Diptera: Empididae)

Diversity and Ecology of Northern Nearctic Rhamphomyia (Diptera: Empididae)

Élodie Anne Vajda Dept. of Natural Resource Sciences McGill University, Montreal August, 2015

A thesis submitted to McGill University in partial fulfillment of the requirements of the degree of Master of Science

ABSTRACT The dance fly genus Rhamphomyia Meigen (Diptera: Empididae) is one of the most dominant and species-rich insect genera in the Nearctic. However, little is known about their spatial distribution and species-richness, making it difficult to assess changes in northern Rhamphomyia as a result of environmental change. This project examines taxonomic (species richness) and ecological (body size) diversity of northern Nearctic Rhamphomyia. An inventory of Rhamphomyia species at 12 sites across northern Canada (4 boreal, 4 low arctic, 4 high arctic) identified 65 species, many of which are undescribed. The highest species richness was in the low arctic, and in particular, in subarctic Yukon Territory. Nearctic Rhamphomyia species usually span the boreal-low arctic biomes, and substantially fewer species are in the high arctic. The first key to the 21 Rhamphomyia species of the Canadian arctic Archipelago, Greenland, and Iceland is provided along with diagnoses of each species. Rhamphomyia vary in body size across latitude. We examined latitudinal and other spatial effects on body size of Nearctic Rhamphomyia. Body size significantly increases with latitude, but only at species-level; smaller species drop out of species assemblages as latitude increases. Rhamphomyia species of the arctic islands are significantly larger than those on the mainland. The Northwest Passage could act as a physical barrier preventing dispersal of smaller Rhamphomyia species to the arctic islands post-glaciation. Other environmental factors of the arctic islands, such as dryness and low temperatures, might also impede smaller Rhamphomyia from successfully colonizing the arctic islands.

RÉSUMÉ Le genre Rhamphomyia (Diptera: Empididae) est un des genre les plus dominant et riche en espèces de l’arctique du Nouveau Monde. Mais l’on connait peu de chose sur leur distirbution géographique et leur richesse en espèces. Il est donc difficile d'évaluer l'évolution des Rhamphomyia du Nord due au changement climatique. Ce projet examine la diversité taxonomique (diversité de l'espèce) et écologique (taille du corps) du genre Rhamphomyia du Nord Arctique. Un inventaire des espèces de Rhamphomyia recueilli dans 12 sites à travers le Nord du Canada (4 boréales, 4 Bas-Arctique, et 4 Extrême-Arctique) rassemble 65 espèces, dont beaucoup ne sont pas décrites. La region du Nord canadien ayant la plus haute richesse en espèce est le bas-arctique, et surtout, le Yukon. Les espèces de Rhamphomyia ont tendance à se concentrer dans les régions boréales et dans le Bas- Arctique, mais se font plus rares dans l’Extrême-Arctique. La première clé d’identification (avec diagnostiques) est écrite pour les 21 espèces Rhamphomyia se trouvant dans les îles de l’Arctique, dans le Groenland et l’Islande. La taille du corp des Rhamphomyia varie avec la latitude. Nous avons examiné les effets de la latitude, ainsi que d’autres effets spatiaux sur la taille du corp des Rhamphomyia du Néarctique. La taille du corp augmente en accord avec la latitude seulement au niveau des espèces; les espèces plus petites sont graduellement perdues avec l’augmentation de la latitude. Aussi, les Rhamphomyia des îles de l’Arctique sont plus larges que les Rhamphomyia du territoire continental. Le Northwest Passage pourrait constituter un obstacle physique empêchant les plus petites espèces de Rhamphomyia de migrer du territoire continental aux îles, à la suite de la fonte des glaces. D’autres facteurs environnementaux, tels que l’aridité et les températures extêmement basses des îles de l’Arctique, pourraient aussi empêcher les plus petites Rhamphomyia de coloniser les îles de l’arctique.

TABLE OF CONTENTS Abstract...... ii Résumé...... iii List of tables...... vi List of figures...... vii List of appendices...... xi Acknowledgements...... xii Preface...... xiii Contributions of authors...... xiv CHAPTER 1. Introduction and literature review...... 1 Arctic and its impacts on arthropod life...... 1 Arctic arthropods and climate change...... 2 Relevance of arthropod studies...... 3 Taxonomy of Empididae and of the genus Rhamphomyia...... 3 Biology of Rhamphomyia...... 4 Functional diversity of Rhamphomyia...... 6 Objectives...... 9 Connecting statement...... 11 CHAPTER 2. Faunal Inventory of Rhamphomyia in Northern Canada...... 12 Materials and methods...... 12 Study sites...... 12 Sampling methods...... 12 Specimen preparation and identification...... 13 Results...... 14 Inventory...... 14 Discussion...... 15 Connecting statement...... 22 CHAPTER 3. Key to species of Rhamphomyia of the Arctic Islands...... 23 Materials and methods...... 24 Results and discussion...... 25 Key to male species of Rhamphomyia of the Arctic Islands (incl. GL, IC)...... 25

Species descriptions and diagnoses...... 29 Connecting statement...... 132 CHAPTER 4. Latitudinal body size patterns in a diverse genus of arctic insects...133 Introduction...... 133 Materials and methods...... 134 Sample sites and collecting...... 135 Specimen identification...... 135 Specimen measurement...... 136 Environmental data...... 136 Analyses...... 137 Results...... 138 Diversity of Rhamphomyia...... 138 Latitudinal trends...... 139 Discussion...... 139 CHAPTER 5. General discussion and conclusions...... 145 References...... 147

LIST OF TABLES Table 2.1. Twelve most abundant morphospecies of Rhamphomyia, number of male specimens identified, and sites at which they were collected by the Northern Biodiversity Program (BI: Banks Island; CB: Cambridge Bay; CH: Churchill; DH: Dempster Hwy; EI: Ellesmere Island; GB: Goose Bay; IQ: Iqaluit; K: Kugluktuk; M: Moosonee; NW: Norman Wells; SC: Schefferville; Y: Yellowknife)...... 18 Table 2.2. Number of Rhamphomyia specimens, species-richness and number of species unique to each Northern Biodiversity Program site. Site codes as in Table 2.1...... 19

LIST OF FIGURES Figure 2.1. Northern Biodiversity Program sampling sites in boreal boreal (circles), low arctic (squares), and high arctic (triangle) biomes. (BI: Banks Island; CB: Cambridge Bay; CH: Churchill; DH: Dempster Hwy; EI: Ellesmere Island; GB: Goose Bay; IQ: Iqaluit; K: Kugluktuk; M: Moosonee; NW: Norman Wells; SC: Schefferville; Y: Yellowknife)...... 20 Figure 2.2. Rhamphomyia species unique to each biome and shared between biomes in Northern Biodiversity Program sampling sites (HA: high arctic; LA: low arctic; B: boreal)...... 21 Figures 3.1-3.4. Rhamphomyia herschelli, posterior and lateral view of projections of pregenital segments (1, 2), Rhamphomyia herschelli, hind tibia, basitarsus, and tarsomeres (3), Rhamphomyia sp.12, hind femur, and tibia (4)...... 103 Figures 3.5-3.7. Rhamphomyia hirtula, hind leg (5), Rhamphomyia hirtula, fore leg (6), Rhamphomyia hirticula, hind tibia, basitarsus, and tarsomeres (7)...... 104 Figures 3.8-3.10. Rhamphomyia (Ctenempis) calvimontis, hind leg (8), fore leg (9), pregenital segments (male) (10)...... 105 Figures 3.11-3.12. Rhamphomyia (Pararhamphomyia) diversipennis sp.1, mid leg (11), Rhamphomyia (Pararhamphomyia) diversipennis sp.1, thorax, dorsal view (12)……………………………………………………………………………...106 Figures 3.13-3.16. Rhamphomyia (Pararhamphomyia) tipularia sp. 1, hind leg, lateral view (13), Rhamphomyia (Pararhamphomyia) tipularia sp. 4, hind leg, lateral view (14), Rhamphomyia (Pararhamphomyia) tipularia sp. 4, male genitalia showing spindle-like structure of epandrium (15), Rhamphomyia (Pararhamphomyia) tipularia sp. 1, fore basitarsus, lateral view (16)...... 107 Figure 3.17. Rhamphomyia (Pararhamphomyia ?) simplex, hind leg, lateral view...... 108 Figures 3.18-3.19. Rhamphomyia (Pararhamphomyia) pusilla sp. 7, hind leg, lateral view (18), Rhamphomyia (Pararhamphomyia) pusilla sp. 7, mid leg, lateral view (19)……………………………………………………………………………...108 Figures 3.20. Rhamphomyia (Pararhamphomyia) fuscula gp. sp.1, hind leg, lateral view……………………………………………………………………………..109 Figures 3.21-3.25. Rhamphomyia (Dasyrhamphomyia) nigrita, hind leg, lateral view (21),

Rhamphomyia (Dasyrhamphomyia) zaitsevi, hind leg, lateral view (22), Rhamphomyia (Dasyrhamphomyia) sp.9, hind tibia, lateral view (23), Rhamphomyia (Dasyrhamphomyia) sp.9, mid leg, lateral view (24), Rhamphomyia (Dasyrhamphomyia) sp.3, mid leg, lateral view (25)...... 110 Figures 3.26. Rhamphomyia (Eorhamphomyia) nigrolineata gp. sp.4, hind leg, lateral view……………………………………………………………………………..111 Figures 3.27-3.30. Rhamphomyia (Pararhamphomyia) lapponica gp. sp.8, hind leg, lateral view (27), Rhamphomyia (Pararhamphomyia) lapponica gp. sp.9, hind leg, lateral view (28), Rhamphomyia (Pararhamphomyia) lapponica gp. sp.11, hind leg, lateral view (29), Rhamphomyia (Pararhamphomyia) hoeli, hind leg, lateral view (30)...... 112 Figures 3.31-3.33. Rhamphomyia (Pararhamphomyia) filicauda, hind leg, lateral view (31), Rhamphomyia (Pararhamphomyia) filicauda, hind femur, tibia, and basitarsus, lateral view (32), Rhamphomyia (Pararhamphomyia) filicauda, mid leg, lateral view (33)...... 113 Figures 3.34-3.36. Rhamphomyia (Pararhamphomyia) ursinella, hind leg, lateral view (34), Rhamphomyia (Pararhamphomyia) caudata gp. sp.7, hind leg, lateral view (35), Rhamphomyia (Pararhamphomyia) ursinella, fore leg, lateral view (36)..114 Figures 3.37-3.39. Rhamphomyia sp.12, lateral view of male genitalia (37), Rhamphomyia herschelli, lateral view of male genitalia (setae not shown) (38), Rhamphomyia herschelli,, lateral view of pregenital segments 7 and 8 (39)...... 115 Figures 3.40-3.41. Rhamphomyia hirtula, lateral view of male genitalia (40), Rhamphomyia hirticula, lateral view of male genitalia (41)...... 116 Figures 3.42-3.43. Rhamphomyia (Ctenempis) calvimontis, lateral view of male genitalia (42), Rhamphomyia (Ctenempis) calvimontis, pregenital segments (male) (43)...... 117 Figure 3.44. Rhamphomyia (Pararhamphomyia) diversipennis sp.1, male genitalia, lateral view...... 118 Figures 3.45-3.46. Rhamphomyia (Pararhamphomyia) tipularia gp. sp. 1, male genitalia, lateral view (45), Rhamphomyia (Pararhamphomyia) tipularia gp. sp. 4, male genitalia lateral view (46)...... 119

Figures 3.47-3.49. Rhamphomyia (Pararhamphomyia ?) simplex, male genitalia, lateral view (47), Rhamphomyia (Pararhamphomyia) pusilla sp.7, male genitalia, lateral view (48), Rhamphomyia (Pararhamphomyia) fuscula gp. sp.1, male genitalia, lateral view (49)...... 120 Figure 3.50-3.52. Rhamphomyia (Dasyrhamphomyia) nigrita, male genitalia, dorsal view (50), Rhamphomyia (Dasyrhamphomyia) sp.9, male genitalia, dorsal view (51), Rhamphomyia (Dasyrhamphomyia) zaitsevi, male genitalia, dorsal view (52)...... 121 Figures 3.53-3.55. Rhamphomyia (Dasyrhamphomyia) nigrita, male genitalia, lateral view (53), Rhamphomyia (Dasyrhamphomyia) zaitsevi, male genitalia, lateral view (54), Rhamphomyia (Dasyrhamphomyia) sp.9, male genitalia, lateral view (55)...... 122 Figures 3.56-3.57. Rhamphomyia (Eorhamphomyia) nigrolineata gp. sp.4, male genitalia, lateral view (56), Rhamphomyia (Eorhamphomyia) nigrolineata gp. sp.4, cerci and phallus tip, dorsal view (57)...... 123 Figures 3.58-3.61. Rhamphomyia (Pararhamphomyia) lapponica gp. sp.8, male genitalia, lateral view (58), Rhamphomyia (Pararhamphomyia) lapponica gp. sp.9, male genitalia, lateral view (59), Rhamphomyia (Pararhamphomyia) lapponica gp. sp.11, male genitalia (60), Rhamphomyia (Pararhamphomyia) hoeli, male genitalia, lateral view (61)...... 124 Figures 3.62-3.64. Rhamphomyia (Pararhamphomyia) filicauda, male genitalia, lateral view (62), Rhamphomyia (Pararhamphomyia) ursinella, male genitalia, lateral view (63), Rhamphomyia (Pararhamphomyia) caudata gp. sp.7 (64)...... 125 Figure 3.65. Rhamphomyia sp.12 (circle), Rhamphomyia herschelli (triangle), distributions...... 126 Figure 3.66. Rhamphomyia hirticula (circle), Rhamphomyia hirtula (triangle), distributions………………………………………………………………..…...126 Figure 3.67. Rhamphomyia (Ctenempis) calvimontis, distribution...... 127 Figure 3.68. Rhamphomyia (Pararhamphomyia) diversipennis sp.1, distribution...... 127 Figure 3.69. Rhamphomyia (Pararhamphomyia) tipularia sp.1 (circle), Rhamphomyia (Pararhamphomyia) tipularia sp.4 (square), distributions...... 128

Figure 3.70. Rhamphomyia (Pararhamphomyia) simplex, distribution...... 128 Figure 3.71. Rhamphomyia (Pararhamphomyia) pusilla gp. sp.7, distribution...... 129 Figure 3.72. Rhamphomyia (Pararhamphomyia) fuscula gp. sp.1, distribution...... 129 Figure 3.73. Rhamphomyia (Dasyrhamphomyia) nigrita (circle), Rhamphomyia (Dasyrhamphomyia) sp.9 (square), Rhamphomyia (Dasyrhamphomyia) zaitsevi (triangle), distributions...... 130 Figure 3.74. Rhamphomyia (Eorhamphomyia) nigrolineata gp. sp.4, distribution...... 130 Figure 3.75. Rhamphomyia (Pararhamphomyia) lapponica gp. sp.8 (circle), Rhamphomyia (Pararhamphomyia) lapponica gp. sp.9 (square), Rhamphomyia (Pararhamphomyia) lapponica gp. sp.11 (triangle), Rhamphomyia (Pararhamphomyia) hoeli (hexagon), distributions...... 131 Figure 3.76. Rhamphomyia (Pararhamphomyia) ursinella (circle), Rhamphomyia (Pararhamphomyia) caudata gp. sp.7 (triangle), Rhamphomyia (Pararhamphomyia) filicauda (square), distributions...... 131 Figure 4.1. Relationship between fore tibia length and overall body size in male Rhamphomyia from the Canadian Arctic...... 142 Figure 4.2. Relationship between Nearctic Rhamphomyia species mean tibiae lengths and sampling locality latitude point...... 143 Figure 4.3. Mean tibia length for each of the twelve sampling localities...... 144

LIST OF APPENDICES Appendix 1. Inventory of Rhamphomyia morphospecies (male specimens only) collected by the Northern Biodiversity Program in 12 sites in northern Canada (2010, 2011). Appendix 2. Mean body size at each site of species spanning four or more sites; a. R. nigrita; b. R. lapponica sp.2; c. R. dispar gp. sp.10; d. R. rivalis; e. R. nigrolineata; f. R. fuscula gp. sp.1; g. R. hirticula gp. sp.2.

ACKNOWLEDGEMENTS I would like to thank many people for their valuable help and support throughout this thesis. This project would not have been possible without my supervisory committee. Thank you to my supervisor, Dr. Terry Wheeler for giving me the opportunity to take on a research project during my undergraduate degree, which later became my Master’s project, and for his help and encouragement throughout my graduate degree. A big thank you to Dr. Brad Sinclair, and Dr. Chris Buddle. Dr. Brad Sinclair was consistently very committed to this project and provided essential help with the taxonomy component of this project. Dr. Chris Buddle provided key advice and support for the ecology component of my thesis. Thank you to my amazing lab mates, Sabrina Rochefort, Heather Cumming, Stéphanie Boucher, Anna Solecki, and Amélie Grégoire-Taillefer, We have worked well together, and have never stopped helping each other out in the lab. Many thanks to Shaun Turney for advice and help with statistical analyses, and to the Biodiversity Institute of Ontario for DNA barcoding. Thank you to the CNC Diptera Unit crew for always making me feel welcomed, and for allowing me to use their equipment and insect collection. Dr. Jeff Cumming provided valuable suggestions and ideas for the taxonomy chapter of my thesis. Dr. Scott Brooks kindly taught me how to use the camera. Fieldwork was carried out under permits from the governments of the Northwest Territories, Nunavut, and Yukon Territory, Parks Canada, Tombstone Territorial Park, and with the permission and cooperation of northern First Nations. Major funding was provided by a Natural Sciences and Engineering Research Council of Canada (NSERC) Strategic Project Grant (to C.M. Buddle, D.C Currie and T.A. Wheeler), NSERC Discovery Grants and Northern Research Supplement to T.A. Wheeler, and the Polar Continental Shelf Project. Finally I wish to address a special thank you to the Cumming family, Shakira Fulton- McLean, Ian Vogel, and to my family, for continuously supporting me, and for always listening to my talking about flies.

PREFACE This thesis consists of 5 chapters. Chapters 2, 3, and 4, are manuscripts to be submitted for publication to peer-reviewed journals Chapter 1 This is an introduction and literature review. Chapter 2 Vajda, É. A., Sinclair, B.J., and Wheeler, T.A. Faunal inventory of Rhamphomyia (Diptera: Empididae) in northern Canada. (for submission) Chapter 3 Vajda, É. A., Sinclair, B.J., and Wheeler, T.A. Key to the Rhamphomyia (Diptera: Empididae) of the Canadian Arctic Islands, Greenland and Iceland. (for submission) Chapter 4 Vajda, É. A., and Wheeler, T.A. Latitudinal body size patterns in a diverse genus of arctic insects (for submission). Chapter 5 This chapter is a general discussion and summary of the thesis with suggestions for future research. Disclaimer: Taxonomic and nomenclatural acts in this thesis are not to be considered as published according to the International Code of Zoological Nomenclature until they are published in the peer-reviewed literature.

CONTRIBUTIONS OF AUTHORS Élodie A. Vajda conducted specimen preparation, identifications, descriptions, drawings and photos, specimen measurements, statistical analyses, and writing. Dr. Brad Sinclair assisted in taxonomic training, specimen identification and drawing techniques. Dr. Terry A. Wheeler provided training, research funding, access to laboratory equipment and work space, in addition to advising and editing of manuscripts.

Chapter 1. Introduction and Literature Review

Arctic and its impacts on arthropod life The strict geographic definition of the Arctic is the area north of the Arctic Circle; at latitude of 66º32’ North (Strathdee & Bale, 1998). However, for this project, the arctic encompasses any region north of the transition zone from boreal to tundra (Strathdee & Bale, 1998). The arctic is a hostile environment for most organisms. Prolonged, cold winters are briefly interrupted by brisk summers. Precipitation is rare, especially at higher latitudes, and only lightly occurs during the warmer months. Average winter temperatures range from -37ºC in the northern regions to -18ºC in the southern regions. Average summer temperatures range from +6ºC in the north to +16ºC in the south. Increasing latitude corresponds to small, yet significant drops in temperature, along with further losses of an already meager flora (Danks, 1981, 1993, 1994). In addition, high latitudes are characterized by imposing cloud cover, extreme exposure, and strong winds (Danks, 1981, 1993, 1994). The most influential environmental aspects on arctic arthropod reproduction, development, and survival are: low temperatures; restricted water availability; dry conditions; reduced yearly heat budget, minimal primary production; extreme and inconsistent weather; absence of regular environmental cues: long daylight during short growing season; relative homogeneity of tundra environment; and acute climatic differences between the boreal zone, the low arctic, and the high arctic (Downes, 1964, Strathdee & Bale, 1998; Nielsen & Wall, 2013). Arthropod inventories in islands of the Canadian high arctic have found rich assemblages including Coleoptera, Lepidoptera, Collembola, Arachnida, Hymenoptera, and Diptera (Oliver, 1963; Danks, 1981; Brodo, 2000). Studies of Canadian arthropod diversity demonstrate that as Coleoptera diversity drops with increasing latitude, Diptera diversity rises, making Diptera a dominant component of high arctic arthropod diversity (Danks, 1994; Brodo, 2000).

Arctic arthropods and climate change The impacts of global climate change are particularly pronounced in the arctic, especially the High Arctic (Strathdee & Bale, 1998; Nielsen & Wall, 2013). The overarching impact on the arctic environment is an increase of both the mean annual temperature and precipitation (Comiso, 2006). This increase leads to a variety of climate change impacts that directly affect arctic arthropod community composition and distribution (Strathdee & Bale, 1998; Nielsen & Wall, 2013). For instance, local warming in the arctic increases plant growth, resulting in proliferation of shrubs and in the northward displacement of the tree line (Callaghan et al., 2011). Arctic winters have been getting warmer over recent decades (Bokhorst et al., 2012). Warmer winters can lead to the loss of vital insulating snow, or to the development of ice layers as the partly thawed snow refreezes (Bokhorst et al., 2012). Such consequences of warmer winters damage both soil structure (e.g.. formation of thermokarst) and plant communities, and thus, arthropod communities (Bokhorst et al., 2012). The extent and results of arctic environmental changes due to climate change are not well understood. However, based on likely environmental consequences, three plausible scenarios can be formulated. Scenario 1: native species outcompeted by temperate species: arctic organisms, particularly arthropods, have developed essential strategies to survive through the harsh conditions of the arctic (Nielsen & Wall, 2013). Strategies such as winter dormancy allow the native biota to thrive under the current arctic conditions (Nielsen & Wall, 2013). However, the corresponding metabolic costs may impede both growth rates and the organism’s competitive capacity under more advantageous arctic conditions brought by climate change (Nielsen & Wall, 2013). Thus, as the arctic climate warms, many native arctic arthropod species might be at risk of being outcompeted by other species (Nielsen & Wall, 2013). Scenario 2: native species keep up and adapt: arthropods have a very short generation time, and hence, respond quickly to environmental changes (Nielsen & Wall, 2013). This implies that these native species could quickly adapt to changing environmental conditions, enabling them to compete more successfully with the arriving new species

(Nielsen & Wall, 2013). Scenario 3: community structure shifts: the third scenario combines elements of both the first and second scenario and assumes a progressive change in the arctic climate rather than an abrupt climatic event (Nielsen & Wall, 2013). The main driver of this scenario is an alteration in arctic vegetation, which will increase the depth of the biologically active layer (Nielsen & Wall, 2013). While some temperate species will migrate northward, some native species will manage to adapt to the changing arctic conditions (Nielsen & Wall, 2013). This movement of species may alter biotic interactions and thus gradually lead to a steady alteration in the arctic ecosystem state (Nielsen & Wall, 2013).

Relevance of arthropod studies The ability to describe faunal patterns reinforces both systematics and ecology (Danks, 1993). Accordingly, studying arthropod community structure and distribution is key to assessing the extent and consequences of ecosystem changes (Timms et al., 2013). Moreover, because species are the operating entities in ecosystems, species level information and identification is key for the study of faunal patterns (Danks, 1993). Understanding the impacts of climate change in the arctic may contribute to forecasting climate change impacts in various other ecosystem types by, for instance, enabling researchers to discern which processes initiate climate change feedbacks (Nielsen & Wall, 2013).

Taxonomy of Empididae and the genus Rhamphomyia The superfamily Empidoidea contains about 11,400 described species. Such species diversity is associated with substantial morphological and structural diversity, especially with regards to the male genitalia. Sinclair and Cumming (2006) recognize five families within Empidoidea: Empididae, Hybotidae, Atelestidae, Brachystomatidae, and Dolichopodidae. The polyphyletic Empididae are distinguished from the other empidoid families by four key characters: vein R4+5 usually branched; prosternum enlarged making precoxal bridge; laterotergite generally with bouquet of setae; small female terminalia equipped with basic cerci (Sinclair & Cumming, 2006).

The subfamily Empidinae, which contains the genus Rhamphomyia, is recognized by five morphological traits (Sinclair & Cumming, 2006): palpi usually curved upwards; broad wings with a well-developed anal lobe; forelegs not raptorial (with the exception of genus Ochtherohilara Frey); empodium often in the form of a bristle; male terminalia typically with grasping cerci or hypandrium laterally flattened or keel-like. The empidines are composed of two tribes: Hilarini and Empidini. The tribe Empidini is a dumping group, that contains genera such as Rhamphomyia, that cannot be placed in the Hilarini (Sinclair & Cumming, 2006).

Biology of Rhamphomyia The genus Rhamphomyia is characterized by elaborate mating displays and swarming behaviors (Sinclair & Cumming, 2006). Males of some species swarm over swarm markers while waiting for a female to pass through (Sinclair & Cumming, 2006). Several authors have observed the opposite situation: females form premating swarms flying in a distinct zig-zag manner, and wait for males to pass through with a nuptial gift (Saigusa, 1963; Downes, 1970; Barták, 2001). Copulating pairs seem to always remove themselves from the premating swarms (Barták, 2002). Swarming markers vary from still water bodies to flower heads (Barták, 2001). Swarming behavior does not appear to be consistent throughout the genus. In fact, Daugeron & Grootaert (2005) found that males of R. (Lundstroemiella) magellens Frey copulate with females on flowers without swarming beforehand and without dispensing a nuptial gift. As opposed to more temperate species of Rhamphomyia, many arctic species are believed not to be active throughout the entire day; swarming only begins in the late morning and ends as soon as temperatures begin to drop (Frohne, 1959). However, Downes (1970) observed Rhamphomyia nigrita Zetterstedt swarming on Ellesmere Island well into midnight because conditions remained suitable. In addition, Downes (1970) noted that some high arctic species, such as R. ursinella Melander predominantly hunt and mate on the ground rather than in the air. There is a sexual differentiation of habits that is unique to the the Empis-Hilara- Rhamphomyia complex: only males hunt for prey; females rely on males to obtain their

protein. Males carry a nuptial gift, which they give to the female while mating. The nuptial gift might be an empty silken ball, seed, or an insect prey, such as a chironomid midge or a mosquito. To date, there is no evidence suggesting that males also feed on insect prey. Moreover, it remains unclear if the female really feeds on the nuptial gift because some Rhamphomyia females only receive an empty silken ball rather than an insect prey. Thus, the prey might just serve as a sign stimulus for courtship, forsaking its purpose as food (Downes & Smith, 1969). Some species of Rhamphomyia, in particular, species of the subgenus Megacyttarus, have peculiar outgrowths on their legs that seem to serve no other purpose than to trick the female into thinking that the swelling is a prey item (Sivinski & Wing, 2008). Other female Rhamphomyia species swallow air to puff up inflatable pouches situated on the abdomen pleura (Funk & Tallamy, 2000). When inflated, these pleural sacs give a swollen appearance to the females, and thus trick males into thinking that they are more fecund (Bussière, et al., 2008). Females further enhance their swollen, saucer- like appearance by extending their often pennate legs around their abdominal margins (e.g. R. longicauda) (Funk & Tallamy, 2000; Bussière, et al., 2008). Thus, in order to obtain their protein, it appears that the swelling of the females has evolved to convey a dishonest message to the mate-seeking males that the eggs are close to maturation (Funk & Tallamy, 2000). Mate selection in Rhamphomyia varies from species to species. While some species demonstrate the more common mate selection system, i.e. the male displays, and the female choses, other species show a less traditional mate selection system. Males of R. longicauda exhibit a “role-reversed” mating system. The male carrying a nuptial gift hover beneath these swarming, inflated females and assess the quality of their potential mates. Upon choosing a mate, the male rises beneath the selected female R. longicauda , hovering until the female drops to meet him. The exchange of the prey is then completed, and copulation occurs outside the swarm. Males without a prey item never infiltrate such female swarms (Funk & Tallamy, 2000). Both male and female Rhamphomyia species appear to be generalists: they feed on insect prey as well as on nectar (Downes & Smith, 1969; Barták & Kubik, 2009). Barták & Kubik (2009) speculated that Rhamphomyia are only predators during mating

activity. Because Nearctic species of Rhamphomyia are extensively anthophilous, it is reasonable to assume that they are key pollinators in the Arctic, especially in the high arctic. A recent study in the subalpine French Alps examined the importance of Diptera, and notably Empidinae pollination in colder environments (Lefebvre et al., 2014). Higher elevations supply advantageous habitats (e.g. moist soils, stream margins) for Diptera larvae, and flies are more efficient than bees at using microhabitats for thermoregulation (Lefebvre et al., 2014). Diptera dominated the flower visitor community assemblage, and Empidine flower visitation rates exceed that of Syrphidae and other Diptera (Lefebvre et al., 2014). The efficiency of Empidinae pollination is probably the result of increased hairiness in Empidines, and their general nectar feeding behavior (Lefebvre et al., 2014). The dominance of empidine flower visitors in alpine habitats is probably mirrored by Rhamphomyia in the Arctic. Larvae of Empididae mostly inhabit soil, decaying wood, leaf litter, mosses, and sometimes running water, although Rhamphomyia larvae do not appear to be truly aquatic (Grootaert, 2004). Larvae are all predators, preying on various arthropods, such as other fly larvae (e.g. dolichopodid larvae) (Barták, 2002).

Functional diversity of Rhamphomyia Rhamphomyia shows a wide variation in body size among species, ranging from 3-13 mm long (T. Saigusa, pers. comm.). To date, the most widely used pattern for broad-scale invertebrate body size distribution patterns is Bergmann’s Rule (Bergmann, 1847; Entling et al., 2010; Shelomi, 2012). Bergmann’s rule argues that closely related homeotherms, and specifically, endotherms become increasingly larger in colder environments (Blackburn et al., 1999; Homburg et al., 2013). Bergmann (1847) suggested that this trend exists due to larger individuals having smaller surface-to-volume ratios, therefore increasing heat conservation. However, Bergmann’s rule and its predictions initially only pertain to body size studies of birds and mammals (Blackburn et al., 1999). Bergmann’s rule does not formulate predictions for ectotherms, and only addresses one process: colder climates versus warmer climates (Blackburn et al., 1999). Bergmann did not equate colder

climates to latitude increase (Cushman et al., 1993; Blackburn et al., 1999). In fact, there is a generally accepted substitute of Bergmann’s original assertion about body size differences between colder and warmer climates with assertions about body size differences across latitude (Entling et al., 2010). There is a large body of literature on arthropod body size patterns across a latitudinal gradient (Shelomi, 2012). Researchers have variously interpreted Bergmann’s argument, resulting in the conception of different types of studies guided by various versions of Bergmann’s rule, and thus, producing conflicting results and conclusions (Shelomi, 2012). Despite such discrepancies in the interpretation of Bergmann’s original rule, such studies use their results to either support or refute Bergmann’s rule (Blackburn et al., 1999). Commonly used arthropods in such studies include spiders, bees, ants, and lepidopterans (Entling et al., 2010). These studies demonstrate that climatic effects on body size of arthropods are extremely varying and hence, do not have a convenient theoretical framework that is comparable and generally applicable (Entling et al., 2010). Cushman et al. (1993) studied latitudinal patterns in body size displayed by European worker ant faunas at the intraspecific level. They solely examined body size trends at the interspecific level. They found that worker ant body length significantly increases with latitude, and suggested that this trend is the result of smaller ant species dropping out of assemblages along with latitude increase (Cushman et al., 1993). Cushman et al. (1993) suggested that resistance to starvation increases with body size because energy reserves increase with size more quickly than metabolic rate. As a result, bigger species are capable of enduring more prolonged periods of harsh conditions before starvation (Cushman et al., 1993). This hypothesis predicts that regions characterized by high seasonality and unpredictable climate are primarily composed of bigger species rather than smaller species (Cushman et al., 1993). Further, Cushman et al. (1993) predict that bigger species should account for a larger fraction of the taxa at elevated latitudes. Entling et al. (2010) examined body size distribution in three datasets of spiders in Europe. They found that the mean body size of spider assemblage decreased from warm/dry environments to colder/moist environments, and argue that this decrease could

be explained by the replacement of large-bodied spider families by small-bodied spider families (Entling et al., 2010). These results are consistent with the predictions of three mechanisms they had considered: metabolic rate, desiccation resistance, and community interaction mechanisms (Entling et al., 2010). Metabolic rate and season duration both increase with temperature, enabling ectothermic species to reach larger adult sizes under warmer conditions (Entling et al., 2010). Desiccation resistance increases with body size on account of a thicker cuticle and a smaller surface area-to-volume ratio (Remmert, 1981). Competition and predation stresses are more prominent in warm environments, thus favouring larger animals (Blackburn et al., 1999). Shelomi (2012) published a comprehensive review of the literature on both inter- and intraspecific insect body size distribution patterns along latitudinal and altitudinal gradients. In general, both Bergmann and converse-Bergman clines are almost equally supported for intraspecific studies. However, similar numbers of intraspecific studies also reported no cline (Shelomi, 2012). Studies at the interspecific level were more frequently reporting no clines rather than Bergmann or converse-Bergmann clines (Shelomi, 2012). Insect order has an influence on the type of clines most frequently reported (Shelomi, 2012). Indeed, Shelomi (2012) reported 83 studies that show that Diptera follow a Bergmann cline, whereas only 14 Coleoptera studies show a Bergmann cline. In interspecific studies, expanding the number of species increases the likelihood of obtaining a converse-Bergmann or a non-cline (Shelomi, 2012). For instance, Blanckenhorn & Demont (2004) hypothesized that bigger and/or usually slower- developing insect orders (e.g. Coleoptera, Orthoptera) would be likely to show converse- Bergmann clines because they are more impacted by high seasonality, whereas faster- developing insect orders (e.g. Diptera) would be more likely to show Bergmann clines. In addition, large spatial-scale studies at the intraspecific levels are more susceptible to exhibit intraspecific studies (Shelomi, 2012). Accordingly, whether Bergmann’s rule pertains to a certain taxon is specific to that taxon and its habitat, as well as extremely contingent on the study’s design (Shelomi, 2012). Intraspecific analyses of species with extensive latitudinal ranges were more likely to exhibit Bergmann clines than analyses at smaller spatial scale (Shelomi, 2012).

Studies that are based on a very small range may actually reflect a local microclimate gradient, thus blurring the broader, latitudinal gradient (Shelomi, 2012). Small spatial scales may reflect only part of a bigger sawtooth cline, thus describing only a gradient at the micro scale but omitting the broader gradient that is actually prompted by a mechanism covarying with latitude (Shelomi, 2012). On the other hand, studies on too large a spatial scale may produce fictitious Bergmann clines, because an excessively large range will cover numerous variables influencing body size such that it can overrun the effect of temperature (Shelomi, 2012). Moreover, studies covering a large latitudinal range but with sparsely located sampling sites might not be including any important clines besides the “pure” latitudinal cline, and hence, it is important to sample as continuous an expanse as possible (Shelomi, 2012).

Objectives Two ways of assessing diversity in communities include: 1) taxonomy (e.g. species richness), and 2) functional diversity (e.g. body size). This project aims to link taxonomy with ecology by describing the species-richness of arctic Rhamphomyia, and thus facilitating analysis of their body size across latitude. The first objective is to conduct an inventory of Rhamphomyia species at multiple sampling sites in northern Canada. The inventory serves as an essential foundation for the subsequent objectives. The inventory also allows us to determine the broad spatial distribution of Rhamphomyia species across the Canadian Arctic biomes. The second objective is to construct a key to the species of Rhamphomyia from the Canadian Arctic Archipelago, Greenland, and Iceland, and to diagnose and describe the species. The third objective is to examine the functional diversity of arctic Rhamphomyia by examining body size patterns. Rhamphomyia species show a wide variation in body size among species, ranging from 3-13 mm long. Most studies of Bergmann’s rule have been focused at the single species-level (Shelomi, 2012), but this study takes a broader approach, examining all species within the genus. To assess the relationship of Nearctic Rhamphomyia body size patterns, we examine latitudinal and other spatial effects. We determine whether the spatial patterns of Rhamphomyia body size in the Nearctic are

driven by temperature decreases corresponding to latitude increase, or by other ecological factors.

Connecting Statement Despite the dominance of Rhamphomyia in the northern Nearctic, little is known about the species-richness, geographic distribution, and ecological patterns in the genus. Therefore, an inventory of the northern Rhamphomyia is necessary baseline data to assess the taxonomic diversity of Rhamphomyia and to assess ecological patterns. The following chapter is an inventory based on standardized sampling at multiple sites from the north boreal to the high arctic in Canada.

Chapter 2. Faunal Inventory of Rhamphomyia in Northern Canada

The Empididae is a widespread, species rich, and dominant Diptera family in the Arctic (Collins & Wiegmann, 2002). The genus Rhamphomyia is one of the most abundant and species-rich genera in the northern Nearctic. Danks (1981) recorded 19 species of Empididae in the northern Nearctic, 14 of which were Rhamphomyia. Six of these Rhamphomyia species occur in the high artic. However, recently collected data and unpublished records in museum collections suggest that Rhamphomyia is far more species-rich, even in the high arctic, than recorded by previous authors (e.g. Downes, 1970, Danks, 1981). The dominance of Nearctic Rhamphomyia, and their role as both pollinators and predators, suggest an important role in arctic food webs. There is a need for baseline data to better understand these assemblages and to monitor future responses to climate and habitat change (Hoye & Sikes, 2013). The Northern Biodiversity Program (NBP) collected arthropods in several target taxa, including Diptera, at multiple sites in northern Canada in an effort to establish baseline data on diversity and as a framework to assess ecological change in northern arthropods. The purpose of this chapter is to compile a species-level inventory of the Rhamphomyia collected by the NBP.

Materials and Methods

Study Sites Specimens were collected from twelve sampling localities across the Canadian north: 4 boreal sites (Moosonee, ON, Goose Bay, NL, Yellowknife, NT, Norman Wells, NT), 4 low arctic mainland sites (Schefferville, QC, Churchill, MB, Dempster Hwy, YT, Kugluktuk, NU), and 4 high arctic island sites (Iqaluit, NU, Cambridge Bay, NU, Banks Island, NT, Ellesmere Island, NU) (Figure 2.1).

Sampling methods Standardized, replicated sampling was used in 2010 (eastern sites), and in 2011 (western sites), using Malaise traps, yellow pan traps, pitfall traps, and sweeping. Sampling grids

were established in each of 3 wet and 3 mesic plots per sampling locality, for a total of 72 sampling plots. We collected at each site for about two weeks during the summer peak of arthropod activity, at 4 day intervals. Kugluktuk was sampled during the entirety of the summer season 2010 and for two weeks in 2011. However, the 2011 sampling season was unsuccessful due to bad weather. Therefore, a subsample of a similar two-week time frame to that of 2011, was selected from the 2010 for inclusion in the inventory.

Specimen preparation and identification Samples were preserved in ethanol in the field and subsequently dried and mounted. All Empididae were extracted from samples and identified to genus. Because most Nearctic Rhamphomyia are undescribed or unidentifiable to named species, all male Rhamphomyia were sorted to morphospecies. This provided the necessary taxonomic resolution to establish overall species richness as well as geographic distribution of species by site and biome. Only males were considered in this study because females are not readily identified to species/morphospecies. Morphospecies sorting of Rhamphomyia males was effective because genitalia are highly species-specific and distinctly modified (Downes, 1970). Chaetotaxy and leg ornamentation were also useful to distinguish morphospecies. When external morphology was not sufficient, genitalia were dissected to examine concealed structures, notably phallic curvature and lamellar lobes. Most Nearctic Rhamphomyia species are undescribed and no key to named Nearctic Rhamphomyia species exists. However, despite this lack of taxonomic resolution, several species were assignable to named species or species groups based published keys to some subsets of the genus (e.g. subgenus Megacyttarus (Barták, 2002)), the Rhamphomyia collection in the Canadian National Collection of Insects (Ottawa, ON), and extensive unpublished taxonomic notes by T. Saigusa (copies deposited in CNC). In addition to sorting based on morphology, DNA barcoding (658 base pairs of 5’ region of the mitochondrial Cytochrome c oxidase subunit I (COI) gene) (Smith, et al., 2005) of representatives of all Rhamphomyia male morphospecies (1 to 4 specimens for each morphospecies) was performed at the Canadian Centre for DNA Barcoding

(University of Guelph, Guelph, ON). DNA barcoding results were used to supplement morphology-based sorting.

Results

Inventory A total of 10,298 male specimens was identified, comprising 67 morphospecies (Appendix 2.1), 44 of which were matched to named species, species group, species number, or subgenus. The remaining 23 morphospecies did not match known taxa and retain an alphabetical morphospecies code. From the 44 morphospecies that were matched, 9 subgenera were identified, two of which are undescribed: Calorhamphomyia Saigusa, Vockerothempis Saigusa, Ctenempis Frey, Dasyrhamphomyia Frey, Eorhamphomyia Frey, Megacyttarus Bigot, Pararhamphomyia Frey, New Subgenus A, and New Subgenus B. The most abundantly collected subgenus is Pararhamphomyia. The twelve most abundant species accounted for 9,486 specimens (Table 2.1). Of the three biomes, the low arctic had the highest species-richness, as well as the highest number of unique species (Figure 2.2). The Dempster Highway site had the highest species richness (26 species) (Table 2.2). On the other hand, the high arctic had both the lowest species-richness and the lowest number of unique species, but the highest species- abundance (Table 2.2). Indeed, of the 10,298 male specimens of Rhamphomyia, 7,066 specimens were collected in the high arctic, contrasting to 2,862 specimens collected in the low arctic. The boreal biome is more species-rich than the high arctic, but has the lowest species-abundance out of the three biomes (370 specimens). Individual species of Rhamphomyia were rarely collected in all three arctic biomes. Instead, species were usually restricted either to the boreal-low arctic biomes or only to the low arctic biome (Figure 2.2). Only four species occurred across all three biomes, only four species spanned the low arctic-high arctic biomes, and no species occurred in both boreal and high arctic biomes. In contrast, 15 species spanned the boreal-low arctic biomes, and 26 species only occurred in the low arctic biome (Figure 2.2).

Discussion The species-richness of northern Nearctic Rhamphomyia is far higher than published records would indicate. The most recent compilation (Danks, 1981) recorded 14 species in arctic Canada, but NBP sampling from only 12 sites collected 65 species. Few species spanned all three biomes. Instead, most species either occurred in only the low arctic biome, or span the boreal-low arctic biomes. The low arctic biome had the highest species-richness, primarily because of the 26 species identified at the Dempster Highway site (Table 2.2). This is likely due to the existence of the Beringian refugium, which included part of the Yukon, during Pleistocene glaciations. Beringia was the region encircling the Bering Strait, the Chuckhi Sea, and the Bering Sea (Elias et al., 1997; Froese et al., 2009). Throughout most of the Pleistocene, the Bering Land Bridge linked Alaska and Siberia and enabled dispersal of terrestrial animals and plants (Elias et al., 1997) Beringia was a mosaic of plant communities that changed over time (Anderson, 1985; Guthrie, 1990; Elias et al., 2000). Broad vegetation patterns in eastern Beringia included steppe-tundra, moist and xeric tundra, fell fields, and herbaceous tundra grasslands (Anderson, 1985; Elias et al., 2000; Froese et al. 2009). Central Beringia, i.e. the Bering Land Bridge, mostly consisted of mesic and shrub tundra (Elias et al., 1997; Lozhkin et al., 2011), and evidence supports the hypothesis that steppe-tundra largely dominated western Beringia (Elias et al., 2000; Alfimov & Berman, 2001). The Beringian climate was favorable to animal life. Coleoptera assemblages from western and inland Alaska dating from the Last Glacial Maximum (LGM) reveal only a 2º–3ºC drop below current in summer mean temperatures, and an 8ºC drop below current mean winter temperatures (Elias, 2000). The herb-dominated vegetation of the full- glacial phase replaced by shrub-tundra, possibly delineates the beginning of climate amelioration in central Beringia (Lozhkin et al., 2011). Indeed, the Alaskan Birch Zone started to expand on the Beringian land bridge as early as 14,000 years ago (the end of the LGM), suggesting a warming of the climate. Towards 11,000 years BP, when the Bering Land Bridge would have begun its submergence, insect assemblages indicate both warmer-than-present summers, but colder-than-modern winters (Elias et al., 1997). The

suitable living conditions of Beringia would have enabled diverse assemblages of Diptera, to thrive and speciate in the Yukon, while the remainder of northern Canada was covered with ice. The high arctic biome had the lowest species-richness, yet the highest species- abundance of Rhamphomyia. The difference in species-richness between the boreal-low arctic biomes and the high arctic is probably primarily driven by the climatic differences between the sub-arctic and the high arctic (Ernst & Buddle, 2015). Climatic factors, such as temperature and precipitation, are key in shaping arthropod species distribution patterns (Ernst & Buddle, 2015). The mean annual temperature of the high arctic hovers around -3ºC, whereas the low arctic biome has a mean summer temperature of 5ºC, and mean winter temperature of -28ºC near the Mackenzie Delta, to -18ºC in northern Quebec (Walker et al., 2005). Although the low arctic is cold, the temperature decrease from the low arctic to the high arctic is significant enough to cause a pronounced drop in species richness in the highest latitudes of the Canadian Arctic (Ernst & Buddle, 2015). In addition to its extremely cold temperatures, the high arctic is the driest, most dynamic and unpredictable environment in Canada. There are drastic differences in day length and sunlight from summer to winter. As a result of low nutrient availability, widespread permafrost, and low soil water content, plant cover drops substantially from the low to the high arctic (Bliss et al., 1973). Most of the archipelago consists of Polar Desert soils (Bliss et al., 1973). In this type of desert, lichens and mosses are common, and vascular plants are extremely scarce. In addition to harsh environmental conditions preventing establishment and survival of species on high arctic islands, the Northwest Passage might also be a physical barrier to species dispersal from the mainland to the arctic islands post-glacially. Because of the salt water barrier and long flight distance, initial colonization of the islands might have been heavily filtered, allowing only a small subset of Rhamphomyia species to cross. Trophic interactions in arctic arthropods are poorly understood (Wirta et al., 2015a, 2015b). Our study of Rhamphomyia in the Nearctic demonstrates the necessity of describing both species-richness and -abundance to gain insight in trophic interactions

(Bolduc et al., 2013). The NBP data reveals the substantially elevated species-abundance and dominance of Rhamphomyia in the high arctic, thus suggesting the crucial role Rhamphomyia have in high arctic biome. Since Rhamphomyia are generalists, this study further confirms that generalism is common in the arctic, especially in predators. In fact, Wirta et al (2015a) find that the primary predators (birds, spiders) are also generalists. In the arctic, generalism is key for animal survival because primary productivity is low and energy resources fluctuate greatly (Hoye & Sikes, 2013), thus limiting the capacity for specialism (Wirta et al., 2015a). Rhamphomyia are both important pollinators and predators (Downes & Smith, 1969), and their elevated abundance in the high arctic insect biomass suggests that they are some of the most important pollinators, predators, and prey. As pollinators, they are frequent visitors of the common high arctic Dryas flowers (Kevan, 1972; Inouye et al., 2015). Although Diptera are considered to be less efficient pollinators than bees (Inouye et al., 2015), authors such as Kearns and Inouye (1994), and Zheng et al (2011), have found that certain flowers visited frequently by less efficient fly pollinators and much less frequently by more efficient bee pollinators, actually rely primarily on the frequent fly pollinators for successful pollination. Therefore, as Diptera are dominant in the high arctic, high arctic flowers are reliant on fly pollination, despite pollination limitation (Inouye et al.., 2015). As predators, Rhamphomyia are known to feed on other insect taxa, such as mosquitos and chironomids (Downes, 1970), and thus, actively participate in the regulation of other high arctic insect populations (Wirta et al., 2015b). In addition, the substantial abundance of the predatory Rhamphomyia in the high arctic suggests that these species strongly shape local prey species assemblages (Wirta et al., 2015b). Rhamphomyia are also likely to serve as an important food source for migratory birds, and for other insect taxa, such as spiders (Danks, 1981; Hoye & Sikes, 2013; Wirta et al., 2015a,b).

Table 2.1. Twelve most abundant morphospecies of Rhamphomyia, number of male specimens identified, and sites at which they were collected by the Northern Biodiversity Program (BI: Banks Island; CB: Cambridge Bay; CH: Churchill; DH: Dempster Hwy; EI: Ellesmere Island; GB: Goose Bay; IQ: Iqaluit; K: Kugluktuk; M: Moosonee; NW: Norman Wells; SC: Schefferville; Y: Yellowknife). Morphospecies Specimens Sites collected R. (Dasyrhamphomyia) nigrita 4690 BI, CB, EI, K R. (Pararhamphomyia) fuscula gp. sp.1 1598 DH, EI, IQ, K, NW R. (Ctenempis) calvimontis 1504 BI, CB, IQ, K R. (Pararhamphomyia) tipularia gp. sp.4 293 BI, CB, DH, NW R. (Pararhamphomyia) filicauda 282 BI, CB R. (Pararhamphomyia) dispar gp. sp.10 229 CH, DH, M, NW, SC R. (Eorhamphomyia) nigrolineata gp. sp.4 174 CB, K, DH, CH, Y R. (Pararhamphomyia) pusilla gp. sp.7 165 DH, K R. (Pararhamphomyia) pusilla gp. sp.1 165 Y, CH, GB, DH, K R. (Dasyrhamphomyia) sp.9 133 BI, CB, K R. (Megacyttarus) praecipua 127 DH R. (New Subgenus A) flexuosa gp. sp.12 126 K

Table 2.2. Number of Rhamphomyia specimens, species-richness and number of species unique to each Northern Biodiversity Program site. Site codes as in Table 2.1 Site Specimens Total species Species unique to site DH 385 26 5 K 2082 17 4 NW 56 14 3 M 282 13 6 CH 265 13 1 Y 23 11 1 SC 130 9 3 GB 9 6 2 CB 2440 6 2 IQ 169 4 0 BI 4447 4 2 EI 10 3 0

CB DH K NW

IQ Y

CH SC GB

Figure 2.1. Northern Biodiversity Program sampling sites in boreal boreal (circles), low arctic (squares), and high arctic (triangle) biomes. (BI: Banks Island; CB: Cambridge Bay; CH: Churchill; DH: Dempster Hwy; EI: Ellesmere Island; GB: Goose Bay; IQ: Iqaluit; K: Kugluktuk; M: Moosonee; NW: Norman Wells; SC: Schefferville; Y: Yellowknife).

HA 4 LA

26 4 4 0 1515

B 12

Figure 2.2. Rhamphomyia species unique to each biome and shared between biomes in Northern Biodiversity Program sampling sites (HA: high arctic; LA: low arctic; B: boreal).

Connecting Statement Although the species inventory of Northern Biodiversity Program Rhamphomyia in Chapter 2 shed light on the overall spatial distribution across the three northern biomes, it also highlighted the lack of named species and the difficulty in identifying species. The absence of identification tools for the Nearctic Rhamphomyia is problematic because it prevents comparison of species- level results across multiple projects in the literature. In order to render this data more usable, it is essential to develop identification tools, to determine which species are undescribed, and to name these unknown species. Although considerable time and resources would be needed to create a key to the Nearctic Rhamphomyia species, partial progress is still desirable. In Chapter 3, I present a key to the Rhamphomyia of the Canadian Arctic Islands, Greenland and Iceland.

Chapter 3. Key to the Rhamphomyia of the Canadian Arctic Islands, Greenland and Iceland

Rhamphomyia Meigen is one of largest genera of the Empididae, comprising over 400 species, mostly in the Holarctic region, but also widespread in other zoogeographical regions (Saigusa, 2012). However, this number is a significant underestimate. Indeed, in Europe, where Rhamphomyia has been more extensively studied, there are 200 known species (T. Saigusa, pers. comm.). Many species remain to be described, especially in the Nearctic, where the species diversity is poorly known (Bartak & Kubik, 2009). In fact, unpublished estimates by T. Saigusa suggest that about 600 Rhamphomyia species occur in the Nearctic, ¼ of which are actually named. Some species might be endemic to the Nearctic, while others are likely Holarctic (Saigusa, 1963). Overall there may be as many as 1500 species of Rhamphomyia. Rhamphomyia is distinguished by: elongate proboscis; mesoscutum disc covered with small hairs; shape and ornaments of legs extremely variable and species specific. However, Rhamphomyia species are subject to within genus variation in a variety of other morphological characters, including: holoptic versus dichoptic eyes, postpedicel and labrum lengths, labella form and setation, chaetotaxy of thorax and legs, depth and angle of axillary incision in the anal lobe of the wing, fading of the vein CuA + CuP (i.e. anal vein), discal cell size and form, shape and size of male genitalic lamellae, and modifications on the pregenital segments (Saigusa, 2012). The subgeneric classification of Rhamphomyia is largely unresolved and incomplete. The subgenera are based predominantly on plesiomorphic morphological traits, such as the complete vein CuA + CuP, which results in paraphyletic and dissimilar groups (Saigusa, 2012). Most morphological characters used to distinguish subgenera seem to have evolved separately in multiple lineages of Rhamphomyia (Saigusa, 2012). For instance, Frey (1922) used discrepancies in the angle of the axillary incision of the anal lobe to recognize Pararhamphomyia Frey + Choreodromia from Dasyrhamphomyia + Rhamphomyia (Saigusa, 2012). However, the pronounced and acute axillary incision as a result of the enlargement of the alula and the anal lobe is common in bigger

Rhamphomyia, whereas the 90º-angled and obtuse axillary incisions are more prominent in smaller Rhamphomyia (Saigusa, 2012). The species of Rhamphomyia in the Nearctic are difficult to identify because there is no key to species. Although a comprehensive Nearctic key would be a massive undertaking, recent collecting effort in northern Canada provided extensive material for partial progress towards this goal. In this chapter, a key to the species of the Canadian Arctic Archipelago, Greenland, and Iceland is written to facilitate the identification of the 21 Rhamphomyia species known from the region.

Materials and Methods This key is based extensively on Northern Biodiversity Program specimens deposited in the Lyman Entomological Museum, McGill University, Ste-Anne-de-Bellevue, QC (LEMQ), and previously identified, as well as newly identified specimens deposited in the Canadian National Collection of Insects, Ottawa, ON (CNC). A species (re)description and diagnosis was written for each species. In addition, because male genitalia, and often legs, of Rhamphomyia are distinctive and species- specific, genitalic illustrations and photographs of distinctive leg features are also provided. Photographs were taken with a Leica camera model DFC425C managed by a Leica Digital Imaging System. The morphological terms of Sinclair & Cumming (2006) are followed. Subgenera, species groups and individual species are included in a single key. Because most species are undescribed, morphospecies codes are used extensively in the key. Some newly recognized synonymies are noted where relevant.

Results and discussion

Key to male Rhamphomyia of the Canadian Arctic Islands, Greenland and Iceland

Remark: There are four species which do not have a published subgeneric classification. All other species are keyed to their respective previously published subgeneric classification.

1. Anal vein distinctly visible and reaching wing margin… 15 - Anal vein faint and usually not reaching wing margin; if reaching wing margin, vein unpigmented, only a faint impression…Pararhamphomyia... 2

2. Anal vein present…R. hoeli Frey [species keys to both sides of couplet because of difficulty in interpreting extend of anal vein] - Anal vein absent or very faintly reaching wing margin as a fold… 3

3. dm-cu crossvein missing (discal cell open); vittae dark and shiny (Fig. 3.12) …R. (P.) diversipennis gp. sp. 1 - dm-cu crossvein present; vittae usually present but not shiny… 4

4. Mid-leg distinctly modified: femur strongly curved upwards; tibia curved inwards, with 3-5 dark, stout mid posterodorsal setae longer than 2x tibial width; basitarsus swollen in middle with several rows of stout, dark mid dorsal, antero- and posterodorsal setae, always shorter than 2x basitarsal width (Fig. 3.19)…R. (P.) pusilla gp. sp. 7 - Mid-leg not distinctly modified; often hind leg with distinctive modifications… 5

5. Tergite 5 bearing cluster of dark lateral setae, stouter than remaining tergal setae and decreasing in length posteriorly… R. (P.) tipularia gp. sp. 4 - Tergite 5 not bearing such cluster of dark, stout lateral setae…6

6. Hind femur with row of dark, stout, spine-like posteroventral setae, shorter than ventral pubescence (Fig. 3.20); apex of phallus expanded with ridge of teeth-like projections (Fig. 3.49)… R. (P.) fuscula gp. sp. 1 - Hind femur lacking posteroventral row of setae shorter than ventral pubescence; tip of phallus lacking teeth-like projections… 7

7. Fore basitarsus swollen, broader than width of fore tibia and mid and hind basitarsi (Fig. 3.16)… R. (P.) tipularia gp. sp. 1 - Fore basitarsus not swollen…8

8. Phallus stout, without curvature, never extending farther than length of genitalia (Fig. 3.47); tergite 8 long, subequal to length of epandrium … R. (P.) simplex Zett - Phallus slim to hair-like, usually extending farther than length of genitalia, and often with curvatures…9

9. Epandrium elongated and slim; cerci constricted in middle; phallus hair-like, always longer than length of genitalia… 10 - Epandrium shorter and broader; cerci not constricted in middle; phallus slender, but not hair-like … 12

10. Apex of epandrium truncate (Fig. 3.64); hind tibia with distinctive inward bend on apical end (Fig. 3.35)…R. (P.) caudata gp. sp. 7 - Apex of epandrium rounded; hind tibia without distinctive inward bend…11

11. Hind basitarsus swollen and increasing in size apically such that apical width larger than hind tibial width (Fig. 3.34); antero- and posterodorsal and dorsal setae of hind basitarsus 2x length of basitarsal width; cerci weakly constricted in middle, posterior half parallel to epandrium (Fig. 3.63)…R. (P.) ursinella Melander - Hind basitarsus not swollen and antero- and posterodorsal and dorsal basitarsal setae not 2x length of basitarsal width (Figs. 3.31, 3.32); cerci strongly constricted in middle, posterior half strongly divergent from epandrium (Fig. 3.62) …R. (P.) filicauda Frey

12. Phallus not forming loops… 13 - Phallus forming loops… 14

13. Hind tibia with some antero- and posterodorsal and dorsal setae more than 2x tibial width (Fig. 3.27)… R. (P.) lapponica gp. sp. 8 - Hind tibia with antero- and posterodorsal and dorsal setae shorter than 2x tibial width (Fig. 3.30)… R. (P.) hoeli Frey

14. Hind tarsomere 2 with antero- and posterodorsal setae 2x width of tarsomere (Fig. 3.28); apex of epandrium gradually tapered and lengthened (Fig. 3.59)… R. (P.) lapponica gp. sp. 9 - Hind tarsomere 2 with antero/posterodorsal setae shorter than 2x the length of tarsomere width, except for antero- and posterodorsal preapical setae (Fig. 3.29); apex of epandrium sharply attenuated with slender extension (Fig. 3.60)…R. (P.) lapponica gp. sp. 11 15. Prosternum clothed entirely with setae …R. (Ctenempis) calvimontis Cockerell - Prosternum bare in middle with bundles of setae only on sides… 16

16. Cerci greatly lengthened, overlapped and curved up above abdomen (Figs. 3.50- 3.55)… Dasyrhamphomyia… 17 - Cerci not greatly lengthened, not overlapping and not extended up above abdomen… 19

17. Cerci folded or wrapped over each other, leaving no opening (Fig. 3.50). Hind tibia and femur with pronounced fine, pale ventral pubescence (Fig. 3.21)… R. (D.) nigrita Zetterstedt - Cerci not completely folded over each other, separated by gap (Fig. 3.51, 3.52); hind tibia lacking fine, pale ventral pubescence… 18

18. Cerci slightly overlapping at tip and base, with wide, elongate opening; cercus cylinder-shaped (i.e., apical width about equal to basal width) and reaching tergite 5 (Fig.

3.52). Hind tibia with anteroventral row of dark, stout, spine-like setae shorter than half tibial width; dorsal setae of hind tibia shorter than tibial width (Fig. 3.23); 3 distinctive acr, and dc, brown thoracic vittae… R. (D.) sp.9 - Cerci overlapping at tips and at base, with small, slim oval-shaped opening; cercus triangular-shaped, and ending before tergite 5 (Fig. 3.51). Hind tibia with dorsal setae longer than tibial width but shorter than 2x tibial width (Fig. 3.22); 2 faint greyish vittae… R. (D.) zaitsevi, Becker

19. Phallus hidden within epandrium, leaving phallus barely visible (Fig. 3.56)…R. Eorhamphomyia nigrolineata gp. sp. 4 - Phallus exposed beyond epandrium, with visible curvatures… 20

20. Cercus with distinctive finger-like projection dorsally near base (Figs. 3.40, 3.41)… … 21 - Upper margins of cercus without finger-like projections…… 22

21. Large species (wing length 12 mm). Phallus curvature occurring before folding into epandrium. Cercal finger-like projection stout (Fig. 3.40). Mid tibia with at least 3 preapical antero- and posteroventral setae dark, stout, spine-like… R. hirtula Zetterstedt - Small species (wing length 7.5 mm). Phallus curvature hidden within epandrium. Cercal finger-like projection small and slim, short, less than half cercal length (Fig. 3.41). Mid tibia with 2 anterodorsals and 2 posterodorsals longer than twice tibial width… R. hirticula Collin

22. Sternite 8 with stout, “horn-like” lateral projections (Figs. 3.1, 3.2, 3.39). Phallus with loop forming an acute angle (Fig. 3.38). Hind basitarsus swollen, wider than hind tibial width (Fig. 3.3)… R. herschelli Malloch - Sternite 8 without projections. Phallus with small in-ward U-shaped loop (Fig. 3.37). Hind basitarsus not swollen… R. sp. 12

Species diagnoses and descriptions

Notes regarding transcription of label information: “/”: signifies a line break f#; m#: signifies female, male, respectively CNC; LEMQ: Canadian National Collection of Insects, Arachnids, and Nematodes, ON; Lyman Entomological Museum of Quebec

Rhamphomyia (Ctenempis) calvimontis, Cockerell, 1916 Figs. 3.8, 3.9, 3.10, 3.42, 3.43, 3.67

Material examined. CANADA. Northwest Territories: Head of Clyde Inlet, Baffin Is., 7.vii.1958, 7.viii.1958, G.E. Shewell (9m#, 6f#, CNC); Head of Clyde Inlet, Baffin Is., 7.viii.1958, J.E.H. Martin, (1m#, 6f#, CNC); Coral Harbour, Southampton Is., 6- 17.vii.1948, G.E. Shewell (28m#, 47f#, CNC); Coral Harbour, 15-16.viii.1959, W.R.N. Mason (2m#, 2f#, CNC); Coral Harbour, 16.viii.1959, J.A. Downes (1f#, CNC); Cambridge Bay, 12.viii.1959, J.A. Downes (1f#, CNC); Padley, 24, 27.vii.1950, R.E. Duckworth (2m#, 1f#, CNC); Masik Riv., Banks Is., 3-29.vii.1968, W.R.M. Mason (17m#, 8f#, CNC); Masik Riv., Banks Is., 7-30.vii.1968, G.E. Shewell (17m#, 18f#, CNC); Victoria Is., 71º17’N. 114ºW, 8-28.vii.1975, G. & M. Wood (1m#, 1f#, CNC); Muskox L., 64º45’N, 108º10’W, 11, 18.vii.1953, Ecological Data, J.G. Chillcott, (1m#, 2f# CNC); Baker Lake, 2, 23.vii.1947, T.N. Freeman (2m#, 4f#, CNC); Baker Lake, 6.viii.1949, R.G., (1m#, 1f#, CNC); Padley, 1.viii.1950, R.A. Hennigar (1f#, CNC); Padley, 27.vii.1950, R.E. Duckworth (2f#, CNC); Eskimo Point, 12, 13.vii.1950, G.G. DiLabio (4f#, CNC); Reindeer Depot, Mackenzie Delta, 10.vii.1948, J.R. Vockeroth (1f#, CNC); Melville Pen., Ross Bay, N.66.51, W.84.53, 19-22.vii.1948, Cody McArthur (1f#, CNC); Spence Bay, 14.vii.1951, J.G. Chillcott (1f#, CNC); Bathurst Inlet, 5,19.vii.1951, C.D. Bird (2f#, CNC); Fraser Lake, 68º45’N, 120º36W, 19.vii.1969, G.E. Shewell (4m#, 4f# CNC); Coppermine, 30.vi., 14, 20.vii.1951, S.D. Hicks (4f#, CNC); Eskimo Point, 17.vii.1950, G.R. Roberts (1m#, CNC); -21 m.e. Tuktoyaktuk, 2-5, 17-21, 20-25.vii.1971, D.M. Wood (7m#, 10f#, CNC); Cockburn PI, Canadian Arctic Expedition, Coll, 904, 3.ix.1914, F. Johansen (1f#, CNC); Aklavik, 28.vi.1956, E.F.

Cashman (1m#, CNC); Aklavik, 29, 30.vi., 3.vii.1956, R.E. Leech (1m#, 2f#, CNC). Nunavut. Cambridge Bay, 69.11993, -105.42065, Malaise, mesic, 7-19.vii.2011, wet, 15-19.vii.2011, Repl. 1-3, NBP Field Party, (161m#, LEMQ); Cambridge Bay, 69.11993, -105.42065, Pan, 15-19.vii.2011, mesic, Repl.2, 3, NBP Field Party (2m#, LEMQ); Cambridge Bay, 69.11993, -105.42065, Sweep, 13-15.vii.2011, wet & mesic, Repl. 1-3, (12m#, LEMQ); Herschel Is., F. Johansen Coll, Canadian Arctic Expedition (1f#, CNC). .Yukon: Herschel Is., 11-30.vii.1953, C.D. Bird (11m#, 24f#, CNC); Herschel Is., 24- 29.vii.1971, D.M. Wood (5m#, 1f#, CNC); Herschel Is., 29,30.vii.1971, W.R.M. Mason (2f#, CNC); Herschel Is., 5-11.vii.1953, J.S. Waterhouse (7m#, 10f#, CNC); North Fork Crossing, Mi.42, Peel Plt. Rd. 3500’ 24.vi.1962, P.J. Skitsko (1f#, CNC); North Fork Crossing, Mi.42, Peel Plt. Rd. 3500’ 24.vi.1962, R.E. Leech (1f#, CNC); Firth River, 13- 17.vii.1956, E.F. Cashman (3f#, CNC); Firth River, 13, 14.vii.1956, R.E. Leech (1m#, 3f#, CNC); British Mts. Firth River, 27.vii.1956, R.E. Leech (1m#, CNC) Dempster Hwy., MI.87, 1-12.vii, 27-30.vi.1973, G. & D.M. Wood (1m#, 2f#, CNC). Quebec: Sugluk, 14-22.vii.1954, H. Huckel (2m#, 4f#, CNC). Saskatchewan: Ross Bay, Melville P, N.66.51, W.84.53, 19.vii.1948, G.E. Shewell (2f#, CNC). USA. Alaska: Shrader Lake, 69º30-N, 145ºW, 3-5.vii.1972, K.W. Philip (1m#, CNC); Cape Thompson, 23.vii.1961, R. Madge (1m#, CNC). Colorado. Summit L. Flats, Mt. Evans, 12800', 7, 10, 24.vii.1961, C.H. Mann (16m#, 25f#, CNC); Summit L., Mt. Evans, 12800', 24.vii.1961, J.G. Chillcott (8m#, 2f#, CNC); Summit L., Mt. Evans, 12800', Marshy lake and stream margin, 24.vii.1961, J.G. Chillcott (2f#, CNC); Mt. Evans, Rocky Slope, 13600', 4.viii.1961, J.G. Chillcott (2m#, 6f#, CNC); Summit L., Mt. Evans,12800', 24.vii.1961, S.M. Clark (1m#, 1f#, CNC); Summit L., Mt. Evans,12800', 24.vii.1961, B.H. Poole (6m#, 6f#, CNC); Mt. Evans, 13000', On tundra, 28.vii.1961, C.H. Mann (2m#, 9f#, CNC); Mt. Evans, 12000', 3.viii.1961, W.R.M. Mason (1f#, CNC); Mt. Evans, 13200', 19.vii.1961, W.R.M. Mason (2m#, 4f#, CNC); Mt. Evans, 14000', 25.vii.1961, B.H. Poole (6m#, 10f#, CNC); Mt. Evans, 14000', 27.vii.1961, J.R. Stainer (1m#, CNC); Mt. Evans, 132000', 28.vii.1961, S.M. Clark (1m#, 2f#, CNC). Diagnosis. Male. Upper occipital setae black, long and stout; lower occipital setae pale, hair-like and as long as dark occipital setae; postgena setae pale or dark. Notopleural setae pale, sometimes with several dark setae; setae on coxae hair-like, and

pale or dark, or both; abdomen sometimes with pale setae (Fig. 3.10). Female. Similar to male except setae shorter than male’s and wings yellow; legs without pennate setae. Dorsal thorax with 2 darker grey stripes, diverging near the start of the scutellum. Thorax with distinct row of thickened, yellow, wide bristles. Description. Wing length 13-13.8 mm. Male. Head dark grey-blue in ground- colour, with greyish pruinescence on face, frons, postgena and occiput; oral margin black and shiny. Holoptic, eyes with ommatidia larger on upper half of eye, smaller on lower half. Frons divergent towards antennae. Joining margins of eyes bearing dark, short setae. Margins of face parallel. Ocellar triangle dark and densely covered with setae: anterior pair of parallel ocellar setae, 2 pairs of posterior setae slightly shorter than anterior ocellar setae, and one pair of postocellar setae. Upper half of occiput bearing row of postocular stout, black setae. Upper occipital setae black, long and stout; lower occipital setae pale, hair-like, and as long as occipital setae of occipital upper half and distinctly dense on postgena. Postpedicel, stylus and scape dark. Scape slightly more than 2× longer than pedicel; postpedicel nearly 3× longer than basal width; stylus length about equal to length of scape. Palpus dark, bearing many long, dark setulae. Labrum dark and glossy; labellum dark and bearing many dark setae, shorter than palpal setae. Thorax dark grey-blue in ground-colour, largely densely grey pruinescent on pleura, less so on scutum. Distinctly glossy and darker grey to black vittae between acr and dc rows extending from pronotal depression to prescutellar depression; brownish tints flank anterior region of vittae separate vittae. Glossiness of scutum vary from specimen to specimen. Posterior corner of postpronotal lobe lustrous and reddish-brown. Pleura with orangish tint along notopleural ridge, as well as on posterior margin of katepisternum; pleural sclerites with varyingly pronounced shadows of brown. Anterior and posterior spiracles distinctly golden. Proepisternum and prosternum with many long, pale setae. Propleuron bearing cluster of hair-like, pale setae shorter than proepisternal setae. Laterals of antepronotum with cluster of long, pale, hair-like setae; middle of antepronotum with many short, pale setae. Postpronotal lobe with crowded cluster of short, hair-like, pale setae. Scutum with dense biserial row of pale, short, fine acr setulae; dc, ial, ad spal setae longer than acr, also fine and hair-like, alternating from pale to dark. Setae become stouter, darker and sparser on prescutellar depression. Anterior notopleuron

with dense cluster of shorter (no shorter than ac), pale, hair-like setae; posterior notopleuron with row of 4 to 6 stout, golden, pale and longer setae. Laterotergite with crowded cluster of pale, hair-like setae. Scutellum with pair of short, dark and converging sctl; pair of subapical scutellar setae, longer and darker; scutellar setae subequal to subapical setae; basal scutellar setae subequal to scutellars. Legs long, entirely dark, darker than thorax ground color. Anterior and lateral regions of mid and hind coxae with numerous pale setae; fore coxae bearing setae only on anterior region. Trochanters also with many shorter, pale, fine, condensed especially on ventral surface of trochanters; fore trochanter bearing few setae. Femora with ventral pubescence (Figs. 3.8, 3.9); femora bearing several rows of antero/posterodorsal, dorsal, antero/posteroventral and ventral setae fine, pale, of varying lengths, but never longer than femoral width, on proximal 1/2, anteroventral setae of apical ½ dark, stouter, half length of femoral width, with cluster of similar setae except shorter on apical 1/5. Mid and fore tibiae (Fig. 3.9) clothed with prominent white pubescence; hind tibiae (Fig. 3.8) with ventral pubescence; mid tibiae with several rows of posterior dark, stout, spine-like setae no longer than pubescence; tibiae with antero/posterodorsal setae dark, stout, subequal to tibial width; mid tibia with posterior preapical cluster of dark, stout, spine- like setae no longer than antero/posterodorsal setae; fore tibia (Fig. 3.9) with several posteroventral and ventral setae dark, stout, as long as antero/posterodorsals. Hind and fore basitarsi similar (Figs. 38, 3.9): posterodorsal, dorsal, and posterior setae dark, stout, as long as 2x basitarsal width, anterodorsal setae also dark and stout, half length of posterodorsals, anterior clothed with many rows of dark, fine setae shorter; antero/posteroventral setae dark, stouter than anterior setae but less than posterodorsals, as long as basitarsal width; mid basitarsus as long as tarsomeres 2-3, with strong ventral pubescence; several anteroventral, ventral, and posteroventral setae dark, stout, spine- like, never longer than basitarsal width; 1 posterodorsal and posterior preapical seta dark, stouter than antero/posterodorsal and dorsal setae, 2x basitarsal width. Wing clear with yellowish veins; all veins complete (except Sc), well sclerotized. Anal vein well-developed and axillary incision acute. Pterostigma distinctly present as darker yellow coloring. Halter pale yellow; stem base darkened; cluster of short, dark setae on base of knob.

Abdomen pale grey; margins of tergites and sternites lighter grey. Abdomen pruinose, covered in long, hair-like, sometimes all pale or sometimes both pale and dark setae, shorter on dorsal middle of tergites. Sternite 6 with dense row of golden, stout setae (Figs. 3.10, 3.43). Tergite 6 swollen such that diameter larger than that of other tergites; setae shorter than rest of abdominal setae; posterior corners of tergite 6 enlarged and curved downwards; margins of tergite 6 brownish. Tergite 7 reduced and U-shaped; sternite 7 also reduced, with two stout, rounded, finger-like projections (Figs. 3.10, 3.43). Tergite 8 slim and barely visible when specimen not dissected. Sternite 8 larger than tergite 8, bearing many ventral long, hair-like, pale setae; sternite 8 and tergite 7 with varyingly dark, lustrous regions. Terminalia (Fig. 3.42) darker than abdomen: epandrium subtriangular and pruinose with long, pale, hair-like setae along outer margin; posterior epandrium joining together with fan-like, blunted tips; middle of epandrium convex. Cercus subtriangular with rounded corners, slightly longer than or subequal to length of epandrium; dorsal margin of cercus with 3 humps, lined with short, fine, dark, erect setae; cercus width less than half width of epandrium. Hypandrium large, visible beneath epandrium, heavily sclerotized, L-shaped, hugging swollen basiphallus, ending in pointed shape; basiphallus dark and glossy; distiphallus also relatively thick; without loops, not extending farther than cercus; dorsal region with black crease running down middle. Ejaculatory apodeme short, not heavily sclerotize, fan-shaped with anterior margin flat, without lateral “wings”. Female. Similar to male except setae shorter than male’s and wings yellow; legs without pennate scales and with setae shorter always shorter than those of male’s. Thoracic setae also shorter than those of males. Remarks. Saigusa (unpublished) further subdivided this species into R. (Ctenempis) sp. 1 and R. (Ctenempis) sp. 2 because of differences in thorax and abdomen setae coloration. However, we consider both species to be the same because of their overlapping geographical distribution and because setae coloration appears to be highly variable and random. Geographical distribution. Nearctic distribution in Figure 3.67.

Rhamphomyia (Pararhamphomyia) diversipennis gp. sp. 1 Figs. 3.11, 3.12, 3.44, 3.68

Material examined. CANADA. British Columbia: Mount Whistler, 6300 m., 27.vii.1973, J.R. Vockeroth (1m#, CNC). Manitoba: Eastern creek nr. Churchill, 2- 30.vii.1952, J.G. Chillcott (23m#, 15f#, CNC). Northwest Territories: Coral Harbour, Southampton Is., 11-20.vii.1948 (13m#, CNC); Kidluit Bay, N. Richards Is., W., 25, 29.vii.1948, J.R. Vockeroth (1m#, 2f#, CNC); Coral Harbour, 15, 16.viii.1959, W.R.M. Mason (3m#, 1f#, CNC); Chesterfield, 19-31.vii., 1-14.viii.1950, J.R. Vockeroth (23m#, 16f#, CNC); Eskimo Point, 18.vii.1950, G.R. Roberts (1m#, CNC); Eskimo Point, 24- 30.vii.1950, G.G. DiLabio (71m#, 37f#, CNC); Tuktoyaktuk, 15.vii.1971, D.M. Wood (2m#, CNC); Near Beechy Lake, 65º14'N, 106º50'W, 18.vii.1966, G.E. Shewell (1m#, CNC); Dubawnt L., 63º18'N, 101º37'W, 2.viii.1966, J.G. Chillcott (2m#, 1f#, CNC); Fraser Lake, 68º45'N, 120º36'W, 19.vii.1969, G.E. Shewell (1m#, CNC). Yukon: Firth River, 14-31.vii., 3.viii.1956, R.E. Leech (6m#, 4f#, CNC); British Mountains, Firth River, 25, 27.vii.1956, R.E. Leech (4m#, 3f#, CNC); British Mountains, Firth River, 25.vii.1956, E.F. Cashman (1m#, CNC). USA. Alaska: Umiat, 10.vii.1959, J.E.H. Martin (2m#, CNC); Umiat, 14.vii.1959, R. Madge (1m#, CNC); Unalakleet, 3-28.vi.1961, B.S. Heming (19m#, 7f#, CNC). Diagnosis. Male. Wing vein dm-cu absent. Vittae dark and shiny (Fig. 3.12). Mid tibia with antero/posterodorsal preapical and proximal setae are as long as 3 times tibial width (Fig. 3.11). Hind femur (Fig. 3.11) with single row of anteroventral setae, subequal to femoral width on proximal and apical third, shorter than femoral width on middle third. Female. Wing cells bm and dm fused and enlarged, almost reaching wing margin, about 3x the length of cup; hind femur glossy with pubescence only on apical third. Description. Wing length 8.3 mm. Male. Head dark in ground-colour, with grey pruinescence on face, frons, postgena and occiput. Holoptic, eye with ommatidia larger on upper half, smaller on lower half. Frons slightly divergent towards antennae; face slightly divergent towards mouthparts; bare, with oral margin dark and shiny. Ocellar triangle with grey pruinescence, with pair of anterior long, stout, dark ocellar setae; 1 pair of posterior ocellar setulae slightly more than 0.5× length of anterior pair and as dark and

as stout as anterior; 3 pairs of dark postocular setae shorter and finer than ocellar setae. Occiput bearing row of postocular setae, fine, black and only present on upper section; longer and finer than stouter and shorter occipital setae; setae on postgena fine, dark, short. Postpedicel, stylus and scape dark, densely pruinose. Scape 2× longer than pedicel; pedicel bulbous; postpedicel about 5× longer than basal width; stylus as long as scape. Palpus grey, densely pubescent, with fine setulae dark. Labrum lustrous and dark reddish- brown; largely yellow; labellum grey, with dark setae. Thorax dark in ground-colour, largely grey pruinescent: posterior corner of postpronotal lobe lustrous and pale brown, npl ridge brown. Scutum with pair of distinct lustrous, dark vittae between acr and dc rows, separated by broad grey pollinose band (Fig. 3.12); and dark, lustrous patch between intalr and spalr. Pleura grey, with brownish shadows of sclerites. Proepisternum grey, with several shorter and some longer, fine, dark setae; prosternum bare, with orange-brown coloring in middle. Antepronotum with row of short, stout, dark setae. Postpronotum with several short, dark, stout setae, with 2 longer, stouter, dark, posterior setae. Scutum with biserial row of fine acr setulae, increasing in length posteriorly, ending before scutellar depression; dc longer and stouter than acr, increasing in length and stoutness posteriorly; spalr setae dark, stout, long, with 1 anterior splr seta stouter and longer, 2 last posterior splr setae stouter and longer; postalar callus bearing 1 dark, stout, long seta and 2-3 shorter, finer, dark setae; npl bearing 4 anterior dark, short, fine setae and 4 posterior dark, longer, stouter setae. Scutellum with pair of long, stout, apical sctl and 1 pair of shorter, stout, dark parallel subapical sctl. Laterotergite with several fine, dark, shorter and longer setae, but never longer than sctl. Anterior and posterior spiracles pale with outer black ring. Legs long, shiny, mostly dark reddish-brown, except coxae dark grey and densely pollinose. Hind and mid coxae with lateral row of dark, short, stout setae, with several scattered lateral finer setae; fore coxae with dorsal row of dark, stout, short setae and with several scattered dorsal, finer setae. Hind and mid trochanters bearing several short, dark, fine, ventral setae; fore trochanter lacking setae. Hind and mid femora with ventral pubescence; hind femur with anteroventral setae dark, fine, never longer than femoral width, shorter in middle of hind femur, decreasing in length posteriorly in fore femur (Fig. 3.11). Mid femur with posteroventral setae dark, subequal to femoral width;

anteroventral setae dark, finer and longer on proximal 1/3, stouter and shorter, spine-like, on apical 2/3. Hind tibia with antero/posterodorsal setae stout, as long as 2x tibial width (Fig. 3.11). Mid tibia with one preapical and one proximal anterodorsal seta stout, as long as 2x tibial width, with antero/posteroventral setae stout, dark, spine-like. Fore tibia with one preapical anterodorsal setae stout, dark, longer than tibial width but shorter than 2x tibial width. Hind tibia with antero/posteroventral setae dark, fine, decreasing in length posteriorly, with several rows of antero/posteroventral and ventral setae on proximal ¼, and shorter, stouter, anterior apical seta dark, stout, longer than ventrals, but shorter than antero/posterodorsals. Hind basitarsi as long as tarsomeres 2-4, with antero/posterodorsal setae dark, fine, subequal to basitarsal width, combined with at least 4 antero/posterodorsal setae stouter and as long as 2x basitarsal width; many rows of posterior setae fine, dark, shorter than antero/posterodorsals; 1 anterior preapical seta and 2 posterior preapical setae, stout, dark, as long as postero/anteroventrals. Mid basitarsi shorter than mid tarsomeres 2-4; postero/anteroventral setae stout, dark, longer than basitarsal width; 1 finer anterodorsal preapical seta longer than postero/anteroventrals. Fore basitarsi similar to mid basitarsi except, antero/posteroventral setae not stout. Wing clear with brownish veins; all veins complete, well sclerotized, except wing vein dm-cu absent, anal vein faint but still reaching wing margin. Anal lobe well- developed; axillary fissure forming rectangular angle. Pterostigma present as darkened area. Halter yellow. Abdomen dark similar to thoracic ground-colour, shiny, with pale segment margins. Margins of tergites lined with long, dark, stout setae; setae on remaining segments finer and slightly shorter; setae on sternites mostly shorter and finer than setae on tergites, although middle seta of sternite segments long and stout. Marginal setae on segment 8 shorter than length of sternite 8. Tergite 8 short. Terminalia (Fig. 3.44) dark: epandrium as large as abdominal width, outer corner pronounced and flattened, bearing many dark, long setae. Epandrium ending in tubercular projection directed inwards and bearing short, dark, spine-like setae. Cercus upright, black, pollinose, short and apex bearing many short, fine setae. Dorsal view of cerci distinctive: forming 2 separate circle-shaped openings, posterior opening smaller than anterior opening. Surstylus slim, short, tear-drop shaped, bearing several short, dark, fine

setae and one long, fine seta at tip. Phallus broad, lustrous and pale orange-brown, hidden within epandrium, sometimes visible sticking out from cerci; phallus tip swollen, almost broader than basiphallus, characterized by dentate projections coming off distiphallus dorsal margin, and tip pointed and curling upwards; some dentate projections might be observed on sides, but might be difficult to discern. Hypandrium short, stout, tubular, hidden within epandrium. Ejaculatory apodeme about as large as cercus, vertical wing almost a 90º angle fan-shape, lateral wings shorter than vertical wing, upper margin steeply sloped; ejaculatory apodeme held outwards posterior to middle of apodeme. Female. Wing narrowed with crossvein bm-cu absent, cells bm and dm fused into enlarged cell, closing at vein CuA1; vein CuA1 short and reaches wing margin at last third of wing length; hind femur glossy with pubescence only on ventral lower half. Geographical distribution. Nearctic distribution in Figure 3.68.

Rhamphomyia (Pararhamphomyia) tipularia gp. sp.1 Figs. 3.13, 3.16, 3.45, 3.69

Material examined. CANADA. Nunavut: Frobisher Bay, 15.viii.1959, W.R. Richards (2f#, CNC); Iqaluit, 63.775122, -68.45927, wet & mesic, Malaise, Repl.3, 17- 25.vii.2010, NBP Field Party (2m#, LEMQ); Kugluktuk, 67.78463, -115.27979, wet & mesic, Malaise, Repl.1-3, 7.vii.-10.viii.2010 (9m#, LEMQ). Yukon: Dempster Hwy. nr. North Fork Pass, 64.59736, -138.31009, wet, Malaise, Repl.2, 24-27.vi.2011, NBP Field Party (1m#, LEMQ). Diagnosis. Male. Fore basitarsus elongated and swollen, broader than width of fore tibia and mid and hind basitarsi (Fig. 3.16). Description. Wing length 15 mm. Male. Head dark grey-blue in ground-colour, with dense grey pruinescence on face, frons, occiput and postgena. Holoptic, eye with ommatidia larger on upper half, smaller on lower half. Frons slightly divergent towards antennae; face slightly divergent towards mouthparts; bare, with oral margin dark and thinly shiny. Ocellar triangle lustrous or only lightly pruinose and brown, with 1 pair of anterior, long, fine, parallel ocellar setae, 1 pair of posterior setae dark, fine, and 1/2 length of anterior ocellars, slightly divergent, and more widely separated than anterior

ocellars, 1 pair of postocellar setae similar to posterior ocellars. Occiput bearing row of postocular setae, dark and longer on upper half of occiput, dark and shorter on lower half of occiput; occipital setae shorter than postocular setae; occipital setae scarce; setae on postgena finer and longer than occipital setae. Postpedicel, stylus and scape mostly lighter than head ground-color; scape and pedicel yellowish; postpedicel darker; pruinose. Scape slightly more than 2× longer than pedicel; pedicel bulbous; postpedicel about 6× longer than basal width and postpedicel only swollen on first 1/2; stylus fine and slightly shorter than basal width. Palpus pale, with short, fine, dark setulae. Labrum lustrous and dark reddish-brown with apex black; labellum grey and bearing fine, short, dark setulae. Thorax dark grey-blue in ground-colour, largely densely grey pruinescent, postpronotum corner lustrous and pale orange-brown, splr ridge pale. Scutum with pair of distinct lustrous dark vittae between acr and dc rows; separated by thick darker grey pollinose band and flanked by lustrous patches, although glossiness of vittae varies and may sometime appear faint and pollinose. Pleura grey, with orange shadows on sclerites, especially concentrated around katepisternum, anepimeron and anepimeron. Proepisternum grey, with 2-3 short, fine, dark setae; prosternum bare and pale orange in middle. Antepronotum with row of short, dark, stout setae. Postpronotum bearing 4 short, dark setae. Scutum with very sparse biserial row of short, fine, dark acr setulae; dc sparse, dark, slightly longer than acr; 1 longer, stouter, dark anterior intlr seta; posterior spalr setae sparse, dark, stout, shorter than intlr; post-alar callus pale and bearing one short, dark seta; npl bearing 1 anterior seta dark, short, and 3 posterior setae dark, longer, stouter. Scutellum with pair of long, convex, converging although never touching, apical sctl; subapical sctl dark, converging and 1/2 length of apical sctl; 2 pairs of sctl similar to subapical sctl although slightly longer but still shorter than apical sctl. Laterotergite with many dark, fine setae. Anterior and posterior spiracles pale, almost white. Legs long, mostly pale orange-brown, except for dark coloring at femoral-tibial junctions and on ventral side of trochanter. Hind coxa with row of dark, stout, lateral setae; mid and fore coxae with several anterior shorter, dark, setae and row of longer, stouter, dark setae at coxal-trochanter junction. Trochanter bearing several fine, dark, short setae on ventral side. Hind femur with anterodorsal setae dark, increasing in length

posteriorly, never longer than femoral width, posteroventrals dark, stout, half length of anteroventrals (Fig. 3.13); posterior setae on proximal 1/3 fine, dark, as long as anteroventrals; antero/posterodorsal and dorsal regions clothed with short, stout setae not erect. Mid femur with 2 rows of anteroventrals, 1 row of longer setae, decreasing in length posteriorly, 1 row of shorter, stouter, setae and 2-3 rows of posteroventral setae, where 1 row has short, stout, dark setae and 2 other rows have finer and longer setae. Fore femur with fine antero/posteroventral setae, decreasing in length posteriorly, antero/posterodorsals similar to those of hind and mid femora. Mid and hind tibiae (Fig. 3.13) with antero/posteroventral setae dark, stout, spine-like; fore tibia with antero/posteroventral surfaces nearly bare; antero/posterodorsal setae as long as at least 2x tibial widths, finer than antero/posteroventrals; fore tibia with antero/posterodorsal setae shorter than half tibial width. Fore basitarsus elongated and swollen, broader than width of fore tibia and mid and hind basitarsi (Fig. 3.16). Mid and hind basitarsi similar, antero/posteroventral setae dark, stouter than antero/posterodorsals and subequal in length to antero/posterodorsals. Wing clear with brownish veins; all veins complete (except anal vein), well sclerotized. Anal vein, although faint and seemingly “broken” in middle, reaches wing margin. Pterostigma present as darkened region. Anal lobe well-developed; axillary fissure forming rectangular angle. Halter white. Abdomen shiny, light brown, lighter than thoracic ground colour; margin of abdominal segments pale. Margins of tergites lined with stouter, longer, dark setae; remaining area of tergites clothed with finer, shorter, dark setae. Middle of sternites bearing several fine, dark, short setae, with 1 longer seta in middle of each sternite. Width of sternite 8 smaller than widths of sternite 1-7; sternite 8 larger than tergite 8; margin of sternite 8 lined with long, dark setae; ventral sternite 8 with cluster of shorter, finer, dark setae; sternite 8 with light pubescence. Margin of tergite 8 bearing many long, fine, dark setae. Terminalia (Fig. 3.45) largely light brown with tip of epandrium pale yellow; cercus pubescent; epandrium base pubescent, bearing several long, dark setae; epandrium width decreasing till constricted in middle; epandrium tip lustrous and rectangular, bearing many dark, stout, short, spine-like setae. Cercus swollen, almost as large as

epandrium width, shorter than epandrium length, ending in slim dorsal finger-like projection, shorter than ½ cercus length; dorsal margin bearing many short, fine, erect setae. Subepandrial lobe narrower than cercus, bifurcate with outer fork about 2x longer than inner fork. Hypandrium short, boot-shaped, triangular tip ending at base of basiphallus; phallus lustrous, orange; basiphallus swollen and more rectangular than rounded; ventral surface of basiphallus bearing short, fine setae; phallus without loops, extending almost farther than epandrium, and curving back in between cerci; distiphallus slimmer than middle phallus. Ejaculatory apodeme small, acutely fan-shaped, with lateral wings shorter than vertical wing. Female. Unknown. Geographical distribution. Nearctic distribution in Figure 3.69

Rhamphomyia (Pararhamphomyia) tipularia gp. sp. 4 Figs. 3.14, 3.15, 3.46, 3.69

Material examined. CANADA. Northwest Territories: Banks Is., Aul. Natnl Park, 73.22412, -119.55255, wet, Malaise, Repl.1, 7-11.vii.2011, (1m#, LEMQ); Norman Wells, 65.29112, -126.62262, wet, Malaise, Repl.1, 14-17.vi.2011, wet, Sweep, Repl.1, 14.vi.2011, NBP Field Party (2m#, LEMQ); Norman Wells, 65.29112, -126.62262, wet, Sweep, Repl.1, 17.vi.2011, NBP Field Party (1m#, LEMQ) Nunavut: Cambridge Bay, 69.11993, -105.42065, mesic, Malaise, Repl.1-3, 7-19.vii.2011, mesic, Pan, Repl.3,2, 7- 15.vii.2011, wet & mesic, Sweep, Repl.1,2, 13.vii.2011, NBP Field Party (11m#, LEMQ); Lake Hazen, Ellesmere Is., 81.83179, -71.44115, wet, Sweep, Repl.1, 17.vii.2011, NBP Field Party (1m#, LEMQ); Iqaluit, 63.76144, -68.57352, wet & mesic, Malaise, Repl.1-3, 17-29.vii.2011, wet & mesic, Pan, Repl.1,2, 17-25.vii.2011, wet & mesic, Sweep, Repl.1,3, 21.vii.2011, NBP Field Party (149m#, LEMQ); Kugluktuk, 67.78508, -115.27573, wet & mesic, Malaise, Repl.1-3, 25-29.vi., 7-26.vii., 2- 10.viii.2010, wet & mesic, Pan, Repl.1,3, 8-19.vii.2010, wet, Sweep, Repl.1, 30.vi.2010, NBP Field Party (1419m#, LEMQ). Yukon: Dempster Hwy, km. 128, 64.9292, - 138.2702, Malaise near pond, 13-15.vii.2013, T. Wheeler, S. Rocherfort, E. Vajda (1m#, LEMQ); Demspter Hwy near North Fork Pass, 64.60629, -138.35637, wet & mesic,

Malaise, Repl.1,2, 21-27.vi.2011, wet, Pan, Repl.2, 21-24.vi.2011, wet, Sweep, Repl.2, 27.vi.2011, NBP Field Party (50m#, CNC). GREENLAND. Nedre Midsommer Sö, T32- 4, 29.vi.1966, T31-5, T32-5, T33-5, T33-6, T34-4, T34-5, M1-8, M1-14, M2-11, 3- 29.vii.1966, Can. Peary Land Expd. (24m#, CNC). Diagnosis. Male. Laterals of abdominal tergite 5 bearing distinctive lateral cluster of stouter and shorter, almost spine-like setae decreasing in length posteriorly. Female. Body darker than that of male; abdominal tergite 5 not bearing distinctive cluster of shorter, stouter setae. Description. Wing length 6.5–7.5 mm. Male. Head grey-blue in ground-colour, with silvery greyish pruinescence on face, frons, postgena and occiput. Holoptic, eye with ommatidia larger on upper half, smaller on lower half. Frons divergent towards antennae; face mostly parallel, although slightly divergent towards mouthparts; bare with oral margin black and shiny. Ocellar triangle dark grey, pruinose, with anterior pair of short, dark, stout divergent ocellar setae and posterior pair of setae similar to anterior ocellars; 2 pairs of postocellar setae similar to ocellars. Occiput bearing row of dark, fine postocular setae, only present on upper half of occiput; occipital setae dark, stouter and slightly shorter than postoculars; postgena setae shorter and finer than occipitals. Postpedicel, scape, pedicel and stylus dark brown with dense grey pruinescence; pedicel more pale brown than postpedicel. Scape nearly 2x length of pedicel; pedicel width greater than scape width; postpedicel basal width greater than lower postpedicel width; postpedicel subequal to basal width; stylus longer than scape but never longer than scape. Palpus dark grey and pruinose, with longer and shorter dark setulae. Labrum lustrous and dark brown; labellum grey with dark setae. Thorax blue in ground-colour, largely lightly silvery-grey pruinescent; posterior corner of postpronotal lobe lustrous and pale brown; brownish shadows on postalar callus; region between posterior spiracle and halter base brown. Scutum with pair of distinct brownish brownish vittae between acr and dc rows, separated by blue band, flanked by posterior brownish patches. Pleura blue with light silvery-grey pruinescence, with brownish shadows on sclerites; margin of katepisternum and meron lustrous and dark. Proepisternum with several short, stout, dark setae; prosternum bare. Antepronotum with row of short, stout, dark setae. Postpronotum with 3-4 short dark setae and 1 long,

stout posterior seta. Scutum with biserial row of stout, dark acr setulae, ending before sctl depression; dc similar to acr, except slightly longer and increasing in length and stoutness posteriorly; intlr setae sparse, stout, no longer than dc; npl with 3 short, dark, stout anterior setae and 2 longer, stouter posterior setae. Scutellum with pair of long, stout, convergent and crossing over apical sctl; subapical sctl stout, slightly shorter and convergent; shorter pair of posterior sctl. Laterotergite with about 7 short, stout, dark setae. Anterior and posterior spiracles orange-brown. Legs long, mostly dark brown, darker than thoracic and abdomen colouring, coxae more grey-blue with distinct brown shadows, trochanter with ventral black coloring, hind and mid tibial-femoral junctions blackened, fore tibial-femoral junction pale. Femoral pruinescence may give silvery reflections. Hind and mid coxae with row of dark, stout, short lateral setae, sparser on hind coxae than on mid coxae; fore coxae with dorsal/anterior rows of short, dark, stout setae; setae shorter and finer closer to thoracic- coxal junction. Trochanter bearing only several shorter, finer ventral setae; fore trochanter nearly bare. Hind femur with ventral pubescence, anteroventral setae dark, increasing in length towards apical 1/3 (Fig. 3.14), setae always shorter than femoral width; mid femur with posteroventral setae dark, subequal to femoral width; fore femur with antero/posteroventral setae dark, finer than setae of mid and hind femora, never longer than femoral width, with 2 stouter anterodorsal preapical setae. Hind tibia with dorsal and ventral white pubescence, with at least 4 antero/posterodorsal setae dark, stout, subequal to 2x antero/posteroventral setae, with preapical cluster of stout, dark, short setae (Fig. 3.14). Mid tibia with 3 anterodorsal setae dark, stout, as long as 3x tibial width, apical 2/3rds of anteroventral row with dark, stout, spine-like setae. Fore tibia with antero/posteroventral setae very fine, ½ tibial width; antero/posterodorsal setae fine, 2x length of antero/posteroventral setae; preapical antero/posterodorsals and antero/posteroventrals stouter. Hind basitarsal width slightly larger than mid and fore basitarsal widths, setae subequal to or shorter than basitarsal widths, respectively; hind basitarsus with 2 preapical setae dark, stouter than remaining basitarsal setae; mid and fore basitarsi slim, clothed with white pubescence on posterior and ventral regions, preapical ring of dark, stouter, short setae. Wing clear with yellowish and whitish veins; all veins complete (except anal vein

faint), well sclerotized. Seta on base of costa longer and stouter than remaining costal setae. Anal lobe well-developed; axillary fissure forming rectangular angle. Pterostigma absent. Halter white. Abdomen pruinose silvery grey-blue, although abdomen less blue than thorax. Margins of sclerites pale. Abdomen covered in dark setae; setae finer and longer on first several tergites, stouter and shorter towards terminalia; laterals of tergite 5 bearing distinctive cluster of stouter and shorter, spine-like setae. Sternite 8 enlarged, tergite 8 smaller than tergite 1-7; margin of sternite 8 bearing stout setae longer than lower abdominal setae; upper corner of sternite 8 lustrous and dark; tergite 8 slim, less than half width of sternite 8; margin of sternite 8 bearing several short, dark setae. Terminalia (Figs. 3.15, 3.46) largely dark grey and pruinose: epandrium broadly triangular-shaped, with apex rounded and indented; epandrium grey and pollinose, except apex lustrous and pale orange-brown; dorsal and ventral margins of posterior epandrium bearing several dark setae, never longer than abdominal setae; dorsal margin of epandrium apex black, bearing many dark, stout setae, conglomerated together, thus forming spindle-like structure, convergent towards “spindle” of other epandrium (Figs. 3.15, 3.46). Cercus grey, pollinose, shorter than epandrium length; dorsal margin of posterior cercus bearing several dark, stout setae, no longer than cercus width; cerci forming tear-drop-shaped opening; surstylus lustrous, pale brown, longer and slimmer than cercus. Subepandrial lobe bifurcate, with two slim finger-like projections; outer finger-like projection darkened, longer than inner finger-like projection, bearing 1 long, fine, black seta; inner finger-like projection bearing single spine-like, stout, short seta, only visible upon dissection. Hypandrium slim, short, rounded L-shaped, triangular tip hugging base of basiphallus; basiphallus broad and pale, thinning out significantly and darkening; phallus without curvatures, never extending farther than terminalia, folded back in between epandrium. Ejaculatory apodeme large, vertical width larger than horizontal width, lateral wings rounded and shorter than vertical wing; gonocoxal apodeme fan-shaped, with distinctive feature. Female. Body darker than that of male; abdominal tergite 5 not bearing distinctive cluster of shorter, stouter setae. Halter yellow. Eyes dichoptic. Geographical distribution. Nearctic distribution in Figure 3.69.

Rhamphomyia (Pararhamphomyia ?) simplex Zett Figs. 3.17, 3.47, 3.70

Material examined. ICELAND. Breddalsvik, moist Luzula empetrum moss pocket, 1360/1/1-, 7.vii.1981, J.A. Downes (12m#, 4f#, CNC); Myvata, road to Skutustadir, 1357/2/15-, 3.vii.1981 (1m#, 3f#, CNC); Laxa River, Myvata, 1357/1/11-, 3.vii.1981, J.A. Downes (11m#, 4f#, CNC). Diagnosis. Male. Body pale, abdomen with light powdery blue pruinescence, with thorax remarkably powdery light blue; tergite 8 long, subequal to length of epandrium; phallus without loops and broad: base as broad as pedicel; phallus with dark suture running through the middle (Fig. 3.47); apex of hind tibia with dark setae no longer than width of hind tibia and projecting out from under pale apical comb on posteroventral side of hind tibia (Fig. 3.17). Female. Similar to male. Description. Wing length 7.3–7.5 mm. Male. Head grey-blue in ground-colour, with dense greyish-silvery pruinescence on face, frons, postgena and occiput. Holoptic, eye with ommatidia larger on upper half, smaller on lower half. Frons divergent towards antennae; face slightly divergent towards mouthparts; bare with oral margin reddish- brown and shiny. Ocellar triangle grey-blue, with greyish-silvery pruinescence, with anterior pair of long, dark, parallel ocellar setae and 2 pairs of shorter, less than ½ length of anterior ocellar setae, dark posterior setae; postocellar setae finer and slightly shorter than posterior ocellar setae. Upper half of occiput bearing row of long, dark postocular setae; occipital setae black, shorter and stouter than postocular setae; postgena setae subequal to postocular setae, but finer than occiptals. Antennae largely orange-brown and densely pruinescent; postpedicel and stylus darker than scape and pedicel. Scape only slightly longer than pedicel; postpedicel about 4× longer than basal width; stylus subequal to scape length. Palpus dark reddish-brown, with dark setulae, some stouter than others. Labrum lustrous and dark reddish-brown; apex pale; labellum grey and with dark setae. Thorax silvery light grey-blue in ground-colour, largely densely grey pruinescent; posterior corner of postpronotal lobe lustrous and yellow-brown, npl ridge pale orange-

yellow, postalar callus yellow-brown; posterior intlr region with brown patch. Scutum with pair of very faint brownish vittae between acr and dc rows, slightly more prominent anteriorly. Pleura light silvery grey-blue, with pale brown on margins of some sclerites, especially margin of katepisternum and meron, region between posterior spiracle and halter base pale brown. Proepisternum with 2-3 long, stout, dark setae; prosternum bare. Antepronotum with row of dark, stout setae, longer on sides, shorter in middle. Postpronotum with several short, stout, dark setae and 1 long seta located posteriorly. Scutum with biserial row of dark, stout acr setulae, increasing in length posteriorly; dc similar to acr, increasing in length posteriorly, and slightly longer than acr; intlr setae dark, stout, longer than dc, increasing in length posteriorly, posterior-most intlr seta longest, stoutest; notopleuron with 3-4 anterior short, fine, dark setae, 1 anterior longer, stouter seta, and with 3-4 posterior npl stouter and more than 2x length of anterior npl; postalar callus with single long, stout seta. Scutellum with pair of long, dark, stout apical sctl, subapical sctl slightly shorter than apical sctl and 1 pair of finer sctl less than 1/3 of subapical sctl. Laterotergite with dense cluster of long, stout, fine setae all of subequal lengths to each other. Anterior and posterior spiracles pale. Legs long, entirely light brown, except coxae grey-blue with brownish colouring and coxae-thoracic junction pale brown. Hind coxae with distinct row of dark, stout lateral and apical setae, mid and fore coxae with anterior long, dark, stout setae, interspersed with finer, shorter setae. Hind trochanter with many ventral dark, stouter and finer setae, mid and fore trochanters with fewer, shorter, and finer ventral setae. Hind femur with ventral pubescence (Fig. 3.17); mid and hind femora with anteroventral setae dark, subequal to femoral widths, and increasing in length posteriorly in hind femur; antero/posteroventral and ventral surfaces of fore femur nearly bare; femora with setae of antero/posterodorsal and dorsal regions dark, shorter than femoral width and not erect. Mid and hind tibiae (Fig. 3.17) with dark, stout antero/posterodorsal setae, about as long as 2x tibial widths, respectively; antero/posteroventrals dark, finer than antero/posterodorsals, shorter than tibial widths. Fore tibia with dark, stouter antero/posterodorsals/ventrals, finer than those of mid and hind tibiae, except preapical antero/posterodorsal setae as stout and as long as antero/posterodorsals of mid tibia. Hind (Fig. 3.17) and mid basitarsi with preapical antero/posterodorsal setae stouter and darker

than remaining basitarsal setae; mid basitarsus with antero/posteroventrals dark, stouter, and longer than or subequal to basitarsal width. Wing clear with yellowish-white veins; all veins complete, well sclerotized (except anal vein faint). Costal seta short, stout, dark, and 1 basal costal seta longer, stouter than remaining costal setae. Anal lobe well-developed; axillary fissure forming rectangular angle. Pterostigma present in form of darker yellow colouring. Halter pale, stems darker yellow, knobs white-yellow. Abdomen darker than thorax, mostly brownish-grey, with silvery pruinescence, with some bluish shadows, especially sternites 1 and 2; sclerites margins paler. Tergites covered in dark setae, longer laterally, shorter towards middle of tergites; sternites with similar setae, although fewer. Sternite 8 larger than tergite 8; sternite 8 width no longer than remaining sternite widths; marginal setae of sternite 8 shorter than lateral tergites setae; posterior corner of sternite 8 enlarged and rounded. Anterior margin of tergite 8 with 3 curves. Terminalia (Fig. 3.47) largely dark reddish-brown: hypandrium lustrous and dark brown, with posterior margin lined thinly with black colouring; epandrium pruinose, swollen and rounded, ending with squared knob-like projection, and sharp, slim, finger- like projection converging inwards; epandrium margins bearing dark setae, decreasing in length and stoutness from base to dorsal margin. Cercus small, stout, rounded, pruinose, dorsal margin bearing fine, erect setae, shorter than cercus length; surstylus larger than cercus, about 2x cercus length, of similar width to cercus, and dorsal margin bearing many long, fine, hair-like setae only visible upon dissection; surstylus ending with sharp spine-like projection at apex. Hypandrium rectangular, short, ending before swollen, pale basiphallus. Basiphallus and middle phallus joined at end of hypandrium, V-shaped, forming acute angle; middle phallus broad, but less broad than basiphallus portion after hypandrium, hidden within epandrium, never extending farther than epandrium; distiphallus darker orange-brown, slimmer than middle phallus, without loops, curving towards cercus at rounded 90º angle. Ejaculatory apodeme almost as large as epandrium, tear-drop-shaped, lateral wings shorter than vertical wing. Female. Similar to male. Geographical distribution. Known distribution in Figure 3.70.

Rhamphomyia (Pararhamphomyia) pusilla gp. sp. 7 Figs. 3.18, 3.19, 3.48, 3.71

Material examined. CANADA. Northwest Territories: Masik River, Banks Is., 4- 83.vii.1968, W.R.M. Mason (18m#, 6f#, CNC); Masik River, Banks Is., 16.vii.1968, G.E. Shewell (1m#, 1f#, CNC); Mackenzie Delta, Hooper Is., 28.vi.1971, D.M. Wood (2m#, CNC); -21 m.e. Tuktoyaktuk, 2-25.vii.1971, D.M. Wood (17m#, 1f#, CNC); -30 m.e. Tuktoyaktuk, 18.vi.1971, D.M. Wood (3m#, CNC); Tuktoyaktuk, 17.vi.1971, D.M. Wood (3m#, CNC); Coppermine, 20.vii.1951, S.D. Hicks (2m#, 1f#, CNC). Nunavut: Kugluktuk, 67.78463, -115.27979, 25.vi.-26.vii.2011, wet & mesic, Malaise, 29.vi.- 3.vii.2011, mesic, Repl.1-3, NBP Field Party (147m#, LEMQ). Yukon: M1.51, Dempster Hwy., 17-27.vi.1973, G. & D.M. Wood (9m#, CNC); Demspter Hwy., nr. North Fork Pass, 64.58353, -138.26843, 24-27.vi.2011, wet & mesic, Malaise, Sweep, Repl.2, NBP Field Party (36m#, LEMQ). Diagnosis. Male. Mid-leg distinctly modified (Fig. 3.19): femur strongly curved upwards; tibia curved inwards, with 3-5 dark, stout mid posterodorsal setae dark, stout, longer than 2x tibial width; basitarsus swollen in middle with several rows of stout, dark mid dorsal antero/posterodorsal setae, always shorter than 2x basitarsal width. Hind tibia with several rows of stout proximal anterodorsal and anterior setae, never shorter than half the tibial width, and never longer than twice the tibial width (Fig. 3.18). Description. Wing length 7.8–8.3 mm. Male. Head dark grey-blue in ground- colour, with greyish pruinescence on face, frons and occiput and postgena. Holoptic, eye with ommatidia larger on upper half, smaller on lower half. Frons divergent towards antennae; face slightly divergent towards mouthparts; bare, with oral margin dark grey- blue and shiny. Ocellar triangle dark, subshiny, with 1 anterior pair of dark, stout, long, parallel ocellar setae, 1 pair of posterior ocellar setulae similar to anterior ocellar setae, with 2nd pair of posterior ocellar setae where setae widely separated from each other, shorter and finer than anterior ocellar setae; 2 pairs of finer, shorter postocellar setae. Occiput bearing row of fine, long, black postocular setae, only present on upper section; occipital setae black, long and fine; postgena bearing dark and long setae finer than

occipital setae. Postpedicel, stylus and scape entirely pruinose, dark and darker than head ground-colour. Scape no longer than 2× longer than pedicel; pedicel bulbous, width larger than scape width; postpedicel nearly 4× longer than basal width; stylus no longer than pedicel length. Labrum lustrous and dark reddish-brown. Labellum grey with many dark, short, fine setulae. Thorax dark, mostly grey-blue in ground-colour, largely densely grey pruinescent. Scutum with pair of distinct brownish, pruinose vittae between acr and dc rows, separated by broad greyish-blue stripe in middle. Pleura largely dark grey-blue, often with brownish tinges on sclerites; posterior corner of postpronotal lobe lustrous and dark orange-brown; region between posterior spiracle and halter base not pruinose and orange- brown; npl ridge thinly pale. Proepisternum grey-blue, pruinose, with several long, dark, stout setae; prosternum bare. Antepronotum with row of short, stout dark setae. Postpronotum with several long, dark setae and 1 longer, stouter, dark seta located posteriorly. Scutum with biserial row of fine, dark acr setulae; dc similar to acr, increasing in length posteriorly, last dc setae stouter; intlr seta similar to ac; spalr setae longer and stouter than dc; postalar callus bearing single anterior stout, long setae; notopleuron bearing 5 anterior dark, long, stout setae grouped together and 3 posterior longer, stouter setae. Scutellum with 1 pair of long, dark, stout apical sctl crossing over each other; 1 pair of dark, stout, convex subapical sctl slightly shorter than apical sctl; 1 pair of sctl dark, stout, long, and slightly convex. Anterior and posterior spiracles pale with outer dark ring. Legs long, entirely dark brown, except for darkened femoral-tibial and femoral- trochanter junctions. Hind coxae with several dark, fine lateral setae, anteriorly and laterally on mid coxae, only present anteriorly on fore coxa. Hind trochanter with several dark, fine setae, increasing in length towards femoral-trochanter junction. Mid and fore coxae nearly bare. Hind femur with single apical anteroventral and posteroventral rows of stout, dark setae longer than dorsal setae (Fig. 3.18); apical single anteroventral row absent on mid femur; mid femur curved upwards (Fig. 3.19), with several rows of anterior proximal setae never longer than tibial width; fore femur with postero/anteroventral setae finer but no shorter than those of mid and hind femora (Figs. 3.18, 3.19). Hind tibia with several rows of proximal anterodorsal and anterior setae

stout, never shorter than half tibial width, and never longer than 2x the width; posteroventral pubescence on lower ½ of hind tibia; anteroventral setae of hind tibia subequal to tibial width (Fig. 3.18). Mid tibia curved; posterodorsal setae stouter than stoutest setae of hind tibiae, no shorter than 2x tibial width; apical anterodorsal setae increasing in length and stoutness posteriorly; 2-3 rows of proximal stout, dark anteroventral setae shorter than anterodorsals; 1 posterior preapical seta dark, stout, longer than antero/posteroventral setae but shorter than anteroventral setae (Fig. 3.19). Fore tibia with anterodorsal setae fine, decreasing in length posteriorly, such that proximal setae slightly longer than tibial width; posterodorsal setae longer than tibial width; ventral pubescence. Mid basitarsus swollen in middle with several rows of stout, dark dorsal and antero/posterodorsal setae, always shorter than 2x basitarsal width (Fig. 3.19). Hind basitarsus and hind tarsomeres with preapical ring of dark, stout setae longer than antero/posterodorsal, dorsal, postero/anteroventral and ventral setae. Wing clear with yellowish veins; all veins complete (except anal vein faint towards wing margin), well sclerotized. Anal lobe well-developed; axillary fissure forming rectangular angle. Base of costa bearing dark, stout seta longer than remaining costal setae. Pterostigma present, darkened yellow colouring, although faint. Halter dark yellow with knob blackened. Abdomen dark grey-blue with pale segment margins. Abdomen covered in dark setae longer on laterals of tergites and in middle of sternites, shorter in middle of tergites, nearly absent on laterals of sternites. Sternite 8 enlarged, margin bearing many longer, stouter setae. Tergite 8 reduced, barely visible, held vertically more than horizontally. Terminalia (Fig. 3.48) entirely dark, more black than grey-blue, largely pruinescent: epandrium decreasing in width anteriorly and curving upwards; posterior corner of epandrium sharp, nearly rectangular; epandrium ventral margin bearing many dark, stout, long setae decreasing in length towards epandrium tip; setae at tip of epandrium numerous, almost spine-like. Cercus divided into anterior and posterior lobes by tight, short constriction; anterior lobe “tear-drop”-shaped, densely pruinose, bearing many short, fine, dark setae, smaller than posterior lobe; posterior lobe more rectangular and less pollinose and bearing numerous fine, short setae. Hypandrium slim, short, not heavily sclerotized, ending at 1/3 of swollen basiphallus; membranous region of

hypandrium visible upon dissection; basiphallus tear-drop-shaped; phallus dark reddish- brown, slim, hair-like, not forming loops, curving back into cerci or epandrium. Ejaculatory apodeme as large as anterior cercus lobe, oval-shaped, with lateral wings subequal to vertical wing. Female. Not examined. Geographical distribution. Nearctic distribution in Figure 3.71.

Rhamphomyia (Pararhamphomyia) fuscula gp. sp.1 Figs. 3.20, 3.49, 3.72

Material examined. CANADA. Northwest Territories: Lake Harbour, Baffin Is., 5, 9.viii.1935, W.J. Brown (2m#, 2f#, CNC); Frobisher Bay, Baffin Is., 10-31.vii.1948, F.G. DiLabio (5m#, 8f#, CNC); Frobisher Bay, Baffin Is., 10.vii.1948, T.N. Freeman (1f#, CNC); Frobisher Bay, 10.viii.1959, J.A. Downes (2m#, 1f#, CNC). Yukon: Dempster Hwy, km.128, 129, 64.9292, -138.2702, Malaise near pond, 13-20.vii.2013, T. Wheeler, S. Rochefort, E. Vajda (26m#, LEMQ); Dempster Hwy near North Fork Pass, 64.59736, - 138.31009, wet & mesic, Malaise, Repl. 2,3, 24.vi.-1.vii.2011, NBP Field Party (28m#, LEMQ). Quebec: Payne Bay, 1-31.viii.1958, E.E. MacDougall (176m#, 131f#, CNC); Payne Bay, 7.vii.1954, E.E. Sterns (1m#, CNC); Payne Bay, 8.vii.1954, H. Huckel (3m#, 14f#, CNC); Payne Bay, 9.vii.1954, R. McCondochie (3m#, CNC); Payne Bay, 15- 26.vii.1954, R. McCondochie (3m#, 5f#, CNC); Payne Bay, 18, 20.vii.1958, E.E. MacDougall (3m#, 1f#, CNC); Point Harrison, 5-27.vii.1949, D.P. Whillans (26m#, 16f#, CNC); Sugluk, 11, 14, 28.vii.1954, H. Huckel (2m#, 1f#, CNC). Labrador: Nutak, 26.vii.1954, E.E. Sterns (1f#, CNC). Diagnosis. Male. Hind femur curved upwards with row of posteroventral setae that are dark, stout, spine-like, and shorter than ventral pubescence. Distiphallus broader than middle phallus; expanded with ridge of teeth-like projections; phallus tip with incisions. Description. Wing length 8.8–9.3 mm. Male. Head blue in ground-colour, with silvery pruinescence; postgena and occiput slivery-blue and pruinose. Holoptic, eye with ommatidia of larger size on upper half, smaller size on lower half. Frons small and

triangular, subequal to width of bulging ocellar triangle. Face parallel-sided towards mouthparts; bare, with oral margin dark and shiny. Ocellar triangle darker grey-blue, subshiny, with 2 pairs of dark, parallel ocellar setae; anterior pair 3x length of posterior pair; postocellar setae subequal to posterior ocellar setae. Occiput bearing row of black, slender, long postocular setae on upper half of occiput; occipital setae also black, but shorter and stouter than postoculars. Antenna dark and pruinose; scape and pedicel light brown. Scape slightly longer than pedicel; pedicel bulbous; postpedicel about 4x basal width; stylus no longer than scape and pedicel. Palpus black, pruinose with dark setulae; labrum largely dark brown and glossy; labellum with dark setae. Thorax blue in ground-colour, with silver pruinescence. Scutum with pair of faint blue, pruinescent vittae between acr and dc rows, flanked with brown patch. Pleura entirely blue, sometimes with paler, orange-brown highlights. Antepronotum blue, pruinose, with row of stout, dark, short setae. Postpronotum with 4-5 anterior short, stout, dark pprn and 1 stout, dark posterior pprn, 2x length of anterior pprn. Scutum with long, dark, and biserial acr and multi-serial dc; dc slightly longer than acr, increasing in length posteriorly; 3 long, stout, dark npl and several shorter dark setae; spalr setae dark, stout, slightly longer than dc. Scutellum with converging apical sctl, and 1 outer pair of sctl setae finer and ¼ of length of apical sctl. Laterotergite with crowded cluster of long dark setae. Anterior and posterior spiracles dark. Legs dark brown, not lustrous. Coxae powdery blue; hind coxa with lateral and anterior rows of dark, stout setae; mid and fore coxae with anterior row of setae similar to those of hind coxa. Hind trochanter with several dark setae finer than coxal setae; mid and fore trochanters bare. Femora clothed with silvery pollinosity; hind femur with anteroventral setae dark, stout, shorter than ½ femoral width, except for apical setae 2x length of remaining anteroventrals (Fig. 3.20); mid and fore femora similar to hind femur except anteroventrals of mid femur shorter and stouter than anteroventrals of hind femur; anteroventrals of fore femur finer than those of hind femur. Tibiae clothed with silvery pollinosity; hind tibia with antero/posterodorsals dark, stout, longer than tibial width but shorter than 2x tibial width; cluster of many short, fine antero/posteroventrals and ventrals on apical 1/3 (Fig. 3.20). Mid tibia with antero/posterodorsals stout and dark, but shorter than tibial width, anteroventral setae shorter than antero/posterodorsals, dark,

stout, except for anteroventrals of proximal 1/5 finer. Fore tibia with antero/posterodorsals dark, stout, longer than those of mid tibia but shorter than those of hind tibia. Basitarsi (Fig. 3.20) of similar lengths to each other, with antero/posteroventrals dark, stout, shorter than basitarsal widths; antero/posterodorsals finer, lighter and shorter than antero/posteroventrals; hind and fore basitarsi with many short, light, fine setae on posterior regions. Terminalia (Fig. 3.49) dark, clothed with silvery pollinosity; phallus lustrous and dark brown. Epandrium subrectangular, with anterior corner elongated and projecting upwards; lateral margin of epandrium tip bearing many fine, short, erect, pale setae; dorsal margin of epandrium tip bearing distinguishing row of dark, stout setae, longer than setae of lateral margin, forming dark comb-like structure. Upon dissection, inner vacuole observed in anterior region of epandrium. Cercus densely pollinose, with “fuzzy” appearance; cercus small, short, not reaching half epandrium length, lobe-shaped. Subepandrial lobe slightly narrower than cercus, short, rounded, bifurcate such that 2 lobes compose subepandrial lobe (mitten-shaped); 2nd lobe directed inwards. Hypandrium short, slim, rectangular, hugging start of basiphallus; basiphallus slightly more swollen than middle phallus; distiphallus cylindrical and more swollen than middle phallus; phallus tip with “flower-like” appearance with pointed short projections; dentate structures observed on these projections only upon dissection. Ejaculatory apodeme slim, small; dorsal margin of vertical wing with 3 distinctive sides; lateral wings shorter than vertical wing. Ejaculatory apodeme thinly fused to phallus. Female. Not examined. Geographical distribution. Nearctic distribution in Figure 3.72.

Sg. Dasyrhamphomyia

Rhamphomyia (Dasyrhamphomyia) nigrita Zetterstedt, 1838 Figs. 3.21, 3.50, 3.53, 3.73

Rhamphomyia nigrita, Zetterstedt, 1838: xx. Rhamphomyia conservativa Malloch, 1918: 49, syn. nov.

Type material examined. PARATYPE: CANADA. Northwest Territories: R. conservatica No.219, Bernard Harbour, 10.vii.1915, Canadian Arctic Expedition, F.D. (1f#, CNC). HOLOTYPE: CANADA. Northwest Territories: R. conservatica, No. 219, Mall., W. of Bernard Is., 19.vii.1916, Coll. Expedition (1m#, CNC). ALLOTYPE: CANADA. R. conservatica, No. 219, Mall., W. of Bernard Is., 14.vii.1916, Canadian Arctic Expedition, F. Joanhansen Coll. (1f#, CNC). Additional material examined. CANADA. British Columbia: Summit Lake, Mi392 Alaska Hwy, 5000’, 6.vii.1959, 4700', 7.vii.1959, R.E. Leech (2f#, CNC). Manitoba. Churchill, 14.vii.1947, C.R. Twinn (2m#, CNC); Churchill, 12.vii.1948, W.R. Richards (1m#, CNC); Churchill, 29.vi.1948, G.E. Shewell (1f#, CNC); Fort Churchill, 4.vii.1952, J.G. Chillcott (1f#, CNC); Eastern Cr. Nr. Churchill, 25.vi.1952, J.G. Chillcott (1f#, CNC); Churchill, 10-20.vi.1930, 47, 48, 52, 54, Lot 298, O. Bryant (8m#, 4f#, CNC). Northwest Territories. Tranquary Fjord, Ellesmere Is., 18.vii.1963, H.K. Rutz (1m#, CNC); 81º49’N,71º18’W, Hazen Camp, Ellesmere Is., 17.vii.1962, R.B. Madge (1m#, CNC); Keewatin, 64º15’N, 89º30’W, 22.vii.1950, J.G. Chillcott (1m#, CNC); Repulse Bay, 6, 23.vii.1950, P.F. Bruggemann (19m#, CNC); Repulse Bay, 4, 21, 26.vii.1950, J.E.H. Martin (1m#, 4f#, CNC); Repulse Bay, 23.vii.1950, P.F. Bruggemann (17f#, CNC); Chesterfield, 6-30.viii.1950, J.R. Vockeroth (9m#, 10f#, CNC); Chesterfield, 6-27.viii.1950, J.G. Chillcott (1m#, 5f#, CNC); Chesterfield, 31.viii.1950, J.G. Chillcott (4m#, 14f#, CNC); Wager Bay, 65º15’N, 88º00’W, 22.vii.1950, J.G. Chillcott (1m#, 1f#, CNC); Spence Bay, 2-30.vii.1951, J.G. Chillcott (31m#, 28f#, CNC); Spence Bay, 14-30.vii.1951, A.E.R. Downes (4m#, 13f#, CNC); Eskimo Point, 12, 31.vii.1950, G.G. DiLabio (4m#, 14f#, CNC); Eskimo Point, 21, 24.vii.2950, G.R. Roberts (1m#, 3f#, CNC); Bernard Harbour, 1914, 17.viii.1915, 10.vii.1916, 1043, Canadian Arctic Expedition, F.D. (2f#, 1m#, CNC); Coppermine, 14-20.vii.1951, S.D. Hicks (2m#, 4f#, CNC); Melville Pen., 19-22.vii.1948, C. McArthur (2m#, CNC); Lady Melville L, 93º15’W, 69º25’N, 3.vii.1951, J.G. Chillcott (1m#, 1f#, CNC); Salmita Mines, 64º05’N, 111º15’W, 4.vii.1953, J.G. Chillcott (1m#, 4f#, CNC); Fraser Lake, 68º45N, 120º36W, 19.vii.1969, G.E. Shewell (5m#, 5f#, CNC); Aklavik, 1-18.vii.1931, Lots 243, 245, 297, Bryant (5f#, CNC); Atkinson Pt., Tukrtoyaktuk Penn., 19.vii.1971,

D.R. Oliver (1m#, CNC); Tuktoyaktuk, 7, 15.vii.1971, D.M. Wood (9m#, 8f#, CNC); Banks Is., Aul. Natnl Park, 73.233284, -119.54719, 7-19.vii.2011, wet & mesic, Pan, 10- 19.vii.2011, wet & mesic, Sweep, Repl. 1-3, NBP Field Party (41m#, LEMQ); Eureka, 6- 30.vii.1954, P.F. Bruggemann (2m#, 5f#, CNC); Eureka, 2, 3.viii.1963, J.R. Vockeroth (4m#, CNC); Frobisher Bay, 2, 8.vii.1948, F.G. DiLabio (2f#, CNC); Kidluit Bay, N. Richard Is., W., 26-31.vii., 29.viii.1948, J.R. Vockeroth (3f#, CNC); Kidluit Bay, N. Richards Is. W., 27.vii.1948, T.J.R. Vockeroth (1m#, 1f#, CNC); 60 mi. N. of Aklavik, Police cable on Moose River, 14.vii.1930, O. Bryant (1m#, 4f#, CNC); Frobisher Bay, Baffin Is., 26.vi.1948, T.N. Freeman (1m#, CNC); Holman, Victoria Is., 1-27.vii.1952, 3.vii.1952, Area A, D.P. Grey (3m#, 6f#, CNC); 81º49'N, 71º18'W, Hazen Camp, 1- 30.vii.1962, R.B. Madge (24m#, 9f#, CNC); Hazen Camp, Ellesmere Is., 31.vii., 5- 17.viii., 20-22.vii.1961, D.R. Oliver (12m#, 2f#, CNC); Hazen Camp, NE. 317, 81º49'N, 71º16'W, 14.vii.1962, D.R. Oliver (1m#, CNC); Lake Hazen, Ellesmere Is., 18.viii.1961, D.R. Oliver (1m#, CNC); Hazen Camp, 81º49'N, 71º16'W, 21.vi.1962, J.F. McAlpine (1m#, CNC); Hazen Camp, 81º49'N, 71º18'W, 1.viii.1963, J.R. Vockeroth Collector (1f#, CNC); Masik Riv., Banks Is., 3-31.vii.1968, 16.vii. 1968 (along rock in stream), 30.vii.1968 (sweeping), G.E. Shewell (55m#, 77f#, CNC); Sachs Harbour, 27-30.vi.1968, W.R. Mason (2m#, 2f#, CNC); Hazen L., Ellesmere Is., 7-13.vii.1963, P.11, N.10, B. Hocking (5f#, CNC); Hazen L., Ellesmere Is., 13.vii.1963, N.10, C.D. Sharplin (3f#, CNC); Hooper Is., Mackenzie Delta, 28.vi., 16.vii.1971, D.M. Wood (3m#, 5f#, CNC); Hot Weather Creek, Forsheim Peninsula, Ellesmere Is., 79º58'N, 84º28'W, 5-17.vii.1990, F. Brodo (8m#, CNC); Hot Weather Creek, Forsheim Peninsula, Ellesmere Is., 79º58'N, 84º28'W, FEN 11, 16, 18, 126, vii.1991, F. Brodo (7m#, 6f#, CNC); Head of Traquary Fjord, Ellesmere Is., 2.viii.1963, J.R. Vockeroth (2f#, CNC); Head of Traquary Fjord, Ellesmere Is., 2.viii.1963, J.R. Vockeroth (2f#, CNC); Head of Clyde Inlet, Baffin Is., 7.viii.1958, J.E.H. Martin (1m#, 5f#, CNC); Victoria Is., 71º17'N, 114ºW, 23-28.vii.1975, G. & M. Wood (2m#, 2f#, CNC). Nunavut: Cambridge Bay, 69.1207, -105.42585, 11- 19.vii.2011, mesic, Malaise, Repl. 2-3, 15.vii.2011, wet & mesic, Sweep, Repl. 1,3, NBP Field Party (49m#, LEMQ); Cambridge Bay, 26.vii.1950 G.K. Sweatman (1f#, CNC); Cambridge Bay, 29.vii.1950, E.H.N. Smith (1f#, CNC); Ellesmere Is., Lake Hazen, 81.82975, -71.32244, 19-25.vii.2010, wet & mesic, Malaise, Repl. 1, 2, 22-27.vii.2010,

wet, Sweep, Repl.1, NBP Field Party (7m#, LEMQ); Kugluktuk, 25-29.vi.2010, wet & mesic, Malaise, Repl.1, 3-26.vii.2010, wet & mesic, Malaise, Repl.1-3, NBP Field Party (26m#, LEMQ); Nettilling Lake, Baffin Is., 7,9.vii.1925, J.D. Soper (2m#, CNC); Coral Harbour, Southampton Is., 1-17.vii.1948, G. E. Shewell (56m#, 64f#, CNC); Coral Harbour, Southampton Is., 18.vii.1952, T.P.R. Ehlich (1m#, CNC); Cambridge Bay, 18- 20.vi., 22-31.vii.1950, E.H.N. Smith (7m#, 9f#, CNC); Cambridge Bay, 20-26.vii.1950, G.K. Sweatman (7m#, 14f#, CNC); Cambridge Bay, 12.viii.1959, W.R. Richards (2m#, 2f#, CNC); Cambridge Bay, 12.viii.1950, W.R.M. Mason Collector (4m#, 5f#, CNC); Axel Heiberg Is., 79º25'N, 90º45'W, Wolf R., 19-31.vii., 4, 25.viii.1963, H.K. Rutz (92m#, 52f#, CNC). Yukon. Firth River, 11-14.vii.1956, R.E. Leech (8m#, 13f#, CNC); Firth River, 11-17.vii.1956, E.F. Cashman (3m#, 2f#, CNC); Herschel Is., 5-30.vii.1953, J.S. Waterhouse (85m#, 38f#, CNC); Herschel Is., 9-30.vii.1953, C.D. Bird (40m#, 40f#, CNC); Hershel Is., 24-28.vii.1971, D.M. Wood (1m#, 2f#, CNC); Herschel Is., 28- 30.vi.1971, D.M. Wood (12m#, 5f#, CNC); Herschel Is., 18-25.vii.1930, O. Bryant (21m#, 30f#, CNC). USA. Alaska: Cape Thompson, 22-29.vii.1961, R. Madge (14m#, 22f#, CNC); Mile 213, Richard Hwy, 17.vi.1951, J.R. McGillis (1f#, CNC). Colorado. Mt. Evans, 12000', 3.viii.1961, W.R.M. Mason (1f#, CNC). GREENLAND. Nedre Midsommer Sö, 1-30.vii.1966, Can. Peary Land Expd. (68m#, 53f#, CNC); Nedre Midsommer Sö, Midsommer Sö, M2-10, 12.vii., T16-8, 11.vii., T13-9, T34-7, 18.vii., M2-15, 22.vii., M2-11, 14.vii., M2-09, 10.vii, M2-06, 4.vii., M2-13, M2-17, T12-9, 18.vii., T34-12, 5.viii., T37-9, 19.vii., T34—5, 3.vii., 8.vii.1966, Can. Peary Land Expd (27m#, 8f#, CNC); Sondrestrom Air Base, 10-14.vi., 18.vii.1952, W.J. Brown (10m#, 7f#, CNC). Diagnosis. Male. Cerci pale and folded or wrapped over each other, leaving no opening (Fig. 3.50). Hind tibia and femur with pronounced ventral pubescence (Fig. 3.21). Hind femora with anteroventral setae only present on anterior 1/3 and decreasing in length posteriorly. Hind tibia with anteroventral setae stout, dark, spine-like, shorter than ventral pubescence (Fig. 3.21). Female: Wings yellow. Halteres yellow and pollinose. Legs pollinose. Hind femora with pronounced crease on ventral side. Thorax hairy and with 2 obvious lighter grey stripes. Scutellum bristles black and sticking

upwards. Lateral side of fore femur: with a row of thin, black, evenly spaced bristles projecting straight outwards,. Description. Wing length 11.6–13.5 mm. Male. Head dark in ground-colour, with greyish pruinescence on face, frons, postgena and occiput; oral margin dark and lustrous. Occiput coloured with blue tinge. Holoptic, eye with ommatidia larger on upper half and smaller on lower half. Frons divergent towards antennae. Face divergent towards mouthparts. Ocellar triangle dark; 1 pair of anterior divergent ocellar setae, 1 pair of posterior ocellar setae, of subequal length to latter; 2-3 pairs of postocellar setulae slightly shorter than ocellar setae. Upper half of occiput bearing row of slender, dark postocular setae; occipital setae black, long and slender. Postgena with many fine, hair- like, long, dark setae. Postpedicel, stylus and scape dark. Scape slightly more than 2× longer than bulbous pedicel; pedicel width larger than scape width; postpedicel cone-like and nearly 4× longer than basal width; stylus subequal to basal width of postpedicel. Palpus dark, with many dark, long, hair-like setulae. Proboscis dark; labrum lustrous and reddish brown with yellow apex; labellum with dark setae. Thorax dark in ground-colour, largely densely grey-blue pruinescent. Pleura dark grey-blue, with dark and lustrous marking on anterior region of anepisternum; lustrous brown colour on posterior corner of postpronotal lobe; notopleural ridge pale. Anterior spiracle pale, posterior spiracle glossy black. Prescutum and scutum with pair of distinct grey-blue vittae between acr and dc rows, separated by broad brownish stripe and flanked by broad brownish tinge; spalr margins grey-blue. Proepisternum with many long, hair- like, dark setae; prosternum bare. Propleuron bearing small cluster of dark setae. Antepronotum with row of short, stout dark setae. Postpronotum with many long, dark setae covering most of postpronotal lobe. Thorax densely clothed with dark setae; scutum with dense row of dark acr shorter than postpronotal setae; dc similar to acr, increasing in length posteriorly; posterior spalr setae dark, longer than acr and dc; many anterior notopleural setae dark, hair-like, and of equal length to postpronotal setae; 4 posterior ntplr setae dark, stouter, slightly longer than anterior npl setae. Postalar callus with single stout, long, dark seta. Scutellum with 2 pairs of apical sctl setae: 1 pair long, 1 pair short; 3 pairs of subapical sctl setae of equal length to apical sctl setae. Laterotergite with crowded cluster of dark setae longer than postpronotal setae.

Legs long, entirely dark. Coxae with numerous dark, fine anterior setae. Hind trochanter with many dark setae shorter than coxal setae, concentrated mostly on ventral surface; fore trochanter nearly bare. Hind femur with pronounced ventral pubescence, with several rows of dark antero/posteroventral, ventral and anterior stout setae of varying lengths but never longer than femoral width, and only present on proximal 1/3 of femur (Fig. 3.21). Mid femur with 2-3 rows of antero/posteroventral setae, anteroventral setae stout, spine-like; posteroventral setae stout, 2x length of anteroventrals. Fore femur with ventral pubescence, width decreasing apically; antero/posteroventral fine setae never dark, subequal to femoral width. Tibiae with strong ventral pubescence, mid tibia with antero/posteroventral rows of dark, stout, spine-like setae no longer than pubescence; mid and fore tibiae with antero/posterodorsals fine, dark, never longer than 2x tibial width; hind tibia with antero/posterodorsals dark, stout, about ½ tibial width (Fig. 3.21). Basitarsi with strong ventral pubescence, setae never longer than basitarsal widths (Fig. 3.21). Wing clear with yellowish veins; all veins complete (except Sc), well sclerotized. Costal setae dark, decreasing towards wing apex. Pterostigma present, although sometimes not obvious. Halter pale with stem slightly darker. Anal lobe well-developed; axillary fissure acute. Abdomen dark grey-blue; middle of tergites and sternites often with brownish tinges; margins of segments pale grey-blue. Coloration may slightly vary from more blue to more brownish, depending on specimen. Except for sternite 1, abdomen covered in long, dark setae; setae shorter on dorsal region of tergites. Sternite 8 large and subtriangular; tergite 8 barely visible because greatly reduced and because genitalic cercal plates fold over abdomen. Setae of margin of tergite 8 usually visible, long, dark; setae of sternite 8 also long, dark. Terminalia (Figs. 3.50, 3.53) large, brown; epandrium separated into basal membranous portion with dark tracing converging into lustrous, yellow sclerotized region. Epandrium large, bearing 1 stout, rounded, finger-like projection, densely pruinose and bearing many dark, longer setae. Hypandrium short, slim, subrectangular, hugging phallus base after phallus V-shaped curvature. Phallus short, lustrous, comprised of 3 sections: swollen yellow basiphallus; shorter, less swollen darker mid-section,

distiphallus slim with 2 small teeth-like projections. Phallus without loops, never extends farther than cerci. Cercus yellow, long, fold over each other such that no opening left (Fig. 3.50). Base of cercus bearing many short, dark, fine, erect setae. Ejaculatory apodeme held low, large, fan-shaped, with lateral wings large but no larger than vertical wing; broadly connected to phallus. Female. Similar to male. Remarks: The synonymy of the two species has not yet been formally published. However, Oliver (1963) treated the species as R. nigrita (Downes, 1970). Geographical distribution. Nearctic distribution in Figure 3.73.

Rhamphomyia (Dasyrhamphomyia) zaitsevi, Beccker Fig.s 3.22, 3.25, 3.51, 3.55, 3.73

Material examined. CANADA. Nunavut: Cambridge Bay, 69.12366, -105.42531, 15- 19.vii.2011, wet, Pan, Repl.3, NBP Field Party (1m#, LEMQ). Northwest Territories: Eskimo Point, 31.vii.1950, G.G. DiLabio (1m#, CNC). Diagnosis. Male. Cerci fold over each other at apex and base, leaving a slim oval- shaped opening; cerci triangular-shaped, ending before reaching tergite 5 (Fig. 3.51). Hind tibia with dorsal setae longer than tibial width but shorter than 2x tibial width (Fig. 3.22); 2 faint greyish vittae. Description. Wing length 11.3–13 mm. Male. Head dark in ground-colour, with dense greyish pruinescence on face, frons and occiput. Holoptic, eye with ommatidia larger on upper half, smaller on lower half. Frons divergent towards antennae. Face divergent towards mouthparts; pruinose with oral margin dark and shiny. Ocellar triangle dark, subshiny, with 4 pairs of dark, parallel ocellar setae; 2 anterior pairs of subequal length and shorter than 3rd and 4th pair of ocellar setae; 3 pairs of distinct postocellar setulae shorter than ocellar setae and decreasing in length posteriorly. Occiput dark, bearing row of postocular setae, long, dark, only present on upper section of occiput; upper occipital setae black, stout, about half length of postocular setae; lower occipital setae finer, longer; postgena bearing dark setae finer than occipital setae. Postpedicel, stylus and scape dark and densely pruinose. Scape no more than 2× longer than pedicel;

pedicel and scape width similar; postpedicel nearly 4× longer than basal width; stylus only slightly shorter than lengths of scape and pedicel together. Palpus dark, with setulae dark. Proboscis largely reddish-brown and lustrous; apex of labrum yellow brown; labellum with dark setae. Thorax dark in ground-colour, largely densely grey pruinescent. Prescutum and scutum with pair of indistinct lighter grey-brownish vittae between acr and dc rows. Post- alar callus light brown. Pleura dark grey, often with darker brownish shadows on sclerites; posterior corner of postpronotal lobe lustrous yellow-orange; notopleural ridge not lustrous, pale yellow-brown; anterior and posterior spiracles dark with pale setae. Proepisternum with many dark, fine setae; prosternum bare. Propleuron bearing cluster of dark, fine setae. Antepronotum with dense row of short, stout, dark setae. Postpronotal lobe with several shorter and longer, dark, stout setae, no longer than anterior intlr and splr setae. Scutum with dense row of short, dark, fine acr setae increasing in length posteriorly; dc similar to acr; splr setae dark, increasing in length and stoutness posteriorly; setae on postalar callus dark and subequal to posterior splr setae; 1 postalar callus seta stouter and longer; anterior npl setae dark, fine, subequal to posterior npl setae; 3-4 posterior npl setae darker than anterior npl setae. Scutellum shorter than width of thorax, with 1 pair of dark, long apical sctl, 2 pairs of subapical sctl as long as apical sctl; 2 pairs of marginal sctlr setae, fine and shorter than apical sctl setae. Legs long, entirely dark, except for lustrous pale yellow-brown ring at hind femur-hind tibia junction. Mid and fore coxae with numerous dark setae located anteriorly; hind coxae with dark setae located laterally. Hind and mid trochanters with many short, dark setae, slightly longer on hind trochanter; fore trochanter nearly bare. Femora with ventral pubescence, several rows of antero/posteroventral and ventral setae on proximal 1/2, dark, of varying lengths but never longer than femoral widths, with several rows of antero/posterodorsal, dorsal and anterior setae similar to antero/posteroventrals and ventrals. Hind tibia with antero/posterodorsals stout, dark, longer than, but less than 2x tibial width; antero/posteroventrals stout, spine-like, dark, as long as ½ tibial width, decreasing in length towards apical ¼, with cluster of antero/posteroventral and ventral setae on proximal ¼ stout, dark, shorter than ½ tibial width; anterior and anteroventral apical setae dark, stout, spine-like, 2x length of

proximal anteroventral setae (Fig. 3.22). Mid (Fig. 3.25) and fore tibiae with antero/posterodorsal setae dark, fine, no longer than tibial width; mid tibia with antero/posteroventral rows of stout, dark, spine-like setae, no longer than ventral pubescence. Hind and mid basitarsi (Figs. 3.22, 3.25) with antero/posteroventral setae dark, stout, as long as basitarsal widths; fore basitarsi without stout antero/posteroventral setae, with strong ventral pubescence. Wing clear with yellowish veins; all veins complete (except Sc), well sclerotized. Costal setae dark, decreasing in length towards wing apex. Pterostigma present. Anal lobe well-developed; axillary fissure forming acute angle. Halter dark. Abdomen dark reddish-brown; abdominal segments with pale margins. Abdomen covered in dark setae of equal lengths to each other, except slightly shorter on middle of tergites. Tergite 8 modified, large, U-shaped and held upwards; dorsal surface with slight bump; sternite 8 broadened at base, triangular-shaped. Terminalia dark (Figs. 3.51, 3.55). Epandrium swollen, oval, enlarged with stout, rounded, finger-like projections covered with short, dark, stout setae; hypandrium well sclerotized, broad with cylindrical portion hugging base of phallus; phallus short, divided into 3 parts: basiphallus broadened, middle of phallus less broad than basiphallus; distiphallus thinning to hair-like width and with 2 small, teeth-like projections. Cercus pale; cercal plates elongated, subequal to length of epandrium; cerci held up over abdomen reaching tergite 6 in triangular shape; cercal plates do not fold over each other, but anterior and posterior ends closed, ovoid opening visible; cercal lobes with inner margins bearing many fine, short, erect dark setae (Fig. 3.51); ejaculatory apodeme larger than R. nigrita, fan-shaped, vertical wing open at 90º angle; broadly connected to phallus base. Female. Not examined. Geographical distribution. Nearctic distribution in Figure 3.73.

Rhamphomyia (Dasyrhamphomyia) sp.9 Figs. 3.23, 3.24, 3.52, 3.54, 3.73 Material examined. CANADA. Nunavut: Cambridge Bay, 69.11993, -105.42065, 7- 19.vii.2011, mesic, Malaise, Repl.1-3, NBP Field Party (8m#, LEMQ); Kugluktuk,

67.83538, -115.20987, wet & mesic, Malaise, Repl. 1-3, NBP Field Party (103m#, LEMQ). Diagnosis. Male. Cerci folded over each other at apex and base, leaving a wide, pointed, oval-shaped opening (Fig. 3.52); cercus cylindrical-shaped, reaching tergite 5. Cercal tip dark. Three distinctive acr and dc, brown thoracic vittae. Hind tibia with anteroventral row of dark, stout, spine-like setae shorter than half tibial width (Fig. 3.23). Description. Wing length 13 mm. Male. Head light grey-blue in ground-colour, with dense greyish pruinescence on face, frons and occiput. Clypeus yellow-brown. Holoptic, eye with ommatidia larger on upper half, smaller on lower half. Frons divergent towards antennae. Face slightly divergent towards mouthparts; bare with oral margin dark and shiny. Ocellar triangle dark grey-blue, with 1 pair of parallel ocellar setae, 1 pair of shorter, finer posterior setae and 2-3 pairs of short, fine postocellar setae. Occiput bearing row of stout, short, dark postocular setae on upper occipital half; occipital setae of upper half similar to postocular setae; setae of lower occiput longer; post-gena bearing finer, longer seta than occipitals. Postpedicel, stylus and scape mostly dark brown; pedicel slightly darker than scape, postpedicel and stylus. Scape length less than 2× longer than pedicel; postpedicel cone-shaped, about 4× longer than basal width; stylus length equal to length of scape. Palpus short, subequal to length of scape and pedicel together; palpus pollinose and light brown, with dark, fine setulae. Labrum reddish-brown and lustrous; apex of labrum yellow; proboscis largely pale grey; labellum with dark setae. Thorax light grey-blue in ground-colour, largely densely grey pruinescent. Prescutum and scutum with pair of distinct shiny, dark grey-blue vittae between acr and dc rows, separated by broad brown stripe and flanked by broad brown tinge. Postalar callus pale. Pleura light grey-blue, with darker shadows on anepisternum, anepimeron and katepisternum; base of wing glossy and dark brown; posterior corner of postpronotal lobe lustrous and brown-yellow; npl ridge pale. Proepisternum with 6 short, dark setae; prosternum bare. Antepronotum with row of short, stout dark setae. Postpronotal lobe with several anterior short, dark setae; 2 posterior longer, dark setae. Propleuron bare. Scutum with biserial, sparse row of stout, dark acr; dc similar to acr, except dc increasing in length and stoutness posteriorly; several short, dark splr setae, although anterior splr setae stouter and longer; postalar callus bearing 1 long, dark, stout seta; anterior npl setae

dark, mostly shorter with 1 longer seta; 3 posterior npl setae long and dark. Scutellum with 1 pair of longer apical sctl, 1 shorter pair of subapical sctl. Laterotergite with numerous fine, long, dark setae. Anterior and posterior spiracles pale. Legs long, largely light yellowish-brown; with femoral-tibial joint paler. Coxae more grey-blue than yellowish-brown; ventral margins of trochanters black and lustrous. Hind and mid coxae with a few short, dark lateral setae; fore coxa with several more dark anterior setae, longer than lateral setae of mid and hind coxae. Trochanter with a few dark, fine setae. Femora with ventral pubescence, with antero/posteroventral setae dark, stout, no longer than 1/2 femoral width; hind femur with several anterior and posterior rows of dark, stout, setae shorter than antero/posteroventrals; mid (Fig. 3.24) and fore femora with 1 anterior and 1 posterior row of dark, stout setae, shorter than antero/posteroventrals. Hind tibia (Fig. 3.23) with anteroventral setae dark, stout, spine- like, shorter than ¼ of tibial width, absent on proximal ¼ of tibia; mid tibia (Fig. 3.24) with antero/posteroventral setae stout, dark, spine-like, 1.5x length of anteroventral setae of hind tibiae; fore tibiae clothed with short, fine, dark setae, except for 1 preapical antero/posterodorsal seta, 3 anterior preapical setae, 2 stouter anteroventral setae, as long as, or subequal to, tibial width. Hind and fore basitarsi subequal to hind and mid tarsomeres 2-4, respectively; mid basitarsi shorter than mid tarsomeres 2-4; basitarsi clothed with short, dark, spine-like setae (Fig. 3.24), except for antero/posteroventral and ventrals fore-basitarsi fine. Wing clear with yellowish veins; all veins complete (except Sc), well sclerotized. Costal setae dark, decreasing in length towards wing apex. Anal lobe well-developed; axillary fissure forming acute angle, although almost seemingly rectangular. Pterostigma present, large and dark yellow. Halter white. Abdomen light brown with distinct white margins of each tergite and sternite. Abdomen covered in dark, fine, short setae, longer on laterals of tergites and in middle of sternites. Sternite 8 triangular, bearing dark, fine setae only at margin; tergite 8 greatly reduced, marginal setae noticeable; tergite 8 modified with dome-shaped flap projecting upwards, like being pushed over by cerci; basal width of tergite 8 wider than apical width of tergite 8, triangular shaped. Terminalia (Figs. 3.52, 3.54) largely dark brown except cerci mostly pale white

and apex brown. Inner margins of cercus densely pubescent; basal cercus bearing several fine, darker, short, erect setae. Epandrium swollen, oval, with stout, rounded, finger-like projections longer than those of R. nigrita and R. “zaitsevi”; with pronounced crease; bearing many short, stout, dark, spine-like setae as well as many finer, longer setae. Upper margin of epandrium darkened and bearing many fine, dark setae. Ventral surface of epandrium also bearing many fine, dark, long setae. Hypandrium short, cylindrical, partly membranous, partly well-sclerotized, lustrous and yellow-brown, hugging basiphallus. Phallus lustrous yellow-brown; basiphallus broadened; dark crease running down middle; phallus short, mostly hidden between epandrial lamellae, never extending further than length of genitalia; distiphallus slimmer than basiphallus and bearing 2 short, teeth-like projections. Cercus distinctly modified (Fig. 3.52): cerci elongated, held up over abdomen, reaching tergite 5; cercal plates folding such that dorsal margins of apex and ventral margin at base touching each other, forming oval-shaped opening. Ejaculatory apodeme small, with lateral wings of equal length to vertical wing, broadly fused to phallus base. Female. Unknown. Geographical distribution. Nearctic distribution in Figure 3.73.

Rhamphomyia (Eorhamphomyia) nigrolineata gp. sp.4 Figs. 3.26, 3.56, 3.57, 3.74

Material examined. CANADA. Alberta: Ptarmigan L.L. Louise, 7600', 26.vii., K.C. Hermann (3f#, CNC) Ontario: Stittsville, unt, from cage under W. spruce, 13.v.1957, H.A. Tripp (1m#, CNC). Northwest Territories: Masik River, Banks Is., 6.vii.1968, W.R.M. Mason (1m#, CNC); -21 m.e. Tuktoyaktuk, 20-25.vi., 2-12.vii.1971, D.M. Wood (21m#, 3f#, CNC); -30 m.e. Tuktoyaktuk, 18.vi.1971, D.M. Wood (3m#, CNC); Head of Clyde Inlet, Baffin Is., 7.viii.1958, G.E. Shewell (1m#, CNC); Aklavik, 28.vi.1956, R.E. Leech (1m#, CNC). Nunavut: Shewell Harbour, Southampton Is., 9.vii.1948, G.E. Shewell (11m#, 14f#, CNC); Shewell Harbour, Southampton Is., Dryas flowers, 10, 26.vii.1948, G.E. Shewell (4f#, CNC). Yukon: Herschel Is., 5-20.vii.1953, J.S. Waterhouse, (21m#, 30f#, CNC); Herschel Is., 9, 22.vii.1953, C.D. Bird (35m#, 12f#,

CNC); Herschel Is., 28,29.vi.1971, D.M. Wood (4m#, 3f#, CNC); Aklavik, 20.vi.1953, C.D. Bird (1m#, CNC). Quebec: Payne Bay, 21.vii.1958, E.E. MacDougall (1f#, CNC). USA. Alaska: Mi. 32 Denali Hwy., 4500', 22.vii.1962, R.E. Leech (2m#, CNC); Cape Thompson, 23.vii.1961, R. Madge (1m#, 5f#, CNC); Cape Thompson, 23.vii.1961, Heming (1f#, CNC); Cape Thompson, CH-149, UA Botany Plot 9, 1076, 1079, 16.vii.1960, W.C. Hanson (1m#, 1f#, CNC); Cape Thompson, CH-161, UA Botany Plot 9, 1098, J.J. Davis (1m#, CNC); Cape Thompson, CH-162, UA Botany Plot 9, 110, 1107, 1105, 1106, 1103, 20.vi.1960, J.J. Davis (2m#, 2f#, CNC); Cape Thompson, CH-163, UA Botany Plot 9, 1070, 1068, 1069, J.J. Davis (1m#, 2f#, CNC); Cape Thompson, CH- 1003, UA Botany Plot 9, 1084, 31.viii.1960, J.J. Davis (1m#, CNC). Diagnosis. Male. Inner faces of cercal lobes with stout, dark, spine-like setae (Fig. 3.57). Cell dm shorter than cell bm. Upper occipital setae short, stout, dark; lower occipital setae longer, finer, dark; postgenal setae fine, hair-like, long, dark. Phallus not visible, hidden within epandrium (Fig. 3.56); epandrium more glossy than pruinose; posterior lower corner with sharp angle, not rounded. Sternites entirely pollinose; tergites partly lustrous. Female. Wings yellow. Abdomen pollinose ventrally and glossy dorsally. Thorax pollinose. Legs not pennate. Labrum black and glossy. Halteres light yellow. Mid and hind tibiae (Fig. 3.26) with distinct pattern of thickened, pointy, black bristles on dorsal side. Thorax with 2 darker grey stripes. Above wing, anteriorly, row of 4 black, thickened bristles. Hairs on scutellum converging. Very long, thick, black and pointy bristle on postalar callus (right above wing). Wing with pterostigma present. Description. Wing length 9.1-10.1 mm. Male. Head dark in ground-colour, with dense greyish pruinescence on face, frons, postgena and occiput; oral margin dark and lustrous. Holoptic, eye with ommatidia on upper half larger than ommatidia on lower half. Frons divergent towards antennae. Face slightly divergent towards mouthparts. Ocellar triangle dark, subshiny; anterior and posterior pair of ocellar setae dark, subequal and parallel; 3 pairs of postocellar setae dark, parallel and shorter than ocellar setae; 3rd pair of postocellar setae ½ length of 1st pair of postocellar setae. Upper half of occiput bearing row of sout, black postocular setae; occipital setae shorter and more slender on lower section; upper occipital setae black, short and stout; lower occipital setae more hair-like, dark and longer. Postgena with many hair-like, dark, long setae. Postpedicel,

stylus and scape dark and pruinose; scape slightly longer than 2x length of pedicel; pedicel bulbous; postpedicel about 4x as long as basal postpedicel width; stylus subequal to basal length of postpedicel. Palpus dark, with setulae dark. Labrum dark and glossy; labellum dark and bearing many dark setae, shorter than palpal setae. Thorax dark in ground-colour, largely grey pruinescent. Scutum with pair of usually distinct blackish, most often more lustrous than pruinose, vittae between acr and dc rows, separated by broad brownish, pruinose stripe. Pleura blackish-brown in ground colour, densely greyish pruinose, with brownish tint at margin of katepisternum, in middle of anepimeron; lustrous brownish tint at corner of postpronotum and along npl ridge. Postpronotal lobe with tint of pruinose brown. Proepisternum with many long, dark setae; prosternum bare. Antepronotum with row of short, stout dark setae; longer at edges, shorter in middle. Postpronotal lobe with cluster of dark, long setae; 1-2 posterior setae longer and stouter. Scutum with biserial row of dense, dark acr setulae, shorter than dc; dc dense, dark, increasing in length posteriorly; splr setae longer than dc, dark, increasing in length posteriorly; 1-2 dark, stout setae on postalar callus slightly longer than posterior splr setae; npl with anterior cluster of fine, short, dark setae; posterior row of 4 dark, long setae; 1 posterior npl seta as short as anterior npl setae. Scutellum with pair of dark, long, converging apical sctl; subequal to pair of dark subapical sctl setae; 2 pairs of sctl setae dark, shorter than apical and subapical sctl; 1 pair of basal sctl setae dark, subequal to sctl. Anterior and posterior spiracles pale. Legs long, dark, lustrous and entirely reddish-brown; coxae densely pruinose. Hind coxa with several dark, stout, lateral setae; mid and fore coxae bearing numerous dark, finer setae on anterior surfaces. Trochanters with ventral cluster of short, dark setae. Hind femur with ventral pubescence (Fig. 3.26), anteroventral setae fine, never longer than femoral width; mid and fore femora with antero/posteroventral setae dark, stout, increasing in length towards apical ¼. Hind tibia with many rows of antero/posteroventral and ventral setae shorter than femoral width, dark, increasing in stoutness towards proximal 2/3, with posterior apical seta stouter and 2x length of ventrals; hind (Fig. 3.26), mid, and fore tibiae with antero/posterodorsal setae stout, dark, longer than femoral width; mid tibia with antero/posteroventrals stout, dark, but less than 2x tibial width, with short, dark, stout, spine-like antero/posteroventrals setae on proximal ¼; fore tibia with

several rows of fine posterior and anterior setae, shorter than tibial width. Hind basitarsi (Fig. 3.26) with posterodorsal seta stout, dark, longer than 2x basitarsal width, 3-4 postero/anteroventral and ventral setae stout, subequal to basitarsal width, mid and fore basitarsi with fine antero/posterodorsal/ventral, dorsal and ventral setae, never longer than basitarsal width, except preapical ring of setae stout, longer than basitarsal width. Wing clear with yellowish veins; all veins complete (except Sc), well sclerotized. Pterostigma present. Anal vein well-developed. Basal costa with 2 stout, dark, long setae. Axillary fissure forming acute angle. Halter yellow. Abdomen dark, brownish-red in ground colour, tergites mostly lustrous, some grey pollinosity, margins pale grey. Sternites entirely and densely pollinose. Middle of tergites bearing many short, dark setae; laterals of tergites bearing many long, dark setae; sternites bearing many long, dark setae. Anterior region of abdomen bearing longer setae than posterior region of abdomen. Sternite 8 larger than tergite 8; ventral surface of sternite 8 bearing many dark, fine, long setae; margin of tergite 8 bearing several short, fine, dark setae. Terminalia dark (Figs. 3.56, 3.57), mostly pollinose, except for lustrous region on epandrium and lustrous hypandrium; epandrium bearing many dark setae, no longer than marginal setae of sternite 8; epandrial setae slightly curved towards opposing epandrium setae; epandrium with ridge on dorsal margin; epandrium subrectangular. Cercus slimmer than half epandrium width; dorsal margin distinctive with three humps, subequal to epandrium length; dorsal margin bearing many short, fine, dark setae. Dorsal view of cercus showing distinctive curvatures. Hypandrium large, heavily sclerotized, visible beneath epandrium, L-shaped, hugging basiphallus, ending in pointed elongation; basiphallus slimmer than middle phallus, which appears bulbous; phallus mostly hidden by epandrium (distiphallus may sometimes extend out from within cercus); within epandrium, phallus without loops, projecting upwards at obtuse angle; side view of phallus tip tear-drop shaped; dorsal view of phallus tip with 4 projections, 2 posterior projections smaller than 2 anterior projections. Ejaculatory apodeme large, but smaller than epandrium, fan-shaped, anterior margin of vertical wing rounded, without lateral wings; phallus broadly fused to ejaculatory apodeme. Female. Similar to male except wings yellow.

Geographical distribution. Nearctic distribution in Figure 3.74.

Rhamphomyia (Pararhamphomyia) lapponica gp. sp. 8 Figs. 3.27, 3.58, 3.75

Material examined. CANADA. Yukon: Herschel Is., 9-26.vii.1953, J.S. Waterhouse (14m#, 5f#, CNC); Herschel Is., 10-23.vii.1953, C.D. Bird (21m#, 33f#, CNC). Diagnosis. Male. Hind tibia with some antero/posterodorsal and dorsal setae longer than 2x tibial width on apical 2/3, shorter or subequal to tibial width on proximal 1/3 (Fig. 3.27). Fore tibia with anterior pubescence; fine posterior setae longer than anterior pubescence. Epandrium with small, slim, finger-like projection and phallus without curvature (Fig. 3.58). Female. Similar to male except setae of dorsal hind tarsi as long as setae of ventral hind tarsi. Description. Wing length 8.8-9 mm. Male. Head dark in ground-colour, with greyish pruinescence on face, frons and occiput. Holoptic, eye with ommatidia larger on upper half and smaller on lower half. Frons divergent towards antennae. Face almost parallel, although slightly divergent towards mouthparts; bare, with oral margin dark and shiny. Ocellar triangle dark, with 1 anterior pair of dark, parallel ocellar setae and 1 pair of posterior setae; anterior pair longer than posterior pair and posterior setae more widely separated from each other than anterior setae; 2 pairs of dark postocellar setae, no longer than posterior ocellar setae. Occiput bearing row of postocular setae only on upper half of occiput; setae dark and stout. Occipital setae black, stout, shorter than postocular setae. Postgena bearing dark setae longer and more hair-like than occipital setae. Postpedicel, stylus and scape dark and heavily pollinose. Scape about 2× longer than pedicel; pedicel bulbous and wider than scape; postpedicel nearly 5× longer than basal width; stylus no longer than scape. Palpus dark and pollinose, with setulae dark. Labrum lustrous and black; labellum dark and bearing several dark setae. Thorax dark, largely densely grey pruinescent. Prescutum and scutum with pair of distinct darker grey vittae between acr and dc rows, flanked with brownish tinge along outer margin. Pleura grey, often with darker brown shadows on sclerites and along edges of sclerites; rarely completely greyish. Posterior corner of postpronotal lobe lustrous and

yellow-brown; region between posterior spiracle and halter stem not pollinose and dark brown. Postalar callus light brown. Propleuron bare. Proepisternum with 3 stout, dark setae; prosternum bare. Antepronotum with row of short, stout dark setae. Postpronotum with several short, stout, dark setae and 1 long, dark, stout posterior seta. Scutum with sparse, biserial row of stout, dark acr; dc similar to acr, increasing in length posteriorly; 1 anterior intlr, 1 anterior and 1 posterior splr seta longer and stouter than other thoracic setae; small anterior cluster of short, stout, dark npl setae; 4-6 posterior npl setae longer, stouter, and darker than anterior npl setae; postalar callus bearing 2 long, stout and dark setae; posteriormost dc setae long, stout and dark. Scutellum with pair of long, stout, dark, converging apical sctl; 1 pair of stout, dark, subapical sctl setae. Laterotergite bearing dense cluster of stout, long, pale setae. Anterior and posterior spiracles golden yellow-brown with black ring on outer margins. Legs long, entirely dark reddish-brown; relatively shiny, although coxae sometimes more grey and pollinose than brown and shiny. Fore coxae with row of stout, dark, short setae on anterior region; mid and hind coxae with several dark lateral setae; mid coxal setae stouter than hind coxal setae. Hind and mid trochanters with ventral dark setae; fore trochanter bare. Femora with short, white ventral pubescence; hind femur with anteroventral setae dark, fine, doubling in length towards proximal 1/3, never longer than femoral width (Fig. 3.27); mid and fore femora with antero/posteroventral setae dark, fine, decreasing in length posteriorly, with proximal setae subequal to femoral width. Hind tibia with antero/posterodorsal setae dark, as long as 2x tibial width; mid and hind tibiae with antero/posteroventral setae no longer than tibial width, decreasing in length and increasing in stoutness towards apical 1/3 (Fig. 3.27); fore tibia with many rows of fine antero/posteroventral, ventral, anterior and posterior setae no longer than tibial width; mid tibia with preapical anterodorsal seta dark, stouter than antero/posterodorsals, subequal to tibial width; preapical anterodorsal, anterior, anteroventral and ventral dark, stout setae no longer than tibial width. Hind basitarsus (Fig. 3.27) with antero/posterodorsal setae dark, stout, longer than basitarsal width but shorter than 2x basitarsal width, middle anterior seta dark, stout, no longer than basitarsal width; mid and hind basitarsi with antero/posteroventral setae stout, shorter than basitarsal width; mid and fore basitarsi with antero/posterodorsals finer than, and subequal to,

antero/posteroventrals, except for dark, stout preapical antero/posterodorsal setae, slightly longer than basitarsal width; fore basitarsus with many fine antero/posteroventral and ventral setae, with dark, stout antero/posteroventral setae, no longer than basitarsal width and posterodorsal setae fine, dark, longer than basitarsal width, dorsal and anterodorsal setae dark, stout, of similar length to posterodorsals. Fore, mid and hind tarsomeres 2-3 similar to fore, mid and hind basitarsi, respectively; fore tarsomeres with strong ventral white pubescence. Wing clear with yellowish veins; all veins complete (except Sc), well sclerotized, except anal vein faint and not reaching wing margin. Anal lobe well-developed and axillary fissure forming rectangular angle. Pterostigma darker yellow, although faint. Halter dark yellow, with knob blackish. Abdomen dark grey, with reddish-brownish tinges on tergites, margins of segments pale. Abdomen clothed with long, hair-like, pale setae; setae longer on laterals of tergites than in middle of tergites; setae of tergites 1-5 longer than setae of tergites 6-8; middle of sternites with pair of setae divergent from each other, longer and stouter; margin of sternite 8 with many stout, long, pale setae; margin of tergite 8 bearing many shorter, finer, pale setae than marginal setae of sternite 8. Tergite 8 length half length of sternite 8. Terminalia (Fig. 3.58) dark; epandrium subtriangular, densely pruinescent, ventral margin with strong pubescence; epandrium bearing many long, pale, fine setae on ventral margin, and many darker, stouter, shorter setae at apical dorsal margin; epandrium with lighter brown tinge on middle of lamella where depression occurs; epandrium ending in tip slightly pointed, appearing finger-like upon dissection. Cercus oval-shaped, shorter than half epandrial length; dorsal margin of cercus lined with fine, short setae. Subepandrial lobe of subequal length to cercus, tear drop-shaped, apex bearing several fine, long setae; hypandrium U-shaped, ending at base of basiphallus; basiphallus swollen, lustrous and yellow-brown; ventral surface of basiphallus bearing many short, fine, setae; distiphallus darker brown, slender;loops lacking, only slight hump; phallus not usually extending farther than length of genitalia; ejaculatory apodeme fan-shaped, almost with 90º angle, small, with lateral “wings” shorter than vertical wing. Female. Similar to male except setae of dorsal hind tarsi as long as setae of

ventral hind tarsi. Geographical distribution. Nearctic distribution in Figure 3.75.

Rhamphomyia (Pararhamphomyia) lapponica gp. sp. 9 Figs. 3.28, 3.59, 3.75

Material examined. CANADA. Yukon: Herschel Is., 11-29.vii.1953, J.S. Waterhouse (25m#, 21f#, CNC); Herschel Is., 20-29.vii.1953, C.D. Bird (13m#, 3f#). USA. Alaska: Cape Thompson, 25,26.vii.1961, R. Madge (1m#, 1f#, CNC); Cape Thompson, 25, 29.vii.1961, B.S. Heming (4m#, CNC); Umiat, 10,15.vii.1959, J.E.H. Martin (2m#, 2f#, CNC). Diagnosis. Male. Hind tibia clubbed-shaped, such that width increases posteriorly; antero/posterodorsal and dorsal setae increasing in length apically, setae always never shorter than 2x tibial width; several rows of posterior setae shorter than dorsals, subequal to tibial width; antero/posteroventral and ventral setae shorter than dorsals, increasing in length towards middle, decreasing in length towards apex, apical cluster of antero/posteroventral and ventral setae short and stouter. Hind tarsomere 2 shorter than half hind basitarsus, with antero/posterodorsal setae 2x width of tarsomere; hind basitarsus with antero/posterodorsal and dorsal setae longer than 2x the basitarsal width (Fig. 3.28); apex of epandrium gradually tapered and lengthened (Fig. 3.59). Female. Similar to male except hind tibiae not swollen. Description. Wing length 8.1-8.6 mm. Male. Head dark in ground-colour, with greyish pruinescence on face, frons, postgena and occiput. Holoptic, eye with ommatidia larger on upper half, smaller on lower half. Frons sides divergent, not parallel-sided; face mostly parallel-sided and only slightly divergent near mouthparts; bare, with oral margin shiny. Ocellar triangle dark, subshiny, with 1 anterior pair of parallel ocellar setae and 1 pair of posterior setulae ½ length of anterior ocellar setae; 2 pairs of postocular setae subequal to length of posterior ocellar setae. Occiput bearing row of postocular setae, only present on upper half of occiput; setae hair-like, dark, longer than occipital setae; remaining occipital setae shorter, stouter and black; postgena bearing hair-like setae as long as postocular setae. Antenna dark, with dense grey pruinescence, pedicel sometimes with inner paler coloration. Scape 2x length of pedicel; pedicel bulbous; margins of

pedicel and scape bearing dark, short setae; postpedicel about 4× longer than basal width; stylus equal to scape length. Palpus dark reddish-brown, with dark setulae. Labrum lustrous and reddish brown; labellum dark brown, bearing many dark setulae. Thorax dark in ground-colour, largely densely grey pruinescent: posterior corner of postpronotal lobe lustrous and yellow-brown; notopleural ridge dark yellow-brown; postpronotum and sclerites on pleura sometimes with yellowish-brown tinge; postalar callus brown. Prescutum and scutum with pair of distinct dark vittae between acr and dc rows; flanked with brown tinge. Proepisternum grey, with several short, stout, dark setae; prosternum bare. Propleuron bare or with 1 short, dark seta. Antepronotum with row of short, stout, dark setae. Postpronotum with several short, stout, dark setae and 1 stout, dark setae longer than other postpronotal setae. Scutum with biserial row of dark, fine acr; dc longer than acr, increasing in length and stoutness posteriorly; 2 anterior npl setae shorter than posterior npl setae; 6 posterior npl longer than anterior npl setae, 2 posterior npl slightly shorter than remaining 4 posterior npl setae. Sparse row of intlr setae, stout, subequal to dc, with 1 anterior intlr seta longer and stouter; sparse row of spalr setae, stout, shorter than dc; 1 stout, long pal and several short, hair-like, pale setae; 1 dark, stout, long postalar callus. Scutellum with 1 pair of dark, stout, converging apical sctl setae; 1 pair of dark, stout, subapical sctl setae slightly shorter than apical sctl setae; 1 pair of sctl setae ½ length of subapical sctl setae. Laterotergite with many long, stout, pale setae. Anterior and posterior spiracles pale with outer black ring. Legs long, entirely dark reddish-brown, except trochanters more black than brown. Hind and mid coxae bearing several lateral dark, stout setae; fore coxae bearing several stout, dark anterior setae. Hind trochanter bearing several short, pale and hair-like setae ventrally; mid trochanter bearing shorter, pale, fine setae ventrally; fore trochanter nearly bare. Mid and hind femora (Fig. 3.28) with ventral pubescence; hind femur with anteroventral setae fine, dark, shorter than femoral width, doubling in length towards apical 1/3 (Fig. 3.28); mid femur with posteroventrals similar to anteroventrals of hind femur; fore femur with antero/posteroventrals sparse, fine, always shorter than femoral width. Hind tibia swollen apically such that tibial width nearly doubles; antero/posterodorsal/ventral and dorsal/ventral setae fine, dark, longer than tibial width and antero/posterodorsal and dorsal setae longer than antero/posteroventral and ventral

setae, with several rows of shorter, fine, dark anterior setae (Fig. 3.28); mid tibia with anterior setae stout, dark, shorter than tibial width, antero/posterodorsal setae dark, finer, subequal to tibial width; fore tibia with fine setae, except antero/posterodorsal preapical setae dark, stout, subequal to tibial width. Hind basitarsus (Fig. 3.28) and hind tarsomeres with antero/posterodorsal and dorsal setae dark, fine, at least 3x length of their respective tarsomere width; antero/posteroventral and ventral setae stout, dark, shorter than tarsomere width; hind basitarsus slightly swollen, slightly larger than width of tarsomere 2, but smaller than hind tibial width; mid and hind basitarsi and tarsomeres slender, fine, antero/posterodorsals no longer than 2x basitarsal width. Wing clear with yellowish brown veins; all veins complete (except Sc), well sclerotized. Anal lobe well-developed; axillary fissure forming rectangular angle. Pterostigma faint, but present and darker yellow coloration. Halter yellow. Abdomen dark with brownish tinges sometimes visible on sternites. Abdomen covered with many hair-like, pale setae, shorter on middle of tergites. Setae on tergite 6-8 shorter. Tergites 5 to 8 sometimes with visible dark brown lateral markings. Sternite 8 triangular with basal width larger than upper width; margin of sternite 8 lined with pale setae projecting outwards, not curved. Tergite 8 very small and reduced, ½ length of sternite 8. Terminalia (Fig. 3.59) dark. Epandrium subtriangular, elongated with apical tip shaped as long finger-like projection bearing many dark, stout, short setae; base of epandrium dark brownish-black; middle of epandrium with yellowish patch where depression occurs; epandrium pruinose with ventral margin bearing many fine, pale setae. Cercus with four rounded corners, shorter than ½ epandrium length; densely pruinose; dorsal margin bearing many fine, short, dark setae. Subepandrial lobe tear drop-shaped, elbowed, with stem slimmer than apex; apex bearing several fine, dark setae, longer than setae on cercus; subepandrial lobe longer than cercus but shorter than 2x cercal length; hypandrium cylindrical, short, ending at base of swollen basiphallus; phallus slender and reddish brown; curved inwards once, forming loop, extending further than length of terminalia, then loops back between epandrial lamellae. Ejaculatory apodeme fan-shaped, small, dorsal margin of vertical wing flattened, 2 short lateral wings at base shorter than vertical wing.

Female. Similar to male except hind tibia not swollen. Geographical distribution. Nearctic distribution in Figure 3.75.

Rhamphomyia (Pararhamphomyia) lapponica gp. sp. 11 Figs. 3.29, 3.60, 3.75

Material examined. CANADA. Nunavut: Coral Harbour, Southampton Is., 13, 20.vii.1948, G.E. Shewell (13m#, 32f#, CNC); Chesterfield, 1,8.viii.1950, J.R. Vockeroth (1m#, 3f#, CNC). Diagnosis. Male. Hind tibia with antero/posterodorsal and dorsal setae increasing in length posteriorly, such that posterior 2/3 as long as 2x tibial width, setae of proximal 1/3 of hind tibia longer than tibial width, but shorter than 2x tibial width; antero/posteroventral and ventral setae never longer than 2x tibial width; hind basitarsus with antero/posterodorsal and dorsal setae finer and longer than antero/posteroventral and ventral setae, and as long as 2x basitarsal width; hind tarsomere 2 with antero/posterodorsal and dorsal setae shorter than tarsomere width, except for preapical antero/posterodorsal setae stouter and longer (Fig. 3.29). Fore coxa with 2 rows of stout, dark anterior setae. Setae of laterotergite pale. Apex of epandrium sharply attenuated with slender extension (Fig. 3.60). Female. Similar to male except as follows: dorsal setae on hind basitarsus not longer than ventral setae; femora glossy and tibiae pruinose. Description. Wing length 9.1 mm. Male. Head dark in ground-colour, with greyish pruinescence on face, frons, occiput and postgena. Holoptic, eye with ommatidia of larger size on upper half and of smaller size on lower half. Frons triangular; face parallel-sided towards mouthparts; bare, with oral margin brown and shiny. Ocellar triangle dark, with 1 pair of anterior parallel ocellar setae and 1 pair of posterior ocellar setae 1/2 length of anterior ocellar setae; ocellar setae dark; anterior ocellar setae more stout than posterior ocellar setae. Occiput bearing row of stout, black postocular setae only present on upper 1/2; occipital setae black, short and stout on upper 1/2, finer on lower 1/2; postgena bearing setae, finer than occipital setae. Antenna dark, with some inner paler brown coloration. Scape about 2x longer than pedicel; pedicel bulbous; postpedicel about 5× longer than basal width; stylus subequal to length of scape. Palpus dark, with setulae dark. Labrum dark reddish-brown; labellum with dark setae.

Thorax dark in ground-colour, largely grey pruinescent: posterior corner of postpronotal lobe lustrous and yellow-brown, splr ridge usually with discrete reddish- brown shadow, npl ridge pale, region between posterior spiracle and halter base dark brown. Prescutum and scutum with pair of varyingly distinct, darker grey, and pollinose vittae between acr and dc rows, flanked by faint brownish tinges. Pleura with brownish shadows on sclerites, region beneath wing base darker brown. Proepisternum with several stout, dark setae; prosternum bare. Propleuron bare. Antepronotum with row of short, stout, dark setae. Postpronotum with several fine, pale setae and with several darker, stouter, longer setae. Scutum with sparse biserial row of fine acr; dc longer, darker and stouter than acr, increasing in length posteriorly; sparse row of dark, stout splr setae, no longer than dc except for postsutural splr seta longer and stouter; 3 presutural intlr seta as stout, dark and long as dc; notopleuron with several dark, stout, subequal setae; postalar callus with 1 stout, dark, long seta. Scutellum with 2 pairs of sctl; 1 apical pair slightly longer and stouter than subapical sctl; both scutellar pairs converging towards each other, not overlapping. Laterotergite with many long, pale setae. Anterior and posterior spiracles dark orange-brown with outer black ring. Legs long, entirely reddish-brown, except coxae dark with grey pruinescence and brownish shadows; hind tibial-femoral junction pale. Hind coxa with numerous dark, stout, short, lateral setae; anterior region of hind coxa with dome-shaped region distinctively orange-brown and lustrous; anterior mid and fore coxae with many dark, stout, short setae. Trochanter with sparse, fine, pale, short ventral setae; ventral setae of fore trochanter shorter and finer. Hind femur with anteroventral setae dark, fine and shorter than femoral width on proximal half, stouter and at least 2x in length on apical half (Fig. 3.29); mid femur with anteroventral setae dark, stout, shorter than femoral width, posteroventral setae dark, stout, also shorter than femoral width, except for at least 2 stouter, longer anteroventrals on apical 1/3, but never longer than femoral width; fore femur with all setae fine and shorter than femoral width, except 2 preapical anterodorsal setae stouter and slightly longer. Hind tibia with apical width 2x proximal width, setae dark and fine, antero/posterodorsal and dorsal setae increasing in length towards apical ½, such that apical setae more than 2x tibial width (Fig. 3.29); antero/posteroventral setae subequal or longer than tibial width, shorter than 2x tibial width, expect setae of apical ¼.

Mid and fore tibiae slim, setae never longer than 2x tibial width, antero/posterodorsals fine; mid femur with antero/posteroventrals stout, dark, spine-like; fore femur with antero/posteroventrals fine. Hind basitarsi slightly swollen, width larger than hind tarsomere width, smaller than hind tibial width (Fig. 3.29); antero/posterodorsal and dorsal setae dark, fine and longer than 2x hind basitarsal width; antero/posteroventral and ventral setae stouter, dark and subequal to basitarsal width; hind tarsomeres similar to hind basitarsi except shorter and slimmer. Mid and fore basitarsi not swollen; antero/posterodorsal and dorsal setae fine, dark, no longer than 2x mid and fore basitarsal width, with antero/posteroventral setae dark, stout, subequal to 2x basitarsal width; preapical ring of dark, stout setae, never longer than 2x basitarsal width. Wing clear with yellowish brown veins; all veins complete (except Sc), well sclerotized. Anal lobe well-developed; axillary fissure rectangular. Pterostigma darker yellow, although very faint. Halter mostly yellow with faintly darkened knob. Abdomen dark grey with distinct brown coloring on tergites and sternites; margins of segments pale. Abdomen clothed with long, fine, pale setae, shorter in middle of tergites, longer on laterals and in middle of sternites. Setae of sternite 6-8 shorter and finer. Sternite 8 subtriangular, larger than tergite 8; margin of sternite 8 with many fine, pale setae. Terminalia (Fig. 3.60) mostly dark; epandrium elongated with basal width larger than width of apex; middle of epandrium marked by depression highlighted by pale colouring of epandrium; epandrium ending in slim finger-like projection, shorter than cercus; epandrium with ventral margin bearing many fine, pale, long setae; finger-like projection bearing several fine, long, pale setae; upper margin of epandrium bearing stouter, pale setae; dorsal margin of epandrium apex bearing dense cluster of stout, spine- like, dark, short setae; cercus with four rounded corners, shorter than subepandrial lobe, apical width smaller than basal width; dorsal margin of cercus bearing many fine, dark, erect setae, subequal to, or slightly longer than, cercus width; subepandrial lobe club- shaped, apex bearing dark, fine, erect setae longer than cercus setae. Hypandrium short, cylindrical, ending at swollen basiphallus; basiphallus turning at 90º angle into distiphallus; distiphallus slim, curved inwards once, forming large loop extending farther than length of terminalia, looping into epandrial lamellae; distiphallus tip cone-shaped.

Ejaculatory apodeme fan-shaped, forming obtuse angle, lateral wings shorter than vertical wing. Female. Similar to male except as follows: setae on dorsal hind basitarsus no longer than setae on ventral hind basitarsus; femora glossy and tibiae pruinose. Geographical distribution. Nearctic distribution in Figure 3.75.

Rhamphomyia (Pararhamphomyia) hoeli, Frey 1949 Figs. 3.30, 3.61, 3.75

Material examined. CANADA. Northwest Territories: Victoria Is., 71º17'N, 114ºW, 1-28.vii.1975, G. & M. Wood (17m#, 10f#, CNC); -21. M.e. Tuktoyaktuk, 20-25.vi.1971, D.M. Wood (1m#, CNC); Ford Lake, 63º11'N, 107º19'W, 27.vi.1966, G.E. Shewell (1m#, CNC). Nunavut: Axel Heiberg Is., 79º25'N, 90º45'W, 19, 21.vii.1963, H.K. Rutz (6f#, CNC); Axel Heiberg Is., 79º25'N, 90º45'W, Gypsum Hill, 20.vii.1963, H.K. Rutz (4f#, CNC); Axel Heiberg Is., 79º25'N, 90º45'W, Wolf Rd., 24.vii.1963, H.K. Rutz (1f#, CNC); Alex Fjord, 78.875, 75.799, Bowl Trap White & Blue, UBC: GH: 2256, 2283, 2603, 2248, 1685, 2162, 1122, 1104, 4081, 4-8.vii.2012, S. Robinson (2m#, 6f#, CNC); Alex Fjord, 78.875, 75.799, Netting, UBC:GH: 2162, 5.vii.2012, G. Galto (1f#, CNC); Hazen Camp, 81º49'N, 71º16'W, 19, 21, 24.vi.1962, J.F.McAlpine (7m#, CNC); 81º49’N, 71º18’W, Hazen Camp, 25.vi., 1.vii.1962, R.B. Madge (26m#, CNC); Hazen Camp, Ellesmere Is., 14-20.vii., 3.viii.1961, D.R. Oliver (10m#, CNC); Hot Weather Creek, Fosheim Peninsula, Ellesmere Is., 79º58'N, 84º28'W, 19, 22.vi., 1-9.vii.1990, F. Brodo (5m#, 13f#, CNC); Ellesmere Is., 79º58'N, 84º28'W, 2.viii.1989, S.A. Edlund (1m#, CNC); Tranquary Fjord, Ellesmere Is., 18.vii.1963, H.K. Rutz (10m#, 16f#, CNC); Hazen Camp, 81º49'N, 71º18'W, 1.viii.1963, J.R. Vockeroth Collector (3m#, CNC); Hazen L., Ellesmere Is., 11, 13.vii.1963, M.10, N.10, C.D. Sharplin (12f#, CNC); Hazen L., Ellesmere Is., 11, 13.vii.1963, T.5, N.10, B. Hocking (5m#, 17f#, CNC); North Fork Crossing, MI. 42 Peel Plt. Rd. 3500', 24.vi.1962, P.J. Skitsko (1m#, CNC). GREENLAND. Nedre Midsommer Sö, 26-29.vii., 1-30.vii.1966, Can. Peary Land Expedition (177m#, 104f#, CNC). Diagnosis. Male. Mid tibia with 3 anterodorsal setae dark, stout, longer than 2x

tibial width; 3 posterodorsal setae dark, stout, as long as 2x tibial width. Hind tibia with antero/posterodorsal and dorsal setae shorter than 2x tibial width (Fig. 3.30). Phallus mostly hidden within epandrium, except for basiphallus visible (Fig. 3.61). Female: Similar to male. Description. Wing length 7.5–8 mm. Male. Head dark greyish-blue in ground- colour, with greyish pruinescence on face, frons, postgena and occiput. Holoptic, eye with ommatidia larger on upper half, smaller on lower half. Face slightly more divergent than strictly parallel-sided towards mouthparts; bare with thin oral margin shiny. Ocellar triangle dark, subshiny, with 1 anterior pair of dark, parallel ocellar setae, with 2-3 pairs of dark, parallel posterior setulae 1/2 length of anterior ocellar setae; 2-3 pairs of postocular setae slightly shorter and finer than posterior ocellar setulae. Occiput bearing row of dark, long postocular setae only present on upper 1/2 of occiput; occipital setae dark, slightly shorter than postocular setae; setae of postgena similar to occipital setae. Antenna dark brown, with scape and pedicel paler than postpedicel. Scape and pedicel subequal to each other; postpedicel about 4x basal width; postpedicel basal width larger than pedicel width; stylus equal to postpedicel basal width. Palpus dark, with dark setulae. Labrum lustrous and dark reddish-brown; labellum dark and bearing dark setulae. Thorax dark greyish-blue in ground-colour, largely densely grey pruinescent: posterior corner of postpronotal lobe lustrous and dark brown; notopleural ridge dark brown; splr ridge with brownish tinge; region between posterior spiracle and halter base not pruinescent and brown-orange. Scutum with pair of indistinct, darker grey vittae between acr and dc rows; flanked by brownish tinges. Pleura mostly grey, with faint brownish shadows on sclerites. Proepisternum greyish, with 4 stout, dark setae; prosternum bare. Antepronotum with row of short, stout dark setae. Postpronotum with 3- 4 short, dark setae and 1 posterior longer and dark seta. Scutum with biserial row of dark acr setulae; dc longer than acr, increasing in length posteriorly; splr setae dark, long, stout, increasing in length posteriorly; postalar callus bearing 1 long, stout, dark seta; 3 anterior dark, stout npl setae; 2 posterior npl setae stouter and longer than anterior npl setae. Scutellum with apical sctl setae long, dark, stout; first pair of subapical scutellars dark, stout, long and convergent, although slightly shorter than apicals and never crossing over each other; 2nd pair of subapicals about ½ length of 1st pair of subapicals, finer than

apicals. Laterotergite with several long, stout, dark setae and a few shorter, finer setae. Anterior and posterior spiracles dark orange-brown with outer black ring. Legs long, entirely dark brown, except for darker rings at trochanter-coxae and trochanter-femur junctions. Coxae with dark lateral setae, longer and stouter along apical anterior margins. Ventral surface of trochanter with several dark setae, shorter and finer than coxal setae. Femora with short, white ventral pubescence; hind (Fig. 3.30) and mid femora with dark, stout anteroventral setae slightly longer on apical ½; fore femur with dark, finer antero/posteroventral setae, decreasing in length apically. Hind (Fig. 3.30) and mid tibiae similar, antero/posterodorsals dark, stout, longer than tibial widths, antero/posteroventrals dark, stout, shorter than tibial width; fore tibia with all setae shorter or subequal to tibial width, fine and clothed with short, silvery pile, giving silvery appearance. Basitarsi with setae dark, stout, shorter than ½ basitarsal widths; fore basitarsi with white ventral pubescence. Wing clear with yellowish brown veins; all veins complete (except Sc), well sclerotized. Pterostigma darker yellow, although faint. Base of costa with 1 longer, stouter, dark seta. Anal lobe well-developed. Axillary fissure forming rectangular angle. Halter yellow. Abdomen greyish-blue with brownish tinge; margins of segments pale; abdomen covered with black, stout setae, shorter on middle of tergites. Sternite 8 subtriangular and margin bearing stout, dark setae projecting outwards, not downwards. Tergite 8 reduced, barely visible. Terminalia (Fig. 3.61) dark grey, barely longer than abdominal width; epandrium with some brown; epandrium margin pubescent; apical tips of epandrium curved inwards towards each other with both tips merged; middle of epandrium bearing 8 dark, stout setae, longer than subepandial lobe, bearing cluster of dark, long stout setae at tips, curving towards each other. Cercus broadly V-shaped, middle bending into obtuse angle; anterior portion of cercus slimmer than posterior portion of cercus; dorsal margin of cercus bearing many short, fine, erect, dark setae. Subepandrial lobe cylindrical, projecting outwards slightly farther than cercus, slimmer than cercus; apex bearing short, fine, dark setae; spine-like setae at apex of lobe might be observed upon dissection. Hypandrium short, boot-shaped, ending in middle of swollen basiphallus; phallus lustrous

and reddish brown to yellow, not forming loops and tucked between epandrial lamellae; phallus short, not usually extending farther than epandrial lamellae; distiphallus tip with shallow S-shaped curvature; ejaculatory apodeme almost as large as epandrial width, fan- shaped, forming acute to subrectangular angle, lateral wings shorter than vertical wing. Female. Similar to male. Remarks: Oliver (1963) listed this species as R. sp. near lamelliseta Ringdahl Geographical distribution. Nearctic distribution in Figure 3.75..

Rhamphomyia (Pararhamphomyia) filicauda Henrikson & Lundbeck, 1917 Figs. 3.31, 3.32, 3.33, 3.62, 3.76

R. similata Malloch, 1919

Type material examined. HOLOTYPE: R. similata, Mall, Bernard Harbor, 18, 19.vi.1915, CNC No.217, Canadian Arctic Expedition (1m#, 1f#, CNC) Additional material examined. CANADA. Northwest Territories: Lady Melville L., 93º15'W, 69º25'N, 3.vii.1951, J.G. Chillcott (3f#, 2m#, CNC); Spence Bay, 6-22.vii.1951, J.G. Chillcott (16f#, 12m# CNC); Aklavik, 16.vi.1956, C.D. Bird (1m#, CNC); Aklavik, 26.vi.1956, R.E. Leech (1f#, CNC); Aklavik, 28, 29.vi.1956, E.F. Cashman (5m#, 1f#, CNC); Fort McPherson, 12.vii.1957, R. Hurley (2f#, CNC); Hazen Lake, Ellesmere Is., 11.viii.1963, M.10, C.D. Sharplin (1m#, CNC); Ellesmere Is., 79º29'N, 75º50'W, 18-24.vii.-1-15.viii.1980, J.S.E.P.M.B. (22m#, CNC); Coral Harbour, Southampton Is., 1-8.vii.1948, G.E. Shewell (29m#, CNC); Cambridge Bay, 18- 25.vii.1950, G.K. Sweatman (15f#, CNC); Cambridge Bay, 19-26.vii.1950, E.H.N. Smith (18f#, CNC); Hazen Camp, 81º49'N, 71º18'W, 14-25.vi.-4-8.vii.1962, R.B. Madge (6m#, 12f#, CNC); Hazen Camp, NE.372, 81º49'N, 71º16'W, 5,8.vii.1962, D.R. Oliver (1f#, CNC); Hazen Camp, Ellesmere Is., 9-22.vii.1961, D.R. Oliver (10m#, 5f#, CNC); Hazen Camp, 81º49'N, 71º16'W, 19-24.vi.1962, J.F. McAlpine (23m#, 23f#, CNC); Eureka, Ellesmere Is., 28.vii.1953, P.F. Bruggemann (10m#, 18f#, CNC); Victoria Is., 71º17'N, 114ºW, 1-30.vii.1975, G. & M. Wood (27m#, 12f#, CNC); Masik River, Banks Is., 3-

29.vii.1968, W.R.M. Mason (26m#, 6f#, CNC); Sachs Harbour, 27-30.vi.1968, W.R.M. Mason (6m#, 9f#, CNC); Hazen Lake, Ellesmere Is., 2-26.vii.1963, P.11, N.10, J.3, N.8, L.4.WHF, K.2, B. Hocking (11m#, 17f#, CNC); Hazen Lake, Ellesmere Is., 9- 17.vii.1963, M.10, A.9, C.D. Sharplin (11m#,19f#, CNC); Hazen Camp, 89º49'N, 71º18'W, K.2, 1-16.vii.1963, H.K. Rutz (3m#, 4f#, CNC); Tranquary Fjord, Ellesmere Is., 18.vii.1963, H.K. Rutz (4m#, 10f#, CNC); Hot Weather Creek, Forsheim Peninsula, Ellesmere Is., 79º58'N, 84º28'W, On Dryas flowers, wet meadows, fen, 18-28.vi., 1- 25.vii.1990, F. Brodo (14m#, 6f#, CNC). Nunavut: Banks Is., Aul. Natnl. Park, 73.22181, -119.56059, 7-19.vii.2011, wet & mesic, Pan, Repl.1-3, (23m#, LEMQ); Axel Heiberg Is., 79º25'N, 90º45'W, 19-23.vii.1963, H.K. Rutz (16m#, 15f#, CNC); Axel Heiberg Is., 79º25'N, 90º45'W, Gypsum Hill, 20-22.vii.1963, H.K. Rutz (10m#, 23f#, CNC); Axel Heiberg Is., 79º25'N, 90º45'W, Colour L., 22.vii.1963, H.K. Rutz (2m#, CNC); Axel Heiberg Is., 79º25'N, 90º45'W, Wolf Rd., 24, 25.vii.1963, H.K. Rutz (2m#, 5f#, CNC); Axel Heiberg Is., 79º25'N, 90º45'W, Wh. Glacier, 26.vii.1963, H.K. Rutz (1m#, CNC). Yukon: British Mts., 69º13'N, 140º05'W, 21-25.1984, G. & D.M. Wood & D. Lafontaine (2m#, CNC); Dempster Hwy., Mi.87, 1-30.vii.1973, G. & D.M. Wood (5m#, 8f#, CNC); North Fork Crossing Mi.42, Peel Plt. Rd., 3500’, 26.vi., 5.vii.1962, R.E. Leech (1m#, 6f#, CNC); Herschel Is, 28-29.vi.1971, D.M. Wood (1f#, CNC). USA. Alaska: Isabel Pass, Mi.206, Richardson Hwy., 2900', 13.vii.1962, R.E. Leech (3m#, CNC); Umiat, 4-12.vii.1959, J.E.H. Martin (2m#, 6f#, CNC); Umiat, 6-18.vii.1959, R. Madge (1m#, 8f#, CNC); Mi.213, Richardson Hwy., 17.vi.1951, W.R.M. Mason (1m#, CNC); Mi.213, Richardson Hwy., 17.vi.1951, J.R. McGillis (1f#, CNC). GREENLAND. Nedre Midsommer Sö, 3-24.vii.1966, T31-05, T34,-38-7, T36,37,38-10, Can. Peary Land Expd. (10m#, CNC); Nedre Midsommer Sö, 3.viii.1966, T9-13, T31-10, T32-12, T33-12, T33-6, T34-6, T34-12, T34-6, T37-12, T38-12, Can. Peary Land Expd., (3m#, CNC), 24.vi.-3,7.vii.-5.viii.1966, Can. Peary Land Expd (9m#, CNC); Nedre Midsommer Sö, 1- 30.vii.-4.viii.1966, Can. Peary Land Exp (92m#, 121f#, CNC). Diagnosis. Male. Tibiae with 2 rows of stout, dark posterodorsal setae, subequal to tibial width; posterior of fore tibia clothed with many fine setae, longer than ventral setae but shorter than dorsals (Figs. 3.31, 3.32, 3.33); hind femur with several rows of antero/posteroventral and ventral setae increasing in stoutness towards apical ½ and

decreasing in length towards apical ¼ (Fig. 3.32). Cerci strongly constricted in middle, posterior ½ strongly divergent from epandrium (Fig. 3.62). Female. Similar to male. Description. Wing length 11.6–11.8 mm. Male. Large species. Head dark in ground-colour, with dense greyish pruinescence on face, frons, postgena and occiput. Holoptic, eye with ommatidia larger on upper half, smaller on lower half. Frons triangular towards antennae. Face triangular sided, with oral margin dark brown and shiny. Ocellar triangle dark, subshiny, with 1 pair of anterior dark, long, stout, parallel ocellar setae and 1 pair of posterior dark, stout, parallel ocellar setae of subequal length to anterior ocellar setae; 3 pairs of postocellar setae dark, finer, shorter than ocellar setae. Occiput bearing dense row of postocular setae, stout, black, long only present on upper occipital section; occipital setae slightly shorter than postocular setae; postgena bearing many setae slender than occipital and postocular setae. Antenna dark brown, with faint inner paler coloration. Scape about 2x pedicel length; pedicel bulbous; pedicel and scape bearing stout, dark, short setae; postpedicel more than 4× longer than basal width; subequal to length of scape. Palpus dark brownish-black, with setulae long and dark. Labrum lustrous and dark reddish-brown with pale apex; labellum with ventral side dark black and bearing dark setulae, with laterals white and bare. Thorax dark in ground-colour, largely densely grey pruinescent, densely covered with setae: posterior corner of postpronotal lobe lustrous and dark yellow-brown, npl ridge orange-brown, margin between anepisternum and anepimeron pale; region around wing base lustrous and reddish-brown; scutum with brownish colouring. Prescutum and scutum with pair of distinct dark vittae between acr and dc rows, flanked on both margins with brownish tinges. Pleura dark grey, with brownish shadows. Proepisternum greyish, with many long, stout, dark setae and with one slightly longer and stouter seta; prosternum bare. Propleuron bare. Antepronotum with dense row of stout and dark setae, shorter in middle, longer on laterals. Postpronotal lobe with dense cluster of long and short, stout and finer setae, with 1 seta slightly longer and stouter. Scutum with dense biserial row of dark, stout acr increasing in length posteriorly; dc slightly longer than acr, increasing in length posteriorly; notopleuron bearing anterior cluster of long, dark, stout setae, and 6 longer, stouter, dark setae; postalar callus with brown colouring, anterior cluster of several short, fine setae and 1 posterior long, stout, dark seta; dense rows of

dark, long, stout, intlr and splr setae slightly finer than posterior dc setae. Scutellum with 1 pair of dark, stout, long, convergent apical sctl; pair of subapical sctl dark, stout, long, convergent; 4 pairs of stout, dark, convergent apical sctl slightly shorter than apicals and subapicals; basal sctl dark, finer and slightly shorter than apicals. Laterotergite bearing dense cluster of dark, some stout, some finer, some longer and some shorter setae. Anterior and posterior spiracles dark. Legs entirely pollinose, long, entirely dark with brownish tinges, especially on coxae. Hind coxa with numerous dark, stout, lateral setae; mid and fore coxae with many long, stout setae on anterior region. Trochanter with short, stout, dark setae mostly concentrated on ventral surface. Hind femur (Fig. 3.32) multiple rows of anteroventrals and ventrals on apical 2/3 numerous, stout, dark, never longer than femoral width; mid femur (Fig. 3.33) with ventral row of dark, stout, short setae, posteroventral setae dark, 2x length of ventrals, antero/posteroventrals and ventrals decreasing in length posteriorly; fore femur with ventrals absent, antero/posteroventral setae dark, finer than those of mid and hind femora. Hind tibia (Fig. 3.32, 3.33) with antero/posterodorsal and dorsal setae dark, stout, no longer than tibial width, anteroventral row of dark, stout setae, shorter than antero/posterodorsals, with cluster of dark, fine setae shorter than anteroventrals on apical ¼. Mid tibia (Fig. 3.33) with antero/posterodorsal and dorsal setae similar to those of hind tibia, except longer than tibial width; antero/posteroventral setae dark, stout, shorter than tibial width; preapical antero/posteroventral and ventral setae dark, stouter than remaining tibial setae and 2x length of antero/posteroventrals. Fore tibia similar to mid tibia except as follows: ventrals absent, many rows of anterior and posterior setae fine, short, dark; antero/posteroventrals and ventrals fine and short. Basitarsi with preapical ring of dark, stout setae never longer than 2x basitarsal width; hind basitarsi longer and larger than mid and fore basitarsi. Mid and hind basitarsi (Fig. 3.31, 3.32) with antero/posteroventrals/dorsals stout, dark; antero/posteroventrals/dorsals of hind basitarsus shorter than width; antero/posterodorsals of mid basitarsus (Fig. 3.33) longer than width, but shorter than 2x width. Fore basitarsi with posterodorsal setae stout, dark, subequal to 2x fore basitarsal width, with stout, dark ventral setae on proximal ½, with remaining setae finer, shorter. Tarsomeres with strong ventral white pubescence.

Wing clear with yellowish brown veins; all veins complete (except Sc), well sclerotized. Anal lobe well-developed; axillary fissure forming rectangular angle. Coastal base bearing one dark seta that is longer and stouter than other costal setae. Pterostigma present and dark yellow. Anal vein not reaching wing margin. Vein junctions sometimes thickened and clouded. Halter stem black; knob pale. Abdomen dark with brown tinges on sclerites and tergites, with pale margins. Abdomen clothed with dark, stout setae. Laterals of tergites bearing longer setae than middle of tergites. Middle of sternites bearing long, stout setae. Margin of sternite 8 bearing many long, stout, dark setae. Sternite 8 subtriangular. Tergite 8 greatly reduced, also bearing dark setae, setae slightly shorter than setae of tergite 8. Pregenital segments dark brown with margins black. Terminalia (Fig. 3.62) dark. Epandrium width constricted in middle, forming elongated, stout, rounded anterior projection; projection as long or slightly longer than basal length of epandrium; epandrium ventral margin bearing many dark, stout setae; apex dorsal margin bearing many fine, short, pale setae; epandrium dark brown with black margins; pale patch on dorsal ½ of anterior projection. Cercus enlarged, almost as long as epandrium length, constricted in middle, forming acute V-shape; cercus bearing many short, dark setae, finer on posterior lobe than anterior lobe. Hypandrium slim, short, ending at base of swollen basiphallus; distiphallus brown, hair-like, with slight inwardly curve towards middle, forming loop extending farther than genitalia and between cerci. Ejaculatory apodeme held downwards, broadly fused to phallus, small, subrectangular, with flat dorsal margin, lateral wings shorter than vertical wings. Female. Similar to male. Geographical distribution. Nearctic distribution in Figure 3.76.

Rhamphomyia (Pararhamphomyia) ursinella Melander Figs. 3.334, 3.36, 3.63, 3.76

Rhamphomyia ursina Malloch, 1919: 46. (preoccupied) Rhamphomyia ursinella Melander, 1928: 209. (replacement name)

Type material examined. HOLOTYPE: m#, “HOLOTYPE/ Rhamphomyia ursinella Mel./ CNC No. 216 [red label]”; “Canadian/Arctic/Expedition/F.J. 1915”; “Sandy Beach/ July, 19”; “Rhamphomyia/ursina/ Mall. Type”. PARATYPES: 2f#, same data as holotype. Additional material examined. CANADA. Northwest Territories: Hazen Camp, Ellesmere Is., 81º49'N, 71º18'W, 6.vii.1962, R.B. Madge (15m#, 34f#, CNC); Mould Bay, Prince Patrick Is., 3.vii.1952, P.F. Bruggemann (4m#, 1f#, CNC); Eureka, Ellesmere Is., 3.viii.1963, J.R. Vockeroth (1m#, 1f#, CNC); Ellesmere Is., 79º58'N, 84º28'W, 7-14.vii.1988, 2.viii.1989, S.A. Edlund (3m#, 5f#, CNC); Aklinck R. S. coast of Bylot, Vial 19, 50, 53, & 55 (1m#, 3f#, CNC); Fosheim Peninsula, Hot Weather Creek, Ellesmere Is., 79º58'N, 84º28'W, 5.vii.1990, F. Brodo (1m#, 1f#, CNC); Lady Melville L., 93º15'W, 69º25'N,3.vii.1951, J.G. Chillcott (2m#, 9f#, CNC). Nunavut: Axel Heiberg Is., 79º25'N, 90º45'W, 19, 23.vii.1963, H.K. Rutz (2f#, CNC); Axel Heiberg Is., 79º25'N, 90º45'W, Gypsum Hill, 20.vii.1963, H.K. Rutz (1m#, 6f#, CNC). Diagnosis. Male. Basitarsi with antero/posterodorsal and dorsal setae 2x the length of hind, mid and fore basitarsi, respectively; hind basitarsi swollen such that proximal width smaller than apical width (Fig. 3.34). Fore tibia with antero/posterodorsal and dorsal setae more than 2x the length of tibial width (Fig. 3.36). Cercus weakly constricted in middle, posterior ½ parallel to epandrium (Fig. 3.63). Female. Similar to male. Re-description. Wing length 7.6-7.8 mm. Male. Head dark brown in ground- colour, with light greyish pruinescence on face, frons and occiput. Holoptic, with ommatidia larger on upper half, smaller on lower half. Frons divergent towards antennae; face parallel, although slightly divergent towards mouthparts; bare with oral margin dark brown and shiny. Ocellar triangle dark, subshiny, with 1 anterior pair of parallel, slightly divergent, dark ocellar setae and 1 posterior pair of divergent, dark ocellar setae slightly shorter than the anterior ocellar setae. Occiput bearing row of dark, fine, long postocular setae, only present on upper half of occiput. Occiput bearing many fine, dark setae shorter than postocular setae; posterior postgena setae long; anterior postgena setae short, about ½ length of posterior setae. Postpedicel, stylus and scape mostly dark brown; pedicel and apex of scape slightly paler than post-pedicel. Scape slightly more than 2×

longer than pedicel; postpedicel about 4× longer than basal width; postpedicel width decreasing towards stylus; stylus slightly shorter than postpedicel basal width; scape and pedicel bearing dark, short setulae. Palpus dark and bearing many dark, fine, long setulae. Labrum lustrous and dark reddish brown; labellum dark and bearing fine, short, dark setulae. Thorax dark reddish-brown in ground-colour, with lustrous patches and sparse grey pruinescence. Scutum with pair of distinct lustrous blackish vittae between acr and dc rows, separated by broad greyish pruinescent stripe; and flanked on sides with greyish pruinescence. Pleura dark reddish brown, with lustrous patch on anepisternum; posterior corner of postpronotal lobe lustrous and orange-brown; postpronotal lobe often paler reddish brown. Proepisternum with yellow-brown colouring, with few short, fine setae and few longer, fine setae; prosternum bare. Antepronotum with row of short, stout, dark setae. Postpronotum with several long, dark, fine setae and a few shorter anterior setae. Scutum with biserial row of fine, long acr setulae; dc similar to acr, increasing in length and stoutness posteriorly; several presutural splr, 1 postsutural splr; 2 stout, long pslr (most distinctive ones), and 1-2 shorter, finer, dark setae; many fine, dark, long npl and 1 posterior seta longer and stouter. Scutellum bearing pair of long, stout, dark apical sctl setae; dark, stout subapical scutellar setae slightly shorter than apicals; 3 pairs of sctl setae slightly finer and shorter than subapicals; 2 pairs of shorter, finer sctl setae about 1/3 length of subapicals. Laterotergite bearing patch of fine, dark setae. Anterior and posterior spiracles dark. Legs long, entirely brown, mostly lustrous. Upper coxae with black colouring. Hind coxa with numerous dark lateral setae; mid and fore coxae with many lateral and dorsal dark setae. Hind trochanter with many ventral dark, short, stout setae; mid and fore trochanter nearly bare. Hind tibia lustrous except for anterior white pubescence; antero/posteroventral setae short, fine, except row of anteroventral setae on apical 1/3 at least 4x length of remaining anteroventrals; posterior setae of apical ¼ as long as femoral width, fine, dark (Fig. 3.34). Mid (Fig. 3.36)and fore tibiae with antero/posteroventrals fine, dark, subequal to mid and hind femoral width, respectively, remaining setae shorter than ½ femoral width; ventral surface with pubescence. Tibiae (Fig. 3.34, 3.36) with antero/posterodorsal and dorsal setae dark, fine, at least as long as 2x tibial widths;

antero/posteroventral and ventrals fine, shorter than tibial widths; hind tibia with apical seta stout, dark, subequal to tibial width (Fig. 3.34). Hind basitarsus swollen apically; (Fig. 3.34) mid and fore basitarsi not swollen; mid basitarsus shorter than fore and hind basitarsi; basitarsi with numerous antero/posterodorsal and dorsal setae fine, dark, at least as long as 3x basitarsal widths; antero/posteroventral and ventral setae shorter than at most ½ basitarsal width. Hind tarsomeres 2 and 3 with numerous, fine antero/posterodorsal and dorsal setae, at least 3x longer than tarsomere width (Fig. 3.34). Wing clear with yellowish veins; all veins complete (except anal vein), well sclerotized. Anal lobe well-developed, axillary fissure forming rectangular angle. Pterostigma nearly absent. Halter dark. Abdomen dark reddish brown, and lustrous. Margins of tergites and sternites pale. Abdomen clothed with dark, fine setae. Tergite setae longer towards thorax, shorter and finer towards terminalia. Sternite 8 subtriangular and larger than tergite 8; margin of sternite 8 bearing many long, dark, stouter, erect setae. Terminalia (Fig. 3.63) entirely dark. Epandrium brown in ground colour and with black tinges; long, slightly constricted in middle, forming two swollen “lobes”; epandrium bearing many dark, fine, long setae, decreasing in length towards epandrium apex. Cercus pollinose, almost as long as epandrium; slightly constricted elongated middle section, forming anterior portion larger than posterior portion; except for middle section; dorsal margin bearing many short, fine setae; inner surface with one short, stubby projection. Hypandrium slim, rectangular, short, ending at start of swollen basiphallus; phallus hair-like, not forming loops, arched back between epandrium or cercus. Ejaculatory apodeme small, acutely fan-shaped, with lateral wings shorter than vertical wing, with dorsal margin of vertical wing blunted. Female. Similar to male except as follows: ground colour more black than brown; tibiae never swollen; hind basitarsi not swollen. Remarks: Rhamphomyia ursinella is assigned to the “caudata” group (most likely defined by T. Saigusa), based on the European species R. caudata (Zetterstedt). This latter species has not been collected in the Nearctic. Geographical distribution. Nearctic distribution in Figure 3.76.

Rhamphomyia (Pararhamphomyia) caudata gp. sp. 7 Figs. 3.35, 3.64, 3.76

Material examined. CANADA. Northwest Territories: Victoria Is., 71º17'N, 114ºW, 23-28.vii.1975, G. & M. Wood (20m#, 10f#, CNC); Masik River, Banks Is., 17, 27.vii.1968, G.E. Shewell (3m#, 1f#, CNC); Coral Harbour, Southampton Is., 6, 7.vii.1948, G.E. Shewell (11m#, CNC); Cockburn PI., Coll. 905, 5.ix.1914, Canadian Arctic Expedition, F. Johansen (1m#, CNC); Netilling Lake, Baffin Is., 9.vii.1925, J.D. Soper (1m#, CNC). Nunavut: Cambridge Bay, 18-26.vii.1950, G.K. Sweatman (17m#, CNC). Diagnosis. Male. Hind basitarsi clubbed shaped, such that proximal width larger than apical width, elongated, about ½ the length of hind tibia, and antero/posterodorsal and dorsal setae 2x basitarsal width; hind tibia with distinctive inward bend (Fig. 3.35). Apex of epandrium truncate; epandrial setae pale (Fig. 3.64). Fore tibia with strong posterior pubescence. Coxae with pale, hair-like, long setae and 2-3 apical setae, stouter and dark. Description. Wing length 11.1–12 mm. Male. Head dark in ground-colour, with greyish pruinescence on face, frons, occiput and postgena. Holoptic, eye with ommatidia larger on upper half, smaller on lower half. Frons divergent towards antennae; upper margins of eyes bearing fine, dark, short setulae; face slightly divergent towards mouthparts; bare, with oral margin dark brownish-black and shiny. Ocellar triangle dark, subshiny, with 1 anterior pair of long, dark, parallel ocellar setae and 1 shorter, posterior pair of dark, finer, divergent posterior setulae; 2 pairs of short, fine, dark postocular setae. Occiput bearing row of postocular setae, fine and dark, only present on upper section; occipital setae black, long and fine; postgena setae fine, hair-like, long and pale. Postpedicel, stylus and scape mostly dark, clothed with dense grey pubescence, with brownish tinge on pedicel. Scape 2× longer than pedicel; pedicel width greater than scape width; pedicel bulbous; postpedicel nearly 5× longer than basal width; stylus shorter than scape, thin and cone-like. Palpus dark, densely pubescent, with setulae long, dark and fine. Labrum lustrous and dark reddish brown; labellum grey with dark setae.

Thorax dark brownish to black in ground-colour, with light grey pruinescence. Scutum with pair of distinct glossy and black vittae between acr and dc rows, separated by distinct grey pubescent stripe and flanked by glossy black patch on outer laterals of vittae. Pleura mostly dark with some reddish-brown tinges on sclerites; posterior corner of postpronotal lobe lustrous and dark orange-brown. Proepisternum dark in ground- colour, clothed with dense grey pruinescence, bearing several fine, pale, slightly shorter than postgena setae; prosternum bare. Antepronotum with row of short, stout, pale golden setae. Postpronotum with several fine, hair-like, pale setae never longer than postgena setae and some setae shorter than others. Scutum with biserial row of sparse, fine, short pale, acr setulae; dc similar to acr, except only slightly longer than dc, with last acr setae stout, black and longer; notopleuron with dense anterior cluster of fine, pale setae, slightly longer than postpronotal setae, 3 stouter, longer, pale posterior setae and 1 black, stout, long posterior seta; sparse row of fine, short, pale splr setae; postalar callus yellowish brown and densely pruinescent, bearing one long, stout, black seta. Scutellum margin densely pubescent; with pair of long, stout, dark apical sctl; 1 pair of dark, stout, slightly shorter than subapicals, with distinct convex curve; 3 pairs of sctl; sctl finer, lighter, shorter. Laterotergite with dense cluster of pale, stout setae; setae no longer than npl setae. Anterior and posterior spiracles dark; middle of anterior spiracle dark orange- brown. Legs long, entirely dark brown, except for inconspicuous darkened area around hind femoral-tibial junction. Hind and mid coxae with many pale, fine lateral and dorsal setae; fore coxa with many pale, fine dorsal setae. Posterior outer side of hind coxa bearing 2 stouter, longer, and black setae. Hind trochanter bearing several fine, pale setae, mostly concentrated on ventral side; mid and fore trochanters bearing fewer short, fine, pale setae, also mostly concentrated on ventral side. Femora lustrous except for white ventral pubescence. Hind femur with anteroventral setae pale, shorter than femoral width and cluster of dark, stouter apical anteroventral setae, 2x length of pale anteroventrals; apical posterodorsal setae pale and of subequal lengths to apical anteroventrals (Fig. 3.35). Mid femur with posteroventral setae dark, stouter than setae of hind femur; mid posteroventrals 2x length of apicals and proximals; fore femur nearly bare. Tibiae with posterior region clothed with many fine, short setae; hind tibia with proximal inward

bend, posterodorsal setae fine, 2x tibial length (Fig. 3.35); mid tibia with anteroventral setae dark, stout, shorter than or subequal to tibial width, anterior region also clothed with many fine, short setae; fore tibia with at least 3 preapical anterior setae stouter than remaining fore tibial setae. Hind basitarsus clubbed such that basitarsal width about 2x to 3x larger than mid and fore basitarsal widths (Fig. 3.35); antero/posterodorsal and dorsal setae 3x as long as basitarsal width, dark, stout antero/posteroventral setae, shorter than basitarsal width and lighter than antero/posterodorsals and dorsals; posterior region of basitarsi clothed with many short, fine, pale setae; mid basitarsus with proximal antero/posteroventrals darker, stouter, longer than remaining mid basitarsal setae, shorter than 2x basitarsal width; fore basitarsus with strong white pubescence on anterior, posterior and ventral regions. Wing clear with yellowish veins; all veins complete and well sclerotized, except anal vein faint towards wing margin. Anal lobe well-developed; axillary fissure forming rectangular angle. Costa near wing base bearing single longer, stouter, dark seta. Pterostigma present: dark orange-brown. Halter mostly yellow with darkened stem base. Abdomen glossy, mostly dark brown, although sternites more blackish, lower abdomen mostly blackish-brown, margins of segments pale. Abdomen covered in fine, pale, long and shorter setae. Marginal setae on sternite 8 longer than length of sclerites and stouter. Tergite 8 constricted in middle, smaller than sternite 8 and other tergites. Terminalia (Fig. 3.64) dark and subequal to length of thorax. Epandrium elongated, middle region constricted, forming 2 “lobes”: posterior lobe pale, larger and longer than anterior lobe; anterior lobe curving inwards, then curving outwards away from opposite anterior lobe, forming S-shaped curvature; epandrium lobes bearing many pale, fine, short and longer setae. Epandrium setae of ventral margin merging towards each other, forming “drape” of setae closing opening left by epandrium; posterior lobe of epandrium bearing distinctive row of at least 5 long, fine setae. Cercus slimmer than epandrium, elongate, shorter than epandrium; middle of cercus constricted and lengthened, also forming two “lobes”; posterior lobe dark, with middle brown and anterior lobe pale, with irregular; dorsal margin of cercus bearing many fine, short, pale setae; anterior lobes of cerci with stubby, short projections. Dorsal view of cerci showing rounded lozenge-shaped opening closed at both ends. Hypandrium short, slim,

rectangular, ending at start of swollen basiphallus; phallus hair-like, without loops, and curving back to epandrium. Ejaculatory apodeme broadly fused to phallus, small, acutely fan-shaped, lateral wings shorter than vertical wing. Female. Not examined. Geographical distribution. Nearctic distribution in Figure 3.76.

Rhamphomyia herschelli Malloch, 1919 Figs. 3.1, 3.2, 3.38, 3.39, 3.65

Type material examined. SYNTYPES 1 f#, “HOLOTYPE/ Rhamphomyia herschelli Mall./ CNC No. 218 [red label]”; “Herschell Isl./ N.W.T.”; F. Johnsen/ Coll.”; Canadian/ Arctic/ Expedition/ July 29, 1916”; “Rhamphomyia/ herschelli/ type” (CNC); 1 f# “PARATYPE/ Rhamphomyia herschelli Mall./ No. 218 [yellow label]”; “Herschell Isl./ N.W.T.”; F. Johnsen/ Coll.”; Canadian/ Arctic/ Expedition/ July 29, 1916” (CNC). Additional material examined. CANADA. Labrador: Tessiujak Bay, Nagvak Fjord Lab, 15.viii.1954, J.F. McAlpine (1♀, CNC). Northwest Territories: Kidluit Bay, N. Richards Is. W. 25, 29.vii.1948, J.R. Vockeroth (1♂, CNC); Coppermine, 6.viii.1951, S.D. Hicks (1♂, CNC). Yukon: Herschel Is., 9, 16, 22, 23, 26, 28, 29, 30.vii.1953, C.D. Bird (15♂, 34♀, CNC); Herschel Is., 11, 22, 23, 26, 28, 29, 30.vii.1953, J.S. Waterhouse (41♂, 33♀, CNC). USA. Alaska: Umiat, 10, 23.vii.1950, R. Madge (2♂, CNC). Diagnosis. Male. This species is distinguished by the horn-like lateral projections on sternite 8 (Figs. 3.1, 3.2); phallus with sharp or acute bend in loop (Fig. 3.38); hind basitarsus swollen, wider than hind tibial width (Fig. 3.3). Female. Mid and hind femora with pennate setae located posteroventrally; pennate setae shorter than width of femur; hind basitarsus swollen, subequal to or slightly more than width of apical hind tibia; body glossy and brown; female slightly smaller or subequal to males. Description. Wing length 10.3–10.5 mm. Male. Head dark in ground-colour, with greyish-blackish pruinescence on frons; dark, glossy face; postgena and occiput greyish-blackish pruinose. Holoptic, eye with ommatidia of larger size on upper half, smaller size on lower half. Frons small and triangular, subequal to width of bulging ocellar triangle. Face parallel-sided towards mouthparts; bare with oral margin shiny.

Ocellar triangle dark, subshiny, with 2 pairs of parallel ocellar setae projecting forward; first pair located anteriorly and longer than second pair located posteriorly; pair of divergent posterior setulae, posterior to ocellars as long as length of anterior ocellar setae. Occiput bearing row of black postocular setae projecting forward and often strongly curved, slender, longer than anterior ocellar setae on upper half; shorter and more slender on lower section; occipital setae black, long and stout. Antenna black and pruinose. Scape slightly longer than pedicel; pedicel bulbous; postpedicel basal width broader than width of pedicel, gradually tapered apically; about 2× longer than stylus; stylus shorter than scape and pedicel combined. Palpus black, pruinose with dark setulae; labrum largely dark brown and glossy; apex of labrum yellowish brown; labellum with dark setae. Thorax dark in ground-colour, with grey pruinescence. Scutum without pair of vittae between acr and dc rows in anterior view; pair of grey pruinescent stripes in posterior view. Pleura entirely dark brownish-black, lacking paler highlights. Proepisternum dark brownish-black, with several long, dark setae; prosternum bare. Antepronotum dark, pruinose, with 5 pairs of dark lateral setae projecting outwards. Postpronotum with 1 stout pprn and nearly 10 dark setae of various lengths. Scutum with long and dark biserial acr and multi-serial dc; dc slightly longer than acr, increasing in length posteriorly; 3 npl and several shorter dark setae; 1 presut spal and several shorter, thinner setae; 1 psut spal and numerous anterior setae; 1 pal and 1-2 short setulae. Scutellum with 2 pairs of convergent sctl, apical pair longer than outer pair. Laterotergite with crowded cluster of long dark setae. Anterior and posterior spiracles dark. Legs long, entirely dark. Coxae with numerous dark lateral setae; trochanter with long, similar-length setae. Femora with white, ventral pubescence; hind femur with at least 2 rows of anteroventral setae dark, shorter than half femoral width, many rows of posterior setae dark, longer on proximal half, shorter on apical half, but never longer than femoral width; mid femur with at least 2 rows of antero/posteroventral setae, dark, shorter than half femoral width, except small cluster of setae 2x length of remaining antero/posteroventrals on proximal ¼; fore femur with sparse cluster of dark, fine setae on proximal ¼ no longer than femoral width. Hind tibia with posterodorsal setae dark, 2x length of antero/posteroventrals, antero/posterodorsal preapical setae stout, dark, subequal to proximal tibial width (Fig. 3.3); mid tibia with posterodorsal setae dark,

stout, as long as 3x tibial width, preapical antero/posterodorsal setae dark, stout, longer than 3x tibial width, antero/posteroventral stouter, dark, never longer than 2x tibial width, preapical posteroventral seta stouter, slightly longer but no longer than 2x tibial width; fore tibia with posterodorsal setae dark, fine, shorter than 2x tibial width, preapical posterodorsal as long as 2x tibial width, antero/posteroventral setae finer, dark, shorter than tibial width, except preapical antero/posteroventral setae stouter and slightly longer but still shorter than tibial width, anterior preapical seta stout, dark, similar to antero/posteroventral preapicals. Hind basitarsus swollen, broader than hind tibial width, setae dark, stout, antero/posteroventral and ventral setae half-length of antero/posterodorsal and dorsal setae, never longer than 2x basitarsal width (Fig. 3.3); fore and mid basitarsi similar, setae fine, dark, never longer than 2x basitarsal width, except antero/posteroventrals of mid basitarsus stout, preapical antero/posterodorsals stouter and slightly longer than antero/posterodorsals. Wing infuscate, clear with yellowish brown veins; all veins complete (except subcostal) and well sclerotized. Pterostigma faint but present. Anal lobe well-developed. Alular incision forming rectangular angle. Halter dark; knob dark brown. Abdomen dark (black) and glossy; tergite marginal setae shorter than dorsal tergite setae. Lateral tergite setae long and dark; mid sternite setae longer than lateral tergite setae. Sternite 7 forming fan-shaped U, heavily sclerotized and broadened such that width larger than other abdominal segments; with deep vertical groove running down length of sternite and located in middle of sternite. Sternite 8 enlarged and modified, laterally with “horn-like” projections (Figs. 3.1, 3.2, 3.39). Terminalia (Fig. 3.38) glossy black, concolourous with abdomen, except phallus dark reddish-brown. Epandrium triangular, subequal in length with cercus. Epandrium anterior margin lined with dark, stout setae, decreasing in length towards epandrium tip; epandrium tip rounded. Cercus slender, less than ½ width of epandrium; cercus cylindrical, rounded, dorsal margin bearing many dark, fine, erect setae shorter, subequal to, or slightly longer than cercus width. Dorsal view of cercal lobes with two finger-like projections joining together forming O-shaped opening. Hypandrium tubular. Basiphallus pale and swollen, S-shaped; phallus darker, slender, with sharp bend forming acute angle. Epandrium triangular, subequal in length with cercus. Ejaculatory apodeme small, fan-

shaped, vertical wing subequal to lateral wings; apodeme stem mostly membranous, and loosely connected to hypandrium. Female. Mid and hind femora with pennate scales located posteroventrally; scales slightly shorter than width of femur; anterodorsal row of setae thickened, less developed than pennate scales. Hind basitarsus swollen, slightly broader than apical hind tibia. Body glossy and brown; female slightly smaller or subequal in size to male, not larger. Geographical distribution. Nearctic distribution in Figure 3.65.

Rhamphomyia sp. 12 Figs. 3.4, 3.37, 3.65

Material examined. CANADA. Northwest Territories: Chesterfield, 2-30.viii.1950, J.R. Vockeroth (33m#, 27f#, CNC); Baker Lake, 26.vii., 8,11.viii.1947, T.N. Freeman (2m#, 3f#, CNC); Bathurst Inlet, 17-20, 27.vii., 1,2.viii.1951, W.I. Campbell (19m#, 17f#, CNC). Nunavut: Frobisher Bay, Baffin Is., 17.vii.1948, T.N. Freeman (1m#, 5f#, CNC); Lake Harbour, Baffin Is., 6, 9.viii.1935, W.J. Brown (6m#, 12f#, CNC). Diagnosis. Male. Phallus with small, tight U-shaped loop (Fig. 3.37); hind tibia with several rows of stout anteroventral setae (Fig. 3.4), shorter than width of tibia, and several rows of fine anteroventral setae on apical 1/3. Female. Hind femur with posteroventral pennate setae, and pennate setae around middle of hind femur as long as width of femur; hind tibia with pennate setae both anterodorsally and anteroventrally, and setae shorter than width of hind tibia; three evenly spaced, dark, and longer than acrostichals, notopleural setae. Description. Wing length 10–11.6 mm. Male. Head dark in ground-colour, with greyish pruinescence on face, frons, postgena, and occiput. Holoptic, eye with ommatidia of larger size on upper half, and smaller size on lower half. Frons small and triangular, subequal to width of bulging ocellar triangle. Face parallel-sided towards mouthparts; bare with oral margin shiny. Ocellar triangle dark, subshiny: 1 pair of anterior parallel ocellar setae about 2x length of ocellar triangle, 1 pair of posterior diverging ocellar setae about 1/3 of length of anterior ocellar setae, with 2 pairs of posterior setulae about 1/2 length of anterior ocellar setae. Occiput bearing row of postocular setae only present on

upper half of occiput, stout, long and black; occipital black, stout and shorter than postocular setae; postgena pruinose and bearing dark, long, finer setae. Antenna dark brown and pruinose. Scape length about double length of pedicel. Pedicel bulbous; postpedicel less than 4× longer than basal width. Stylus no longer than length of scape and pedicel combined. Palpus dark brown, pruinose, with setulae dark. Labrum glossy and dark reddish-brown; labellum with dark setae. Thorax dark black in ground-colour, with grey pruinescence; posterior corner of postpronotal lobe lustrous and brown; notopleural ridge yellowish orange. Scutum with indistinct pair of vittae lateral to dc and above pronotal depression. Pleura black with dark reddish-brown shadows on sclerites; sclerite margins sometimes pale. Proepisternum dark, with several fine, short, dark setae; prosternum bare. Antepronotum dark, pruinose, bearing row of stout, dark setae. Postpronotal lobe bearing several anterior dark, short setae, and 1-2 stouter posterior setae, at least 2x length of anterior postpronotal setae. Postalar callus bearing 1 stout, dark, long seta. Scutum with biserial row of dark, long acr setulae; dc of equal length to acr; 3-4 supra-alar setae as long as acrostichals; 3 dark, long, stout notopleural setae; scutellum with apical scutellar setae crossing over one another, dark and longer than subapical scutellar setae. No discal scutellar setae or basal scutellar setae. Laterotergite with crowded cluster of long dark setae. Posterior and anterior spiracles pale. Legs long, entirely dark reddish-brown. Coxae pruinose and with numerous dark lateral setae of subequal lengths; hind coxae with anterior glossy patch, with posterior setae longer than anterior coxal setae that project downwards. Trochanter with curved spine-like setae confined to ventral medial patch and subequal to coxal setae. Hind femur (Fig. 3.4) with several antero/posteroventral and ventral setae dark, fine, shorter than femoral width, except several stouter setae on proximal ½, longer, but never longer than femoral width. Mid femur with strong ventral pubescence, anteroventral setae dark, stout, shorter than ½ femoral width, spine-like; 3 proximal anteroventrals 2x length of remaining anteroventrals; posteroventral setae dark, stout, 2x length of anteroventrals; second row of posteroventral setae on proximal 1/3 stout, dark, half length of longer posteroventrals. Fore femur similar to mid femur, except anteroventrals not stout. Hind tibia (Fig. 3.4) with several rows of antero/posterodorsal setae, consisting of rows of fine

setae shorter than tibial width, and row of stout setae as long as 2x tibial width; anteroventral seta stout, dark, spine-like, shorter than half tibial width; ventral setae finer, dark, 3x length of anteroventrals; preapical anteroventral cluster of dark, short setae, and posterior apical seta stout, dark, as long as tibial width. Mid tibia with ventral pubescence, at least 3 dorsal setae dark, stout, as long as 3x tibial width; several anterior rows of short, dark, fine setae; anteroventral setae dark, stout, spine-like, barely longer than pubescence; posteroventral setae stout, dark, about 4x length of anteroventrals; antero/posteroventral preapical setae dark, stout, no longer than tibial width. Fore tibia with many rows of posterior, postero/anteroventral, ventral, and anterior setae short, fine; antero/posterodorsal setae fine, dark, as long as tibial width. Hind basitarsus subequal to tarsomeres 2-5 combined, with proximal width slightly larger than apical width but less than 2x as large; several rows of antero/posterodorsal and dorsal setae shorter than basitarsal width, dark, fine; antero/posterodorsal rows of stouter, dark setae as long as proximal basitarsal width; posterior clothed with fine, short, lighter setae; antero/posteroventral setae light-colored, stout, shorter than basitarsal width; mid basitarsus similar to hind basitarsus except as follows: shorter than tarsomeres 2-4, and preapical antero/posterodorsal setae 2x length of basitarsus; fore basitarsus clothed with fine setae, no longer than basitarsal width. Wing clear with yellowish brown veins; all veins complete (except Sc), well sclerotized. Anal lobe well-developed with axillary incision forming acute angle.

Pterostigma faint but present. Vein R2+3 heavily sclerotized, especially at base. Halter yellow. Abdomen dark reddish-brown; tergal marginal setae shorter than ventral and dorsal abdominal setae. Margins of segments pale. Setae on laterals of tergites longer than setae of tergite margins; sternite setae shorter and finer than tergal setae, arranged in clusters located in middle of each sternite, with one seta longer than other sternal setae. Tergite and sternite 8 length shorter than lengths of tergites and sternite 1-7; margin of sternite 8 bearing many dark, stout setae, longer than length of sternite 8; ventral surface of sternite bearing several short, fine, dark setae. Margin of tergite 8 bearing shorter, fewer setae than margin of sternite 8 (setae of tergite 8 only visible upon genitalia dissection).

Terminalia (Fig. 3.37) dark reddish-brown. Middle of subrectangular epandrium lustrous; margins lightly pruinose and bearing many dark, stout, long setae, decreasing in length towards apex of epandrium. Cercus subtriangular, about as long as epandrium, and of about ½ as wide as epandrium width; cercus more densely pruinose than epandrial margins; dorsal margin of cercus lined with many fine, dark, erect, short setae. Hypandrium slim, L-shaped, with tubular stem, with tentacle-like tips hugging swollen base of phallus. Basiphallus swollen rather stout, yellow-orange, and lustrous; phallus tapers towards distiphallus; phallus forming small, U-shaped loop, sometimes hidden within epandrium; phallus extending slightly farther than length of terminalia. Ejaculatory apodeme larger than cercus; subrectangular, with lateral wings shorter than vertical wing; posterior margin of vertical wing distinctively S-shaped. Female. Hind femur with posteroventral pennate scales, and scales around middle of hind femur as long as width of femur; hind tibia with pennate scales both anterodorsally and anteroventrally, and scales shorter than width of hind tibia; three notopleural setae evenly spaced, dark, and longer than acrostichals. Geographical distribution. Nearctic distribution in Figure 3.65

Rhamphomyia hirtula Zetterstedt, 1842 Figs. 3.5, 3.6, 3.40, 3.66

Material examined. ICELAND. Myvata, sweeping west side of Vindbelgjarfjall, 1.vii.1981, 1355/1/3-500, J.A. Downes (2m#, 2f#, CNC); Myvata, Rd. to Skutustadir, 1357/2/15, 3.vii.1981, J.A. Downes (1m#, CNC); Asheidi, Kelduhverfi Nord- Pingeyiarsysia, 13/52/3/1-2, ex. with Sim. vittatum prey, O.K Nielsen & J.O. Hilmarsson (1m#, CNC). Diagnosis. Male. Cerci bifurcate such that upper margin has a prominent, stout, finger-like projection (Fig. 3.40); phallus with visible curvature below epandrium (Fig. 3.40). Apex of epandrium with cluster of fine, dark, short setae curved upwards (Fig. 3.40). Hind basitarsi and hind tarsomeres 2-4 with several rows of antero/posterodorsal setae longer than twice basitarsal and tarsomere width (Fig. 3.5). Mid tibia with at least 3 preapical antero/posteroventral setae dark, stout, spine-like. Female. Similar to male

except as follows: dorsal tarsomeres of mid and hind legs without setae longer than length of ventral tarsomere setae; middle of abdominal sternites without setae as long as setae of abdominal tergites; abdominal setae short, fine, fewer. Description. Wing length 12 mm. Male. Body densely covered with setae. Head dark in ground-colour, with greyish pruinescence on face, frons, and occiput. Holoptic, eye with ommatidia larger on upper half, smaller on lower half. Frons divergent towards antennae; face only slightly divergent towards mouthparts; oral margin dark brown and shiny. Ocellar triangle dark, subshiny, with 3 pairs of ocellar setae; anteriormost pair shorter than posterior pair, and posterior ocellar setae dark, longer than anterior and parallel. Three pairs of postocellar setae dark, shorter than posterior ocellar setae. Occiput bearing row of postocular setae, dark, long; occipital setae black, shorter and stouter; setae on postgena dark, finer than occipital setae. Postpedicel, stylus and scape dark. Scape 2× longer than pedicel; pedicel bulbous; first 1/3 of postpedicel swollen, and thinning out towards tip; postpedicel more than 5× longer than basal width; stylus as long as scape length. Palpus dark, with many long, fine, dark setulae. Labrum lustrous and reddish brown, apex black; labellum lighter brown than labrum, with dark, hair-like setae. Thorax dark, largely dark grey pruinescent. Prescutum and scutum with pair of faint, light grey vittae between acr and dc rows. Pleura dark greyish-black, often with reddish-brown shadows on katepisternum, anepisternum, and anepimeron; rarely completely dark greyish-black. Posterior corner of postpronotal lobe lustrous light brown; notopleural ridge light brown; coxal sockets light brown; region between posterior spiracle and halter base light brown to reddish-brown; supra-alar ridge with distinct light brown tint. Postalar callus brown. Proepisternum with many short, dark, hair-like setae; prosternum bare. Propleuron with cluster of fine, dark setae. Antepronotum with dense row of short, stout, dark setae. Postpronotal lobe with many short, dark setae and 1 longer, stout seta. Scutum with biserial, dense, dark, row of fine acr; dc similar to acr, except slightly longer and increasing in length posteriorly; posterior supra-alar setae dark, stout, and longer than dorsocentrals; setae on postalar callus long, stout, dark. Anterior notopleural setae dark, stout, shorter than posterior notopleural setae; posterior notopleurals stouter and longer than anterior notopleurals. Scutellum with apical scutellar setae dark, stout, and convergent; many subapical scutellar setae of subequal lengths to

each other and to apical scutellar; discal scutellar setae present and as long as subapical scutellars; basal scutellar setae present. Anterior and posterior spiracles pale light brown. Legs long, densely covered with setae, mostly reddish brown, coxae more grey than brown. Trochanter-coxa junction pale yellow-brown. Mid and hind coxae with many lateral hair-like, long, dark setae; anterior fore coxae bearing many hair-like, long, dark setae. Trochanter bearing many short, hair-like, dark setae. Femora with short white ventral pubescence; hind and mid femora with many rows of antero/posterodorsal, dorsal, anterior, antero/posteroventral, and ventral setae dark, anterodorsal and dorsal setae longer than antero/posteroventral, ventral and anterior setae, but no longer than 2x their lengths; fore femur similar, except postero/anteroventral setae 2x length of antero/posterodorsal and dorsals. Hind tibia (Fig. 3.5) with many rows of antero/posterodorsal and dorsal dark setae, longer than tibial width, but shorter than 2x tibial width, antero/posteroventral and ventral setae similar to dorsals, except decreasing in length towards proximal 1/4 and proximal ½ of anteroventral setae stouter than posteroventrals; mid and fore tibiae clothed with white pubescence, antero/posterodorsal and dorsals subequal to tibial width; mid tibia with apical antero/posteroventrals short, dark, stouter, spine-like; fore tibia (Fig. 3.6) with many rows of fine, short, posterior antero/posteroventrals and ventrals; mid tibia with many rows of posterior setae similar to antero/posterodorsals, antero/posterodorsals and dorsals stout, dark, shorter than tibial width but longer than half tibial width. Hind (Fig. 3.5) and mid basitarsi and tarsomeres 2-5 with antero/posterodorsals and dorsals more than 2x their respective widths; antero/posteroventrals subequal to their respective widths, dark, stout; preapical antero/posteroventrals of tarsomeres 2-5 2x lengths of proximal antero/posteroventrals; fore basitarsus and fore tarsomeres 2-5 with strong ventral white pubescence, with antero/posterodorsals subequal to their respective widths; fore basitarsi with finer antero/posteroventrals, stouter than antero/posterodorsals, slightly longer than basitarsal width; fore tarsomeres 2-5 without antero/posteroventrals, except preapicals stouter, dark, spine-like and subequal to respective widths. Wing clear with yellowish veins; all veins complete (except Sc), well sclerotized. Pterostigma distinct, dark yellow. Anal lobe well-developed; axillary fissure forming acute angle. Halter dark yellow.

Abdomen reddish brown with grey pruinescence; sternite and tergite margins pale. Laterals of tergites and sternites covered with long, stout dark setae; middle of tergites bearing shorter and finer setae; setae of tergite 6 short and fine; setae of tergite 7 conglomerated only in middle of tergite. Tergite 8 reduced, smaller than sternite 8 and bearing several short, dark setae. Sternite 8 margin bearing many dark, stout setae short or longer with apex curved. Terminalia (Fig. 3.40) dark reddish-brown; epandrium ovoid-shaped, with black tinge, bearing many dark, long setae; epandrium tip rounded, bearing dense cluster of hair-like, dark setae projecting slightly downwards, then outwards and slightly shorter than setae of epandrium margin; cluster of short, fine, dark, erect setae on dorsalapical margin of epandrium. Cercus and epandrium of subequal width, epandrium slightly larger than cercus; cercus bifurcate such that upper margin has prominent, stout, finger-like projection; cercus clothed with many fine, short, dark setae; posterior base of cercus with cluster of dark, fine, erect setae; cercus ventral anterior corner with distinctive cluster of short, fine, dark setae. Hypandrium slim, cylindrical, membranous, short, ending before basiphallus; basiphallus swollen, tapered to hair-like phallus; phallus curvature forming one lower, smaller lobe, and one upper, larger lobe. Ejaculatory apodeme smaller than cercus, vertical wing with dorsal margin somewhat flattened, subequal to lateral wings. Female. Similar to male except as follows: dorsal tarsomeres of mid and hind legs without setae longer than length of ventral tarsomere setae; middle of each abdominal sternites without setae as long as setae of abdominal tergites; abdominal setae short, fine, fewer. Remarks. Saigusa considers the hirtula group a subgroup of the hirticula group: halteres black, dorsal projection of cercus protruding from basal to sub-basal region of cercus. Geographic distribution. Distribution in Figure 3.66

Rhamphomyia hirticula Collin, 1937 Figs. 3.7, 3.41, 3.66

Material examined. CANADA. Manitoba: Churchill, 58.73034, -93.79509, 1- 5.vii.2010, wet, Malaise, Repl.2, 1-13.vii.2010, wet & mesic, Pan, NBP Field Party (71m#, LEMQ). Northwest Territories: Clyde, Baffin Is., 1-14.vii.1958, G.E. Shewell (31m#, 25f#, CNC). Nunavut: Ellesmere Is., Lake Hazen, 81.83179, -71.44115, 17.vii.2010, wet, Sweep, Repl.3, NBP Field Party (2m#, LEMQ); Iqaluit, 63.75117, - 68.45898, 17-21.vii.2010, wet & mesic, Malaise, Repl.3, 17-29.vii.2010, wet & mesic, Pan, Repl. 1,3, NBP Field Party (15m#, LEMQ); Iqaluit, 63.75117, -68.45898, 21- 25.vii.2010, mesic, Sweep, Repl.3, NBP Field Party (2m#, LEMQ). Yukon: Dempster Hwy., nr. N. Fork Pass, 64.58353, -138.26843, 27.vi.-1.vii.2011, mesic, Malaise, Repl.3, NBP Field Party (6m#, LEMQ). Diagnosis. Male. Mid tibia with 2 anterodorsals and 2 posterodorsals longer than twice tibial width, and stouter. Apex of epandrium with cluster of fine, dark, short setae curved upwards (Fig. 3.41). Tergite 8 with 2 lateral rounded triangular, horn-like, rounded projections. Description. Wing length 7.5 mm. Male. Head dark in ground-colour, with greyish pruinescence on face, frons and occiput. Holoptic, eye with ommatidia larger on upper half, smaller on lower half. Frons divergent towards antennae; face slightly divergent towards mouthparts; bare, with oral margin dark and shiny. Ocellar triangle dark, with 2 pairs of parallel ocellar setae: anterior and posterior pair both dark and of equal lengths; 3 pairs of dark postocellar setae shorter than ocellar setae. Occiput bearing row of postocular setae, slender and black; longer on upper section; shorter on lower section; occipital setae black, short and stout; setae on postgena longer than occipital setae. Postpedicel, stylus and scape dark and pruinose; pedicel bulbous. Scape shorter than 2x length of pedicel; postpedicel cone-like and nearly 5× longer than basal width; stylus slender and subequal to basal width of postpedicel. Palpus dark, with setulae long and dark. Labrum dark reddish brown and apex light brown; labellum pale yellowish- brown, with dark setae. Thorax dark grey in ground-colour, with dark grey pruinescence. Prescutum and scutum with pair of faint, lighter grey vittae between acr and dc rows. Pleura dark grey, often with reddish brown shadows on anepimeron, katepisternum, and anepisternum; posterior corner of postpronotal lobe lustrous and light brown; notopleural ridge light

grey; supra-alar ridge with distinct light brown tinge; region between posterior spiracle and halter base light orange-brown; anterior and posterior spiracle dark. Postalar callus light brown. Propleuron with small cluster of fine, dark setae. Proepisternum with many short, dark, hair-like setae; prosternum bare. Antepronotum with row of short, stout dark setae. Postpronotal lobe with many short, dark setae and 1 long seta located posteriorly. Scutum with biserial row of fine, short, dark acr; dc similar to acr, increasing in length posteriorly; anterior seta on postalar callus short, stout and dark, posterior seta dark, stouter and more than twice as long as anterior postalar callus seta; anterior npl cluster of setae with shorter and longer setae all stout and dark; 3 posterior npl setae as long and as stout as longer and stouter anterior npl setae. Scutellum: apical scutellars widely separated and longer than any other thoracic setae; 1 pair of subapical scutellar setae also dark and long, although shorter than apical scutellars; 2nd pair of subapical scutellars shorter and finer than other scutellars. Legs long, entirely dark reddish brown. Coxae with several dark lateral seta. Trochanters short and bearing several short, dark setae. Hind femur with antero/posteroventral setae dark, fine, shorter than femoral width on proximal 2/3, stouter, longer than femoral width on apical 1/3; mid femur with short white ventral pubescence, with posteroventrals similar to those of hind femur; fore femur with antero/posteroventral and ventral setae dark, fine, subequal to femoral width on proximal 1/3, shorter than half femoral width on apical 2/3. Hind (Fig. 3.7) and mid tibiae with antero/posterodorsals and dorsals dark, stout, longer than 2x tibial width; anteroventral setae dark, stout, increasing in length towards middle tibia; anteroventrals of apical 1/3 short, stout, dark, spine-like; anterior preapical seta dark, stout, subequal to tibial width. Fore tibia with antero/posterodorsals stout, dark, subequal to tibial length; anterior setae lighter, short, stout; antero/posteroventral and ventral setae fine, short, light-colored. Hind (Fig. 3.7) and mid basitarsi with antero/posteroventral setae of equal lengths to antero/posterodorsals, antero/posteroventrals stouter than antero/posterodorsals, 2 preapical posterodorsal setae as long as 3x basitarsal width; hind tarsomeres similar to hind basitarsus; mid basitarsus with anterior setae dark, stout, longer than basitarsal width, but shorter than 2x basitarsal width. Wing clear with yellowish veins; all veins complete (except Sc), well sclerotized;

1 basicostal seta longer and stouter than remaining costal setae. Pterostigma present as darkened yellow coloring. Anal lobe well-developed; axillary fissure rectangular, not acute. Halter: knob black, stem dark yellow. Abdomen dark reddish-brown with silvery pruinescence. Tergite and sternite margins more pale grey than brown. Abdomen covered in dark, stout setae, shorter in middle of tergites; longer, stouter on laterals of tergites and in middle of sternites. Sternite 8 lustrous; margins with greyish pruinescence. Tergite 8 shorter than sternite 8, with 2 lateral horn-like, rounded projections. Terminalia (Fig. 3.41) dark reddish-brown. Epandrium glossy in middle, flanked with silvery pruinescence, bearing many long, stout, straight, dark setae; epandrium subrectangular; tip of epandrium slender and rounded, colored with light brownish tinge, bearing cluster of densely packed hair-like, dark setae curving downwards then upwards together; clusters of setae converging towards each other such that tips of setae barely meet. Cercus shorter than length of epandrium, densely pruinose, bearing several short, dark, stout setae; cercus bifurcate; posterior dorsal margin forming slim, rounded, finger- like projection curved anteriorly; dorsal margin of cercus base bearing several dark, slim, erect setae. Cercus and epandrium of subequal widths. Hypandrium short, slender, sclerotized, triangular tip hugging base of swollen basiphallus; basiphallus lustrous, swollen; distiphallus with small S-shaped curve, color changing from light brown to darker reddish-brown; remaining phallus slender, not forming any loops, and extends beyond cercus. Ejaculatory apodeme large, fan-shaped, vertical wing forming acute angle, lateral wings shorter than vertical wing. Female. Unknown. Geographic distribution. Known distribution in Figure 3.66.

1 2

Figures 3.1-3.4. 1, 2, Rhamphomyia herschelli, posterior and lateral view of projections of pregenital segments; 3, Rhamphomyia herschelli, hind tibia, basitarsus, and tarsomeres; 4, Rhamphomyia sp.12, hind femur, and tibia.

Figure 3.5-3.7. 5, Rhamphomyia hirtula, hind leg; 6, Rhamphomyia hirtula, fore leg; 7, Rhamphomyia hirticula, hind tibia, basitarsus, and tarsomeres.

Figure 3.8-3.10. 8, Rhamphomyia (Ctenempis) calvimontis, hind leg; 9, fore leg; 10, pregenital segments (male).

11 12

Figure 3.11-3.12. 11, Rhamphomyia (Pararhamphomyia) diversipennis sp.1, mid leg; 12, Rhamphomyia (Pararhamphomyia) diversipennis sp.1, thorax, dorsal view.

13 14

15 16

Figure 3.13-3.16. 13, Rhamphomyia (Pararhamphomyia) tipularia sp. 1, hind leg, lateral view; 14, Rhamphomyia (Pararhamphomyia) tipularia sp. 4, hind leg, lateral view; 15, Rhamphomyia (Pararhamphomyia) tipularia sp. 4, male genitalia showing spindle-like structure of epandrium; 16, Rhamphomyia (Pararhamphomyia) tipularia sp. 1, fore basitarsus, lateral view.

Figure 3.17. Rhamphomyia (Pararhamphomyia ?) simplex, hind leg, lateral view.

Figure 3.18-3.19. 18, Rhamphomyia (Pararhamphomyia) pusilla sp. 7, hind leg, lateral view; 19, Rhamphomyia (Pararhamphomyia) pusilla sp. 7, mid leg, lateral view.

Figure 3.20. Rhamphomyia (Pararhamphomyia) fuscula gp. sp.1, hind leg, lateral view.

21 22

24 25

Figure 3.21-25. 21, Rhamphomyia (Dasyrhamphomyia) nigrita, hind leg, lateral view; 22, Rhamphomyia (Dasyrhamphomyia) zaitsevi, hind leg, lateral view; 23, Rhamphomyia (Dasyrhamphomyia) sp.9, hind tibia, lateral view; 24, Rhamphomyia (Dasyrhamphomyia) sp.9, mid leg, lateral view; 25, Rhamphomyia (Dasyrhamphomyia) zaitsevi, mid leg, lateral view.

Figure 3.26. Rhamphomyia (Eorhamphomyia) nigrolineata gp. sp.4, hind leg, lateral view.

27 28

29 30

Figure 3.27-3.30. 27, Rhamphomyia (Pararhamphomyia) lapponica gp. sp.8, hind leg, lateral view; 28, Rhamphomyia (Pararhamphomyia) lapponica gp. sp.9, hind leg, lateral view; 29, Rhamphomyia (Pararhamphomyia) lapponica gp. sp.11, hind leg, lateral view; 30, Rhamphomyia (Pararhamphomyia) hoeli, hind leg, lateral view.

32 33

Figure 3.31-3.33. 31, Rhamphomyia (Pararhamphomyia) filicauda, hind leg, lateral view; 32, Rhamphomyia (Pararhamphomyia) filicauda, hind femur, tibia, and basitarsus, lateral view; 33, Rhamphomyia (Pararhamphomyia) filicauda, mid leg, lateral view.

35 36

Figure 3.34-3.36. 34, Rhamphomyia (Pararhamphomyia) ursinella, hind leg, lateral view; 35, Rhamphomyia (Pararhamphomyia) caudata gp. sp.7, hind leg, lateral view; 36, Rhamphomyia (Pararhamphomyia) ursinella, fore leg, lateral view.

38 39

Figure 3.37-3.39. 37, Rhamphomyia sp.12, lateral view of male genitalia; 38, Rhamphomyia herschelli, lateral view of male genitalia (setae not shown), 39, Rhamphomyia herschelli, lateral view of pregenital segments 7 and 8.

Figure 3.40-3.41. 40, Rhamphomyia hirtula, lateral view of male genitalia; 41, Rhamphomyia hirticula, lateral view of male genitalia.

43 Figure 3.42-3.43. 42, Rhamphomyia (Ctenempis) calvimontis, lateral view of male genitalia; 43, Rhamphomyia (Ctenempis) calvimontis, pregenital segments (male).

Figure 3.44. Rhamphomyia (Pararhamphomyia) diversipennis sp.1, male genitalia, lateral view.

Figure 3.45-3.46. 45, Rhamphomyia (Pararhamphomyia) tipularia gp. sp. 1, male genitalia, lateral view; 46, Rhamphomyia (Pararhamphomyia) tipularia gp. sp. 4, male genitalia lateral view.

47 48

Figure 3.47-3.49. 47. Rhamphomyia (Pararhamphomyia ?) simplex, male genitalia, lateral view. 48. Rhamphomyia (Pararhamphomyia) pusilla sp.7, male genitalia, lateral view. 49. Rhamphomyia (Pararhamphomyia) fuscula gp. sp.1, male genitalia, lateral view. 50 51 52

Figure 3.50-3.52. 50, Rhamphomyia (Dasyrhamphomyia) nigrita, male genitalia, dorsal view; 51, Rhamphomyia (Dasyrhamphomyia) zaitsevi, male genitalia, dorsal view; 52, Rhamphomyia (Dasyrhamphomyia) sp.9, male genitalia, dorsal view..

Figure 3.53-3.55. 53, Rhamphomyia (Dasyrhamphomyia) nigrita, male genitalia, lateral view; 54, Rhamphomyia (Dasyrhamphomyia) sp.9, male genitalia, lateral view; 55, Rhamphomyia (Dasyrhamphomyia) zaitsevi, male genitalia, lateral view.

Figure 3.56-3.57. 56, Rhamphomyia (Eorhamphomyia) nigrolineata gp. sp.4, male genitalia, lateral view; 57, Rhamphomyia (Eorhamphomyia) nigrolineata gp. sp.4, cerci and phallus tip, dorsal view.

Figure 3.58-3.61. 58, Rhamphomyia (Pararhamphomyia) lapponica gp. sp.8, male genitalia, lateral view; 59, Rhamphomyia (Pararhamphomyia) lapponica gp. sp.9, male genitalia, lateral view; 60, Rhamphomyia (Pararhamphomyia) lapponica gp. sp.11, male genitalia; 61, Rhamphomyia (Pararhamphomyia) hoeli, male genitalia, lateral view. 62

Figure 3.62-3.64. 62, Rhamphomyia (Pararhamphomyia) filicauda, male genitalia, lateral view; 63, Rhamphomyia (Pararhamphomyia) ursinella, male genitalia, lateral view; 64, Rhamphomyia (Pararhamphomyia) caudata gp. sp.7. Figure 3.65. Rhamphomyia sp.12 (circle), Rhamphomyia herschelli (triangle), distributions.

Figure 3.66. Rhamphomyia hirticula (circle), Rhamphomyia hirtula (triangle), distributions.

Figure 3.67. Rhamphomyia (Ctenempis) calvimontis, distribution.

Figure 3.68. Rhamphomyia (Pararhamphomyia) diversipennis sp.1, distribution. Figure 3.69. Rhamphomyia (Pararhamphomyia) tipularia sp.1 (circle), Rhamphomyia (Pararhamphomyia) tipularia sp.4 (square), distributions.

Figure 3.70. Rhamphomyia (Pararhamphomyia) simplex, distribution. Figure 3.71. Rhamphomyia (Pararhamphomyia) pusilla gp. sp.7, distribution.

Figure 3.72. Rhamphomyia (Pararhamphomyia) fuscula gp. sp.1, distribution. Figure 3.73. Rhamphomyia (Dasyrhamphomyia) nigrita (circle), Rhamphomyia (Dasyrhamphomyia) sp.9 (square), Rhamphomyia (Dasyrhamphomyia) zaitsevi (triangle), distributions.

Figure 3.74. Rhamphomyia (Eorhramphomyia) nigrolineata gp. sp.4, distribution. Figure 3.75. Rhamphomyia (Pararhamphomyia) lapponica gp. sp.8 (circle), Rhamphomyia (Pararhamphomyia) lapponica gp. sp.9 (square), Rhamphomyia (Pararhamphomyia) lapponica gp. sp.11 (triangle), Rhamphomyia (Pararhamphomyia) hoeli (hexagon), distributions.

Figure 3.76. Rhamphomyia (Pararhamphomyia) ursinella (circle), Rhamphomyia (Pararhamphomyia) caudata gp. sp.7 (triangle), Rhamphomyia (Pararhamphomyia) filicauda (square), distributions. Connecting Statement In the process of conducting the species-level inventory in Chapter 2, there was an emerging pattern that more northerly specimens of Rhamphomyia were generally larger than specimens from southern sites. Body size gradients across latitude is the subject of many studies because body size has a direct impact on every aspect of an arthropod’s life. Such studies have found conflicting results, and the relationship between latitude and arthropods remains unclear. To gain a better understanding of the Rhamphomyia body size pattern across the Nearctic, we analyze body size of Rhamphomyia across the latitudinal gradient of the Northern Biodiversity Program sampling sites.

Chapter 4: Latitudinal body size patterns in a diverse genus of arctic insects

Introduction Body size has a direct impact on every aspect of an organism’s life (Lawton, 1990) such as its relationship with the environment, and the pace of its physiological processes (Cushman et al., 1993; Entling et al., 2010). Functional traits help ecologists better understand the dynamics of ecological communities, which in turn, help gain insight in species turnover resulting from environmental shifts (McGill et al., 2006; Ho et al., 2010). To date, the most widely cited pattern for broad-scale invertebrate body size distribution patterns is Bergmann’s Rule (Entling et al., 2010; Shelomi, 2012). Bergmann’s rule argues that closely related homeotherms, and specifically, endotherms become increasingly larger towards colder environments (Blackburn et al., 1999; Homburg et al., 2013). Bergmann (1847) suggested that this trend exists due to larger individuals having smaller surface-to-volume ratios, therefore increasing heat conservation. However, Bergmann’s rule and its predictions initially only pertained to body size studies of birds and mammals (Blackburn et al., 1999). Bergmann’s rule does not formulate predictions for ectotherms, and only considered one process: colder climates versus warmer climates (Blackburn et al., 1999). Bergmann did not equated colder climates to latitude increase (Cushman et al., 1993; Blackburn et al., 1999). In fact, there is a generally accepted substitute of Bergmann’s original assertion about body size differences between colder and warmer climates with assertions about body size differences across latitude (Entling et al., 2010). In the Canadian Arctic, arthropods are the dominant terrestrial animals. As latitude increases, arthropod diversity decreases, but Diptera become increasingly dominant as other Orders decline. One of the most abundant and species-rich Nearctic families of terrestrial Diptera are the dance flies (Empididae). The empidid genus Rhamphomyia Meigen appears to be one of the most species-rich and species-abundant Diptera genera in the Arctic. In fact, the terrestrial Diptera assemblages of the arctic islands are mostly composed of Rhamphomyia species. Rhamphomyia species are

important Arctic pollinators and predators, and show a wide variation in body size among species, ranging from 3-13 mm long (T. Saigusa, pers. comm). Recent study of a large species assemblage of Rhamphomyia across a latitudinal gradient in northern Canada suggested the body size of Rhamphomyia specimens collected from the high arctic is, in general, larger than specimens collected in subarctic or boreal sites. Most studies of Bergmann’s rule have been focused at the single species-level (Shelomi, 2012). Because of the high species-richness of Rhamphomyia at the same sites in northern Canada, we chose to examine body size patterns at the species level of the entire assemblage rather than at the level of individuals within a species. To assess the relationship of northern Nearctic Rhamphomyia body size patterns, we examined latitudinal and other spatial effects to determine whether the spatial patterns of Rhamphomyia body size in the Nearctic are driven by temperature decreases corresponding to latitude increase, or by other ecological factors. This chapter examines whether the body size of Rhamphomyia increases along with latitude increase, and if such a pattern does occur, whether this pattern occurs within or among species. We then discuss the possible drivers of the observed body size pattern.

Materials and Methods

Sample sites and collecting Specimens were collected from twelve sampling localities across northern Canada: 4 boreal sites (Moosonee, ON, Goose Bay, NL, Yellowknife, NT, Norman Wells, NT), 4 low arctic mainland sites (Schefferville, QC, Churchill, MB, Dempster Hwy, YT, Kugluktuk, NU), and 4 high arctic island sites (Iqaluit, NU, Cambridge Bay, NU, Banks Island, NT, Ellesmere Island, NU). Standardized, replicated sampling was used in 2010 (eastern sites), and in 2011 (western sites), using Malaise traps, yellow pan traps, pitfall traps, and sweeping. Sampling grids were set up in each of 3 wet and 3 mesic plots per sampling locality, for a total of 72 sampling grids. We collected at each site for about two weeks during the summer peak of arthropod activity, at 4 day intervals. Kugluktuk was sampled during the entire summer season 2010 and for two weeks in 2011. Because of

poor weather during the 2011 sampling period at Kugluktuk, a two-week subsample of the 2010 sampling season was selected from the same time period and used in analyses . Goose Bay, Yellowknife, Norman Wells, and Ellesmere Island were omitted because of insufficient data (i.e. five or fewer specimens of each species). Thus, analyses are based on the remaining eight localities. However, for the statistical analyses of spatial effects on body size other than latitude, all twelve sampling localities were included. The six sampling grids within each site were generally within 2 km of one another and latitudinal analyses were based on a single mean value of the latitudes of all grids within each site.

Specimen identification All Empididae were extracted from the samples and identified to genus. Rhamphomyia specimens were sorted by sex and analyses based on male specimens only because females are often impossible to identify to species, and because females and males are significantly different in size within a species. Male specimens were sorted to morphologically distinct morphospecies because most Nearctic Rhamphomyia are undescribed. Species-level resolution was necessary because of size differences between species. Male Rhamphomyia are easily sorted to morphospecies because genitalia are species-specific and distinctly modified (Downes, 1970). Chaetotaxy and leg ornamentation were also useful to distinguish morphospecies. When external morphology was not sufficient, genitalia were dissected to examine hidden structures, especially aedeagus curvature and lamellae lobes. Reference material in the Canadian National Collection of Insects, Arachnids and Nematodes (Ottawa, Ontario) (CNC), published keys to some subsets of species (e.g., Barták, 2002) and unpublished notes by empidid taxonomist T. Saigusa (deposited in CNC), allowed several species to be named or assigned to species groups. DNA barcoding of 1–4 specimens of each Rhamphomyia male morphospecies was used to supplement and confirm morphospecies identification. DNA barcoding, based on the standard 658 bp region of the mitochondrial CO1 gene, was carried out at the Canadian Center for DNA Barcoding (University of Guelph) .

Specimen measurement Fore tibia length as proxy for body size. To test the validity of fore tibia as a proxy for full body size, five undamaged specimens of each species represented by five or more specimens in each of the eight sampling localities were selected for measurement. Fore tibia length and full body size (measured from antennal base to tip of abdomen, excluding genitalia) were measured using an eyepiece micrometer on a stereomicroscope (at 43.2X magnification). For consistency, all measurements were taken by one individual (E A.V.) using the same microscope. We used a linear regression, in which body size was a function of tibia length, to test the validity of fore tibia length as a proxy for body size. This linear model was run on RStudio (R Core Team, 2014) using the function lm in the stats package. A total of 181 specimens were used for this test. We obtained a significant positive correlation between tibia length and full body size (β=; p<0.0001; Adj. R2=0.8843; Figure 4.1).

Latitudinal variation of body size. To test for latitudinal effects, we included tibia lengths of all males for all species in each of the eight sampling localities (1301 specimens). However, because one species had over 1000 specimens, 122 randomly selected specimens were measured. Tibia lengths were measured using a scope magnification of 68.4X. All measured species were considered, including species only represented by a single specimen. For species collected at four or more sampling sites, we plotted the mean body size against the latitude point for each site for each of these species.

Environmental data We extracted climate normals computed by using 15 years of data recorded from 1981 to 2010 at the nearest weather station to each of the twelve sampling localities to acquire mean annual temperature (ºC), mean annual precipitation (mm), and mean growing degree days above 5ºC (Meteorological Service of Canada, 2012). To test whether these three factors were significantly correlated with latitude increase, we ran a linear regression, in which mean temperature, mean annual precipitation, or mean annual growing degree days were a function of mean latitude.

Analyses We determined if the mean annual temperature significantly decreases as latitude increases. To test for latitudinal effects, we assessed whether the mean tibia length of the Rhamphomyia assemblage changes with latitude and whether the mean tibia length of the assemblage is a function of community composition. To address other spatial affects, we assessed whether mean tibia length differs between sampling localities and if there a difference in the mean tibia length between the subarctic mainland and the high arctic islands? Finally, we determined if the increase in latitude significantly corresponds to a decrease in mean annual precipitation, and mean annual growing degree days.

Latitudinal effects. To examine whether there was a trend across latitude of mean tibia length, we ran a linear mixed-effects model in R using the function lme from the nlme package (Pinheiro et al., 2014). Sampling locality was included as a random effect. We also tested whether species identity had an effect on body size by running the linear mixed-effects model where tibia length is a function of species and where species identity is a fixed-effect. We modeled tibia length (the response variable) as a function of latitude. To avoid violating the linear model assumption that all observations are independent, we specify the sampling localities as our grouping structure (i.e. the random effect). We ran this model a second time, adding another “fixed-effect”, species, while site remained as a random effect. By doing so, tibia length became a function of latitude and of species.

Other spatial effects. To determine if mean tibia length significantly differed between sampling localities, we tested an ANOVA model, where tibia length was the response variable and sampling locality was the explanatory variable. We then conducted a post- hoc Tukey’s Honest Significance Difference (HSD) test to discern which pairs of sampling localities showed a significant difference in mean tibia length. This next analysis comprised all species collected, including species represented by less than 10 specimens, and all twelve sampling localities. We assigned the sampling localities to “mainland” or “island”. Latitude points were not considered in this analysis as we were attempting to compare distributions between the mainland and the islands:

Iqaluit is at a lower latitude than Kugluktuk, even though Kugluktuk is on the mainland (i.e. subarctic), and Iqaluit is on the Arctic islands (i.e. high arctic). We ran a linear mixed-effects model, with tibia length as the response variable, the grouping (mainland versus island) as the fixed-effect, and sampling sites as the random effect.

Latitude and other ecological factors. To test whether mean annual temperature, mean annual precipitation, and mean annual growing degree days are significantly correlated with latitude increase, we then ran a linear regression, in which mean temperature, mean annual precipitation, or mean annual growing degree days were a function of mean latitude

Results

Diversity of Rhamphomyia A total of 3733 Rhamphomyia male specimens were sorted to 65 morphospecies. The number of species per sampling locality varied greatly. Banks Island and Ellesmere Island had the lowest species richness (3 species); Dempster Highway had the highest species richness (26 species). The second most species-rich sampling locality was Kugluktuk (18 species). Abundance was lowest at Ellesmere Island, with 10 specimens collected, and highest at Dempster Highway, with a total of 385 collected specimens. Species did not frequently occur across all three biomes. Species were usually restricted to either the boreal and low arctic biomes, or to the low arctic and high arctic biomes. More rarely, species might be restricted to the boreal biome, or to the high arctic biome. The most abundant and widespread species was R. (Pararhamphomyia) fuscula gp. sp.1 which was collected in boreal (Norman Wells), low arctic (Dempster Hwy, Kugluktuk), and high arctic (Iqaluit, Ellesmere Island) sites.

Latitudinal trends Six species could not be use for the body size analyses because the specimens were damaged and could not be measured. Therefore, the body size analyses includes 59 species. Mean annual temperature, along with mean annual precipitation and mean growing degree days significantly decreased as latitude increases (p<0.001, Adj- R2=0.8674; p=0.0002, Adj-R2=0.7272; p=0.004, Adj-R2=0.5366, respectively). These correlations are important to consider in arthropod body size studies because temperature, precipitation, and growing degree days are factors that are known to affect arthropod body size (Strathdee & Bale, 1998; Gray & Bradley, 2005). A significant increase in mean tibia length with latitude increase was confirmed using a linear mixed effects model (β=0.0293107; p=0.0054) (Figure 4.2). When species identity was included in the mixed effects model, latitude was no longer significantly correlated with tibia length (β= 0.0027; p=0.2658), demonstrating that the trend in body size increase is not intraspecific (Appendix 4.1). Species identity was significantly correlated with tibia length (p<0.001). There was a significant correlation of sampling locality with tibia length (β=119.3; p<2e-16) where body size differences significantly differed among sites. Moreover, there was a significant difference in the tibia length distribution of mainland (sub-arctic) versus islands (high arctic) (β=-0.3452485; p-value=0.0048) (Figure 4.3), in which specimens from the high arctic were significantly larger than specimens from the low arctic.

Discussion The sorting of Rhamphomyia based on their key morphological traits reveals a Nearctic Rhamphomyia species richness much higher than previously recorded (Downes, 1970; Danks, 1981). Concurrently, our findings showed a great variation in body size within Rhamphomyia, in that specimens collected in the high arctic were larger than in the subarctic. Temperature decrease was significantly correlated with latitudinal increase, and hence, according to Bergmann’s premise, we would expect Rhamphomyia to increase in

body size intraspecifically moving northward from Moosonee to Ellesmere Island. Yet, the latitudinal increase in Rhamphomyia body size (Figure 4.2) was due to a shift in the community composition, not body size increase within species, as latitude increases: smaller species drop out of the community assemblages as latitude increases. Accordingly, the trend observed is interspecific, not intraspecific. We found a significant increase in mean tibia length between the sub-arctic specimens and the high arctic specimens. In addition to mean annual temperature, mean annual precipitation and mean growing degree days also significantly decrease as latitude increases. The body size distribution pattern of the Nearctic Rhamphomyia might be the result of a combination of environmental factors such as the salt water barrier separating the mainland from the arctic islands, temperature, precipitation and growing degree days. The Northwest Passage separating the Canadian Arctic Archipelago from the Canadian mainland might be a physical barrier hindering smaller Rhamphomyia species from colonizing the islands post-glaciation. Bigger Diptera may be stronger flyers than smaller Diptera because larger flies have a more substantial energy reserve to power flight activity, and hence, larger flies can endure longer travel time (Coulson et al., 2002). Based on this premise, larger Rhamphomyia may have been more capable of successfully crossing the salt-water barrier to the islands. The dryness of the air masses in the high arctic (Maxwell, 1981; Coulson et al., 2002) might also contribute to this observed body size distribution. Larger insects are more desiccation-resistant than smaller insects, because a larger body enables a higher initial water content than a smaller body (Chown, 1993; Renault & Coray, 2004; Gray & Bradley, 2005). Smaller arthropods have a lower initial water content, and therefore, must feed more often to avoid fatal desiccation (Renault & Coray, 2004; Gray & Bradley, 2005). Consequently, the long dispersal distance across the Northwest Passage is likely to favor desiccation-resistant arthropods (i.e. larger arthropods) (Coulson et al., 2002). In the event of smaller-bodied Rhamphomyia species surviving dispersal across the Northwest Passage, by, for instance, riding the wind currents, the next challenge is successful establishment and reproduction in the high arctic (Coulson et al., 2002). The Canadian Archipelago is the driest region in Canada (Maxwell, 1981). Small Rhamphomyia species in the high arctic would arguably have to feed frequently to

replenish their body water content. However, energy resources are scarce and sporadic in the high arctic (Strathdee & Bale, 1998). In addition, larger arthropods have a higher fasting-endurance because energy reserves grow with body size faster than metabolic rate (Cushman et al., 1993). Thus larger Rhamphomyia are more likely than smaller Rhamphomyia to survive sudden drops in food supplies (Cushman et al., 1993). There is a possibility that the observed body size pattern is a result of island gigantism through founder effects caused by mainland Rhamphomyia migrating to the arctic islands (Lomolino, 2005). However, if this were so, there would be intraspecific body size increase from the mainland to the islands (i.e. low arctic to high arctic), and species such as R. nigrita, spanning low arctic and high arctic sites, would be significantly larger at their high arctic sites than at their low arctic sites. This was not the case (Appendix 4.1). Instead, species spanning low arctic and high arctic sites show no particular trend in body size (Appendix 4.1). Another potential explanation is that larger Rhamphomyia in the arctic islands are a result of reduced resource competition in the high arctic (Palkovacs, 2003; Lomolino, 1985). However, resource competition may not be a strong factor affecting insect body size in the arctic, especially with regards to generalist predators, such as Rhamphomyia. The arctic has a substantial biomass, and prey items like chironomids and mosquitos do not appear to significantly fluctuate across latitude. These results, together with the results of other arthropod body size studies (Blackburn et al., 1999; Ho et al., 2010; Pincheira-Donoso, 2010), emphasize four points regarding studies of arthropod body size distribution patterns: 1. the relevance of examining more than a single environmental factor that may affect arthropod body size; 2. the necessity of knowledge of the biology, life-history, and survival needs of the taxa in question; 3. the pertinence of studying how different environmental factors may interact with the survival needs of the taxa in question; and 4. the suitability of the body part used as a proxy for body size.

Figure 4.1. Relationship between fore tibia length and overall body size in male Rhamphomyia from the Canadian Arctic.

Figure 4.2. Relationship between Nearctic Rhamphomyia species mean tibia lengths and sampling locality latitude point.

Figure 4.3. Box plots of mean tibia length, at the assemblage level, of Rhamphomyia for each of the twelve sampling localities.

Chapter 5. General Discussion and Conclusions

The inventory of Rhamphomyia species established for this project demonstrates that the genus is far more species-rich across the Nearctic than previously recorded (Downes, 1970; Danks, 1981). The inventory also sheds light on the poorly known spatial distribution of Rhamphomyia: (1) same species rarely span all three biomes; most species occur in the low arctic biome and/or across the boreal and low arctic biomes, and substantially fewer species are found in the high arctic biome, (2) there are few common species and many rarer species, (3) Dempster Highway has the highest species-richness in addition to the highest number of unique species to its site; likely due to the persistence of species in Beringia during the last glaciation, and (4) the high arctic has the highest species-abundance, therefore indicating the critical role Rhamphomyia play in the high arctic insect biomass as pollinators, predators, and prey. The first key to the species of the Canadian Archipelago, Greenland, and Iceland is presented, along with preliminary diagnoses and descriptions, of the species, most of which are as yet undescribed. Because the key is based on males only, emphasis is placed on genitalia and leg characters, since these are the morphological traits which most vary interspecifically in male Rhamphomyia. We are currently continuing to determine which species are new and previously described. The body size study across the latitudinal gradient shows that Rhamphomyia body size increases at species-level at higher latitudes; smaller species gradually drop out of the assemblages as latitude increases. Moreover, Rhamphomyia species of the arctic islands are significantly larger than Rhamphomyia species of the mainland. We suggest two plausible reasons to explain this discrepancy: (1) bigger Rhamphomyia are stronger fliers than smaller Rhamphomyia, and can therefore successfully cross the Northwest Passage to colonize the arctic archipelago post-glaciation, and (2) bigger Rhamphomyia have a higher fasting-endurance, and are more desiccation-resistant than smaller Rhamphomyia, and hence, can endure the extreme environmental conditions of the arctic islands. The results of the body size study, in consideration with the results brought by other arthropod body size studies (Blackburn et al., 1999; Ho et al., 2010; Pincheira- Donoso, 2010; Ernst & Buddle, 2015), emphasize the following four points regarding

studies of arthropod body size distribution patterns: (1) the relevance to examine more than a single environmental factor that may affect arthropod body size across latitude, (2) the necessity of in-depth knowledge of the biology, life-history, and survival needs of the arthropod taxa in question, (3) the pertinence of studying how different environmental factors may interact with the survival needs of the arthropod taxa in question, and (4) the body part that is being measured as a proxy for body size. This project serves as a first step, and a foundation, to address other questions about northern Nearctic Rhamphomyia. For instance, the inventory could help with examining temporal patterns, such as the emergence and seasonal activity patterns of Rhamphomyia during the short arctic activity season. Danks (1981) proposed that Rhamphomyia species emerge simultaneously at the same point in time. However, our preliminary results suggest that instead, emergence times for different Rhamphomyia species differ. However, more data needs to be collected in order to conduct an accurate study about Rhamphomyia emergence times. The high species-richness documented in an inventory based on only 12 sample sites in the Canadian north suggests that the actual species-richness of northern Nearctic Rhamphomyia is still much higher than documented to date. Although it would be an ambitious project, these species should be described and a key to the Nearctic species of Rhamphomyia should be written to facilitate identification of species. This essential taxonomic work will enable authors to connect research done on Nearctic Rhamphomyia with that done on Rhamphomyia from other geographical regions, because species-level resolution, as opposed to morphospecies codes, is necessary for comparative work across projects. Finally, the overarching objective of this project was to emphasize the undeniable link between the fields of arthropod taxonomy and arthropod ecology. Had the Rhamphomyia specimens in this project been sorted only to genus-level, as is commonly done in arthropod biodiversity studies, there would be no story to tell, and Chapter 2 would have been nothing more than a map of the Canadian Arctic speckled with dots indicating the presence of Rhamphomyia.

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"+%"+5 "%&$)* 75+6:847 ,249+4211: % 35 ( 3122 38 ( 3122 )& 3 47 - . "+%"+5 "%&$)* 75+6:847 ,249+4211: % 35 ( 3122 38 ( 3122 )& 3 7 - . "+%"+5 "%&$)* 75+6::26 ,249+41722 % 35 ( 3122 38 ( 3122 % 3 2 - . "+%"+5 "%&$)* 75+6:847 ,249+4211: 32 ( 3122 35 ( 3122 )& 3 2 - . "+%"+5 "%&$)* 75+6:847 ,249+4211: )" 38 ( 3122 38 ( 3122 )& 3 2 - . "+%"+5 !$ % 76+3:223 ,237+73373 % 25 ( 3122 28 ( 3122 )& 2 2 - . "+%"+5 !$ % 76+3:223 ,237+73373 )" 25 ( 3122 25 ( 3122 )& 2 2 3:4 - . "+%"+2 "%&$)* 75+6:847 ,249+4211: % 35 ( 3122 38 ( 3122 )& 3 2 - . "+%"+2 #'& 74+86233 ,79+56:38 % 32 ( 3121 36 ( 3121 )& 4 2 - . "+%"+2 #'& 74+86228 ,79+569:9 28 ( 3121 32 ( 3121 % 4 2 - . "+%"+2 ''&' 78+89574 ,226+38:8: % 36 ( 3121 3: ( 3121 % 2 2 - . "+%"+2 ''&' 78+94649 ,226+31:98 % 4 ( 3121 8 ( 3121 % 3 2 - . "+%"+2 ''&' 78+88547 ,226+41843 % 8 ( 3121 23 ( 3121 % 4 2 - . "+%"+2 ''&' 78+8874: ,226+41993 % 8 ( 3121 23 ( 3121 )& 4 2 - . "+%"+2 ''&' 78+89574 ,226+38:8: % 24 ( 3121 2: ( 3121 % 2 4 - . "+%"+2 ''&' 78+89574 ,226+38:8: % 5 ( 3121 21 ( 3121 % 2 2 - . "+%"+2 !) 73+61825 ,224+4:554 % 26 ( 3122 29 ( 3122 % 4 2 22 - . %% 84+33292 ,22:+6716: % 8 ( 3122 22 ( 3122 % 2 2 2 %"+ !$ % 76+3:315 ,237+74836 % 25 ( 3122 28 ( 3122 % 2 2 2 %"+ '$ 69+84462 ,:4+8:8:3 % 6 ( 3121 : ( 3121 % 2 4 %"+ "%&$)* 75+6:847 ,249+4211: % 35 ( 3122 38 ( 3122 )& 3 3 %"+ "%&$)* 75+68:53 ,249+39323 % 38 ( 3122 2 ( 3122 )& 4 2 7 %"+ "%&$)* 75+7173: ,249+46748 % 32 ( 3122 35 ( 3122 )& 2 29 %"+ "%&$)* 75+6::26 ,249+41722 % 35 ( 3122 38 ( 3122 % 3 3 %"+ "%&$)* 75+69464 ,249+37954 % 38 ( 3122 2 ( 3122 % 4 27 %"+ "%&$)* 75+68:53 ,249+39323 % 38 ( 3122 2 ( 3122 )& 4 2 %"+ ''&' 78+944:7 ,226+3248 )" 4 ( 3122 4 ( 3122 )& 3 2 %"+ !$ % 76+377:5 ,237+83938 % 22 ( 3122 25 ( 3122 % 3 3 51 %"+ !!%! 62+35733 ,91+78392 % 29 ( 3121 32 ( 3121 % 2 22 %"+ !!%! 62+35577 ,91+78878 % 32 ( 3121 35 ( 3121 % 3 26 %"+ !!%! 65+96373 ,77+78878 34 ( 3121 37 ( 3121 % 3 2 38 %"+ !!%* 64+3273 ,71+56112 % 26 ( 3121 29 ( 3121 )& 4 2 %"+ !!%* 64+3273 ,71+56112 27 ( 3121 29 ( 3121 )& 4 2 &$, #%!" & 65,30494 -126,70873 *$ 11 ) 2011 11 ) 2011 !& 3 3 &$, #%!" & 65,30494 -126,70873 *$ 14 ) 2011 14 ) 2011 !& 2 2 &$, %) 54,8535 -66,67043 & 30 ) 2010 3 ) 2010 *' 3 9 &$, %) 54,85262 -66,66879 & 3 ) 2010 6 ) 2010 *' 2 6 22 &$, #%!" & 65,30494 -126,70873 *$ 14 ) 2011 14 ) 2011 !& 3 2 2 &$, !$&'% *+ 64,59736 -138,31009 & 24 ) 2011 27 ) 2011 *' 2 2 &$, %) 54,90526 -67,15049 & 6 ) 2010 9 ) 2010 !& 1 1 3 &$, %) 54,8535 -66,67043 & 30 ) 2010 3 ) 2010 *' 3 1 &$, %) 54,7597 -66,7112 & 6 ) 2010 9 ) 2010 *' 1 2 &$, %) 54,7597 -66,7112 & 6 ) 2010 9 ) 2010 *' 1 2 5 &$, !$&'% *+ 64,60629 -138,35637 & 21 ) 2011 24 ) 2011 *' 1 31 &$, !$&'% *+ 64,57942 -138,28212 & 27 ) 2011 1 ) 2011 *' 3 1 &$, !$&'% *+ 64,58353 -138,26843 & 27 ) 2011 1 ) 2011 !& 3 1 &$, !$&'% *+ 64,605 -138,36143 & 21 ) 2011 24 ) 2011 !& 1 8 &$, !$&'% *+ 64,60629 -138,35637 *$ 27 ) 2011 27 ) 2011 *' 1 1 &$, ##&#" 51,24466 -80,67767 & 21 ) 2010 24 ) 2010 !& 2 1 &$, #%!" & 65,28901 -126,84077 & 7 ) 2011 11 ) 2011 *' 3 1 &$, #%!" & 65,25226 -126,66128 & 11 ) 2011 14 ) 2011 *' 2 6 &$, #%!" & 65,29204 -126,63725 *$ 14 ) 2011 14 ) 2011 !& 1 1 &$, #* " 62,5107 -113,39479 & 7 ) 2011 11 ) 2011 !& 1 2 &$, #* " 62,51591 -113,39146 *$ 10 ) 2011 10 ) 2011 *' 3 1 54 &$, #%!" & 65,30494 -126,70873 " 7 ) 2011 11 ) 2011 !& 3 4 &$, #%!" & 65,30494 -126,70873 " 11 ) 2011 14 ) 2011 !& 3 3 &$, #%!" & 65,30494 -126,70873 " 14 ) 2011 17 ) 2011 !& 3 4 11 &$, %) 54,90526 -67,15049 & 6 ) 2010 9 ) 2010 !& 1 38 &$, %) 54,7597 -66,7112 & 6 ) 2010 9 ) 2010 *' 1 3 41 &$, ##&+ 53,21199 -60,45052 & 21 ) 2010 24 ) 2010 *' 1 2 2 &$, (% 58,73351 -93,79792 & 5 ) 2010 9 ) 2010 !& 1 1 1 &$, ( ( '( 67,77639 -115,30882 & 7 ) 2010 12 ) 2010 *' 3 7 &$, ( ( '( 67,77639 -115,30882 & 14 ) 2010 19 ) 2010 *' 3 1 &$, ( ( '( 67,77639 -115,30882 & 26 ) 2010 2 ) 2010 *' 3 4 12 &$, (% 58,73573 -93,79789 & 5 ) 2010 9 ) 2010 *' 1 1 &$, ( ( '( 67,78508 -115,27573 & 25 ) 2010 29 ) 2010 *' 1 4 & ""!" 01&2--3+ '++/&,+.+ - # ,*+* 1 # ,*+* $! , +. & ""!" 01&2-/-2 '++/&,*321 - # ,*+* 1 # ,*+* , ++ & ""!" 01&11.-0 '++/&-*1-, 1 # ,*+* +, # ,*+* - , & ""!" 01&110-3 '++/&-*22, 1 # ,*+* +, # ,*+* $! - + -- & " /2&1-/1- '3-&13123 / # ,*+* 3 # ,*+* $! + . & " /2&0/1-/ '3-&2-+*. + # ,*+* / # ,*+* $! - + & " /2&1-/1- '3-&13123 $ / # ,*+* / # ,*+* $! + / & " /2&0/1-/ '3-&2-+*. $ 3 # ,*+* 3 # ,*+* $! - 1 & ""!" 01&110-3 '++/&-*22, 1 # ,*+* +, # ,*+* $! - +0 & ""!" 01&110-3 '++/&-*22, - # ,*+* 1 # ,*+* $! - + & ""!" 01&110-3 '++/&-*22, 2 # ,*+* +. # ,*+* $! - 2 & ""!" 01&110-3 '++/&-*22, +. # ,*+* +3 # ,*+* $! - + & ""!" 01&110-3 '++/&-*22, ,* # ,*+* ,0 # ,*+* $! - + & ""!" 01&2--3+ '++/&,+.+ ,3 # ,*+* . # ,*+* $! , + & ""!" 01&2-.,1 '++/&,*23/ $ -* # ,*++ -* # ,*++ $! - + & ""!" 01&2--30 '++/&,+-1 $ - # ,*++ - # ,*++ $! , , & $ 0,&/+/3+ '++-&-3+.0 +/ # ,*++ +2 # ,*++ $! - + .3 & ""!" 01&12/*2 '++/&,1/1- ,/ # ,*+* ,3 # ,*+* $! + + & ""!" 01&2--3+ '++/&,+.+ - # ,*+* 1 # ,*+* $! , , & ""!" 01&2-/-2 '++/&,*321 - # ,*+* 1 # ,*+* , + & ""!" 01&2--3+ '++/&,+.+ +3 # ,*+* ,0 # ,*+* $! , + / & ""!" 01&2-/-2 '++/&,*321 - # ,*+* 1 # ,*+* , + & ""!" 01&12.0- '++/&,1313 +- # ,*+* +3 # ,*+* + + & ""!" 01&2--3+ '++/&,+.+ +3 # ,*+* ,0 # ,*+* $! , + - & " /2&1-,-/ '3-&13*,2 + # ,*+* / # ,*+* 1 +1 & " /2&1-*-. '3-&13/*3 + # ,*+* / # ,*+* $! , , & " /2&1--/+ '3-&1313, / # ,*+* 3 # ,*+* + ,, & " /2&1-/1- '3-&13123 / # ,*+* 3 # ,*+* $! + - & " /2&1--/+ '3-&1313, $ 2 # ,*+* 2 # ,*+* + + & ! $% 0.&/31-0 '+-2&-+**3 ,. # ,*++ ,1 # ,*++ $! , + & ! $% 0.&/33+/ '+-2&-*0++ ,. # ,*++ ,1 # ,*++ , + .1 & 0/&,003. '+,0&1,2,1 ++ # ,*++ +. # ,*++ , + & $ 0,&/+/3+ '++-&-3+.0 +/ # ,*++ +2 # ,*++ $! - + )% ($(,$ $$& &,$') $( %#$$ $+ %#$$ & $ & )' (,*&) $&+ &$##, %' %#$$ %* %#$$ % $ $ (& &$+, )# %,(,' $+ %#$# %$ %#$# $ $ $ $#%,+ Appendix 2. Mean body size at each site of species spanning four or more sites; a. R. nigrita; b. R. lapponica sp.2; c. R. dispar gp. sp.10; d. R. rivalis; e. R. nigrolineata; f. R. fuscula gp. sp.1; g. R. hirticula gp. sp.2.