPE male, labelled: “DUTCH N. GUINEA, Cyclops Mtns., Sabron, 930 ft., V-1936, L. E. Cheesman, BM 1936-271” (BMNH). Description. — Length 5. 3-6.0 mm. Antennal Segments I-XI punctate; head distinctly longer than wide; median lobe oval with two to six coarse punctures, tip obtusely pointed; frontal space wider than long; margins moderately curved; medial angles acute, contiguous; posteriomedial margin oblique; posteriolateral margin evenly curved; occipital angle absent; antennal groove narrow; orbital groove rudimentary, ended just posterior to anterior margin of eye; temporal lobe with 15-26 fine punctures; one temporal seta; postorbit flat, pollinose; suborbital tubercle inconspicuous, about 0.2 as long as eye; eye very large, round, prominent. Pronotum moderately elongate, length/greatest width 1.20; widest near middle; base distinctly narrowed; apex markedly narrowed; lateral margins curved; margin not sinuate anterior to hind angle; outer carina slightly narrower than inner one at middle; medial margin of outer carina scarcely sinuate anterior to base; outer carina broadest at anterior 0.33, narrowed to point anteriorly; inner carina with base narrow, attenuate; outer carina with 25-30 fine punctures; inner carina with about six fine punctures; pronotum without setae; prosternum without precoxal carinae. Elytron with striae shallowly impressed; strial punctures coarse, shallow; base of Stria IV with longitudinal pollinose scarp; Stria IV with one seta near apex; subapical stride with one seta; Stria VII with several setae near apex; metasternum completely punctate; abdominal Sterna 111-V with coarse, scattered punctures, these slightly confluent near lateral margin; male with shallow lateral pit on Sternum IV; male with ventral tooth on anterior femur; middle calcar minute, obtuse, almost absent; hind calcar triangular, its apex acute, both margins straight; female unknown. Among the species which have suborbital tubercles, this one has the smallest tubercle. O. pulvinatus and O. asetatus have the tubercles conspicuously larger, and have the posterior margin of the temporal lobe margined with pollinosity, while O. sedlaceki has a much broader pronotum which is not narrowed at the base. Range. - North coast of New Guinea at low elevations. In addition to the holotype, we have Studied the following specimens: one male, labelled “NEW GUINEA (NW), Ifar, Cyclops Mts, 300-500 m„ 23-25-VI-1962, J. Sedlacek” (BPBM); one male, labelled “Maffin Bay, Dutch New Guinea, V 1 1-8- 1 944,coll. E. S. Ross” (CAS). Omoglymmius (sensu stricto ) asetatus new species Figs. 221, 227 Type material. - HOLOTYPE male, labelled: “Madang, N. Guinea, Nov. 196 , Hornabrook coll.” (NMNZ). Revision of Omoglymmius Ganglbauer 251 Description. — Length 6.0 mm. Antennal Segments I-X coarsely punctate; Segment XI impunctate; head distinctly longer than wide; median lobe impunctate, oval, tip obtuse; frontal space twice as wide as long; nearly U-shaped, its margins markedly curved; medial angles obtuse, nearly contiguous; posteriomedial margin oblique, posteriolateral margin evenly curved; occipital angle absent; posterior margin of temporal lobe bordered by pollinosity; antennal groove rather broad; orbital groove short, ended opposite anterior 0.3 of eye; temporal lobe with 22-27 very fine punctures; temporal seta absent; postorbit convex, pollinose; suborbital tubercle both longer and deeper than those of O. cheesmanae , length 0.4 of that of eye; eye very large, round, prominent. Pronotum moderately long, length/greatest width 1.20; widest slightly anterior to middle; base slightly narrowed; apex markedly narrowed; lateral margins curved; margin sinuate anterior to hind angle; outer carina about 0.6 as wide as inner Carina at middle; outer carina with medial margin distinctly sinuate just anterior to base; outer carina with width just anterior to base and width at anterior 0.33 equal; inner carina long, oval, its base narrowly rounded; outer carina with 12-13 fine punctures; inner carina with three to four fine punctures; pronotum without setae; prosternum without precoxal carinae. Elytron with striae shallowly impressed; strial punctures coarse, shallow; base of Stria IV with longitudinal pollinose scarp; Stria IV with one seta near apex; subapical stride with one seta; Stria VII with several setae near apex; metasternum completely punctate; abdominal Sterna III-V coarsely punctate, punctures in irregular transverse row on each sternum; punctures markedly coalescent near lateral margins, in form of indistinct transverse sulci; that of Sternum IV dilated to form indistinct lateral pit; with small obtuse ventral tooth on anterior femur; middle calcar almost absent; hind calcar small, triangular, its apex obtuse; female unknown. This species is similar to O. cheesmanae , but differs in having the posterior face of the temporal lobe pollinose, the suborbital tubercle larger, the temporal seta absent, and the lateral margin of the pronotum sinuate just anterior to the hind angle. Omoglymmius (sensu stricto) biroi new species Figs. 228, 234 Type material. - HOLOTYPE female, labelled: “N. GUINEA Biro’ 1899. Sattelberg, Huon Gulf’ (MNHN). The specimen is labelled as a type of R. pulvinatus , but is not from the type locality of the latter species, Hatam, and is not mentioned in the original description. It is not conspecific with Grouvelle’s type series. PARATYPE female, labelled: “D. N. Guinea, Sattelberg”, collector and data not indicated (MNHB). Description. — Length 6.0-7. 7 mm. Antennal Segments I-X coarsely punctate; Segment XI impunctate; head distinctly longer than wide; median lobe impunctate, suboval, tip pollinose, obtusely rounded frontal space broader than long; margins curved at middle of length; medial angles obtuse, slightly lobate; moderately separated; posteriomedial margin oblique; posteriolateral margin rather suddenly bent near temporal seta, posterior portion nearly transverse; posterior margin of temporal lobe margined with pollinosity; occipital angle obtuse; antennal groove broad; orbital groove broad, ending near middle of eye; temporal lobe with 15-20 fine punctures; one temporal seta; postorbit flat, pollinose; suborbital tubercle small, about 0.3 as long as eye; eye large, round, prominent. Pronotum moderately elongate; length/greatest width 1.21; widest slightly anterior to middle; base slightly narrowed; apex moderately narrowed; lateral margins weakly curved; margins shallowly sinuate anterior to hind angle; outer carina about 0.6 as wide as inner carina at middle; medial margin of outer carina curved, not sinuate anterior to base; outer carina widest at anterior 0.25, narrowed to apex, narrowed near base, but base widened; inner carina with base narrowly truncate; outer carina with 30-35 fine punctures; inner carina with 10-16 fine punctures; pronotum without setae; prosternum without precoxal carinae. Elytron with striae shallowly impressed, intervals flat; strial punctures fine, shallow; base of Stria IV with pollinose longitudinal scarp; transverse basal scarp pollinose from Stria IV to Stria I; Stria IV with one seta near apex; subapical stride with one seta; Stria VII with several setae near apex; metasternum with very fine punctures along lateral margins and in midline, otherwise impunctate; abdominal Sterna III-V with coarse punctures in form of irregular transverse row on each sternum; female with shallow semicircular lateral pit on Sternum IV; female with obtuse ventral tooth on anterior femur; male unknown. Among those species with suborbital tubercles, this one is recognized by the elongate, subquadrate form of the pronotum, and nearly truncate tip of the median lobe. Quaest. Ent., 1982, 18 (1-4) 252 Bell and Bell Omoglymmius (sensu stricto ) pulvinatus (Grouvelle) Figs. 229, 235 Rhysodes pulvinatus Grouvelle 1903: 115-116. Omoglymmius (sensu stricto ) pulvinatus (Grouvelle) Bell and Bell 1978 Type material. - LECTOTYPE (here designated) male, labelled: “Hatam, VII, Beccari, 1875, typus, 6881” (GEN). It bears a label “ Rhysodes pulvinatus ” in Grouvelle’s handwriting. PARALECTOTYPE: one male, same label as lectotype (GEN). A female, with the same labels as the lectotype (GEN), is not this species, but is the species described herein as O. denticulatus new species. Description. — Length 6.3 mm. Antennal Segments I-IX coarsely punctate; Segments X, XI more finely punctate, head distinctly longer than wide; median lobe impunctate, suboval, tip obtusely rounded; frontal space broader than long; margins curved at middle of length; medial angles obtuse, moderately separated; posteriomedial margin oblique; posteriolateral margin evenly curved, margined with pollinosity; occipital angle obtuse; antennal groove broad; orbital groove absent; temporal lobe with 11-13 fine punctures; one temporal seta; postorbit flat, nearly glabrous; suborbital tubercle small, about 0.3 as long as eye; eye large, round, less prominent than in O. biroi. Pronotum rather short; length/greatest width 1.13; widest near middle; base slightly narrowed, apex markedly narrowed; lateral margins rather strongly curved; margin sinuate anterior to hind angle; outer carina about 0.4 as wide as inner carina at middle; medial margin of outer carina curved, scarcely sinuate anterior to base; marginal groove strongly dilated; inner carina narrowed to base; outer carina with 10-15 moderately coarse punctures; inner carina impunctate; pronotum without setae; prosternum without precoxal carinae. Elytron with striae shallowly impressed, intervals flat; strial punctures fine, shallow; base of Stria IV with pollinose longitudinal scarp; transverse basal scarp pollinose from Stria IV to Stria I; Stria IV with one seta near apex; subapical stride with one seta; Stria VII with several setae near apex; metasternum punctate near margins and in midline, otherwise impunctate; abdominal Sterna 1II-V with numerous scattered punctures; with small lateral pit on Sternum IV; anterior femur with ventral tooth; middle calcar minute; hind calcar triangular, obtuse; female unknown. This species differs from others with suborbital tubercles in having the outer carina very narrow, and the marginal groove dilated. O. biroi appears to be the closest related species, but it differs from the latter in having the abdominal punctures numerous and scattered, and in having the pronotum shorter, with more rounded lateral margins. Omoglymmius (sensu stricto ) sedlaceki new species Figs. 230-233, 236, 237 Type material. - HOLOTYPE male, labelled: “NEW GUINEA:(NE), Wau, Morobe Distr. 1200 m., 14-IX-61, coll. J. Sedlacek & native collector” (BPBM). PARATYPES one male, two females, same data as holotype. The following paratypes all have the same locality data as holotype but with the following dates, elevations and collectors: two males, 1400 m., 27-VIII-1961, J. Sedlacek; two males, 1300 m., 27-VII-1961 and 22-XII-1961, J. & J. H. Sedlacek; one female, 1050-1100 m., 1 5-XII- 1 96 1 , G. Monteith, J. Sedlacek; one female, 1650 m., 5-XII- 1961, J. Sedlacek; one female, 1700-1800 m., 17-XI-1961, J. Sedlacek; one male, 1200 m., 1-1-1962, J. & M. Sedlacek; one female, 1200-1700 m., 1 0- VIII- 1 962, J. Sedlacek; one male, 1200-1300 m., 6-IV-1963, J. Sedlacek (ALL BPBM); one female, New Guinea (NE): Wau, Mt. Kaindi, 1500-1800 m., 12-14- VI-1963, coll. P. Shanahan (BPBM); one female, New Guinea (NE), Wau, Kunai Creek, 1500 m., 28-30-V-1963, coll. P. Shanahan (BPBM). Description. — Length 6. 8-8. 4 mm. Outer antennal segments finely punctate; head distinctly longer than wide; median lobe moderately to very broad, its tip obtusely rounded to slightly pointed; frontal space nearly as long as broad; its margin shallowly curved to nearly oblique; medial angles nearly rectangular, well separated; posteriomedial margin deeply emarginate; posteriolateral margin curved; occipital angle distinct, in some specimens lobate; posterior margin between occipital angle and posteriolateral margin oblique in some specimens, emarginate in others; antennal groove moderately Revision of Omoglymmius Ganglbauer 253 broad; orbital groove absent; temporal lobe with nine to 13 fine punctures, mostly near lateral margin; one temporal seta; postorbit flat, glabrous; suborbital tubercle inconspicuous, 0.3 of length of head; eye large, round, prominent. Pronotum rather short, length/greatest width about 1.10; widest near middle, subquadrate; base slightly narrowed, extreme apex markedly narrower; lateral margin feebly curved except near apex, where markedly curved; margin feeble to markedly sinuate anterior to hind angles; width of outer carina subequal to inner one at middle; outer carina sinuate on medial margin just anterior to base; outer carina with two equal points of greatest width, one just anterior to base, another anterior to middle; inner carina with base narrow, attenuate, pointed; outer carina varying in punctuation; inner carina impunctate; pronotum without setae; prosternum without precoxal carinae. Elytron with striae shallowly impressed; strial punctures shallow, rather coarse; base of Stria IV with longitudinal pollinose scarp; transverse basal scarp pollinose between bases of Stria 1-1 V; Stria IV with one seta near apex; subapical stride with one seta; Stria VII with several setae near apex; metasternum with punctures near margins, its disc impunctate; punctures of abdominal Sterna II1-V coarse, sparse, tending to form one transverse row on each Sternum, interrupted at midline; semicircular lateral pit present on Sternum IV in both sexes, deeper in female; male with prominent ventral tooth on anterior femur; female without such tooth; middle calcar scarcely developed; hind calcar triangular, obtuse, its proximal margin nearly straight, its distal margin emarginate between tip of calcar and spur. In this species, the punctures of the metasternum are limited to margins, the postorbit is flat and glabrous, and the outer carina is as wide just anterior to the base as it is at the anterior third. Variations. - We provisionally include in this species a series of allopatric samples which differ from one another in shape of the pronotum, median lobe, and posterior margin of the temporal lobe, as well as in the number of punctures on the pronotum. Possibly these forms are subspecifically or specifically distinct, although it is also possible that there are clinal variations in these characters. We believe that we can distinguish five allopatric forms. Form A, from Wau and vicinity, includes the type series. The posterior margin of the temporal lobe is sinuate near the medial angle but oblique lateral to it, or with sinuation barely suggested. The outer carina has about 20 punctures in its lateral half. Form B, from Kassem, has the temporal lobe as in the preceding, but lacks punctures in the outer carina. Form C, from the Eastern Highlands Province (Kamira, Okapa, Doulo Pass, Mount Michael), has the posterior margin of the temporal lobe twice sinuate, with a lobate occipital angle. The outer carina has only two or three punctures. Form D, from Mount Otto, differs from all others in having a very broad median lobe with a distinct angle at its tip. It has the posterior margin of the temporal lobe twice sinuate, and the outer carina with many punctures. Form E, from Sepalakambang, has the median lobe narrower than in any other form. The posterior margin of the temporal lobe is once sinuate, and the pronotol carinae are impunctate. Range. - Highlands of Northeast New Guinea. In addition to the type series, we provisionally include the following specimens: (FORM B) one male, one female, Kassem Pass, E. Highlands, 24-2-74, coll. R. Hornabrook (NMNZ); (FORM C) two females, Doulo Pass, Asato-Chimbu Divide, 16-1-72 & 13-3-72, coll. R. Hornabrook (NMNZ; one female, Mt. Michael, Lufa, Feb. -71, coll. R. Hornabrook; one male, one female, Okapa, Kamira, 2-9-1964 & 24-6-1965, coll. R. Jornabrook (NMNZ); (FORM D) one female, Mt. Otto, 2200 m„ June 24-1955, coll. J. L. Gressitt (BPBM); (FORM E) one female, Sepalakambang, Salawaket Range, 1920 m., IX- 1 5-56, E. J. Ford, jr. (BPBM). SUBSTITUTIONS FOR PREOCCUPIED GENERIC NAMES Two of the generic names proposed by us in Part I. (Bell and Bell 1978) are preeoccupied. Tangaroa Bell and Bell 1978 is preoccupied by Tangaroa Lehtinnen 1967. We propose the substitute name Tangarona New Name. The type species and only member of the genus is Tangarona pensus (Broun) NEW COMBINATION. Kupea Bell and Bell 1978 is preoccupied by Kupea Philpott 1930. We propose the substitute name Kupeus New Name. The type species and only member of the genus is Kupeus arcuatus Quaest. Ent., 1982, 18 (1-4) 254 Bell and Bell (Chevrolat) NEW COMBINATION. We thank Dr. Norman Platnick for pointing out the first homonymy, and Dr. P. Basilewsky for pointing out the second one. ACKNOWLEDGEMENTS We wish to thank the numerous curators and collectors whose aid has made this work possible. Many of them were mentioned in Parts I and II. We are especially indebted to Dr. Roberto Poggi for detailed drawings and descriptions of some of the types in the Genoa Museum, and for the loan of valuable specimens from the latter museum. Dr. F. Hieke of Humboldt University (Berlin) also made valuable types available to us. We are grateful to Dr. O. Kryzhanovskij of the Academy of Sciences, Leningrad, for sending us samples of species from the U.S.S.R., and for the loan of specimens from other regions. We also acknowledge our debt to Mrs. Joyce Murray for her care in the long and difficult task of typing this manuscript. SUPPLEMENTARY REFERENCES3 Bell, R. T. and J. R. Bell. 1981. Coleoptera: Rhysodidae. Insects of Micronesia 15(2): 51-67. Nakane, T. 1978. New or little-known Coleoptera from Japan and its adjacent regions, XXIX (In English), Reports of the Faculty of Science, Kagoshima University (Earth Sciences and Biology) 11: 129-134. NOTE ADDED IN PROOF Because two specimens designated as holotypes were badly damaged in the mail, replacements are herewith designated: Omoglymmius modicus: HOLOTYPE male, labelled “6-4-62-1455, Savo, Solomon Islands” (BMNH). This is listed on p. 233 as a paratype. Omoglymmius regius: HOLOTYPE female, labelled “Isabela, Tatumba. 24-8-1963, P. Greenslade, 9587” (BMNH). This is listed on p. 235 as a paratype. 3This supplements the list of references found in Part I and Part II. Revision of Omoglymmius Ganglbauer 255 INDEX TO NAMES OF TAXA (Synonyms in italics) FAMILY GROUP TAXA Buprestidae, 206 Cerambycidae, 206 Clavicornia, 206 Omoglymmiina, 131, 164 Rhysodidae, 221 Rhysodini, 162, 232, 233, 235, 238 GENERA AND SUBGENERA Boreoglymmius new subgenus, 132, 133, 140 Caeconavitia new subgenus, 132, 166, 167, 168 Carinoglymmius new subgenus, 132, 176, 177 Dhysores Grouvelle, 164 Hemiglymmius Bell and Bell, 132, 135, 137, 138, 140, 141, 161, 164, 177 Indoglymmius new subgenus, 132, 157, 161, 167 Kupea Bell and Bell, 253 Kupea Philpott, 253 Kupeus New Name, 253 Laminoglymmius new subgenus, 132, 133, 146, 156, 157, 162, 164, 167 Navitia Bell and Bell, 132, 164, 167 Nitiglymmius Bell and Bell, 132, 166, 167, 168, 180 Omoglymmius (sensu stricto), 140, 167, 169, 180, 207,208,211,224, 225 Omoglymmius Ganglbauer, 130, 131, 132, 137, 146, 157, 162, 164, 243 Orthoglymmius Bell and Bell, 132, 133, 161, 164, 169, 177, 180 Pyxiglymmius Bell and Bell, 130, 131, 132, 143, 146, 154, 155, 157 Rhyzodiastes Fairmaire, 177, 227 Tangaroa Bell and Bell, 253 Tangaroa Lehtinnen, 253 Tangarona New Name, 253 SPECIES AND SUBSPECIES actae new species, Omoglymmius, 132, 146, 156, 157, 160, 161 africanus (Grouvelle), Omoglymmius, 133, 135 africanus Grouvelle, Rhysodes , 132 alticola (Grouvelle), Omoglymmius, 170, 173, 175 alticola Grouvelle, Rhysodes , 173 americanus (Castelnau), Omoglymmius, 133, 141, 143, 144, 206 americanus Castelnau, Rhysodes , 140, 144 americanus Reitter, Rhysodes , 194 amplus new species, Omoglymmius, 183, 189, 211, 212 aratus Chevrolat, Rhysodes , 143, 194 aratus Newman, Rhysodes , 144 arcuatus (Chevrolat), Kupeus, 254 armatus (Arrow), Omoglymmius, 147, 150, 153, 154 armatus (Arrow), Pyxiglymmius , 154 armatus Arrow, Rhysodes , 1 54 asetatus new species, Omoglymmius, 186, 193,250 aterrimus (Chevrolat), Omoglymmius, 153 aterrimus Chevrolat, Rhysodes , 153, 154 auratus new species, Omoglymmius, 187, 193,246, 247,250 batchianus (Arrow), Omoglymmius, 184, 190, 223,225,226, 228 batchianus Arrow, Rhysodes , 223 bicarinatus new species, Omoglymmius, 181, 189, 230 biroi new species, Omoglymmius, 186, 193,251,252 borneensis (Grouvelle), Omoglymmius, 133,214 bouchardi new species, Omoglymmius, 184, 189,212,216, 221,222 bucculatus (Arrow), Omoglymmius, 181, 190, 229, 237 Quaest. Ent., 1982, 18 (1-4) 256 Bell and Bell bucculatus Arrow, Rhysodes, 229 caelatus Bell and Bell, Omoglymmius, 183, 194, 232 capito (Grouvelle), Omoglymmius, 187, 194, 249 capito Grouvelle, Rhysodes , 249 carinatus (Grouvelle), Omoglymmius, 177, 179 carinatus Grouvelle, Rhysodes , 176, 179 cavea new species, Omoglymmius, 184, 192, 242, 243 cavifrons (Grouvelle), Omoglymmius, 170, 172, 174, 175 cavifrons Grouvelle, Rhysodes , 174 cheesmanae (Arrow), Omoglymmius, 186, 193, 250, 251 cheesmanae Arrow, Rhysodes , 250 classicus new species, Omoglymmius, 183, 192, 232, 238,239 coelebs new species, Omoglymmius, 185, 189, 209, 215 consors new species, Omoglymmius, 185, 189,214,216, 221 continuus new species, Omoglymmius, 180, 190, 226 coomani (Arrow), Omoglymmius, 170, 174, 175, 176, 179 coomani Arrow, Rhysodes , 176 crassicornis new species, Omoglymmius, 183, 188,211 crassiusculus (Lewis), Omoglymmius, 146, 150, 151 crassiusculus (Lewis), Pyxiglymmius, 151 crassiusculus Lewis, Rhysodes , 143, 151 crenatus (Grouvelle), Omoglymmius, 170, 175, 176 crenatus Grouvelle, Rhysodes , 175 cristatus new species, Omoglymmius, 146, 150, 152, 153 data new species, Omoglymmius, 185, 188, 209,217 denticulatus new species, Omoglymmius, 187, 193,246, 247, 252 duplex new species, Omoglymmius, 180, 184, 189, 209,219, 220 ephemeris new species, Omoglymmius, 185, 192, 244 evasus new species, Omoglymmius, 185, 189,212,213,214,215 exaratus Erichson, Rhysodes , 194 exaratus Serville, Rhysodes , 144 feae (Grouvelle), Omoglymmius, 170, 172, 174, 176 feae Grouvelle, Rhysodes , 172 follis new species, Omoglymmius, 186, 193,245,246 fraudulentus new species, Omoglymmius, 185, 189,212,214,215,217,222 fringillus new species, Omoglymmius, 182, 193,224, 227, 240, 241 fulgens Bell and Bell, Omoglymmius, 168 germaini (Grouvelle), Omoglymmius, 133, 136, 137, 138 germaini Grouvelle, Rhysodes , 137 germari (Ganglbauer), Omoglymmius, 141, 180, 187, 194, 206 germari Ganglbauer, Rhysodes , 131, 180, 194 gorgo new species, Omoglymmius, 155, 158, 160, 163 gracilicornis (Grouvelle), Omoglymmius, 185, 192, 230, 244 gracilicornis Grouvelle, Rhysodes , 244, 245 greensladei Bell and Bell, Omoglymmius, 168 gurneyi new species, Omoglymmius, 182, 191,227,232, 235,236 hamatus (Leconte), Omoglymmius, 133, 141, 143, 145 hamatus Leconte, Rhysodes , 143 hemipunctatus new species, Omoglymmius, 133, 136, 137, 138 hesperus new species, Omoglymmius, 147, 150, 153, 154 hexagonus (Grouvelle), Omoglymmius, 177, 178, 179 hexagonus Grouvelle, Rhysodes , 178 hiekei new species, Omoglymmius, 185, 189, 209,218 Quaest. Ent., 1982, 18 (1-4) Revision of Omoglymmius Ganglbauer 257 hornabrooki Bell and Bell, Omoglymmius, 168 humeralis (Grouvelle), Omoglymmius, 184, 190, 224 humeralis Grouvelle, Rhysodes , 224 ichthyocephalus Lea, Rhysodes , 180 impletus Bell and Bell, Omoglymmius, 186, 194, 231 imugani new species, Omoglymmius, 184, 188, 209,210,211,218 inaequalis new species, Omoglymmius, 132, 157, 158, 160, 161, 167, 177, 178 ineditus (Dajoz), Omoglymmius, 135, 136, 138, 139, 140 ineditus Dajoz, Rhysodes , 138 inermis new species, Omoglymmius, 135, 139, 140 insularis (Grouvelle), Omoglymmius, 146, 157, 160, 162 insularis (Grouvelle), Pyxiglymmius , 162 insularis Grouvelle, Rhysodes , 156, 162 intrusus (Grouvelle), Omoglymmius, 164, 166 intrusus Grouvelle, Rhysodes , 164, 166 iridescens new species, Omoglymmius, 187, 193,245,247 javanicus (Grouvelle), Omoglymmius, 133, 136, 137, 138 krikkeni new species, Omoglymmius, 146, 150, 153, 156 laticeps Bell, Omoglymmius, 181, 188, 207 lederi (Lewis), Omoglymmius, 146, 150, 152, 157 lederi (Lewis), Pyxiglymmius , 152 lederi Lewis, Rhysodes , 152 lentus new species, Omoglymmius, 187, 194, 249 lewisi (Nakane), Omoglymmius, 133, 141, 143, 151,206 lewisi Nakane, Rhysodes , 141 lindrothi new species, Omoglymmius, 183, 192, 231,232, 233 lineatus (Grouvelle), Omoglymmius, 167 lineatus Grouvelle, Rhysodes , 167 longiceps (Grouvelle), Omoglymmius, 170, 173, 174, 176 longiceps Grouvelle, Rhysodes , 173 lustrans Bell and Bell, Omoglymmius, 168 malabaricus (Arrow), Omoglymmius, 181, 187, 207,208 malabaricus Arrow, Rhysodes , 208 malaicus (Arrow), Omoglymmius, 185, 187,214,216 malaicus Arrow, Rhysodes , 215 manni new species, Omoglymmius, 183, 191,232, 234, 235 massa new species, Omoglymmius, 187, 193,246, 247, 249 microtis new species, Omoglymmius, 170, 172, 175 modicus new species, Omoglymmius, 183, 191,232, 233,234 modiglianii new species, Omoglymmius, 183, 189,212 morditus new species, Omoglymmius, 183, 190, 228 mycteroides new species, Omoglymmius, 185, 191,237 nasalis new species, Omoglymmius, 184, 190, 228,237 nemoralis new species, Omoglymmius, 185, 189,213,214,215,222 nicobarensis (Grouvelle), Omoglymmius, 133, 177 nicobarensis Grouvelle, Rhysodes , 161, 111 oberthueri (Grouvelle), Omoglymmius, 133, 157, 160, 162 oberthueri Grouvelle, Rhysodes , 162 occultus new species, Omoglymmius, 133, 136, 138 oceanicus Bell and Bell, Omoglymmius, 184, 194, 231 offafinus Bell and Bell, Omoglymmius, 168 opticus new species, Omoglymmius, 184, 190, 225,226 oroensis new species, Omoglymmius, 182, 193,224, 240, 241 patens new species, Omoglymmius, 186, 192, 231,242, 243 258 Bell and Bell pectoralis new species, Omoglymmius, 185, 189,216,218,219 pensus (Broun), Tangarona, 253 philippensis (Chevrolat), Omoglymmius, 184, 188, 209, 210, 211, 212, 213, 215, 216,217, 220, 231 philippensis (Chevrolat), Rhysodes, 209 philippensis Chevrolat, Rhyzodes, 209 philippinensis (Chevrolat), Rhysodes , 209 pilosus (Grouvelle), Omoglymmius, 146, 150, 153, 155, 156, 161 pilosus (Grouvelle), Pyxiglymmius, 155 pilosus Grouvelle, Rhysodes , 155 planata Fab., Uleiota, 206 planiceps new species, Omoglymmius, 187, 194, 231,248,249 politus new species, Omoglymmius, 181, 184, 188, 189, 210, 218, 230 princeps new species, Omoglymmius, 183, 191, 232, 235, 236, 239 pulvinatus (Grouvelle), Omoglymmius, 186, 193,250, 252 pulvinatus Grouvelle, Rhysodes , 251, 252 puncticornis new species, Omoglymmius, 182, 192, 241 quadraticollis (Arrow), Omoglymmius, 186, 190, 222, 225, 226 quadruplex new species, Omoglymmius, 180, 185, 189,219, 220 regius new species, Omoglymmius, 183, 192, 232, 233,235 renutus new species, Omoglymmius, 184, 191,232, 236, 239 repetitus new species, Omoglymmius, 186, 190, 222, 223 rimatus new species, Omoglymmius, 133, 136, 139 rojasi Chevrolat, Clinidium, 137 rugosus (Grouvelle), Omoglymmius, 146, 147, 157, 160, 163 rugosus (Grouvelle), Pyxiglymmius , 160 rugosus Grouvelle, Rhysodes , 160 rusticus new species , Omoglymmius, 192 rusticus new species, Omoglymmius, 183, 232, 233,234 sakuraii (Nakane), Omoglymmius, 181, 188,207 sakuraii Nakane, Rhysodes , 207 scopulinus new species, Omoglymmius, 182, 191, 232, 235, 237 sectatus new species, Omoglymmius, 185, 192, 242, 243,244 sedlaceki new species, Omoglymmius, 186, 193,250, 252 semioculatus new species, Omoglymmius, 168, 169 semperi new species, Omoglymmius, 182, 188,217,218 silvatica L., Fagus, 206 solitarius (Arrow), Omoglymmius, 132, 178, 186, 187,208 solitarius Arrow, Rhysodes , 208 strabus (Newman), Omoglymmius, 146, 147, 150, 151, 152, 153, 154 strabus (Newman), Pyxiglymmius , 153 strabus Newman, Rhysodes , 146, 153 stylatus new species, Omoglymmius, 164, 165, 166' subcaviceps (Grouvelle), Omoglymmius, 146, 150, 151 subcaviceps (Grouvelle), Pyxiglymmius , 150 subcaviceps Grouvelle, Rhysodes , 1 50 sulcicollis (Lewis), Omoglymmius, 170 sulcicollis Germar, Dechomus, 206 sulcicollis Lewis, Rhysodes , 169, 170 summissus new species, Omoglymmius, 181, 188,212,216,217 sus new species, Omoglymmius, 187, 194, 248 tabulatus new species, Omoglymmius, 186, 191, 238 thoracicus new species, Omoglymmius, 185, 189,216,219 toxopei Bell and Bell, Omoglymmius, 168 trepidus new species, Omoglymmius, 184, 192, 242, 243,249 trisinuatus new species, Omoglymmius, 158, 160, 163, 164 vadosus new species, Omoglymmius, 181, 190, 227,228 Quaest. Ent., 1982, 18 (1-4) Revision of Omoglymmius Ganglbauer 259 vicinus (Grouvelle), Omoglymmius, 182, 192, 239, 240, 241,245 vicinus Grouvelle, Rhysodes, 239 viduus new species, Omoglymmius, 182, 190, 225, 226, 241 wittmeri new species, Omoglymmius, 182, 190, 227 zimmermani Bell and Bell, Omoglymmius, 166, 167, 168 CLASSIFICATION, PHYLOGENY AND ZOOGEOGRAPHY OF THE NORTH AMERICAN SPECIES OF SIPHONA MEIGEN (DIPTERA: TACHINIDAE) J.E. O’Hara Department of Entomology University of Alberta Edmonton, Alberta, Canada T6G 2E3 Quaestiones Entomologicae 18:261-380 1982 ABSTRACT Members of the cosmopolitan genus Siphona are small tachinid flies with a long geniculate proboscis. Their known hosts are Tipula spp. (Tipulidae) and Microlepidoptera ( especially Noctuidae ). Siphonopsis Townsend, with the following three species, is removed from synonymy with Siphona: type-species Siphonopsis plusiae (Coquillett) ( California ), Siphonopsis brasiliensis Townsend ( Brazil ) and Siphonopsis conata Reinhard (California). Two species, Crocuta alticola Mesnil (Burma) and Crocuta crassulata Mesnil (Burma), are placed in Ceranthia s. lat. Siphona diluta Wulp (Mexico) is transferred to Clausicella. Crocuta malaisei Mesnil (Burma) and Crocuta pellex Mesnil (Burma) are placed in Actia. Phantasiosiphona Townsend is synonymized with Siphona, with type-species P. tropica Townsend (Mexico) and P. kuscheli Cortes (Juan Fernandez Islands). Twenty-one Siphona species of North America (America north of Panama-Colombia border) are recognized and described, including 11 new species. Three species described from the Palearctic Region represent new New World records: S. cristata ( Fabr .), S. hokkaidensis Mesnil and S. maculata Staeger. A species described from western United States, S. lurida Reinhard, is a new Palearctic record (from Japan). Two keys are provided for identification of adult specimens, one to species of America north of Mexico and the other to Middle American species. Male and female head profiles and male genitalia are illustrated for all species. Known distributions of North American species are mapped. Six species groups are recognized in North America on the basis of a cladistic analysis of 15 characters of adults. A cladogram is presented for each species group, showing hypothetical phylogenetic relationships among North American species. The S. maculata group is inferred to be the most primitive of the North American species groups. Available data are insufficient to determine phylogenetic relationships among the other species groups. Analysis of Palearctic Siphona species illustrates the general applicability of North American species groups to species in other regions. The following six species groups with their included species are: (1) S. maculata group: S. floridensis n. sp. (type-locality Orlando, FA), S. intrudens (Curran), S. lurida Reinhard, S. maculata Staeger and S. pacifica n. sp. (type-locality Dartford, WA), (2) S. cristata group: S. cristata (Fabr.), S. lutea (Townsend), S. multifaria n. sp. (type-locality mi. 87, Dempster Hwy., Y.T.), S. oligomyia n. sp. (type-locality Keremeos, B.C.) and S. pisinnia n. sp. (type-locality 21km. n. Silver City, NM), (3) S. geniculata group: S. hokkaidensis Mesnil, S. geniculata (De Geer) and S. medialis n. sp. (type-locality Lockeport, N.S.), (4) S. futilis group: S. brunnea n. 262 O’Hara sp. ( type-locality Zontehuitz, Chiapas, Mex.j, S. futilis Wulp and S. illinoiensis Townsend, (5) S. tropica group: S. akidnomyia n. sp. ( type-locality Zontehuitz, Chiapas, Mex.), S. longissima n. sp. ( type-locality lOmi. ne. San Cristobal de las Casas, Chiapas, Mex.), S. rizaba n. sp. ( type-locality Orizaba, Veracruz, Mex.), and S. tropica (Townsend), and (6) S. macronyx group: S. macronyx n. sp. ( type-locality Pullman, WA). S. tenuis Curran is synonymized with S. lutea (Townsend), S. ceres (Curran) with S. futilis Wulp and S. silvarum Herting with S. hokkaidensis Mesnil. Lectotypes are designated for S. futilis Wulp, S. illinoiensis Townsend and S. lutea (Townsend). Three general distribution patterns are evident within species groups of Siphona: (1) America north of Mexico and the Palearctic Region - S. maculata, S. cristata and S. geniculata groups, (2) America north of Mexico and Middle America - S. futilis group, and (3) Middle America and Afrotropical Region - S. tropica group. Within North America, distribution patterns of Siphona species are classed as follows: (1) transcontinental - S. cristata, S. hokkaidensis, S. intrudens, S. lutea, S. maculata, S. medialis and S. multifaria, (2) western - S. geniculata ( introduced ), S. lurida, S. macronyx, S. oligomyia, S. pacifica and S. pisinnia, and (3) Mexican - S. akidnomyia, S. brunnea, S. futilis, S. longissima, S. rizaba and S. tropica. A historical zoogeographic analysis seeks an explanation to these distribution patterns. The patterns are considered in the context of current paleoclimatic and paleogeographic hypotheses to arrive at several conclusions about the historical zoogeography of Siphona species: (1) Siphona is hypothesized to have originated in Africa between the Late Cretaceous and Late Eocene, (2) a S. tropica group taxon was first to enter the New World, following a northern paratropical route between Eurasia and North America, no later than the Late Eocene, (3) the first taxa of the S. cristata, S. geniculata and S. maculata groups migrated to North America through Beringia after the Tertiary climatic optimum, probably no earlier than the Miocene, (4) exchange between Old and New Worlds continued within the S. cristata, S. geniculata and S. maculata groups into the Pleistocene, and (5) three species pairs represent Pleistocene vicariant events: S. maculata and S. floridensis, S. lutea and S. oligomyia, and S. futilis and S. illinoiensis. RESUME Le genre Siphona comprend de petites mouches tachinides caracterisees par un long proboscis genicule. Les holes connus incluent des Tipula spp. (Tipulidae) et des Microlepidopteres (principalement des Noctuidae). Siphonopsis Townsend, contenant les trois especes suivantes, n’est plus considere comme synonyme de Siphona: Siphonopsis plusiae ( Coquillett ) (de Calif ornie), F espece-type du genre, Siphonopsis brasiliensis Townsend ( du Bresil), et Siphonopsis conata Reinhard (de Californie). Les especes Crocuta alticola Mesnil et Crocuta crassulata Mesnil (toutes deux de Birmanie ) sont placees dans le genre Ceranthia s. lat. Siphona diluta Wulp (du Mexique) est transferee dans le genre Clausicella. Crocuta malaisei Mesnil et Crocuta pellex Mesnil (toutes deux de Birmanie ) sont placees dans le genre Actia. Phantasiosiphona Townsend, incluant P. tropica Townsend (du Mexique ), V espece-type, et P. kuscheli Cortes (des lies Juan Fernandez ), est considere comme synonyme de Siphona. Vingt-et-une especes de Siphona, dont II nouvelles, sont reconnues et decrites pour FAmerique du Nord (au nord de la frontiere panamo-colombienne). Trois especes connues precedemment de la region palearctique represented de nouvelles additions a la fauna nearctique: S. cristata (Fabr.), S. hokkaidensis Mesnil et S. maculata Staeger. S. lurida Reinhard, connue de Vouest des Etats-Unis, est nouvellement mentionnee dans la region palearctique (Japon). Deux cles permettent Tidentification des adultes, une pour les especes se trouvant au nord du Mexique, et V autre pour les especes d’Amerique Centrale. Le profit de la tete des males et des femelles, de me me que Torgane genital des males sont illustres pour chaque espece. Des cartes illustrant la repartition geographique connue des especes nord-americaines sont fournies. Une analyse cladistique portant sur 15 caracteres des adultes permet de reconnoitre six groupes d’especes nord-americaines. Pour chaque groupe, les relations phyletiques hypothetiques entre les especes constituantes sont representees par un cladogramme. Le groupe de S. maculata represente probablement le group d’especes North American species of Siphona Meigen 263 nord-americaines le plus primitif. Les donnees disponibles sont insuffisantes pour etablir les relations phyletiques entres les autres groupes d’especes. L' analyse des especes palearctiques de Siphona illustre I'applicabilite generate des regroupements bases sur les especes nord-americaines aux especes des autres regions. Les six groupes, ainsi que leurs especes constituantes, sont: (1) le groupe de S. maculata, comprenant S. floridensis, nouvelle esp'ece (localite du type: Orlando, Floride), S. intrudens (Curran), S. lurida Reinhard, S. maculata Staeger, et S. pacifica, nouvelle espece ( localite du type: Dartford, Washington); (2) le groupe de S. cristata, incluant S. cristata (Fabr.), S. lutea (Townsend), S. multifaria, nouvelle espece (localite du type: mille 87, autoroute Dempster, Yukon), S. oligomyia, nouvelle espece (localite du type: Keremeos, Colombie-Britannique), et S. pisinnia, nouvelle espece (localite du type: 21 km. au nord de Silver City, Nouveau- Mexique); (3) le groupe de S. geniculata, comprenant S. hokkaidensis Mesnil, S. geniculata (De Geer), et S. medialis, nouvelle espece (localite du type: Lockeport, Nouvelle-Ecosse); (4) le groupe de S. futilis, comprenant S. brunnea, nouvelle espece (localite du type: Zontehuitz, Chiapas, Mexique), S. futilis Wulp, et S. illinoiensis Townsend; (5) le groupe de S. tropica, incluant S. akidnomyia, nouvelle espece (localite du type: Zontehuitz, Chiapas, Mexique), S. longissima, nouvelle espece (localite du type: 10 mi. au nord-est de San Cristobal de las Casas, Chiapas, Mexique), S. rizaba, nouvelle espece (localite du type: Orizaba, Veracruz, Mexique), et S. tropica (Townsend); (6) le groupe de S. macronyx, incluant S. macronyx, nouvelle espece (localite du type: Pullman, Washington). S. tenuis Curran est consideree comme synonyme de S. lutea (Townsend), S. ceres (Curran), comme synonyme de S. futilis Wulp, et S. silvarum Herting, comme synonyme de S. hokkaidensis Mesnil. Des lectotypes sont designes pour S. futilis Wulp, S. illinoiensis Townsend et S. lutea (Townsend). Les groupes d’especes de Siphona presentent trois patrons de distribution: (I) des groupes repartis en Amerique du Nord au nord du Mexique et dans la region palearctique, c’est-a-dire ceux de S. maculata, S. cristata et S. geniculata; (2) le groupe de S. futilis, reparti en Amerique du Nord et en Amerique Centrale; (3) le groupe de S. tropica, reparti en Amerique Centrale et dans la region afrotropicale. En Amerique du Nord, les patrons de distribution des especes de Siphona se repartissent comme suit: (I) especes transcontinentales, incluant S. cristata, S. hokkaidensis, S. intrudens, S. lutea, S. maculata, S. medialis et S. multifaria; (2) especes de I'Ouest, avec S. geniculata (introduite), S. lurida, S. macronyx, S. oligomyia, S. pacifica et S. pisinnia; (3) especes mexicaines, avec S. akidnomyia, S. brunnea, S. futilis, S. longissima, S. rizaba et S. tropica. Une analyse historique de la biogeographie tente d’expliquer ces patrons de distribution. En examinant ceux-ci a la lumiere des hypotheses cour antes sur les paleoclimats et la paleogeographie, Thistoire biogeographique des especes de Siphona semble avoir inclu les evenements suivants: (I) Siphona supposement eut son origine en Afrique entre le Cretace superieur et I’Eocene superieur; (2) un taxon du groupe de S. tropica atteignit le premier le Nouveau Monde en suivant un itineraire septentrional a climat paratropical entre VEurasie et I'Amerique du Nord, et ce, avant I’Eocene superieur; (3) les premiers taxons des groupes de S. cristata, S. geniculata et S. maculata atteignirent I’Amerique du Nord via Beringia apres la periode climatique la plus chaude du Tertiaire, soit probablement au plus tot durant le Miocene; (4) les groupes de S. cristata, S. geniculata et S. maculata continuerent d’echanger des elements entre I’Ancien et le Nouveau Monde jusque durant le Pleistocene; (5) finalement, trois paires d’especes, soit S. maculata et S. floridensis, S. lutea et S. oligomyia, et S. futilis et S. illinoiensis, evoluerent a la suite de phenomenes de vicariance qui eurent lieu durant le Pleistocene. Table of Contents Introduction 264 Materials and Methods 265 Materials 265 Methods . . . 267 Genus Siphona Meigen 272 World list of Siphona species 277 Keys to adults of North American Siphona species 281 Key to adults of Siphona species of Canada and United States 281 Key to adults of Siphona species of Middle America 284 The S. maculata group 286 The S. cristata group 292 The S. geniculata group 299 The S. futilis group 305 The S. tropica group 309 The S. macronyx group 312 Quaest. Ent., 1982, 18 (1-4) 264 O’Hara Phylogenetic interpretation 314 Zoogeographic considerations 327 Acknowledgements 336 Literature cited 337 Figures 345 Index 377 INTRODUCTION Specimens of Siphona species are small (3-6mm long) tachinids of cosmopolitan distribution. Like most other tachinids, they spend their three larval instars as internal parasitoids of other insects. Chiefly parasitic on larval Microlepidoptera in general and the Noctuidae in particular (if the meagre published data are any indication), a few species parasitize their distant dipteran relatives, Tipula species (Tipulidae). As parasitoids of other insects, tachinids are highly regarded for their role in the natural environment and their potential for use as control agents against pest insects. When the introduced pest Tipula paludosa Meigen began damaging lawns and meadows in parts of Newfoundland, Cape Breton Island and the lower Fraser Valley of British Columbia, a Palearctic parasitoid was sought for its control. Siphona geniculata (De Geer) was recommended for introduction and became part of a biological control program on both coasts (Wilkinson, 1971). Though little success has been achieved in limiting T. paludosa populations by S. geniculata (Wilkinson, per. comm.), the program illustrates the desirability of an alternative to long term chemical control of a pest species. Whether New World Siphona species are economically important, or have the potential to be so, is unknown. Until now even taxonomic aspects have been neglected, so a framework has not been developed upon which to build a more comprehensive knowledge of the group. This paper represents a step in that direction by providing a revision to North American1 Siphona species. The history of North American Siphona classification has been primarily one of individual species descriptions under varied generic names. Curran (1932) tried to improve the situation through publication of a key to North American Siphona species, but evidently based the key mainly on descriptions. As a result, an assemblage of species belonging to three genera were keyed according to untrustworthy and insignificant characteristics. Furthermore, two described Siphona species were apparently overlooked, for they were not treated in the key. Judged from a historical perspective, the list of eight Siphona species compiled by Sabrosky and Arnaud (1965) represented a significant contribution. It was from this source that my first impression of the number of species and constitution of the genus in the Nearctic Region was gleaned. I was soon to discover the inadequate state of Siphona classification - a result of lack of attention, not expertise. Entwined with species problems was the more basic question of how to define the genus. In answering this question my revision was extended to include Middle America, for it was chiefly through study of Siphonopsis species and allied genera from America north of Panama-Colombia border North American species of Siphona Meigen 265 southwestern United States and Mexico that I recognized the fundamental difference between those taxa and Siphona. A major portion of this paper is concerned with classificatory aspects of North American Siphona species. Also considered are phylogenetic and zoogeographic aspects, with reference to species from other regions. Due to the excellent taxonomic contributions of Dr. L.P. Mesnil (1960, 1961, 1964 and 1965), it has been possible to study interrelationships between North American (particularly Nearctic) and Palearctic Siphona species. Species groups are recognized for the first time in Siphona. Based on cladistic relationships, it is hoped that these North American species groups will have general applicability. MATERIALS AND METHODS Materials This study was based on examination of approximately 4000 adult specimens of Siphona and related siphonines, borrowed from the following institutions and individuals. Abbreviations are those used in the text to indicate deposition of specimens. Institutional loans. - AMNH Department of Entomology, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024; P. Wygodzinsky. BMNH Department of Entomology, British Museum (Natural History), Cromwell Road, London, England SW7 5BD; J.P. Dear. CAES Connecticut Agricultural Experiment Station, 123 Huntington Street, Box 1 106, New Haven, CT 06504; K.A. Welch. CAS Department of Entomology, California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118; P.H. Arnaud, Jr. CFA Coleccion de la Facultad de Agronomia, Universidad de Chile, Casilla 1004, Santiago, Chile; R. Cortes. CNC Biosystematics Research Institute, Central Experimental Farm, K.W. Neatby Building, Ottawa, Ontario K1A 0C6; D.M. Wood. CSU Department of Zoology and Entomology, Colorado State University, Fort Collins, CO 80523; H.E. Evans. CUI Department of Entomology, Cornell University, Comstock Hall, Ithaca, NY 14853; L.L. Pechuman. FSCA Florida State Collection of Arthropods, Division of Plant Industry, Florida Department of Agriculture, Gainesville, FL 32602; H.V. Weems, Jr. INHS Illinois Institute of Natural Resources, Section of Faunistic Surveys and Insect Identification, Illinois Natural History Survey, 172 Natural Resources Building, Urbana, IL 61801; D.W. Webb. KSU Department of Entomology, Kansas State University, Waters Hall, Manhattan, KS 66506; H.D. Blocker. KUR Biological Laboratory, College of General Education, Kyushu University, Ropponmatsu, Fukuoka 810, Japan; H. Shima. MCZ Museum of Comparative Zoology, Harvard University, Cambridge, MA 02138; M.K. Thayer. MSU Department of Entomology, Michigan State University, East Lansing, MI 48824; R.L. Fischer. Quaest. Ent., 1982, 18 (1-4) 266 O’Hara MSUB Department of Biology, Montana State University, Bozeman, MT 59717; S. Rose. NCSU Department of Entomology, Entomology Museum, North Carolina State University, Box 5215, Raleigh, NC 27650; C.S. Parron. NDSU Department of Entomology, North Dakota State University, Fargo, ND 58102; E.U. Balsbaugh, Jr. NSM Nova Scotia Museum, 1747 Summer Street, Halifax, Nova Scotia, B3H 3A6; B. Wright. OKSU Department of Entomology, Oklahoma State University, 501 Life Sciences West, Stillwater, OK 74074; W.A. Drew. OSU Department of Entomology, Oregon State University, Corvallis, OR 97331; M.D. Schwartz. OUCO Department of Entomology, Ohio State University, 103 Botany and Zoology Building, 1735 Neil Avenue, Columbus, OH 43210; C.A. Triplehorn. PSU Frost Entomological Museum, Department of Entomology, Pennsylvania State University, University Park, PA 16802; S.W. Frost. ROM Royal Ontario Museum, 100 Queen’s Park, Toronto, Ontario M5S 2C6; G.B. Wiggins. SMNS Staatl. Museum fur Naturkunde, Stuttgart, D-714 Ludwigsburg, Arsenalplatz 3, West Germany; B. Herting. UAF Department of Entomology, University of Arkansas, Fayetteville, AR 72701; R. Chenowith. UASM Department of Entomology, Strickland Museum, University of Alberta, Edmonton, Alberta T6G 2E3; G.E. Ball. UCD Department of Entomology, University of California, Davis, CA 95616; R.O. Schuster. UCR Department of Entomology, University of California, Riverside, CA 92521; S.I. Frommer. UGA Department of Entomology, University of Georgia, Athens, GA 30602; C.L. Smith. UGG Department of Environmental Biology, University of Guelph, Guelph, Ontario NIG 2W1; S. Marshall. UKL Department of Entomology, Snow Entomology Museum, University of Kansas, Lawrence, KS 66045; C.D. Michener and G.W. Byers. UMW Department of Entomology, University of Manitoba, Winnipeg, Manitoba R3T 2N2; T.D. Galloway. USNM Systematic Entomology Laboratory, SEA, U.S. Department of Agriculture, United States National Museum, Washington, DC 20560; C.W. Sabrosky. USP Museu de Zoologia, Universidad de Sao Paulo, Avenida Nazare, 481, Caixa Postal 7172, 04263, Sao Paulo, SP, Brazil; J.H. Guimaraes. WSUP Department of Entomology, James Entomological Collection, Washington State University, Pullman, WA 99164; W.J. Turner. ZMUC Zoologisk Museum, Universitetsparken 15, Dk 2100, Copenhagen, Denmark; S. Andersen. Loans from private collections. - DMW D.M. Wood, c/o CNC. North American species of Siphona Meigen 267 WLD W.L. Downes, c/o P.L. Fischer, MSU. JEOH designates specimens in my personal collection. Methods Criteria for ranking of taxa. - My concept of a bisexual species is essentially that of Mayr (1969: 26), whereby reproductive isolation of populations is the basic criterion upon which the concept is based. Seldom is it practical, or even possible, for a systematist to field test this hypothesis with phena under revision. Rather, reproductive isolation is inferred from other data. I used two criteria to aid in recognition of species. One was distributional data and the other phenetic divergence. Widely sympatric phena were assumed to represent species, for otherwise interbreeding would presumably have led to deterioration of phenetic difference (barring ecological or temporal barriers). For phenetically similar species like S. cristata and S. multifaria , and S. maculata and S. intrudens, this criterion was especially useful. The degree of phenetic divergence exhibited among sympatric species provided an indication of interspecific variation within Siphona. Along with studies of intraspecific variation, it became possible to distinguish which allopatric phena were sufficiently unique phenetically as to represent species. The subspecific category (as discussed by Mayr, 1969: 187-197) has not been used in this work. It was considered for the polymorphic populations of S. hokkaidensis, but rejected as inappropriate for reasons given under “Variation” of that species. Geographical distributions and structural characteristics of other phena were such that the subspecific category was not a viable alternative. Species sharing one or more derived states and representing apparently monophyletic (Ashlock’s (1971) holophyletic) lineages were arranged under the informal category of species groups. Decisions at the generic level do not figure prominently in this work, so criteria governing generic decisions are only briefly mentioned here. Monophyletic genera are preferred, though paraphyletic genera are considered acceptable. Polyphyletic genera are considered unacceptable. The following are regarded as important generic considerations: (1) distinctiveness from other taxa, (2) degree of intrageneric morphological difference, and (3) number of taxa included (see Mayr, 1969, for discussion). These three parameters are believed to influence the acceptability of paraphyletic taxa in a classification. Cladistic classifications such as those expounded by Eldridge and Cracraft (1980) and Nelson and Platnick (1981) are not used here. Specimen examination. - Pinned, dried specimens were examined with a Wild M5 stereoscopic microscope with standard light source, at magnifications up to 50X. Genitalia were studied with a Leitz SM-LUX compound microscope at magnifications up to 400X. Measurements. - Body length was measured dorsally at SOX through a Wild M5 stereoscopic microscope with an ocular micrometer (100 divisions) calibrated to a slide micrometer (100 divisions to the millimeter). Body length was taken as length from pedicel to scutellum plus abdominal length from attachment to thorax to apex of tergum 5. This measurement eliminated differences due to position of antennae and abdomen relative to rest of body. Quaest. Ent., 1982, 18 (1-4) 268 O’Hara Head measurements of eye and head height and length of first flagellomere were taken in profile, as shown in Fig. 1. Proboscis length was taken as combined length of prementum and labella. Prementum was measured basally from ventral edge of sclerotization to most distal point. Labellar length was measured from dorsal base of labella to labellar tip. Head measurements were taken at 50X with an ocular micrometer, and expressed in text and Hubbs-Hubbs diagrams in ratios. Illustrations are provided with scale bars, which were obtained by projecting an image of a slide micrometer through a camera lucida (see “Illustrations” below) onto the drawing. Hubbs-Hubbs diagrams. - Hubbs-Hubbs diagrams representing intraspecific variation in height of compound eye, lengths of first flagellomere and proboscis (Figs. 13-15), and ratio of length of first flagellomere: eye height (Fig. 16) were prepared. Male specimens were chosen for representation in Hubbs-Hubbs diagrams because their eye and first flagellomere display greater interspecific difference than females. Proboscis length is not sexually dimorphic so measurements from female specimens were included in Fig. 15 for species with few male specimens. Hubbs-Hubbs diagrams are arranged as follows. Species names are along the left margin, abbreviated to their first four letters and listed in order of proboscis length, from shortest to longest. S. hokkaidensis is divided into samples “a” and “b”, as discussed under variation of that species. Number of specimens measured is given along the right margin. The appropriate ratio is along the X-axis. Illustrations. - All illustrations were made with the aid of a camera lucida, as detailed below, and later inked to mylar drafting film for reproduction. Rough head drawings were made with the aid of a camera lucida attached to a Wild M5 stereoscopic microscope, at a magnification of 50X. Rough drawings were redrawn before inking so parts could be reoriented and details altered to best represent the species, and to show characteristics of each species in comparable positions. For male head figures, setae of the ocellar triangle and postcranium were not drawn. To simplify the task of drawing male and female heads of all species, setae were drawn on male head illustrations only. With large variation in eye height and first flagellomere and proboscis lengths within species, head figures should be consulted in conjunction with the Hubbs-Hubbs diagrams to avoid reliance on the figured characteristics of each species. The rostrum and proboscis are shown in a variety of positions. The rostrum is externally membranous so its shape has no taxonomic value. The prementum of the proboscis is shown as a straight cylindrical structure, for in life it is inflexible. This is not true of the labella, the distal half of which is highly manoeuverable in live specimens. This flexibility is evident in its twisted nature in a few drawings, but generally the labella is shown fairly straight to keep drawings standardized. Proboscises as drawn are meant to convey relative differences in length, not shape or structure. Genitalia were temporarily mounted on slides in glycerine jelly2 for illustrative purposes. Once drawings were completed genitalia were washed free of jelly in hot water and returned to glycerine for storage. 2formula of Pantin (1969). North American species of Siphona Meigen 269 Genitalia should not be left for extended periods in glycerine jelly or their complete extraction becomes difficult; ie. the jelly loses its ability to mix freely with hot water and remains coated to the specimen. A Leitz SM-LUX compound microscope with drawing tube was used at a magnification of 100X for rough drawings of all genitalic figures. Rough drawings were redrawn and reoriented for better representation of parts, as mentioned for heads. I did not detect setal differences in male and female genitalia useful for distinguishing species, so setae were not illustrated. All sclerites of female Siphona genitalia from sternum 5 to cerci are shown in Fig. 12. Note that the dorso-longitudinally interrupted tergum of segment 8 is all that remains of terga 6 to 10. Sternum 9 is either lost, or fused with sternum 10. Figures of male genitalia show only those structures associated with segments 9 to 11. In addition to vestiture, the ejaculatory apodeme and paramere (postgonite) are excluded from illustrations. I do not recommend comparison of figured male genitalia on the basis of structures illustrated but not discussed in the text, for differences may be misleading. For instance, hypandrial and aedeagal apodemes and epandrium are considerably varied intraspecifically, so their figured appearance cannot be used for identification purposes. Genitalia were drawn as shown to represent taxonomically useful characters in their proper relative positions as parts of a functional complex. Thoracic and abdominal dorsa (Figs. 2, 7-8) were drawn as described for heads. Hairs, which sparsely cover both regions, were not drawn. Acropods (Figs. 4-6) and the wing (Fig. 3) were drawn with the aid of a drawing tube attached to a SM-LUX compound microscope. Acropods were suspended in glycerine jelly and drawn at a microscope magnification of 400X. Short hairs on claws, pulvilli and tarsomere 5 are not shown. The wing (Fig. 3) was mounted in Canada balsam and drawn at a microscope magnification of 100X. Costal setulae and wing hairs were not illustrated. Terms. - A variety of terms apply to the same structures of adult flies. This developed partly from gradual introduction of new terms by successive generations of dipterists, and partly by establishment of terms unique to particular groups. With the recent publication of the “Manual of Nearctic Diptera, Volume 1”, an attempt was made to standardize terms for structures of adult Diptera (McAlpine, 1981). The depth and detail that pervade this work prompted me to abandon the more familiar terms for tachinids of Crosskey (1973, 1976a) and others in favour of it. Certain of McAlpine’s terms are controversial, and for that reason unacceptable to all, yet acceptance of his work in principle would lead to establishment of a unified system of names for structures. Most structures to which I refer in the text are labelled in Figs. 1-12. Where possible, abbreviations correspond to those in the “Manual of Nearctic Diptera”. The Manual should be consulted for structures not figured. In a few instances I included a familiar term in parenthesis after a Manual term if the feature was not figured. One of my two deviations from terms in the Manual involves a few thoracic setae. I retained use of “humeral” and “presutural” setae. The former was termed “postpronotal” in the Manual, but this seems incongruous without a replacement name for the posthumeral seta, which was not mentioned in the Manual. I see no reason to incorporate the presutural seta into the supra-alar row, so I disregarded this change. Quaest. Ent., 1982, 18 (1-4) 270 O’Hara Certain descriptive terms in the text require explanation to avoid misinterpretation. First of these is colouration, which is difficult to describe qualitatively. This is compounded in Siphona, as in most calyptrate flies, by the effect pruinosity has on ground colour; where pruinosity (pollinosity) refers to “a nap-like covering to any part of the body formed by closely aggregated ultramicroscopic pubescence” (Crosskey, 1973: 19), and ground colour simply refers to the hue of sclerites. Both pruinosity and ground colour contribute to Siphona colouration, but separately described impart little meaning. Therefore I describe “surface colour”, which I define as the hue or hues perceived as a result of light reflected from pruinosity as seen against ground colour. This is essentially synonymous with “colour” of most authors, but here permits distinction between ground and surface colour. In the text, where neither surface nor ground colour is specified, surface colour is implied. Wing colour is varied specifically, but appears very faint from dorsal aspect. To obtain wing colour used in descriptions, I viewed wings from an acute angle to increase colour density and permit differentiation of yellow, brown and cinereous hues. Less ambiguous than colour but requiring explanation are terms used in describing macrotrichia. Here I diverge from the Manual again. I classify macrotrichia as hairs, setulae and setae (bristles). Hairs are macrotrichia of approximately equal diameter along their lengths. Setulae and setae are macrotrichia with thick bases and slender tips, inserted in conspicuous sockets. Setulae and setae are qualitatively separated on basis of size; setulae being small setae. The Manual equated macrotrichia with setae, reserving bristles, setulae and hairs as subordinate categories. This is untenable on two counts. Firstly, “seta” is Latin for “bristle”, so it is incorrect to consider the latter subordinate to the former. Similarly, a setula is by definition a small seta, so a setula should in no instances be larger than a seta. (In the sense of the Manual, a seta can refer to a hair, which is smaller than a setula.) I make use of the terms “weak” and “strong” setae in descriptions. This avoids repeated reference to the length and thickness aspects of setal size, both of which are implied by these terms. Leg setae are named for their positions relative to body axis when a leg is fully extended to the side. Abbreviations used in description of the genus are as follows: a - anterior ad - antero-dorsal av - antero-ventral d - dorsal p - posterior pd - postero-dorsal pv - postero-ventral v - ventral Notes on descriptive format. - Genus and species descriptions are presented in a uniform format, with same style and headings. Differences concern degree of coverage. The aim of the genus description was to describe in detail the attributes of Siphona , without differentiation between character states shared with other genera and states unique to Siphona. Siphona species descriptions do not repeat non-varied attributes of the genus. In most instances a character state or set of states is described in full for each species unless it is shared with a closely related species. For a few characters in which one state is shared by most species. North American species of Siphona Meigen 271 that state is described as average, and average is described in the genus description. Certain characteristics are more useful than others for identification purposes, so a diagnosis is presented prior to each description under the heading “Recognition”, summarizing those attributes by which the species is most reliably recognized. Also included under this heading is a comparison of the described species with other species with which it may be confused. Label data for holotypes and allotypes of new species, and examined type material of described species are presented in a uniform manner. Labels are cited in full, following closely the style of Arnaud (1979). Under this system labels are listed from the top down, with data from each label enclosed in quotation marks. Original spelling and punctuation are preserved, and label lines are delimited by a slash mark(/). Square brackets signify information not included on label. Deposition of each specimen is given in parenthesis. Paratype data of new species were treated less stringently than the above. A standardized format was adopted to increase readability and reduce likelihood of errors. Countries and provinces (or states) are arranged alphabetically; counties, cities, etc. are not. Counties (if given on label) are followed by locality, date of collection, collector and number of males and/or females examined. Specimen repositories are given at the end of each paratype list. Date of collection is given in order of day, month, year, with month in Roman numerals and year abbreviated to the last two digits (though written in full if earlier than 1900). Specimens collected in the same locality on different dates by the same collector are cited together, with commas between dates of collection. Inclusive dates of collection are indicated by dashes. For example, 5,7-9. VII.81 signifies that specimens were collected on the 5th of July and again between the 7th and 9th of July, 1981. I have not presented data about examined, non-type specimens of described species, considering their value insignificant in light of the distribution maps and chart of adult seasonal occurrence. For the sake of posterity a list of these specimens was prepared and deposited in the Archives of the University of Alberta. The list format follows that used for paratypes of new species. Distribution maps. - Localities represented by specimens I personally examined are indicated by dots on the maps. I excluded published locality records because of their unreliability as to species collected. Only New World records of species described herein are mapped. Holarctic species are indicated as such in figure captions and under heading “Distribution” following descriptions. Genitalia dissections. - Dissections were required to permit examination of Siphona male and female genitalia. The technique described below refers to dissection of male genitalia from dried specimens, though it works as well with female and fresh material, with obvious minor changes. This method differs slightly from those reported elsewhere (eg. Beneway, 1963; Wilder, 1979), and was developed to its present form during the course of this study. Dissection of male genitalia involved removal and slight clearing of the abdomen, extraction and clearing of the pregenital ring (sternum 5 and segments 6-8) and genitalia (segments 9 to 1 1), separation of ring and genitalia, and replacement of abdomen on specimen. Details of this procedure follow. Removal of an abdomen from a dried specimen was relatively easy because it naturally tends to break at its connection to the thorax when gently pushed from below. Gentle prodding from above and from side to side helped detach the abdomen cleanly and evenly. The abdomen was placed in a solution of about 10% NaOH until the cuticle became flexible (about 10 minutes, but varied from specimen to specimen). Air trapped in the abdominal cavity Quaest. Ent., 1982, 18 (1-4) 272 O’Hara was released during this time (by gentle squeezing) so that membranes around the genitalia would soften from within as well as from without. This was done carefully and after some flexibility had been attained, for otherwise preabdominal sclerites (segments 1-5) pulled apart. The abdomen was then transferred to 20% acetic acid to neutralize the base. Dissection of the genitalia was done in the acetic acid solution. The bent and blunted tip of an insect pin, attached to a handle, was used to hold the abdomen ventral side up against the dish bottom. In this position only a few minutes were required to slice around syntergosternum 7 + 8 and sternum 5, and pull them and enclosed genitalia free. This operation worked well with a very sharp tungsten steel needle. If tissues were not sufficiently cleared, the preabdominal terga were easily torn; if overcleared there was danger of the terga separating. The genitalia were returned to the NaOH solution until attached membranous tissue changed from translucent to transparent. This step would take from 5 to 20 minutes, depending on the specimen. The preabdomen was taken through 10 minute washes of 70% and 95% ethyl alcohol to toluene. After 20 minutes it was removed, air dried (a matter of seconds) and affixed with shellac glue to the thorax of the specimen from whence it came. Slight shrivelling of the preabdomen usually occurred, but I have been unable to devise a procedure that eliminates this. The toluene step (for which similar solvents can be substituted) yielded better results than the traditional method of alcohol steps only. Sufficiently cleared genitalia were transferred through 5-10 minute washes of 20% acetic acid, 70% and 95% alcohol to glycerine. The high viscosity of glycerine facilitated separation of the epandrium from syntergosternum 7 + 8. This accomplished, the genitalia were easily removed from the ring-like pregenital segments. During the course of this study more than 400 male and about 25 female genitalia of Siphona and related siphonines were dissected. Since study of male genitalia figured prominently in development of my Siphona species concepts, they needed to be readily accessible. To suit this end small vials 1cm in diameter were cut to a height of 7mm and stored in specially constructed trays measuring about 6. 5x6. 5x1. 5cm (outside dimensions). Each tray was fitted with a lid and held 20 vials in 4 rows. To the pin of each dissected specimen a numbered label was attached, and a corresponding number placed with the genitalia in the vial of glycerine. Genitalia were not placed in microvials and stored with their respective specimens until the study was completed. GENUS SIPHONA MEIGEN Aspects of life history Siphona immatures. - Larvae of Siphona species live as internal parasitoids of certain other larval insects. Adult females do not produce many eggs (50-100 in those for which this information is known), which are retained in an elongate, coiled uterus until they have transformed into active larvae {S. [l]cristata, Roubaud, 1906 and Pantel, 1910; undetermined species, Townsend, 1934; S. multifaria and S. maculata , personal observation). They are then deposited onto hosts. Larvae bore through the host integument and feed within the haemocoel, meanwhile procuring air via a posterior connection to a host trachial tube (Roubaud, 1906; Pantel, 1910). When fully mature, Siphona larvae abandon their hosts to seek pupation sites in the ground. North American species of Siphona Meigen 273 Hosts. - Hosts of Siphona species are inadequately known. S. geniculata has been recorded from Tipula paludosa Meigen and T. oleracea L. in Europe (Rennie and Sutherland, 1920). Also recorded from tipulid larvae are S. hokkaidensis ( T . irrorata Meigen, Mesnil, 1965) and S. cristata ( T . maxima Poda [as T. gigantea Schrank], Roubaud, 1906). In addition to the tipulid host, S. cristata has been recorded from ca. 15 noctuid species (see Herting, 1960). However, S. cristata has been confused with S. flavifrons Staeger (and possibly others), so all records are suspect. Siphona maculata has been reared from larvae of Agrotis obelisea and A. candelisequa (Herting, 1960). Also recorded from noctuids are the European species S. collini and S. confusa (Mesnil, 1965), the Mexican species S.futilis (as Bucentes ceres (Curran, 1932)), and S. pseudomaculata Blanchard from South America (Blanchard, 1963). A parasite-host list for European Siphona species is given in Andersen (in press “a”). Adult seasonal occurrence. - A chart of adult Siphona seasonal occurrence is presented in Fig. 17. Most species span three or more months, though I suspect this is in most instances an artifact of supraposition of data from the entire geographic range of each species, rather than an indicator of long adult life or overlapping generations. Indeed, populations near Edmonton with which I am familiar have one generation per year, and adults are seen for only about one month (S. cristata , S. maculata and S. multifaria) . S. geniculata , the only species for which generation time has been reported, is bivoltine (Rennie and Sutherland, 1920) or trivoltine (Andersen, in press “a”). Fig. 17 indicates trends; that is, which species are likely to be found in early spring, the fall, etc. As such the chart doubles as a check of identifications, although its value in this respect is limited for species with inadequate data. Reclassification Siphona is remarkably distinct for a genus with more than 80 described species. The long geniculate proboscis is generally a reliable generic characteristic that permits Siphona specimens to be identified as such with relative ease. However, possession of a Siphona- like proboscis has led to inclusion of a few species in Siphona that belong elsewhere. In this section I discuss removal from Siphona of eight species that superficially resemble Siphona but are more closely related to other taxa3, and I synonymize one genus with Siphona. In 1890 Wulp described Siphona diluta and S. futilis. The syntype series of both species were examined, and S. futilis is retained as a valid Siphona species. S. diluta Wulp is only distantly related to Siphona , belonging to Clausicella Rondani. I remove the genus Siphonopsis Townsend from Siphona , with type-species S. plusiae (Coq.), and two other species, S. brasiliensis Tnsd. and S. conata Rnh. (both originally described under Siphonopsis but later transferred to Siphona by Sabrosky and Arnaud (1965) and Guimaraes (1971)). The proboscises of members of Siphonopsis are slightly elongate, with labella folded back against, but shorter than, prementum, and shorter than eye height. The labella bear pseudotracheae to their base, unlike the Siphona condition wherein the basal half of the labella is fused into a cylindrical sheath. Excluded from consideration are Siphona species not examined during this study; see world list below for species examined. Quaest. Ent., 1982, 18 (1-4) The male genitalia of Siphonopsis species are similar to Siphona except the aedeagus is long and thin and the antero-lateral arms of sternum 5 are hooked inward. A very few Siphona species have similar male fifth sterna ( S . oligomyia n. sp. (Fig. 11), S. cuthbertsoni Curran, and perhaps others I did not examine), but these are almost certainly apotypic and developed independently of Siphonopsis. Recognition of Siphonopsis is expedient. The three described species are closely related, but the generic description does not easily accommodate numerous Mexican and Neotropical species that are yet undescribed. The problem lies not with separation of Siphonopsis and Siphona , but in elucidation of the relationships among Siphonopsis, Pseudosiphona Tnsd., Ceranthia R.D., Aphantorhapha Tnsd. and Asiphona Mesnil. Especially in the New World this group is inadequately known and in need of revision. Further study may reveal that Siphonopsis and perhaps others are best regarded as subgenera of Ceranthia. Two species described from Burma, Crocuta malaisei Mesnil and Crocuta pellex Mesnil, belong to a group of Actia species whose adults possess a long geniculate proboscis (including A. siphonosoma Malloch, A. jocularis Mesnil, A. triseta Mesnil and perhaps others). In this group the length and shape of the proboscis of adults is amazingly like that in Siphona , representing an excellent example of parallel evolution. In other features these Actia species are characteristic of that genus. Mesnil recognized his error and briefly mentioned C. malaisei and C. pellex as Actia species in his description of A. jocularis (1957: 47), and again in his treatment of Palearctic Actia (1963: 813). Crosskey examined the types of both C. malaisei and C. pellex and placed both species back into Siphona (1976a: 214). I have examined a male and female specimen of C. malaisei and I agree with Mesnil’s placement of this species in Actia. From the description of C. pellex and Mesnil’s remarks concerning it, I am confident it, too, belongs in Actia. Two other species described from Burma, Crocuta alticola Mesnil and Crocuta crassulata Mesnil, are placed in Ceranthia s. lat. Adults of both species have prementum and labella elongate, as in Siphona species, but the labella bear pseudotracheae along their lengths, as in Siphonopsis adults. The male genitalia are distinctly Ceranthia-Wke, with broad surstyli (slender in Siphona species) and a single short posteriorly directed seta on each gonopod (long in other examined Ceranthia spp., absent from all examined Siphona spp.). I refer to Ceranthia in the wide sense because the genus has traditionally been comprised only of adults with an average-sized proboscis and very reduced (apotypic) palpus (ie. Ceranthia s. str.). The palpi of C. alticola and C. crassulata adults are average-sized, but I consider the male terminalia with its complex structures more suitable for generic decisions. I include Asiphona in Ceranthia s. lat., for I am unaware of any synapotypies that unite Asiphona species. Asiphona is apparently comprised of species that are phylogenetically close to, but do not belong in, Ceranthia s. str., (cf. Andersen, in press “b”). To Siphona I add one genus, Phantasiosiphona Tnsd., with type-species P. tropica Tnsd. and one other species, P. kuscheli Cortes. Given the spectrum of specific attributes within Siphona, P. tropica is not extraordinary, and does not rank even subgeneric status. P. kuscheli also conforms to the generic concept of Siphona. In summary, Siphonopsis is removed from synonymy with Siphona , along with species Siphonopsis plusiae (Coq.) (1895: 125) (the type-species), S. brasiliensis Tnsd. (1929: 374) and S. conata Rnh. (1959: 162). New combinations for other species removed from Siphona are as follows: North American species of Siphona Meigen 275 Siphona alticola (Mesnil), 1953: 1 10 ( Crocuta (Siphona)), Burma = Ceranthia alticola. Siphona crassulata (Mesnil), 1953: 112 ( Crocuta (Siphona)), Burma = Ceranthia crassulata. Siphona diluta Wulp, 1890: 126 (Siphona), Mexico = Clausicella diluta. Siphona malaisei (Mesnil), 1953: 1 10 (Crocuta (Siphona)), Burma = Actia malaisei. Siphona pellex (Mesnil), 1953: 1 1 1 (Crocuta (Siphona)), Burma = Actia pellex. Generic synonymy of Siphona Meigen A complete list of references to each of the following names was not attempted. Rather, only major catalogues and literature with keys or descriptions pertaining to New World Siphona species are cited. Published state records are largely inaccurate and are not included. Crocuta Meigen, 1800: 39. Type-species, Musca geniculata De Geer (Coquillett, 1910: 528). Suppressed by I.C.Z.N., 1963: 339 (Opinion 678).- Townsend, 1919: 584. Siphona Meigen, 1803: 281. Type-species, Musca geniculata De Geer, 1776, by designation of I.C.Z.N., 1974: 157 (Opinion 1008) (see Sabrosky, 1971, for discussion of the history of Siphona nomenclature).- Wulp, 1890: 1 25. — Townsend, 1891: 368.- Coquillett, 1897: 75.- Aldrich, 1905: 444.- Curran, 1933: 10.— Curran, 1934: 455.- Aldrich, 1934: 108.- Townsend, 1936: 150.- Townsend, 1940: 292.- Reinhard, 1943: 20.- Sabrosky and Arnaud, 1965: 1063.- Cole, 1969: 519, 570.- Cortes and Hichins, 1969: 57.- Cortes and Campos, 1970: 98.- Guimaraes, 1971: 170.- Arnaud, 1978: 458. Bucentes Latreille, 1809: 339. Type-species, B. cinereus Latreille cinereus Latreille, Bucentes (monotypy) = S. geniculata (DeGeer).— Curran, 1932: 13. Phantasiosiphona Townsend, 1915: 93. Type-species P. tropica Townsend (original designation).— Townsend, 1936: 150, 1940: 286.- Cortes, 1952: 1 10.— Guimaraes, 1971: 169. New synonymy. Generic description of Siphona Meigen The following generic diagnosis and description are based on study of adult specimens of North American Siphona species. A few species from other regions do not fit this generic description in all characteristics. Recognition. - Small flies with narrow clypeus, subquadrangular head in profile and a long, geniculate proboscis. Labella slightly longer than prementum, in most species longer than eye height. Second aristomere elongate, at least 2.5X longer than wide. Anal vein extended to wing margin at least as fold. R4+5 setulose dorsally between bifurcation of R2+3 and R4+5 and crossvein r-m (beyond crossvein in S. lutea). Three katepisternal (sternopleural) setae, antero-ventral as strong as or stronger than antero-dorsal. Most species with light coloured pruinosity, particularly on preabdomen, few species darker but not black. Femur and tibia predominantly yellow in most species, tarsi brown to black. Preabdomen without discal setae, T1+2 with or without one pair each median and lateral marginal setae. Female genitalia unmodified (Fig. 12). Male genitalia as in Figs. 1,60-80. Description. - Length: 3. 0-6. 5mm. Male. Head (Figs. 1,18-40). Front: wider than high, 1.1 -1.3 head height; clypeus narrowed, parallel-sided; distance between vibrissae less than half head width (0.33-0.45). Profile: subquadrangular; higher than wide, width at antennal axis 0.60-0.75 head height; frons slightly convex; face vertical to slightly retreated, lower margin protruded beyond vibrissal angle; angle at scape between frons and facial ridge in most species sharply defined, in few species broadly rounded; width of parafacial and gena wide or narrow, inversely proportional to eye size. Colouration: frontal vitta orange or yellow to testaceous, brown or reddish-brown; face, parafacial, gena and lower third or less of postcranium light in ground colour, surface white or light yellow to light brown; ground colour of fronto-orbital plate and upper postcranium light to dark, surface colour varied markedly, from yellow or gold to brown or dark brown, darkest on orbital plate; ocellar triangle dark in ground colour, surface brown to dark brown; antenna highly varied, yellow to black; palpus entirely yellow in most species, apex infuscate in few species; proboscis highly varied, yellow to black. Eye: bare or nearly so; size highly varied, small to large, slender to broad, widest at or above center, narrowed below center or evenly rounded along anterior margin; eye height 0.627-0.91 1 head height (Fig. 13), average height for genus between ca. 0.73-0.82 head height. Quaest. Ent., 1982, 18 (1-4) 276 O’Hara Macrotrichia: five frontal setae, medio-reclinate, anteriormost seta lateral to insertion of first flagellomere; 2nd and 4th frontal setae stronger than 1st, 3rd and 5th; 2 reclinate orbital setae in line with and posterior to frontal setae, anterior seta intermediate in strength between largest frontal seta and inner vertical seta, posterior seta latero-reclinate, in strength intermediate between shortest and longest frontal setae; 2 proclinate orbital setae, subequal in size, not larger than largest frontal seta, lateral to frontal and reclinate orbital setae; anterior reclinate orbital seta from halfway between proclinate orbital setae to lateral to posterior proclinate orbital seta (position intraspecifically varied); inner vertical seta well-developed, subequal in size to vibrissa; outer vertical seta latero-reclinate, subequal in size to lower reclinate orbital seta [appears shorter in most figures because of its inclination]; ocellar triangle with 1 pair latero-proclinate ocellar setae and several hairs; postocellar setae divergent or parallel; paravertical seta subequal in size to setulae of postocular row; postcranium uniformly and sparsely haired; upper parafacial and fronto-orbital plate sparsely haired with less than 10 hairs in most species, to 15 or 20 in S. lurida and S. geniculata\ vibrissal angle with 2-10 setae in addition to vibrissa, and 1 to several hairs, average for genus 1 seta lateral to or above vibrissa and 1 below, and 2 or 3 hairs; 1 subvibrissal seta on antero-ventral margin of gena; aristomere 3 micropubescent; palpus with 2 or 3 setulae on lower surface, and several hairs; proboscis sparsely haired. Antenna: first flagellomere extremely varied in length among species, 0.403-0.740 head height (Fig. 14); shape of first flagellomere in profile from short and slender or broad, to long and slender or broad; length of aristomere 2 proportional to size of first flagellomere, in most species between 2. 5-6.0 times longer than wide, up to 8 times longer than wide in species with very long first flagellomeres (eg. S. tropica, S. pisinnia and S. lurida)-, length of aristomere 3 0.50-0.65 head height in most species, shorter (0.35-0.45) in most specimens of S. hokkaidensis, S. geniculata and S. medialis ; in all species except S. hokkaidensis, aristomere 3 not thickened beyond basal half, distally tapered to a fine tip, in S. hokkaidensis aristomere 3 thickened almost to tip in most specimens. Mouthparts: maxillary palpus slightly clavate, length 0.35-0.55 head height; labella elongate, straight at least on basal half, apical 0.25 (on species with a long proboscis) to 0.5 (species with a short proboscis) flexible in live specimens, twisted on most dried specimens; proboscis length (combined lengths of prementum and labella) varied from 1.3-3. 2 head height (Fig. 15), average for genus between ca. 1.8-2. 5 head height. Thorax (Fig. 2). Colouration: average colouration for genus: ground colour black, with humeral callus and apex of scutellum light in ground colour in most specimens; dorsum blue-gray or gray anteriorly, mixed with brown on posterior prescutum, scutum and scutellum; amount of brown varied within and among species, in most species 3 faint to distinct brown vittae visible on posterior prescutum and scutum; ground colour more apparent on pleura than dorsum, such that light reflected from light gray, blue-gray, yellow or light brown pruinosity and black ground colour produces a sheen - pleura alternately appearing light or dark as specimen rotated. A few species differ from average colouration in possessing yellow thorax (in ground and surface colour). Wing tegula yellow to black; wing membrane yellowish, cinereous or brownish hyaline. Legs, average for genus: femora yellow, or predominantly yellow and distally reddish-brown on mid- and hindlegs and varied amounts of reddish-brown on dorsal and posterior surfaces of foreleg (intraspecifically varied); tibia yellow, or predominantly yellow and basally reddish-brown when femur reddish-brown distally. A few species with greater than average amounts of reddish-brown on femora and tibia, or fuscous overall. Tarsi brown to black. Macrotrichia: setae on prescutum and scutum: acrostichal (weak) 2-3 (presutural), 3-4 (postsutural); dorsocentral 2- 3, 3-4; intra-alar 1,3; supra-alar 3; humeral 2; posthumeral 1; presutural 1; notopleural 2; postalar 2. Scutellum: strong basal, lateral and subapical, weak apical and discal. Pleura: proepisternal (propleural) 1; proepimeral (prostigmatic) 1; anepisternal (mesopleural) row; anepimeral (pteropleural) 1 (weak); katepisternal (sternopleural) 3, postero-dorsal very strong, others weaker, antero-ventral slightly longer than antero-dorsal; meral (hypopleural) row of 3- 4; katepimeron (barette) and anatergite (mediotergite) bare. Sterna: prosternum with a single pair of setulae in most specimens, bare in few (or single setula on one side only). Foreleg: femur, pd row of 3-5 setae; pv surface with 1 to several setae, strongest seta subadjacent to distal pd seta. Tibia, ad and pd surfaces each with row of 2-5 setae; p seta ca. 0.4 tibial length from apex; preapical setae, well-developed d and pv, pd and v weak or absent. Midleg: femur, 2 adjacent a setae at midpoint; ca. 0.1 femoral length from apex, well-developed pd and p setae, weak ad seta; sparse row of setulae on pv surface, with 1 strong setula or seta 0.25 femoral length from base. Tibia, ad seta slightly distad of midpoint; 2 pd setae, ca. 0.3 and 0.6 tibial length from base; v seta ca. 0.6 tibial length from base; preapical setae, well-developed ad, v and pv, other setae, if present, weak. Hindleg: femur, ad row of setae, increasing in strength apically; very strong av seta ca. 0.25 femoral length from apex; 2 v setae basad of midpoint; pd seta ca. 0.1 femoral length from apex. Tibia, ad and pd surfaces each with row of 4-6 setae, alternating weak and strong; 3-5 av setae from ca. 0.4-0. 7 tibial length from base; preapical setae, d, ad, pd and av. Acropod (Figs. 4-6): claws and pulvilli varied in size, subequal on all legs; length of each claw and pulvillus from ca. width of tarsomere 5 (ie. averaged-sized, Figs. 4,6) to ca. length of tarsomere 5 (ie. large, Fig. 5). Wing (Fig. 3): short and broad; Sc incised adjacent to subcostal break; apices of R4+5 and M closely approximated or contiguous in wing margin, slightly anterior to wing tip; anal vein (A,) reaching wing margin at least as fold; R, bare dorsally except for few species with 1 or 2 setulae on bend of R,; R4+5 dorsally with row of 3-10 setulae between bifurcation of R2+3 and R4+5 and crossvein r-m; setulae extended beyond crossvein r-m in most specimens of S. lutea\ CuA, bare dorsally except for 1-3 setulae in very few specimens of S. lutea-, single setula ventrally at bifurcation of R2+3 and R4+5. North American species of Siphona Meigen 277 Preabdomen (Figs. 7-8): Ovoid to slightly elongate in dorsal view; middorsal depression extended to middle of T1+2. Colouration: highly varied intra- and interspecifically; most species dorso-medially vittate, average pattern T4 and T5 black in ground colour, vitta on T]+2 and T3 (width averaging distance between median marginals on T3), rest of preabdomen yellow in ground colour; areas black in ground colour with surface blue-gray, gray, brownish-gray or brown, dark brown to black around macrotrichial insertions; areas yellow in ground colour with surface yellow, light brown or tawny. Preabdomen entirely black in ground colour in a few species, at least dorsally and laterally, or with very reduced areas of yellow ground colour on extreme lateral edge of T] + , and antero-lateral edge of T3; preabdomen entirely yellow in ground and surface colour in a few other species. Macrotrichia: discals absent; T1+2 without median marginals in most species, 1 seta on one side or 1 pair present in few species; T,+2 with 1 pair strong lateral marginals in most species (ca. 0.75 length of lateral marginals on T3 or longer), weak pair in few species (less than 0.5 length of lateral marginals on T3), 2 pair strong lateral marginals in few specimens of few species; T3 with 1 pair each strong median and lateral marginals in most species, few specimens of few species with 2 pair lateral marginals; T4 with 3 pair marginals; T5 with 3-5 pair marginals. Genitalia (Figs. 9,60-80). Epandrium in profile saddle-shaped, slightly higher than long, antero-lateral margin with medially-angled flange; hypandrial apodeme articulated with antero-ventral margin of epandrium; gonopod sub-triangular, with a flange produced ventro-laterally; paramere lobe-like, ca. twice as long as wide; surstylus narrow, apex even with or extended beyond apex of cerci; cerci fused medially along lower half, long and narrow, smoothly tapered with slight inflexion at midpoint, apex curved anteriorly; epiphallus not developed. Aedeagus: distiphallus in profile triangular to quadrangular, surface Finely dentate, antero-ventral margin from dentate to bearing well-developed hooks; angle of distiphallus from even with angle of basiphallus (eg. S. illinoiensis) to sharply bent downward (eg. S'. intrudens)\ apex in profile from narrow (eg. S. tropica) to broad (eg. S. medialis). Sternum 5 (Figs. 10-11): posterior margin U-shaped (not V-shaped as in most Actia species); notched postero-medially; all North American species except S. oligomyia with postero-lateral arms smoothly curved around posterior margin, S. oligomyia with arms hooked inward. Female. As described for male except as follows. Head (Figs. 41-59). Eye and first flagellomere smaller than in male; female palpus of few species longer than in male. Thorax. Acropod: smaller than in males; most species with large claws and pulvilli in male have only slightly larger than average claws and pulvilli in female. Preabdomen. Colouration: same range as described for male; female darker than male in most species; females of species with vittate males in most species with more extensive areas of black ground colour than males, vitta shorter and broader than in males, or absent. Genitalia (Fig. 12). Unmodified; structurally very similar among Siphona females; terga 6, 7, 9 and 10 lost; tergum 8 dorso-longitudinally interrupted; sternum 9 lost, or fused with sternum 10 (ref. Herting, 1957); cerci well developed; two spermathecae. World list of Siphona species This list is inclusive of all Siphona species described before August, 1981, plus those described as new herein4. The genus in which each species was described is given only if not Siphona. A capital letter to left of species name denotes the region from which the species was described with, for a few species, a second letter denoting presence of the species in another region. Species of which specimens were examined during the course of this revision are so indicated in the list. Unless otherwise stated, specimens examined were not type material. Status of species not examined follows recent revisionary or catalogue placements. New species, synonymies, combinations and lectotype designations are noted. Types of nominal species of Siphona described by Robineau-Desvoidy (1830, 1850) are lost (Herting, per. comm.), and although most were probably conspecific with S. geniculata, their status cannot be verified so the names are treated as nomina dubia. Tour new species of European Siphona are described by Andersen (in press “a”). They are not listed here to ensure the new names are first published in his paper. Quaest. Ent., 1982, 18 (1-4) 278 O’Hara Regional code: A Afrotropical, sensu Crosskey and White, 1977. N North America (America north of Panama-Colombia border). O Oriental, sensu Crosskey, 1976a. P Palearctic, as limited by Afrotropical and Oriental regions in their present interpretation. S South America. I have examined specimens of a new species of Siphona from Australia, but as yet no Siphona have been described from that region. The tribe Siphonini is apparently not represented in New Zealand (Dugdale, 1969). A abbreviata (Villeneuve), 1915: 199 ( Bucentes ) [examined], N akidnomyia new species. A albocincta (Villeneuve), 1942: 55 ( Bucentes ) [examined], A amoena (Mesnil), 1952: 12 ( Crocuta ) [examined]. A amplicornis Mesnil, 1959: 21. syn. nigrohalterata Mesnil, 1959: 22 (as ssp.). A angusta Mesnil, 1959: 22. A antennalis (Mesnil), 1952: 9 ( Crocuta ) [examined], A atricapilla Mesnil, 1959: 20. A bevisi Curran, 1941: 7 [holotype examined]. A bilineata (Mesnil), 1952: 10 ( Crocuta ) [examined]. P boreata Mesnil, 1960: 190 [holotype examined], N brunnea new species. A capensis Curran, 1941: 7. P collini Mesnil, 1960: 188 [holotype examined]. P confusa Mesnil, 1961: 201 [holotype examined]. A cothurnata (Mesnil), 1952: 17 ( Crocuta ) [examined]. A creberrima (Speiser), 1910: 142 {Crocuta). P,N cristata (Fabricius), 1805: 281 ( Stomoxys ) [lectotype examined], syn. chaetolyga Rondani, 1865: 194. palpina Zetterstedt, 1859: 6064. A cuthbertsoni Curran, 1941: 7 [holotype examined]. P efflatouni Mesnil, 188 [holotype examined]. P flavifrons Staeger in Zetterstedt, 1849: 321 1 [examined]. N floridensis new species. O foliacea (Mesnil), 1953: 1 13 {Crocuta {Siphona )) [examined]. A fuliginea Mesnil, 1977: 77. ssp. cerina Mesnil, 1977: 76. ssp. rubea Mesnil, 1977: 77. N futilis Wulp, 1890: 125 [lectotype designated]. syn. ceres (Curran), 1932: 14 ( Bucentes ) [holotype examined, new synonymy]. O gedeana Wulp, 1896: 109. syn. nigripalpis (de Meijere), 1924: 223 {Bucentes). P,N geniculata (De Geer), 1776: 20 {Musca) [examined], syn. analis Meigen, 1824: 157. North American species of Siphona Meigen 279 cinerea Meigen, 1824: 156. cinerea (Latreille), 1809: 339 ( Bucentes ). meigenii (Lepeletier and Serville in Latreille, et al.), 1828: 501 {Bucentes). minuta (Fabricius), 1805: 282 ( Stomoxys ). nigrovittata Meigen, 1824: 157. tachinaria Meigen, 1824: 156. urbanis (Harris), 1780: 153 ( Musca ). A gracilis (Mesnil), 1952: 13 ( Crocuta ) [examined], P grandistyla Pandelle, 1894: 108 [examined]. P,N hokkaidensis Mesnil, 1957: 36 [holotype examined], syn. silvarum Herting, 1967: 9 [holotype examined, new synonymy]. N illinoiensis Townsend, 1891: 368 [lectotype designated]. N intrudens (Curran), 1932: 14 ( Bucentes ) [holotype examined], A janssensi (Mesnil), 1952: 4 ( Crocuta ) [examined]. S kuscheli (Cortes), 1952: 1 10 ( Phantasiosiphona ) [examined, new combination]. A laticornis Curran, 1941: 9. A lindneri Mesnil, 1959: 22. N longissima new species. N,P lurida Reinhard, 1943: 20 [holotype examined], N lutea (Townsend), 1919: 584 ( Crocuta ) [lectotype designated]. syn. tenuis Curran, 1933: 10 [holotype examined, new synonymy]. N macronyx new species. P,N maculata Staeger in Zetterstedt, 1849: 3212 [holotype examined]. ssp. griseola Mesnil, 1970: 1 18 [holotype examined]. P maculipennis Meigen, 1830: 365 (unrecognized). N medialis new species. A melania (Bezzi), 1908: 58 {Bucentes). A melanura Mesnil, 1959: 23. N multifaria new species. A munroi Curran, 1941: 6. A murina (Mesnil), 1952: 15 {Crocuta) [examined]. P nigricans (Villeneuve), 1930: 100 {Bucentes) [holotype examined]. A nigroseta Curran, 1941: 8. O nobilis (Mesnil), 1953: 1 12 {Crocuta {Siphona)). A obesa (Mesnil), 1952: 8 {Crocuta) [examined], A obscuripennis Curran, 1941: 8 [holotype examined]. N oligomyia new species. N pacifica new species. P paludosa Mesnil, 1960: 188 [examined]. A patellipalpis (Mesnil), 1952: 10 {Crocuta) [examined], P pauciseta Rondani, 1865: 193 [examined]. syn. delicatula Mesnil, 1960: 190. oculata Pandelle, 1894: 108. A phantasma (Mesnil), 1952: 7 {Crocuta) [examined]. A pigra Mesnil, 1977: 78. Quaest. Ent., 1982, 18 (1-4) 280 O’Hara N pisinnia new species. S pseudomaculata Blanchard, 1963: 233. A reducta (Mesnil), 1952: 18 ( Crocuta ) [examined]. ssp. ludicra Mesnil, 1977: 78. N rizaba new species. P rossica Mesnil, 1961: 202 [examined]. A rubrapex Mesnil, 1977: 79. A rubrica (Mesnil), 1952: 11 ( Crocuta ) [examined]. A setinerva (Mesnil), 1952: 16 ( Crocuta ) [examined], P setosa Mesnil, 1960: 191 [examined], P seyrigi Mesnil, 1960: 189 [holotype examined]. A simulans (Mesnil), 1952: 18 {Crocuta). S singulars (Wiedemann), 1830: 335 ( Tachina ) (unrecognized). A sola Mesnil, 1959: 21 [examined]. A spinulosa (Mesnil), 1952: 12 ( Crocuta ) [examined], A trichaeta (Mesnil), 1952: 18 ( Crocuta ) [examined]. N,S tropica (Townsend), 1915: 93 ( Phantasiosiphona ) [paratype examined, new combination]. A unispina (Mesnil), 1952: 14 {Crocuta) [examined]. syn. infuscata (Mesnil), 1952: 14 ( Crocuta , as ssp.). A vittata Curran, 1941:8 [holotype examined]. A vixen Curran, 1941: 9 [holotype examined]. A wittei (Mesnil), 1952: 5 {Crocuta) [examined]. Nomina Dubia analis Robineau-Desvoidy, 1830: 92. clausa Robineau-Desvoidy, 1850: 209. consimilis Robineau-Desvoidy, 1850: 205. fuscicornis Robineau-Desvoidy, 1850: 205. humeralis Robineau-Desvoidy, 1850: 207. melanocera Robineau-Desvoidy, 1850: 206. pusilla Robineau-Desvoidy, 1830: 92. syn. persilla. Misspelled reference to pusilla Robineau-Desvoidy (Coquillett, 1897: 76). quadrinotata Robineau-Desvoidy, 1850: 203. syn. quadricincta. Misspelled reference to quadrinotata Robineau-Desvoidy (Rondani, 1859: 10). silvatica Robineau-Desvoidy, 1850: 208. testacea Robineau-Desvoidy, 1850: 207. tristis Robineau-Desvoidy, 1850: 203. Nomina Nuda anthomyformis Lynch Arribalzaga in Brauer, 1898: 505 (1898: 13). dorsalis Brauer and Bergenstamm, 1891: 410. pauciseta Mesnil, 1964: 856 (as ssp. of geniculata). taiwanica (Baranov in Hennig), 1941: 195 {Crocuta). North American species of Siphona Meigen 281 Keys to adults of North American Siphona species Siphona species are not sufficiently different and are too intraspecifically varied to permit construction of a key based solely on external, diagnostic characteristics. Therefore I found it necessary to frequently employ mensural characteristics (based on data presented in Hubbs-Hubbs diagrams Figs. 13-16) and features of male genitalia in the following keys to North American Siphona species. As a consequence, female specimens do not key as accurately as males, and should only be used if male associates are unavailable. Even males of S. cristata and 5*. multifaria cannot always be separated (couplet 19"), while a few specimens of other species may be outside the recorded range of variation. Given certain conditions it should be possible to identify most Siphona specimens. Firstly, males are more reliably keyed than females. Secondly, it is advisable to check characteristics of several male specimens of the same species against values given in the Hubbs-Hubbs diagrams to confirm a species identification. Thirdly, the “Recognition” section under each species treatment has been included to assist identifications by summarizing key features of a description and contrasting similar species. To simplify keying of North American Siphona specimens, species were divided into two geographical groups and keyed separately: (1) species of Canada and the United States, and (2) species of Middle America. S. pisinnia is shared between regions and is keyed twice. S. multifaria is recorded almost to the Mexican border in the United States, so it is included in the key to Middle American species. The Siphona fauna of Middle America is inadequately known. As more specimens of known species become available the key will probably require modification to incorporate greater intraspecific variation. Similarly, the key will have to be modified for inclusion of currently undiscovered species. Most terms are as used in the “Manual of Nearctic Diptera, Volume 1” (McAlpine, 1981). See “Terms” section and genus description for information regarding Siphona characters. Figure numbers of male and female head profiles are cited following each species name. Male genitalia of all species are shown in Figs. 9, 60-80. The geographical distribution of each species is given as an aid to identification of specimens. Key to adults of Siphona species of Canada and United States 1 T1+2 with one pair strong median marginal setae (Fig. 8) 2 V T1+2 without median marginal setae (Fig. 7), or one strong seta on one side only, or one pair scarcely differentiated from adjacent marginal hairs 6 2 (1) Surstylus with apex even with or extended slightly beyond apex of cerci (Fig. 72); distiphallus slightly or sharply bent downward from angle of basiphallus; specimens from Washington, Idaho, Oregon and California with femora and tibiae reddish-brown to fuscous, specimens from other areas with dorsal and posterior surfaces of fore femur reddish-brown to fuscous in few specimens, other specimens with legs average (ie. femoral-tibial articulation reddish-brown, rest of femur and tibia yellow); transcontinental, most widely recorded from western USA (Fig. 94) S. medialis n. sp. (Figs. 29, 52), p. 303 2' Surstylus with apex extended beyond apex of cerci by more than preapical width of surstylus (Figs. 70-71); slope of distiphallus even with that of basiphallus; leg colouration average 3 Quaest. Ent., 1982, 18 (1-4) 282 O’Hara 3 (2') Male 4 3' Female 5 4 (3) Distiphallus parallel-sided in profile, slender in anterior view (Fig. 70); aristomere 3 tapered to fine tip; first flagellomere short and narrow, length 0.42-0.48 head height; introduced to, and only recorded from lower, Fraser Valley of British Columbia (Fig. 91) S. geniculata (De G.) (Fig. 28), p. 299 4' Distiphallus slightly broader at base than apex in profile, average Siphona width in anterior view (Fig. 71); aristomere 3 thickened to near tip in most specimens; first flagellomere varied, from short and narrow to long and broad, length 0.43-0.67 head height; transcontinental (Fig. 81) S. hokkaidensis Mesn. (in part) (Figs. 30-32), p. 301 5 (3') First flagellomere with apex truncate in most specimens; transcontinental (Fig. 81) S. hokkaidensis Mesn. (in part) (Fig. 51), p. 301 5' First flagellomere with apex rounded along ventral margin; introduced to, and only recorded from, lower Fraser Valley of British Columbia (Fig. 91) S. geniculata (De G.) (Fig. 50), p. 299 6 (T) Body pale yellow overall in ground colour 7 6 ' Body reddish-brown to black in ground colour at least on thoracic dorsum 8 7 (6) Male sternum 5 with antero-lateral arms not hooked inward (Fig. 10); proboscis shorter than 2.2X head height; most specimens with R4+5 setulose beyond crossvein r-m on at least one side; transcontinental (Fig. 89) S’, lutea (Tnsd.) (Figs. 26,48), p. 296 1' Male sternum 5 with antero-lateral arms hooked inward (Fig. 11); proboscis longer than 2.3X head height; R4+5 not setulose beyond crossvein r-m; western North America (Fig. 90); female unknown . S. oligomyia n. sp. (Fig. 27), p. 297 8 (60 Proboscis shorter than 1.8X head height 9 8' Proboscis longer than 1.8X head height 13 [S. intrudens keys through both halves of couplet] 9 (8) Vibrissal angle with 5-10 setae in addition to vibrissa and several hairs; eye very small, in male less than 0.68 head height; T1+2 with strong lateral marginal setae; western USA (Fig. 86) S. lurida Rnh. (Figs. 21, 41), p. 290 9' Vibrissal angle with 2-3 setae in addition to vibrissa and several hairs; male eye larger than 0.70 head height; T1+2 with weak or strong lateral marginal setae 10 10 (9') Four postsutural dorsocentral setae; distiphallus dentate along antero-ventral margin (Fig. 61); Florida and southern Georgia (Fig. 84) S. floridensis n. sp. (Figs. 19, 43), p. 288 10' Three postsutural dorsocentral setae; distiphallus with several distinct, small or large hooks along antero-ventral margin (Figs. 60, 62) 11 11 (100 Male 12 1 V Female S. maculata Staeg. (Fig. 42), p. 286 and S. intrudens (Cn.) (Fig. 44), p. 288 [separation difficult without male associates; see discussion under S. maculata, “Recognition”] 12 (11) Distiphallus U-shaped in anterior view, with 3-4 large hooks along antero-ventral margin (Fig. 62); preabdomen black in ground colour overall, or North American species of Siphona Meigen 283 vittate and yellow in ground colour laterally on T1+2 and T3; transcontinental (Fig. 85) S. intrudens (Cn.) (in part) (Fig. 20), p. 288 12' Distiphallus V-shaped in anterior view, with several small hooks (Fig. 60); preabdomen vittate, yellow in ground colour laterally on T1+2 and T3; transcontinental (Fig. 82) S. maculata Staeg. (Fig. 18), p. 286 13 (8') Distiphallus U-shaped in anterior view, with 3-4 large hooks along antero-ventral margin (Fig. 62); 3 postsutural dorsocentral setae; T1+2 with weak lateral marginal setae in most specimens; proboscis shorter than 2. OX head height; transcontinental (Fig. 85) S. intrudens (Cn.) (in part) (Figs. 20, 44), p. 288 13' Distiphallus V-shaped in anterior view, anteroventral margin dentate or with small hooks; 3 or 4 postsutural dorsocentral setae; T1+2 with weak or strong lateral marginal setae, if weak (some specimens of S. pacifica ), then 4 postsutural dorsocentral setae; proboscis longer than 1.85X head height 14 14 (130 Male 15 14' Female 15 15 (14) Surstylus with apex extended beyond apex of cerci by more than preapical width of surstylus (Fig. 71); short, thick aristomere 3 (0.35-0.45 head height) in most specimens; most specimens from eastern North America with broad, truncate first flagellomere (Fig. 31), specimens from elsewhere with first flagellomere varied (Figs. 30,32); transcontinental (Fig. 81) S. hokkaidensis Mesn. (in part) (Figs. 30-32), p. 301 15' Surstylus with apex even with or extended slightly beyond apex of cerci; aristomere 3 of average length (0.50-0.65 head height), tapered to fine tip; first flagellomere with shape varied 16 16 (15') Length of first flagellomere less than 0.50 head height; western USA 17 16' Length of first flagellomere greater than 0.50 head height; widely distributed . . 18 17 (16) Three postsutural dorsocentral setae; tarsal claws large (as in Fig. 5); vibrissal angle with 3-6 setae and several setulae in addition to vibrissa and hairs; distiphallus sharply bent downward from basiphallus, with small hooks on antero-ventral margin (Fig. 75); head with characteristic habitus in profile; recorded from Washington and western Idaho (Fig. 88) S. macronyx n. sp. (Fig. 33), p. 313 17' Four postsutural dorsocentral setae; tarsal claws of average size (as in Figs. 4,6); vibrissal angle with 2-3 setae in addition to vibrissa and hairs; distiphallus bent slightly downward from angle of basiphallus, dentate on antero-ventral margin (Fig. 64); recorded from California and Washington (Fig. 87) S. pacifica n. sp. (Fig. 22), p. 291 18 (17') Proboscis shorter than 2.3X head height; length of first flagellomere greater than 0.76 eye height; first flagellomere fuscous to black; southwestern USA and Mexico (Fig. 83) S. pisinnia n. sp. (Fig. 23), p. 298 18' Proboscis longer than 2.25X head height, or from central or eastern North America ( S . illinoiensis ); length of first flagellomere less than 0.85 eye height; first flagellomere varied in colour, testaceous to black 19 19 (18') Distiphallus narrow in profile, apex sloped sharply postero-ventrally in most specimens (Fig. 9); slope of distiphallus even with that of basiphallus in most Quaest. Ent., 1982, 18 (1-4) 284 O’Hara specimens, slightly bent in few; proboscis shorter than 2.5X head height; length of first flagellomere less than 0.69 eye height; central and eastern North America (Fig. 92) S. illinoiensis (Tnsd.) (Fig. 1), p. 307 19' Distiphallus broad in profile, apex not sloped sharply postero-ventrally in most specimens (Figs. 66-67); distiphallus bent sharply downward from angle of basiphallus; proboscis longer than 2.25X head height; length of first flagellomere greater than 0.65 eye height; transcontinental (Figs. 93, 95) S. cristata (Fabr.) (Fig. 24), p. 292 and S. multifaria n. sp. (Fig. 25), p. 293 [see “Recognition” under S. cristata for discussion concerning separation of these species] 20 (14r) First flagellomere truncate apicaily; aristomere 3 short (0.35-0.45 head height) in most specimens; transcontinental (Fig. 81) S. hokkaidensis Mesn. (in part) (Fig. 51), p. 301 20' First flagellomere rounded distally along ventral margin; aristomere 3 average length (0.50-0.65 head height) 21 21 (20') Vibrissal angle with 3-6 setae and several setulae in addition to vibrissa and hairs; 3 postsutural dorsocentral setae; recorded from Washington and western Idaho (Fig. 88) S. macronyx n. sp. (Fig. 56), p. 313 2 1' Vibrissal angle with 2-3 setae in addition to vibrissa and hairs; 3-4 postsutural dorsocentral setae 22 22 (2F) Proboscis shorter than 2.3X head height; western USA 23 22 ' Proboscis longer than 2.25X head height, or from central or eastern North America (S. illinoiensis) 24 [for specimens from western USA within or near overlap in proboscis length, check distribution maps and see “Recognition” under appropriate species] 23 (22) Four postsutural dorsocentral setae; recorded from California and Washington (Fig. 87) S. pacifica n. sp. (Fig. 45), p. 291 23' Three postsutural dorsocentral setae in most specimens; recorded from southwestern USA and Mexico (Fig. 83) S. pisinnia n. sp. (Fig. 49), p. 298 24 (220 Eye large (Fig. 54); proboscis shorter than 2.5X head height; central and eastern North America (Fig. 92) S. illinoiensis (Tnsd.) (Fig. 54), p. 307 24' Eye of average size (Figs. 46-47); proboscis longer than 2.25X head height; transcontinental (Figs. 93, 95) S. cristata (Fabr.) (Fig. 46), p. 292 and S. multifaria n. sp. (Fig. 47), p. 293 [see “Recognition” under S. cristata for discussion concerning separation of these species] Key to adults of Siphona species of Middle America 1 Male 2 V Female 9 [female of S. longissima unknown] 2 (1) Length of first flagellomere less than 0.52 head height 3 2' Length of first flagellomere greater than 0.58 head height, or not south of Tropic of Cancer (5. multifaria) 4 North American species of Siphona Meigen 285 3 (2) Preabdomen light coloured with narrow vitta; 3 postsutural dorsocentral setae in all but very few specimens; palpus yellow in most specimens, slightly infuscate apically in few; distiphallus triangular and pointed in profile (Fig. 73); widely distributed, from central Mexico to Costa Rica (Fig. 96) S.futilis Wulp (Fig. 34), p. 305 3' Preabdomen dark brown, not vittate, with narrow light coloured bands across anterior of T3, T4 and T5; 4 postsutural dorsocentral setae; palpus fuscous or black on apical half; distiphallus slender, apex truncate in profile (Fig. 74); Chiapas (Fig. 97) S. brunnea n. sp. (Fig. 35) p. 308 4 (20 Preabdomen broadly vittate (vitta wider than distance between median marginal setae on T3), dark coloured; palpus infuscate apically; distiphallus short and broad in profile with small hooks along antero-ventral margin (Fig. 76); southern Mexico (Fig. 99) S. akidnomyia n. sp. (Fig. 36), p. 31 1 4' Preabdomen average vittate (vitta not wider than distance between median marginal setae on T3), light coloured; palpus yellow; distiphallus not shaped as above, with or without small hooks along antero-ventral margin 5 5 (4') Proboscis shorter than 2.3X head height; distiphallus slightly bent downward from basiphallus, shaped as in Fig. 65; southwestern USA to Mexico state, Mexico (Fig. 83) S. pisinnia n. sp. (Fig. 23), p. 298 5' Proboscis longer than 2.6X head height, or if shorter ( S . multifaria 2. 25-3. OX) then distiphallus sharply bent downward from basiphallus and not south of Tropic of Cancer 6 6 (5") North of Tropic of Cancer; distiphallus as in Fig. 67; widely distributed in Canada and USA (Fig. 93), presence in northern Mexico likely S. multifaria n. sp. (Fig. 25), p. 293 6' South of Tropic of Cancer 7 7 (60 Length of first flagellomere less than 0.75 eye height; surstylus with apex extended slightly to far beyond apex of cerci (Fig. 79); distiphallus with apex rounded in profile; bend of R] bare; Veracruz (Fig. 98) S. rizaba n. sp. (Fig. 39), p. 310 T Length of first flagellomere greater than 0.75 eye height; surstylus with apex even with or extended slightly beyond apex of cerci (Figs. 77-78); distiphallus with apex pointed in profile; bend of R, with single setula or bare 8 8 (70 Eye height greater than 0.86 head height; bend of R, with single setula on at least one side in most specimens; slope of distiphallus even with that of basiphallus; distiphallus dentate along antero-ventral margin (Fig. 78); Veracruz to Costa Rica (Fig. 101) S. tropica (Tnsd.) (Fig. 38), p. 309 8' Eye height less than 0.85 head height; bend of R, bare; distiphallus bent sharply downward from angle of basiphallus; distiphallus with small hooks along antero-ventral margin (Fig. 77); Chiapas (Fig. 100) S. longissima n. sp. (Fig. 37), p. 31 1 9 (10 Proboscis longer than 2.7X head height 10 9' Proboscis shorter than 2.7X head height 12 [N. multifaria keys through both halves of couplet] 10 (9) North of Tropic of Cancer; widely distributed in Canada and USA (Fig. 93), presence in northern Mexico likely Quaest. Ent., 1982, 18 (1-4) 286 O’Hara S. multifaria n. sp. (in part) (Fig. 47), p. 293 10' South of Tropic of Cancer 11 11 (100 Bend of with a single setula on at least one side in most specimens; Veracruz to Costa Rica (Fig. 101) S. tropica (Tnsd.) (Fig. 57), p. 309 1 V Bend of Rj bare; Veracruz (Fig. 98) S. rizaba n. sp. (Fig. 58), p. 310 [male associates required for positive identification] 12 (90 Proboscis longer than 2.5X head height; wing brownish hyaline; southern Mexico (Fig. 99) S. akidnomyia n. sp. (Fig. 59), p. 31 1 12' Proboscis shorter than 2.5X head height; wing yellowish or brownish hyaline 13 13 (120 Palpus with apical half fuscous or black; 4 postsutural dorsocentral setae; body dark brown; Chiapas (Fig. 97) S'. brunnea n. sp. (Fig. 55), p. 308 13' Palpus yellow; 3 postsutural dorsocentral setae in all but very few specimens, or not south of Tropic of Cancer; body blue-gray with brown areas, not extensively dark brown 14 14 (130 Head profile as shown in Fig. 53; eye relatively large, first flagellomere relatively short; widely distributed, from central Mexico to Costa Rica (Fig. 96) S.futilis Wulp (Fig. 53), p. 305 14' Head profiles as shown in Figs. 47,49; eye relatively small; first flagellomere relatively long 15 15 (140 Proboscis shorter than 2.3X head height; southwestern USA to Mexico state, Mexico (Fig. 83) S. pisinnia n. sp. (Fig. 49), p. 298 15" Proboscis longer than 2.25X head height; widely distributed in Canada and USA (Fig. 93), presence in northern Mexico likely S. multifaria n. sp. (in part) (Fig. 47), p. 293 The S. maculata group This group is characterized by a short to very short proboscis and a slender first flagellomere broadly rounded apically along ventral margin. Weak lateral marginal setae on T1+2 are presumed symplesiotypic for the group, but this state is not expressed in all specimens (and not at all in Siphona lurida). There is a trend toward a smaller than average eye. Except for Siphona pacifica, adults are seen in spring and early summer. Siphona maculata Staeger Siphona maculata Staeger in Zetterstedt, 1849: 3212- Mesnil, 1965: 871 (redescription). Type material examined. - Lectotype (by designation of Andersen, in press “a”), male, labelled: “<$/ Chariot-/ tenlund [Denmark] [handwritten]”; “Coll./ Staeger [yellow label]”; “TYPE [red label]”; “Macula/ ta Staeg. [handwritten]”; “TYPE [red label]”; “Lectotype 6 [on left side of label]/ Siphona maculata/ Staeger in Zett. [handwritten]/ det. Stig. Andersen” (ZMUC). Recognition. - Average-sized eye and first flagellomere, very short proboscis. Most distinctive features are weak lateral marginal setae on T1+2 and small but distinct hooks on antero-ventral margin of distiphallus. Colouration of preabdomen average, vittate. Three postsutural dorsocentral setae. Similar to S. floridensis , from which it can be separated by number of postsutural dorsocentrals, structure of distiphallus and geographical distribution. North American species of Siphona Meigen 287 It is most easily confused with S. intrudens. Males are separated by structure of distiphallus. There are several trends which are useful for distinguishing between S. intrudens and S. maculata, these apply to both male and female specimens, though more conspicuous in former: S. maculata has a shorter proboscis, smaller tarsal claw, larger eye and longer first flagellomere than S. intrudens. With respect to females, preabdomen of most specimens of S. intrudens is black in ground colour overall, while preabdomen of S. maculata in most specimens has a short, broad vitta, with yellow ground colour laterally on T1+2 and antero-laterally on T3. Description. — Specimens examined: 263 males, 117 females from North America; ca. 20 European specimens. Length: 3. 5-5. 0mm. Male. Head (Fig. 18). Colouration: frontal vitta yellow to testaceous; gena and parafacial white to light yellow; frontal plate light yellow to light brown; scape and pedicel yellow to fuscous, first flagellomere in most specimens fuscous or black, testaceous in few light coloured specimens; palpus yellow; proboscis reddish-brown to fuscous. Eye: size average for genus, 0.755-0.825 head height (Fig. 13), slender, widest at center, evenly rounded along anterior margin. Macrotrichia: average for genus. Antenna: first flagellomere 0.564-0.640 head height (Fig. 14); length of aristomere 3 average for genus. Mouthparts: proboscis short, 1 .30-1 .74 head height (Fig. 1 5). Thorax. Colouration: dorsum average, 3 brown vittae distinct in light coloured specimens, dorsum predominantly brown to dark brown in dark coloured specimens; tegula testaceous, light brown to brown; wing yellowish hyaline; legs average. Macrotrichia: postsutural dorsocentrals, 44 specimens with three, 1 with four [males and females], Acropod: claws and pulvilli average-sized. Preabdomen. Colouration: faintly vittate; most specimens black in ground colour on T4 and T5 and medially on Tl+2 and T3, yellow in ground colour laterally on T1+2 and T3; areas black in ground colour with surface blue-gray or gray, dark brown or black around macrotrichial insertions, especially around median marginals on T3 and T4; areas yellow in ground colour with surface yellow to light brown; extent of black ground colour varied among specimens, in few greater than average (posterior portion of T3 black in ground colour) or less than average (anterior portion of T4 yellow in ground colour). Macrotrichia: median marginals absent from T, +2; lateral marginals on T]+2 weak, in most specimens scarcely or not at all differentiated from marginal hairs. Genitalia (Fig. 60). Thirty-eight examined, including 2 from European specimens. Apex of surstylus even with or extended slightly beyond apex of cerci. Aedeagus: distiphallus bent sharply downward from angle of basiphallus; antero-ventral margin V-shaped in anterior view, armed with several well-developed hooks (though smaller than hooks on distiphallus of S. intrudens ); in profile tapered to narrow apex. Female. As described for male except as follows. Head (Fig. 42). Palpus longer and thicker than in males in few specimens. Preabdomen. Colouration: highly varied, most specimens darker than average male; average specimen yellow in ground colour laterally on T|+2 and antero-laterally on T3, rest black in ground colour, dorsal vitta short and broad; range from entirely black in ground colour (vitta absent), to average condition described for male (vitta present); dark specimens with large dark brown or black areas around macrotrichial insertions. Geographical distribution (Fig. 82). - Europe and North America. Transcontinental in North America; most records within a rather narrow belt along southern limit of boreal forest. Habitat. - I have collected specimens of this species in mixed forest dominated by maple (Gatineau Co., Que.) or aspen (Edmonton, Alta.). Individuals flew close to the ground in a zig zag pattern, apparently searching for mates or ground-dwelling hosts. Chorological affinities. - S. maculata and S. intrudens are widely sympatric over much of their apparent ranges. A few examined series of specimens contain both S. intrudens and S. maculata , suggesting their ecological requirements are similar. A closely related species, S. floridensis, is allopatric in distribution. Phylogenetic relationships (Fig. 103). - The phenetic similarity between S. maculata and S. floridensis suggests they might be sister species, though a synapotypy has not been found to substantiate this. Of the North American taxa of the S. maculata group, only S. maculata and S. intrudens share the derived condition of hooks on the antero-ventral margin of the distiphallus (Figs. 60, 62). However, on the basis of other characters, this condition is interpreted as derived at the base of the lineage terminating in S. maculata , S. intrudens and S. lurida (Fig. 103), Quaest. Ent., 1982, 18 (1-4) 288 O’Hara precluding a sister group relationship between S. maculata and S. intrudens. Siphona floridensis new species Holotype. - Male, labelled: “Orlando,/ Fla. [Florida, USA] III.3 . [ 1 9] 46 / F. Snyder” (AMNH). Genitalia in microvial on pin below specimen. Allotype. - Female, same data as holotype (AMNH). Derivation of specific epithet. - With only one record outside Florida, this species is named for its restricted known distribution. Recognition. - Average-sized eye and first flagellomere, very short proboscis. Lateral marginal setae on T1+2 weak or strong. Distiphallus average in structure. Preabdomen vittate or black in ground colour overall. Four postsutural dorsocentral setae. Externally very similar to the allopatric S. maculata. The proboscis of both species is very short. S. floridensis has a slightly larger eye and slightly shorter first flagellomere than S. maculata. Males are separated by lack of hooks on antero-ventral margin of distiphallus (small hooks present in S. maculata) and number of postsutural dorsocentrals (three in S. maculata , four in S. floridensis). Description. - Length: 3. 5-5. 5mm. Male. Head (Fig. 19). Colouration: as described for S. maculata. Eye: size average for genus, 0.783-0.839 head height (Fig. 13), broad, widest at center, evenly rounded along anterior margin. Macrotrichia: average for genus. Antenna: first flagellomere slightly shorter and thinner than in S. maculata , 0.548-0.574 head height (Fig. 14); length of aristomere 3 average for genus. Mouthparts: proboscis short, within range for S. maculata , 1.37-1.59 head height (Fig. 15). Thorax. Colouration: dorsum average, 3 light brown vittae visible in most specimens, scutum and scutellum predominantly brown in few specimens (generally somewhat lighter than in S. maculata ); tegula testaceous; wing yellowish hyaline; legs average. Macrotrichia: postsutural dorsocentrals, all specimens with four [males and females]. Acropod: claws and pulvilli average-sized. Preabdomen. Colouration: specimens in “Orlando” series vittate, as described for average S', maculata ; other males black in ground colour, surface light blue-gray or gray, dark brown to black around macrotrichial insertions. Macrotrichia: median marginals absent from T1 + 2; lateral marginals on T]+2 weak in 2 males and 4 females from “Orlando” series, strong in others. Genitalia (Fig. 61). Four examined. Apex of surstylus extended slightly beyond apex of cerci. Aedeagus: distiphallus bent very slightly downward from angle of basiphallus in 3 specimens, bent more sharply (as in S. maculata) in one specimen; antero-ventral margin dentate (without hooks); apex broad in profile. Female. As described for male except as follows. Head (Fig. 43). Palpus longer and thicker than in males. Preabdomen. Specimen from Lake Placid mottled black and yellow in ground colour, weakly vittate; other specimens black in ground colour, surface light blue-gray or gray. Geographical distribution (Fig. 84). - Known only from Florida and southern Georgia. Chorological affinities. - This is the only Siphona species restricted to extreme southeastern United States, and the only member of the S. maculata group recorded from that area. S. multifaria is the only Siphona species known to be sympatric with this species. Phylogenetic relationships (Fig. 103). - This species is plesiotypic in almost all states, so its placement beyond species group cannot be clarified. It might be the sister species to S. maculata , as inferred from phenetic similarity and zoogeographic analysis. Paratypes. — Four males, 7 females. USA: Florida: Orlando, 3,15.111.46, F. Snyder, 2M, 4F; Wakulla Co., Ochlockonee R. St. Pk„ 29. IV. 70, W.W. Wirth, 1M; Highlands Co., Archbold Biological Station, 14.111.62, S.W. Frost, 1 M; Archbold Biological Sta., L. Placid, 10. IV. 66, R.W. Dawson, IF; Biscayne Bay, IF. Georgia: Liberty Co., St. Catherines Island, 24-28. IV. 72, Thompson and Picchi, 1 F. (Deposited in AMNH, CNC, PSU, USNM and WSUP.) Siphona intrudens (Curran) Bucentes intrudens Curran, 1932: 14. North American species of Siphona Meigen 289 Siphona intrudens : Sabrosky and Arnaud, 1965: 1064. Type material examined. - Holotype, male, labelled: “Bucentes/ intrudens/ Curran <3 [red label]”, “Castle Rock/ IV- 1 7- [ 1 9]08 Pa [Pennsylvania, USA]” “Accession]. 5642 A.J. Weidt coll.”; “Bucentes/ intrudens/ Curran” (AMNH). Allotype , female, same data as holotype except without fourth label (AMNH). Recognition. - Small to average-sized eye, average-sized first flagellomere and short proboscis. Most specimens with weak lateral marginal setae on T1+2, large tarsal claws and a dark blue-gray preabdomen with dorsal vitta. A few specimens very similar to S. maculata , but the U-shaped distiphallus (in anterior view), with 3-4 large hooks along antero-ventral margin, is unique to S. intrudens. Three postsutural dorsocentral setae. S. intrudens can be confused with either S. maculata or S. lurida. For discussion concerning former, see “Recognition” under that species. Specimens of S. lurida with a first flagellomere of average Siphona length are similar in general appearance to S. intrudens. The dark colouration of these species, with blue-gray, non-vittate preabdomens, contributes to their similarity and with other characteristics attests to their close phylogenetic relationship. S. intrudens specimens with strong lateral marginals on T1+2 can be separated from S. lurida by the larger eye, presence of only 2-3 setae on vibrissal angle (besides vibrissa) and by the unique structure of the distiphallus. Description. — Specimens examined: 408 males, 338 females. Length: 3. 5-5. 5mm. Male. Head (Fig. 20). Colouration: as described for S. maculata. Eye: small to average for genus, 0.703-0.817 head height (Fig. 13), slender, widest at center, evenly rounded along anterior margin. Macrotrichia: vibrissal angle with 2 or 3 setae and several hairs, setae in most specimens 2 in number, 1 laterad or above vibrissa and 1 below. Antenna: first flagellomere 0.474-0.593 head height (Fig. 14); length of aristomere 3 average for genus. Mouthparts: proboscis short, 1 .45-2.00 head height (Fig. 1 5). Thorax. Colouration: dorsum average, blue-gray to blue with varied amounts of brown; 3 brown vittae not clearly differentiated in most specimens ( S . intrudens generally with more blue on dorsum than S. maculata , and darker than S. floridensis)', tegula testaceous to fuscous; wing yellowish to cinereous hyaline; legs average to darker than average, femur of hindleg predominantly reddish-brown in few specimens. Macrotrichia: postsutural dorsocentrals, 68 specimens examined, all with three [males and females]. Acropod: claws and pulvilli large in most specimens (Fig. 5), average in few. Preabdomen (Fig. 7). Colouration: highly varied, from black in ground colour, surface predominantly blue-gray or gray, to vittate condition described for S. maculata\ most specimens from western North America dark form, both dark and light (vittate) forms well-represented in examined material from prairies to eastern North America. Macrotrichia: median marginals absent from T]+2; lateral marginals on T,+2 in most specimens weak, slightly stronger than surrounding hairs, in few specimens not differentiated, in few strong. Genitalia (Fig. 62). Thirty-seven examined. Apex of surstylus even with or extended slightly beyond apex of cerci. Aedeagus: distiphallus bent sharply downward from angle of basiphallus; antero-ventral margin U-shaped in anterior view, with several large, well developed hooks (larger than in S. maculata)\ in profile tapered to narrow apex. Female. As described for male except as follows. Head (Fig. 44). Palpus longer and thicker than in males in most specimens. Preabdomen. Colouration: most specimens black in ground colour, surface predominantly blue-gray or gray; few specimens vittate, as described for average specimen of female S', maculata. Geographical distribution (Fig. 85). - Transcontinental species; records mostly south of boreal forest, and particularly well recorded from northeastern United States. Chorological affinities. - Among S. maculata group members, this species is evidently parapatric or narrowly sympatric with S. lurida and S. pacifica. Also see notes under S. maculata. Phylogenetic relationships (Fig. 103). - This is a highly derived species, closely related to S. lurida. A decision on whether these species are sister species awaits a cladistic analysis of related (undescribed) Oriental species of Siphona. See also notes under S. maculata. Quaest. Ent., 1982, 18 (1-4) 290 O’Hara Siphona lurida Reinhard Siphona lurida Reinhard, 1943: 20-- Sabrosky and Arnaud, 1965: 1064. Type material examined. - Holotype , male, labelled [lines not recorded]: “Rainier, Ore. [Oregon, USA] III.3. [ 1 9] 30”; “R.E. Dimick Coll.”; “HOLOTYPE Siphona lurida Reinhard [red label]”; “Siphona lurida Rnh R’43 [Reinhard det. label]” (CNC). Allotype , female [abdomen missing], labelled: “Union Hills/ 11-20-1940/ R.E. Rieder, col.”; “ALLOTYPE/ Siphona/ lurida/ Reinhard [red label]”; “Siphona/ lurida/ Rnh. R. [Reinhard det. label]”; “PARATYPE/ Siphona/ lurida/ Reinhard [yellow label]” (FSCA). Paratypes. — , 4 males, top two labels: “Union Hills/ Marion Co[unty], Ore. [Oregon, USA]/ 11-20-1940/ R.E. Rieder, col.”; “Paratype/ Siphona/ lurida/ Reinhard [red label]” (2 in CNC, 1 each in FSCA, OSU). Recognition. - Eye smaller than any other species, first flagellomere long in most specimens. Proboscis short. Cluster of 5-10 setae on vibrissal angle besides vibrissa and hairs, and above average number of hairs on parafacial and frontal plate (up to ca. 15). Dark coloured species, preabdomen black in ground colour overall, apex of palpus infuscate in most specimens. Tarsal claws large in most specimens. Wing cinereous hyaline. Lateral marginal setae on T1+2 strong. Three postsutural dorsocentral setae. Adults are seen in very early spring to mid-summer; one southern record from December. A few specimens are very similar to S. intrudens, as discussed under “Recognition” of that species. S. lurida specimens with a very small eye and large first flagellomere are easily identified. The holotype and paratypes are of this form, as are examined specimens of S. lurida from Japan. It is uncertain how prevalent specimens with a short first flagellomere are, for the species has been inadequately collected. Description. — Specimens examined: 1 1 males, 9 females from North America; 1 male, 2 females from Japan. Length: 4. 5-5. 5mm. Male. Head (Fig. 21). Colouration: frontal vitta bright orange to reddish-brown; gena and lower parafacial orange or yellow to gray, upper parafacial and frontal plate yellow or blue-gray to brown; orbital plate light to dark brown; scape, pedicel and a portion of first flagellomere above aristal insertion orange to fuscous, rest of first flagellomere fuscous to black; palpus yellow, apex infuscate in most specimens; proboscis reddish-brown to fuscous. Eye: very small, 0.627-0.677 head height (Fig. 13), slender, widest at center, evenly rounded along anterior margin. Macrotrichia: number and length of hairs on upper parafacial and fronto-orbital plate above average for genus in most specimens, up to 15 present; vibrissal angle with tight cluster of 5-10 setae and several hairs in addition to vibrissa. Antenna: first flagellomere highly varied in length, 0.508-0.677 head height (Fig. 14); length of aristomere 3 average for genus. Mouthparts: proboscis short, 1.45-2.00 head height (Fig. 15). Thorax. Colouration: dorsum average, blue-gray or blue with 3 clearly differentiated brown vittae in most specimens, few specimens with dorsum predominantly brown (very close to S. intrudens)-, tegula fuscous; wing cinereous hyaline; legs average to darker than average, femur of foreleg predominantly reddish-brown in few specimens. Macrotrichia: postsutural dorsocentrals, 20 specimens examined, all with three [males and females]. Acropod: claws and pulvilli large in most specimens, average in few. Preabdomen. Colouration: not vittate; black in ground colour, surface varied, from predominantly blue-gray to predominantly dark brown mixed with black with blue-gray restricted to bands on anterior third of T3, T4, and T5; dark brown or black around macrotrichial insertions. Macrotrichia: median marginals absent from T1+2; lateral marginals on T1 + 2 strong. Genitalia (Fig. 63). Six examined, including one Japanese specimen. Apex of surstylus even with or extended slightly beyond apex of cerci. Aedeagus: distiphallus bent sharply downward from angle of basiphallus; antero-ventral margin V-shaped in anterior view, dentate; in profile tapered to rounded apex. Female. As described for male except as follows. Head (Fig. 41 ). Palpus longer and thicker than in males in most specimens. Preabdomen. Colouration: black in ground colour; surface varied, from predominantly blue-gray or gray to predominantly dark brown with or without narrow bands of blue-gray on anterior margins of T3, T4, and Ts. Geographical distribution (Fig. 86). - Japan (new record) and western United States. North American species of Siphona Meigen 291 Chorological affinities. - In North America, this species is apparently parapatric or narrowly sympatric with S. maculata and the closely related S. intrudens. The range of S. pacifica coincides with that of another S. maculata group member, S. lurida. Phylogenetic relationships (Fig. 103). - Like S. intrudens , this species is highly derived. It might be sister species to S. intrudens , as mentioned under that species. Siphona pacifica new species Holotype. - Male, labelled: “Dartford, WASH. [Washington, USA]/ Malaise trap”; “R.D. Gray/ VI. 10-11 1969” (WSUP). Allotype. - Female, labelled: “Dartford, Wash/ VII-5- 1 970”; “Malaise trap/ R.D. Gray” (WSUP). Derivation of specific epithet. - This species is named in recognition of its western distribution. Recognition. - Proboscis length and eye height average, first flagellomere short. Preabdomen light blue-gray or gray, with or without dorsal vitta. Lateral marginal setae on T , _|_ 2 weak or strong. Distiphallus bent slightly downward from angle of basiphallus, apex broad in profile. Four postsutural dorsocentral setae. This species has a unique set of characteristics that distinguish it from other species. However, taken singly these features are unremarkable, and S. pacifica is therefore difficult to recognize. It is placed in the S. maculata group because the first flagellomere is slender and broadly rounded apically along ventral margin, lateral marginal setae on T)+2 are weak in some specimens, and general colouration is typical of the group. Description. - Length: 3.5-5.0mm. Male. Head (Fig. 22). Colouration: frontal vitta orange, testaceous to reddish-brown; gena and parafacial white, light yellow to light brown; fronto-orbital plate light brown to brown; scape and pedicel and in few specimens portion of first flagellomere above aristal insertion yellow; first flagellomere fuscous to black; palpus yellow; proboscis reddish-brown. Eye: size average to large for genus, 0.796-0.850 head height (Fig. 13), broad, widest at center, evenly rounded along anterior margin. Macrotrichia: vibrissal angle with 1 seta above vibrissa (or laterad) and in most specimens 2 setae below, and several short hairs. Antenna: first flagellomere short, 0.431-0.472 head height (Fig. 14); length of aristomere 3 average for genus. Mouthparts: proboscis length average for genus, 1.87-2.15 head height ( Fig. 1 5). Thorax. Colouration: dorsum average, blue-gray or gray finely mixed with brown in most specimens, 3 brown vittae visible in most specimens; tegula testaceous; wing yellowish hyaline; legs average. Macrotrichia: postsutural dorsocentrals, four in all specimens. Acropod: claws and pulvilli average-sized. Preabdomen. Colouration: varied, vittate in few specimens; most specimens predominantly or entirely black in ground colour, few specimens with yellow ground colour laterally on T)+2 and T3 or less; surface light coloured, areas of black ground colour light blue-gray or gray in surface colour, brown to black around macrotrichia! insertions; areas yellow in ground colour with surface yellow or light brown. Macrotrichia: median marginals absent from T]+2; lateral marginals on T|+2 differentiated from marginal hairs, weak or strong. Genitalia (Fig. 64). Two examined. Apex of surstylus even with or extended slightly beyond apex of cerci. Aedeagus: distiphallus bent slightly downward from angle of basiphallus; antero-ventral margin dentate; in profile anterior edge straight, apex broad. Female. As described for male except as follows. Head (Fig. 45). Length of palpus slightly longer than in males. Preabdomen. Colouration: not vittate; black in ground colour, surface blue-gray or gray. Geographical distribution (Fig. 87). - Recorded from several localities in California and one in Washington. Chorological affinities. - The western distribution of this species is very similar to that of S. lurida. Two other species of the S. maculata group, S. maculata and S', intrudens , are evidently parapatric or narrowly sympatric with this species. Phylogenetic relationships (Fig. 103). - S. pacifica is interpreted as distantly related to most other North American members of the S. maculata group. This species possesses the Quaest. Ent., 1982, 18 (1-4) 292 O’Hara synapotypic feature of the group - a slender, pointed first flagellomere - as well as the symplesiotypies, weak lateral marginal setae on T1+2 and a relatively short proboscis. Paratypes. — Seven males, 4 females. USA: California: Sugar Loaf Mt., 28.V.46, J. Sperry, 1M; Marin Co., Mill Valley, IX. 51, E.S. Ross, 1M; Placer Co., Brockway, VII. 41, G.E. Bohart, 1M. Washington: Spokane Co., Dartford, 6-9, 20-31. VI, 2-3. VII, 2-3.VIII.69, 5.VII.70, R.D. Gray, 4M, 3F; ibid., 23.VI.70, IF. (Deposited in CAS, USNM and WSUP.) The S. cristata group Characteristics varied; most species with an average to long proboscis, average-sized eye, broad first flagellomere rounded or slightly truncate apically along ventral margin (not distinctly pointed), and an anterior indentation on the distiphallus. The three predominantly to entirely yellow North American Siphona species belong to this group. Siphona cristata (Fabricius) Stomoxys cristata Fabricius, 1805: 281. Siphona cristata: Mesnil, 1964: 861 (redescription).— Curran, 1932: 13 (in key; probable misidentification). Siphona chaetolyga Rondani, 1865: 194.- Herting, 1969: 192. Siphona palpina Zetterstedt, 1859: 6064.- Mesnil, 1964: 861. Mesnil (1964) and Crosskey (1976b) followed earlier authors and listed Siphona oculata Pandelle as a synonym of S. cristata. Herting recently examined the type of S. oculata and found it conspecific with Siphona pauciseta Rondani (Herting, 1978: 6). Type material examined. - Stomoxys cristata Fabricius, HOLOTYPE, female, labelled: “TYPE [red label]”; “S. cristata/ [locality indecipherable; Denmark] [handwritten]”; “Holotype 2/ Siphona/ cristata (F.) [handwritten]”/ det. “Stig Andersen” (ZMUC). I cannot with certainty confirm that the type of S. cristata is conspecific with the North American species I identify as such. The type is a female with features characteristic of (my) S. cristata and S. multifaria. Therefore my interpretation of S. cristata is based primarily on Mesnil’s description (1964) and male and female specimens determined by Drs. Mesnil and Herting. Recognition. - Eye height and first flagellomere length average, proboscis longer than average. Colouration highly varied; in few specimens thoracic pleura yellow in ground colour, dorsum dark in ground colour, rest of specimens with thorax predominantly dark in ground colour. Preabdomen of both sexes varied, from entirely yellow, to yellow with narrow median vitta, to average vittate with predominantly tawny surface colour. Very few specimens with a single or 1 pair median marginal setae on T1+2 (3% each). Distiphallus bent sharply downward from angle of basiphallus, in most specimens with anterior edge curved inward ca. 0.33 of its length from ventral edge. Postsutural dorsocentral setae three (35%) or four (65%). Due to varied nature of this species, identification can be difficult. A few specimens resemble S. lutea (as discussed under “Recognition” of that species), but do not pose a serious identification problem. Separation of S. cristata and S. pisinnia is discussed under the latter species. Specimens of S. cristata can be very difficult to separate from those of S. multifaria , for they are similar in most characteristics. Colouration is useful, but not always trustworthy. The yellow form of S. cristata is very distinct from S. multifaria , but specimens with vittate preabdomen are not always readily separable from S. multifaria specimens. In Edmonton, Alberta, both species occur at the same time in the same place. S. cristata specimens are slightly larger and have a tawny, vittate preabdomen. The preabdomen of S. North American species of Siphona Meigen 293 multifaria specimens is yellow laterally (not tawny as in S. cristata ) and has a narrow brown pruinose vitta medially, overlying a indistinct light blue-gray vitta. Differences in the distiphallus are recognizable, but do not hold for all specimens. Since both S. cristata and S. multifaria are quite varied, the differences seen in Edmonton populations are not suitable for species discrimination in all areas. For this reason I have not separated these species in the key. Most male specimens should be identifiable on the basis of the foregoing remarks, though female specimens will not meet with equal success. Description. — Specimens examined: 78 males, 112 females from North America, ca. 10 European specimens. Length: 4. 5-5. 0mm. Male. Head (Fig. 24). Colouration: as described for S', lutea except for antenna and proboscis; scape, pedicel and portion of first flagellomere above aristal insertion yellow in most specimens, fuscous in few; rest of first flagellomere testaceous to fuscous, in few specimens black; proboscis brown to reddish-brown. Eye: size average to above average for genus, highly varied, 0.754-0.855 head height (Fig. 13), slender to broad, widest at, or slightly above, center, narrowed or rounded below. Macrotrichia: average for genus. Antenna: first flagellomere 0.547-0.672 head height (Fig. 14), resembling first flagellomere of S. lutea but more varied in length and shape; first flagellomere broad, varied from truncate to broadly rounded along ventral margin; length of aristomere 3 average for genus. Mouthparts: proboscis length above average for genus, highly varied, 2.25-3.00 head height (Fig. 1 5). Thorax. Colouration: black in ground colour in most specimens, light brown in ground colour on pleura in few specimens; dorsum from yellow or light brown finely and evenly mixed with blue-gray or gray to entirely yellow, golden or brown; 3 brown or golden vittae dorsally in very few specimens; tegula testaceous; wing yellowish hyaline; legs average. Macrotrichia: postsutural dorsocentrals, 20 specimens with three, 37 with four [males and females]. Acropod: claws and pulvilli average in size. Preabdomen. Colouration: one form predominantly yellow in ground colour, with or without narrow dark vitta, surface yellow to bright tawny; pattern varied from yellow form to average Siphona vittate form, with T5, most of T4, triangular area on T3, and T,+2 medially black in ground colour, surface gray to brown; dark brown to black around macrotichial insertions; rest of preabdomen yellow in ground colour, surface yellow to tawny. Macrotrichia: most specimens without median marginals on T1+2, few with 1 seta on one side (6 specimens) or 1 pair (5 specimens); lateral marginals on T1+2 strong. Genitalia (Fig. 66). Thirty-two examined, including 2 from European specimens. Apex of surstylus even with or extended slightly beyond apex of cerci. Aedeagus: distiphallus bent sharply downward from angle of basiphallus; antero-ventral margin dentate; in profile anterior edge varied, in most specimens indented ca. 0.33 distiphallus length from ventral edge, straight in few specimens; apex broad in profile. Female. As described for male except as follows. Head (Fig. 46): Palpus longer and thicker than in males in most specimens. Geographical distribution (Fig. 95). - Wide ranging in the Palearctic and Nearctic Regions, though sparsely recorded from latter over much of its apparent range. Habitat. - I have collected specimens of S. cristata , S. lutea and S. multifaria on flowers of the composite, Erigeron philadelphicus L., in river valley of the North Saskatchewan R., Edmonton, along shaded bank of a small stream in vicinity of poplar and scattered spruce. Chorological affinities. - In North America this wide ranging species is sympatric with the closely related species S. multifaria and S. lutea , in addition to most other Siphona species of America north of Mexico. As noted above, specimens of S. multifaria and S. lutea have been caught in the same locality as those of S. cristata. In southwestern New Mexico, specimens of S. pisinnia were collected with specimens of this species. Phylogenetic relationships (Fig. 104). - Close to S. multifaria among North American species, though the sister species might be a Palearctic species. Siphona multifaria new species Holotype. - Male, labelled: “MI. 87 [also known as km. 140.5], Y.T. [Yukon Territory, Canada]/ Dempster Hwy ./ 8-12.VII.1973/ G.&D.M. Wood” (CNC). Allotype. - Female, same data as holotype except date 16-17.VII.1973 (CNC). Quaest. Ent., 1982, 18 (1-4) 294 O’Hara Derivation of specific epithet. - From the Latin multifarius, meaning “having great variety”. S. multifaria is so named because it is a widely distributed species of varied appearance, being particularly difficult to identify because of its resemblance to S. cristata. Recognition. - Eye height and length of first flagellomere average, proboscis longer than average. Wing yellowish hyaline in most specimens, cinereous in few. Male preabdomen varied, in most specimens light coloured, vittate, with a very narrow brown pruinose vitta (unassociated with vitta produced by ground colour), few specimens predominantly black in ground colour. Female preabdomen darker than in male, not vittate. Very few specimens with 1 median marginal seta on one side on T1+2 (2%). Distiphallus bent sharply downward from basiphallus, anterior edge slightly or not at all curved inward ca. 0.33 of its length from ventral edge. Most specimens (65%) with three postsutural dorsocentral setae. Most difficult to separate from S. cristata , as discussed under “Recognition” of that species. S. multifaria and S. pisinnia are narrowly sympatric in southwestern United States. Specimens from that region should be checked carefully against data in the Hubbs-Hubbs diagrams, for colouration is essentially the same in both species, and differences between distiphalli are not diagnostic. The data base for S. multifaria and S. pisinnia is large enough that measurements cited in key, especially with reference to male specimens, should be representative of the two species. Description. - Length: 3. 5-5. 0mm. Male. Head (Fig. 25). Colouration: frontal vitta orange or yellow to testaceous; gena and lower parafacial white to light yellow; upper parafacial and fronto-orbital plate light yellow to brown, darkest on orbital plate, in few specimens fronto-orbital plate bright yellow (these specimens resembling S. cristata ); scape, pedicel, and in few specimens portion of first flagellomere above aristal insertion yellow to testaceous, in few specimens reddish-brown or fuscous; first flagellomere reddish-brown, fuscous or black; palpus yellow; proboscis light brown to reddish-brown or fuscous. Eye: size average to above average for genus, highly varied, 0.755-0.854 head height (Fig. 13), slender, widest at center, evenly rounded along anterior margin. Macrotrichia: vibrissal angle with 2 or 3 setae and several hairs, setae in most specimens 2 in number (average for genus), 1 laterad or above vibrissa and 1 below. Antenna: first flagellomere highly varied in length, 0.531-0.675 head height (Fig. 14); shape of first flagellomere varied, as described for S. cristata ; length of aristomere 3 average for genus. Mouthparts: proboscis length above average for genus, highly varied, 2.31-3.02 head height (Fig. 15). Thorax. Colouration: dorsum average, blue-gray or gray finely mixed with brown in most specimens, 3 brown vittae visible in few specimens; tegula testaceous; wing yellowish hyaline in most specimens, cinereous hyaline in few; legs average. Macrotrichia: postsutural dorsocentrals, 68 specimens with three, 37 with four [males and females]. Acropod: claws and pulvilli average-sized. Preabdomen. Colouration: faintly or clearly vittate; most specimens black in ground colour on T5, most or all of T4, and medially on T]+2 and T3, surface blue-gray, gray or light brownish-gray; dark brown to black around macrotrichial insertions; surface of most specimens with a faint or distinct narrow median brown pruinose vitta, narrower than vitta formed by black ground colour; areas yellow in ground colour with surface yellow or light brown; in few specimens black ground colour more extensive, covering most of T3, all of T4 and T5, with wide vitta T1+2. Macrotrichia: most specimens without median marginals on T1+2, 7 with 1 on one side (1 specimen from Labrador, 6 from western North America); lateral marginals on T, +2 strong. Genitalia (Fig. 67). Fifty-six examined. As described for S. cristata , except anterior edge of distiphallus in profile straight or slightly indented in most specimens, very few specimens with indentation as strong as in average S. cristata. Female. As described for male except as follows. Head (Fig. 47). Palpus longer and thicker than in males in most specimens. Preabdomen. Colouration: darker than male; most specimens not vittate, black in ground colour except laterally on T, +2; surface blue-gray, mixed with brown, in few specimens very light, in few very dark. Geographical distribution (Fig. 93). - Very widely distributed throughout eastern and western North America; records absent for central plains. Habitat. - See S. cristata. Chorological affinities. - Due to the ubiquitous nature of this species, it is sympatric with all other Siphona species of America north of Mexico. Specimens of S. cristata and S. lutea have been caught together with those of this species. North American species of Siphona Meigen 295 Phylogenetic relationships (Fig. 104). - Because this species is wide ranging in North America and recorded from northern localities, I consider its presence in the Palearctic Region likely. Therefore its sister species is not necessarily a species of Nearctic distribution. S. cristata is evidently the closest Siphona species to S. multifaria in North America. Paratypes. — One hundred and eigthy-nine males, 128 females. CANADA. Alberta: George L., 53°57'N 114°06'W, 13-17.V1II.66, P. Graham, 1 M; Edmonton, 19.V.47, E.H. Strickland, IF; ibid., 1932, O. Bryant, 1M; ibid., 29.V1.80, J.F. Landry, 2M; ibid., 7,8.V11.80, 30.VI, 3,6.VII.81, J.E. O'Hara, 13M; Mildred L„ 57°03'N 111°35'W, 11-15.V1I.79, J. Ryan and G. Hilchie, 1M; Banff, 3.VI1.49, E.H. Strickland, IF; Banff N.P., 8. VII. 55, R. Coyles, 1 M; Frank, 15.VI.62, W.R.M. Mason, 1 M; Medicine Hat, 22. VI. 74, J.F. McAlpine, 1 M; Cypress Hills, 10.VIII.49, E.H. Strickland, 3F. British Columbia: Summit L., mi. 392, Alaska Hwy., 4500', 29-30. VI, 2-4,1 1-14. VII. 59, R.E. Leech, 2M, IF; ibid., 4200', 21, 31. VII, 19-21 VIII.59, R.E. Leech, 6M, IF; ibid., 19-21. VIII. 59, E.E. MacDougall, 2F; Lakelse L. bog, s. ofTerrace, 1 1,27.VI.60, B. Heming, 2M; ibid., 11.VI1.60, C.H. Mann, 1M; Lac La Hache, 15.VI1.73, H.J.Teskey, 1M; Keremeos, 18.VI.23, C.B. Garrett, 1M; Atlin, 2200', 3.VII.55, H.J. Huckel, 1M; Horsefly, 14.VII.73, H.J. Teskey, 1M; Ketchum L„ 58°22'N 131°45'W, 3600', 23.VIII.60, W.W. Moss, IF; ibid., R. Pilfrey, 2F; McQueen L„ 10 mi. n. Kamloops, 2. VII. 73, H.J. Teskey, IF. New Brunswick: Kouchibouquac N.P., 10, 24. VI. 77, J.R. Vockeroth, 3M, 6F; ibid., 7. VII. 77, J.F. McAlpine, 1M; St. Andrews, 3. VIII. 57, G.E. Shewed, 1M. Newfoundland: St. John's, 21.VII.67, J.F. McAlpine, IF; Cartwright, Labrador, 19.VII.55, E.E. Sterns, IF. Northwest Territories: Norman Wells, 8, 16. VIII. 69, G.E. Shewed, 2F. Ontario: Maynooth, 9. VIII. 74, D.M. Wood, 1M; Mer Bleu, 5 mi. e. Ottawa, 3,11. VI. 66, D.D. Munroe, 2M; Mink L., nr. Maynooth, 27. VII. 55, J.F. McAlpine, 1M; Forestry Station, Petawawa, 28.V.59, J.R. Vockeroth, 1M; Petawawa, 28.V.59, J.R. Vockeroth, 3F; Go Home Bay, 8 mi. w. Bala, 22.V.59, J.G. Chillcott, 1M; Foxboro, 10.V.63, A.F. Johnson, 1M. Quebec: Gatineau Co., Masham Twp., 25-31. VII. 74, D.M. Wood, 2M; Lac Larouche, La Verendrye Pk„ 30.VII.72, D.M. Wood, 2M, 2F; Mistassini Post, 1 8. VII. 56, J.R. Lonsway, 1M;4 mi. n. Eardley, 20,25. VI 1 1.7 1 , D.M. Wood, 1M, 3F; Kam. Co., Parke Reserve, 28.VII.57, G.E. Shewed, 1M; Duncan L., nr. Rupert, 28.VI1, 9.VIII.70, 31. VII, 14. VIII. 71, J.F. McAlpine, 3M, 2F. Yukon Territory: 14 mi. e. Dawson, 1300', 4. VIII. 62, P.J. Skitsko, IF; 17 km. wnw. Burwash Flats, 1250m, 10. VI 1.80, Wood and Lafontaine, IF; mi. 40, Dempster Hwy., 1-6. VI 1.73, G. and D.M. Wood, 1M; mi. 87, Dempster Hwy., 27-30.VI, 1-12, 18-27. VII, 4-8.V1II.73, G. and D.M. Wood, 37M, 23F; mi. 51, Dempster Hwy., 18-27.VII.73, G. and D.M. Wood, 1 M; km. 140.5, Dempster Hwy., 900m, 27-29.VI1, 1.VIII.80, Wood and Lafontaine, 1 M, 7F; km. 1 55, Dempster Hwy., 950m, 29.V1-3.VII.80, Wood and Lafontaine, 2M, 3F. USA: Alabama: Eufaula, 18. VI. 54, R.L. Fischer, IF. Alaska: Richard Hwy., Donnelly Dome, 25. VI. 51, W.R.M. Mason, 1 M; Cape Thompson, 1 .VI 1 1.61 , R. Madge, 1 F; Unalakleet, 1 7. VI 1, 1 1 .VI 11.61 , B.S. Heming, 2F; Kodiak, IX. 17, J.S. Hine, IF; Douglas, 4. VIII. 01, E. Jenne, IF; Katmai, VIII. 17, J.S. Hine, 1M; King Salmon, Naknek R., 10. VII, 10.VIII.52, W.R. Mason, 1 M, IF; ibid., 2-3. VIII. 52, J.B. Hartley, 3F; Naknek, 18. VII, 8. VIII. 52, W.R. Mason, 3F. Arizona: Sunnyside Canyon, Huachuca Mts., 9. VI 1.40, D.E. Hardy, 2F; Cochise Co., Southwestern Research Station, 8 km. w. Portal, 1650m, 21.VII1.62, 2M; ibid., 27.X.64, IF; ibid., 3.XI.64, V.D. Roth, IF; ibid., 14.VIII.65, 2M; ibid., 9. VII, 27.VIII.65, V.D. Roth, 2M; ibid., 5-25.IX.65, C.W. Sabrosky, 3M, IF; ibid., 23.IX.66, P.H. Arnaud, Jr., 1M; ibid., 26. IX. 66, V.D. Roth, IF. California: Mono Co., Leavitt Meadow, 7200', 12. VIII. 63, H.B. Leech, IF; Trinity Co., Butter Creek, 3450', ca. 12 mi. se. Hyampom, 21, 22. VII. 68, H. Leech, 4M. Colorado: Mt. Evans, Doolittle Ranch, 9800', 22,23. VII, 3. VIII. 61, J.G. Chillcott, 3M; ibid., 3.VIII.61, W.R.M. Mason, 4M, IF; ibid., C.H. Mann, 1M; ibid., B.H. Poole, 1M, IF; Mt. Evans, Echo L., 10,600', 12. VIII. 61, C.H. Mann, IF; Boulder, 1922, 1M; Pingree Pk., 17. VIII. 32, IF; Mesa Co., Glade Pk., 8. VI 1.53, Gurney, IF. Florida: Inverness, Robertson, 1M; S. Florida, Robertson, 2M, 2F; Lake Co., Paisley, 18.11.66, C.L. Felshaw, 1M. Georgia: Dallas, 2. VI. 40, P.W. Fattig, IF; Lumpkin Co., 15mi. nw. Dahlonega, 25. VI. 69, F. Santana, 1M; Oconee Co., 4mi. s. Farmington, 20.111.75, B.F. Freeman, 1M. Idaho: Newman L., 16.V.25, A.L. Melander, IF; Viola, 26. VI. 12, J.M. Aldrich, IF. Massachusetts: Chester, 3. VIII, C.W. Johnson, 1M; Essex, 20. VII. 11, IF; Cuttyhunk Is. and Elisabeth Is., 21. VIII. 71, C.T. Parsons, IF. Michigan: Thayers, 16.V.55, J.R. Vockeroth, IF; Montcalm Co., Flat River Game Area, 14.V.55, R. L. Fischer, 1M; Emmet Co., 27.V.60, R. and K. Dreisbach, 6M; Charlevoix Co., 31.V.60, R. and K. Dreisbach, 3M, IF; Iron Co., 26. V 1 1 1.59, R. and K. Dreisbach, 1M; Hillsdale Co., 21 .V.60, R. and K. Dreisbach, 1 M, 2F; Saginaw Co., 24.V.60, R.R. Dreisbach, 1 M. Montana: Granite Co., Elkhorn Ranch, 9 mi. s. Clinton, 3800', 26.VI.71, J.R. Powers, 1M. New Hampshire: Flume, n. Woodstock, 19. VI 1.35, Blanton and Borders, IF; Franconia, 1M; Halfway House, to Gorham, 20. VII. 35, Blanton and Borders, 1M; Gorham, 19. VIII. 56, W.A. Drew, IF; ibid., R.W. Hodges, 1M; ibid., 14.VI1I.58, J.R. Vockeroth, 1M. New Jersey: Avalon, 8.VI, 2M; ibid., 30.VI, IF. New York: Lake Placid, 2000', 19.VII.62, J.R. Vockeroth, 3M, IF; ibid., J.G. Chillcott. 3M, IF; Keene Valley, 10.V111.16, E.L. Diven, 1M, IF; Dug Mt., 8.VII1.12, D.B. Young, 1M; Olean, 5. IX. 13, IF; Babylon, L.I., 9. VI. 35, 28.VIII.36, Blanton and Borders, 8M, 12F; ibid., 5.VIII.37, F.S. Blanton, IF. North Carolina: Highlands, 3-5000', IV. 36, R.C. Shannon, 1M; ibid., 3800', 29.V.57, J.R. Vockeroth, IF; Cumberland Co., Fort Bragg, 16.V, 27.IX-3.X.67, J.D. Birchim, 1 M, IF. Oregon: Deschutes Co., Meadow Cpgd., Bend, 3840', 29.VII.70, P.H. Arnaud, Jr., 1M. Pennyslvania: West Co., Pitcairn, 17.V.65, J.G. Chillcott, 1M. Tennessee: Gatlinburg, 22. VI. 40, R.C. Osburn, 1M. Utah: Grand Co., Warner Ranger Station, 28 mi. ese. Moab, 9200', 2. V 1 1 1 .60, F., P. and B. Rindge, 1M. Vermont: Chittenden, Rutland, 1-15. VIII. 16, J. Bequaert, 3M; Norwich, 7. VII, C.W. Johnson, IF. Washington: Glacier Pk., Avalanche L., 14. VII. 35, A.L. Melander, 1 M; Asotin Co., Fields' Spr. St. Pk., 15. VI. 72, W.J. Turner, 1M; Yakima Co., 8 mi. sw. Tieton R.S., Bear Creek, 16, 28. VII. 72, W.J. Turner and W.B. Garnett, 7M. Wisconsin: Polk Co., VII, Baker, IF. Quaest. Ent., 1982, 18 (1-4) 296 O’Hara Wyoming: Sweetwater Co., (Old) Hwy. 187, 11.5 mi. s. Eden, 23. VIII. 65, H.B. Leech, 1M. (Deposited in AMNH, CAS, CNC, CUI, FSCA, INHS, JEOH, MCZ, MSU, OUCO, UASM, UGA, UGG, UKL, USNM, and WSUP.) Siphona lutea (Townsend) Crocuta lutea Townsend, 1919: 584. Siphona lutea : Sabrosky and Arnaud, 1965: 1064. Siphona tenuis Curran, 1933: 10.-- Sabrosky and Arnaud, 1965: 1064. New synonymy. Notes about synonymy. - Curran did not include S. lutea (Townsend) in his 1933 key to adults of North American Siphona species. This suggests he was unaware of Townsend’s description of this species when he described S', tenuis in 1933. I have examined the types of both nominal species and regard them conspecific. Type material examined. - Crocuta lutea Townsend, 2 syntypes. One female specimen here designated LECTOTYPE, labelled: “TD 4519 [handprinted]"; “Franconia/ NH [New Hampshire, USA]’’; “CHT Townsend/ coll"; “21. VII, [19] 1 5“[handprinted]; “Type/ No. U.S.N.M. [red label; no number]’’; “Crocuta/ lutea/ T. [handprinted]/ Det CHTT" (USNM). My lectotype label “LECTOTYPE/ Crocuta/ lutea Tnsd./ O’Hara designation/ Selected 1982 [red bordered, handprinted label]” has been attached to this specimen. One paralectotype, female, same locality data as lectotype except collected 20.VII.1915 (USNM). Siphona tenuis Curran, HOLOTYPE, male [left wing, 4 legs missing], labelled: “Siphona/ tenuis/ Curran 6 [red type label]’’; “Timagami, Ont. [Ontario, Canada]/ 1. VIII. 1932/ A.W.A. Brown“; “Siphona/ tenuis/ Curran” (AMNH). Recognition. - Particularly distinct because of yellow ground and surface colour overall. Most specimens examined (78%) with R4+5 setulose beyond crossvein r-m on at least one side. Similarly, Rj distally with 1-2 setulae on at least one side in half the specimens (53%). Most specimens (94%) with four postsutural dorsocentral setae. Shares with S. oligomyia the synapotypic character state of yellow ground and surface colouration. Differs from S. oligomyia in possessing a shorter proboscis and four postsutural dorsocentrals and lacking a modified male sternum 5. The only other species resembling S. lutea is S. cristata. The preabdomen of most specimens of S. cristata is yellow with a thin dark median vitta; in few specimens it is darker, or entirely yellow as in S. lutea. The darker thorax of S. cristata serves to separate specimens of that species from S. lutea where abdominal colouration is ambiguous. Also, most specimens of S. lutea are setulose on R4+5 beyond crossvein r-m, unlike other North American species. Description. — Specimens examined: 41 males, 39 females. Length: 4. 0-5. 5mm. Male. Head (Fig. 26). Colouration: frontal vitta orange or yellow to testaceous; gena and parafacial white to yellow; fronto-orbital plate yellow, bright yellow to gold; scape, pedicel, palpus and proboscis yellow; first flagellomere yellow to testaceous. Eye: size average for genus, 0.760-0.821 head height (Fig. 13), slender, widest at center, in most specimens evenly rounded along anterior margin, in few slightly narrowed below center. Macrotrichia: average for genus. Antenna: first flagellomere 0.561-0.643 head height (Fig. 14), broad and slightly truncate; length of aristomere 3 average for genus. Mouthparts: proboscis slightly shorter than average for genus, 1.83-2.14 head height (Fig. 15). Thorax. Colouration: yellow in ground colour and pruinosity; tegula yellow; wing yellowish hyaline; femora and tibiae yellow. Macrotrichia: postsutural dorsocentrals, 2 specimens with three, 31 specimens with four [males and females]; wing, 32 males and females examined, R4+5 setulose beyond crossvein r-m on at least one side in 25 specimens, R, with at least 1 setula on apical third on at least one side in 17 specimens, CuA, with 1 to 3 setulae in 4 specimens. Acropod: claws and pulvilli average-sized (Fig. 6). Preabdomen. Colouration: yellow in ground and surface colour; few specimens with red or dark areas posteriorly, where dark coloured internal structures are visible through translucent exoskeleton. Macrotrichia: median marginals absent from T, +2; lateral marginals on T, +2 strong. Genitalia (Fig. 68). Nine examined. Apex of surstylus extended slightly beyond apex of cerci. Aedeagus: distiphallus bent sharply downward from angle of basiphallus in most specimens; antero-ventral margin dentate; in profile anterior edge slightly indented ca. 0.33 distiphallus length from ventral edge in most specimens, straight in few; apex broad or North American species of Siphona Meigen 297 narrow in profile. Sternum 5 (Fig. 10): average for genus. Female. As described for male except as follows. Head (Fig. 48). Palpus longer and thicker than in males in most specimens. Geographical distribution (Fig. 89). - Transcontinental, in narrow belt south of boreal forest. Habitat. - Label data indicates specimens of this species have been collected from sphagnum bogs. Also see S. cristata. Chorological affinities. - This species is evidently parapatric with sister species S. oligomyia. The range of this species is broadly sympatric with ranges of closely related S. cristata and S. multifaria , as well as with ranges of S. hokkaidensis, S. maculata, S. intrudens and S. medialis. Phylogenetic relationships (Fig. 104). - This species and S. oligomyia are interpreted as sister species on the basis of their highly derived colouration and setula on bend of R,. Together I interpret these species as the sister lineage to the S. cristata - S. multifaria lineage. Siphona oligomyia new species Holotype. - Male, labelled: “Keremeos, B.C. [British Columbia, Canada]/ 20. VI 1923/ C.B. Garrett” (CNC). Genitalia in microvial on pin below specimen. Derivation of specific epithet. - Formed from oligos and myia , Greek for “few” and “fly” respectively. This combination was chosen because the type-series is comprised of but two specimens, collected more than 1000 kilometers apart. Recognition. - Yellow ground and surface colour as in S. lutea. This is the only North American species with antero-lateral arms of sternum 5 hooked inward. R, distally with one setula on only one side in both specimens. Three postsutural dorsocentral setae. Female of species unknown. This species is compared with S. lutea under “Recognition” of that species. The characterization of S. oligomyia remains incomplete until more specimens are collected and examined and female of the species recognized. In particular, usefulness of proboscis length as a means for distinguishing S. oligomyia from S. lutea needs testing, and wing setular characteristics require additional data to establish trends. Description. - Length: 4.0-4. 5mm. Male. Head (Fig. 27). Colouration: as described for S. lutea. Eye: size average for genus though smaller than eye of S. lutea , 0.741-0.750 head height (Fig. 13), slender, widest at center, evenly rounded along anterior margin. Macrotrichia: average for genus. Antenna: first flagellomere 0.574-0.596 head height (Fig. 14), broadly rounded distally along ventral margin; length of aristomere 3 average for genus. Mouthparts: proboscis longer than in S. lutea, average for genus, 2.30-2.35 head height (Fig. 15). Thorax. Colouration: yellow in ground colour and pruinosity; tegula yellow; wing yellowish hyaline; femora and tibiae yellow. Macrotrichia: postsutural dorsocentrals, both specimens with three; wing: R4 + 5 not setulose beyond crossvein r-m (ie. average, cf. S. lutea); R, with single setula on apical third on one wing only in both specimens. Aeropod: claws and pulvilli slightly larger than average. Preabdomen. Colouration: yellow in ground and surface colour. Macrotrichia: median marginals absent from T]+2; lateral marginals on T, +2 strong. Genitalia (Fig. 69). Both holotype and paratype examined. Apex of surstylus ca. even with apex of cerci. Aedeagus: distiphallus slightly bent downward from angle of basiphallus; antero-ventral margin dentate; in profile anterior edge convex, not indented (as it is in most S. lutea); apex broad in profile. Sternum 5 (Fig. 1 1): antero-lateral margins hooked inward. Female. Unknown. Geographical distribution (Fig. 90). - Known only from Lake Tahoe, Calif., and Keremeos, B.C. Quaest. Ent., 1982, 18 (1-4) 298 O’Hara Chorological affinities. - Though this species has been inadequately collected, it is apparently parapatric with sister species S. lutea. Among other S', cristata group members, it is sympatric with S. cristata and S. multifaria. Similar in distribution are S. lurida and S. pacifica of the S. maculata group. Phylogenetic relationships (Fig. 104). - This species is sister species to S. lutea, as mentioned under that species. S. cuthbertsoni Curran (described from Salisbury, Zimbabwe) and Siphonopsis species have the antero-lateral arms of male sternum 5 hooked inward, as in S. oligomyia. S. oligomyia is not closely related to S. cuthbertsoni , and neither is closely related to Siphonopsis species. Paratype. — One male. USA: California: Lake Tahoe, 20. Vi. 1 5, A.K. Fisher (USN M). Siphona pisinnia new species Holotype. - Male, labelled: “USA N.M. [New Mexico] Grant Co./ Gila Nat. For. Cherry/ Ck. Campgrd. 21km.n./ Silver City 2250m./ 3-4.VIII.1980/ J.E. & W.M. O’Hara” (CNC). Allotype. - Female, same data as holotype (CNC). Derivation of specific epithet. - To my knowledge “pisinnia” has no classical meaning, and was chosen as an arbitrary combination of letters. Recognition. - Eye height and proboscis length average, first flagellomere long and broad in most specimens (the most distinctive external feature). Male preabdomen light coloured, faintly vittate. Female preabdomen darker, not vittate. Distiphallus slightly bent downward from angle of basiphallus, shape not very distinctive; anterior edge straight or slightly .rounded, apex broad or narrow in profile. Three postsutural dorsocentral setae in most specimens (87%). In Mexico male and female specimens are easily distinguished by head characteristics. Male genitalia of S. pisinnia are quite different from those of Mexican species. In southwestern United States S. cristita specimens have been caught with specimens of S. pisinnia. Differences in colouration, eye size, first flagellomere and proboscis lengths will in most instances separate specimens of these species. Most S. cristita specimens are slightly larger than those of S. pisinnia. Very similar to S. pisinnia is S. multifaria. Differentiation between the two is discussed under “Recognition” of S. multifaria. Description. - Length: 3. 0-5. 0mm. Male. Head (Fig. 23). Colouration: frontal vitta yellow to brown or reddish-brown; gena and lower parafacial white to light yellow; upper parafacial and fronto-orbital plate light yellow to brown, darkest on orbital plate; scape and pedicel testaceous to fuscous; first flagellomere fuscous to black; palpus yellow; proboscis reddish-brown or fuscous. Eye: size average for genus, 0.725-0.811 head height (Fig. 13), slender, widest at center, evenly rounded along anterior margin. Macrotrichia: average for genus. Antenna: first flagellomere highly varied, in most specimens long and broad, 0.577-0.740 head height (Fig. 14), in most specimens rounded distally along ventral margin, in few specimens slightly truncate; length of aristomere 3 average for genus. Mouthparts: proboscis length average for genus, 1.87-2.32 head height (Fig. 15). Thorax. Colouration: dorsum average, blue-gray or gray, finely mixed with brown in most specimens, 3 brown vittae visible in few 'specimens; tegula testaceous to fuscous; wing yellowish hyaline; legs average. Macrotrichia: postsutural dorsocentrals, 53 specimens with three, 8 with four [males and fema-les]. Acropod: claws and pulvilli average-sized. Preabdomen. Colouration: faintly vittate; most specimens black in ground colour on T5, T4 and medially on T)+2 and T3, rest of preabdomen yelldw in ground colour; surface blue-gray, gray or light brownish-gray on areas of black ground coloiir, dark brown to black around macrotrichial insertions; areas yellow in ground colour with surface yellow or light browp; in few specimens black ground colour extensive, over most of T3, all of T4 and T5, with broad vitta on T1+2. Genitalia (Fig. 65). Thirteen examined. Apex of surstylus even with or extended slightly beyond apex of cerci. Aedeagus: distiphallus slightly bent downward from angle of basiphallus; antero-ventral margin dentate; in profile anterior edge straight or slightly rounded; apex broad or narrow in profile. North American species of Siphona Meigen 299 Female. As described for male except as follows. Head (Fig. 49). Palpus longer and thicker than in males in most specimens. Preabdomen. Colouration: darker than male, not vittate; black in ground colour in most specimens, yellow laterally on T, _j_2 *n few specimens; surface blue-gray or gray, mixed with light brown. Geographical distribution (Fig. 83). - A western species, ranging from southwestern United States to central Mexico. Habitat. - I have collected specimens from one locality each in western New Mexico and eastern Arizona. At the former, in the pine-juniper zone, a large series was collected from flowers of a short geranium (unidentified), while a few specimens were caught on white sweet clover ( Meli lotus alba). In eastern Arizona, in the pine-spruce zone, specimens were swept from grass under cover of pine trees. Chorological affinities. - Uniquely distributed throughout southwestern United States and western Mexico, this species is narrowly sympatric with S. multifaria and S. cristita along the northern limit of its range. It is allopatric to other S. cristita group members. Phylogenetic relationships (Fig. 104). - Though not well characterized, the S. cristita group is considered monophyletic, and S. pisinnia is tentatively regarded as sister species to the other members of the group. Phenetically, S. pisinnia is close to S. multifaria. ParatypeS. — Sixty-eight males, 73 females. USA: Arizona: Rustler Pk., Portal, 8200', 3. VIII. 55, R.R. Dreisbach, 1M; Santa Catalina Mts., 14. VII. 50, L.D. Beamer, 1M; Dripping Springs, Organ Pipe, 31.111.67, D.M. Wood, 1 M; Sunnyside Canyon, 11. VII. 40, D.G. Hall, 1M; ibid., Huachuca Mts., 9. VI 1.40, E.E. Kenaga, IF; ibid., D.E. Hardy, 2F; Chiricahua Mts., 6. VIII. 33, Bryant, 1M; ibid., 7. VIII. 41, R.H. Beamer, 1M; Cochise Co., Chiricahua Mts., Flys Peak, 85-9700', 5. VIII. 27, J.A. Kusche, 1 M; Cochise Co., Southwestern Research Station, 8 km. w. Portal, 1650m, 1 3. VII. 56, E. Ordway, 1M; ibid., 3.IX.59, D.D. Linsdale, 1M; ibid., 9,22.X.64, P.H. Arnaud, Jr., 3F; ibid., 27.VII.65, 1M; ibid., 2. VIII. 65, IF; ibid., 22.IX.65, IF; ibid., 2.X.65, IF; ibid., 5-25.IX.65, C.W. Sabrosky, 2M, 13F; ibid., 6.VIII.65, 8, 26. IX. 66, V.D. Roth, 1M, 9F; ibid., 1 1,19, 22, 24. IX. 66, P.H. Arnaud, Jr., 1 M, 1 1 F. California: Snow Creek, 1500', 7. III. 55, W.R. Richards, IF; Garnet, 4.IV.45, A.L. Melander, IF; Indio, 13.XII.44, A.L. Melander, 2F; nr. Indio, 23.11.49, A.L. Melander, 1 M; Riverside, 2.V.35, A.L. Melander, 1 M; Riverside Co., Deep Canyon, 23.11.64, M.E. Irwin, IF; ibid., P.L. Boyd Desert Research Center, s. Palm Desert, 16-17. X, 4-6.XI.69, S. Frommer and R. Worley, 4F; ibid., 3 1 .1- 10.11.70, S. Frommer, R. Worley and L. La Pre, 3M, IF; ibid., 4- 1 1 .11.73, A. Tabet, 7F; ibid., 1 2.IV.75, J.B. Tucker, 1 M; ibid., J.B. Turner, 1 F. Nevada: Austin, 12.VIII.40, D.E. Hardy, 1 M. New Mexico: Grant Co., Gila Nat. For., Cherry Creek Cpgd., 21 km. n. Silver City, 2250m, 3-4.VIII.80, J.E. and W.M. O'Hara, 23M, 9F; Socorro Co., Magdalena Mts., 19.V1II.51, E.L. Kessel, 2F. MEXICO: Chihuahua: Mesa del Huracan, 108°15'N 30°04'W, 7400', 21-25. VII. 64, J.E.H. Martin, 2M. Durango: Rio Chico, 20 mi. w. Durango, 7000', 10. VIII. 64, J.F. McAlpine, 1 M; 10 mi. w. El Salto, 9000', 21.VI.64, J.F. McAlpine, 1 M; 6500', VII 1.64, J.F. McAlpine, 1 M; 24 mi. w. La Ciudad, 7000', 28.VI.64, W.R.M. Mason, 2M. Mexico: Pedregal de San Angel, 5-8.VII.78, G. and M. Wood, 16M, 2F. Sinaloa: El Palmito, 6400', 2.VII.64, J.F. McAlpine, 1 M. (Deposited in AMNH, CAS, CNC, DMW, JEOH, MSU, UASM, UCR, UKL and USNM.) The S. geniculata group North American species of the S. geniculata group have a short aristomere 3 (0.35-0.45 head height), average length proboscis and at least one of the following two characteristics: one pair of median marginal setae on T1+2 (Fig. 8), and/or a long surstylus which extends beyond apex of cerci by more than preapical width of surstylus (Figs. 70-71). Siphona geniculata (De Geer) Musca geniculata De Geer, 1776: 38. Crocuta geniculata'. Coquillett, 1910: 518. Siphona geniculata-. Mesnil, 1965: 866 (redescription).- Following New World records are based on misidentifications. Coquillett, 1897: 76.— Aldrich, 1905: 444.— Aldrich, 1934: 109.— Sabrosky and Arnaud, 1965: 1 064. — Cortes and Hichins, 1969: 57. -Cortes and Campos, 1970: 98.- Arnaud, 1978: 458. Siphona analis Meigen, 1824: 157.— Herting, 1972: 3. Siphona cinerea Meigen, 1824: 156.-- Herting, 1972: 4. Bucentes cinerea Latreille, 1809: 339. Quaest. Ent., 1982, 18 (1-4) 300 O’Hara Siphona meigenii (Lepeletier and Serville in Latreille, et al.), 1828: 501 ( Bucentes ). Stomoxys minuta Fabricius, 1805: 282. Siphona nigrovittata Meigen, 1824: 157.— Herting, 1972: 11. Siphona tachinaria Meigen, 1824: 156.-- Herting, 1972: 13. Musca urbanis Harris, 1780: 153. The type of Musca geniculata De Geer is lost (Herting, per. comm.). Nevertheless the species now recognized as S. geniculata (whether or not the original one) is readily distinguishable so a neotype is not required. Recognition. - Specimens are relatively large for Siphona. Eye height and proboscis length average, first flagellomere and aristomere 3 short. Most specimens with above average number of hairs (10-20) on upper parafacial and frontal plate. This is one of two species with both a long surstylus and 1 pair of strong median marginal setae on T1+2 (see below). Abdominal vitta present, broader than average, and obscured by dense pruinosity. Four postsutural dorsocentral setae. With median marginal setae on T1+2 and a long surstylus, S. geniculata is unlike any North American species other than S. hokkaidensis. Generally, shape and length of male first flagellomere permits separation of specimens of these species, since most S. hokkaidensis specimens with median marginals on T1+2 have large first flagellomeres (as in Fig. 32). Description. — Specimens examined: 6 males, 3 females from North America, 16 males and 11 females from Europe. Length: 4. 5-6. 5mm. Male. Head (Fig. 28). Colouration: frontal vitta testaceous, brown to reddish-brown; gena and lower parafacial white to light yellow; fronto-orbital plate and upper parafacial yellow to dark brown, darkest on orbital plate; scape and pedicel testaceous to fuscous; first flagellomere fuscous to black; palpus yellow, apex slightly infuscate in most specimens; proboscis reddish-brown or fuscous. Eye: size average for genus, 0.786-0.833 head height (Fig. 13), broad, widest at center, evenly rounded along anterior margin. Macrotrichia: cluster of 10-20 hairs on upper parafacial and frontal plate in most specimens; vibrissal angle average in most specimens, more heavily setose than average in few specimens with 2 or 3 setae and several setulae in addition to vibrissa and several hairs. Antenna: first flagellomere short, 0.420-0.484 head height (Fig. 14), rounded distally along ventral margin; aristomere 3 short for genus, in most specimens 0.35-0.45 head height. Mouthparts: proboscis length average for genus, 2.02-2.50 head height (Fig. 15). Thorax. Colouration: dorsum average, blue-gray or gray finely mixed with brown, not vittate; tegula testaceous to reddish-brown; wing yellowish hyaline; legs average. Macrotrichia: postsutural dorsocentrals, 25 specimens examined, all with four [males and females]. Acropod: claws and pulvilli average to slightly larger than average. Preabdomen. Colouration: dark, faintly vittate; T4, T5 and broad vitta on T, +2 and T3 black in ground colour, rest of preabdomen yellow in ground colour; surface dark gray or brownish-gray on areas of black ground colour, dark brown to black around macrotrichial insertions; areas of yellow ground colour with surface light brown or tawny. Macrotrichia: 1 pair median marginals on T1+2; 1 or 2 pair strong lateral marginals on T1+2 and T3. Genitalia (Fig. 91). Four examined, including 3 from European specimens. Apex of surstylus extended beyond apex of cerci by more than preapical width of surstylus. Aedeagus: slope of distiphallus even with angle of basiphallus (not bent); antero-ventral margin dentate; in profile parallel-sided and elongate; apex truncate or nearly so. Female. As described for male except as follows. Head (Fig. 50). Palpus equal or slightly longer than in males. Preabdomen. Colouration: dark, not vittate; black in ground colour, surface gray finely mixed with light brown. Geographical distribution (Fig. 91). - A wide ranging species in the Palearctic Region; introduced and established in the lower Fraser Valley of British Columbia. It has not, to date, been recorded elsewhere in North America. It is unlikely S. geniculata will spread beyond its restricted range unless its introduced host, Tipula paludosa , extends its range, or S. geniculata adapts to a native Tipula species. Chorological affinities. - At present S. geniculata occupies a narrow range in North America, and it is uncertain whether this species interacts with other Siphona species (such as closely related S. hokkaidensis ) in the area. Phylogenetic relationships (Fig. 105). - The sister species to this species is probably a Palearctic species. In North America, S. hokkaidensis is most closely related to it. North American species of Siphona Meigen 301 Siphona hokkaidensis Mesnil Siphona hokkaidensis Mesnil, 1957: 36.— Mesnil, 1965: 869.- Herting, 1967: 9 (comparison of S. hokkaidensis Mesnil and S'. silvarum Herting). Siphona silvarum Herting, 1967: 9. New synonymy. Type material examined. - Siphona hokkaidensis Mesnil, HOLOTYPE, female, labelled: “Obihiro [Hokkaido, Japan]/ S. Takano [handwritten label]”; “295 [green label]”; “Siphona/ hokkaidensis Mesnil/ L.P. Mesnil det., 1970”; “Type [red label]”; “EX/ L.-P. MESNIL/ COLLECTION 1970” (CNC). Siphona silvarum Herting, HOLOTYPE, male, labelled: “ex larvae/ Tipula/ irrorata Mg. [handwritten label]”; “Mooswald/./ Freiburg [West Germany] i.B./ IV. 1958 [handwritten label]”; “Siphona/ silvarum Hert./ L.P. Mesnil det., 1970”; “TYPE [red label]”; “EX/ L.-P. MESNIL/ COLLECTION 1 970” [slash immediately following Mooswald is on label] (CNC). Two puparia accompany type on same pin. Recognition. - Eye height, first flagellomere and proboscis lengths varied. Aristomere 3 short and thick in most specimens. Most eastern specimens without marginal setae on T]+2 and with first flagellomere subtriangular (Fig. 31). Specimens from Alberta, Saskatchewan and Manitoba with short slender first flagellomere (Fig. 30). Northwestern specimens varied, few with 1 or pair of median marginal setae on T1+2 and very large first flagellomere (Fig. 32). Few specimens with above average number of setae on vibrissal angle, up to 4 present. Apex of surstylus extended far beyond apex of cerci, distiphallus truncate or nearly so. Most specimens with four postsutural dorsocentral setae. Specimens with median marginal setae on T1+2 are similar to S. geniculata, and are discussed under “Recognition” of that species. Eastern specimens of S. hokkaidensis form a homogeneous group, readily distinguished in most instances by the subtriangular (male), apically truncate (both sexes) first flagellomere and short (and in most males, thick) aristomere 3. S. hokkaidensis specimens from other regions varied, as discussed below. In all regions male genitalia permit separation of this species from others, because of the very long surstylus and characteristic distiphallus (Fig. 71). Description. — Specimens examined: 271 males, 144 females. Length: 3. 5-5. 5mm. Male. Head (Figs. 30-32). Colouration: frontal vitta testaceous, brown to reddish-brown; gena and parafacial light yellow to light brown; fronto-orbital plate brown to dark brown; scape, pedicel, and in few specimens portion of first flagellomere above aristal insertion yellow, testaceous, or (in few specimens) fuscous; first flagellomere reddish-brown, fuscous to black; palpus yellow, in few specimens apex infuscate; proboscis testaceous to fuscous. Eye: size highly varied, 0.71 1-0.820 head height (Fig. 13), slender and narrowed below center to broad and evenly rounded along anterior margin. Maerotrichia: vibrissal angle with 2-4 setae and several hairs, in addition to vibrissa. Antenna: length and shape of first flagellomere extremely varied, 0.426-0.667 head height (Fig. 14), treated below under geographic variation; aristomere 3 short for genus, in most specimens 0.35-0.45 head height and thickened almost to tip. Mouthparts: length of proboscis highly varied, 1 .96-2.53 head height (Fig. 1 5). Thorax. Colouration: dorsum average, blue-gray or gray finely mixed with brown in most specimens, 3 brown vittae visible in few specimens; tegula testaceous to fuscous; wing yellowish hyaline; legs average. Maerotrichia: postsutural dorsocentrals, 7 specimens with three, 91 with four. Acropod: claws and pulvilli average-sized. Preabdomen. Colouration: extremely varied, vittate in all but very few specimens; eastern specimens, black ground colour on T5, vittate or entirely black on T4, vittate on T1+2 and T3, rest of preabdomen yellow in ground colour; areas black in ground colour with surface blue-gray, dark brown to black around macrotrichial insertions; areas yellow in ground colour with yellow or light brown surface; specimens from Alberta, Saskatchewan and Manitoba with ground colour as for eastern specimens, surface predominantly light coloured, brownish-gray on areas of black ground colour, yellow or light brown on areas of yellow ground colour; vitta visible though faint through light coloured pruinosity; dark areas around macrotrichial insertions very reduced; pattern in western specimens highly varied, most specimens similar to eastern specimens, few similar to specimens from Alberta, Saskatchewan and Manitoba, and few specimens unique - more extensively black in ground colour, yellow ground colour restricted to extreme lateral edge of T1 + 2 and antero-lateral edge of T3, not vittate; surface predominantly blue-gray, wide area around macrotrichial insertions dark brown to black. Maerotrichia: median marginals on T1+2 present or absent, data recorded in Fig. 81; 1 or 2 pair strong lateral marginals Quaest. Ent., 1982, 18 (1-4) 302 O’Hara on T1 + 2 and T3, data recorded for T3 in Fig. 81. Genitalia (Fig. 71). Twenty-seven examined, including 1 from a European specimen. Apex of surstylus extended beyond apex of cerci by more than preapical width of surstylus. Aedeagus: slope of distiphallus even with that of basiphallus, or distiphallus very slightly bent downward; antero-ventral margin dentate; in profile nearly parallel-sided; apex truncate or nearly so. Female. As described for male except as follows. Head (Fig. 51). Length of palpus subequal or slightly longer than in males. Preabdomen. Colouration: in all geographic regions ground colour pattern very similar, not vittate, yellow ground colour restricted to extreme lateral edge of T1+2 and antero-lateral edge of T3; eastern specimens very dark, surface blue-gray to gray, wide areas around macrotrichial insertions dark brown to black, black ground colour very evident through light coloured pruinosity; specimens from Alberta, Saskatchewan and Manitoba with surface brownish-gray, dark areas around macrotrichial insertions reduced, ground colour obscured by light coloured pruinosity; specimens from Washington, British Colombia, Northwest Territories, Yukon and Alaska with surface highly varied, from light gray or brownish-gray with light appearance as in Alberta, Saskatchewan and Manitoba specimens, to very dark like most eastern specimens. Variation, with notes about synonymy. - Polymorphism in S. hokkaidensis was investigated in some detail to provide the basis for a taxonomic decision regarding morphs. Two character systems in particular were amenable to analysis. These were shape of first flagellomere and number and position of abdominal setae. These characteristics were at least in part geographically distributed, as shown in Fig. 81. Three basic shapes of first flagellomere were recognized in males of S. hokkaidensis , permitting specimens to be sorted into three groups on that basis. The three antennal types, designated 1, 2 and 3, are illustrated in Figs. 30-32. Intermediate forms were few and limited to specimens from western North America in regions where more than one antennal type were present. Specimens with intermediate first flagellomeres were assigned the antennal type they most closely resembled. Females of S. hokkaidensis have smaller first flagellomeres than males, so the three antennal types recognized in males could not be applied to females. Females were treated in a fourth catagory (“f” in Fig. 81). Except for one sample from northern Quebec (Indian House Lake), all male specimens examined from Ontario eastward had a type 2 antenna. In other characteristics this group varied little. Males from Alberta, Saskatchewan and Manitoba also formed a homogeneous group, sharing type 1 antenna, a slightly larger eye and shorter proboscis than average for the species, and similar colouration. This group was called S. hokkaidensis “a” and treated separately from other specimens of the species ( S . hokkaidensis^ b”) in the Hubbs-Hubbs diagrams. Locality records for specimens of this group are represented by squares on the distribution map (Fig. 81). Throughout most of the Northwest Territories type 3 antenna predominated among specimens examined. In Alaska and the Yukon all three antennal types were represented in localities in relatively close proximity to one another, with mixed series recorded in two samples. Abdominal setae reflected a similar geographic pattern to the one exhibited by antennal variation (Fig. 81). Median marginal setae on T1+2 were present in a greater percentage of specimens with type 3 antenna than any other. However, in northwestern North America specimens with antennal types 1 and 2 had a higher occurrence of median marginals than they did anywhere else. The number of lateral marginals on T3 also conformed to this pattern (Fig. 81). My interpretation of these data is as follows. Whatever events produced polymorphism in S. hokkaidensis , they apparently did not result in permanent interruption of gene flow between North American species of Siphona Meigen 303 morphs, as exemplified by the heterogeneous populations in northwestern North America. Therefore I regard all morphs as members of a single species, S. hokkaidensis. Most distinctive of all were specimens with type 1 antenna from Alberta, Saskatchewan and Manitoba, and for this reason they were treated separately in the Hubbs-Hubbs diagrams. Unfortunately, there were too few specimens of S. hokkaidensis from the western provinces to permit detailed interpretation of the relationship between S. hokkaidensis “a” and “b”. Primarily because specimens with a type 1 antenna were found in northwestern North America with other characters varied, I concluded that S. hokkaidensis “a” does not represent a new species or subspecies. Furthermore, two autapomorphies support the present interpretation of S. hokkaidensis as one species: male genitalia are unique and almost unvaried over the North American range of the species, and the arista is thickened in most specimens irrespective of antennal type. Differences between specimens with antennal types 2 and 3 are slight, so there is little reason to suspect they represent more than morphs of a single species. Dr. Herting noted (per. comm.) that both antennal types are represented in European specimens of “ S . silvarum", so the situation is geographically widespread. Mesnil first described S. hokkaidensis from a female specimen from Hokkaido, Japan (1957), then redescribed the species from European specimens in his treatment of Siphona in Lindner’s “Die Fliegen der palaearktischen Region” (1965). Herting (1967) compared European and Japanese specimens and noted the former lacked median marginals on T1+2 and differed in colouration from the latter. At the time neither form was known from North America, and their distributions appeared disjunct in the Palearctic Region. There was sufficient reason for believing the forms represented two species, so Herting (1967) named S. hokkaidensis of Mesnil, 1965 (not 1957) Siphona silvarum. From data presented herein the varied nature of abdominal setae in S. hokkaidensis is documented. The type of S. silvarum conforms in all respects to the present interpretation of S. hokkaidensis - bearing a type 2 antenna and lacking median marginals on T]+2 - so it has been synonymized with S. hokkaidensis. Geographical distribution (Fig. 81). - Holarctic, in the Palearctic Region ranging over Europe, USSR and Japan; in the Nearctic Region widely distributed in Canada, Alaska and extreme northeastern United States. This species has a more northern distribution in Canada than any other Siphona species. Chorological affinities. - This species is sympatric with another S. geniculata group member, S. medialis, along the southern edge of its range. It is widely sympatric with S. maculata, S. intrudens, S. lutea, S. multifaria and S. cristata. Phylogenetic relationships (Fig. 105). - Among North American Siphona species, this species is most closely related to S. geniculata. However, several European species resemble S. hokkaidensis , and one of these is probably its sister species. Siphona medialis new species Holotype. - Male, labelled: “Cranberry I. [Island]/ Lockeport, N.S. [Nova Scotia, Canada]/ 25.VII.1958/ J.R. Vockeroth” (CNC). Allotype. - Female, same data as holotype (CNC). Derivation of specific epithet. - S. medialis is the only wide ranging North America species that is characterized by strong median marginal setae on T1+2. S', medialis is named in Quaest. Ent., 1982, 18 (1-4) 304 O’Hara recognition of this attribute. Recognition. - A dark coloured species with a very large eye, short first flagellomere and \ average-sized proboscis. Aristomere 3 short in most specimens. Legs of western specimens very dark, specimens from other areas with legs average to darker than average. Preabdomen black | in ground colour (not vittate), with 1 pair median marginal setae on T1+2. Three or four postsutural dorsocentral setae. This is one of the more distinctive species, with its dark colouration, large eye, median ! marginals on T1+2 and (in most specimens) dark legs. It can only be confused with the two other species with median marginals on T1+2. Male specimens of S’, hokkaidensis have a smaller eye, longer surstylus and are lighter j coloured than S. medialis. In addition, most specimens of S. hokkaidensis with median marginals on T1+2 are from northwestern North America and have large first flagellomeres (as in Fig. 32). S. medialis is most easily distinguished from S. geniculata by colouration, relative position of the apices of surstylus and cerci, and in almost all instances by range differences. Description. - Length: 4. 0-5. 5mm. Male. Head (Fig. 29). Colouration: frontal vitta testaceous to reddish-brown; gena and parafacial white to light brown; fronto-orbital plate light to dark brown, darkest on orbital plate; scape and pedicel black in most specimens, fuscous in few; first flagellomere black; palpus yellow, in few specimens apex infuscate; proboscis fuscous to black. Eye: very large for genus, 0.786-0.833 head height (Fig. 13), broad, widest at center, evenly rounded along anterior margin. Macrotrichia: vibrissal angle average in most specimens, few specimens more heavily setose than average with 2-4 setae in addition to vibrissa and several hairs. Antenna: first flagellomere on average slightly longer than, but similar in shape to, that of S. geniculata , 0.439-0.533 head height (Fig. 14); aristomere 3 short for genus, in most specimens 0.35-0.45 head height. Mouthparts: proboscis length average for genus, 2.00-2.51 head height (Fig. 15). Thorax. Colouration: dorsum average, blue-gray or gray finely mixed with brown in most specimens, 3 brown vittae visible in few specimens; tegula reddish-brown to black; wing yellowish to cinereous hyaline; leg colouration geographically varied; specimens examined from Washington, Idaho, Oregon and California with femora and tibiae fuscous in most specimens, reddish-brown in few; specimens from other areas average to darker than average, reddish-brown or fuscous portion covering dorsal and posterior surfaces of fore femur in few specimens, reddish-brown portions on mid and hind femora not covering more than apical two-thirds, most extensive dorsally, tibiae predominantly yellow. Macrotrichia: postsutural dorsocentrals, 1 1 specimens with three, 14 with four. Acropod: claws and pulvilli average-sized. Preabdomen. Colouration: dark; most specimens not vittate, black in ground colour, surface blue-gray, gray or brownish-gray, dark brown to black around macrotrichial insertions; in very few specimens T]+2 and T3 laterally yellow in ground colour, surface light brown. Macrotrichia: one pair median marginals on T1+2; lateral marginals on T1+2 strong. Genitalia (Fig. 72). Seven examined. Apex of surstylus even with or extended slightly beyond apex of cerci. Aedeagus: distiphallus bent slightly or sharply downward from angle of basiphallus; antero-ventral margin dentate; apex broad in profile. Female. As described for male except as follows. Head (Fig. 52). Palpus equal or slightly longer than in males. Preabdomen. Colouration: not vittate; as described for average male. Geographical distribution (Fig. 94). - Transcontinental; widely distributed in western United States, but restricted to region along Canada-United States border in central and eastern North America. Habitat. - I have collected specimens of this species only once, in southwestern Utah. Adults were swept from a damp meadow of sedge and grass. Specimens collected by J.F. McAlpine and R.L. Hurley in Ninette, Manitoba, were caught in Poa- sedge meadows (label data). Chorological affinities. - This species is sympatric with S. hokkaidensis along the northern edge of the formers’ range. In addition to being transcontinental, S. medialis is widely distributed in western United States, and is sympatric with all Siphona species of America north of Mexico except S. pisinnia, S. illinoiensis and S. floridensis. Phylogenetic relationships (Fig. 105). - This species is related to the S. geniculata - S. hokkaidensis lineage of the S. geniculata group. North American species of Siphona Meigen 305 Paratypes. - Twenty-six males, 40 females. CANADA: Alberta: 15mi. e. Morley, 23. VI. 62, W.R.M. Mason, 1M; 13mi. n. Banff, Banff-Jasper Hwy., 4500', 26. VII. 55, R. Coyles, IF. Manitoba: Ninette, 14. VI. 58, J.F. McAlpine, 1M; ibid., R.L. Flurley, IF; 2mi. e. Douglas, 27. VI 1.58, N.B. Chillcott, 1 F. New Brunswick: Kouchibouguac N.P., 7,1 3. VI 1.77, J.F. McAlpine, 2M, IF; Birch Cove, nr. Chamcook, 4. VII. 65, G.E. Shewell, 4M. Nova Scotia: Cranberry I., Lockeport, 18,20,24-25, 27-28, 30.VII.58, J.R. Vockeroth, 2M, 6F; Halifax Co., Lawrencetown, 19-20.VII.67, D M. Wood, 2M, IF. Prince Edward Island: Green Gables, Cavendish Beach, 22. VII. 67, D M. Wood, 2F. Quebec: mi. 139, rte. 58, La Verendrye Prov. Pk„ 29. VI, 1 VI 1.65, D.M. Wood, 2M, 3F. USA: California: Nevada Co., Sagehen, 5mi. nw. Hobart Mills, 20. VI. 54, M.T. James, 2F; ibid., 25. VI. 54, R.M. Bohart, IF; ibid., 2.VII.54, E.I.Schlinger, IF; ibid., 22. VI. 72, R.A. Belmont, IF; ibid., 15.VII.64, M.E. Irwin, 1 M, IF; Merced Co., Dos Palos, 7.V.49, IF; Sierra Co., 4.8mi. se. Sierraville, 14. VI. 59, G.W. Byers, 1M. Colorado: Estes Pk., 7500', 2. VII. 61, C.H. Mann, 2M, IF; Hartsel, 20.VI.40, A. L. Melander, 2M, 2F; Tenn. Pass, 26.VII.17, J.M. Aldrich, IF; 2061, IF. Idaho: Moscow, 30.V.1 1, J.M. Aldrich, 1M. Maine: Sagahadoc Co., Popham Beach, 15. VII. 71, P. Ward, IF. Michigan: Iosco Co., 30. VII. 48, R.R. Dreisbach, IF. Minnesota: Eaglesnest, 30. VI. 59, W V. Balduf, IF. Montana: Bozeman, 20. VI. 06, 1M; Powell Co., Nigger Hill, VII, W.M. Mann, IF. Oregon: Klamath Co., Mare's Egg Spring, 12. VI. 64, J. Schuh, 2M; ibid., Fort Klamath, 26. VI. 52, E.I. Schlinger, IF. Utah: Grizzly Ridge Camp, 30mi. n. Vernal, 8400', 8. VII. 61, J.G. Chillcott, IF; Kane Co., Dixie Nat. For., Duck Ck. Cpgd., 50km. se. Cedar City, rte. 14, 2620m, 16-17.VIII.80, J.E. and W.M. O'Hara, 1 M, 5F. Washington: Pullman, 23.V.18, 4.VI.22, A.L. Melander, 1 M, IF. (Deposited in CNC, INHS, JEOH, MSUB, UASM, UCD, UCR, UKL, USNM and WSUP.) The S.futilis group Members have an average length proboscis, large eye and narrow, short to average length first flagellomere. S. brunnea is a provisional member. Siphona futilis Wulp Siphona futilis Wulp, 1890: 1 25.— Aldrich, 1905: 444.— Guimaraes, 1971: 170. Bucentes futilis : Curran, 1932: 13 (in key). Bucentes ceres Curran, 1932: 14. New synonymy. Siphona ceres’. Guimaraes, 1971: 170.— Arnaud, 1978:458. Notes about synonymy. - Curran included S. futilis in his 1932 key to adults of North American Siphona species. However, Curran’s key appears to have been written primarily from published species descriptions. There is no indication that Curran examined specimens of S’. futilis prior to his description of S. ceres. I regard the type of -S', ceres as conspecific with the lectotype of S.futilis. Type material examined. - Siphona futilis , 14 syntypes from BMNH, 2 syntypes from USNM. This series includes specimens of 2 species. One species is Siphonopsis plusiae (Coquillett), and I have chosen to retain this name as valid because this species is the type-species of Siphonopsis , and its life history has been investigated under that specific epithet (Bloeser, 1914). The other species has been described by Curran under the name Bucentes ceres , subsequent to the description of S. futilis. The lectotype here designated for Siphona futilis is conspecific with B. ceres , and replaces that name. LECTOTYPE, male, labelled: “Co-/ type [round yellow-ringed label]”; “Omilteme,/ Guerrero, [Mexico]/ 8000 ft./ July. H.H. Smith.”; “B.C.A. Dipt. II./ Siphona/ futilis,/ v.d.W.“; “(5”; “Central America./ Pres, by/ F.D. Godman./ O. Salvin./ 1903-172“ (BMNH). My lectotype label ’’LECTOTYPE/ Siphona/ futilis/ van der Wulp/ O’Hara designation/ Selected 1981 [red bordered, handprinted label]” has been attached to this specimen. PARALECTOTYPES, 7 specimens of Siphonopsis plusiae (Coq.) (6 BMNH, 1 USNM). Eight specimens (7 BMNH, 1 USNM) conspecific with lectotype: MEXICO, DISTRITO FEDERAL: Mexico City, V.1888, H.H.S., 1M. GUERRERO: same data as lectotype, 2M, 2F; Xucumanatlan, 7000ft, VII, H.H. Smith, 2M. VERACRUZ: Orizaba, XII. 1887, H,H.S. & F.D.G., IF. [BMNH specimens damaged in transit, most in very poor condition.] To these specimens both paralectotype and determination labels have been attached. Quaest. Ent., 1982, 18 (1-4) 306 O’Hara Bucentes ceres , HOLOTYPE, female, labelled: “Antigua, Guatemala]./ 5-V-1931/ D.M. Bates 506”; “Bucentes/ TYPE/ ceres 9/ Curran/ No. [red label]”; “Bucentes/ ceres / Curran [handwritten det. label]” (AMNH). My determination label “Siphona/ futilis / van der Wulp/ det./ J.E. O’Hara 1981” has been attached to this specimen. Recognition. - Eye large, first flagellomere short and proboscis length average. Scape and pedicel fuscous to black. Palpus yellow in most specimens, slightly infuscate apically in few specimens. Male preabdomen light coloured, vittate. Female preabdomen darker than in male; without vitta. Distiphallus bent sharply downward from angle of basiphallus, narrow in profile, apex sloped sharply postero-ventrally in most specimens. Three postsutural dorsocentral setae in most specimens (95%). Eye and proboscis sizes are very similar between S. futilis and S. illinoiensis. S. futilis has a shorter first flagellomere. There are also slight colour differences: scape and pedicel are dark coloured in S. futilis and light coloured in S. illinoiensis ; female preabdomen broadly vittate in S. illinoiensis , entirely dark in ground colour in S. futilis. There is a marked similarity between distiphalli of these species, attesting to their close relationship. The main difference is that the distiphallus is bent sharply downward from angle of basiphallus in S. futilis and is even with basiphallus in S. illinoiensis. Since S. futilis and S. illinoiensis are allopatric, the only species likely to be confused with the former is S. brunnea. Head characteristics are similar, but dark colouration of S. brunnea in combination with other features readily separate these species (see S. brunnea , “Recognition”). Description. — Specimens examined: 66 males, 53 females. Length: 4. 0-5. 0mm. Male. Head (Fig. 34). Colouration: frontal vitta testaceous to reddish-brown; gena and parafacial white, light yellow to light brown; fronto-orbital plate light to dark brown, darkest on orbital plate; scape and pedicel fuscous to black; first flagellomere black; palpus yellow, slightly infuscate at apex in few specimens; proboscis reddish-brown or fuscous. Eye: large for genus, 0.847-0.911 head height (Fig. 13), broad, widest at center, evenly rounded along anterior margin. Macrotrichia: average for genus. Antenna: first flagellomere 0.450-0.517 head height (Fig. 14), slender, rounded distally along ventral margin; length of aristomere 3 average for genus. Mouthparts: proboscis length average for genus, 2.10-2.41 head height (Fig. 15). Thorax. Colouration: dorsum average, blue-gray or gray finely mixed with brown in most specimens, 3 brown vittae visible in few specimens; tegula testaceous to reddish-brown; wing yellowish hyaline; legs average. Macrotrichia: postsutural dorsocentrals, 39 specimens with three, 2 with four. Acropod: claws and pulvilli average-sized. Preabdomen. Colouration: light coloured, vittate; yellow ground colour laterally on T1+2, T3 and anterolaterally on T4, rest black in ground colour; vitta long and narrow; areas black in ground colour with surface blue-gray, gray or brownish-gray, dark brown to black around macrotrichial insertions; areas yellow in ground colour with surface yellow to light brown; pattern varied in few specimens, slightly greater or lesser than average amounts of black ground colour. Macrotrichia: median marginals absent from T1+2; lateral marginals on T]+2 strong. Genitalia (Fig. 73). Seven examined. Apex of surstylus even with or extended slightly beyond apex of cerci. Aedeagus: distiphallus bent sharply downward from angle of basiphallus; antero-ventral margin dentate; in profile narrow, apex sloped sharply postero-ventrally in most specimens. Female. As described for male except as follows. Head (Fig. 53). Palpus longer than in males in few specimens. Preabdomen. Colouration: darker than male, without vitta; most specimens black in ground colour, very few laterally yellow in ground colour on T, +2; surface as described for male. Geographical distribution (Fig. 96). - Recorded from central Mexico to Costa Rica. Chorological affinities. - The range of this species encompasses most of the range of S. tropica , and the entire known ranges of S. brunnea, S. rizaba, S. akidnomyia and S. longissima. The ranges of S. futilis and S. pisinnia overlap in central Mexico. S. futilis is allopatric to sister species S. illinoiensis. Phylogenetic relationships (Fig. 107). - This species is sister species to 5. illinoiensis , from which it differs only slightly. North American species of Siphona Meigen 307 Siphona illinoiensis Townsend Siphona illinoiensis Townsend, 1891: 368.- Coquillett, 1 897: 76 (as syn. of S. geniculata).- Aldrich, 1 905: 444 (as syn. of S', geniculata).- Bezzi and Stein, 1907: 382 (as syn. of S. geniculata).- Sabrosky and Arnaud, 1965: 1064. Type material examined. - One syntype, here designated LECTOTYPE, male [not female], labelled: “9”; “Robertson/ S. Illinois”; “Siphona/ illinoiensis/ Tws”; “Type”; “COTYPE/ Siphona/ illinoiensis/ Tns. [red label]” (UKL). My lectotype label “LECTOTYPE/ Siphona/ illinoiensis Tnsd./ O’Hara designation/ Selected 1981 [red bordered label]” has been attached to this specimen. Townsend described S. illinoiensis from 2 specimens. The UKL collection has only one specimen labelled “COTYPE”; the one herein designated lectotype. However, the UKL collection has a male specimen of S. illinoiensis with same locality label as the type. It is probably the missing syntype. Recognition. - Eye large, first flagellomere and proboscis lengths average. Scape and pedicel yellow to testaceous. Male preabdomen light coloured, vittate. Female preabdomen slightly darker than male, with short broad vitta. Slope of distiphallus even with that of basiphallus. Distiphallus narrow in profile, apex sloped sharply postero-ventrally in most specimens. Three postsutural dorsocentral setae in most specimens (86%). Similarities and differences between S. illinoiensis and S. futilis are discussed under “Recognition” of the latter species. Within central and eastern North America, specimens of S. cristita and S. multifaria may be mistaken for S. illinoiensis. Most male specimens of S. illinoiensis can be distinguished by characteristics given in key and by comparing specimen measurements with those presented in the Hubbs-Hubbs diagrams. Male genitalia of S. illinoiensis are relatively unvaried, and sufficiently different from S. cristita and S. multifaria to aid in identifications. Not all female specimens of S. illinoiensis can be distinguished from those of S. cristita and S. multifaria. Eye and proboscis sizes can yield ambiguous results. In most specimens the light brownish-gray, non-vittate female preabdomen of S. illinoiensis can be distinguished from the abdominal colouration of S. cristita and S. multifaria. However, decisions regarding subtle colour differences are untrustworthy if specimens of only one species are at hand. Description. — Specimens examined: 217 males, 163 females. Length: 3. 0-5. 0mm. Male. Head (Fig. 1 ). Colouration: frontal vitta yellow, testaceous to reddish-brown; gena and parafacial white to light yellow; fronto-orbital plate light yellow to light brown or brown, darkest on orbital plate; scape, pedicel, and in few specimens portion of first flagellomere above aristal insertion yellow to testaceous; first flagellomere fuscous; palpus yellow; proboscis testaceous to reddish-brown. Eye: large for genus, 0.807-0.909 head height (Fig. 13), broad, widest at center, evenly rounded along anterior margin. Macrotrichia: average for genus. Antenna: first flagellomere 0.500-0.596 head height (Fig. 14), slender, rounded distally along ventral margin; length of aristomere 3 average for genus. Mouthparts: proboscis length average for genus, 1.93-2.49 head height (Fig. 15). Thorax. Colouration: dorsum average, blue-gray or gray finely mixed with brown in most specimens, 3 brown vittae visible in few specimens; tegula testaceous; wing yellowish hyaline; legs average. Macrotrichia: postsutural dorsocentrals, 91 specimens with three, 15 with four. Acropod: claws and pulvilli average-sized (Fig. 4). Preabdomen. Colouration: light coloured, vittate; most specimens black in ground colour on T4, T5 and medially on Ti+2 and T3, rest of preabdomen yellow in ground colour; areas black in ground colour with surface brownish-gray, brown to black around macrotrichial insertions; areas yellow in ground colour with surface light yellow or light brown to tawny; pattern varied in few specimens, areas black in ground colour slightly more, or less, extensive. Macrotrichia: median marginals absent from T, +2; lateral marginals on T, +2 strong. Genitalia (Fig. 9). Twenty examined. As described for S. futilis, except slope of distiphallus even with that of basiphallus in most specimens, slightly bent downward in few. Female. As described for male except as follows. Head (Fig. 54). Palpus longer and thicker than in males in most specimens. Preabdomen. Colouration: average specimens slightly darker than male, with yellow ground colour restricted to lateral edge of T1+2 and antero-lateral edge of T3, dark vitta short and broad; surface colour and variation as decribed for male. Quaest. Ent., 1982, 18 (1-4) 308 O’Hara Geographical distribution (Fig. 92). - Widely distributed in eastern United States. Chorological affinities. - This uniquely distributed species is parapatric or narrowly sympatric with S. hokkaidensis, S. maculata, S. lutea, S. medialis and S', cristata along the northern edge of its range. Widely sympatric with S. illinoiensis are S. intrudens and S. multifaria. S. illinoiensis is allopatric to sister species S.futilis. Phylogenetic relationships (Fig. 107). - This species is sister species to the Mexican species S. futilis. Siphona brunnea new species Holotype. - Male, labelled: “MEX. [Mexico] Chis. [Chiapas] 9600ft./ Zontehuitz, nr. S. Crist. [San Cristobal de las Casas]/ 17 May 1969/ W.R.M. Mason” (CNC). Allotype. - Female, same data as holotype (CNC). Derivation of specific epithet. - Brunneus is Latin for brown. This species is named for its very dark brown colouration. Recognition. - Eye large, first flagellomere short and proboscis length average. A dark brown species, with apical half of palpus fuscous or black, wing brownish hyaline and legs in some specimens darker than average. Male tarsal claws slightly longer than average. Male preabdomen not vittate, surface dark brown with a light coloured band across anterior portion of each tergite. Female abdominal colouration varied. Surstylus slightly longer than cerci. Distiphallus bent sharply downward from basiphallus, narrow and truncate in profile. Four postsutural dorsocentral setae. An easily recognized species, quite different from members of the Mexican S. tropica group. Similar in head characteristics to S.futilis , but distinguishable on basis of colouration, number of postsutural dorsocentrals and in particular by structure of distiphallus. Description. - Length: 4.5-5.0mm. Male. Head (Fig. 35). Colouration: frontal vitta dark brown to reddish-brown; gena and parafacial light brown; fronto-orbital plate dark brown; antenna black; palpus yellow basally, apical half fuscous or black; proboscis fuscous or black. Eye: large for genus, 0.833-0.843 head height (Fig. 13), broad, widest above center, slightly narrowed below center. Macrotrichia: average for genus. Antenna: first flagellomere 0.463-0.510 head height (Fig. 14), short and distally rounded; length of aristomere 3 average for genus. Mouthparts: proboscis length average for genus, 2.35-2.44 head height (Fig. 15). Thorax. Colouration: dorsum dark brown; tegula black; wing brownish hyaline; legs average to darker than average, reddish-brown regions covering dorsal and posterior surfaces of fore femur and distal half of mid and hind femora in two specimens. Macrotrichia: postsutural dorsocentrals, four in all specimens. Acropod: claws and pulvilli slightly larger than average. Preabdomen. Colouration: dark, not vittate; black in ground colour except for extreme lateral edge of T1+2 and T3; surface dark brown with light bands across anterior 0.25 of T3, T4 and T5. Macrotrichia: median marginals absent from T, + 2; lateral marginals on T, +2 strong. Genitalia (Fig. 74). One examined. Apex of surstylus extended slightly beyond apex of cerci. Aedeagus: distiphallus bent sharply downward from angle of basiphallus; antero-ventral margin dentate; in profile narrow, apex truncate. Female. As described for male except as follows. Head (Fig. 55). Length of palpus subequal in males and females. Preabdomen. Colouration: light bands slightly wider than in male; one female with same ground colour pattern as male, other female vittate, with yellow ground colour dorso-laterally on T, +2. Geographical distribution (Fig. 97). - Known only from Chiapas, Mexico. Chorological affinities. - From label data this species has evidently been collected from the same locality as S. akidnomyia, though at a different time of year. Phylogenetic relationships (Fig. 107). - Due to the highly derived nature of this species, it is difficult to place with certainty. It is here interpreted as a member of the S.futilis group on the basis of its large eye and general habitus. North American species of Siphona Meigen 309 Paratypes. — One male, 1 female. MEXICO: Chiapas: Zontehuitz, nr. San Cristobal de las Casas, 9600', 17.V.69, W.R.M. Mason, 1M, IF. (Deposited in CNC.) The * S', tropica group Group characteristics are: long to very long proboscis and a very long first flagellomere, the latter in profile apically wide and broadly rounded along ventral margin. Most species have a large eye. Siphona tropica (Townsend) Phantasiosiphona tropica Townsend, 1915: 93.— Townsend, 1936: 150 (in key to genera of the Siphonini), Townsend, 1940: 286 (redescription of genus).- Guimaraes, 1971: 169. Bucentes tropica-. Curran, 1932: 13 (in key). Type material examined. - PARATYPE, male [not female], labelled: “Granada / Nicaragua/ Coll. Baker”; “V/ 77 [tiny handwritten label]”; “Paratype No./ 19954/ U.S.N.M. [red label]”; “Phantasiosiphona/ tropica/ Tns. det/ Sabrosky” (USNM). Recognition. - Eye large, first flagellomere and proboscis long. Wing brownish hyaline, with single setula on bend of Rj in most specimens (89%). Preabdomen light coloured, vittate in both sexes. Distiphallus distinctive, slope even with that of basiphallus, apex narrow and pointed in profile. Three postsutural dorsocentral setae. Similar to S. longissima and S. rizaba in proboscis length, and to latter in eye height. Most specimens of S. tropica are readily identified by presence of a setula distally on R,. No other Middle American species is known to exhibit this character state. Structure of the male distiphallus is particularly diagnostic for this species. Description. — Specimens examined: 14 males, 6 females. Length: 4. 0-5. 0mm. Male. Head (Fig. 38). Colouration: frontal vitta testaceous to dark brown; gena and parafacial white, light yellow to light brown; fronto-orbital plate light to dark brown, darkest on orbital plate; antenna fuscous; palpus yellow; proboscis testaceous, reddish-brown to fuscous. Eye: large for genus, 0.860-0.891 head height (Fig. 13), slender, widest above center, narrowed below center. Macrotrichia: average for genus. Antenna: first flagellomere very long and broad for genus, 0.654-0.745 head height (Fig. 14), rounded distally along ventral margin and in most specimens produced slightly forward distally along dorsal margin; length of aristomere 3 average for genus. Mouthparts: proboscis long for genus, 2.76-3.02 head height (Fig. 15). Thorax. Colouration: dorsum average, blue-gray or gray finely mixed with brown in most specimens, 3 brown vittae visible in few specimens; tegula fuscous to black; wing brownish hyaline; legs average. Macrotrichia: postsutural dorsocentrals, three in all 19 specimens examined; wing, 19 specimens examined: one seta on both wings in 14 specimens; 1 seta on one wing only in 3 specimens; setae absent, 2 specimens. Acropod: claws and pulvilli average-sized. Preabdomen. Colouration: vittate; varied, yellow ground colour ranging from laterally on T1+2 and antero-laterally on T3 to laterally on T]+2, T3 and antero-laterally on T4, surface yellow, light brown to tawny; areas black in ground colour with surface blue-gray, gray to brown, darker around macrotrichial insertions. Macrotrichia: median marginals absent from T, +2; lateral marginals on T, +2 strong. Genitalia (Fig. 78). Four examined. Apex of surstylus even with or extended slightly beyond apex of cerci. Aedeagus: slope of distiphallus even with that of basiphallus (not bent); antero-ventral margin dentate; apex narrow and pointed in profile. Female. As described for male except as follows. Head (Fig. 57). Length of palpus subequal in males and females. Preabdomen. Colouration: vittate; yellow ground colour laterally on T1 + 2 and antero-laterally on T3, otherwise as described for male. Geographical distribution (Fig. 101). - Recorded from southern Mexico to western Colombia. Habitat. - Recorded from cloud forest at 5100' in Puebla, 5 mi. ne. of Teziutlan. In Veracruz (Fortin de las Flores) specimens of this species were collected between 2500-3000' (from label data). Quaest. Ent., 1982, 18 (1-4) 310 O’Hara Chorological affinities. - Though widely distributed throughout Central America and southeastern Mexico, records do not show specimens of this species as having been collected with those of other Middle American Siphona species. Phylogenetic relationships (Fig. 106). - Relationships among S. tropica group members are unclear, for reliable synapotypies have not been discovered. S. tropica and S. rizaba are tentatively grouped as sister species on the basis of a very large eye and the loss of hooks on the distiphallus. Siphona rizaba new species Holotype. - Male, labelled: “Orizaba, Ver[acruz].,/ Mex[ico]. 8/12-22/61/ R&K Dreisbach” (MSU). Genitalia in microvial on pin below specimen. Allotype. - Female, same data as holotype, with Reinhard determination label: “ Bucentes / tropica/ Tm. R. [Reinhard initial]” (MSU). Derivation of specific epithet. - This species is named after the type-locality, Orizaba, from which the first letter has been removed. Recognition. - Eye large, first flagellomere and proboscis long. Wing yellowish to brownish hyaline. Preabdomen light coloured, vittate in both sexes. Externally without good diagnostic characteristics. Apex of surstylus extended slightly to far beyond apex of cerci. Distiphallus slightly bent downward from basiphallus, apex rounded in profile. Three postsutural dorsocentral setae. This species is the least distinctive of the S. tropica group. It shares the general characteristics of the other members of the group, but lacks good diagnostic features. S. tropica specimens without a single setula on R] can be confused with this species. Dissection of male genitalia is advisable where ratios of first flagellomere to eye and head heights are near upper values for S. rizaba and lower values of S. tropica and S. longissima (Figs. 14,16). Description. - Length: 4. 0-5. 0mm. Male. Head (Fig. 38). Colouration: frontal vitta testaceous to brown; gena and lower parafacial white, light yellow to light brown; upper parafacial and frontal plate light brown to dark brown; orbital plate brown to dark brown; scape and pedicel testaceous to reddish-brown or fuscous; first flagellomere fuscous; palpus yellow; proboscis testaceous to reddish-brown. Eye: large for genus, 0.846-0.870 head height (Fig. 13), slender, widest at center, slightly narrowed below center. Macrotrichia: average for genus. Antenna: first flagellomere long and broad, 0.593-0.648 head height (Fig. 14), broadly rounded distally along ventral margin; length of aristomere 3 average for genus. Mouthparts: proboscis long for genus, 2.74-3.03 head height (Fig. 15). Thorax. Colouration: dorsum average, blue-gray or gray finely mixed with brown in most specimens, 3 brown vittae visible in few specimens; tegula reddish-brown to fuscous; wing yellowish to brownish hyaline; legs average. Macrotrichia: postsutural dorsocentrals, three in all specimens. Acropod: claws and puivilli average-sized. Preabdomen. Colouration: vittate; T4, T5 and vitta on T1+2 and T3 black in ground colour, surface gray, brownish-gray or brown, darker around macrotrichial insertions; T1+2 and T3 laterally yellow in ground colour, surface yellow to light brown. Macrotichia: median marginals absent from T1+2; lateral marginals on T]+2 strong. Genitalia (Fig. 79). Four examined. Apex of surstylus extended slightly beyond to more than preapical width of surstylus beyond apex of cerci. Aedeagus: distiphallus bent slightly downward from angle of basiphallus; antero-ventral margin dentate; apex rounded in profile. Female. As described for male except as follows. Head (Fig. 58). Palpus subequal in length in males and females. Preabdomen. Colouration: short vittate; yellow ground colour restricted to T1+2, otherwise as described for male. Geographical distribution (Fig. 98). - Recorded from two localities in Veracruz, Mexico. Chorological affinities. - This species is evidently sympatric with S. futilis and S. tropica , but specimens of it have not been caught together with those of any other species. Phylogenetic relationships (fig. 106). - See this heading under S. tropica. Paratypes. - Three males, 2 females. MEXICO: Veracruz: Orizaba, 12-22. VIII. 61, R. and K. Dreisbach, 1M, 2F; Cordoba, 13,20.VII.66, J.S. Buckett, M R. and R.C. Gardner, 2M; (Deposited in CNC, MSU and UCD.) North American species of Siphona Meigen 311 Siphona longissima new species Holotype. - Male, labelled: “MEX. [Mexico] Chis. [Chiapas] lOmi NE./ San Cristobal [de las Casas]/ 13.V.69 7500'/ H.J. Teskey” (CNC). Genitalia in microvial on pin below specimen. Derivation of specific epithet. - From longissimus, Latin for “longest”. S. longissima has the longest proboscis of the species described in this paper. Recognition. - Eye large and first flagellomere long. Proboscis very long, one specimen with longest proboscis of any recorded North American species (3.2X head height). Wing brownish hyaline. Tarsal claws intermediate in size between average and large. Preabdomen vittate in male. Distiphallus distinctive, bent sharply downward from basiphallus, antero-ventral margin with small hooks, apex narrow and pointed in profile. Three postsutural dorsocentral setae. Female of species unknown. Most similar to S. tropica and S. rizaba. Other than characteristic sizes of eye, first flagellomere and proboscis, S. longissima can be distinguished by shape of distiphallus and the small hooks (as in S. macronyx and S. akidnomyia) along its antero-ventral margin. Description. - Length: 4.5-5.0mm. Male. Head (Fig. 37). Colouration: frontal vitta brown; gena and parafacial light brown; fronto-orbital plate brown to dark brown, darkest on orbital plate; antenna fuscous to black; palpus yellow; proboscis fuscous. Eye: slightly larger than average for genus, 0.796-0.850 head height (Fig. 13), slender, widest above center, narrowed below center. Macrotrichia: average for genus. Antenna: first flagellomere very long and broad for genus, 0.650-0.667 head height (Fig. 14), rounded distally along ventral margin; length of aristomere 3 average for genus. Mouthparts: proboscis long for genus, 2.74-3.20 head height (Fig. 15). Thorax. Colouration: dorsum darker than average, dark brown; tegula fuscous to black; wing brownish hyaline; legs average. Macrotrichia: postsutural dorsocentrals, three in all specimens. Acropod: claws and pulvilli intermediate in size between average and large. Preabdomen. Colouration: vittate; yellow ground colour laterally on T1+2 and T3, surface yellow; areas black in ground colour with surface gray to brown, darker around macrotrichial insertions. Macrotrichia: median marginals absent from T| +2; lateral marginals on T, +2 strong. Genitalia (Fig. 77). Two examined. Apex of surstylus even with or extended slightly beyond apex of cerci. Aedeagus: distiphallus sharply bent downward from angle of basiphallus; antero-ventral margin with small hooks (similar to S. macronyx and S. akidnomyia)-, apex narrow and pointed in profile. Female. Unknown. Geographical distribution (Fig. 100). - Recorded from southern Mexico. Chorological affinities. - Specimens of this species have been caught in the same area as those of S.futilis and S. akidnomyia , but on different dates (from label data). Phylogenetic relationships (Fig. 106). - The position of this species within the S. tropica group is tentatively as sister species to the highly derived S. akidnomyia. Paratypes. - Two males. MEXICO: Chiapas: 3mi. n. San Cristobal de las Casas, 7000', 29.V.69, H.J. Teskey, 1M. Distrito Federal: Desierto de Los Leones, X.65, N.L.H. Krauss, 1 M. (Deposited in CNC and USNM.) Siphona akidnomyia new species Holotype. - Male, labelled: “MEX. [Mexico] Chis. [Chiapas] 9600ft./ Zontehuitz, nr. S. Crist. [San Cristobal de las Casas]/ 25 June 1969/ W.R.M. Mason” (CNC). Genitalia in microvial on pin below specimen. Allotype. - Female, labelled: “MEXICO, Chiapas/ Mt. Tzontehuitz/ 9400' 12mi. NE San/ Cristobal [de las Casas] 27. V./ 1969 B.V. Peterson” (CNC). Derivation of specific epithet. - Derived from the Greek for feeble ( akidnos ) fly ( myia ). This is in reference to Siphona in general, which lack the size and strength of their larger relatives. Quaest. Ent., 1982, 18 (1-4) 312 O’Hara Recognition. - Eye size average, first flagellomere and proboscis long. Palpus long, apex infuscate in most specimens. Wing brownish hyaline. Male tarsal claws large. Male preabdomen dark, broadly vittate, female preabdomen average vittate. Distiphallus unique, short and broad in profile with small hooks on antero-ventral margin. Three postsutural dorsocentral setae. Closest in appearance to, yet readily distinguished from, S. tropica , S. rizaba and S. longissima. S. akidnomyia is the darkest of these species (though lighter than S. brunnea ), and has a shorter proboscis, smaller eye and longer tarsal claws than the others. The infuscate palpus is very large for Siphona , especially in the female. The distiphallus has small hooks along its antero-ventral margin (as in S. macronyx and S. longissima ), and a very distinctive shape. Description. - Length: 4. 0-5. Omm. Male. Head (Fig. 36). Colouration: frontal vitta brown; gena and parafacial white to light brown; fronto-orbital plate brown to dark brown, darkest on orbital plate; antenna fuscous to black; palpus yellow, apex infuscate; proboscis fuscous. Eye: size average for genus, 0.781-0.804 head height (Fig. 13), slender, widest at center, slightly narrowed below center. Macrotrichia: average for genus. Antenna: first flagellomere long and broad for genus, 0.585-0.660 head height (Fig. 14), broadly rounded distally along ventral margin; length of aristomere 3 average for genus. Mouthparts: palpus longer than average for genus; proboscis long for genus, 2.49-2.66 head height (Fig. 15). Thorax. Colouration: dorsum average to dark brown; tegula fuscous; wing brownish hyaline; legs average. Macrotrichia: postsutural dorsocentrals, three in all specimens. Acropod: claws and pulvilli large. Preabdomen. Colouration: broadly vittate; laterally on T|+2 and laterally or antero-laterally on T3 yellow in ground colour, surface yellow; areas black in ground colour with surface blue-gray, gray to brown, darker around macrotrichial insertions. Macrotrichia: median marginals absent from T1+2; lateral marginals on Tl+2 strong. Genitalia (Fig. 76). Five examined. Apex of surstylus even with or extended slightly beyond apex of cerci. Aedeagus: distiphallus slightly or sharply bent downward from angle of basiphallus; narrow section between basiphallus and distiphallus longer than average; antero-ventral margin with small hooks (similar to 5. macronyx and S. longissima ); in profile distiphallus short and broad, apex pointed. Female. As described for male except as follows. Head (Fig. 59). Palpus larger than in males, very long for genus, apex infuscate in one female. Preabdomen. Colouration: vitta average; surface colour as described for male, except slightly lighter. Geographical distribution (Fig. 99). - Recorded from southern Mexico. Chorological affinities. - Evidently sympatric with S. longissima, S. brunnea and S.futilis. Specimens of this species have been recorded from the same localities as the former two species, but were not collected on the same dates. Phylogenetic relationships (Fig. 106). - This is the most highly derived species of the S. tropica group. Intragroup relationships are difficult to interpret without additional characters. S. longissima is tentatively placed as sister species to S. akidnomyia on the basis of larger than average tarsal claws. Paratypes. - Four males, 1 female. MEXICO; Chiapas: Mt. Tzontehuitz, 9400', 12mi. e. San Cristobal de las Casas, 27.V.69, B.V. Peterson, 1M; 8mi. ne. San Cristobal de las Casas, 7500", 15.V.69, H.J. Teskey, 1M; San Cristobal de las Casas, 7087', 13, 28. VI. 69, B.V. Peterson, 1M,1F. Distrito Federal: Desierto Leones, 111, IV, V. 65, N.L.H. Krauss, 1M. (Deposited in CNC and USNM.) The S. macronyx group This group contains one highly autapotypic species, S. macronyx. The characteristics of this species (see “Recognition” below) are such that it cannot, at this time, be placed in one of the other North American Siphona species groups. Further study and additional characters might indicate that S. macronyx is a derived member of one of the other recognized groups. If so, then it should be placed in that group. North American species of Siphona Meigen 313 Siphona macronyx new species Holotype. - Male, labelled: “Pullman, Wash. [Washington, USA]/ Malaise Trap.”; “R.W. Dawson/ May 6 1969” (WSUP). Genitalia in microvial on pin below specimen. Allotype. - Female, same data as holotype except date May 13, 1969 (WSUP). Derivation of specific epithet. - From the Greek words for large claw, makros and onyx, after this attribute of the species. Recognition. - Head with characteristic habitus due to broad eye, above average number of setae and setulae on vibrissal angle, short first flagellomere and dark colouration. Proboscis average length. Male tarsal claws large. Preabdomen dark, in male broadly vittate, in female entirely black in ground colour. Distiphallus bent sharply downward from angle of basiphallus with several small hooks along antero-ventral margin. Three postsutural dorsocentral setae. The derived features of this species are such that specimens are not closely similar to those of any other species. Description. - Length: 4.0-5. 5mm. Male. Head (Fig. 33). Colouration: frontal vitta orange, testaceous to reddish-brown; gena and parafacial light brown in most specimens, light yellow in few; fronto-orbital plate brown to dark brown; scape, pedicel, and in few specimens portion of first flagellomere above aristal insertion yellow, testaceous to reddish-brown; first flagellomere fuscous to black; palpus yellow; proboscis reddish-brown to fuscous. Eye: size average for genus, 0.742-0.804 head height (Fig. 13), broad, widest at center, evenly rounded along anterior margin. Macrotrichia: vibrissal angle in most specimens with cluster of 3-6 setae and several setulae in addition to vibrissa. Antenna: first flagellomere short, 0.403-0.500 head height (Fig. 14), rounded distally along ventral margin; length of aristomere 3 average for genus. Mouthparts: proboscis length average for genus, 2.00-2.34 head height (Fig. 15). Thorax. Colouration: dorsum average, blue-gray or gray finely mixed with brown in most specimens, 3 brown vittae visible in few specimens; tegula testaceous to reddish-brown; wing brownish hyaline; legs average. Macrotrichia: postsutural dorsocentrals, 60 specimens with three, 1 with four [males and females]. Acropod: claws and pulvilli large. Preabdomen. Colouration: dark, broadly vittate; T4, T5 and vitta on T1+2 and T3 black in ground colour, surface gray to brown, black around macrotrichial insertions; yellow ground colour laterally on Tl+2 and T3 in most specimens, surface yellow to light brown; in 2 specimens yellow ground colour very reduced. Macrotrichia: median marginals absent from T1+2; lateral marginals strong. Genitalia (Fig. 75). Two examined. Apex of surstylus even with or extended slightly beyond apex of cerci. Aedeagus: distiphallus bent sharply downward from angle of basiphallus; antero-ventral margin with small hooks (similar to S. longissima and 5. akidnomyia)', apex broad in profile. Female. As described for male except as follows. Head (Fig. 56). In most specimens gena yellow, parafacial and frontal plate bright yellow tinged with brown (near golden); length of palpus slightly longer than in male in few specimens. Preabdomen. Colouration: not vittate; black in ground colour except for faint traces of yellow ground colour laterally on T1+2 in few specimens; surface light blue-gray, gray or brownish-gray, mixed with brown medially. Geographical distribution (Fig. 88). - Known only from Washington and western Idaho. Chorological affinities. - The small known range of this species is sympatric with the northern limits of the ranges of western species S. lurida , S. oligomyia and N. pacifica. S. macronyx is parapatric or sympatric with S. hokkaidensis, S. maculata, S. intrudens, S. lutea, S. multifaria, S. medialis and S. cristita, though specimens of it have not been collected with specimens of any of these species (from label data). Phylogenetic relationships (Fig. 103). - See discussion under species group heading above. Paratypes. - Eleven males, 49 females. USA: Idaho: Moscow Mt., 5. VII. 19, A.L. Melander, 1M. Washington: Whitman Co., Almota, 27. IV. 72, W.J. Turner, 1 M, 2F; Jefferson Co., falls outside Olympic Nat. Pk. entrance, 15km. ne. Quinault, P.H. Arnaud, Jr., IF; Asotin Co., Fields’ Spr. St. Pk., 15. VI. 72, W.J. Turner, 3F; Dewatto, 7. VI. 06, J.M. Aldrich, IF; Chelan Co., Squillchuck St. Pk., 9mi. se. Wenatchee, 17, VI. 73, M. Jackson, IF; Lyle Grove, Pullman, 3. VI. 69, H.S. Telford, IF; ibid., 10. VI. 11, A.L. Melander, IF; Pullman, WSU Campus-bee area, 3.V.71, W.J. Turner, 1M; Pullman, nr. SCS Pond, WSU, 29.V.71, W.J. Turner, IF; Pullman, 9,1 1-12, 14, 19,28, 30.V, 1,4,6.VI.65, R.D. Akre, 4M, 9F; ibid., 7,18. V.66, 5-6,8, 15-17, 22,25-29.V, 1.VI.69, R.W. Dawson, 4M, 29F. (Deposited in CAS, USNM and WSUP.) Quaest. Ent., 1982, 18 (1-4) 314 O’Hara PHYLOGENETIC INTERPRETATION Introduction In the ensuing pages phylogenetic aspects of Siphona species are considered. The approach I have taken is cladistic, wherein character states are evaluated and polarized according to their inferred plesiotypic (ancestral) or apotypic (derived) nature. Assuming only apotypic states to be analyzable, these states are arranged in a nested pattern of hierarchies to produce a cladogram (a diagram depicting a succession of apotypic character states representing hypothetical phylogenetic relationships) most parsimonious with the data available. Since publication of Hennig’s English version of his book “Phylogenetic Systematics” (1966a), numerous practical accounts of cladism have been published (eg. Griffiths, 1972; Ross, 1974; Eldredge, 1979; Wiley, 1981). The reader is referred to these sources for a detailed exposition of the cladistic method of phylogenetic analysis used here. The term monophyly is subject to different interpretations. As used here it excludes paraphyly, being monophyly in Hennig’s sense (equivalent to holophyly of Ashlock, 1971). Within the Siphonini a monophyletic group of genera is distinguished on the basis of two shared derived character states (cf. Andersen, in press “b”): 1) anal vein extended to wing margin at least as fold (ground plan for the Tachinidae is anal vein terminating in wing membrane)5, and 2) three katepisternal setae, of which postero-dorsal is strongest and antero-ventral is longer than antero-dorsal (groundplan for at least the Siphonini is antero-dorsal seta longer than antero-ventral). This group is here referred to as the Siphona group, including the Old World genera recognized by Andersen (in press “b”) and 3 genera endemic to the New World. Genera included are: Aphantorhapha Townsend, Asiphona Mesnil, Ceranthia Robineau-Desvoidy, Pseudosiphona Townsend, Siphona Meigen and Siphonopsis Townsend. Peribaea Robineau-Desvoidy is excluded from this group, though it possesses state (1) above. If Peribaea is related to this group, it is evidently sister genus to all the others. Andersen (in press “b”) hypothesizes that Peribaea is the most primitive Old World siphonine genus, and this possibility is not excluded here. With the exception of Siphona, the genera of the Siphona group form an assemblage of species of unclear relationships. Of these genera I have examined many undescribed species from Middle and South America and suspect other inadequately collected areas also harbour new species. Recognized genera do not bear well such scrutiny, and I predict a generic revision will significantly alter their limits. Due to their similarities I refer to them collectively as Ceranthia s. lat. to distinguish them from Siphona and to focus attention on their unsettled status. Siphona is probably monophyletic, as evidenced by proboscis structure. The proboscis is long and geniculate, with the labella at least as long as prementum and at least (and in most species considerably more than) eye height in length (Fig. 1,18-59). Considered by Andersen (in press “b”) to be the ground plan state for the Siphonini, with subsequent loss in the Actia group ( Goniocera B. & B., Ceromya R.D. and Actia R.D.). North American species of Siphona Meigen 315 Most species of Ceranthia s. lat. have a labellar disc similar to that of a housefly. In North American species in which it is elongate ( Siphonopsis and Pseudosiphona species in particular), it can be distinguished by characteristics given in the “Reclassification” section. Similarity between the proboscis of several Oriental and eastern Palearctic Actia species and Siphona is also discussed in the “Reclassification” section. A detailed comparison of male genitalia of Siphona species with those of species of Ceranthia s. lat. will probably corroborate the monophyletic origin of Siphona species. Currently, states of the major characters of male genitalia within taxa of the Siphona group have not been polarized, so a cladistic analysis cannot be conducted. However, preliminary studies suggest most genitalic features are plesiotypic in Siphona, even though the combination of features that contribute to the general habitus of the male genitalia in that genus is unique. A detailed study of the male genitalia of Ceranthia s. lat. taxa (including many undescribed species) is needed to determine the synapotypic features of this character complex in Siphona species. Such a study might also provide the necessary framework upon which to build a classification for taxa of Ceranthia s. lat. Character analysis General notes. - I have recognized 15 characters as suitable for phylogenetic analysis of North American Siphona species. Each is discussed below in relation to delimitation and polarization of states. Difficulties peculiar to delimitation of mensural characters are discussed under Character 1, eye height. Out-group and in-group comparisons have been conducted to polarize character states. The out-group is Ceranthia s. lat. (defined in the previous section). The in-group is world Siphona species, as represented by specimens made available to me during the course of this study (see world list for described species examined). A character weighting system has not been used, as explained in the “Reconstructed phylogeny” section. One character useful for identifications has been excluded from the analysis. This character, number of postsutural dorsocentral setae, is either 3 or 4, and in many species the number is of diagnostic value. Both states are widely distributed in the out-group and in-group, so consequently the apotypic state could not be determined; if it had been, then the number of reversals necessary to account for its present distribution would probably have negated its usefulness. In the following section and in the character matrix (Fig. 102) an a represents the inferred plesiotypic state. The first apotypic state is given an A. Other apotypic states not assignable to a transformation series with the first are denoted by successive capitals letters; B, C, D, etc. The order in which these are named refers only to their position in the matrix, starting from the top. Transformation series are denoted by superscripts, negative numbers being used only with mensural characters in which the plesiotypic state is inferred to be an average value (ie. eye height). Characters and states Character 1 . Male eye height This character and proboscis length are difficult to analyse because of their complex pattern of inter- and intraspecific distribution. As such they represent transformation series of phylogenetic importance, albeit without the relatively unambiguous delimitation of states Quaest. Ent., 1982, 18 (1-4) 316 O'Hara common to meristic characters. I have employed a practical solution to this problem: delimitation of states based on apparent groupings about mean values (refer to Figs. 13,15), with consideration of concordance with other characters for species of ambiguous placement. The theoretical danger here is introduction of circularity into the analysis by allowing perceived relationships among certain characters to affect delimitation of others. To offset this departure from standard phylogenetic practice, I discuss doubtful species placements under each species group. Despite this drawback I regard the characters eye height and proboscis length as important to the elucidation of Siphona phylogenetics, and their treatment in the manner indicated necessary for their interpretation. The ranges given below for each eye height state are based only on consideration of species under revision, in accordance with criteria discussed above. The plesiotypic state is inferred to be an average-sized eye, as this state is the most widespread throughout the genus and across species groups. Both small and large eyes are considered apotypic. a - average-sized, mean between 0.76-0.82 head height. A*2 - very small eye of S. lurida. A-1 - small, though larger than eye of S. lurida , and less than 0.76 head height. A1 - larger than average, between 0.82-0.85 head height. A2 - very large, greater than 0.85 head height. Character 2. Male first flagellomere Six independent states are recognized in size and shape of male first flagellomere. The states of this character are too varied in taxa of Ceranthia s. lat. to permit their polarization by out-group comparisons. Since no state is distributed among examined species of world Siphona in a manner suggestive of plesiotypy, all states are presumed apotypic. Relationships between states are unknown (other than divisions of A, B and C), so they cannot be treated as a transformation series. A1 - narrow, average length to long, broadly rounded apically along ventral margin such that dorsal edge is apically pointed. A2 - shaped as in A1 though shorter. B1 - relatively broad in profile, average length to long, dorsal and ventral surfaces parallel, rounded or slightly truncate apically along ventral margin, not pointed as in A. B2 - shaped as in B1 but very long. C1 - narrow, short, rounded along ventral margin, not pointed. C2 - shaped as in C1 but of average length. D - triangular, apically truncate. Here S. hokkaidensis is treated separately according to the shape of first flagellomere of eastern North American specimens (Fig. 31), though other morphs resemble closely states C and F. E - short and apically rounded. F - very long, wide apically in profile and broadly rounded along ventral margin. Character 3. Proboscis length The plesiotypic state for Siphona at one time must have been a short proboscis, perhaps of the sort exhibited by Siphonopsis species. To my knowledge no extant Siphona species has a proboscis shorter than S. floridensis or S. maculata , so the truly ancestral state of the genus is North American species of Siphona Meigen 317 no longer represented. I have treated proboscis length as a transformation series from very short (plesiotypic) to very long (apotypic), as polarized by out-group comparisons. A regression from a derived to less derived state (ie. a reduction in proboscis length) is only inferred to have occurred in S', lutea and S. akidnomyia. An alternative exists to this polarization scheme. The S. maculata group is monophyletic, based on shape of first flagellomere, and contains two character states interpreted as symplesiotypies: a short proboscis and weak lateral marginal setae on Tl+2. Since both states are virtually limited to that group, either of two explanations account for their restricted distribution: (1) the S. maculata group branched from the main lineage of Siphona early in the history of the genus, and retained to the present ancestral states which became modified in the main Siphona lineage, or (2) the S. maculata group is relatively young, and states limited to it are apotypic. Due to lack of corroborative evidence for the second hypothesis I consider the first more parsimonious with available data. Criteria for delimitation of character states is discussed under eye height. a - very short, less than 1.55X head height. A1 - short, mean between 1.55-1.8X head height. A2 - slightly shorter than average, 1.8-2. IX head height. A3 - average-sized, 2.1-2.5X head height. A4 - long, 2.5-2.7X head height. A5 - very long, greater than 2.7X head height. Character 4. Aristomere 3 This character is represented by a transformation series which is easily polarized by in-group comparisons. Species with the derived states are apparently monophyletic in Siphona, for the states are in accord with other data. However, a few species in the ex-group (eg. Ceranthia flavipes (Coquillett) and C. abdominalis (Robineau-Desvoidy)) also possess state A1. a - average length for Siphona , 0.50-0.65 head height. A1 - shorter than average, 0.35-0.45 head height. A2 - short as in A1 and thickened to near tip. Character 5. Vibrissal angle States were determined by in-group and out-group comparisons. Although the states seem to represent a transformation series, no other evidence suggests the bearers of A1 and A2 ( S . macronyx and S. lurida respectively) are closely related. a - average condition, 2-3 setae and several hairs in addition to vibrissa. A1 - 3-6 setae and several setulae and hairs in addition to vibrissa. A2 - heavily setose, up to 10 setae and several setulae and hairs in addition to vibrissa. Character 6. Male colouration Colouration within the out-group is too varied to be useful in polarization of states in Siphona. Quaest. Ent., 1982, 18 (1-4) 318 O’Hara The most prevalent male colour pattern among Siphona species, and that least in accordance with other characters, is predominantly blue-gray over most of thorax (where it is mixed with light brown) and abdomen, the latter medially vittate and laterally yellow in ground colour on T1+2 and T3. This state is regarded as plesiotypic, as are slight deviations from it. a - given above. A - abdomen black in ground colour, not vittate, deep blue-gray as basic pattern. B1 - thorax dark in ground colour at least dorsally, abdomen partially to entirely yellow. B2 - body yellow in ground and surface colour. C - abdomen black in ground colour, not vittate, surface predominantly brownish-gray. D1 - brown pruinosity over most of dorsal surfaces. D2 - as in D1 only darker brown. Character 7. Wing setulae on R With varied frequency, R, wing setulae are present among species of a number of siphonine genera. Except in a few genera their presence is evidently autapotypic. This is as true of the genus Siphona as any other. There appears to be a predisposition for expression of wing setulae on R, among these genera, so species sharing this trait are not necessarily closely related. Among siphonines setulae on R] are expressed in one of three ways: (1) vein setulose along its length, (2) limited to a row along the bend, or (3) represented by one or two setulae near the bend. Most non -Siphona species with R, setulose belong to the first two categories. Siphona bevisi Curran and S. setinerva Mesnil (both African) are in the first category, other examined Siphona species belong to the third category - S. lutea, S. oligomyia, S. tropica and S. grandistyla Pandelle (the last from the Pyrenees), a - setulae absent. A - one or two setulae near bend of R1. Character 8. Wing setulae on R4+5 Wing setulae on R4+5 between bifurcation of R2+3 and R4+5 and crossvein r-m is characteristic of the Siphonini, and represents the plesiotypic state in Siphona. As with R, however, the apotypic state in Siphona has been independently derived in other genera and presumably within Siphona as well. Of the species under revision only S. lutea has R4+5 setulose beyond r-m. Other Siphona species with this condition (of species examined) are S. bevisi Curran, S. setinerva (Mesnil) and S. unispina (Mesnil) (all from Africa). The African species resemble one another and further analysis may indicate they belong in the same group. They do not share characteristics other than wing setulae with S. lutea , so are probably not closely related to it. a - vein R4+5 setulose between bifurcation of R2+3 and R4+5 and crossvein r-m. A - vein R4+5 setulose beyond r-m. Character 9. Male tarsal claws The groundplan for Siphona is small tarsal claws. Large tarsal claws of males are present in very few species of other Siphona group genera (eg. state A1 in Asiphona brunnescens (Vill.)). These observations suggest the apotypic states, especially A1, are easily acquired and seldom indicative of a monophyletic origin. North American species of Siphona Meigen 319 a - average-sized, length of claw subequal to width of tarsomere 5 (Figs. 4,6). A1 - intermediate in size between a and A2. A2 - large claw, length subequal to length of tarsomere 5 (Fig. 5). Character 10. Median marginal setae on T1+2 I am unaware of any Siphona group species not belonging to Siphona that possess a pair of median marginals on T1+2. Nevertheless, consideration of more distantly related tachinids reveals that median marginals on T1+2 are widespread, indicating their absence from most members of the Siphona group almost certainly results from suppression of this attribute. This may explain why a very few specimens of S. cristata and S. multifaria have median marginals on T1+2, and why a few specimens of other species have slight development of a median marginal hair on T1+2 such that it is distinctly stronger than others to either side. a - absent. A1 - a significant proportion of specimens with median marginals on T]+2 (here interpreted as a regression from A2). A2 - virtually all specimens with one pair median marginal setae on T, +2. Character 1 1. Lateral marginal setae on T1+2 Most species of the Siphona group other than those belonging to Siphona have weak lateral marginal setae on T1+2, although more distantly related tachinids in most species have strong setae in this position. As with median marginals on T1+2, weak lateral marginals probably result from suppression. In Siphona weak lateral marginals on T1+2 are almost exclusive to species of the S. maculata group. Since this state is widespread in the out-group, it is interpreted as symplesiotypic in the S. maculata group (also see discussion under character 3). a - setae absent or one pair weakly developed. A - one pair strong setae. B - two pair strong lateral marginal setae on T1+2 in significant number of specimens. Character 12. Male sternum 5 Of male specimens examined, only two unrelated Siphona species, S. oligomyia and S. cuthbertsoni Curran, possess a sternum 5 in which the antero-lateral arms are hooked inward. This condition is characteristic of Siphonopsis species, which form either the sister group to Siphona or branch from the Ceranthia lineage. Be that as it may, there is no other indication that the hooked state of sternum 5 is plesiotypic in Siphona. In fact, the most parsimonious explanation is independent evolution of the hooked state among Siphona species rather than its loss in all but very few. a - antero-lateral margins smoothly rounded (Fig. 10). A - antero-lateral margins hooked inward (Fig. 1 1). Character 13. Antero-ventral margin of distiphallus By far the most prevalent condition of the distiphallus by out-group and in-group comparisons is a dentate antero-ventral margin. This is presumed plesiotypic. The difference Quaest. Ent., 1982, 18 (1-4) 320 O’Hara between the dentate state and one in which small hooks are present (A1) is slight, hence is more likely to occur in unrelated taxa than are states A2 and A3. a - dentate. A1 - small hooks (Figs. 75-77). A2 - hooks medium in size (Fig. 60). A3 - large hooks, margin U-shaped in anterior view (Fig. 62). Character 14. Shape of distiphallus The plesiotypic shape of the distiphallus in Siphona is unknown. I have chosen to regard all shapes without apparent apotypic features as plesiotypic. This conservative approach should ensure that species with the same apotypic shape have a high probability of sharing a uniquely derived state. A transformation series could not be discerned from distiphallus shapes exhibited, so each apotypic state is treated independently. a - without recognizable apotypic shape (ie. not necessarily plesiotypic, but treated as such). A - subtriangular, apex sharply pointed in profile (Figs. 60,62). B - anterior margin of distiphallus slightly indented in profile about one third of its length from apex (Figs. 66-68). C - anterior and posterior surfaces of distiphallus nearly parallel in profile, apex truncate (Figs. 70-71). D - narrow in profile, sloped postero-ventrally (Figs. 9, 73). E - narrow in profile, apically truncate (Fig. 74). F - broad and short in profile (Fig. 76). G - narrow in profile, subtriangular, apex pointed (Figs. 77-78, 80). Character 15. Surstylus length The most prevalent condition of the surstylus and one least in agreement with other character states is presumed plesiotypic. a - apex of surstylus even with or extended slightly beyond apex of cerci. A - apex of surstylus extended beyond apex of cerci by more than preapical width of surstylus. Reconstructed phylogeny Introduction. - In reconstructing the phytogenies depicted in Figs. 103-107, I have not employed a character weighting system. I recognize its advantages when applied to certain types of data, as lucidly demonstrated by Hecht and Edwards (1976, 1977) for example, but such a system is not appropriate to all situations. With respect to Siphona, the characters analyzed are inadequate to permit elucidation of relationships among most species groups, so where character weighting would have been most useful it cannot be applied. Furthermore, with relatively few characters of broad import, the most parsimonious reconstructed phytogeny for each species group has few alternatives. I have depicted only one phylogenetic scheme for each species group, but I discuss problem species placements below, and consider the strengths and weaknesses of the character states involved. Under these circumstances I consider the utility of a character weighting system negated. North American species of Siphona Meigen 321 Figs. 103-107 illustrate inferred relationships among North American Siphona species. Exclusion of Palearctic species from these diagrams is not indicative of more distant relationships between regions, for indeed the Siphona faunas of the Nearctic and Palearctic Regions are intimately associated. My analysis of Palearctic Siphona species has not been sufficiently detailed to warrant species placements beyond group. In general, Siphona species from other regions have been considered to an even lesser extent. The approach I have taken is thus: 1) inferred relationships among North American species are discussed under each species group; 2) alternative phylogenies are considered; 3) the 1 1 described Palearctic Siphona species6 (excluding Holarctic species and S. seyrigi Mesnil of the Canary Islands) are used to test my hypotheses concerning delimitation of species groups; and 4) Siphona species from the Neotropical and Afrotropical Regions are considered where their character states are relevant to the discussion. The S. maculata group. - The ancestor to extant members of the S. maculata group is inferred to have had a narrow, pointed first flagellomere, short proboscis and weak lateral marginal setae on T,+2 (Fig. 103). The latter two states are interpreted as symplesiotypies (as discussed above under polarization of states of proboscis length (character 3)), and identify the S. maculata group as sister group to the other species groups. Except for the apotypic shape of first flagellomere upon which the S. maculata group is based (Figs. 18-22), there are only two synapotypies below the level of species pair. Therefore relationships within this group are not well understood. The only resolution is among S. maculata, S. intrudens and S. lurida. The latter two species share four apotypies, clearly attesting to their close phylogenetic relationship. There are a number of differences between S. intrudens and S. lurida , mostly because of the highly derived condition of the latter. S. maculata is interpreted as closely related to the S. intrudens - S. lurida lineage, because of complex modifications of the distiphallus shared between S. maculata and S. intrudens (characters 13 and 14 (Fig. 102), Figs. 60,62). Since S. lurida lacks these modifications (Fig. 63), the plesiotypic states of its distiphallus evidently represent losses (thus the coding of 13-a and 14-a as derived in S. lurida in Fig. 103). Distribution of character states are inadequate to resolve the trichotomy depicted for lineages of the S. maculata group (Fig. 103). The autapotypic states of S. pacifica result in its general similarity to plesiotypic members of other Siphona groups. However, there is no evidence to question placement of S. pacifica in the S. maculata group. The phylogenetic position of S. floridensis within the S. maculata group is also enigmatic. This species has the fewest recognized apotypic states of species analyzed. Therefore, the overall similarity between S. maculata and S. floridensis in external characteristics is the result of symplesiotypies, which are unanalyzable cladistically. Also, to consider 5. maculata and S. floridensis as sister species, one must regard the plesiotypic states of the distiphallus in the latter (Fig. 61, cf. Fig. 60) as losses (as inferred for S. lurida ). Nonetheless, a close relationship between these species is tenable on zoogeographic grounds. A zoogeographic The new, as yet unpublished, Siphona species of Andersen (in press “a”) were not examined. Quaest. Ent., 1982, 18 (1-4) 322 O’Hara scenerio whereby S. floridensis might have evolved from a S. maculata population during the Quaternary is outlined in the “Zoogeographic” section. Pending further morphological evidence, I leave the trichotomy in Fig. 103 unresolved. Two Palearctic species have been identified as members of the S', maculata group: S. collini Mesnil and S. efflatouni Mesnil. They are the only Palearctic Siphona species with weak lateral marginal setae on T1+2. They possess an A1 type first flagellomere and are the only Palearctic species with a short proboscis (not longer than A2). The distiphallus of S. collini has small hooks along the antero-ventral margin (state A1). (Only the female holotype of S. efflatouni was examined, so the distiphallus of this species was not studied.) In general habitus S. collini and S. efflatouni also appear to belong to the S. maculata group. Despite the plesiotypic nature of weak lateral marginal setae on T1+2 and a short proboscis, both states are well correlated with the apotypic shape of first flagellomere in the S. maculata group. As such these characteristics are useful indicators of S. maculata group members, even though they are interpreted as symplesiotypies. A test I have not had an opportunity to perform to date involves use of undescribed Oriental Siphona species H. Shima notes (per. comm.) several species similar in appearance to S. lurida from the Orient, and study of these would undoubtedly cast light on the phylogeny presented here. It will be particularly interesting to note whether these species are more closely related to S. lurida or S. intrudens, since the implications critically affect the zoogeographic interpretation as well as the reconstructed phylogeny of the S. maculata group. The S. cristata group. - The S. cristata group is based on one apotypic state, shape of first flagellomere (Fig. 104). S. oligomyia is included in this group, though its first flagellomere is not characteristic of the group (possessing state 2- A instead of 2-B). Justification for inclusion of S. oligomyia in the S. cristata group is based on two synapotypies between S’, lutea and S. oligomyia , as discussed below. At the next higher level on the cladogram (Fig. 104) is shape of distiphallus (state 14-B), uniting the S. cristata - S. multifaria and S. lutea - S. oligomyia lineages. Again, S. oligomyia does not share the derived state of the other members. S. pisinnia is treated as sister species to the other species of the S. cristata group, though it is actually not as distantly removed phenetically from the rest of the group as it appears in the cladogram. Only three states separate it from S. multifaria , and one of these, distiphallus shape (Fig. 65; cf. Fig. 67), is a consequence of the system I employed to delimit states. I have recognized the indentation of the anterior margin of the distiphallus as an apotypic state, while necessarily relegating forms near this shape without an indentation to the plesiotypic state, for lack of distinguishing features. Because of the highly derived appearance of S’, lutea and S. oligomyia , S. pisinnia is phenetically closer to S. multifaria and some specimens of S. cristata than are the others. Within the S. cristata group I have depicted S. multifaria and S. cristata as sister species (Fig. 104). Under this scheme I have not followed the apparent transformation series of character 6 (colouration), which suggests S. cristata is closer to S. lutea than to S. multifaria. The distiphalli of S. cristata and S. lutea are also very close (Figs. 66, 68). Nevertheless, characters 3 and 7 argue in favour of the depicted arrangement. Additional support for the reconstruction comes from the Palearctic species S. flavifrons Staeger. It is similar in colouration to the yellow form of S. cristata , yet probably belongs to the S. geniculata group. This is evidence that the B1 colouration of S. cristata need not have shared North American species of Siphona Meigen 323 an ancestry with B2 colouration of S. lutea and S. oligomyia. Character state 6-B2 is highly apotypic. Through concordance with state 7-A, S. lutea and S. oligomyia are well established as sister species. S. oligomyia is distinctive because of many autapotypic features; had it been of average colouration its species group placement would be uncertain. A Palearctic species of dubious placement is S', setosa Mesnil. It has median marginal setae on T1+2> but none of the other S. geniculata group apotypic characteristics. With its 2-B first flagellomere, it might belong to the S. cristata group. Other characteristics are inconclusive: distiphallus type 14-a, proboscis and eye average-sized, colouration average, etc. I leave S. setosa unplaced. Two Palearctic species are easily recognized as members of the S. cristata group. One, S. confusa Mesnil, has an average-sized eye and proboscis and a first flagellomere varied between states A1 and B1. The one distiphallus examined is close to state 14-B, but without an indentation anteriorly. A few specimens have a tawny abdomen as in S. cristata , while others are of average colouration. The other Palearctic species, S. pauciseta Rondani, has an average-sized to A1 eye and a proboscis of average length. First flagellomere and distiphallus possess the characteristics of the group, with states 2-B1 and 14-B. S. confusa and S. pauciseta do not share the apotypic states of S. lutea and S. oligomyia. Their phylogenetic position relative to S. pisinnia , S. multifaria and S. cristata is unclear. As members of the S. cristata group, S. confusa and S. oligomyia illustrate a deficiency in the distinction between states A and B of first flagellomere. These states are not markedly different, and both species indicate that 2-A is derivable from 2-B. Unfortunately, without additional characters there is no alternative to 2-B1 as a synapotypy for the group. S. confusa also illustrates a problem in delimitation of character 14; distiphallus shape. S. pauciseta supports recognition of 14-B as a good synapotypic state, yet S. confusa , like S. pisinnia , lacks this state. To incorporate S. confusa and S. pisinnia , S. medialis, S. floridensis and S. pacifica would also have to be included, thereby considerably reducing the usefulness of this uniquely derived state. A subjective element thus enters the analysis, for resemblance of a type 14-a to a 14-B distiphallus is not without significance. Regardless of the difficulties inherent in recognition of state 14-B, it is possible that relegation of S. pisinnia to its depicted position is phylogenetically inaccurate. An alternative is to disregard the slight distiphallus differences beween S. pisinnia and its relatives, and examine the consequences to the cladogram. A trichotomy results, with the S. pisinnia lineage on the same node as the S. cristata - S. multifaria and S. lutea - S. oligomyia lineages. Given this possibility, similarity between S. pisinnia and S. multifaria suggests 51. pisinnia might be more closely related to the S. cristata - S. multifaria lineage than to the other. To support this alternative hypothesis (and refute the relationship shown in Fig. 104) a synapotypy is required, and one has not yet been found. The S. cristata group would benefit more than others from analysis of additional characters. Possibilities include larvae, ethological and host data, differences in the reproductive system of females, electrophoresis and cytotaxonomic data. As illustrated above, external features and male genitalia are not particularly amenable to phylogenetic analysis. The S. geniculata group. - In North America the S. geniculata group includes three species (Fig. 105), one of which ( S . geniculata ), has been introduced by man to this continent. These species share the derived states of one pair median marginal setae on T1 + 2 (albeit not all S. Quaest. Ent., 1982, 18 (1-4) 324 O’Hara hokkaidensis specimens) and a short aristomere 3 (Figs. 28-32). The apotypic states shared between S. geniculata and S. hokkaidensis attest to their close phylogenetic relationship, even though most specimens of the latter are without either median marginals on T1+2 or a type 2-A1 first flagellomere (= antennal type 1, Fig. 30). Five Palearctic species are assignable to the S. geniculata group, though their characteristics are varied. Only two species have median marginal setae on T1+2, S. flavifrons Staeger and S. rossica Mesnil. S. rossica is evidently close to S. hokkaidensis , with its medium-sized eye and proboscis, first flagellomere near type 2-D, two pair lateral marginal setae on T3 and a short aristomere 3. I did not have an opportunity to dissect a male specimen, but I predict the male genitalia of this species has a long surstylus. I recognize S. flavifrons as a member of the S. geniculata group by its joint possession of median marginals on T,+2 (most specimens) and a long surstylus. Its distiphallus is apically truncate, but slightly too wide basally to be classed type 14-C. S. flavifrons does not have a shortened aristomere 3 and its first flagellomere and colouration are as in S. cristata. In light of other character state differences I view these similarities as convergences. The characteristic shape of first flagellomere for the S. cristata group has already been shown to be weakly based. I regard the average-sized aristomere 3 as a significant departure from the condition characteristic of the S. geniculata group, for to my knowledge S. flavifrons is the only species with median marginals on T1+2 and a long surstylus for which the aristomere 3 is not shortened. S. paludosa Mesnil lacks median marginals on T1+2, but has a short aristomere 3, type 14-B distiphallus and a 15-A surstylus, strongly supporting its placement in the S. geniculata group. I also include two other Palearctic species without median marginals on T1+2, S. grandistyla Pandelle and S. nigricans (Villeneuve). Both have a first flagellomere resembling the 2-D state of S. hokkaidensis , and a short aristomere 3. Unfortunately I did not have an opportunity to dissect male specimens of these, which would have provided a good test of their placement. The only Palearctic species with median marginal setae on T1+2 that I exclude from the S. geniculata group is S. setosa Mesnil. It is discussed under the S. cristata group. I have examined specimens from Ecuador of three undescribed Siphona species with median marginal setae on T1+2. In other respects these species are quite unlike members of the S. geniculata group (in particular, they lack states 4-A and 15-A), so I exclude them. Discussion of these species and their affinities more appropriately awaits a revision of South American Siphona species. I mention them here to demonstrate that not all species with median marginals on T1+2 belong to the S. geniculata group. Examined Afrotropical and Oriental Siphona species do not have median marginals on T1+2 (see world list for species examined). Despite lack of unequivocal character states for the S. geniculata group, concordance among states functions well for distinguishing between included and excluded species. Consideration of Palearctic species illustrates that the S. geniculata group is a complex assemblage, yet evidently monophyletic. The S. futilis group. - The apotypic character state for this group is a large eye, best exemplified by the 1-A2 state in S. futilis and S. illinoiensis (Figs. 1,34). Similarity between S. futilis and S. illinoiensis is such that I can polarize but one difference, length of first flagellomere. North American species of Siphona Meigen 325 I cannot with certainty place S. brunnea in the S'. futilis group. In general habitus it resembles S. futilis and S. illinoiensis despite its autapotypies, so I include S. brunnea in this group until such time as its relationship is clarified (Fig. 107). With S. brunnea a provisional member, and only one apotypic (and not unique) state for the group, consideration must be given to states of characters 1, 2 and 14 in placement of non-North American species (Fig. 102). The Palearctic species S. boreata Mesnil might belong to the S. futilis group. It has a large eye, average length proboscis (characteristic of the group, but evidently plesiotypic), and average colouration. However, first flagellomere is near type 2-F. I did not have an opportunity to study male genitalia of this species, thereby limiting my analysis. Examined specimens of at least two (possibly undescribed) South American species probably belong to the S. futilis group, study of which might help define the group and clarify the position of S', brunnea. The S. tropica group. - This group is particularly noteworthy because it is well represented in the Afrotropics. Of Afrotropical species examined, S. unispina (Mesnil) and S. gracilis (Mesnil) are the most easily recognized as members. I hesitate to assign to this group other Afrotropical species, since the characteristics that readily distinguish members in Middle America are widespread among Afrotropical Siphona species and not restricted to the S. tropica group. I have used character states of S. unispina to help polarize states within the S. tropica group. I have assumed the Middle American species to be more closely related among themselves than to S. unispina. As a result, states of characters 13 and 14 have been polarized differently than they otherwise would have been (Fig. 106). In conflict with my understanding of the S. tropica group is character state 8- A: setulae on R,. A single setula is present near bend of R, in S. tropica and S. unispina. Because it is absent from examined specimens of S. rizaba, S. longissima and S. akidnomyia , I suggest it has been independently derived as in other Siphona species ( S . lutea + S. oligomyia , S. grandistyla). S. tropica could be postulated as closer to S. unispina than to the others, but this would not explain the presence of hooks on the distiphallus of S. longissima and S. akidnomyia (Figs. 76-77), nor their other S. tropica group characteristics. Two separate ancestors could be invoked to explain the anomaly, but are not justified on the basis of a single low weight apotypy. The shared characteristic between S. longissima and S. akidnomyia is a weak synapotypy. Additional data are needed to elucidate relationships among S. tropica group members. The S. tropica group is apparently absent from the Nearctic and Palearctic Regions. In the Neotropics it is most diverse in Middle America. Other than S. tropica , I have examined one species (probably undescribed) of this group from South America (Chile) and one from the Philippines. The Siphona fauna of Middle America is inadequately known. I have examined a single specimen of a new species belonging to the S. tropica group that I have not described due to its poor condition. I suspect several other Siphona species are undiscovered or undescribed from this region as well. The S. macronyx group. - S. macronyx is placed in a monobasic species group because its phylogenetic position relative to other North American Siphona species has not been clarified (Fig. 103). Quaest. Ent., 1982, 18 (1-4) 326 O’Hara The large tarsal claws and above average number of setae and setulae on vibrissal angle of S. macronyx are derived states shared with S. lurida of the S. maculata group. This species also has hooks on the distiphallus, a derived state for the S. maculata, S. floridensis, S. intrudens and S', lurida lineage. Nevertheless, a sister species relationship between S. macronyx and S. lurida is untenable in light of their overall dissimilarity (Figs. 21,33) and the synapotypies between S. intrudens and S. lurida (notably a small eye, short proboscis and similar colouration). Furthermore, S. macronyx lacks the slender, pointed first flagellomere, which is apotypic for the S. maculata group. There is a slight similarity between S. macronyx and the S. geniculata group (characters 2,3 and 9 (Fig. 102)), but S. macronyx lacks two of three apotypic states upon which the group is based. Evolutionary trends The most satisfactory method for obtaining information about most characters of varied structure in Siphona adults is to observe living specimens under natural conditions. Though I did this intermittently, I did not pursue it intensely. It has been my experience that adults of Siphona searching for a mate or host are usually seen fleetingly, and in many instances rarely. On the other hand, adults seeking nourishment can be observed for some time as they probe deeply into flowers with their long proboscises, in quest of nectar. As a result, the functional significance of most derived characteristics can only be gleaned indirectly through study of dead specimens and by analogy with other taxa. It is a basic tenet in evolutionary studies that most modifications in structure result from selective pressures and have functional significance. If so, the most derived and varied feature of Siphona adults, the proboscis, must have attained its present variety because of advantages conferred on its bearers. The only known function of the proboscis in Siphona adults is for procurement of food; chiefly nectar from flowers. Unlike large tachinines, adults of most siphonines are too small to force apart flower components to reach deep nectaries, so the benefits of a long proboscis are obvious. Not only did adults of Siphona apparently evolve an elongate prementum and labella to aid feeding, but so too did others, such as adults of certain species of Ceranthia s. lat. and Actia. Adults of Clausicella achieved essentially the same end through elongation of the prementum, albeit with less maneuverability distally than that provided by a geniculate proboscis. In fact, an elongate prementum is found in various groups throughout the Tachinidae, though an elongate labella is apparently unique to the Siphonini. It is not possible to correlate development of a geniculate proboscis in adult Siphona with success of the taxon. For one reason, certain genera like Ceromya, Peribaea and most species of Actia have a short labellar disc, yet approach the diversity of Siphona. For another, too little is known about parasite-host interactions to dismiss as minor, developments in that area. However, it is possible that individuals with a long proboscis have a selective advantage over others in varied environments, for they are physically less dependent upon certain kinds of flowers than are their relatives with shorter proboscises. Few Siphona species other than members of the primitive S. maculata group have a short proboscis, so once attained a long proboscis seems seldom to have shortened (I have inferred just two such reductions; in S. oligomyia and S. akidnomyia). Therefore a definite trend in adults of Siphona species from a short to long proboscis is evident (cf. Fig. 15). North American species of Siphona Meigen 327 There are no other trends in Siphona as unidirectional as that of proboscis length. Greatly varied is eye size, but from the inferred plesiotypic state of an average-sized eye, both small and large eyes are necessarily derived. Shape and size of first flagellomere are also considerably varied (even within species), and a trend is not apparent. The significance of varied eye and first flagellomere sizes among species is unknown, and speculation is best deferred until more information on behaviour and environment of individual species is available. Within species, males have the larger eye and first flagellomere, implying both are used in the search for a mate. Females are probably mostly host oriented, and the slightly longer palpus of females in most species might be involved in determining host suitability. Notable in the evolution of Siphona species is the similarity among species groups in mean number of apotypic states (Fig. 102). Though the S. maculata group is interpreted as the most primitive, two included species ( S . intrudens and S. lurida) are very derived, so even this group is not markedly removed from the others in degree of apotypy. This pattern implies that evolution within Siphona has progressed at a relatively constant rate, without invasion of, and diversification in, new adaptive zones. ZOOGEOGRAPHIC CONSIDERATIONS Diversity and general paterns I prefer not to use the terms Nearctic and Neotropical in discussing New World Siphona species distributions. Most Middle American Siphona species are not derived from extant Nearctic or South American species groups, so to emphasize this point I. discuss distribution patterns in terms of America north of Mexico, Middle America (Mexico to Panama) and South America. Only two species are not confined within boundaries of one of these areas. S. pisinnia (Fig. 83) ranges widely in America north of Mexico and Middle America. The distribution of S. tropica (Fig. 101) extends into extreme northwestern South America, but for present purposes it will be regarded as a Middle American species. My use of terms Palearctic, Oriental and Afrotropical were discussed in the preamble to “World list”. I treat S. geniculata as a Palearctic species in the ensuing discussion because in North America it is an introduced species. The 79 described Siphona species7 are distributed as follows, by region in order of decreasing diversity: Afrotropical - 40 species. Palearctic - 17 species. America north of Mexico - 14 species. Middle America - 7 species. South America - 3 species. Oriental - 3 species. Madagascar is included in the Afrotropical Region. It has three endemic species and five others that are shared with the mainland. The Canary Islands off the northwest coast of Africa, with endemic species S. seyrigi , is considered Palearctic. So too is Africa north of the Sahara, from whence S. nigricans was described (from Algeria). Excluding S. maculipennis Meigen, an unrecognized Palearctic species that is probably conspecific with a recognized species. Quaest. Ent., 1982, 18 (1-4) 328 O’Hara Four species (excluding introduced species S. geniculata ) are known to be shared between America north of Mexico and the Palearctic Region: S. cristata, S. hokkaidensis, S. lurida and S. maculata. The Palearctic Region is not known to share species with any other region. America north of Mexico has S. pisinnia in common with Middle America. The Siphona fauna of South America is inadequately known. I estimate from the little material I have examined from the region that at least ten species live there, possibly more. I have examined adults of one undescribed species of Siphona from Australia. Siphona species from the Oriental Region are not well known and are probably more diverse than indicated by three described species. In the previous section a cladistic analysis helped define six North American Siphona species groups. Within each group there is a general pattern of geographic distribution involving at least two regions. Three general interregional patterns are recognizable among the groups: 1) America north of Mexico - Palearctic Region. This pattern is evidenced by a close relationship between species of both regions, with a high proportion of Holarctic distributions for species with northern records (three of six New World species). Representing this pattern are the S. maculata, S. cristata and S. geniculata groups. 2) America north of Mexico - Middle America. Here belongs the S', futilis group, wherein closely related S. illinoiensis and S. futilis are in adjacent regions. There is evidently a closer affinity to the fauna of South America than the Old World, but this has not been firmly established. 3) Middle America - Afrotropical Region. In the New World the S. tropica group is most diverse in Middle America. No species has been assigned to the group from America north of Mexico or the Palearctic Region. Although represented by at least one species in South America and one in the Philippines, the zoogeographically most significant relationship is between Middle American and African members of the group. Within North America another set of distribution patterns, those involving species rather than species groups, is evident. To a large extent these reflect biotic changes that have taken place in the region since the Middle Tertiary. These are: 1) Transcontinental (mostly cool to cold temperate) - S. cristata, S. hokkaidensis, S. intrudens, S. lutea, S. maculata, S. medialis and S. multifaria. 2) Western - S. geniculata (introduced), S. lurida, S. macronyx, S. oligomyia, S. pacifica and S. pisinnia. 3) Eastern - S. floridensis and S. illinoiensis. 4) Mexican - S. akidnomyia, S. brunnea, S. futilis, S. longissima, S. rizaba and S. tropica. Historical zoogeography It is premature to speculate in detail about historical events responsible for present day Siphona species distributions in North America. The three main distribution patterns identified in the previous section make obvious the fact that the North American assemblage of Siphona species is polyphyletic. Indeed, interregional relationships are so complex that the monobasic S. macronyx group is the only one that is exclusively North American. The best zoogeographic hypothesis is one that stems from a thorough cladistic analysis of a group. Only then are all requirements of a scientific hypothesis met: explanatory power, internal consistency, predictive power and testability (falsification) (Ball, 1976; Noonan, North American species of Siphona Meigen 329 1979). As a consequence the cladograms in Figs. 103-107 (representing only North American species) are inadequate for detailed interpretation of the historical zoogeography of North American Siphona species. Siphona species groups, on the other hand, are suitable for zoogeographic analysis because they are based on inclusive and not (as for North American species) exclusive data. Siphona species groups are monophyletic units, members of which need not be cladistically arranged for the groups to be zoogeographically analyzed, provided regional group relationships are known (Hennig, 1966b). For instance, the very close phylogenetic relationships among taxa of the S. maculata group between America north of Mexico and the Palearctic Region are indicative of past faunal exchange between regions. This presumed interchange satisfies the four requirements of a scientific hypothesis, so a zoogeographic analysis seeking a historical explanation for the interchange is well founded. Ideally all Siphona species groups and data about intergroup affinities should be included in a zoogeographic treatment of the genus. It may be many years before sufficient information on Siphona cladistics is gathered to realize this goal. Therefore I judge it timely to interpret aspects of Siphona history from the data available now. In the absence of a fossil record there is an alternative means for deriving age of a taxon. The minimum age is that of the earliest demonstrable vicariant event within the taxon. Maximum age is that already determined for a higher taxon to which the taxon in question belongs, or as evidenced by the taxon’s global distribution pattern (provided the taxon is widespread). For the Calyptratae in general the early fossil record is inadequate; they have been assumed pre-Tertiary in age for many years (eg. Rohdendorf, 1974), but not until 1970 was the first calyptrate fly from the Mesozoic (Upper Cretaceous, 70 mybp') recorded (McAlpine, 1970). To date, a fossil tachinid has not been discovered that has a significant bearing on the age of origin of this family (van Emden, 1959), much less Siphona. It has become possible in the past decade to outline in a broad sense major geologic and climatic events of the earth since early Mesozoic time. Concomitant with this knowledge has developed a general willingness on the part of biogeographers to interpret species distributions according to established geologic and climatic theory. Fossils remain valuable means of hypothesis testing, but are no longer central to most biogeographical analyses unless the fossil record is unusually complete. Recent studies of Diptera reflect this trend. For example, Bowden (1973) suggests a Late Cretaceous origin of Dischistus (Bombyliidae) based on distributional data. Papavero (1977), places origin of the Oestridae between Late Jurassic and Early Cretaceous on the basis of a Gondwanian distribution. Quite remarkable is McAlpine’s (1977) suggestion of a Gondwanian distribution for an extant piophilid genus, Protopiophila (the first Gondwanian distribution recognized for a genus of the Schizophora). By analogy with studies on other Schizophora the Tachinidae are apparently pre-Tertiary in origin. Provided distributional data suggest a more recent origin of Siphona than analogy has for the Tachinidae, then there is no age conflict between them9. Although Siphona species are cosmopolitan, they show no indication of a Late Cretaceous or earlier origin. Had Siphona evolved in the Southern Hemisphere during the Late Cretaceous 8million years before present. 9as yet a time scale has not been proposed for origin of tachinid subfamilies and tribes, so these taxa cannot be discussed here. Quaest. Ent., 1982, 18 (1-4) 330 O’Hara a Gondwanaland distribution would be expected (assuming extinction was not a factor). By Late Cretaceous Africa was drifting north, while South America, Antarctica and Australia were still connected via at least an archipelagic route (Dietz and Holden, 1970; Smith, Briden and Drewry, 1973; Howarth, 1981). In fact, the southern route through Antarctica may have been in use by terrestrial animals as late as Early Oligocene (Raven and Axelrod, 1975). An adequately supported argument against a Gondwanaland distribution for Siphona must await a cladistic analysis of African, South American and Australian species. However, the few Siphona species in South America and especially Australia, relative to Africa and the northern continents, suggests dispersal of Siphona species was not taking place between southern continents in the Late Cretaceous. In the Northern Hemisphere during the Late Cretaceous, high sea levels had a greater isolating effect on continents than did plate positions. Across the North Atlantic and through Beringia, land (?filter) bridges were apparently in operation, though north-south seaways hindered intracontinental faunal movements (Kurten, 1969; Hallam, 1973; Matthews, 1979; Howarth, 1981). If Siphona species were in the Northern Hemisphere during the Late Cretaceous I would expect a more pronounced difference between Siphona faunas of America north of Mexico and the Palearctic Region than evident in the S. maculata, S. cristata and S. geniculata groups. Presumably Siphona species could have dispersed readily between northern regions had they been there during the Late Cretaceous. Yet present faunal similarities are better explained by Late Cenozoic exchange. Based on available evidence and without comparable studies within the Tachinidae, I hypothesize a maximum age of Siphona as Late Cretaceous. As mentioned earlier, minimum age necessarily precedes the earliest demonstrable vicariant event. Before discussing how I apply this principle to Siphona , it is appropriate to consider region of origin of the genus. This simplifies the discussion further on. I emphasize the term region of origin, not center of origin. Thus I am not concerned with the controversy between centers of origin in the Darwinian sense and vicariance biogeography (Croizat et al., 1974). In a region of continental size, taxa can evolve under either paradigm and produce equal diversity, so the distinction is not important here. Siphona probably arose in the Old World. The basis for this inference is twofold. First, Old World Siphona species and species of siphonines related to Siphona ( Actia , Ceromya and Peribaea spp.) considerably outnumber New World taxa. Secondly, an invasion of North America from the Old World during the Tertiary by Siphona is compatible with New World species distributions, while it is inconceivable without a number of assumptions that invasion of Siphona into the Palearctic Region from North America would lead to the present patterns of diversity and distribution in the Old World. To account most parsimoniously for Old World Siphona patterns, Africa must be interpreted as region of origin for the genus. Diversity favours this choice, but does not negate the possibility of adaptive radiation in Africa with an origin elsewhere. More important than diversity is the pattern of diversity, as determined through comparison of species numbers among regions. An African origin of Siphona is compatible with the observed decrease in number of species from a high in the Afrotropical Region to a low in the Oriental and Australian Regions. If the Palearctic Region were postulated as area of origin of Siphona , the observed pattern of diversity would be unparsimonious. During most of the Tertiary, Africa was isolated from North American species of Siphona Meigen 331 other regions by the Tethys Sea (Cooke, 1972; Hallam, 1973; Azzaroli and Guazzone, 1979; Howarth, 1981). It is unlikely Palearctic Siphona individuals crossed this barrier enough times to attain the present Afrotropical diversity in number of species and species groups without similarly invading the more accessible Oriental Region. Returning to the question of minimum age of Siphona , I limit my analysis to species groups in which cladistic relationships between regions have been determined. These are the S. maculata, S. cristata, S. geniculata and S. tropica groups. The first three do not reflect a necessarily old distribution pattern. The S. tropica group is disjunct between Middle America and Africa, and it is from this group I hypothesize minimum age of Siphona. Concomitant with historical biogeographic hypotheses are assumptions. Assumptions should be formalized, for they enhance the central hypothesis by clearly distinguishing the assumed from the inferred (cf. Lewis, 1980: 554). Furthermore, while some assumptions are untestable (and are, for that reason, assumptions) others result from insufficient data and become falsifiable as new data are added. Assumptions of the latter type should be accompanied by statements about their testability, to make the central hypothesis more scientific. Assumptions upon which I base my analysis of the disjunction of the S. tropica group between Middle America and Africa are: 1) The cladistic relationship between Middle American members of the S. tropica group and S. unispina in Africa (Zaire) has been correctly interpreted. (Falsifiable by cladistic study of additional taxa or reinterpretation of existing data.) 2) Members of the S. tropica group in South America are derived from Middle American species; they are not Gondwanaland relicts. (Relationships testable by cladistic analysis.) 3) Africa was region of origin of Siphona and the S. tropica group. (Subject to corroboration or refutation through study of world species groups of Siphona and closely related genera.) 4) Disjunction in distribution of the S. tropica group is not a result of long distance dispersal between Middle America and Africa. The dispersal ability of Siphona adults is attested to by their presence on islands: Madagascar, Canaries, Juan Fernandez, Tierra del Fuego, Philippines and others. There are eight species recorded from Madagascar, of which five are shared with Africa. Presumably the Mozambique Channel (500km wide) has not been an efficient barrier to dispersal of Siphona individuals. There is no evidence, however, that individuals cross wide barriers. For example, Siphona is not recorded from Hawaii. I suspect water gaps are more easily crossed than other physical or climatic barriers. 5) Current hypotheses of paleoclimate and paleogeography relevant to my interpretation of past events are accurate. (Testable by results of future research.) 6) Climate and habitat requirements of S. tropica group taxa have not changed significantly throughout their existence. (Discovery of a S. tropica group species in a cold temperate environment would cast doubt on this assumption.) An explanation is sought that accounts for the geographically disjunct nature of the S. tropica group. Assuming the genus originated in Africa subsequent to the Late Cretaceous, direct dispersal across the Atlantic is considered improbable because South America and Africa were widely separated by that time. The only available land route was northward to Europe or Asia and into North America through Beringia or across a North Atlantic land or filter bridge. First consider the ecological aspects involved in a Northern Hemisphere route. The S. tropica group is now restricted to the Tropics, so existence of northern land bridges is not sufficient to account for Siphona migration into North America. Had a S. tropica group taxon Quaest. Ent., 1982, 18 (1-4) 332 O’Hara invaded the Northern Hemisphere from Africa during a relatively cool period, descendants would presumably still be there. If instead there was a period of significant warming in the north, a S. tropica group taxon could have entered Eurasia and spread to North America without a major shift in way of life. If deterioration of the warm trend took place soon after the ancestral New World Siphona taxon arrived in North America,' then retreat to Middle America could have occurred without leaving northern, cool adapted descendants. These events could not have occurred later than the Late Eocene (34 mybp) (Wolfe, 1978, 1980). For many years the Geoflora10 concept dominated paleobotanical interpretations. Paleocene and Eocene climates of northern North America were interpreted as subtropical to warm temperate with gradual cooling from the Oligocene to Quaternary (Dorf, 1964). The Geoflora concept has recently come under incisive criticism with the realization that plant species respond individualistically to climatic shifts (Graham, 1972; Wolfe, 1978). Nowhere is this better exemplified than in late Quaternary macrofossil records of packrat middens (Wells, 1979; Van Devender and Spaulding, 1979). Recent studies substantiate fluctuating warm periods from the Early Tertiary to Late Eocene but envision much warmer conditions than the Geoflora model, followed by rapid climatic deterioration (Wolfe, 1978, 1980; Matthews, 1979). Paratropical" conditions are thought to have prevailed at higher latitudes over most of the Northern Hemisphere (ibid.). During this period the most suitable conditions of the Tertiary must have existed for entry of a S. tropica group taxon into North America. Physical aspects of a S. tropica group taxon dispersing from Africa into Eurasia and then to North America prior to the Late Eocene is complicated by two factors. First, the Tethys Sea isolated Africa from Eurasia during most of the Tertiary, such that faunal exchange was primarily limited to brief periods of (?filter) connection in the Late Oligocene, Middle and Late Miocene and Late Pliocene (Cooke, 1972; Hallam, 1973; Azzaroli and Guazzone, 1979). It is not precluded that other connections of very short duration occurred during which time Siphona could have entered Eurasia. Alternately, Siphona might have crossed the water barrier directly, by chance dispersal (a sweepstake route, such as between Madagascar and Africa). By whatever means a S. tropica group taxon reached Eurasia, timing was apparently not coincident with a period of substantial mammalian exchange. More important to my hypothesis than Siphona crossing the Tethys Sea at the most opportune time is a S. tropica group taxon reaching the climatic corridor before it closed in the Late Eocene. The second complication pertaining to a Siphona taxon reaching North America is the Turgai Strait separating Europe and Asia during the Early Tertiary (Adams, 1981). Siphona could have entered Europe by a route through Gilbraltar and crossed a North Atlantic land bridge (Matthews, 1979) without ever reaching Asia. Conversely, dispersal could have been via Asia and Beringia. At least the latter route seems highly probable because the existence of a S. tropica group member in the Oriental Region (the Philippines) implies that the Turgai Strait did not prevent the group from dispersing into Asia. At the end of the Late Eocene, the climate in northern regions cooled rapidly (Wolfe, 1978, 1980, as the “terminal Eocene event”). Because S. tropica group taxa are not known from America north of Mexico or the Palearctic Region, I suggest they were unable to adapt and so 10a Geoflora is a flora that undergoes little compositional change over a long period of time, ’‘intermediate to tropical and subtropical. North American species of Siphona Meigen 333 moved southward as conditions changed. This movement could have been in response to shifts in hosts’ ranges, but I doubt this. Even though Siphona hosts are inadequately known, the general distribution pattern of Siphona species - habitation of islands and continents far and wide - suggests to me that hosts are ubiquitous and not crucial to the understanding of Siphona zoogeography. It is unfortunately premature to attempt to cover circumstances by which the S. tropica group became established in Middle America since hosts, habitats and other aspects of extant species’ way of life are unknown. Habitat information is vital, for the history of Tertiary floras is complex (Graham, 1972; Axelrod, 1975, 1979; Rosen, 1978). It is significant, however, that the S. tropica group is not represented in eastern North America. Apparently the S. tropica group did not become associated with the temperate rain forest widely distributed in the United States and Middle America by the Middle Tertiary (Graham, 1972; Axelrod, 1975, 1979). Little data are available for inference of the history of the S', maculata, S. cristata and S. geniculata groups in the Old World. The former is recognized as the most primitive Siphona species group, so it presumably arose in Africa prior to all others. Examined specimens of Siphona taxa from Africa did not include S. maculata group members, so the group may have been virtually replaced in that region by more derived taxa. In America north of Mexico the S. maculata, S. cristata and S. geniculata groups have the same pattern of distribution, so probably have similar histories there. These histories are explored below. Species distributions are primarily northern and western in America north of Mexico, and from this several inferences can be drawn. It is likely the S. maculata, S. cristata and S. geniculata groups have not had a long history in the New World. The distributional pattern does not indicate an Eocene association with paratropical conditions as shown in the S. tropica group. Neither are eastern relicts of the Middle Tertiary temperate rain forest evident. Why the most primitive species group, the S. maculata group, shows a more recent distribution pattern than the derived S. tropica group, is an anomaly without a satisfactory zoogeographic explanation. During the Miocene a cooling trend was evident, as the climate slowly began to approximate our own. By late Middle Miocene higher latitudes were covered with a diverse microthermal coniferous forest with a least some floral similarity between Eurasia and North America (Wolfe, 1980). Similarly, a distinctive flora associated with Upper Tertiary orogenies was developing in western North America, while drier conditions were experienced in central regions and the temperate rain forest was retreating eastward and southward (Graham, 1972). It was probably during this time that the first members of the S. maculata and S. cristata groups entered North America, across the Bering land bridge12. A Miocene or younger Siphona invasion of North America might account for certain western elements in the distribution patterns of the S. maculata and S. cristata groups. It is possible both S. pacifica and S. pisinnia (or their ancestors) represent Late Miocene - Early Pliocene associations with young Rocky Mountain environments, particularly with their inferred status as primitive members of the S. maculata and S. cristata groups, respectively. The latter might be the older, for its distribution implies a warm temperate to subtropical adaptation. Neither species seems to represent a center of origin for North American members 12North Atlantic land bridge(s) broken by that time (Matthews, 1979; Eldholm and Thiede, 1980). Quaest. Ent., 1982, 18 (1-4) 334 O’Hara of its group, so I infer that subsequent migrations of S. maculata and S. cristata group taxa from the Palearctic Region took place. A western distribution of possibly more recent age is that of S. oligomyia. As sister species to -S. lutea, it almost certainly arose in America north of Mexico. S. lutea is cool temperate in adaptation and transcontinental in distribution (Fig. 89), and it is apparently parapatric or narrowly sympatric with the range of the more southerly distributed S. oligomyia (Fig. 90). Southward shifts in range of S. lutea (or ancestor) during Pleistocene glaciations perhaps lead to a vicariance of parental range in the west, with new species, S. oligomyia , developing a unique set of adaptations and structural characteristics. As mentioned in the “Phylogenetic interpretation” section, the relationship between S. maculata and S. floridensis is cladistically unresolved (Fig. 103), though these species might represent sister species, as inferred from their geographic distributions. S. maculata is widespread across northern North America (Fig. 82), while S. floridensis is only recorded from Florida and adjacent areas. Assuming S. maculata reached North America prior to the Holocene, it would have experienced a southward displacement during Pleistocene glaciations. Under full glacial conditions of the Wisconsin (18,000ybp) (or an earlier glaciation), S'. maculata might have reached northern Florida, as did some other boreal and cool temperate elements of the biota (reviewed in Wright, 1981). Florida was apparently too dry to support mesic forest during the Wisconsin full-glacial, but xeric oak-pine forest was present (Watts and Stuiver, 1980). Whatever the exact nature of the biotas, community structures must have been unique. Perhaps as the Laurentide ice sheet retreated and the climate warmed, most populations of S. maculata migrated northward, while others became stranded in pockets of suitable habitat in the south. Though most of the latter became extinct, S. floridensis might be a living descendant of one population that survived and adapted as conditions changed. (Similarly, apparently disjunct populations of S. maculata in Nevada and southern Colorado (Fig. 82) might also be relicts of the once extensive southern range of this species, though they have not undergone the phenetic change of S. floridensis.) Considerable similarity between North American and Palearctic members of the S. maculata and S. cristata groups indicate faunal exchanges have continued into the Quaternary period. At least three species in common between regions almost certainly reflects Pleistocene exchange during interglacials. The S', geniculata group as well could have entered North America as early as the Miocene, but there is no indication of this. S. medialis is more southerly distributed than S. hokkaidensis and more atypical of the group, so its arrival in North America presumably predates that of S. hokkaidensis. I interpret the extraordinarily varied appearance of S. hokkaidensis as primarily a Quaternary phenomenon, whereby populations were isolated and diverged. Further consideration of this taxon must await study of Palearctic members. The relationship between S. futilis and S. illinoiensis of the S. futilis group is very close, but other S. futilis group affinities are nebulous. Perhaps the group is very old; in the New World as old or older than the S. tropica group, since cool temperate taxa are evidently lacking. As sister species, S. futilis and S. illinoiensis conform to a distributional pattern of noted zoogeographic importance (Martin and Harrell, 1957; Axelrod, 1975; Rosen, 1978; Allen and Ball, 1980). S. futilis and S. illinoiensis are, respectively, found in Mexico (Fig. 96) and eastern United States (Fig. 92). It is generally agreed, on the basis of abundant data, that very North American species of Siphona Meigen 335 little biotal exchange has occurred between Mexico and eastern United States since the Miocene, when the Neotropical corridor between these areas deteriorated (Martin and Harrell, 1957; Rosen, 1978; Allen and Ball, 1980). The corridor was apparently not re-established even during Pleistocene full-glacials; at least not with mesic forest elements (reviewed in Wright, 1981). Despite the apparent rarity of post-Miocene biotal exchange between Mexico and eastern United States, I am reluctant to propose a Miocene vicariance for S. futilis and S. illinoiensis. By analogy with diversification of the S. tropica group in Mexico and inferred recent vicariances of S. lutea and S. oligomyia, and S. maculata and S. floridensis, a time span of ca. 15 million years or more for the vicariance (without major change) of S. futilis and S. illinoiensis seems too long. A more recent (Pleistocene) vicariance or dispersal is suggested by phenetic evidence. If this explanation is valid, this example is one of the few that demonstrates a post-Miocene connection between Mexico and eastern United States for mesic adapted taxa. Siphona affinities between Middle and South America have not been closely examined. Most species appear derived from northern elements, with a few evidently autochthonous species. I expect exchange prior to the Pliocene connection of North and South America occurred, though infrequently, leading to a small endemic Siphona fauna in South America. To reiterate, the following are the main conclusions drawn from a historical zoogeographic analysis of species groups and North American species of Siphona : 1 ) Siphona originated in Africa since the Late Cretaceous, 2) a S. tropica group taxon entered North America through a northern paratropical corridor connecting the Old and New Worlds before the climate cooled in the Late Eocene, 3) minimum age of Siphona is Late Eocene, 4) the S. maculata, S. cristata and S. geniculata groups entered North America via Beringia since the Miocene and periodic exchange continued into the Pleistocene, and 5) three species pairs represent Pleistocene vicariant events: S. maculata and S. floridensis, S. lutea and S. oligomyia , and S. futilis and S. illinoiensis. The major historical zoogeographic events concerning Siphona species within North America are summarized in Fig. 108. One of the elements of a zoogeographic hypothesis founded in part on cladistic relationships is its predictive power (Ball, 1976). Explicit predictions, like assumptions, contribute to the clarity and significance of hypotheses. The assumptions listed earlier are testable and therefore render their own predictions. To those can be added another. I do not expect taxa belonging to the S. maculata, S. cristata and S. geniculata groups to be found in South America, for I interpret the ancestors of North American members of these groups northern in origin and temperate to cold temperate in adaptation. Thus, representation of one or more of these groups in South America would imply either an earlier incursion into North America than I hypothesized (ie. during a warm period) or a shift in adaptation of a member taxon from temperate to subtropical or tropical. CONCLUDING REMARKS It should be apparent from the foregoing text that much remains to be learned about the genus Siphona. The basic classificatory work is inadequate for such regions as South America, Australia and the Orient, and until the required studies have been completed, it will continue to hinder attempts such as mine to delve into the historical (phylogenetic and zoogeographic) Quaest. Ent., 1982, 18 (1-4) 336 O’Hara aspects of the genus. With few unique and reliable character states of adults by which to distinguish Siphona species, identifications are commonly difficult. Since most keys are based exclusively on morphological differences among species, only discovery of new morphological characters are likely to improve them. Nevertheless there are other avenues of investigation into which I have not ventured that might prove useful for species delimitation and phylogenetic analysis. These include larval characters, host records, electrophoresis and cytological characters. I have written little about the ecological requirements of Siphona species, for indeed there is little known. Hosts are unrecorded for most North American species and additional information (such as habitat preference) is similarly lacking. I can but hope others will examine the aspects of Siphona species that I have not. ACKNOWLEDGEMENTS This paper was originally written in partial fulfillment of a Master’s degree. It is a pleasure to thank my supervisor, Dr. G.E. Ball, for his thoughtful advice and suggestions throughout the course of the study. In addition, his careful review of the first draft of the thesis led to significant improvements in both substance and form. Financial aid from Natural Sciences and Engineering Research Council grant A- 1399, held by Dr. G.E. Ball, is gratefully acknowledged. For first kindling my interest in tachinids, and suggesting Siphona as a thesis topic, I thank Dr. D.M. Wood of the Biosystematics Research Institute, Ottawa. His knowledge of, and enthusiasm for, the Tachinidae made discussions with him both informative and pleasurable. Dr. Wood reviewed the manuscript and provided useful suggestions for its improvement. Four other tachinid workers also contributed in various ways to this study. Dr. C.W. Sabrosky, recently retired from the Systematic Entomology Laboratory, USDA, Washington, was always prompt and helpful in my many dealings with him, and provided useful notes on Siphona types under his care. Dr. B. Herting of the Staatl. Museum fur Naturkunde, Stuttgart, provided specimens of Palearctic species of Siphona , and information on the Palearctic Siphona fauna. Dr. H. Shima of Kyushu University, Ropponmatsu, sent Japanese specimens of Siphona and commented on affinities among certain species. Special thanks are extended to Dr. S. Andersen of the Zoologisk Museum, Copenhagen, who provided manuscript copies of his “Revision of European species of Siphona Meigen” and “Phylogeny and classification of Old World genera of Siphonini”. These arrived while the thesis was being prepared for publication, so reference to them has been less than would otherwise have been the case. My wife, Wendy, assisted in many ways. As a companion on field trips, she helped collect tachinid specimens. She also prepared locality data lists and distribution maps, and typed portions of the manuscript into the computer. I thank fellow students at the University of Alberta for enlightening discussions of systematic theory and zoogeography, in particular J.S. Ashe, J.M. Cumming, G.A.P. Gibson and R.E. Roughley. I also thank G.A.P. Gibson for testing, and recommending changes to, my keys to adults of North American Siphona species. I also thank an anonymous reviewer and members of my Advisory Committee (Drs. D.A. Craig and J.S. Nelson) for critically reviewing the penultimate draft of the thesis, and suggesting improvements. J.-F. Landry kindly prepared the French abstract. 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Abbreviations: ar, arista; eye, compound eye; eye hgt, eye height; fr pit, frontal plate; fr s, frontal setae; gn, gena; gn dil, genal dilation; hd hgt, head height; i vt s, inner vertical seta; lab, labella; lgt flgm 1, length of first flagellomere; orb pit, orbital plate; o vt s, outer vertical seta; pafc, parafacial; pc orb s, proclinate orbital setae; pip, palpus; pocr, postcranium; prem, prementum; prob, proboscis; re orb s, reclinate orbital setae; rst, rostrum; subvb s, subvibrissal seta; vb, vibrissa. Quaest. Ent., 1982, 18 (1-4) 346 O’Hara Fig. 2. Structure and terms of Siphona thoracic nota, as illustrated by S. macronyx. Abbreviations: Notal divisions: psct, prescutum; set, scutum; sctl, scutellum. Setae: acr, acrostichal; apl, apical; bs, basal; dc, dorsocentral; ds, discal; hm, humeral; ial, intra-alar; ipal, intrapostalar; It, lateral; npl, notopleural; pal, postalar; phm, posthumeral; psl, presutural; sbap, subapical; spal, supra-alar. North American species of Siphona Meigen 347 3 Fig. 3. Right wing of S. maculata, illustrating average bristling on R4+5 and Siphona venation. Figs. 4-6. Male foreleg acropods, dorsal aspect. Setae shown on one side only. Claw hooks directed ventrally, hence not visible in figures. Figs. 4 and 6 illustrate average-sized claws and pulvilli, Fig. 5 large claws and pulvilli. Fig. 4. S. illinoiensis. Fig. 5. S. intrudens. Fig. 6. S. lutea. Figs. 7-8. Preabdomens, dorsal aspect. [Median marginal setae foreshortened in figures; subequal in length to laterals.] Fig. 7. S. intrudens , illustrating weak marginal setae and absence of median marginal setae on T1+2. Fig. 8. S. medialis, illustrating strong lateral marginal setae and presence of median marginal setae on T1+2. Abbreviations: Veins: A,, A2, branches of anal; CuAp cubitus; M, media; R,, R,+3, R4+5, branches of radius; Sc, subcosta. Crossveins: dm-cu, discal medial-cubital; r-m, radial-medial. Abdomen: lat marg, lateral marginal seta; med marg, median marginal seta; T]+2, T3, T4, T5, abdominal terga. Quaest. Ent., 1982, 18 (1-4) 348 O’Hara 12 Fig. 9. Structure and terms of Siphona male genitalia, as illustrated by S. illinoiensis. Fig. 9a. Anterior view of distiphallus. Fig. 9b. Lateral view of genitalia. Fig. 10. Sternum 5 of S. lutea, illustrating average condition of postero-lateral margins. Fig. 1 1 . Sternum 5 of S. oligomyia , illustrating hooked condition of postero-lateral margins. Fig. 12. Structure and terms of Siphona female genitalia, as illustrated by S. hokkaidensis. Fig. 12a. Ventral view. Fig. 12b. Dorsal view. Fig. 12c. Lateral view. Abbreviations: Male genitalia: aed apod, aedeagal apodeme; bsph, basiphallus; cere, cerci; dsph, distiphallus; epand, epandrium; gon, gonopod; hypd apod, hypandrial apodeme; sur, surstylus. Female genitalia: cere, cerci; sg 8, segment 8; st 5, 6, 7 and 10, abdominal sterna. Scale bars = 0.1mm. North American species of Siphona Meigen 349 Quaest. Ent., 1982, 18 (1-4) Fig. 13. Flubbs-Hubbs diagram illustrating variation among specimens of Siphona species; eye height: head height. The range for each species is indicated by a horizontal line and the mean by a short vertical line. A hollow rectangle represents two standard errors to either side of the mean and a solid rectangle represents 1.5 standard deviations to either side of the mean. Only range and mean values are shown for samples with less than ten specimens. See “Methods" section for further information. .44 350 O’Hara Fig. 14. Hubbs-Hubbs diagram illustrating variation among specimens of Siphona species; length of first flagellomere: head height. See caption for Fig. 13 and “Methods” section for further information. 2.0 2.5 North American species of Siphona Meigen 351 « s <£> m t- O^N’- tM r CM *■ in ^ N O 00 CN CO r- o ^ in (O i- « (0 | "Quaest . Ent., 1982, 18 (1-4) Fig. 15. Hubbs-Hubbs diagram illustrating variation among specimens of Siphona species; proboscis length: head height. See caption for Fig. 13 and “Methods" section for further information. 06 352 O’Hara oo CM cm oo in co eo co at Tt in o co o> 10 a> vt at CM o o> C O 00 CO N; CO N; oo ID. CO CD CO o CO CO CO in 'T in CM in o in o UJ CC Fig. 16. Hubbs-Hubbs diagram illustrating variation among specimens of Siphona species; length of first flagellomere: eye height. See caption for Fig. 13 and “Methods” section for further information. North American species of Siphona Meigen 353 Quaest. Ent., 1982, 18 (1-4) 354 O’Hara Figs. 18-21. Male head profiles of Siphona species. Fig. 18. S. maculata\ Fig. 19. S. floridensis\ Fig. 20. S. intrudens\ Fig. 21. S. lurida ; Scale bars = 0.5mm. North American species of Siphona Meigen 355 Figs. 22-25. Male head profiles of Siphona species. Fig. 22. S. pacifiea\ Fig. 23. S. pisinnia\ Fig. .24. 5. cristata: ; Fig. 25. S. multifaria\ Scale bars = 0.5mm. Quaest. Ent., 1982, 18 (1-4) 356 O’Hara Figs. 26-29. Male head profiles of Siphona species. Fig. 26. S. lutea ; Fig. 27. S. oligomyia\ Fig. 28. S. geniculata ; Fig. 29. S. medialis\ Scale bars = 0.5mm. North American species of Siphona Meigen 357 Figs. 30-33. Male head profiles of Siphona species. Fig. 30. S. hokkaidensis , antennal type 1; Fig. 31. S'. hokkaidensis, antennal type 2; Fig. 32. S', hokkaidensis , antennal type 3; Fig. 33. S. macronyx\ Scale bars = 0.5mm. Quaest. Ent., 1982, 18 (1-4) 358 O’Hara Figs. 34-37. Male head profiles of Siphona species. Fig. 34. S. futilis ; Fig. 35. S. brunnea\ Fig. 36. S. akidnomyia; Fig. 37. S. longissima. Scale bars = 0.5mm. North American species of Siphona Meigen 359 Figs. 38-40. Male head profiles of Siphona species. Fig. 38. S. tropica ; Fig. 39. S. rizaba\ Fig. 40. S. unispina., Fig. 41. Head profile of female S. lurida\ Scale bars = 0.5mm. Quaest. Ent., 1982, 18 (1-4) nw mv£ 360 O’Hara Figs. 42-50. Female head profiles of Siphona species. Fig. 42. S. maculata; Fig. 43. S. floridensis\ Fig. 44. S. intrudens\ Fig. 45. S. pacifica\ Fig. 46. S. cristata ; Fig. 47. S. multifaria\ Fig. 48. S. lutea\ Fig. 49. S'. pisinnia ; Fig. 50. S. geniculata\ Scale bars = 0.5mm. North American species of Siphona Meigen 361 Figs. 51-59. Female head profiles of Siphona species. Fig. 51. S. hokkaidensis\ Fig. 52. S. medialis\ Fig. 53. S. futilis\ Fig. 54. S. illinoiensis\ Fig. 55. S. brunnea\ Fig. 56. S. macronyx; Fig. 57. S. tropica ; Fig. 58. S. rizaba\ Fig. 59. S. akidnomyia\ Scale bars = 0.5mm. Quaest. Ent., 1982, 18 (1-4) 362 O’Hara Figs. 60-65. Male genitalia of Siphona species: a. Anterior view of distiphallus; b. Lateral view of genitalia. Fig. 60. S. maculata\ Fig. 61. S. floridensis\ Fig. 62. 5". intrudens\ Fig. 63. S. lurida\ Fig. 64. S. pacifica\ Fig. 65. S. pisinnia ; Scale bars = 0.1mm. North American species of Siphona Meigen 363 Figs. 66-71. Male genitalia of Siphona species: a. Anterior view of distiphallus; b. Lateral view of genitalia; c. Lateral view of second distiphallus. Fig. 66. S. cristata\ Fig. 67. S. multifaria\ Fig. 68. 5. lutea\ Fig. 69. S. oligomyia ; Fig. 70. S. geniculata\ Fig. 71. S. hokkaidensis\ Scale bars = 0.1mm. Quaest. Ent., 1982, 18 (1-4) 364 O’Hara Figs. 72-77 . Male genitalia of Siphona species: a. Anterior view of distiphalius; b. Lateral view of genitalia. Fig. 72. S. medialis ; Fig. 73. S. futilis'. Fig. 74. S. brunnea\ Fig. 75. S. macronyx\ Fig. 76. S. akidnomyia\ Fig. 77. S. longissima-. Scale bars = 0.1mm. North American species of Siphona Meigen 365 Figs. 78-80. Male genitalia of Siphona species: a. Anterior view of distiphallus; b. Lateral view of genitalia. Fig. 78. S', tropica ; Fig. 79. S. rizaba ; Fig. 80. S. unispina\ Scale bars = 0.1mm. Quaest. Ent., 1982, 18 (1-4) 366 O’Hara Fig. 81. Known North American distribution of Holarctic species Siphona hokkaidensis , with data on antennal type and abdominal setae. Dots denote locality records for S. hokkaidensis “b” (Figs. 13-16), and female specimens for which there were no male associates. Squares denote locality records for male specimens with antennal type 1, and associated females, from Alberta, Saskatchewan and Manitoba ( S . hokkaidensis" a”). Rings signify locality records for which data were lacking. Abbreviations: Antennal type 1, Fig. 30; type 2, Fig. 31; type 3, Fig. 32; f, female, Fig. 51. N, number of specimens in sample. No. of med. marg. on T,+2, number of specimens with 0, 1 or 2 median marginal setae on T1+2: 0, setae absent; 1, one seta on one side only; 2, 1 pair of setae. >1 pr. lat. marg. T3, number of specimens with more than 1 lateral marginal seta on at least one side on T3. North American species of Siphona Meigen 367 Fig. 82. Known North American distribution of the Holarctic species Siphona maculata Staeger. Fig. 83. Known distribution of Siphona pisinnia n. sp. Fig. 84. Known distribution of Siphona floridensis n. sp. Quaest. Ent., 1982, 18 (1-4) 368 O’Hara Fig. 85. Known distribution of Siphona intrudens (Curran). Fig. 86. Known North American distribution of the Holarctic species Siphona lurida Reinhard. Fig. 87. Known distribution of Siphona pacifica n. sp. Fig. 88. Known distribution of Siphona macronyx n. sp. North American species of Siphona Meigen 369 Fig. 89. Known distribution of Siphona lutea (Townsend). Fig. 90. Known distribution of Siphona oligomyia n. sp. Fig. 91. Known North American distribution of S. geniculata (De Geer) (introduced from the Palearctic Region). Fig. 92. Known distribution of Siphona illinoiensis Townsend. Quaest. Ent., 1982, 18 (1-4) 370 O’Hara Fig. 93. Known distribution of Siphona multifaria n. sp. North American species of Siphona Meigen 371 Fig. 94. Known distribution of Siphona medialis n. sp. Fig. 95. Known North American distribution. of the Holarctic species Siphona cristata (Fabricius). Quaest. Era., 1982, 18 (1-4) 372 O’Hara Figs. 96-101. Known distributions of Siphona species. Fig. 96. Siphona futilis van der Wulp. Fig. 97. Siphona brunnea n. sp. Fig. 98. Siphona rizaba n. sp. Fig. 99. Siphona akidnomyia n. sp. Fig. 100. Siphona longissima n. sp. Fig. 101. Siphona tropica (Townsend). North American species of Siphona Meigen 373 Character Siphona N. species N 1 2 3 4 5 6 7 8 9 10 12 13 14 15 No. of apotypies Flagel lomere 1 Proboscis Aristomere 3 Vibrissa! angle Colouration Wing setulae R-| in CO CD 25 § s Tarsal claws Median marginals Lateral marginals Sternum 5 Dentition on distiphallus Distiphallus Surstylus maculata a A1 a a a a a a a a a a A2 A a 3 floridensis a A1 a a a a a a a a a a a a a 1 intrudens A-1 A1 A1 a a A a a A2 a a a A3 A a 7 lurida A-2 A1 A1 a A2 A a a A2 a A a a a a 7 pacifica A1 A2 A2 a a a a a a a a a a a a 3 cristata a B1 A4 a a B1 a a a a A a a B a 5 multifaria a B1 A4 a a a a a a a A a a B a 4 lutea a B1 A2 a a B2 A A a a A a a B a 7 oligomyia A'1 A A3 a a B2 A ◦ A1 a A A a a a 8 pisinnia a B2 A3 a a a a a a ◦ A a a a a 3 geniculata a C1 A3 A1 a a a a A1 A2 B a a C A 8 hokkaidensis a D A3 A2 a a a a a A1 B a a C A 7 medialis A2 C1 A3 A1 a C a a a A2 A a a a a 7 futilis A2 C1 A3 a a a a a a a A a a D a 5 illinoiensis A2 c2 A3 a a a a a a a A a a D a 5 brunnea A1 E A3 a a D2 a a A1 a A a a E a 7 tropica A2 F A5 a a a A a a a A a a G a 6 rizaba A2 F A5 a a a a a a a A a a a A 5 longissima A1 F A5 a a a a a A1 a A a A1 G a 7 akidnomyia a F A4 a a a ◦ a A2 a A a A1 F a 6 unispina a F A5 a a a A A a a A a A1 G a 7 macronyx a C1 A3 a A1 D1 a a A2 a A a A1 a a 7 Fig. 102. Character matrix presenting the distribution of character states among North American Siphona species and African species S. unispina (Mesnil). Characters and states are discussed in “Phylogenetic interpretation” section. Quaest. Ent., 1982, 18 (1-4) 374 O’Hara ° "< < u. Fig. 103. Cladogram illustrating inferred relationship between the S. maculata group and the other North American Siphona species groups, with a hypothetical phylogeny of North American species of the S. maculata species group. Dots denote apotypic character states, rings plesiotypic states. For explanation and discussion of characters and states see Fig. 102 and “Phylogenetic interpretation” section. North American species of Siphona Meigen 375 Figs. 104-107. Cladograms illustrating hypothetical phylogenetic relationships among North American species of Siphona species groups. Dots denote apotypic character states, rings plesiotypic states. For explanation and discussion of characters and states see Fig. 102 and “Phylogenetic interpretation” section. Fig. 104. The S. cristata group; Fig. 105. The S. geniculata group; Fig. 106. The S. tropica group; Fig. 107. The S. futilis group. Quaest. Ent., 1982, 18 (1-4) 376 O’Hara Siphona species groups. \ \ "7 ? <2° 0 - S. tropica gp. (D - S. maculata gp. Lata Tartiary'O^' 1 /T / C to Ploistocono T\ ■ S. cristata gp. ■H i : ' % . : M* i S. geniculata gp. (D - S. futilis gp. 108 I T ® \ Fig. 108. Generalized historical zoogeography of North American species of Siphona, as inferred from a cladistic analysis and species distributions. (1) 5. tropica group - migration into North America through Beringia or across a North Atlantic land bridge prior to the Late Eocene during Tertiary climatic optimum, with subsequent diversification in Mexico, (a) Late Tertiary to Pleistocene migration of taxa into Central and South America. (2) S. maculata, S. cristata and S. geniculata groups - first representatives arrived in North America during the Miocene; exchange continued into Pleistocene, (a) Oldest taxa of these groups might have become associated with the developing Rocky Mountain biota (?Late Miocene - Pliocene), (b) Transcontinental, cool to cold temperate taxa, indicative of later invasions (?Pliocene - Pleistocene), (c) Vicariance of S. lutea and 5. oligomyia (Pleistocene), (d) Vicariance of S. maculata and S. floridensis (Pleistocene). (3) S. futilis group - place and age of origin uncertain. Vicariance of S. futilis and S. illinoiensis (Pleistocene). North American species of Siphona Meigen 377 INDEX TO NAMES OF TAXA (Synonyms in italics) FAMILIES AND TRIBES Bombyliidae, 329 Noctuidae, 264 Oestridae, 329 Piophilidae, 340 Siphonini, 278, 309, 314 Tachinidae, 314, 326, 329, 330, 336 Tipulidae, 264 GENERA Aphantorhapha Townsend, 274, 314 Asiphona Mesnil, 274, 314 Bucentes Latreille, 275, 278 Ceranthia Robineau-Desvoidy, 274, 314, 319 Ceranthia 5. lat., 274, 314, 315, 316, 326 Ceranthia 5. str., 274 Crocuta Meigen, 275 Dischistus Loew, 329 Peribaea Robineau-Desvoidy, 314 Phantasiosiphona Townsend, 274, 275 Protopiophila Duda, 329 Pseudosiphona Townsend., 314, 315 Siphonopsis Townsend, 273, 274, 314, 315 SPECIES AND SUBSPECIES , cinereus Latreille, Bucentes, 275 abbreviata (Villeneuve), Siphona, 278 abbreviata Villeneuve, Bucentes, 278 abdominalis Robineau-Desvoidy, Ceranthia, 317 akidnomyia new species, Siphona, 278, 285, 286, 306, 308, 311, 312, 313, 317, 325, 326, 328 albocincta (Villeneuve), Siphona, 278 albocincta Villeneuve, Bucentes, 278 alticola (Mesnil), Ceranthia, 275 alticola (Mesnil), Siphona, 275 alticola Mesnil, Crocuta, 214, 275 amoena (Mesnil), Siphona, 278 amoena Mesnil, Crocuta, 278 amplicornis Mesnil, Siphona, 278 analis Meigen, Siphona, 278, 299 analis Robineau-Desvoidy, Siphona, 280 angusta Mesnil, Siphona, 278 antennalis (Mesnil), Siphona, 278 antennalis Mesnil, Crocuta, 278 anthomyformis Lynch Arribalzaga, Siphona, 280 atricapilla Mesnil, Siphona, 278 bevisi Curran, Siphona, 278, 318 bilineata (Mesnil), Siphona, 278 bilineata Mesnil, Crocuta, 278 boreata Mesnil, Siphona, 278, 325 brasiliensis Townsend, Siphonopsis, 273, 274 brunnea new species, Siphona, 278, 285, 286, 305,306,312,325,328 brunnescens (Villeneuve), Asiphona, 318 candelisequa Schiffermiiller, Agrotis, 273 capensis Curran, Siphona, 278 ceres (Curran), Siphona, 278, 305 ceres Curran, Bucentes, 278, 305 chaetolyga Rondani, Siphona, 278, 292 cinerea (Latreille), Siphona, 219 cinerea Latreille, Bucentes, 279, 299 cinerea Meigen, Siphona, 219 clausa Robineau-Desvoidy, Siphona, 280 collini Mesnil, Siphona, 278, 322 conata Reinhard, Siphonopsis, 273, 274 confusa Mesnil, Siphona, 273, 278, 323 consimilis Robineau-Desvoidy, Siphona, 280 cothurnata (Mesnil), Siphona, 278 cothurnata Mesnil, Crocuta, 278 crassulata (Mesnil), Ceranthia, 274, 275 crassulata (Mesnil), Siphona, 275 crassulata Mesnil, Crocuta, 274, 275 creberrima (Speiser), Siphona, 278 creberrima Speiser, Crocuta, 21 8 cristata (Fabricius), Siphona, 267, 272, 273, 278, 281, 284, 292, 293, 294, 295, 296, 297, 298, 299, 303, 307, 308, 313, 319, 322, 323, 324, 328, 330, 331, 333, 334, 335 cristata Fabricius, Stomoxys, 278, 292 Quaest. Ent., 1982, 18 (1-4) 378 O’Hara cuthbertsoni Curran, Siphona, 274, 278, 298, 319 delicatula Mesnil, Siphona , 279 diluta (Wulp), Clausicella, 275 diluta Wulp, Siphona , 273, 275 dorsalis Brauer and Bergenstamm, Siphona , 280 efflatouni Mesnil, Siphona, 278, 322 flavifrons Staeger, Siphona, 273, 278, 322, 324 flavipes (Coquillett), Ceranthia, 317 floridensis new species, Siphona, 278, 282, 286, 287, 288, 289, 316, 321, 322, 323, 326.328.334.335 foliacea (Mesnil), Siphona, 278 foliacea Mesnil, Crocuta , 278 fuliginea cerina Mesnil, Siphona, 278 fuliginea Mesnil, Siphona, 278 fuliginea rubea Mesnil, Siphona, 278 fuscicornis Robineau-Desvoidy, Siphona, 280 futilis Wulp, Siphona, 273, 278, 285, 286, 305, 306, 307, 308, 310, 311, 312, 324, 325.328.334.335 gedeana Wulp, Siphona, 278 geniculata (De Geer), Siphona, 264, 273, 275, 276, 277, 278, 282, 299, 300, 301, 303, 304, 307, 322, 323, 324, 326, 327, 328.330.331.333.334.335 geniculata De Geer, Musca, 275, 299, 300 gigantea Schrank, Tipula, 273 gracilis (Mesnil), Siphona, 279, 325 gracilis Mesnil, Crocuta , 279 grandistyla Pandelle, Siphona, 279, 318, 324, 325 hokkaidensis Mesnil, Siphona, 267, 268, 273, 276, 279, 282, 283, 284, 300, 301, 302, 303, 304, 316, 323, 324, 328, 334 humeralis Robineau-Desvoidy, Siphona, 280 illinoiensis Townsend, Siphona, 277, 279, 283, 284, 304, 306, 307, 308, 324, 325, 328, 334, 335 infuscata (Mesnil), Siphona , 280 infuscata Mesnil, Crocuta , 280 intrudens (Curran), Siphona, 267, 277, 279, 282, 283, 287, 288, 289, 290, 291, 297, 303, 308, 321, 322, 326, 327, 328 intrudens Curran, Bucentes, 279, 288 irrorata Meigen, Tipula, 273 janssensi (Mesnil), Siphona, 279 janssensi Mesnil, Crocuta , 279 jocularis Mesnil, Actia, 274 kuscheli (Cortes), Siphona, 279 kuscheli Cortes, Phantasiosiphona , 274, 279 laticornis Curran, Siphona, 279 lindneri Mesnil, Siphona, 279 longissima new species, Siphona, 279, 285, 306, 309, 310, 311, 312, 313, 325, 328 lurida Reinhard, Siphona, 276, 279, 282, 286, 287, 289, 290, 291, 298, 316, 317, 321,322,326,327,328 lutea (Townsend), Siphona, 275, 276, 279, 282, 292, 293, 294, 296, 297, 298, 303, 308, 313, 317, 318, 322, 323, 325, 328, 334,335 lutea Townsend, Crocuta , 279, 296 macronyx new species, Siphona, 279, 283, 284, 311, 312, 313, 317, 325, 326, 328 maculata griseola Mesnil, Siphona, 279 maculata Staeger, Siphona, 267, 272, 273, 279, 282, 283, 286, 287, 288, 289, 291, 297, 298, 303, 316, 317, 319, 321, 322, 326, 327, 328, 329, 331, 333, 334, 335 maculipennis Meigen, Siphona, 279 malaisei (Mesnil), Actia, 275 malaisei (Mesnil), Siphona , 275 malaisei Mesnil, Crocuta, 274, 275 maxima Poda, Tipula, 273 medialis new species, Siphona, 276, 277 , 279, 281, 303, 304, 308, 313, 323, 328 meigenii (Lepeletier and Serville), Siphona , 279, 300 meigenii Lepeletier and Serville, Bucentes, 279 melania (Bezzi), Siphona, 279 melania Bezzi, Bucentes, 279 melanocera Robineau-Desvoidy, Siphona, 280 melanura Mesnil, Siphona, 279 minuta (Fabricius), Stomoxys, 279, 300 Quaest. Ent., 1982, 18 (1-4) North American species of Siphona Meigen 379 multifaria new species, Siphona, 267, 272, 273, 279, 281, 284, 285, 286, 288, 292, 293, 294, 295, 297, 298, 299, 303, 307, 308,319,322,323,328 munroi Curran, Siphona, 279 murina (Mesnil), Siphona, 279 murina Mesnil, Crocuta, 279 nigricans (Villeneuve), Siphona, 279, 324, 327 nigricans Villeneuve, Bucentes , 279 nigripalpis (de Meijere), Siphona , 278 nigrohalterata Mesnil, Siphona , 278 nigroseta Curran, Siphona, 279 nigrovittata Meigen, Siphona , 279, 300 nobilis (Mesnil), Siphona, 279 nobilis Mesnil, Crocuta , 279 obelisea Schiffermiiller, Agrotis, 273 obesa (Mesnil), Siphona, 279 obesa Mesnil, Crocuta , 279 obscuripennis Curran, Siphona, 279 oculata Pandelle, Siphona , 279, 292 oleracea L., Tipula, 273 oligomyia new species, Siphona, 274, 277, 279, 282, 296, 297, 298, 313, 318, 319, 322, 323,325,326,328,334,335 pacifica new species, Siphona, 279, 283, 284, 286, 289, 291, 298, 313, 321, 323, 328,333 palpina Zetterstedt, Siphona , 278, 292 paludosa Meigen, Tipula, 264, 273 paludosa Mesnil, Siphona, 279, 324 patellipalpis (Mesnil), Siphona, 279 patellipalpis Mesnil, Crocuta , 279 pauciseta Mesnil, Siphona , 280 pauciseta Rondani, Siphona, 279, 292, 323 pellex (Mesnil), Actia, 275 pellex (Mesnil), Siphona , 275 pellex Mesnil, Crocuta , 274, 275 persilla Coquillett, Siphona , 280 phantasma (Mesnil), Siphona, 279 phantasma Mesnil, Crocuta , 279 pigra Mesnil, Siphona, 279 pisinnia new species, Siphona, 276, 280, 281, 283, 284, 285, 286, 292, 293, 294, 298, 299, 306, 322, 323, 327, 328, 333 plusiae (Coquillett), Siphonopsis, 273, 274, 305 pseudomaculata Blanchard, Siphona, 280 pusilla Robineau-Desvoidy, Siphona, 280 quadricincta Rondani, Siphona , 280 quadrinotata Robineau-Desvoidy, Siphona, 280 reducta (Mesnil), Siphona, 280 reducta ludicra Mesnil, Siphona, 280 reducta Mesnil, Crocuta , 280 rizaba new species, Siphona, 280, 285, 286, 306, 309, 310, 311, 312, 325, 328 rossica Mesnil, Siphona, 280, 324 rubrapex Mesnil, Siphona, 280 rubrica (Mesnil), Siphona, 280 rubrica Mesnil, Crocuta , 280 setinerva (Mesnil), Siphona, 280, 318 setinerva Mesnil, Crocuta , 280 setosa Mesnil, Siphona, 280 seyrigi Mesnil, Siphona, 280, 321, 327 silvarum Herting, Siphona , 279, 301 silvatica Robineau-Desvoidy, Siphona, 280 simulans (Mesnil), Siphona, 280 simulans Mesnil, Crocuta , 280 singularis (Weidemann), Siphona, 280 singulars Wiedemann, Tachina, 280 siphonosoma Malloch, Actia, 274 sola Mesnil, Siphona, 280 spinulosa (Mesnil), Siphona, 280 spinulosa Mesnil, Crocuta , 280 tachinaria Meigen, Siphona , 279, 300 taiwanica (Baranov), Siphona, 280 taiwanica Baranov, Crocuta , 280 tenuis Curran, Siphona , 279, 296 testacea Robineau-Desvoidy, Siphona, 280 trichaeta (Mesnil), Siphona, 280 trichaeta Mesnil, Crocuta , 280 triseta Mesnil, Actia, 274 tristis Robineau-Desvoidy, Siphona, 280 tropica (Townsend), Siphona, 276, 277 , 280, 285, 286, 308, 309, 310, 311,312, —318, 325, 327, 328, 331, 332, 333, 334, 335 tropica Townsend, Phantasiosiphona , 274, 275, 280, 309 unispina (Mesnil), Siphona, 280, 318, 325, 331 380 O’Hara unispina Mesnil, Crocuta , 280 urbanis (Harris), Siphona , 279 urbanis Harris, Musca, 279, 300 vittata Curran, Siphona, 280 vixen Curran, Siphona, 280 wittei (Mesnil), Siphona, 280 wittei Mesnil, Crocuta , 280 Quaest. Ent., 1982, 18 (1-4) 381 BOOK REVIEW SPENCER, K.A. 1981. A revisionary study of the leaf-mining flies (Agromyzidae) of California. Division of Agricultural Sciences, University of California, Special Publication 3273. iv + 489 pages containing 655 figures. $20 U.S. This work is essentially a regional fauna, with description of new species and revisionary comments included as necessary. It raises the number of agromyzid species recorded in California to 252, of which 132 are described as new. Californian dipterists will find Spencer’s treatment a sound foundation for further studies, and I hope they will be inspired to make it out of date as soon as possible. At 252 species the total of known Californian species is surely still underdocumented in comparison with the well over 300 species now known in the British Isles. I would expect the real total of Californian species to be in excess of 500 in view of the great diversity of climates and habitats represented in that state. Two new taxa of the genus-group are proposed in this work: Galiomyza (type-species Agromyza morio Brishke) for certain leaf-miners of Rubiaceae formerly included in a heterogenous concept of Praspedomyza, and Annimyzella (type-species Agromyza maculosa Malloch) as a subgenus of Amauromyza. These proposals seem to me justified. In general the taxonomic treatment is up-to-date, but the synonymy of Metopomyza griffithsi Sehgal with M. scutellata (Fallen) proposed on page 336 has been refuted in the special study of Metopomyza contained in Tschirnhaus’ (1981) work “Die Halm- und Minierfliegen im Grenzbereich Land-Meer der Nordsee” (Spixiana, suppl. 6). Spencer’s figure of the aedeagus of a Californian specimen (Fig. 487) agrees with that of M. griffithsi described from Alberta. The true M. scutellata has not been demonstrated to occur in North America. The generic position of the new species described as Phytomyza minutissima Spencer needs further study. This species does not run to Phytomyza in Spencer’s generic key because of the lack of orbital setulae. Reduction of the aedeagal sclerotization makes the relationship of the species difficult to interpret. On the basis of the information given, it could alternatively belong to Paraphytomyza or Gymnophytomyza as well as to Phytomyza. The 655 figures were mostly prepared by the author’s wife Ann and are of her usual good quality. Some figures have been reproduced from previous publications, but most are original. Scale lines have unfortunately been omitted. While the content of this work meets with my approval and I have no hesitation in recommending it to students of North American Diptera, there are some shortcomings in the presentation. Offset printing has been used without reduction on pages of the standard size for typing paper (8.5 x 1 1 inches). This results in an unnecessarily large volume which is too heavy in relation to the strength of its (paper) binding. Libraries and regular users of the work will need to have their copies rebound with hard covers. It would have been to the benefit of all concerned if the page size had been reduced and stronger binding used. The figures were intended for 1 /3 reduction, and considerable reduction in the space occupied by text could have been achieved by use of single spacing. The proof reading has also been remiss, as evidenced by the tantalizing references to “p. ” which appear frequently in the text. Page 7 has been duplicated (appearing again as p. 12). More careful editing of future publications in this series is called for. Graham C.D. Griffiths Department of Entomology The University of Alberta Edmonton, Alberta T6G 2E3 EDITOR’S ACKNOWLEDGEMENTS In times of financial uncertainty, such as these, editors, and especially publishers, need to scrutinize their operations- for obvious reasons. But it would seem that, at this time, examination of editorial procedures and consciences is a more pressing need. Exponential increase in scientific publication has reached crisis proportions, seemingly as a result of factors having rather little to do with desire of authors to add to our store of knowledge (see Broad, W.J., 1982, Bioscience 32 (8): 645-647). Associated with this increase has been an increase in cost, an increase in presentation of false results, an increase in plagiarism, an increase in repetition of content, an increase in multiple authorship with a corresponding decrease in responsibility for material reported, a decrease of information per publication (the ultimate being publication of “least publishable units”, or LPU’s), a possible decrease in quality of content, and major problems for working scientists to cope with the volume of published pages. Mr. Broad, a reporter for Science, concludes that scientific papers seem to have become more tools for careerists to advance their own causes, rather than principal vehicles for transmission of knowledge. He refers to the situation as “the disease of paper inflation”. It seems to me that responsibility for this state of affairs is partly that of editors, and this is why an examination of conscience is called for. Each editor can contribute to the needed cure for paper inflation by first, identification of how his journal might have contributed to the disease, and second, by a vigorous attempt to expunge those procedures deemed undesirable. While not wishing to be self-righteous, I hasten to point out that Quaestiones Entomologicae cannot be accused of contributing to paper inflation by encouraging publication of “LPU’s”, and I am confident that our authors are not guilty of the other types of offenses listed. In general we have sought extensive, original, treatments of varied entomological topics. I think this volume is a fair sample of the sorts of papers published over the years, and I thank the authors for choosing this journal. As I write these lines, preparation of Volume 18 is nearing completion, and by resorting to the expediency of publishing the entire year’s offering between two covers only, it seems likely that we will achieve our goal of again issuing an entire volume in the same year that is designated on the cover. Mrs. Suseela Subbarao, Publications Manager, is in large part responsible for this, and I am grateful to her for her dedicated effort. Although an editor is responsible for what appears in his journal, he seeks from others the advice he must have about quality and accuracy of content of submitted manuscripts. He is not compelled to accept all of the advice given, though it is probably in his best interests and that of his journal to do so. Whether or not a reviewer’s advice is accepted, it is appropriate to acknowledge assistance rendered. It is in that spirit that the following are acknowledged, and the appearance of their names here does not imply endorsement of the contents of this volume. Be that as it may, I am grateful for the assistance of: H. E. Evans, Department of Zoology and Entomology, Colorado State University, Fort Collins, Colorado; J. R. Spence, Department of Entomology, University of Alberta; D. D. Wilder, Insect Identification laboratory, Agriculture Research Service, U. S. D. A., United States National Museum, Washington, D. C.; D. M. Wood, Biosystematics Research Institute, Research Branch, Agriculture 384 Canada, Ottawa, Ontario; G. C. Williams, Center for Advanced Study in the Behavioral Sciences, Stanford, California. D. A. Craig served as editor during part of the summer, and undertook a substantial portion of the chores associated with this volume. J. F. Landry prepared French translations of several of the abstracts. J. H. Cumming and D. J. Williams read proof. I am grateful to all of these individuals, and I look forward to receiving their cooperation in preparation of future volumes. George E. Ball September, 1982. INDEX TO VOLUME 18 385 Acilius sulcatus (L.), 3 adusta Navas, Povilla , 43, 45, 52 Afronurus Lestage, 33 albopilosus Cresson, Philanthus, 94, 95, 96, 98,99, 103, 104, 105, 106, 112, 113, 116, 118, 119, 120, 121, 123, 124 albrighti Traver, Ametropus, 44 album (Say), Ephoron, 43 Alcock, J., 93, 99, 105 Allen, R.K., 21,30 Alouf, N .J., 44 alpinus (Pictet), Baetis, 26 Ameletus Eaton, 43, 44 Ameletus inopinatus Eaton, 43, 53 americana Edmunds and Traver, Dolania, 44 Ametropodidae, 44 Ametropus albrighti Traver, 44 Ametropus ammophilus Allen and Edmunds, 44 Ametropus eatoni Brodskij, 44 Ametropus fragilis Albarda, 44 Ametropus neavei McDunnough, 44 ammophilus Allen and Edmunds, Ametropus , 44 Andersen, T., 46 Anderson, N.H., see Lehmkuhl, D.M., 34 Andrenidae, 93, 104 Aneletris Edmunds, 43 Anepeorus McDunnough, 33, 54 Aphilanthops frigidus (F. Smith), 103, 106 Aphilanthops Patton, 103 Armitage, K.B., 93, 103, 104, 105 Arthroplea Bengtsson, 33 Ashwell, I.Y., 95 Atalophlebiodes Phillips, 34 atratus McDunnough, Tricorythodes, 44 aurivillii Bengtsson, Ephemerella, 31 Axelrod, D.I., 122 Baetidae, 23, 27, 45, 47, 54 Baetiella, 27 Baetis alpinus (Pictet), 26 Baetis fuscatus (L.), 26 Baetis Leach, 20, 23, 27, 45, 54 Baetis macani bundyae, 26 Baetis macani Kimmins, 26 Baetis muticus (L.), 26 Baetis rhodani (Pictet), 23, 26 Baetis scambus Eaton, 26, 45 Baetis tricaudatus Dodds, 26 Baetis vagans McDunnough, 26 Baetis vernus (Curtis), 26 Baetisca bajkovi Neave, 44 Baetisca obesa (Say), 44 Baetisca rogersi Berner, 44 Baetisca Walsh, 44 Baetiscidae, 44 Baetodes Needham and Murphy, 27 bajkovi Neave, Baetisca , 44 Bartholomae, P.G., 43 basale (Walker), Siphloplecton , 34 Bates, H.W., 8 Behningiidae, 44 belgica Lestage, Torleya, 31 Bembix , 1 1 2 Bembix littoralis Turner, 1 12 Bembix texana Cresson, 1 12 Benech, V., 23 Benke, A.C., 47, 48 Berner, L., , 35 see also Edmonds, G.F., 21 see also Hall, R.J., 44 see also Scott, D.C., 43 see also Trost, L.M.B., 52 Bidwell, A., 43 bifidum (Bengtsson), Procloeon, 27 bilineata (Say), Hexagenia, 32, 49 Bohart, R.M., 98, 103 Bohle, H.W., 31,46 borealis Eaton, Metretopus, 35 Brachycercus Curtis, 29 Bretschko, G., 27, 43 Brittain, J.E., 27, 34 Brown, W.L.jr., 96, 119 Bryson, R.A., 95 Butler, A.G., 8 C.A., Menzie, 48 Caenidae, 27, 29, 47 Caenis horaria (L.), 29 Caenis macrura Stephens, 52 386 Index to Volume 18 Caenis rivulorum Eaton, 47 Caenis robust a Eaton, 52 Caenis Stephens, 27, 29, 54 Callibaetis Eaton, 27, 54 Callibaetis floridanus Banks, 52 Campsuarus Eaton, 54 canadense (Walker), Stenacron , 33 Cazier, M.A., 93 Centroptilum Eaton, 27 Centroptilum elsa Traver, 47 Centroptilum luteolum (Muller), 27 Centroptilum pennulatum (Eaton), 52 Cerceris Latareille, 103 Chadwick, H.W, 94 Chapman, R.N, 102 Choroterpes mexicanus Allen, 34 Christiansen, E.A., see Whitaker, S.H., 93 Cicindela , 123 Cicindelidae, 123 Cinygma Eaton, 33, 54 Cinygmula McDunnough, 33 Clayton, J.S., see Mitchell, J., 95 Clifford, H.F., 27, 34, 46, 47, 52, 54 Cloeon dipterum (L.), 27 Cloeon Leach, 27, 45 Cloeon simile Eaton, 27 Coleman, M.J., , 47 see also Hynes, H.B.N., 47 Colletidae, 104 Coloburiscus humeralis (Walker), 44 Cook, E.F., see Hall, R.J., 44 Corbet, P.S., 43 Corbet, S.A., 43, 52 Coupland, R.T., , 94 see Hulett, G.K., 94 crabroniformis Smith, Philanthus , 93, 99 cryptandrus A. Gray, Sporobolus , 94 cupida (Say), Leptophlebia, 34, 47, 53 D’Orville, H., 8 dalecarlica Bengtsson, Heptagenia, 33 Dalke, P.D., see Chadwick, H.W., 94 danica Muller, Ephemera , 31, 52 Dannella Edmunds, 54 Darwin, C., 2,5,6, 9, 10, 12 Darwin, E., 4 Darwin, F., 7, 9 Dawkins, R., 12 de Beer, G., 5 debilis (Walker), Paraleptophlebia, 34, 45 deficiens (Morgan), Serratella, 47 Demoulin, G., 52 diaphana Navas, Rithrogena , 32 dipterum (L.), Cloeon , 27 dispar (Curtis), Ecdyonurus, 33 Dix, R.L., see Hulett, G.K., 94 Dolania americana Edmunds and Traver, 44 dorothea Needham, Ephemerella, 47 Drunella Needham, 31 Dumas, J., see Lavandier, P., 21, 27 eatoni Brodskij, Ametropus, 44 Ecdyonurus dispar (Curtis), 33 Ecdyonurus Eaton, 32 Ecdyonurus lateralis (Curtis), 33 Ecdyonurus quadrilineatus Landa, 47 Ecdyonurus submontanus Klapalek, 33 Ecdyonurus torrent is Kimmins, 33 Ecdyonurus venosus (Fabricius), 33 Edmunds, C.H., see Edmunds, G.F. Jr., 52 Edmunds, F.H., 95 Edmunds, G.F., 21 Edmunds, G.F. Jr., 44, 52 Elassoneuria insulicola Demoulin, 35 Elliott, J.M., 30,31,46 elsa Traver, Centroptilum , 47 Epeorus Eaton, 33 Ephemera danica Muller, 31, 52 Ephemera Linneaeus, 31, 32 Ephemera simulans Walker, 31 Ephemerella , 21 Ephemerella aurivillii Bengtsson, 31 Ephemerella dorothea Needham, 47 Ephemerella ignita (Poda), 23, 30, 31, 45, 46 Ephemerella mucronata Bengtsson, 31, 47 Index to Volume 1 8 387 Ephemerella notata Eaton, 47 Ephemerella subvaria McDunnough, 47 Ephemerella Walsh, 30, 31, 45, 54 Ephemerellidae, 29, 30, 45, 47, 54 Ephemeridae, 31, 45, 54 Ephoron album (Say), 43 Ephoron virgo (Oliver), 43 Eucerceris Cresson, 103 Eumenidae, 104 Eurylophella funeralis (McDunnough), 31 Eurylophella Tiensuu, 31 Eurylophella versimilis (McDunnough), 47 Euthypolciidae, 44 Evans, H.E., 93, 96, 99, 102, 103, 104, 105, 106, 112 Fiance, S.B., 31 Fisher, R.A., 12 Fjellheim, A., see Andersen, T., 46 Flannagan, J.F., 32 Flint, R.F., 94, 95, 121 floridanus Banks, Callibaetis, 52 Flowers, R.W., 53 Formica Linnaeus, 103 fragilis Albarda, Ametropus, 44 Freitag, R., 93, 94, 122 Fremling, C.R., , 32 see also Thomforde, L.L., 32 Frenzel, B., 94 Frey, D.G., see Wright, H.E. jr., 94 frigidus (F. Smith), Aphilanthops, 103, 106 funeralis (McDunnough), Eurylophella , 31 fuscatus (L.), Baetis , 26 fuscogrisea (Retzius), Heptagenia, 47 George, W.A., 8 germanica Eaton, Rhithrogena , 47 gibbosus (Fabricius), Philanthus , 93, 94, 99 gloriosus Cresson, Philanthus , 94 Gordon, E.L., 34 Grissell, E.E., see Bohart, R.M., 98, 103 Habroleptoides modesta (Hagen), 34 Habroleptoides Schoenemund, 54 Habrophlebia Eaton, 34 Habrophlebia lauta McLachlan, 34 Halictidae, 93, 104 Halictus rubicundus Christ., 103, 106 Hall, R.J., 44 Hamilton, A.L., 48 Hamilton, H., see also Clifford, H.F., 30, 34, 47 Hartland-Rowe, R., 43 Harvey, R.S., 44 Headley, F.W., 9 Heptagenia dalecarlica Bengtsson, 33 Heptagenia fuscogrisea (Retzius), 47 Heptagenia lateralis (Curtis), 33 Heptagenia sulphurea (Muller), 33, 53 Heptagenia Walsh, 33 Heptageniidae, 32, 33, 45, 47, 54 hercynia Landa, Rhithrogena , 47 Heterocloeon McDunnough, 54 Hexagenia bilineata (Say), 32, 49 Hexagenia limbata Serville, 32 Hexagenia Walsh, 31, 32, 45, 54 Hilsenhoff, W.L., see Flowers, R.W., 53 Hirsch, A., see Scott, D.C., 43 Homoeoneuria Eaton, 54 horaria (L.), Caenis, 29 Hudson, P.L., 32 Huff, B.L., see McCafferty, W.P., 33 Hulett, G.K, 94 humeralis (Walker), Coloburiscus, 44 Humpesch, U.H., 33, 46 Hunt, B.P., 32 Huxley, J.S., 1 1 hymenoides Ricker, Oryzopis , 94 Hynes, H.B.N., 47 Hynes, J.D., 21 lersel, J.J.A. van, 1 12 ignita (Poda), Ephemerella , 23, 30, 31, 45,46 388 Index to Volume 18 lilies, J., 26, 31 incertus (Traver), Tortopus, 43 inopinatus Eaton, Ameletus, 43, 53 insulicola Demoulin, Elassoneuria, 35 interpunctatum (Say), Stenacron, 33 Ironodes nitidus (Eaton), 33 Irvine, W., 4 Isonychia Eaton, 43, 54 J., Hunter, 3 Jensen, S.L., see also Edmonds, G.F., 21 Kelly, E.G., see Chapman, R.N., 102 Killins, B.A., see Clifford, H.F., 30, 34 Killnis, B.A., see Clifford, H.F., 47 King-Hele, D., 4 Kirby, W., 3, 4, 5 Klassen, R.W., 95 Kuhn, T., 5 lacustris (Eaton), Siphlornurus , 43 lanceolata Pursh, Psorealea , 94 Landa, V., 21, 31, 33,-43, 47 Lapchin, L., see Neveu, A., 23, 26, 31, 32 Larsen, R., see Andersen, T., 46 lateralis (Curtis), Ecdyonurus , 33 lateralis (Curtis), Heptagenia , 33 lauta McLachlan, Habrophlebia, 34 Lavandier, P., 21, 27 Lehmkuhl, D.M., 33, 34, 47, 53 Leptohyphes Eaton, 54 Leptophlebia cupida (Say), 34, 47, 53 Leptophlebia marginata (L.), 34, 47, 53 Leptophlebia vespertina (L.), 34, 49, 53 Leptophlebia Westwood, 34, 53 Leptophlebiidae, 33, 34, 45, 47, 54 Lesser, M., 5 limbata Serville, Hexagenia , 32 Lin, C.S., see Evans, H.E., 104 Lithobranchia McCafferty, 54 littoralis Turner, Bembix , 1 12 Love, K., 94, 95 Lorenz, K., 93 loyalae Navas, Rhithrogena, 32 luteolum (Muller), Centroptilum, 27 Macan, T.T., 17 Macan, T.T., 27, 52 macani bundyae, Baetis, 26 macani Kimmins, Baetis , 26 MacArthur, R.H., 118 Macdunnoa Lehmkuhl, 33 Mackey, A.P., 52 macrura Stephens, Caenis, 52 maculatus Kimmins, Tricorythus , 44 major (Klapalek), Torleya, 31 manuelito new subspecies, Philanthus, 121, 122, 123 Marchant, J., 7 marginata (L.), Leptophlebia , 34, 47, 53 Martin, P.S., 94, 122 Masaki, S., 46 Matthews, R.W., see Evans, H.E., 96, 112 Maudsley, R., see Macan, T.T., 27 Mayer-Oakes, W.J., 95 McCafferty, W.P., , 33, 43 see also Morihara, D.K., 20 McClure, R.G., 34 McLean, J.A., 44 Meier, P.G., see Bartholomae, P.G., 43 Mengel, R.M., 94 Menke, A.S., see Bohart, R.M., 103 Metretopodidae, 34, 54 Metretopus borealis Eaton, 35 Metretopus Eaton, 35, 54 mexicanus Allen, Choroterpes, 34 Mickel, C.E., see Chapman, R.N., 102 Microbembex, 104, 123 Microbembex monodonta (Say), 104 Miller, G.E., see Chapman, R.N., 102 Minshall, G.W., see Newell, R.L., 44 minutus Traver, Tricorythodes, 44 Index to Volume 18 389 Mitchell, J., 94, 95 Mivart, St. G., 9, 1 1 modest a (Hagen), Habroleptoides, 34 monodonta (Say), Microbembex, 104 Montgomery, T., 10, 1 1 Morgan, T.H., 1 1 Morihara, D.K., 20 morrisoni (Banks), Rhithrogena, 47 Mortenson, M.A., see Cazier, M.A., 93 Moss, H.C., see Mitchell, J., 95 mucronata Bengtsson, Ephemerella, 31, 47 muticus (L.), Baetis, 26 myops (Walsh), Potamanthus , 43 Nageli, C., 12 neavei McDunnough, Ametropus , 44 Neoephemera McDunnough, 54 Neoephemeridae, 44, 54 Neoephemeropsis Ulmer, 44 Neveu, A., 23,31,32 Newell, R.L., 44 nitens (Banks), Philanthus, 93, 113 nitidus (Eaton), Ironodes, 33 notata Eaton, Ephemerella , 47 obesa (Say), Baetisca, 44 occidentalis Eaton, Siphlornurus , 47 Oligoneuriella rhenana (Imhoff), 35, 46 Oligoneuriidae, 35, 45, 54 Oniscigaster wakefieldi McLachlan, 43 ornatum Tshernova, Procloeon, 27 oronti Alouf, Prosopistoma , 44 Oryzopis hymenoides Ricker, 94 Otto, C., see Andersen, T., 46 pacificus Cresson, Philanthus , 93 Paley, W„ 4 Palingenia Burmeister, 31, 44, 54 Palingeniidae, 44, 54 Paraleptophlebia debilis (Walker), 34, 45 Paraleptophlebia Lestage, 34, 54 Paraleptophlebia submarginata (Stephens), 34 Parameletus Bengtsson, 43 Parker, J.R., see Chapman, R.N., 102 Parvella , 104 Peckham, E.G., see Peckham, G.W., 1 1, 93 Peckham, G.W., 11,93 pennulatum (Eaton), Centroptilum , 52 Pentagenia Walsh, 54 Pescador, M.L., 35, 44 Peters, W.L., see also Pescador, M.L., 35, 44 Petr, T., 43 Philanthus , 93, 94, 98, 103, 104, 124 Philanthus albopilosus Cresson, 94, 95, 96, 98,99, 103, 104, 105, 106, 112, 113, 116, 118, 119, 120, 121, 123, 124 Philanthus crabroniformis Smith, 93, 99 Philanthus Fabricius, 103, 113 Philanthus gibbosus (Fabricius), 93, 94, 99 Philanthus gloriosus Cresson, 94 Philanthus manuelito new subspecies, 121, 122, 123 Philanthus nitens (Banks), 93, 1 13 Philanthus pacificus Cresson, 93 Philanthus politus Say, 113 Philanthus psyche Dunning, 94, 123 Philanthus pulcher Dalla Torre, 93 Philanthus solivagus Say, 94 Philanthus triangulum Fabricius, 93 Philanthus ventilabris Fabricius, 94 picteti Sowa, Rithrogena, 32 Pleskot, G., 26 Poliomintha, 104 politus Say, Philanthus , 113 Polymitarcidae, 43, 54 Potamanthidae, 43 Potamanthodes Ulmer, 43 Potamanthus myops (Walsh), 43 Potamanthus Picket, 43 Poulton, E.B., 12 Povilla adusta rsfavas, 43, 45, 52 Pritchard, G., 54 Proboscidoplocia sikorai Vayssiere, 44 Procloeon bifidum (Bengtsson), 27 Procloeon ornatum Tshernova, 27 390 Index to Volume 18 Prosopistoma Latreille, 44, 54 Prosopistoma oronti Alouf, 44 Prosopistomatidae, 44, 54 Pseudiron .McDunnough, 33, 54 Pseudocloeon Klapalek, 27 Psorealea lanceolata Pursh, 94 psyche Dunning, Philanthus, 94, 123 pulcher Dalla Torre, Philanthus, 93 Puthz, V., 21 quadrilineatus Landa, Ecdyonurus , 47 Ramanankasina, R.E., 35, 44 Rau, N., see Rau, P., 93 Rau, P., 93 Reinhard, E.G., 93, 99, 104 rhenana (Imhoff), Oligoneuriella , 35, 46 Rhithrogena Eaton, 32, 33, 54 Rhithrogena germanica Eaton, 47 Rhithrogena hercynia Landa, 47 Rhithrogena loyalae Navas, 32 Rhithrogena morrisoni (Banks), 47 Rhithrogena semicolorata (Curtis), 32 rhodani (Pictet), Baetis , 23, 26 Richardson, M.Y., 33 Rithrogena diaphana Navas, 32 Rithrogena picteti Sowa, 32 rivulorum Eaton, Caenis, 47 Robertson, M.R., see Clifford, H.F., 52 robust a Eaton, Caenis, 52 rogersi Berner, Baetisca, 44 Romanes, J.G., 9, 10 Rosenberg, D., 17 Ross, H.E., 94, 122 rubicundus Christ., Halictus, 103, 106 Rutter, R.P., 32 scambus Eaton, Baetis, 26, 45 Scott, D.C., 43 Sellick, R.D., see Corbet, S.A., 43, 52 semicolorata (Curtis), Rhithrogena, 32 Serratella deficiens (Morgan), 47 Shapiro, A.M., 52 sikorai Vayssiere, Proboscidoplocia, 44 simile Eaton, Cloeon, 27 simulans Walker, Ephemera, 31 Siphlonisca Needham, 43 Siphlonuridae, 43, 47, 54 Siphloplecton basale (Walker), 34 Siphloplecton Clemens, 34 Siphloplecton speciosum Traver, 34, 35 Siphlornurus Eaton, 43 Siphlornurus lacustris (Eaton), 43 Siphlornurus occidentalis Eaton, 47 Smith, H.T.U., , 122 see also Thorp, J., 122 solivagus Say, Philanthus, 94 Sowa, R., 26, 32 speciosum Traver, Siphloplecton, 34, 35 Spence, W., see Kirby, W., 3, 4, 5 Sphecidae, 104 Spinadis Edmunds and Jensen, 54 Sporobolus cryptandrus A. Gray, 94 Steinbeck, J., 7 Stenacron canadense (Walker), 33 Stenacron interpunctatum (Say), 33 Stenacron Jensen, 54 Stenonema Traver, 33, 54 Stenonema tripunctatum (Banks), 33 Stewart, K.W., see McClure, R.G., 34 Stolzmann, J., 10, 1 1 Strnadtmann, R.W., 98 submar ginata (Stephens), Paraleptophlebia, 34 submontanus Klapalek, Ecdyonurus, 33 subvaria McDunnough, Ephemerella, 47 sulcatus (L.), Acilius, 3 sulphurea (Muller), Heptagenia, 33, 53 Svensson, B., 31, 52 Swanson, G.A., , 32 see also Hudson, P.L., 32 Sweeney, B.W., , 47 see also Harvey, R.S., 44 Tamarix, 104 Tarter, D.C., see Richardson, M.Y., 33 Tauber, C.A., see Tauber, M .J., 19 Tauber, M .J., 19 texana Cresson, Bembix, 1 12 Index to Volume 18 391 Thalerosphyrus Eaton, 33 Thibault, M., 23,31,32, 47 Thomforde, L.L., 32 Thorp, J., 122 tiger beetles, 123 Tinbergen, N., 93, 104, 105 tinctus Kimmins, Tricorythus, 44 Tjonneland, A., 44 Torleya belgica Lestage, 31 Torleya major (Klapalek), 31 torrentis Kimmins, Ecdyonurus, 33 Tortopus incertus (Traver), 43 Tortopus Needham and Murphy, 54 Traver, J. R., 27 Traverella Edmunds, 54 triangulum Fabricius, Philanthus, 93 tricaudatus Dodds, Baetis, 26 Tricorythidae, 44, 45, 54 Tricorythodes atratus McDunnough, 44 Tricorythodes minutus Traver, 44 Tricorythodes Ulmer, 44 Tricorythus maculatus Kimmins, 44 Tricorythus tinctus Kimmins, 44 tripunctatum (Banks), Stenonema , 33 Trost, 52 Ulfstrand, S., 26, 33, 35, 53 vagans McDunnough, Baetis , 26 Van Dyck, W., 10, 11 Vannote, R.L., see also Sweeney, B.W., 47 see Harvey, R.S., 44 see Sweeney, B.W., 47 venosus (Fabricius), Ecdyonurus , 33 ventilabris Fabricius, Philanthus , 94 vernus (Curtis), Baetis, 26 versimilis (McDunnough), Eurylophella , 47 vespertina (L.), Leptophlebia , 34, 49, 53 Vignes, J.C., see Neveu, A., 23,26,31,32 virgo (Oliver), Ephoron , 43 Vorzimmer, P., 7 Vuilleumier, B.S., 118 wakefieldi McLachlan, Oniscigaster, 43 Wallace, A.R., , 5, 8, 10, 11 see also Darwin, C., 5, 6 Wallace, Alex, 8 Waters, T.F., 19,47,48 Weir, J., 8 Weismann, A., 12 Whelan, K.F., 31,53 Whitaker, S.H., 93 Whitehead, D.R., 120 Williams, G.C., 12 Willis, H.J., 120, 123 Willoughby, N.G., see Corbet, S.A., 43, 52 Wilson, E.O., see Brown, W.L. jr., 95, 96, 1 19 see MacArthur, R.H., 1 18 Winterbourn, M.J., 34 Wise, E.J., 33 Wisely, B„ 44 Wissing, T.E., see Rutter, R.P., 32 Wright, H.E.jr., 94, 95 Zelt, K.A., see Clifford, H.F., 52 Publication of Quaestiones Entomologicae was started in 1965 as part of a memorial project for Professor E. 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