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Home , Agelenidae, Blabomma, Cybaeidae, Dictynoidea, Pimoidae, Pirate spider

Vol. 18, No. 1 16

Canadian Diversity and Systematics

Robb Bennett British Columbia Ministry of Forests, 7380 Puckle Road, Saanichton BC V8M 1W4 Introduction collected. Recent revisions have added consid- It can be stated accurately that erable numbers of to some groups, par- are to be found everywhere (with apologies to ticularly those with predominantly cryptic, William Morton Wheeler who originally made ground-dwelling members (e.g. Gnaphosidae – this claim for ants). Spiders, unique among all 63 species (Dondale 1979) versus 100 species organisms in their modes of silk production and (Platnick and Dondale 1992)). Such groups usage and of reproduction, are common (if of- have historically been passed over in favour of ten inconspicuous) predatory in all those with larger, showier, or otherwise more terrestrial and many aquatic ecosystems obvious members (Alderweireldt and Jocqué throughout Canada. From marine intertidal 1993). Many genera within some cryptic spider zones to tundra and rocky peaks, and all points groups with considerable numbers of Nearctic between, the ardent araneologist is always close species such as , , and to a wealth of species and numerous specimens Amaurobioidea still await serious revision and of great intrinsic and biological interest. within these groups may lie numbers of un- Among all organisms, spiders form the seventh known species with Canadian representatives. largest . Furthermore, spiders are ruthless Still, the number of spider species known or es- storm troops in the matriarchal anarchy that is timated to be awaiting discovery in Canada is the world: theirs is the most diverse, relatively unchanged from Dondale’s estimate female-dominated, entirely predatory order on – there are probably at most 1,500 spider spe- the face of the earth. As such, spiders are key cies in Canada. This is about 35% and 4% of the components of all ecosystems in which they world total numbers of spider families (108) live. and described species (nearly 40,000) respec- tively, not a bad representation for a land mass Twenty years ago, Dondale (1979) listed repeatedly bulldozed in the Pleistocene. 33 spider families with 1256 known Canadian Coddington and Levi (1991) have estimated a species. He estimated another 144 unrecorded world total of 170,000 spider species and that or undescribed species were likely part of the 20% and 80% respectively of the world and Canadian fauna. Araneology is blessed with Nearctic spider fauna have been described. some of the very best systematists in the world and in the intervening years spider Although Canada’s araneofauna is rea- and systematics have been highly active fields sonably well known, precious little is known of (e.g. see the Introduction in Platnick 1997). the behaviour, ecology, habitat associations and Canada’s spider record now stands at 38 fami- other aspects of the biology of the majority of lies: two families have dropped out of the list species. Furthermore, in spite of a rapidly grow- (one does not occur in Canada and the other is ing interest world-wide in many aspects of spi- generally considered a subfamily), five families der biology (e.g. ecology, sexual selection, be- have been added through the apparent resolu- haviour, taxonomy, and systematics) as well as tion of polyphyletic and paraphyletic groups, in habitat-based spider inventories (see Intro- one was apparently missed or perhaps duction in Platnick 1997), there is currently not included in , and one family one professional araneologist in Canada and the (Bennett and Brumwell 1996) has been newly vast majority of Canadian habitats have yet to Vol. 18, No. 1 17

Fig. 1. Male multnoma Chamberlin and Ivie (RTA , Dictynoidea, ). Scale bar=2mm.

Individuals of about two dozen species of this Holarctic are dominant but poorly known generalist predators in the forest floor arthropod community of the Pacific Northwest, especially in coastal regions. In Canada, six species are known to occur in British Columbia (with one species apparently endemic to the Rocky Mountains of BC and Alberta), one species may occur in southeastern Canadian forests. Many species of Cybaeus (most notably in Japan and unglaciated regions of the Pacific Northwest) have ex- tremely restricted ranges and are known from only a few specimens.

be seriously sampled for spiders. Biologists sects and houses a large, world- seeking reliable identifications of Canadian spi- spider collection and taxonomic library. The ders are forced to use the services of a small federal government has effectively declared handful of amateur specialists located in On- spider studies to be irrelevant to the agricultural tario, Saskatchewan, Alberta, and British Co- needs of the nation (C. D. Dondale pers. lumbia or seek help outside the country. Under comm.) and little use is now made of this re- the auspices of Agriculture and Agri-Food Can- source. ada, the Canadian National Collection of In- 18 Newsletter of the Biological Survey of Canada (Terrestrial Arthropods)

Table: Classification of spider families with Canadian representatives (modified from Dondale 1979 and following Coddington and Levi 1990 and Platnick 1997) and estimated numbers of species in Canada. With greater knowledge of forest litter and canopy spiders, the number of species found in Canada should approach 1,500.

Species* Species* ORDER ARANEAE Suborder “RTA Clade” (cont’d) Infraorder Fornicephalae 7 Atypidae 1 18 2 Clubionidae 35 Tuberculotae 11 Mecicobothriidae 1 Gnaphosidae 100 1 Zoridae 1 Infraorder 47 Neocribellatae 68 Araneoclada Salticidae 110 Amaurobioidea “Scytodoidea” 30 Scytodidae 1 4 1 11 3 Segestriidae 1 3 1 Nesticidae 2 Theridiidae 100 Mimetidae 6 Theridiosomatidae 1 1 1 1 “RTA Clade” Linyphiidae >500 Tetragnathidae 23 Lycosidae 110 Araneidae 74 Pisauridae 7 Oxyopidae 2 TOTAL ~1400 Dictynoidea 75-80 Cybaeidae 11 Hahniidae 16

*Approximate number of species in Canada (1999 estimate)

All spiders produce and use silk and can dominate the former grouping. The latter is a di- be divided roughly into two behavioural group- verse assemblage dominated by the “RTA ings based on this usage: those that use their silk clade” with major families including the to snare prey and those that don’t. salticids (jumping spiders), lycosids (wolf spi- Orbicularians, especially the araneoids (primar- ders), thomisids (one family of crab spiders), ily sheet-web, orb-web, and cob-web weavers) and gnaphosids (ground spiders). However, us- Vol. 18, No. 1 19 age of silk for prey capture is not phylogeneti- der family in Canada in terms of numbers of cally constrained and numerous groups span the species and often are extremely abundant. On gap. For instance many amaurobioid and the world stage, they are second in diversity dictynoid species (common and widespread in only to salticids. Many years ago, Bristowe Canada) utilize greatly reduced webs (e.g. see (1938) made a well reasoned estimate of over Bennett 1985) and rely upon sit, wait, and 2.25 million spiders (mostly linyphiids) per pounce strategies. Other species within these acre of grassy meadow in late summer in Eng- groups build large and characteristic capture land. He also cited a study which produced webs. Among the fascinating but mostly un- 159,000 spiders per acre from the soil alone of a common Canadian kin similar habitat. (These figures should be kept (Mygalomorphae) atypids (purse-web spiders), from the poor arachnophobes who recoil from diplurids (funnel-web ), and the small handful of spiders they encounter an- mecicobothriids (no common name) use silk in nually.) The bulk of linyphiid diversity is made prey capture. Antrodiaetid mygalomorphs up of erigonine species. Dondale (1979) listed (folding-trapdoor spiders) sit in the entrance of over 300 erigonine linyphiids in Canada with an excavated burrows and wait for hapless crea- estimate of 50 remaining to be recorded or de- tures to come near enough to be grabbed and scribed. These species are characterized by dragged deep into the dim recesses of their re- their undistinguished appearance and small to treats and repeatedly stabbed to death (big-city minute size: not only are a majority of downtown terror is far exceeded by the nightly erigonines tiny, most of them look alike. Usable mayhem on a seemingly peaceful suburban taxonomic keys are virtually non-existent and British Columbia lawn). Linyphiidae (includ- most genera and species are very difficult to de- ing Erigonidae of various authors) and termine even for the specialist. Identification is Salticidae, the major family in each behavioural based largely upon microscopic characters of grouping on a world-wide basis, are very diffi- the very complex male genitalia; in contrast, fe- cult taxonomically and have many genera in males usually are unremarkable morphologi- need of revision and large numbers of undes- cally and often are unidentifiable if not col- cribed species. Following is a summary of the lected with male conspecifics. diversity, abundance, and systematics of Can- Few comprehensive spider lists have ada’s araneofauna. been published for any habitat in Canada except Araneoidea some peatlands (but note that only pitfall trap- The superfamily Araneoidea includes ping was used in most of those studies and many of the spiders most familiar to the therefore the resultant lists likely are incom- layperson including the orb-weaving garden plete) but a casual glance at the generated spe- spiders (Araneidae) and the cob-web weaving cies lists shows a preponderance of linyphiids widow spiders and house spiders (Theridiidae). (especially erigonines): 51 of 106 species col- However, the vast majority of araneoids, and lected (Aitchison-Benell 1994); 28 of 49, 37 of the most numerous, are linyphiids (sheet-web 108, 27 of 59, 32 of 97, 26 of 76, and 22 of 77 weavers), only a few of which are encountered (Dondale and Redner 1994; Blades and Mar- or collected regularly by the uninitiated genera- shall 1994); and 95 of 169 (Koponen 1994). No list. On the world scale, Araneoidea is the most other family comes close in diversity. This pat- diverse and phylogenetically best understood tern is common throughout the Holarctic region superfamily of spiders (Griswold et al. 1998). and erigonines increasingly dominate the spider fauna the farther north one travels: Dondale et Linyphiidae al. (1997) recorded 154 known or expected Linyphiidae accounts for over one third linyphiid species (34+3 linyphiines, 102+5 of all Canadian spider species (445 known Ca- erigonines) in the Yukon fauna of 336 spider nadian species with probably over 70 unknown species. Fully 56% of all Yukon arctic spider – Dondale 1979). Linyphiids are the largest spi- species are erigonine. The world is bottom 20 Newsletter of the Biological Survey of Canada (Terrestrial Arthropods) heavy in terms of plant and diversity: Araneidae and Tetragnathidae most species are found in the tropics and south Araneids are probably the most familiar temperate areas (Platnick 1991). However, spiders to the average Canadian. Almost all the among spiders, linyphiids are most diverse in araneids as well as the closely related the north and predominate in the Holarctic tetragnathids (long-jawed orb-weavers) build fauna (Coddington and Levi 1991). characteristic, radially symmetric orb webs and Theridiidae together these two families have about 100 spe- Theridiidae is the second most diverse cies with Canadian representatives. As with the araneoid family in Canada with about 100 spe- theridiids, the Nearctic tetragnathid and araneid cies (Dondale 1979). Overall, theridiids are as genera have been well revised by Levi and oth- diverse as gnaphosids; both share fourth place ers (references in Roth 1993). A field guide to in Canada after the hunting salticids and Canadian species is being readied for publica- lycosids. Theridiids also are a dominant group tion by volunteers on a part-time basis (C. D. on the world scale although, as in most spider Dondale pers. comm.). In Canada, Araneidae is families other than Linyphiidae, theridiid diver- about as diverse as Dictynidae with which it sity primarily occurs south of the Holarctic re- shares honours for sixth and seventh largest gion. Taxonomically, the Nearctic theridiids families after Linyphiidae, Salticidae and are well known and most genera have been re- Lycosidae (equally diverse), and Theridiidae vised recently by Levi and co-authors (refer- and Gnaphosidae (equally diverse). In the ences in Roth 1993). A field guide to Canadian world, Araneidae is the third largest family theridiids is in preparation but is on hold be- (Griswold et al. 1998), a reflection of its great cause of shifting government funding priorities diversity in the tropics. (C. D. Dondale pers. comm.). Araneids and tetragnathids usually are Theridiids are common and abundant abundant in fields, open woodlands, and other spiders throughout southern Canada in most similar habitats (but note that a significant num- habitats but are relatively minor components of ber of araneids are litter inhabitants). the fauna of the northern Nearctic region. They Tetragnathids often are especially common are rather more conspicuous than linyphiids as around water or in wet areas. Several species most build their webs above ground level and are abundant around homes and outbuildings therefore are more often collected by the gener- and in gardens. The classic araneid orb web was alist. However, a number of genera are - like for many years considered to be a highly de- erigonines - tiny, inconspicuous members of rived structure but now is generally accepted to the soil and deep-litter fauna (Dondale 1990). be a relatively primitive structure, and theridiid Several genera are regularly and easily mis- cob-webs and linyphiid sheet-webs are derived identified by inexperienced workers and often from it (see papers in Shear 1986). are found in collections labelled as unidentified RTA clade clubionids and agelenids. Two theridiid genera, Sister to Orbiculariae (Araneoidea plus Steatoda and Achaearanea, are very common Deinopoidea, a superfamily poorly represented in Canadian homes where they kill most other in Canada) is the large and diverse “RTA clade” arthropods unfortunate enough to become en- (Coddington and Levi 1991). (The name is de- tangled in their cob-webs and thus are particu- rived from the probable synapomorphy of a dis- larly efficient, if not particularly welcome, tinctively placed apophysis on the male palpal house-cleaners. These synanthropic species are tibia.) Resolving the familial and superfamilial routinely misidentified as black widows relationships within the RTA clade is the last (Latrodectus spp., which are closely related major hurdle for spider systematics. RTA clade theridiids) or the infamous (and completely families with Canadian representatives are non-Canadian) brown recluse (Loxosceles sp., placed in Dionycha, Lycosoidea, Dictynoidea, ). and Amaurobioidea. Species in the former two Vol. 18, No. 1 21 groupings, Dionycha and Lycosoidea, are com- the transportation by mature females of egg mon and familiar components of the Canadian sacs on their (giving rise to apocry- araneofauna. Dictynoidea and Amaurobioidea phal tales of spiders with three body sections) are also common components but mostly unfa- and of young spiderlings dorsally on their abdo- miliar to the majority of Canadian biologists. mens (giving rise, among arachnophobes who Dionycha encompasses families of have bravely dispatched a large, fuzzy spider two-tarsal-clawed, primarily sedentary (am- only to watch countless small spiders magically bushing) or active hunters. Dominant Canadian appear and scatter in all directions, to night- groups are Salticidae, Clubionidae (sac spi- mares and apocryphal tales of spontaneous gen- ders), Gnaphosidae (ground spiders), eration). Thomisidae, and Philodromidae (philodromid Two other lycosoid families are com- crab spiders). Lycosoidea is represented in mon but have few species in Canada. Pisaurids Canada primarily by the diverse and extremely (nursery-web spiders) often are encountered abundant lycosids. In Canada, Dictynoidea and along the edges of freshwater systems where Amaurobioidea are dominated by their family they hunt actively above and below the water namesakes. The Nearctic lycosoid and surface. Two pisaurid species are the largest na- dionychan genera are taxonomically well tive spiders in Canada and probably have struck known (except for the salticids) and excellent fear into countless cottagers in Ontario and keys exist for the Canadian genera and species Quebec. Some oxyopids (lynx spiders) may oc- (Dondale and Redner 1978, 1982, 1990; cur in large numbers in agricultural fields and Platnick and Dondale 1992). Some Nearctic have been the subject of studies on natural pest amaurobioid and dictynoid genera have been control (see references in Young and Lockley revised (e.g. Bennett 1987, 1991; Chamberlin 1985). and Gertsch 1958; Leech 1972) but many others are poorly known and no comprehensive guides Salticidae to the Canadian fauna exist. Salticids are the charismatic “cute/fuzzies” of the spider world: many are Lycosidae highly photogenic and, deservedly, they feature In Canada, Lycosidae has just one fifth regularly on the pages of glossy natural history as many species as Linyphiidae but shares with magazines. Jumping spiders are renowned for Salticidae the honour of being the second most their often fantastic degree of sexual dimor- diverse spider family here. Dondale (1979) esti- phism and colouration and wild and interesting mated a Canadian fauna of 110 spe- range of behaviours. Much of this is due to the cies with roughly 10% of these undescribed or highly developed visual sense in salticids: the unrecorded. On the world scale, lycosids make huge main eyes are complex, high-resolution up the fifth largest family (Coddington and Levi structures capable of focusing on objects by ret- 1991). Although lycosid diversity is much inal movement. Throughout the twentieth cen- greater elsewhere, the abundance of individual tury salticids have been the focus of many opti- species often is astounding in Canada. Pitfall cal studies (see references in Foelix 1982, traps in any sort of open habitat are regularly 1996). Jumping spiders are agile, visually ori- swamped with wolf spiders (see abundance ta- enting, day-active predators commonly found bles in Blades and Marshall 1994, Dondale and among foliage in sunny locations. Some are Redner 1994, and Koponen 1994). Fields, tun- synanthropic. dra, shorelines, and open wooded areas may lit- World-wide, Salticidae is the largest erally be alive with wolf spiders (often a single family of spiders: well over 10% (about 5,000) species) particularly in the spring. of all described spider species are jumping spi- Lycosids are diverse and abundant in the far ders. Salticid taxonomy and systematics are north of the Holarctic and account for 18% of very active fields of study (see current review the known Canadian arctic araneofauna by D. B. Richman at: (Dondale et al. 1997). Lycosids are unique in 22 Newsletter of the Biological Survey of Canada (Terrestrial Arthropods)

http://dns.ufsia.ac.be/Arachnology/Pages/Doc- lished data) collected hundreds of specimens of uments/Salticid.html) but primarily with tropi- Gnaphosa snohomish Platnick and Shadab, a cal groups. Some excellent revisions of Nearc- large species previously known only from a pair tic groups have been published recently (e.g. of specimens taken in central Washington. Af- Maddison 1996 and references in Roth 1993) ter lycosids, one or a couple of species of but the bulk of the genera found in Canada are gnaphosids may be among the numerically badly in need of revision. Taxonomically they dominant and visually conspicuous spiders pit- are a trying group with identifications difficult fall-trapped in bogs (see Table 2 in even for specialists. In Canada, Salticidae is Aitchison-Benell 1994; Table 1 in Dondale and about as diverse as Lycosidae with about 110 Redner 1994). Typical of groups with a prepon- species including close to 20% of this number derance of tropical species, Canadian yet to be described or recorded (Dondale 1979). gnaphosids become increasingly scarce with in- As with most spiders except lycosids and creasing latitude. However, Gnaphosidae is still linyphiids, salticids are more diverse and abun- a dominant group in subarctic and alpine re- dant in Canada’s southern latitudes than in the gions in Canada although few species occur in far north. the far north (Dondale et al. 1997). Gnaphosidae Clubionidae Gnaphosidae is the sixth largest spider The “old” Clubionidae was paraphyletic family in the world (Coddington and Levi (Coddington and Levi 1991) and the family was 1991) and a major player in spider diversity in relimited after Dondale’s (1979) estimate of a the tropics. In Canada, Gnaphosidae, with Canadian fauna of about 64 species was made. about 100 species known or expected to live At least 29 of these species have been trans- here (Platnick and Dondale 1992), is about as ferred to Corinnidae and Liocranidae which to- diverse as Theridiidae and slightly less so than gether may be sister to Gnaphosidae and its rel- Salticidae and Lycosidae. In general, atives (Coddington and Levi 1991). The re- gnaphosids are common ground-dwelling hunt- maining species are likely most closely related ers and are important in the natural control of to Salticidae and Anyphaenidae. Although a forest and agricultural insects (Platnick and field guide to Canada’s (paraphyletic) Dondale 1992). However, few are encountered clubionids (and our few species of by general collectors because most are incon- anyphaenids) has been prepared (Dondale and spicuously coloured and nocturnal hunters. Redner 1982), the ground-dwelling genera Members of one genus (Herpyllus) are found placed now in Liocranidae are poorly known. In regularly in buildings. Canada, the relimited Clubionidae is comprised Over the last quarter of this century, primarily of nocturnal species actively hunting Platnick and others (references in Roth 1993) on herbs, shrubs, trees, and other plants. A very have revised many of the world’s gnaphosoid few species occur in litter or under stones. The genera including all of those with Canadian rep- majority of Canadian clubionids are found in resentatives. As a direct consequence, an excel- southern regions with a few species ranging lent taxonomic guide to the Canadian into the subarctic. gnaphosids (Platnick and Dondale 1992) has Thomisidae and Philodromidae been published and few new Nearctic species The two crab spider families with Cana- probably await discovery. However, the biol- dian representatives, Thomisidae and ogy of the majority of gnaphosid species is not Philodromidae, are dionychans of uncertain af- well known, a reflection of our relatively poor finities (Coddington and Levi 1991). grasp of ground-dwelling spiders in general. Thomisidae is seventh in terms of world spider For instance, in a 1998 study of arthropods of diversity with a majority of its species occur- cranberry bogs in an area with a high concentra- ring in the tropics. Together, about 115 species tion of Canadian entomologists (southwestern in the two families occur in Canada. Thomisids, BC), J. Troubridge and S. Fitzpatrick (unpub- Vol. 18, No. 1 23 with their two anterior pairs of long legs and ha- families lack clear diagnosis (except for bitual sideways gait, are distinctly more Amaurobiidae – Griswold 1990). No doubt, if crab-like than philodromids. Both are this area remains active, major changes are yet dorso-ventrally flattened spiders; thomisids to come. tend to be sparsely bristled and slow moving, Dictynidae philodromids hairy and quick. Crab spiders are With the addition of two genera from the primarily diurnal ambush hunters. Thomisids agelenids, the Canadian dictynid araneofauna are more common on the ground, philodromids has grown from about 65 (Dondale 1979) to on foliage. Some thomisids are famous for perhaps 75-80 species, making the regional di- waiting in flowers to grab visiting nectar and versity of this family on a par with Araneidae. pollen feeding insects. Both families are well The cribellate dictynids were revised by represented in Canada’s boreal north and some Chamberlin and Gertsch (1958) but they remain species range into the Arctic (Dondale et al. exceedingly difficult to identify without com- 1997) but the bulk of thomisid and philodromid parison to voucher specimens. Most species are diversity is encountered in the south. quite small and cryptic. The ecribellate taxa Dictynoidea and Amaurobioidea need revision and there are no usable species Although generally accepted as valid level keys for the family. Both cribellate and superfamilies, Dictynoidea and Amaurobioidea ecribellate dictynids weave tangled, relatively remain poorly defined and their family level shapeless webs although some species build systematics is in disarray (Coddington and Levi specialized tube-like structures (Bennett 1985). 1991). Classically, all members of both groups The ecribellate species are found primarily un- were considered to be cribellate (the der objects on the ground in wooded areas. is a distinctive silk production organ found in Rotting logs may harbour considerable num- some spiders and derived from the ancestral an- bers of individual webs. Many of the cribellate terior median spinnerets). Lehtinen’s (1967) species are distinctly arboreal, building their exposure of “Cribellatae” as paraphyletic and small webs on branch tips. The current growing defined by a plesiomorphic character (the interest in canopy arthropod surveys may boost cribellum) marked the beginning of the current the number of dictynid species recorded from era of spider systematics which has resulted in Canada. Several dictynids are widespread in massive restructuring of the classification of Canada’s Boreal zone forests and a very few are Araneae from family to subordinal levels. found in Arctic and Subarctic regions (Dondale Dictynidae and Amaurobiidae are now seen to et al. 1997). The majority of Canada’s dictynids contain both cribellate and ecribellate genera, have southern distributions. many of the latter transferred there after the Other Dictynoidea sundering of the old, paraphyletic Agelenidae Hahniidae and Cybaeidae together have (e.g. and are ecribellate about 30 species with Canadian representatives. dictynids; , , and are Both groups are very common but infrequently ecribellate amaurobiids). The two collected litter inhabitants. Cybaeids are forest superfamilies have Canadian representatives in floor spiders; hahniids may be found in a vari- Dictynidae, Cybaeidae, and Hahniidae ety of litter habitats. Hahniidae has been con- (Dictynoidea) and Amaurobiidae, Agelenidae, siderably augmented by genera transferred and Titanoecidae (Amaurobioidea). The taxon- from Agelenidae. Cybaeidae is comprised en- omy and systematics of amaurobioids and tirely of taxa formerly placed in that family. dictynoids has been studied actively in recent The Nearctic cybaeids have been revised in re- years (e.g. Bennett 1987, 1988, 1991, 1992; cent years (references in Roth 1993) but much Griswold 1990) but little has been well resolved of this work remains unpublished and the fam- above the genus level, various genera are ily is not clearly defined. Most of the Nearctic placed only tentatively or uncertainly within hahniid taxa have also been revised (references one or the other superfamily, and most of the 24 Newsletter of the Biological Survey of Canada (Terrestrial Arthropods) in Roth 1993) but all the genera transferred all true agelenid genera need revision. from Agelenidae need major work. A few Titanoecidae, a cribellate group extracted from hahniids are found in the far north (Dondale et Amaurobiidae, is represented in Canada by al. 1997) but most Canadian species in both only four forest floor inhabiting species. families have southern distributions. Some hahniids are very abundant in peatlands (see Mygalomorphs and other spider Table 1 in Dondale and Redner 1994; Koponen groups 1994). Cybaeids and coelotine amaurobiids are All other spider families with Canadian similar morphologically and ecologically but species are much more diverse elsewhere and, have different areas of endemism in North for the most part, rarely encountered here. America. Coelotines are abundant in eastern Pimoidae (previously considered part of forests and none is found in the west. Cybaeids Linyphiidae) has one species in Canada which are diverse in the west but are nearly absent is abundant on the west coast. Mysmenids, from eastern North America. nesticids, and theridiosomatids are small araneoids with a single rare species each in the Amaurobiidae southernmost reaches of Canada. Three species The cribellate amaurobiids were revised of uloborids (small and interesting cribellate, by Leech (1972) and had 25 species with Cana- deinopoid orb-web weavers) are widespread dian representatives (Dondale 1979). Since but not abundant across southern Canada. One then, the Canadian fauna has lost three species zorid (small, lycosid-like dionychans) was re- to Titanoecidae but has gained eight ecribellate cently discovered in disturbed areas on south- species from Agelenidae for a total of about 30. ern Vancouver Island. A small handful of Two of the three ecribellate genera in Canada mimetid pirate-spider species are the only Ca- need revision. Most Canadian amaurobiids are nadian representatives of Palpimanoidea (an in- forest floor dwellers and build rather amor- teresting group, probably sister to Orbiculariae phous, tangled webs under various objects in + RTA clade, containing many specialized woodlands, especially the loose bark of trees araneophagous species). One ecribellate and logs. Spiders of this family often are abun- oecobiid is an uncommon synanthrope across dant but only the larger species are found com- southern Canada. Haplogynae, originally be- monly in general collections. Most species have lieved to be paraphyletic, is probably sister to southern distributions in Canada but a few are all the “higher” spiders (Entelegynae) including widespread in the Boreal zone and into alpine all those discussed above. In Canada, and subarctic areas (Dondale et al. 1997). Haplogynae is represented by one synanthropic Other Amaurobioidea species each of Scytodidae (spitting spiders) In Canada, Dondale (1979) estimated and Dysderidae, a single tiny and very rare spe- there to be some 65 agelenid species. cies of Telemidae found deep in forest leaf litter Agelenidae has long been an acknowledged in British Columbia, a single segestriid species dump heap for amaurobioid and dictynoid gen- each in the southern reaches of eastern and era of uncertain affinities and the family has un- western Canada and three or four pholcids (two dergone major relimitation in recent years (see of which are common in cellars) in various above). Today in Canada, Agelenidae is a mere southern localities. Centres of diversity for all shadow of its former self with perhaps 11 spe- these groups are austral or elsewhere other than cies in two common and one rare genus. Several north temperate. of these species are very well known because of Four mygalomorph families their abundance and/or synanthropic habits. (Mygalomorphae is sister to Araneomorphae) True agelenids all spin distinctive large, each have one or two species found in Canada, non-sticky sheet webs narrowing to a funnel at mostly in the west. One atypid purse-web spi- one edge wherein the builder is usually found der used to be found on the Niagara Peninsula waiting for prey to become entangled. Virtually but this species has not been collected recently Vol. 18, No. 1 25

Figs. 2-4 Cybaeus constrictus Chamberlin and Ivie, Figs. 5-7 C. conservans Chamberlin and Ivie; female genitalia. 2, 5 epigynum (external), ventral views; 3, 6 vulva (internal), dorsal views; 4, 7 same, ventral views. Scale bars = 0.1 mm.

Figs. 8-11 C. angustiarum L. Koch, Figs. 12-13 C. tetricus (C. L. Koch); characters of male left . 8 genital bulb, ventral view; 9, 13 apical apophysis, retrolateral views; 10 carinate retrolateral tibial apophysis, retrolateral view; 11, 12 patellae, dorsal views. Scale bars = 0.1 mm.

Figures: In spiders, species within genera are normally very similar in general morphology and usually only can be distinguished by careful comparison of sexual characters with published drawings or expertly identified voucher specimens. 26 Newsletter of the Biological Survey of Canada (Terrestrial Arthropods) and may have disappeared from Canada. One gram, effectively mothballing its collection of tiny funnel-web weaving diplurid is found at a close to 150,000 specimens and the only com- single montane locality in southern British Co- plete research library in the country. lumbia. A disjunct population of one Basic taxonomic and systematic re- mecicobothriid sheet-web weaver is found on search provides the foundation for all other as- southwestern Vancouver Island. One pects of biological research and will continue to antrodiaetid trapdoor spider species is common do so long into the future. The alpha-level tax- and abundant in reasonably undisturbed habi- onomy (i.e. basic descriptive work, compara- tats along the entire British Columbia coast. tive morphology) of the vast majority of arthro- Another antrodiaetid is occasionally collected pod groups including most spiders is in rela- in the dry southern interior valleys of that prov- tively poor shape. Research dollars tend to be ince. The vast bulk of mygalomorph diversity is directed at the macroflora and macrofauna at found in the tropics and subtropics. the tips of terrestrial ecosystem icebergs. The Conclusion organisms, primarily arthropods and fungi, “This is an exciting time in the study of comprising the huge and invisible foundations spiders” (C. E. Griswold in Dippenaar- of these icebergs (vital to the existence of the Schoeman and Jocqué 1997). Major advances tips) are too often ignored except for those taxa are being made in the taxonomy and systemat- aspiring to “pest” status. Can we expect to un- ics of spiders at all classification levels. Publi- derstand the functioning of any ecosystem cation of taxonomic papers has been very high without basic knowledge of the hidden organic over the last several decades and shows no sign and inorganic network supporting the visible of levelling off (see Platnick 1997). The Smith- component? “Can’t see the forest for the trees” sonian Institution and the American Museum of is an apt expression: trees obscure our vision of Natural History support active graduate pro- the real machinery of the system. Within eco- grams that are training new systema- system machinery, spiders are diverse and im- tists. These programs have taken up the slack portant cogs. With luck, in the new millenium, from Harvard University as its highly regarded Canada will rediscover the inherent value of its program winds down (Coddington et al. 1990). declining cadre of systematic araneologists (to And there is much work available for spider say nothing of entomologists) and ecosystem systematists in North America: ecological in- research biologists will no longer have to rely terest in spiders is at an all-time high and many upon the volunteer services of accomplished families are in urgent need of revision, particu- amateurs to provide names for their data points. larly Linyphiidae and Salticidae but also most References families with cryptic, ground-dwelling genera. Alderweireldt, M. and R. Jocqué. 1993. in Af- Yet there is little in the way of funding or rica and Europe: the case of spiders (Araneae). Biol. Jb. support for systematists beyond the graduate Dodonaea 61:57-67. level. Trained arachnid systematists are finding Aitchison-Benell, C.W. 1994. Bog arachnids (Araneae, it increasingly difficult to find paid work and Opiliones) from Manitoba taiga. pp. 21-31 in A.T. Finnamore and S.A. Marshall (Eds.), Terrestrial Arthro- often when work is found, research is not a pods of Peatlands, with Particular Reference to Canada. component of it (Coddington et al. 1990). Many Memoirs of the Entomological Society of Canada 169. of the North American arachnid systematists 289 pp. are retired or aging and there is little incentive Bennett, R.G. 1985. Taxonomy and natural history of for younger researchers to enter the field: nearly Cicurina bryantae Exline (Araneae, Agelenidae). Jour- nal of 13:87-96. three-quarters of all active workers receive Bennett, R.G. 1987. Systematics and natural history of nothing or less than $1,000 annually in support Wadotes Chamberlin (Araneae, Agelenidae). Journal of of their research (Coddington et al. 1990). Here Arachnology 15:91-128. in Canada, Agriculture and Agri-Food Canada Bennett, R.G. 1988. The spider genus Cybaeota (Araneae, has shut the doors on its arachnid research pro- Agelenidae). Journal of Arachnology 16:103-119. Vol. 18, No. 1 27

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