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Hemiptera of Canada
expand article infoRobert G. Foottit, H. Eric L. Maw, Joel H. Kits, Geoffrey G. E. Scudder§
‡ Agriculture and Agri-Food Canada, Ottawa, Canada
§ University of British Columbia, Vancouver, Canada
Open Access

Abstract

The Canadian Hemiptera (Sternorrhyncha, Auchenorrhyncha, and Heteroptera) fauna is reviewed, which currently comprises 4011 species, including 405 non-native species. DNA barcodes available for Canadian specimens are represented by 3275 BINs. The analysis was based on the most recent checklist of Hemiptera in Canada (Maw et al. 2000) and subsequent collection records, literature records and compilation of DNA barcode data. It is estimated that almost 600 additional species remain to be discovered among Canadian Hemiptera.

Keywords

Barcode Index Number (BIN), biodiversity assessment, Biota of Canada, DNA barcodes, Hemiptera, true bugs

The order Hemiptera, the true bugs, is a relatively large order. Worldwide there are an estimated 106,970 described species (Henry 2017, Bartlett et al. 2018, Hardy 2018). The recognised Canadian Hemiptera fauna (Table 1) has been greatly expanded since the review by Scudder (1979), with an increase of 937 species above the 3079 then known. The checklist of Canadian Hemiptera (Maw et al. 2000) provided comprehensive lists and distributions for all species recognised at that time. Here we present updated totals, including a number of additional unpublished records represented by specimens in the Canadian National Collections of Insects, Arthropods and Nematodes (CNCI) in Ottawa. Gwiazdowski et al. (2015) presented a detailed analysis of all Hemipteran DNA barcodes available for the Canadian fauna.

Census of Hemiptera in Canada.

Taxon1 No. species reported in Scudder (1979)2 No. species currently known from Canada3 No. BINs4 available for Canadian species Est. no undescribed or unrecorded species in Canada General distribution by ecozone4A Information sources5
Suborder Sternorrhyncha
Superfamily Psylloidea CNCI
Aphalaridae 6 50 37 (1) 19 10 all ecozones but Arctic
Calophyidae 6 ? 1 0 0 Mixedwood Plains
Liviidae 6 4 14 (3) 6 2 all ecozones but Arctic
Psyllidae 6 35 52 (5) 65 20 all ecozones but Arctic
Triozidae 18 25 (1) 13 10 all ecozones but Arctic
Superfamily Aleyrodoidea
Aleyrodidae 3 13 (4) 40 40 all ecozones south of taiga CNCI
Infraorder Aphidomorpha
Superfamily Adelgoidea
Adelgidae 22 18 (5) 14 1 all ecozones but Arctic
Superfamily Aphidoidea
Aphididae 650 847 (164) 758 100 all ecozones CNCI
Superfamily Phylloxeroidea
Phylloxeridae 6 8 (2) 11 4 Pacific Maritime, Mixedwood Plains, Boreal Plains (1 sp.)
Infraorder Coccomorpha
Superfamily Coccoidea
Asterolecaniidae 1 2 (2) 0 0 Pacific Maritime, Mixedwood Plains
Coccidae 15 26 (12) 7 5 all ecozones, mostly south of Arctic
Cryptococcidae ? 2 (2) 0 0 Mixedwood Plains, Atlantic Maritime
Dactylopiidae 1 1 0 0 Prairies
Diaspididae 16 30 (10) 4 10 all ecozones, mostly south of taiga
Eriococcidae 3 3 (2) 2 0 Pacific Maritime, Mixedwood Plains, Atlantic Maritime
Kermesidae 0 4 0 1 Mixedwood Plains
Margarodidae 4 7 0 0 0
Matsucoccidae 7 ? 2 0 0 Mixedwood Plains, Atlantic Maritime
Ortheziidae 8 3 5 2 0 all ecozones but Arctic
Pseudococcidae 13 9 25 (9) 11 10 all ecozones, mostly south of taiga Newton et al. 2011; CNCI
Putoidae 9 ? 2 0 0 Montane Cordillera, Boreal Cordillera
Rhizoecidae 0 1 0 2 Pacific Maritime CNCI
Steingeliidae 7 ? 1 (1) 0 0 Mixedwood Plains
Xylococcidae 7 ? 1 0 0 Atlantic Maritime
Total Sternorrhyncha 844 1120 (223) 955 215
Suborder Auchenorrhyncha
Superfamily Fulgoroidea
Acanaloniidae 10 ? 2 2 0 Mixedwoods Plains
Achilidae 17 19 15 3 all ecozones but Arctic
Caliscelidae 7 11 7 0 mostly south of boreal
Cixiidae 25 11 31 14 5 all ecozones but Arctic
Delphacidae 81 138 (1) 102 30 all ecozones
Derbidae 14 21 17 3 all ecozones south of taiga; most in Mixedwood Plains
Dictyopharidae 12 4 8 5 1 widespread south of taiga
Flatidae 1 3 3 0 Mixedwoods Plains
Issidae 3 10 2 1 0 Mixedwoods Plains
Kinnaridae 11 ? 1 0 0 Western Interior Basin
Infraorder Cicadomorpha
Superfamily Cicadoidea
Cicadidae 9 21 10 7 mostly south of taiga, Taiga Plains (1 sp.)
Superfamily Cercopoidea
Aphrophoridae 13 ? 23 (4) 19 2 all ecozones but Arctic
Cercopidae 33 13 1 0 0 Mixedwoods Plains
Clastopteridae 13 ? 12 8 2 all ecozones south of taiga
Superfamily Membracoidea
Cicadellidae 800 1097 (76) 1144 150 all ecozones Dmitriev 2018, specimens in CNCI
Membracidae 70 101 (1) 65 20 widespread south of boreal, few in Boreal Shield and Boreal Plains Dmitriev 2018
Total Auchenorrhyncha 1064 1491 (82) 1412 223
Suborder [?] Heteroptera Scudder 2008 and references included therein, Scudder 2012, Roch 2017
Infraorder Enicocephalomorpha
Superfamily Enicocephaloidea
Aenictopecheidae 0 1 0 0 Boreal Plains (single collection)
Enicocephalidae 14 1 1 2 1 Mixedwood Plains, Atlantic Maritime
Infraorder Dipsocoromorpha
Superfamily Dipsocoroidea
Ceratocombidae 15 1 1 7 5 Pacific Maritime, Mixedwood Plains
Schizopteridae 0 1 (1) 0 0 Pacific Maritime
Infraorder Gerromorpha
Superfamily Gerroidea
Gerridae 19 22 13 1 all ecozones but Arctic
Veliidae 6 8 6 0 all ecozones south of taiga
Superfamily Hebroidea
Hebridae 4 5 2 0 all ecozones south of taiga
Superfamily Hydrometroidea
Hydrometridae 1 1 1 0 Pacific Maritime, Prairies, Mixedwood Plains, Atlantic Maritime
Superfamily Mesovelioidea
Mesoveliidae 2 2 2 0 all ecozones south of taiga
Infraorder Nepomorpha
Superfamily Corixoidea
Corixidae 72 79 57 2 all ecozones but Arctic
Superfamily Naucoroidea
Naucoridae 0 1 1 0 Mixedwood Plains
Superfamily Nepoidea
Belostomatidae 3 4 2 0 all ecozones but Arctic
Nepidae 7 4 6 1 Mixedwood Plains, southern Boreal Shield
Superfamily Notonectoidea
Notonectidae 12 12 10 0 all ecozones but Arctic
Pleidae 2 1 1 0 Mixedwood Plains, southern Boreal Shield
Superfamily Ochteroidea
Gelastocoridae 1 1 1 0 Pacific Maritime, Prairies, Mixedwood Plains
Ochteridae 0 1 0 0 Mixedwood Plains
Infraorder Leptopodomorpha
Superfamily Saldoidea
Saldidae 36 38 23 2 All ecozones, mostly south of Arctic
Infraorder Cimicomorpha
Cimicoidea
Anthocoridae 4116 39 (7) 36 1 widespread south of taiga
Cimicidae 4 7 3 0 widespread south of taiga
Lasiochilidae 16 ? 1 0 0 Mixedwood Plains
Lyctocoridae 16 ? 6 2 0 all ecozones south of taiga
Superfamily Naboidea
Nabidae 12 22 (3) 19 2 all ecozones south of Arctic
Superfamily Microphysoidea
Microphysidae 0 3 (3) 1 0 near Pacific and Atlantic ports of entry
Superfamily Miroidea
Miridae 60117 706 (57) 414 100 mostly south of Arctic, widespread
Tingidae 46 52 (6) 30 10 all ecozones south of taiga
Superfamily Reduvioidea
Reduviidae 2618 29 (3) 20 0 all ecozones south of taiga
Infraorder Pentatomomorpha
Superfamily Aradoidea
Aradidae 47 19 51 13 2 all ecozones but Arctic
Superfamily Coreoidea
Alydidae 10 9 7 0 all ecozones but Arctic
Coreidae 11 15 14 0 all ecozones but Arctic
Rhopalidae 9 19 (1) 14 2 all ecozones but Arctic
Superfamily Lygaeoidea
Artheneidae 0 1 (1) 1 0 all ecozones south of taiga
Berytidae 20 3 5 (1) 6 0 all ecozones south of boreal
Blissidae 21 ? 6 4 0 all ecozones south of boreal
Cymidae 21 ? 5 4 0 all ecozones south of taiga
Geocoridae 21 ? 10 5 0 all ecozones but Arctic
Heterogastridae 21 ? 2 (1) 0 0 Pacific Maritime
Lygaeidae 100 21 27 (1) 18 2 all ecozones, most south of Arctic
Oxycarenidae 21 ? 5 (1) 1 0 all ecozones but Arctic
Pachygronthidae 21 ? 3 4 1 all ecozones but Arctic
Piesmatidae 1 4 1 0 all ecozones south of taiga
Rhyparochromidae 21 ? 71 (10) 52 7 all ecozones but Arctic
Superfamily Pentatomoidea
Acanthosomatidae 5 5 (1) 3 0 all ecozones but Arctic
Cydnidae 7 12 (1) 13 3 Pacific Maritime, Mixedwood Plains
Pentatomidae 63 22 77 (1) 68 5 all ecozones but Arctic
Scutelleridae 9 13 9 2 all ecozones but Arctic, mostly south of taiga
Thyreocoridae 23 9 11 12 2 all ecozones but Arctic, mostly south of taiga
Total Heteroptera 1171 1400 (100) 908 151
Total Hemiptera 3079 4011 (405) 3275 589

A further 590 species are estimated to occur in the country, with the majority expected in the large families Aphididae, Cicadellidae, and Miridae. Estimates of the number of unrecorded species for the less diverse families is based mainly on known but undescribed species and presence of species in adjacent climatologically and ecologically similar parts of the United States, and known distributions of host plants. Molecular data and analysis of host plant usage provides evidence of additional cryptic diversity in the more speciose phytophagous groups. For some families, presence of unnamed clusters in the DNA barcode data suggests additional species, assuming that in most cases Barcode Index Numbers (BINs), as defined by Ratnasingham and Hebert (2013), correspond to one or more species (Gwiazdowski et al. 2015).

The classification used here follows Psyl’list (Ouvrard 2018) for Psylloidea, Aphid Species File (Favret 2018) for Aphidomorpha, ScaleNet (García Morales et al. 2016) for Coccoidea, Bartlett et al. (2018) for higher classification of Auchenorrhyncha, Bartlett et al. (2014) for species level delimitation in Fulgoromorpha, Dmitriev (2018) for species level delimitation in Cicadomorpha, and Henry (2017) for higher level classification of Heteroptera. There have been several changes in higher level classification since Scudder (1979). Homoptera, no longer recognised as a formal taxon, is now treated as two suborders, namely Sternorrhyncha and Auchenorrhyncha. Although not followed here, some authors (after Sorensen et al. 1995) separate the Auchenorrhyncha into suborders Clypeorrhyncha and Archaeorrhyncha. Among Sternorrhyncha, many new families of scale insects have been erected; species in Canada formerly included in Margarodidae are now dispersed among Matsucoccidae, Steingeliidae and Xylococcidae; Putoidae was formerly included in Pseudococcidae. Schemes for the subdivision of family Aphididae, such as that of Heie (1980) (and used in the Hemiptera checklist of Maw et al. 2000), have been proposed, but largely ignored in the absence of a clear consensus on the relationships among aphid subgroups. Within the Fulgoroidea, the Acanaloniidae, Dictyopharidae, and Kinnaridae are now recognised in Canada, with their species removed from the Issidae, Fulgoridae, and Cixiidae, respectively. The broadly constituted Cercopidae has been divided, with most Canadian species now placed in the Clastopteridae and Aphrophoridae. In the Heteroptera, most former lygaeid subfamilies have been given family status so that Lygaeidae of Scudder (1979) is now represented by eight families (Henry 1997); the further segregation of Ischnorhynchidae and Orsillidae (Sweet 2000) from Lygaeidae is not recognised here. Lyctocoridae and Lasiochilidae have been separated from Anthocoridae. On the other hand, Aradidae now includes Meziridae, Miridae includes Isometopidae, and Reduviidae includes Phymatidae and Ploiariidae.

The 419 non-native species of Hemiptera represents a significant proportion of the total fauna. In Aphididae, about 19% of the species are non-native. An overview of the non-native aphid fauna of North America was provided by Foottit et al. (2006) and updated by Skvarla et al. (2017). The non-native Heteroptera of Canada (about 7% of the total fauna) were treated by Scudder and Foottit (2006).

Sternorrhyncha

Worldwide, the Sternorrhyncha are represented by about 18,690 species (Hardy 2018), with about 2950 species in North America (approximate composite number based on Foottit et al. 2006, García Morales et al. 2016, Skvarla et al. 2017, Mallory 2018, Ouvrard and Martin 2018). Currently, 1120 species of Sternorrhyncha are known from Canada compared to 834 in 1979, and it is expected that a further 215 species will be eventually found in the country (Table 1).

In Canada, the Aphididae and Adelgidae are relatively well known. Foottit and Maw (1997, 2014) contributed syntheses of the Yukon and grassland faunas of Aphididae. Aphids and adelgids are well represented by DNA barcodes (Foottit et al. 2008, 2009a, b), and, in general, barcode diversity in these groups corresponds well to morphological species concepts. However, several currently recognised aphid species are represented by more than one BIN, as defined by Ratnasingham and Hebert (2013). In two such cases, subsequent morphological analysis and addition of other genetic loci has resulted in the recognition of new species (Foottit et al. 2010, Foottit and Maw 2018). Conversely, members of several aphid species groups are not distinguishable by COI sequence divergence (Foottit et al. 2008).

The Psylloidea have not been subjected to extensive taxonomic or faunal analysis in Canada, except for work by Hodkinson (e.g., Hodkinson 1976) in British Columbia and Alberta. The identifiable forms of Coccoidea (adult female) and of Aleyrodidae (immatures) are sessile or subterranean and thus not captured by the usual general collecting methods. Consequently, the national fauna and regional distribution of species in these two groups are poorly known, and even limited efforts can yield new records. Kozár et al. (1989) identified several species of scale insects new to Canada based on brief collecting efforts in southern British Columbia. In a recent study of ant–sternorrhynch associations at a single grassland site in Alberta, two of the four species of Pseudococcidae found were newly recorded for Canada (Newton et al. 2011). The number of available BINs (see Table 1), largely based on untargeted sampling by the Biodiversity Institute of Ontario (University of Guelph), has indicated that current knowledge greatly under-represents the true fauna of Psyllidae and Aleyrodidae if BIN diversity can be considered a good approximation of species diversity in these groups.

Auchenorrhyncha

Worldwide there are about 43,024 species of Auchenorrhyncha (Bartlett et al. 2018), but an estimate for North America is currently not available. In Canada, Hamilton (1997, 2014) analysed the cicadellid fauna of the Yukon and the Canadian Prairie Ecozone, respectively, Gareau (2008) documented that of Quebec, and Wilson (1997) treated the Yukon Delphacidae. The taxonomic status of several auchenorrhynch groups has been updated. A number of papers by Hamilton (e.g., Hamilton 1983, 1994, 1998) have revised many groups of Cicadellidae. As well, the large and difficult tribe Erythroneurini has been completely revised by Dmitriev and Dietrich (2007, 2009, 2010). The Cercopoidea were completely revised and a handbook published by Hamilton (1982). Progress on the Fulgoroidea includes a review of the North American fauna (Bartlett et al. 2014), which provides illustrated keys to all genera including the first comprehensive key to delphacid genera in the region. The known diversity of Canadian Auchenorrhyncha has increased since Scudder (1979), mostly as a result of taxonomic progress and improved knowledge of distributions. Currently, 1491 species are known from Canada compared to 1060 in 1979, and it is expected that a further 223 species will be eventually found in the country (Table 1). Most of the increases are in line with estimates of unrecorded species provided by Scudder (1979). The highest proportional increase is among the Cicadidae from nine to 21 recorded species; this increase is entirely due to improved knowledge of distribution (Hamilton 2010, Sanborn and Phillips 2013) as little taxonomic work has been done on the family in Canada. The highest numerical increase is among the Cicadellidae with an increase of 297 species; this is due to a combination of significant taxonomic research, greatly increased knowledge of distributions, and a small number of recently introduced species. We expect this family to hold the largest number of still unrecorded species (estimated at 150 species), particularly among the under-studied and under-collected subfamily Typhlocybinae.

Significant progress has been made in DNA barcoding of the Canadian Auchenorrhyncha. Data for 691 species have been released (Foottit et al. 2014, Gwiazdowski et al. 2015) and unpublished data for additional species is available in Barcode of Life Data System (Ratnasingham and Hebert 2007). A simple comparison between the number of BINs and recorded species suggests that more than half the Canadian species have been barcoded in most families, and an impressive 91% of the highly diverse Cicadellidae. However, caution is required in interpreting these numbers. Single BINs have been shown to include multiple morphologically distinct species in a number of cicadellid genera (Foottit et al. 2014). Conversely, preliminary examination of BINs for the Typhlocybinae suggests that single species may be represented by multiple BINs. Thus, the number of BINs may not be predictive of the number of distinct species within these groups.

Heteroptera

There are about 45,254 described species of Heteroptera in the world. The most recent published comprehensive catalog for the group in North America (Henry and Froeschner 1985) includes 3834 species. About 1400 species are currently known to occur in Canada compared to 1171 in 1979, and it is expected that a further 151 species will be eventually added (Table 1). Most families of this suborder are relatively well known in the country. However, representatives of two families have been found in Canada only recently: Schizopteridae in 2010 (Scudder 2010a) and Aenictopecheidae in 2016 (Scudder and Štys 2016). Roch (2017) recently documented the Heteroptera of Quebec. Detailed analyses of the faunas of the Yukon, grasslands, Atlantic Maritime Ecozone and Montane Cordillera Ecozone have appeared (Scudder 1997, 2010b, 2011, 2014), and the aquatic and semiaquatic Heteroptera of Canadian peatlands and marshlands and the aquatic Heteroptera of the prairies and parklands were documented by Scudder (1987) and Scudder et al. (2010). Kelton (1980) provided a handbook of the Miridae of the Prairie Provinces. A major on-line database of the mirid fauna of North America (Schuh 2002–2013) includes records for a major portion of the holdings of the CNCI. DNA barcodes for the 334 species of Heteroptera drawn from CNCI (mainly Canadian species) were presented by Park et al. (2011) and reanalysed by Gwiazdowski et al. (2015).

The predicted number of Miridae constitutes the bulk of the estimate for Heteroptera overall, but this number is speculative and may be an underestimate if there are a significant number of undetected cryptic species within the more speciose genera.

Summary and opportunities

Despite the significant increase in knowledge of Hemiptera in Canada since 1979, a substantial amount of the country’s biodiversity still awaits discovery. Some groups of Hemiptera are relatively well documented in Canada, while others are quite poorly known. However, even in the well-studied, but highly diverse phytophagous families (such as Aphididae and Miridae), there is probably unrecognised cryptic diversity associated with host plants and geographic variation. Several large genera in these families, such as Lygus, continue to present taxonomic difficulties (Schwartz and Foottit 1998) and opportunities for application of new approaches and technologies. Because many species of Hemiptera are current or potential pests, continuing research on detection, identification, quarantine and management of these groups will be required.

Acknowledgments

The express their thanks to Jeremy deWaard for updates to the number of available Barcode Index Numbers. We thank David Langor and Cory Sheffield for their critical comments.

References

  • Bartlett CR, Dietz LL, Dmitriev DA, Sanborn AF, Soulier-Perkins A, Wallace MS (2018) The Diversity of the True Hoppers (Hemiptera: Auchenorrhyncha). In: Foottit RG, Adler PH (Eds) Insect Biodiversity: Science and Society. Volume II. Wiley Blackwell, Chichester, UK, 501–590. https://doi.org/10.1002/9781118945582.ch19
  • Bartlett CR, O’Brien LB, Wilson SW (2014) A review of the planthoppers (Hemiptera: Fulgoroidea) of the United States. Memoirs of the American Entomological Society 50: 1–286.
  • Dmitriev DA, Dietrich CH (2007) Review of the New World Erythroneurini (Hemiptera: Cicadellidae: Typhlocybinae). I. Genera Erythroneura, Erasmoneura, Rossmoneura, and Hymetta. Illinois Natural History Survey Bulletin 38: 59–128. http://hdl.handle.net/2142/26473
  • Dmitriev DA, Dietrich CH (2009) Review of the New World Erythroneurini (Hemiptera: Cicadellidae: Typhlocybinae). III. Genus Erythridula. Illinois Natural History Survey Bulletin 38: 215–334. http://hdl.handle.net/2142/26003
  • Dmitriev DA, Dietrich CH (2010) Review of the New World Erythroneurini (Hemiptera: Cicadellidae: Typhlocybinae). IV. Genus Eratoneura. Illinois Natural History Survey Bulletin 39: 79–258. http://hdl.handle.net/2142/25964
  • Foottit RG, Floate KD, Maw HEL (2010) Molecular evidence for sympatric taxa within Pemphigus betae (Hemiptera: Aphididae: Eriosomatinae). The Canadian Entomologist 142: 344–353. https://doi.org/10.4039/n10-028
  • Foottit RG, Halbert SE, Miller GL, Maw E, Russell LM (2006) Adventive aphids (Hemiptera: Aphididae) of America North of Mexico. Proceedings of the Entomological Society of Washington 108: 583–610.
  • Foottit RG, Maw E (1997) Aphids (Homoptera: Aphidoidea) of the Yukon. In: Danks HV, Downes JA (Eds) Insects of the Yukon. Biological Survey of Canada Monograph Series 2, 387–404.
  • Foottit RG, Maw E (2014) Aphids (Hemiptera: Aphidoidea) of the Prairies Ecozone of Canada. In: Cárcamo HA and Giberson DJ (Eds) Arthropods of Canadian Grasslands (volume 3): Biodiversity and Systematics Part 1. Biological Survey of Canada, Ottawa, 347–369.
  • Foottit RG, Maw HEL (2018) Cryptic species in the aphid genus Melaphis Walsh (Hemiptera: Aphididae: Eriosomatinae). The Canadian Entomologist 150: 35–65. https://doi.org/10.4039/tce.2017.57
  • Foottit RG, Maw HEL, Havill NP, Ahern RG, Montgomery ME (2009a) DNA barcodes to identify species and explore diversity in the Adelgidae (Insecta: Hemiptera: Aphidoidea). Molecular Ecology Resources 9 (Suppl. 1): 188–195. https://doi.org/10.1111/j.1755-0998.2009.02644.x
  • Foottit RG, Maw HEL, Pike KS (2009b) DNA barcodes to explore diversity in aphids (Hemiptera: Aphididae and Adelgidae). Redia 92: 87–91.
  • Gareau A (2008) Catalogue des cicadellides du Québec. Entomofaune du Québec Inc. , Saguenay, QC, 261 pp.
  • García Morales M, Denno BD, Miller DR, Miller GL, Ben-Dov Y, Hardy NB (2016) ScaleNet: A literature-based model of scale insect biology and systematics. http://scalenet.info [accessed 12.IV. 2018].
  • Hamilton KGA (1982) The spittlebugs of Canada. The Insects and Arachnids of Canada, part 10. Biosystematics Research Institute, Ottawa, ON, 102 pp.
  • Hamilton KGA (1983) Revision of the Macropsini and Neopsini of the New World (Rhynchota: Homoptera: Cicadellidae), with notes on intersex morphology. Memoirs of the Entomological Society of Canada No. 115(S123), 5–223. https://doi.org/10.4039/entm115123fv
  • Hamilton KGA (1994) Evolution of Limotettix Sahlberg (Homoptera: Cicadellidae) in peatlands, with descriptions of new taxa. Memoirs of the Entomological Society of Canada No. 126(S169), 111–133. https://doi.org/10.4039/entm126169111-1
  • Hamilton KGA (1997) Leafhoppers (Homoptera: Cicadellidae) of the Yukon: dispersal and endemism. In: Danks HV, Downes JA (Eds) Insects of the Yukon. Biological Survey of Canada Monograph Series 2, Ottawa, ON, 337–375.
  • Hamilton KGA (1998) The species of the North American leafhoppers Ceratagallia Kirkaldy and Aceratagallia Kirkaldy (Rhynchota: Homoptera: Cicadellidae). The Canadian Entomologist 130: 427–490. https://doi.org/10.4039/Ent130427-4
  • Hamilton KGA (2014) Canadian Grasslands and their Endemic Leafhoppers (Hemiptera: Auchenorrhyncha: Cicadellidae). In: Cárcamo HA, Giberson DJ (Eds) Arthropods of Canadian Grasslands (volume 3): Biodiversity and Systematics Part 1. Biological Survey of Canada (Terrestrial Arthropods), Ottawa, 311–345.
  • Hardy NB (2018) The Biodiversity of Sternorrhyncha: Scale Insects, Aphids, Psyllids, and Whiteflies. In: Foottit RG, Adler PH (Eds) Insect Biodiversity: Science and Society. Volume II. Wiley Blackwell, Chichester, 591–625. https://doi.org/10.1002/9781118945582.ch20
  • Heie O (1980) The Aphidoidea (Hemiptera) of Fennoscandia and Denmark. I. General Part. The families Mindaridae, Hormaphididae, Thelaxidae, Anoeciidae, and Pemphigidae. Fauna Entomologica Scandinavica 9: 1–236.
  • Henry TJ (1997) Phylogenetic analysis of family groups within the infraorder Pentatomomorpha (Hemiptera: Heteroptera), with emphasis on the Lygaeoidea. Annals of the Entomological Society of America 90: 275–301. https://doi.org/10.1093/aesa/90.3.275
  • Henry TJ (2017) Biodiversity of Heteroptera. In: Foottit RG, Adler PH (Eds) Insect Biodiversity: Science and Society. Volume I, Second Ed. Wiley Blackwell, Chichester, UK, 279–335. https://doi.org/10.1002/9781118945568.ch10
  • Henry TJ, Froeschner C (Eds) (1985) Catalog of the Heteroptera, or True Bugs, of Canada and the Continental United States. EJ Brill, New York, 958 pp.
  • Kelton LA (1980) The Plant Bugs of the Prairie Provinces of Canada. Heteroptera: Miridae. The Insects and Arachnids of Canada. Part 8. Research Branch, Agriculture Canada, Publication 1703, Ottawa, ON, 408 pp.
  • Kozár F, Humble LM, Foottit RG, Otvos IS (1989) New and little known scale insects (Homoptera: Coccoidea) from British Columbia. Journal of the Entomological Society of British Columbia 86: 70–77.
  • Langor DW (2019) The diversity of terrestrial arthropods in Canada. In: Langor DW, Sheffield CS (Eds) The Biota of Canada – A Biodiversity Assessment. Part 1: The Terrestrial Arthropods. ZooKeys 819: 9–40. https://doi.org/10.3897/zookeys.819.31947
  • Maw HEL, Foottit RG, Hamilton KGA, Scudder GGE (2000) Checklist of the Hemiptera of Canada and Alaska. NRC Research Press, Ottawa, ON, 220 pp.
  • Newton JS, Glasier J, Maw HEL, Proctor HC, Foottit RG (2011) Ants and subterranean Sternorrhyncha in a native grassland in east-central Alberta, Canada. The Canadian Entomologist 143: 518–523. https://doi.org/10.4039/n11-034
  • Roch JF (2017) Liste des punaises du Québec et des régions adjacentes (Hemiptera : Heteroptera). Entomofaune du Québec, Document Faunique no. 27 (version 2.2), 38 pp.
  • Sanborn AF, Phillips PK (2013) Biogeography of the cicadas (Hemiptera: Cicadidae) of North America, north of Mexico. Diversity 2013: 166–239. https://doi.org/10.3390/d5020166
  • Schwartz MD, Foottit RG (1998) Revision of the Nearctic species of the genus Lygus Hahn, with a review of the Palaearctic species (Heteroptera: Miridae). Memoirs on Entomology, International, Volume 10. Associated Publishers, Gainesville, FL, 428 pp.
  • Scudder GGE (1987) Aquatic and semiaquatic Hemiptera of peatlands and marshes in Canada. In: Rosenberg DM, Danks HV (Eds) Aquatic Insects of Peatlands and Marshes in Canada. Memoirs of the Entomological Society of Canada No. 140, 65–98. https://doi.org/10.4039/entm119140065-1
  • Scudder GGE (1997) True bugs (Heteroptera) of the Yukon. In: Danks HV, Downes JA (Eds) Insects of the Yukon. Biological Survey of Canada Monograph Series 2, Ottawa, ON, 241–336.
  • Scudder GGE (2008) New provincial and state records for Heteroptera (Hemiptera) in Canada and the United States. Journal of the Entomological Society of British Columbia 105: 3–18.
  • Scudder GGE (2010a) The Schizopteridae (Hemiptera): a family new to Canada. Journal of the Entomological Society of British Columbia 106: 3–9.
  • Scudder GGE (2010b) True bugs (Heteroptera) of the Atlantic Maritime Ecozone. In: McAlpine DF, Smith IM (Eds) Assessment of species diversity in the Atlantic Maritime Ecozone. NRC Research Press, Ottawa, ON, 381–403.
  • Scudder GGE (2012) Additional provincial and state records for Heteroptera (Hemiptera) in Canada and the United States. Journal of the Entomological Society of British Columbia 109: 55–69.
  • Scudder GGE (2014) The Heteroptera (Hemiptera) of the Prairie Ecozone of Canada. In: Cárcamo HA, Giberson DJ (Eds) Arthropods of Canadian Grasslands (volume 3): Biodiversity and Systematics Part 1. Biological Survey of Canada, Ottawa, 283–304.
  • Scudder GGE, Alperyn MA, Roughly RE (2010) Aquatic Hemiptera of the prairie grasslands and parklands. In: Shorthouse JD, Floate KD (Eds) Arthropods of the Canadian Grasslands (Volume 1): Ecology and interactions in grassland habitats. Biological Survey of Canada Monograph 3, 303–323. https://doi.org/10.3752/9780968932148.ch14
  • Scudder GGE, Foottit RG (2006) Alien true bugs (Hemiptera: Heteroptera) in Canada: composition and adaptations. The Canadian Entomologist 138: 24–51. https://doi.org/10.4039/n05-801
  • Scudder GGE, Štys P (2016) Aenictopecheidae (Hemiptera: Heteroptera: Enicocephalomorpha): a family new to Canada. Proceedings of the Entomological Society of Washington 118: 588–593. https://doi.org/10.4289/0013-8797.118.4.588
  • Skvarla MJ, Halbert S, Foottit RG, Jensen AS, Maw E, Miller GL (2017) An update to the adventive aphids of America north of Mexico, with notes on intercepted species Proceedings Entomological Society Washington 119: 90–111. https://doi.org/10.4289/0013-8797.119.1.90
  • Sorensen JT, Campbell BC, Gill RJ, Steffen-Campbell JD (1995) Non-monophyly of Auchenorrhyncha (“Homoptera”), based upon 18s rDNA phylogeny: eco-evolutionary and cladistics implications within pre-Heteroptodea Hemiptera (s.l.) and a proposal for new monophyletic suborders. Pan-Pacific Entomologist 7: 31–60.
  • Sweet MH II (2000) Seed and chinch bugs (Lygaeoidea). In: Schaefer CW, Panizzi AR (Eds) Heteroptera of Economic Importance. CRC Press, Boca Raton, FL, 143–264. https://doi.org/10.1201/9781420041859.ch6
  • Wilson SW (1997) Delphacid planthoppers (Homoptera: Fulgoroidea: Delphacidae) of the Yukon. In: Danks HV, Downes JA (Eds) Insects of the Yukon. Biological Survey of Canada Monograph Series 2, Ottawa, ON, 378–385.