124

Linyphiidae
Chapter 35

Final version

LINYPHIIDAE
157 genera, 952 species
Common name Sheet-web spiders. Erigoninae are sometimes called dwarf spiders or money spiders. Similar families Many small ground-dwelling spiders in other families have been mistaken for linyphiids, including Dictynidae (Lathys, p. 94), Mysmenidae (p. 175, Microdipoena), Theridiidae (p. 235, Robertus, Thymoites, Pholcomma, Theonoe), Tetragnathidae (p. 232, Glenognatha), Leptonetidae p. 121, Theridiosomatidae p. 242, and especially Pimoidae p. 195, which is the sister taxon to Linyphiidae (Hormiga 1994a, 2003). Diagnosis Minute to small (<17 mm) three-clawed, ecribellate, entelegyne spiders belonging to the superfamily Araneoidea. They may be distinguished from other Araneoids, except Pimoidae, by the following combination of characters: male palp with discrete, intersegmental paracymbium (rarely lost); stridulatory striae on the ectal side of the chelicerae (occasionally absent) and autospasy of the legs at the patella-tibia junction. Linyphiids and pimoids share cheliceral striae and patella-tibia autospasy. Linyphiids differ from pimoids in having a discrete paracymbium rather than one integrated with the cymbium, having the embolus part of a complex embolic division rather than attached directly to the tegulum, and by lacking a retrolateral cymbial sclerite attached to the cymbium via a membrane, which is a pimoid synapomorphy. Characters body size: from less than 1.0 to 7 mm; most are 1-4 mm long. color: white, tan, brown to black. Occasionally shades of orange, red, and yellow. Some species (especially larger arboreal species) may have dorsal abdominal stripes, chevrons, or markings; or banded legs. carapace: longer than wide; the carapace of many male linyphiids is modified in various ways. Eye region commonly elevated, sometimes exhibits bizarre forms. A pair of lateral pits may be present, opening near the posterior median eyes. Various clypeal protuberances or modifications, unusual setae or clusters of setae, or teeth around the carapace margin may be observed. sternum: longer than wide, projecting behind coxae IV. eyes: usually eight subequal eyes in two rows. Rarely reduced or absent (in cavernicolous species). Eyes of males with modified carapaces often in unusual configurations. chelicerae: relatively robust for Araneoid spiders, armed with teeth on both margins. Most with a stridulatory file on the lateral face. Some males have a mastidion or similar tooth-like apophysis on the anterior cheliceral face (e.g., Zornella). mouthparts: endites are parallel, rarely armed with apophyses in males (e.g., some Erigone spp.). legs: rather long and thin, with macrosetae. Autospasy occurs at the patella-tibia joint. All have a single meta-

Michael L. Draney Donald J. Buckle

Fig. 35.1 Ceraticelus fissiceps (O. PICKARD-CAMBRIDGE 1874)

tarsal trichobothrium on legs I-III, and some also have one on leg IV. Setation on legs of males often somewhat reduced compared to females. abdomen: usually ovoid to somewhat elongate. Some species have abdominal scuta (dorsal and sometimes ventral) of varying size. spinnerets: colulus present, with setae. Spinnerets are of typical araneoid form. respiratory system: one pair of book lungs, and a tracheal system opening onto a single spiracle anterior to the spinnerets, anteriorly displaced in some species (e.g., Tennesseellum). Most members of the subfamily Linyphiinae have unbranched tracheae confined to the abdomen while most members of the Erigoninae have branched tracheae extending into the cephalothorax. genitalia: entelegyne; female epigynum highly variable; usually fairly simple, plate-like sclerotized structure with a dorsal and a ventral sclerite associated with paired copulatory openings, leading via complex ducts to spermathecae. These sclerotized structures are sometimes, particularly in the Linyphiinae, modified into a posteriorly directed scape; male palps nearly always have an intersegmental paracymbium (=flexibly attached). The embolus is part of a complex structure called the embolic division, attached by a stalk to the suprategulum. This and several other sclerites associated with the tegulum, subtegulum, and suprategulum produces structurally complex palp. Erigonines nearly always have a retrolateral tibial apophysis of varying size and shape, sometimes consisting of multiple projections.

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Distribution Worldwide with its greatest diversity in the north temperate regions. Well represented in North America. Natural history Linyphiids spin semi-permanent webs consisting of one or more horizontal sheet of silk often supported above and below by auxiliary support lines. The spider generally hangs upside down beneath the sheet, attacking prey from below. Most species live in leaf litter or on the ground surface, but some (especially larger species) make webs in vegetation. Both myrmecophiles and cavernicoles are known. Linyphiids generally feed on soft-bodied insects, especially springtails (Collembola) and flies (Diptera). They occupy almost all habitats but are most diverse and abundant in more mesic to hydric (i.e., wetlands) areas, and are prominent in northern peatlands. Most species are univoltine or annuals (one generation/year), but multivoltine (>1 generation/year) and merovoltine (>1 year/generation) strategies are common. Most species occur as adults for only a short period each year, but some multivoltine and merovoltine species are active as adults during most or all of the year. Most species overwinter as juveniles or adults, - some are even active under snow in winter - but a few overwinter as eggs. Taxonomic history and notes Linyphiidae is a large, poorly known, and taxonomically difficult family. Several genera and many species remain undescribed. While a number of North American genera have recently been revised, most have not. Their species are known from original descriptions, scattered through 150 years of literature, and from subsequent references under a variety of genus and species names. Eugen von Keyserling, Octavius Pickard-Cambridge, and Nathan Banks were prolific early describers of North American species, each adding over 30 currently valid species. The first, and only, comprehensive survey of North American linyphiids was published by James Emerton in 1882. He subsequently produced a series of papers describing many new species. Cyrus Crosby and Sherman Bishop published a series of revisionary studies in the 1920s and 1930s. Wilton Ivie, often working with Ralph Chamberlin, described many western species and revised several genera before his untimely death in 1969. Between 1968 and 1981 Peter van Helsdingen, revised Linyphia, Neriene, Microlinyphia, Stemonyphantes and Allomengea on a worldwide basis and Centromerus and Oreonetides for North America. In the 1980s, Frank Millidge, revised many North American genera, including Walckenaeria, Spirembolus and Eperigone, and described over 100 species. Buckle et al. (2001) provides an annotated catalog of North American taxa, and Hormiga et al. (2003) contains illustrations of all valid linyphiid genera worldwide. Paquin & Duprr (2003) provide illustrations of all species known from Quebec. This an invaluable reference for anyone working on linyphiids from northern and eastern North America. For a significant number of species it provides the first good illustrations or the first illustration of previously undocumented males or females. The higher level classification of the linyphiids is not presently clear. While the subfamily Erigoninae is considered to be monophyletic (Hormiga 2000) the Linyphiinae, in its classical sense, is not (Millidge 1984, Hormiga 2000). We evade this problem by considering everything not included in the Erigoninae to be a linyphiine. An additional problem is defining the limits of the Erigoninae.

Typical linyphiines and erigonines can be distinguished by several characters. However, there are a dozen or so genera which possess some but not all of the defining characters, in various assortments. These genera have, over the years, been shuffled back and forth between the subfamilies by various authors. Here we follow Hormiga (2000) who considers most to be basal erigonines. This key includes all linyphiid males and linyphiine females. Insufficient information is available at this time to permit construction of a key to erigonine females. There is no simple, straightforward way to distinguish between female linyphiines and erigonines. Almost all linyphiines have 2 dorsal spines on tibia IV, while many erigonines have only 1 or none. Many linyphiines have prolateral spines on tibia I and/or femur I while most erigonines do not. Many linyphiines have epigynal scapes, while most erigonines do not. Erigonines are small, while linyphiines may be small or medium sized; any linyphiid over 3.5 mm is likely to be a linyphiine. Species with patterned abdomens are common in the Linyphiinae, unusual in the Erigoninae. Females of the larger linyphiine genera have palpal claws while those of the erigonines and smaller linyphiines do not. Clitolyna electa and Masikia are not included in the key because males are unknown. The male of Pacifiphantes magnificus is unknown so Pacifiphantes is omitted from the male section of the key. Styloctetor keys out with Ceratinopsis because we were unable to find a way to consistently separate the two taxa. We key out two common western species which probably form an undescribed genus (Spirembolus vasingtonus, nomen dubium and Collinsia wilburi). We also include an undescribed western genus containing Spirembolus approximatus and other undescribed species, currently being examined and defined by J. Miller and T. Prentice, and another containing Linyphia rita, Linyphia catalina and Linyphia tauphora. Generic treatment here differs in various small ways from the current World Catalog. Meioneta is considered to be a synonym of Agyneta, following Saaristo (1973), Collinsia is treated as a synonym of Halorates (see Buckle et al. 2001) and Nanavia monticola CHAMBERLIN & IVIE, which was erroneously synonymized with Leptorhoptrum robustum (Westring 1851) by Eskov and Marusik (1994), has been restored to Nanavia. The recent deconstruction of Lepthyphantes into many small genera by Tanasevitch and Saaristo generated serious problems in the creation of this key. Only about half of the American Lepthyphantes species have yet been placed in the new genera, and some of the genera are based on characters possessed by one sex only. The only viable solution was to retain Lepthyphantes in the key in its old, broad sense. Students wishing to apply the Tanasevitch & Saaristo names can follow an identification technique long used for American linyphiids -thumb through the literature, looking at illustrations, until the species, or a close relative is located, then check a catalog to determine its current placement. Definitions used in this key The following characters are useful for separating the vast number of species into smaller, more manageable groups. However, using these requires separating continuously varying characters into discrete groups, and is necessarily a somewhat subjective process. We provide the following precise definitions to minimize the subjectivity. Some questionable taxa also key out in more than one place, when the interpretation of these characters is inherently difficult.

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Metatarsal trichobothria: A trichobothrium (Tm) is a long, very fine hair emerging from a pit (which looks like a distinct circle) on the dorsal surface of a leg. They often stand nearly perpendicular to the leg surface. Linyphiids have 1 trichobothrium on each metatarsus, but it may be absent from metatarsus IV. The relative position of TmI is sometimes useful as a character; we indicate this position as the decimal fraction of a/b, as shown in Fig. 35.2. More accurate measurements will be obtained if the entire metatarsus is in focus under the microscope. See Roberts (1987) for further explanations and figures of trichobothria. Length of palpal tibia: The minimum distance parallel to the long axis of the tibia, between the point where the tibia and patella contact, and the point where the tibia contacts the tarsus (i.e., a palpal structure distal to the tibia, Figs. 35.3-35.5b). Width of palpal tibia: The minimum diameter of the tibia, perpendicular to the long axis of the tibia, in the dorsal or ectal view (Figs. 35.4-35.5b). Length of tibial apophysis: This is determined in dorsal or ectal view, whichever view the apophysis appears longest. In this view, the apophysis length is the straight line distance from the apical-most point to a point which appears to be midway along a line drawn across the base of the apophysis. This base is somewhat subjective, but the point at which the apophysis begins to expand (Figs. 35.3-35.5b).

Normal versus modified cephalic region: The normal or unmodified linyphiid carapace is slightly raised or lobed in linyphiids (Figs. 35.6, 35.20, 35.120, 35.121), but a modified carapace is noticeably more raised (Fig. 35.57, 35.58) or raised in a distinctive way -the angle of the posterior declivity may be more acute than normal in lateral view, for example, producing a relatively sharp (but not particularly high) lobe (Fig. 35.165), or the entire carapace may be somewhat raised, or the posterior declivity of the cephalic region may be in an unusually posterior position, which is also considered modified (Fig. 35.203). Any noticeable protrusion or protuberance on the cephalic or clypeal region is also considered a modified carapace. Pits or excavations of an otherwise normal shaped carapace are not considered a cephalic modification, nor are any spines or projections of the lateral margins of the carapace (e.g., some Erigone species). Note on linyphiid palps: Many characters (such as the size and shape of the tibial apophysis) are best viewed from a certain orientation, and the best orientation often varies between species. Removing a palp from the body allows easy examination from various orientations, and can facilitate identification in many cases. Note that the left palpus is generally illustrated throughout this work, but many of the revisions of American linyphiids illustrate right palpi.

b a

TA

len gt

T length

35.3

35.2

TA

TA length
len gt h

TA length

T length

T length

T length

T width

T width
T width

35.4

35.5b

35.5a

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Genera LINYPHIINAE Agyneta HULL 1911a, Allomengea STRAND 1912b, Anibontes CHAMBERLIN 1924a, Aphileta HULL 1920, Arcuphantes CHAMBERLIN & IVIE 1943, Bathyphantes MENGE 1866, Bolyphantes C.L. KOCH 1837g, Centromerita F. DAHL 1912b, Centromerus F. DAHL 1886, Drapetisca MENGE 1866, Estrandia BLAUVELT 1936, Florinda O. PICKARD-CAMBRIDGE 1896a, Frontinella F.O. PICKARD-CAMBRIDGE 1902a, Graphomoa CHAMBERLIN 1924a, Helophora MENGE 1866, Jalapyphantes GERTSCH & DAVIS 1946, Kaestneria WIEHLE 1956, Lepthyphantes MENGE 1866, Linyphantes CHAMBERLIN & IVIE 1942a, Linyphia LATREILLE 1804b, Macrargus F. DAHL 1886, Maro O. PICKARD-CAMBRIDGE 1906a, Microlinyphia GERHARDT 1928, Microneta MENGE 1869, Neriene BLACKWALL 1833c, Oaphantes CHAMBERLIN & IVIE 1943, Oreonetides STRAND 1901b, Oreophantes ESKOV 1984a, Pacifiphantes ESKOV & MARUSIK 1994, Pityohyphantes SIMON 1929, Poeciloneta KULCZYN SKI in CHYZER & KULCZYN SKI 1894, Porrhomma SIMON 1884a, Saaristoa MILLIDGE 1978b, Stemonyphantes MENGE 1866, Tapinopa WESTRING 1851, Taranucnus SIMON 1884a, Tennesseellum PETRUNKEVITCH 1925c, Wubana CHAMBERLIN 1919a ERIGONINAE Anacornia CHAMBERLIN & IVIE 1933a, Annapolis MILLIDGE 1984a, Anthrobia TELLKAMPF 1844, Arcterigone ESKOV & MARUSIK 1994, Baryphyma SIMON 1884r, Blestia MILLIDGE 1993a, Carorita DUFFEY & MERRETT 1964, Caviphantes OI 1960a, Ceraticelus SIMON 1884r, Ceratinella EMERTON 1882, Ceratinops BANKS 1905a, Ceratinopsidis BISHOP & CROSBY 1930, Ceratinopsis EMERTON 1882, Cheniseo BISHOP & CROSBY 1935c, Cnephalocotes SIMON 1884r, Coloncus CHAMBERLIN 1949, Coreorgonal BISHOP & CROSBY 1935c, Dactylopisthes SIMON 1884r, Dicymbium MENGE 1868, Dietrichia CROSBY & BISHOP 1933b, Diplocentria HULL 1911a, Diplocephalus BERTKAU in FRSTER & BERTKAU 1883, Disembolus CHAMBERLIN & IVIE 1933a, Dismodicus SIMON 1884r, Entelecara SIMON 1884r, Eperigone CROSBY & BISHOP 1928a, Epiceraticelus CROSBY & BISHOP 1931, Eridantes CROSBY & BISHOP 1933b, Erigone AUDOUIN 1826, Erigonella F. DAHL 1901c, Eskovia MARUSIK & SAARISTO 1999, Eulaira CHAMBERLIN & IVIE 1933a, Floricomus CROSBY & BISHOP 1925b, Glyphesis SIMON 1926, Gnathonargus BISHOP & CROSBY 1935c, Gnathonarium KARSCH 1881a, Gnathonaroides BISHOP & CROSBY 1938, Gonatium

MENGE 1868, Goneatara BISHOP & CROSBY 1935c, Gongylidiellum SIMON 1884r, Grammonota EMERTON 1882, Halorates HULL 1911a, Hilaira SIMON 1884a, Horcotes CROSBY & BISHOP 1933b, Hybauchenidium HOLM 1973, Hypomma F. DAHL 1886, Hypselistes SIMON 1894a, Islandiana BRAENDEGAARD 1932, Ivielum ESKOV 1988b, Jacksonella MILLIDGE 1951, Lessertia SMITH 1908, Lophomma MENGE 1868, Masikia MILLIDGE 1984a, Maso SIMON 1884r, Masoncus CHAMBERLIN 1949, Masonetta CHAMBERLIN & IVIE 1939a, Mecynargus KULCZYN SKI in CHYZER & KULCZYN SKI 1894, Metopobactrus SIMON 1884r, Micrargus F. DAHL 1886, Microctenonyx F. DAHL 1886, Mythoplastoides CROSBY & BISHOP 1933b, Nanavia CHAMBERLIN & IVIE 1933a, Oedothorax BERTKAU in FRSTER & BERTKAU 1883, Origanates CROSBY & BISHOP 1933b, Ostearius HULL 1911a, Paracornicularia CROSBY & BISHOP 1931, Pelecopsidis BISHOP & CROSBY 1935b, Pelecopsis SIMON 1864, Perregrinus TANASEVITCH 1992, Perro TANASEVITCH 1992, Phanetta KEYSERLING 1886b, Pocadicnemis SIMON 1884r, Procerocymbium ESKOV 1989b, Satilatlas KEYSERLING 1886b, Savignia BLACKWALL 1833a, Sciastes BISHOP & CROSBY 1938, Scirites BISHOP & CROSBY 1938, Scironis BISHOP & CROSBY 1938, Scolopembolus BISHOP & CROSBY 1938, Scotinotylus SIMON 1884r, Scylaceus BISHOP & CROSBY 1938, Scyletria BISHOP & CROSBY 1938, Semljicola STRAND 1906a, Silometopoides ESKOV 1990a, Sisicottus BISHOP & CROSBY 1938, Sisicus BISHOP & CROSBY 1938, Sisis BISHOP & CROSBY 1938, Sisyrbe BISHOP & CROSBY 1938, Sitalcas BISHOP & CROSBY 1938, Smodix BISHOP & CROSBY 1938, Soucron CROSBY & BISHOP 1936b, Souessa CROSBY & BISHOP 1936b, Souessoula CROSBY & BISHOP 1936b, Sougambus CROSBY & BISHOP 1936b, Souidas CROSBY & BISHOP 1936b, Soulgas CROSBY & BISHOP 1936b, Sphecozone O. PICKARD-CAMBRIDGE 1870d, Spirembolus CHAMBERLIN 1920a, Styloctetor SIMON 1884r, Subbekasha MILLIDGE 1984a, Symmigma CROSBY & BISHOP 1933b, Tachygyna CHAMBERLIN & IVIE 1939a, Tapinocyba SIMON 1884r, Thaleria TANASEVITCH 1984a, Thyreosthenius SIMON 1884r, Tibioplus CHAMBERLIN & IVIE 1947b, Tiso SIMON 1884r, Tmeticus MENGE 1868, Traematosisis BISHOP & CROSBY 1938, Tunagyna CHAMBERLIN & IVIE 1933a, Tusukuru ESKOV 1993, Tutaibo CHAMBERLIN 1916, Typhochrestus SIMON 1884r, Vermontia MILLIDGE 1984a, Wabasso MILLIDGE 1984a, Walckenaeria BLACKWALL 1833a, Zornella JACKSON 1932a, Zygottus CHAMBERLIN 1949

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Key to genera North America North of Mexico

2(1)

Male ................................................................................... 2 Female (LINYPHIINAE only) ............................................ 197 With 2 dorsal spines on each tibia, or 1 dorsal spine on tibia IV and a prolateral spine on femur I; usually without a retrolateral apophysis on palpal tibia ........................... 3 With 1 or no dorsal spines on tibia IV and no prolateral spine on femur I; usually with a retrolateral apophysis on palpal tibia ...................................................................... 51

Linyphiinae and larger Erigoninae portion

3(2)

4(3)

Prolateral spine present on tibia I .................................... 4 No prolateral spine on tibia I ......................................... 37 Some or all metatarsi with spines (check all legs) .......... 6 No metatarsi with spines .................................................. 5 Femur I with prolateral spine ........................................ 25 Femur I without prolateral spine ................................... 28 Abdomen with conspicuous caudal tubercle (Fig. 35.6); body red (fading in alcohol) except for palps, eye region and caudal tubercle which are black; palp as shown (Fig. 35.7) .................................................................... Florinda
Div. 1 species: Florinda coccinea (HENTZ 1850a) Dist. southeastern USA Ref. Blauvelt 1936

35.7 35.8 35.9

5(4)

6(4)

35.11 35.10

Abdomen without caudal tubercle; color brown or gray .. ............................................................................................ 7 Carapace pale with dark stripe bifurcating anteriorly (Figs. 35.8-35.9) ................................................................ 8 Carapace markings otherwise .......................................... 9 Dorsal stripe on carapace narrow (Fig. 35.8); terminal apophysis of palp twisted in a thick, tight spiral (Fig. 35.10); palp without patellar apophysis ........... Linyphia
Div. 1 introduced species: Linyphia triangularis (CLERCK 1757) Dist. ME Refs. Helsdingen 1969, Jennings et al. 2003

7(6)

8(7)

Dorsal stripe wider (Fig. 35.9); terminal apophysis not spiral (Fig. 35.11); palp with large patellar apophysis (Fig. 35.12) ................................................... Pityohyphantes
Div. 16 species Dist. n and w NA Ref. Buckle et al. 2001

35.13

35.12

9(7)

Cymbium of palp with dorso-basal horn tipped with tuft of hairs (Figs. 35.13-35.14) ........................ Allomengea
Div. 3 species Dist. n NA Ref. Helsdingen 1974 Note 2 of the species have matatarsal spines only on legs III & IV

10(9)

Cymbium otherwise ....................................................... 10 Embolus forming an apically directed loop (Fig. 35.15); lamina large and shield-like ...... Bathyphantes, in part
Div. 26 species, 2 key out here: Bathyphantes keeni (EMERTON 1917a) and B. reprobus (KULCZYNSKI 1916) Dist. widespread Ref. Ivie 1969

Not so .............................................................................. 11

35.14

35.15

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11(10)

12(11)

Metatarsi with ventral spines ......................................... 12 Metatarsi without ventral spines ................................... 17 Tibia I with 2-5 pairs of ventral spines (Fig. 35.16) ...... 13 Tibia I with one pair of ventral spines or 1 to 4 single ones (Fig. 35.17) ...................................................................... 16 Terminal apophysis of palp twisted in a thick, tight spiral (Fig. 35.18) ........................................................... Neriene Terminal apophysis not spiral ........................................ 14 Embolus long and threadlike, making loop beyond confines of cymbium (Fig. 35.19); carapace elongate in cephalic region; chelicerae elongate, angled posteriorly (Fig. 35.20) ....................................................... Microlinyphia
Div. 4 species Dist. n and w NA Ref. Helsdingen 1970

35.16

35.17

13(12)

Div. 9 species Dist. widespread Ref. Helsdingen 1969 Note some authors consider Neriene to be a synonym of Linyphia

14(13)

Embolus remaining within confines of cymbium (Figs. 35.21-35.22), carapace and chelicerae normal .............. 15 Carapace pale with black median and marginal stripes: metatarsus I swollen (Fig. 35.23); palp as shown (Fig. 35.21) ................................................. Stemonyphantes
Div. 1 species: Stemonyphantes blauveltae GERTSCH 1951 Dist. n NA Ref. Helsdingen 1968

35.18

35.19

15(14)

35.20

Carapace uniform brown; metatarsus I not swollen; palp as shown (Fig. 35.22)...................... Undescribed genus
Div. Linyphia catalina GERTSCH 1951, Linyphia rita GERTSCH 1951 and perhaps Linyphia tauphora CHAMBERLIN 1928a Dist. sw USA Ref. Gertsch 1951

16(12)

Tibia I with pair of ventral spines near middle; palp with lamella of embolic division large and U-shaped (Fig. 35.24), Fickerts gland absent ........................ Helophora
Div. 4 species Dist. n and w NA Ref. Buckle et al. 2001

Tibia I with single ventral spine near base or 2-4 scattered ventral spines (Fig. 35.17); palp with lamella otherwise, Fickerts gland present (Fig. 35.25) ..................................... .................................................... Lepthyphantes, in part
Div. 29 species + several undescribed: a few of the larger species key out here Dist. widespread Refs. Zorsch 1937, Buckle et al. 2001

35.21

35.22

35.24

17(11)

AME larger than other eyes; carapace low, about height of AME; anterior eye row strongly recurved (Fig. 35.26); cymbium with long, angled basal apophysis (Figs. 35.2735.28) ................................................................ Tapinopa
Div. 2 species: Tapinopa bilineata BANKS 1893a, Tapinopa hentzi GERTSCH 1951 Dist. eastern NA Refs. Kaston 1981, Gertsch 1951

35.23

AME same size or smaller than other eyes; clypeus height more than twice diameter of AME; anterior eye row nearly straight (Fig. 35.29); cymbium otherwise ....................... 18

35.25

35.27 35.26

35.29

35.28

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18(17)

Prolateral face of chelicerae with 3 or 4 pairs of stout spines (Fig. 35.30), cheliceral teeth long; trichobothrium on leg IV present; palp as shown (Figs. 35.31-35.32) ........ ......................................................................... Drapetisca
Div. 2 species: Drapetisca alteranda CHAMBERLIN 1909a, Drapetisca oteroana GERTSCH 1951 Dist. n NA, NM Refs. Kaston 1981, Gertsch 1951

Prolateral face of chelicerae without stout spines; cheliceral teeth relatively short; trichobothrium on leg IV absent .......................................................................................... 19 Tibia I with ventral spines .............................................. 20 Tibia I with no ventral spines ........................................ 23 Tibia I with 3-5 pairs of ventral spines (Fig. 35.33); bulge on base of palpal tibia with brush of hairs (Fig. 35.34); palp as shown (Figs. 35.35-35.36); abdomen concolorous .................................................................... Centromerita
Div. 1 introduced species: Centromerita bicolor (BLACKWALL 1833c) Dist. NF, NS, BC, WA Ref. Roberts 1987

35.30

35.32

35.31
19(18)

20(19)

35.33

35.34

Tibia with 1 or 2 pairs of spines or a few scattered ones; abdomen patterned or concolorous .............................. 21 Palp with distinctive embolic division (Fig. 35.37); paracymbium with simple U-shape, distal arm very thin; abdomen with dorsal folium or uniformly black (Fig. 35.38) ................................................................ Estrandia
Div. 1 species: Estrandia grandaeva (KEYSERLING 1886b) Dist. n NA Ref. Blauvelt 1936

35.35

21(20)

35.36

35.37

22(21)

Palp and paracymbium otherwise ................................. 22 Fang often stout and sinuous (Fig. 35.39); distal arm of paracymbium often with bifid tip (Fig. 35.40); Fickerts gland present (Fig. 35.41) ......................... Arcuphantes
Div. 7 species Dist. w NA Ref. Chamberlin & Ivie 1943

35.38

Fang normal; distal arm of paracymbium without bifid tip; Fickerts gland absent; palp as shown (Figs. 35.4235.43) ........................................................ Oreophantes
Div. 1 species: Oreophantes recurvatus (EMERTON 1913a) Dist. n NA Refs. Helsdingen 1973b, 1981a

35.39

35.40

23(19)

With short, stout legs (femur I no longer than carapace); basal segment of chelicera with anteriolateral row of short bristles (Fig. 35.44); body concolorous brown, palp as shown (Figs. 35.45-35.46) ......................... Centromerus
Div. 9 species Dist. e and n NA Ref. Helsdingen 1973c

35.41

With longer, thinner legs (femur I 1.1 to 1.5 times length of carapace); basal segment of chelicera without row of bristles; abdomen frequently patterned ........................ 24
35.44

35.42 35.43

35.45

35.46

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24(23)

Palp with long curved embolus supported by a large, broad suprategular apophysis (Fig. 35.47); simple elbowed paracymbium; Fickerts gland absent .......... Taranucnus
Div. 1 species: Taranucnus ornithes (BARROWS 1940) Dist. e NA from QC s to NC and TN Ref. Helsdingen 1973b

Palp with embolus and suprategular apophysis otherwise (Fig. 35.48); paracymbium usually more complex, with teeth (Fig. 35.49); Fickerts gland present (Fig. 35.25) ...... ................................................... Lepthyphantes, in part
Div. 29 species + several undescribed: most of the described species key out here Dist. widespread Refs. Zorsch 1937, Buckle et al. 2001

35.47

35.48

35.49

25(5)

Carapace with medial dark shield-shaped mark (Fig. 35.50); legs long, femur I significantly longer than carapace; metatarsus IV with trichobothrium; palp as shown, similar to Lepthyphantes (Figs. 35.51-35.52) ...................... ....................................................................... Poeciloneta
Div. 10 species Dist. n and w NA Ref. Buckle et al. 2001

Carapace without dark shield-shaped mark; legs shorter, femur I no longer than carapace; metatarsus IV without trichobothrium ............................................................... 26 Embolus forming distal-facing coil (Figs. 35.53-35.54) .... ...................................................... Bathyphantes, in part
Div. 26 species, 24 key out here Dist. widespread Ref. Ivie 1969

26(25)

35.50

35.51

35.52

27(26)

Embolus otherwise (Figs. 35.55-35.56) ......................... 27 Embolus curved and coming to fine point, pointing ventrad (Fig. 35.55) ........................................... Porrhomma
Div. 9 species Dist. widespread Ref. Buckle et al. 2001

Embolus straight and thick with blunt tip, pointing anteriad (Fig. 35.56) ............................................. Kaestneria
Div. 3 species Dist. n NA Ref. Ivie 1969

28(5)

29(28)

Carapace elevated behind eyes (Figs. 35.57-35.58) ....... 29 Carapace not elevated behind eyes ................................ 30 Trichobothria on metatarsus IV absent; palpal tibia with distinctive branched tibial apophysis (Figs. 35.59), palp as in Fig. 35.60 .................................................. Sougambus
Div. 2 species: Sougambus bostoniensis (EMERTON 1882) and Erigone mentasta CHAMBERLIN & IVIE 1947b Dist. n NA Refs. Crosby & Bishop 1936b, Hackman 1954

35.53

35.54

35.55

35.57

Trichobothria on metatarsus IV present; tibial apophysis otherwise (Fig. 35.61), palp as in Fig. 35.62 ......... Hilaira

Div. 24 species Dist. n NA Refs. Buckle et al. 2001, Saaristo & Marusik 2004a Note Saaristo & Marusik (2004a) placed 15 North American species, 10 of them new, in Oreoneta KULCZYNSKI in CHYZER & KULCZYNSKI 1894. They suggested that, of the remaining species, a few closely related to the type should be left in Hilaira and the remaining ones placed in new genera

35.56

35.58

35.59

35.61

35.60

35.62

132

Linyphiidae

30(28)

Palpal tibia with stout, thick, blunt tipped dorsal spine (Fig. 35.63); paracymbium a rectangular plate fused to cymbium (Fig. 35.64); palp with small cymbium; embolic division a flat plate, narrowed basally, and with a short, straight embolus extending distad (Fig. 35.65) ................. ......................................................................... Frontinella
Div. 2 species Dist. Frontinella pyramitela (WALCKENAER 1841), widespread and Frontinella huachuca GERTSCH & DAVIS 1946, AZ Refs. Blauvelt 1936, Gertsch & Davis 1946 Note Frontinella pyramitela is referred as Frontinella communis (HENTZ 1850a) in Platnick (2005)

35.63 35.64

Palpal tibia without stout, thick, blunt tipped dorsal spine; paracymbium U-shaped sclerite, palp otherwise ......... 31 Embolus an anteriad facing spiral (Fig. 35.66) .................. ...................................................................... Linyphantes Embolus otherwise ......................................................... 32
35.66

35.65

31(30)

Div. 19 species Dist. Pacific northwest Ref. Chamberlin & Ivie 1942a

32(31)

Carapace broad; abdomen dark with paired light spots (Fig. 35.67); prolateral spine on palpal tibial large and thick (Fig. 35.68) ......................................... Graphomoa
Div. 1 species: Graphomoa theridioides CHAMBERLIN 1924a Dist. LA, TN Ref. Chamberlin 1924a Note according to Gustavo Hormiga (pers. comm.) Graphomoa theridioides is a synonym of Pocobletus coroniger SIMON 1894a and Graphomoa a synonym of Pocobletus SIMON 1894a

35.67 35.68

33(32)

Without the above combination of characters ............. 33 Long threadlike embolus extending laterally far past the edge of the cymbium (Fig. 35.69); palpal tibia with ventral bump tipped with tuft of hairs (Fig. 35.70) ....................... .................................................................. Jalapyphantes Without the above combination of characters ............. 34
35.69

Div. 1 species: Jalapyphantes cuernavaca GERTSCH & DAVIS 1946 Dist. AZ Ref. Gertsch & Davis 1946

34(33)

35.70

Trichobothrium present on tibia IV, palp as shown (Figs. 35.71-35.72) .................................................... Microneta
Div. 1 species: Microneta viaria (BLACKWALL 1841) Dist. n and w NA Refs. Kaston 1981, Saaristo 1974a, Paquin & Duprr 2003

No trichobothrium present on tibia IV; palp otherwise (Fig. 35.73) ...................................................................... 35

35(34)

Embolic division with radix (R), embolus (E) and lamella (L) not fused (Figs. 35.76-35.77) ......... Agyneta, in part

Div. 45 species and perhaps as many more undescribed: a few western species key out here Dist. widespread Refs. Buckle et al. 2001, Paquin & Duprr 2003 Note some authors treat Meioneta HULL 1920 as a distinct subgenus or genus. Agyneta sensu stricto has a trichobothrium on metatarsus IV while Meioneta does not

Embolic division of palp a compact unit with fused sclerites (Figs. 35.73, 35.74, 35.75) ......................................... 36

35.72

35.73

35.74

E R L L

35.75

35.76

35.77

Spiders of North America

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133

36(35)

Palp as shown (Figs. 35.74 -35.75, 35.78-35.79) ................ ......................................................... Oreonetides, in part Palp as shown (Figs. 35.73, 35.80) ................... Saaristoa

Div. 6 species + several undescribed: 3 described species key out here Dist. n and w NA Ref. Helsdingen 1981a Div. 1 species: Saaristoa sammamish (LEVI & LEVI 1955) Dist. nw NA Ref. Levi & Levi 1955

37(3)

38(37)

Chelicerae with mastidion (Figs. 35.81-35.82) ............. 38 Chelicera without mastidion ......................................... 42 Palpal tibia with a long straight dorsal spur (Fig. 35.83) .. ............................................................................ Tibioplus

35.78

35.80 35.79

Div. 1 species: Tibioplus nearcticus CHAMBERLIN & IVIE 1947b Dist AK, YT Ref. Chamberlin & Ivie 1947b

39(38)

Not so .............................................................................. 39
35.81
35.82
35.83

Abdomen red (fading to pale brown in alcohol) with black rump (Fig. 35.84); palp as shown (Figs. 35.85-35.86) ....... ........................................................................... Ostearius

Div. 1 introduced species: Ostearius melanopygius (O. PICKARD-CAMBRIDGE 1879d) Dist. ne NA Refs. Millidge 1985, Roberts 1987

Abdomen brown or gray, usually concolorous; palp different .................................................................................. 40 Endite with single large setose tubercle (Fig. 35.87); legs relatively long, femur I length 1 1/3 times that of carapace; leg spines short and thin, barely distinguishable from setae; eyes small, especially AME; carapace higher than ocular area; palp as shown (Figs. 35.88-35.89) .................. ......................................................................... Oaphantes
Div. 1 species: Oaphantes pallidulus (BANKS 1904b) + 2 undescribed Dist. CA to BC Ref. Chamberlin & Ivie 1943

40(39)

35.84 35.85 35.86

Endite with several tiny setaceous tubercles or none; legs shorter, femur I no longer than carapace; leg spines of normal size; eyes larger; carapace no higher than ocular area .................................................................................. 41 Mastidion located anteriomesally on chelicera, above anterior teeth (Fig. 35.81); cephalothorax reddish, abdomen charcoal; epigastric plates not striate; palp as shown (Figs. 35.90-35.91) ......................................... Macrargus
Div. 1 species: Macrargus multesimus (O. PICKARD-CAMBRIDGE 1875c) Dist. n NA Ref. Kaston 1981

41(40)

35.87 35.88 35.89

Mastidion located anteriolaterally on chelicera, just in front of stridulating ridges (Fig. 35.82); cephalothorax amber, abdomen brownish gray; epigastric plates striate (Fig. 35.92); palp as shown (Figs. 35.93-35.94) ................. ...................................................................... Maro, in part
Div. 2 species, 1 keys out here: Maro amplus DONDALE & BUCKLE 2001 Dist. n NA Ref. Dondale & Buckle 2001

35.90

35.91

35.92
35.93

35.94

134

Linyphiidae

42(37)

Carapace elevated behind ocular area to a point which is adorned with one or more forward curving spines (Fig. 35.95); paracymbium with long basally extending branch (Fig. 35.96); palpal tibia modified (Fig. 35.96); palp as shown (Figs. 35.96-35.97) .................................. Wubana
Div. 7 species Dist. n and w NA Refs. Chamberlin & Ivie 1936b, Buckle et al. 2001

35.95

Carapace not elevated; paracymbium of usual U-shape, palpal tibia unmodified .................................................. 43
35.96 35.97

43(42)

Clypeus bulging (Fig. 35.98); embolus long and straight, extending from base to apex of cymbium (Fig. 35.99) ...... .............................................................................. Thaleria
Div. 1 species: Thaleria leechi ESKOV & MARUSIK 1992a Dist. AK Ref. ESKOV & MARUSIK 1992a

35.98

44(43)

Clypeus normal, embolus otherwise ............................. 44 Chelicerae with anterio-lateral row of denticles (Fig. 35.100); carapace narrowed anteriorly (Fig. 35.101); antmimic appearance .......................................... Anibontes
Div. 2 species: Anibontes longipes CHAMBERLIN & IVIE 1944 and Anibontes mimus CHAMBERLIN 1924a Dist. se USA Refs. Chamberlin 1924a, Chamberlin & Ivie 1944

35.100

45(44)

Chelicerae and carapace normal .................................... 45 Palpal femur with ventral teeth (Fig. 35.102), spiracle forward by 1/4 to 1/3 distance between spinnerets and epigastric furrow (Fig. 35.103); palp is similar to that of Agyneta (Figs. 35.102, 35.104) .............. Tennesseellum Palpal femur without ventral teeth, spiracle just in front of spinnerets (Fig. 35.105) .................................................. 46
35.102

35.99

35.101

Div. 2 species Dist. Tennesseellum formicum (EMERTON 1882) widespread and an undescribed species from CA Ref. Kaston 1981

46(45)

Tiny spiders (1.0-1.5 mm) with large, distinctive palps (Figs. 35.106-35.107) having complex embolic divisions .. ................................................................................. Sisicus

35.103

Div. 2 species: Sisicus apertus (HOLM 1939b) and Sisicus penifusifer BISHOP & CROSBY 1938 Dist. n NA Ref. Paquin & Duprr 2003

47(46)

Without the above combination of characters ............. 47 Palp with distinctive paracymbium (Fig. 35.108) and simple embolic division (Fig. 35.109) .............. Aphileta Without the above combination of characters ............. 48

35.104

Div. 1 species: Aphileta misera (O. PICKARD-CAMBRIDGE 1882b) Dist. n NA Refs. Bishop & Crosby 1938, Paquin & Duprr 2003

35.105

35.106

35.107

35.108

35.109

Spiders of North America

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135

48(47)

Palpal bulb and cymbium relatively small (Fig. 35.110); embolic division a simple plate (Fig. 35.111); palpal tibia one to two times length of cymbium; abdomen with dark heart mark and posterior chevrons .................... Nanavia
Div. 1 species: Nanavia monticola CHAMBERLIN & IVIE 1933a + 1 undescribed Dist. UT, CA Ref. CHAMBERLIN & IVIE 1933a

Palpal bulb and cymbium larger (Figs. 35.112, 35.113); embolic division complex; palpal tibia shorter than cymbium; abdomen concolorous ......................................... 49 Palpal bulb with distinct arrangement of embolus and radix (Figs. 35.112, 35.114); abdomen concolorous or patterned ............................................... Agyneta, in part
Div. 45 species and perhaps as many more undescribed: most key out here Dist. widespread Refs. Buckle et al. 2001, Paquin & Duprr 2003 Note some authors treat Meioneta HULL 1920 as a distinct subgenus or genus. Agyneta sensu stricto has a trichobothrium on metatarsus IV while Meioneta does not

49(48)

35.111

35.112

Palp otherwise (Figs. 35.115, 35.113, 35.116, 35.117); abdomen concolorous .................................................... 50
35.110

50(49)

Body under 1.7 mm in length; stridulating ridges on epigastric plates (Fig. 35.118); palp as shown (Figs. 35.113, 35.115) ........................................................ Maro, in part
Div. 2 species, 1 keys out here: Maro nearcticus DONDALE & BUCKLE 2001 Dist. n NA Ref. Dondale & Buckle 2001

Body usually over 1.7 mm in length; no stridulating ridges on epigastric plates; palp as shown (Figs. 35.116-35.117) . ........................................................ Oreonetides, in part

35.113

35.114

Div. 6 species + several undescribed: 3 key out here Dist. n and w NA Ref. Helsdingen 1981a

35.115

35.116

35.118 35.117

136

Linyphiidae

Erigoninae portion, 107 genera

With a pair of distinct pits or excavations on the cephalothorax, usually just behind PLE (Fig. 35.119) ............ 52 Without distinct pits on the cephalothorax (Figs. 35.12035.121) ............................................................................ 53 Cephalic region raised, or modified in the eye or clypeus region with one or more fairly distinct lobes, large setae or processes (Figs. 35.122-35.126) ..................................... 54 Cephalic region normal (Figs. 35.120-35.121, although these illustrations lack cephalic pits) unmodified although the cephalic region is commonly slightly higher than the posterior portion of the cephalothorax, and some species have spinose carapace margins ...................................... 93 Cephalic region raised, or modified in the eye or clypeus region with one or more fairly distinct lobes, large setae or processes (Figs. 35.127-35.131) ..................................... 98 Cephalic region normal, unmodified (Figs. 35.12035.121) although some species have spinose carapace margins ......................................................................... 135
With pits and modified cephalic region, 41 genera

35.119

35.120

35.121

52(51)

35.122

35.123

35.124

35.123 35.125 35.126

53(51)

35.127

35.128

35.129

54(52)

Abdomen with sclerotized scutum or shield (Fig. 35.132); reduced in some species to a small region at anteriormost portion of abdominal dorsum........................ Pelecopsis
Div. 6 species Dist. widespread except southwest USA Refs. Buckle et al. 2001, Paquin & Duprr 2003

35.130

35.131

55(54)

Without a scutum ........................................................... 55 Cephalic region with both a clypeal and a distinct cephalic lobe or protuberance (Figs. 35.122, 35.133) ................. 56 Cephalic region with either a clypeal or one or two cephalic lobes, which may be quite small (Figs. 35.125-35.126, 35.134) ............................................................................ 57
35.132

35.133

56(55)

Cephalic lobe produced into a secondary protuberance that lies directly over the clypeal protuberance, giving the impression of stacked protuberances (Fig. 35.122); embolic division lacking obvious spiraled tailpiece .................... ............................................................................. Savignia
Div. 1 species: Savignia birostra (CHAMBERLIN & IVIE 1947b) Dist. Widespread except southwest USA Ref. Chamberlin & Ivie 1947b

35.134

Cephalic lobe without a secondary protuberance and not stacked over the clypeal protuberance (Fig. 35.133); embolic division with obvious spiraled tailpiece (at least half as long as tegulum, T in Fig. 35.135) ........................... ........................................................ Scotinotylus, in part
Div. 34 species, 5 key out here Dist. CAN, n and w USA Refs. Millidge 1981a, Paquin & Duprr 2003

T
57(55)

With cephalothoracic lobe mostly posterior of eyes (not bearing eyes, Fig. 35.136) OR with pits (not just excavated area surrounding pit) distinctly larger than PME (Fig. 35.138) ............................................................................ 58 Lobe not mainly posterior to eyes (often bearing one or more pairs) AND with pits smaller than PME, although surrounding excavation may be larger than PME ........ 64

35.136

35.135

Spiders of North America

Linyphiidae

137

58(57)

Cephalothorax and legs orangish or reddish and abdomen black ................................................................ Hypomma Body coloration otherwise ............................................. 59

Div. 3 species Dist. CAN, AK, n USA Refs. Crosby & Bishop 1933b, Chamberlin & Ivie 1947b, Paquin & Duprr 2003

59(58)

Carapace with reticulate texture (like lines scratched with a needle), visible without highest magnification (visible at 30X); lobe mostly posterior of eyes and pits of normal size ................................................... Tapinocyba, in part
Note 2 undescribed species from CA (T. Prentice, pers. comm.)

Carapace smooth (except under very high magnification) ......................................................................................... 60 With cephalic lobe mostly posterior of eyes and pits twice or more the diameter the PME (Fig. 35.137) ..................... ...................................................................... Oedothorax

35.137

60(59)

cavity

Div. 7 species Dist. widespread Refs. Buckle et al. 2001, Paquin & Duprr 2003 Note many N. American Oedothorax probably do not belong in that genus

61(60)

With either of the above characters ............................... 61

Embolic division lacking spiraled tailpiece; cavity surrounding pit usually more than 3 times larger than diameter of PME (Fig. 35.138) .................... Hybauchenidium
Div. 4 species Dist. AK, CAN, northern USA Ref. Buckle et al. 2001, Paquin & Duprr 2003

Embolic division with obvious spiraled tailpiece (at least half as long as tegulum); cavity surrounding pit less than 3 times the diameter of PME; usually with lobe mostly posterior of eyes ............................................................. 62 Tibial apophysis with thick basoectal macroseta (Fig. 35.139); tip of embolus retroverted or pointing more or less toward proximal end of palp (Fig. 35.139) .................. .......................................................... Disembolus, in part
Div. 24 species, 9 key out here Dist. widespread, more common in west Ref. Millidge 1981b

62(61)

35.139

Tibial apophysis lacking thick basoectal macroseta; tip of embolus not retroverted, often terminates in wide coil at end of palp ...................................................................... 63 Tibal apophysis more than twice as long as length of tibia; paracymbium elongate ................. Spirembolus, in part
Div. 43 species, 4 key out here Dist. western NA Ref. Millidge 1980b

63(62)

Tibial apophysis less than twice the length of the tibia; paracymbium of normal length, c-shaped ......................... ........................................................ Scotinotylus, in part
Div. 34 species, 12 key out here Dist. CAN, n and w USA Refs. Millidge 1981a, Paquin & Duprr 2003 Note some members of an undescribed genus (see couplet 66) will key out here as well, including Spirembolus approximatus CHAMBERLIN 1949 and four undescribed species

35.140

35.141

64(57)

65(64)

Apical portion of tibial apophysis pointing ectally (Figs. 35.140-35.142) ................................................................ 65 Tibial apophysis otherwise ............................................. 68 Tibial apophysis with one apical portion, which points to the ectal side (Fig. 35.140) ........ Diplocephalus, in part Tibial apophysis with two or more apical portions which point to the ectal side (Figs. 35.141-35.142) ................. 66

Div. 4 species, 3 key out here Dist. CAN, AK, northern USA Ref. Buckle et al. 2001, Paquin & Duprr 2003

35.142

138

Linyphiidae

66(65)

Embolus coiled at end of palp; tailpiece spiraled .............. ............................................ undescribed genus, in part Without the above combination of characters ............. 67

Div. Ceratinopsis palomara CHAMBERLIN 1949 + undescribed species Dist. CA and OR Ref. Chamberlin 1949

67(66)

Embolus longer than cymbium, forms nearly a complete coil (Fig. 35.143) ................................................ Horcotes
Div. 3 species Dist. widespread Refs. Buckle et al. 2001, Paquin & Duprr 2003

Embolus shorter than cymbium, curves but does not form a coil (Fig. 35.144) ......................................... Entelecara

Div. 3 species Dist. CAN, AK, WA Refs. Buckle et al. 2001, Paquin & Duprr 2003

68(64)

69(68)

Tibial apophysis definitely longer than tibial length (Figs. 35.145, 35.148) ............................................................... 70 Tibial apophysis not longer than tibial length .............. 69 Tibial apophysis definitely shorter than width of narrowest part of tibia (Figs. 35.146-35.147) ..................... 78 Tibial apophysis not shorter than width of narrowest part of tibia (Figs. 35.149-35. 150) ........................................ 86 Tibial apophysis with conspicuous dorsoectal row of 3 or more long, thick macrosetae (Fig. 35.151) ......................... ........................................................................... Glyphesis

35.143

35.144

35.145

70(68)

Div. 2 species: Glyphesis idahoanus (CHAMBERLIN 1949) and Glyphesis scopulifer (EMERTON 1882) Dist. CAN and n USA Refs. Buckle et al. 2001, Paquin & Duprr 2003

35.146 35.148

Tibial apophysis lacking conspicuous row of macrosetae . .......................................................................................... 71

71(70)

Cephalic lobe bearing eyes terminates in a rather sharp, forward-pointing cone (Fig. 35.152) ........... Origanates
Div. 1 species: Origanates rostratus (EMERTON 1882) Dist. Eastern NA Ref. Crosby & Bishop 1933b

72(71)

Cephalic lobe, if present, is rounded ............................. 72 Embolus very long, forming a large coil and a smaller terminal coil (Fig. 35.153); tibial apophysis clearly bifid with two long sharp points ............................... Traematosisis
Div. 1 species: Traematosisis bispinosus (EMERTON 1911) Dist. Eastern NA Ref. Bishop & Crosby 1938

35.149 35.151 35.150

Embolus not forming a large and small double coil; tibial apophysis otherwise ....................................................... 73 Tibial apophysis consists of one long process (Fig. 35.145) which terminates in 2-3 sharp points at extreme apical end (not visible in Fig. 35.145) .............................. Dietrichia
Div. 1 species: Dietrichia hesperia Crosby & Bishop 1933b Dist. CAN, CA Ref. Crosby & Bishop 1933b

73(72)

Tibial apophysis otherwise ............................................. 74

35.152

35.153

Spiders of North America

Linyphiidae

139

74(73)

Tibial apophysis with 3 apical points (Fig. 35.154) OR embolus much longer than cymbium, making a spiral around entire palpal bulb, its long thin apical end held away from rest of palp (Fig. 35.155) ................................... .......................................................... Baryphyma, in part
Div. 4 species, 2 key out here: Baryphyma gowerense (LOCKET 1965) and Baryphyma trifrons affine (SCHENKEL 1930b) Dist. CAN, n USA Ref. Roberts 1987, Buckle et al. 2001, Paquin & Duprr 2003

Tibial apophysis with one or two apical points; embolus usually shorter than cymbium, apical end of embolus not as above ........................................................................... 75 Tailpiece of embolic division curved, but does not coil or spiral (Fig. 35.156-35.157); tibial apophysis terminates in one or two apical processes ............................................ 76 Tailpiece of embolic division coils or spirals at least one turn; tibial apophysis terminates in one apical process (though an additional basal process may be present, Fig. 35.158 Spirembolus) ........................................................ 77 Cephalic region with small secondary lobe anterior to main lobe (Fig. 35.159), OR tibial apophysis with two separate apical processes, each longer than tibial width (Fig. 35.160); paracymbium normal, c-shaped; tarsal claws I-III strongly pectinate (Fig. 35.161) ........................ ..................................................... Walckenaeria, in part
Div. 68 species, 2 key out here: Walckenaeria castanea (EMERTON 1882), Walckenaeria atrotibialis (O. PICKARD-CAMBRIDGE 1878a), Dist. widespread [Walckenaeria atrotibialis: CAN, AK, n USA Refs. Millidge 1983, Paquin & Duprr 2003

75(74)

35.154 35.155

76(75)

35.158

35.159

With one cephalic lobe and tibial apophysis not as above; paracymbium reduced, triangular rather than c-shaped (P in Fig. 35.156); tarsal claws not pectinate .......................... .............................................................. Eridantes, in part
Div. 2 species: Eridantes erigonoides (EMERTON 1882) and Eridantes utibilis CROSBY & BISHOP 1933b Dist. eastern NA, AZ Refs. Crosby & Bishop 1933b, Paquin & Duprr 2003 Note both species may key out here

35.156

35.157

77(75)

Tibial apophysis with thick basoectal macroseta; tip of embolus retroverted or pointing more or less toward proximal end of palp (Fig. 35.162) ..................................... .......................................................... Disembolus, in part
Div. 24 species, 11 key out here Dist. widespread, more common in west Ref. Millidge 1981b

35.161 35.160

Tibial apophysis lacking thick basoectal macroseta; tip of embolus not retroverted, often terminates in wide coil at end of palp; sometimes with unusually long paracymbium (Fig. 35.158); sometimes with abdominal pattern of chevrons .............................................. Spirembolus, in part
Div. 43 species, 6 key out here Dist. western NA Ref. Millidge 1980b

78(69)

Cymbium with row of stout tubercles across dorsum; embolus longer than cymbium, emerges toward proximal part of palp and hangs free well ventral of palp (Fig. 35.163) ................................................... Cnephalocotes
Div. 1 species: Cnephalocotes obscurus (BLACKWALL 1834b) Dist. CAN Refs. Roberts 1987, Paquin & Duprr 2003

35.162

Cymbium without row of stout dorsal tubercles; embolus of variable length, but not as above ............................... 79 Eyes in front of rather tall lobe (about as tall as basal width of lobe); pit located directly above PLE (Fig. 35.164); cephalothorax and legs orangish, abdomen black ............. ........................................................................ Dismodicus
Div. 4 species Dist. CAN, n USA Refs. Buckle et al. 2001, Paquin & Duprr 2003

79(78)

Without the above combination of characters ............. 80

35.164

140

Linyphiidae

80(79)

Clypeus hairy (Fig. 35.165) and with a row of long spines on venter of tibia and metatarsus I and II (Fig. 35.166). Often found in wetlands ..................... Satilatlas, in part
Div. 6 species, several may key out here Dist. CAN and n USA to CO Refs. Millidge 1981c, Paquin & Duprr 2003

81(80)

Without this combination of characters ....................... 81

82(81)

Embolus rather short, does not spiral around palp ........... .......................................................................................... 82 Embolus long, spirals around end of palp .................... 84 Carapace and sternum smooth ...... Tapinocyba, in part

Div. 15 species, 10 key out here Dist. widespread Refs. Buckle et al. 2001, Paquin & Duprr 2003

Carapace (and sometimes sternum) at least somewhat textured (rough, wrinkled, or reticulate, as visible even under lower magnification), at least along cervical grooves .................................................................................... 83 Carapace and sternum often with rugose or reticulate texture (visible without highest magnification; embolus usually with a single terminal projection (except C. crenatus, which has undulating lateral margins of carapace, Fig. 35.167); TmI located at 0.40 or less .................................... ........................................................... Ceratinops, in part
Div. 8 species, 6 key out here Dist. widespread Refs. Crosby & Bishop 1933b, Paquin & Duprr 2003

35.166

83(82)

35.167

Carapace usually only textured along cervical grooves; embolus with two terminal projections (Fig. 35.168); TmI located at greater than 0.35, usually greater than 0.40 ...... ........................................................................ Lophomma
Div. 6 species Dist. eastern NA Ref. Crosby & Bishop 1933b, Holm 1960b, Paquin & Duprr 2003

84(81)

Tibial apophysis broader apically than basally (Fig. 35.169) .............................................................. Tusukuru
Div. 1 species: Tusukuru hartlandianus (EMERTON 1913a) Dist. eastern NA Ref. Crosby & Bishop 1933b

85(84)

Tibial apophysis with one or more sharp points .......... 85 Cymbium somewhat Y-shaped in dorsal view, much broader distally (Fig. 35.170); TmI located at 0.85-0.90; TmIV present ....................................................... Pocadicnemis
Div. 3 species Dist. CAN and n USA to NM Ref. Millidge 1976

Cymbium of normal shape in dorsal view; TmI located at 0.30-0.42; TmIV absent ..................... Micrargus, in part

35.169 35.168

Div. 2 species, 1 keys out here: Micrargus longitarsus (EMERTON 1882) Dist. CAN, AK, and n USA Refs. Crosby & Bishop 1933b, Paquin & Duprr 2003 Note Millidge (1977), without explicitly stating this to be a new combination, placed Baryphyma longitarsum in Micrargus, saying it was closely related to Micrargus herbigradus. Platnick (2005) retains it in Baryphyma

86(69)

Clypeus hairy (Fig. 35.165) and with a row of long spines on venter of tibia and metatarsus I and II (Fig. 35.166). Often found in wetlands ..................... Satilatlas, in part
Div. 6 species, several may key out here Dist. CAN and n USA to CO Refs. Millidge 1981c, Paquin & Duprr 2003

Without this combination of characters ....................... 87

35.170

Spiders of North America

Linyphiidae

141

87(86)

Cephalic lobe hourglass shaped (constricted at sides, wider at top) when viewed from the front (Fig. 35.173); cephalothorax and legs reddish, abdomen black; tibial apophysis projects dorsally away from tibia; TmIV present; TmI at 0.87-0.84 ........................................... Hypselistes
Div. 4 species Dist. CAN, AK, n USA Ref. Buckle et al. 2001, Paquin & Duprr 2003

Cephalic lobe rarely so constricted; coloration otherwise; tibial apophysis does not project dorsally away from tibia (unless there are multiple apophyses: Mythoplastoides); TmIV absent; TmI at 0.38-0.70 ...................................... 88 Embolic division long and relatively thin (>10 times longer than greatest width), and at least partly coiled or spiralled (Figs. 35.174-35.175) ....................................... 89 Embolic division rather thick (<10 times longer than greatest width), and does not coil or spiral, though it may be bent (Figs. 35.176-35.177) ............................................. 91 Tibial apophysis consists essentially of a single process, although it may be bifid with two short processes apically (Fig. 35.178) ........................................ Eridantes, in part
Div. 2 species: Eridantes erigonoides (EMERTON 1882) and Eridantes utibilis CROSBY & BISHOP 1933b Dist. eastern NA, AZ Ref. Crosby & Bishop 1933b, Paquin & Duprr 2003 Note both species may key out here

35.173

88(87)

89(88)

35.174

35.175

Tibial apophysis consisting of more than one process joined near the base, or a single, distinctly bifid process (Fig. 35.179) .................................................................... 90 Apical end of embolus thick and truncate (not sharply pointed) (Fig 35.174); end of embolus protrudes beyond cymbium somewhat (Fig 35.174) ................ Symmigma Embolus thin and pointed at end, and does not protrude beyond cymbium (Fig. 35.175) ......... Mythoplastoides

90(89)

Div. 1 species: Symmigma minimum (EMERTON 1923) Dist. AK, northwestern NA Ref. Crosby & Bishop 1933b

Div. 2 species: Mythoplastoides erectus (EMERTON 1915b) and Mythoplastoides exiguus (BANKS 1892a) Dist. widespread except southwestern USA Ref. Crosby & Bishop 1933b

35.176 91(88)

Cephalic lobe somewhat bifid longitudinally (Fig 35.180) ................................................................ Thyreosthenius

35.177

Div. 1 species: Thyreosthenius parasiticus (WESTRING 1851) Dist. CAN, n USA Refs. Crosby & Bishop 1933b as Hormathion limnatum CROSBY & BISHOP 1933b, Roberts 1987

92(91)

Cephalic lobe not bifid ................................................... 92 Embolus recurved, apical end points toward proximal portion of palp (Fig. 35.176) ................ Microctenonyx

Div. 1 species: Microctenonyx subitaneus (O. PICKARD-CAMBRIDGE 1875e) Dist. introduced to MA Refs. Crosby & Bishop 1933b, Roberts 1987

Embolus not recurved, apical end usually points to distal portion of palp (Fig 35.177) .......... Tapinocyba, in part
Div. 15 species, 5 key out here Dist. widespread Refs. Buckle et al. 2001, Paquin & Duprr 2003

35.178

35.179

With pits but unmodified cephalic region, 6 genera

93(52)

With prominent mastidion on cheliceral face; body length > 3 mm; TmIV present ....................................... Zornella
Div. 2 species: Zornella cryptodon (CHAMBERLIN 1920a) and Zornella cultrigera (L. KOCH 1879C) Dist. AK, CAN, n USA Refs. Buckle et al. 2001, Paquin & Duprr 2003

Without a cheliceral mastidion; body length < 3 mm; TmIV absent ................................................................... 94

35.180

142

Linyphiidae

94(93)

Carapace and sternum somewhat rugose (visible without highest magnification); embolus short and spinelike ....... ........................................................... Ceratinops, in part

Div. 8 species, 2 key out here: Ceratinops annulipes (BANKS 1892a) and Ceratinops rugosus (EMERTON 1909) Dist. widespread Ref. Crosby & Bishop 1933b

95(94)

Carapace and sternum smooth; embolus variable ....... 95 Embolus bifurcate terminally, shorter than width of cymbium, spinelike; often associated with ants ... Masoncus
Div. 4 species Dist. Widespread Refs. Cushing 1995, Buckle et al. 2001

Embolus not bifurcate terminally; embolus longer than width of cymbium; not associated with ants ................ 96 Embolus recurved, points toward proximal portion of palp; palpal tibia with dorsoectal macroseta ...................... .......................................................... Disembolus, in part

96(95)

35.181

Div. 24 species, 1 keys out here: Disembolus alpha (CHAMBERLIN 1949) Dist. CO Ref. Millidge 1981b

Embolus not recurved, palpal tibia lacking dorsoectal macroseta ........................................................................ 97 Embolus a large thick coil with a smaller apical coil (see Fig. 35.285 of Sisis rotundus, a similar species); tibial apophysis longer than length of tibia ................ Sisis, in part
Div. 2 species, 1 keys out here: Sisis plesius (CHAMBERLIN 1949) Dist. UT WY Ref. Chamberlin 1949

97(96)

Embolus coils, forming two loops subequal in size (Fig. 35.181); tibial apophysis shorter than length of tibia ........ ............................................................. Micrargus, in part
Div. 2 species, 1 keys out here: Micrargus longitarsus (EMERTON 1882) Dist. n NA Ref. Paquin & Duprr 2003

35.182

Modified cephalic region, no pits, 33 genera

98(53)

Most conspicuous carapace modification being one or two very large and thick forward-pointing macrosetae near apex of cephalic region (Figs. 35.182-35.184) ...... 99 Without one or two very large macrosetae near apex of cephalic region, although various smaller setae may be present ........................................................................... 101
35.183

99(98)

With one macroseta at apex of cephalic region (Fig. 35.182) ............................................... Islandiana, in part
Div. 14 species, 1 keys out here: Islandiana unicornis IVIE 1965 Dist. TX Ref. Ivie 1965

With two macrosetae at apex of cephalic region (Figs. 35.183-35.184) .............................................................. 100 Cephalic macrosetae nearly as long as width of eye area (Fig. 35.183); embolic division lacks obviously spiraled tailpiece ................................................ Paracornicularia
Div. 1 species: Paracornicularia bicapillata CROSBY & BISHOP 1931 Dist. MO, MS Ref. Crosby & Bishop 1931

100(99)

Cephalic macrosetae much shorter than width of eye area (Fig. 35.184); embolic division with obviously spiraled tailpiece ......................................... Scotinotylus, in part
Div. 34 species, 1 keys out here: Scotinotylus eutypus (CHAMBERLIN 1949) Dist. BC, OR, WA Ref. Millidge 1981a

35.184

Spiders of North America

Linyphiidae

143

101(98)

Abdomen with sclerotized scutum or shield (Fig. 35.185); reduced in some species to a small region at anteriormost portion of abdominal dorsum ..................................... 102 Abdomen lacking a scutum ......................................... 104

102(101) Embolic division with conspicuous tailpiece that is elon-

gate (more than twice as long as wide), long (more than half as long as palp width), and oriented with its long axis parallel to the long axis of the palp (Fig. 35.186) .............. ........................................................... Ceraticelus, in part Embolic division with tailpiece inconspicuous or otherwise than above ................................................... 103

35.185

Div. 41 species, 20 key out here Dist. Widespread Refs. Crosby & Bishop 1925c, Buckle et al. 2001, Paquin & Duprr 2003

35.187 103(102) Clypeus or clypeal projection covered with dense setae or

with variously modified setae (Fig. 35.187) ....................... ......................................................................... Floricomus

35.186

Div. 14 species Dist. widespread Refs. Buckle et al. 2001, Paquin & Duprr 2003

Clypeal projection with sparse or no setae (Fig. 35.188); membrane accompanying embolus with a sharp point midway along its length; this point projects toward the end of the palp ............................................ Pelecopsidis
Div. 1 species: Pelecopsidis frontalis (BANKS 1904a) Dist. southeastern USA to CT Ref. Bishop & Crosby 1935b

35.188

104(101) Palpal patella with ventral apical apophysis longer than

width of segment (Fig. 35.189) ............. Erigone, in part Palpal patella without ventral apical apophysis .......... 105

Div. 40 species, several may key out here Dist. widespread Refs. Buckle et al. 2001, Paquin & Duprr 2003

35.189

105(104) With distinct knoblike or proboscis-like process (not bearing eyes) emerging from eye region (Fig. 35.190) ..... 106

35.190

Without such a process in the eye-region ................... 107

106(105) Tibial apophysis longer than length of tibia; tarsal claws

I-III strongly pectinate (Fig. 35.191) .................................. ...................................................... Walckenaeria, in part

Div. 68 species, 25 key out here Dist. widespread Refs. Millidge 1983, Paquin & Duprr 2003

Tibial apophysis a darkened knob, shorter than length of tibia; tarsal claws not pectinate ......... Micrargus, in part

Div. 2 species, 1 keys out here: Micrargus pacificus (EMERTON 1923) Dist. AK, BC, WA Ref. Buckle et al. 2001 Note the placement of thisspecies in Micrargus is doubtful

35.192

107(105) With two cephalic lobes, both bearing eyes and many setae (Figs. 35.192-35.194) .................................................... 108

One or two cephalic lobes, only one lobe bearing eyes ..... ........................................................................................ 110

35.193

108(107) Posterior cephalic lobe much smaller than anterior lobe

(Fig. 35.192); tibial apophysis emerges ectally and tip points mesally, without long ectally-bent terminal spine (Fig. 35.195) ........................................... Dactylopisthes
Div. 1 species: Dactylopisthes video (CHAMBERLIN & IVIE 1947b) Dist. AK, YT Ref. Chamberlin & Ivie 1947b

Cephalic lobe subequal in size; tibial apophysis terminating in long spine bent abruptly to the ectal side ........ 109

35.194

35.195

144

Linyphiidae

109(108) Clypeal and cephalic lobes indistinctly separated (Fig.

35.196); embolus a loose coil at end of palp; (Figs. 35.197) tibia as in Fig. 35.198) .................................. Dicymbium
Div. 2 species: Dicymbium elongatum (EMERTON 1882) and Dicymbium nigrum (BLACKWALL 1834a) Dist. UT, WA Ref. Buckle et al. 2001

35.196

With two distinctly separated lobes (Fig. 35.199); embolus a bent spine rather than a coil ............................................ ..................................................... Diplocephalus, in part

Div. 4 species, 1 keys out here: Diplocephalus cristatus (BLACKWALL 1833a) Dist. CAN n USA Refs. Bishop & Crosby 1935c, Roberts 1987, Paquin & Duprr 2003

35.198 35.199 35.197

110(107) With either a distinct projection in the clypeal area (Figs.

35.200-35.202) or with the cephalic/clypeal region divided into two lobes [though they may be closely appressed, as with the subocular sulci that separate the clypeus from the ocular area in Blestia (Fig. 35.203) Figs. 35.204-35.208] ... ........................................................................................ 111 With only one elevation or lobe in the cephalic region, or with the carapace broadly raised (Figs. 35.209-35.210) .... ........................................................................................ 121

35.200

35.201

35.202

111(110) With distinct proboscis-like projection in the clypeal region (Figs. 35.200-35.202) ........................................ 112

With cephalic region divided into two lobes .............. 114

35.203

35.204
AME PME

35.205
posterior lobe

112(111) Clypeal projection with a conspicuous sclerotized cap

(Fig. 35.200) .................................... Baryphyma, in part

Div. 4 species, 2 keys out here: Baryphyma groenlandicum (HOLM 1967) and Baryphyma kulczynskii (ESKOV 1979) Dist. CAN, n USA. Refs. Buckle et al. 2001, Paquin & Duprr 2003

35.206

35.207

35.208

Clypeal projection without sclerotized cap, although setae may be present .............................................................. 113

113(112) Clypeal projection shorter than length of cheliceral bases,

faintly resembling a human nose (Fig. 35.201); paracymbium of normal size .................................... Perregrinus
Div. 1 species: Perregrinus deformis (TANASEVITCH 1982) Dist. CAN Refs. Buckle et al. 2001, Paquin & Duprr 2003

35.209

35.210

Clypeal projection longer than length of cheliceral bases, phallus-like (Fig. 35.202); paracymbium with a long and apically pointed distal arm ..................... Gnathonargus
Div. 1 species: Gnathonargus unicorn (BANKS 1892a) Dist. NY Ref. Bishop & Crosby 1935c

114(111) With the anterior cephalic lobe bearing eyes (Figs. 35.20435.206) .......................................................................... 115

With the posterior cephalic lobe bearing one or more pairs of eyes (Figs. 35.207-35.208) ........................................ 119
35.211 35.212

115(114) Embolic division with conspicuous tailpiece that is elon-

gate (more than twice as long as wide) and long (more than half as long as palp width) (T in Figs. 35. 21135.212) .......................................................................... 116 Embolic division with tailpiece inconscpicious, absent or otherwise than above (Figs. 35.213-35. 214) ............... 117

35.213

35.214

Spiders of North America

Linyphiidae

145

116(115) Tailpiece of embolic division twisted into a spiral (T in

Fig. 35.215); posterior cephalic lobe usually quite tall and placed over the anterior cephalic lobe, with a considerable space between them (Fig. 35.216; except Spirembolus mirus MILLIDGE 1980b with two small lobes); embolus coiled at end of palp ..................... Spirembolus, in part

35.216

Div. 43 species, 3 key out here Dist. Western NA Ref. Millidge 1980b

Posterior cephalic lobe comprised of a series of appressed longitudinal lobes (Fig. 35.217); tailpiece of embolic division not twisted into a spiral; embolus recurved (pointing toward proximal portion of palp) (Fig. 35.218) ................ ....................................................... Grammonota, in part
Div. 24 species, 1 keys out here: Grammonota gigas (BANKS 1896a) Dist. Western NA Refs. Bishop & Crosby 1933, Paquin & Duprr 2003

35.217 35.218

35.215

35.219

117(115) Clypeus densely setose (Fig. 35.219); embolus rather short

and not coiled ................................................ Goneatara

Div. 4 species Dist. eastern NA Refs. Bishop & Crosby 1935c, Barrows 1943, Paquin & Duprr 2003

Clypeus with at most a few setae; embolus long and curved or coiled ............................................................ 118

35.222
AME PME

118(117) Embolus forming more or less a complete coil, tip points

toward apical end of palp (Fig. 35.220) .............................. .................................................... Typhochrestus, in part

Div. 3 species, 2 may key out here: Typhochrestus latithorax (STRAND 1905) and Typhochrestus pygmaeus (SRENSEN 1898) Dist. CAN, AK, n USA Refs. Holm 1960a, 1967, Paquin & Duprr 2003

Embolus recurved, tip points mesally (Fig. 35.221) .......... .............................................................................. Souessa

35.223 35.220 posterior lobe

35.221

Div. 1 species: Souessa spinifera (O. PICKARD-CAMBRIDGE 1974b) Dist. e NA Refs. Crosby & Bishop 1936b, Paquin & Duprr 2003

119(114) Posterior cephalic lobe bearing all eight eyes (Fig. 35.222);

tibial apophysis shorter than width of tibia ......... Blestia

Div. 1 species: Blestia sarcocuon (CROSBY & BISHOP 1927) Dist. NC, PA, VA Ref. Millidge 1993a

35.224

35.225

Posterior cephalic lobe bearing one or two pairs of eyes; tibial apophysis longer than width of tibia ................. 120

120(119) Posterior cephalic lobe bearing AME and PME (Fig.

35.223); tibial apophysis with stout dorsoectal macroseta; tarsal claws not pectinate ........................... Coreorgonal
Div. 3 species Dist. western USA, western CAN Ref. Millidge 1981a

Posterior cephalic lobe bearing PME (Fig. 35.224); tibial apophysis lacks a stout dorsoectal macroseta; tarsal claws I-III strongly pectinate (Fig. 35.225) .................................. ...................................................... Walckenaeria, in part

Div. 68 species, about 13 species key out here (the Walckenaeria tricornis group) Dist. widespread Refs. Millidge 1983, Paquin & Duprr 2003

121(110) Embolic division with conspicuous tailpiece that is elon-

35.226

35.227

35.228

gate (more than twice as long as wide) and long (more than half as long as palp width) (T in Figs. 35.215, 35.22635.227) .......................................................................... 122 Embolic division with tailpiece inconspicuous, absent, or otherwise than above (Figs. 35.228-35.230) ................ 127

122(121) Embolus spinelike, much shorter than tailpiece; tibial apo-

physis bifid ................................................... Arcterigone

Div. 1 species: Arcterigone pilifrons (L. KOCH 1879c) Dist. AK, NT, YT Refs. Leech & Ryan 1972, Buckle et al. 2001

Embolus similar in length to much longer than, tailpiece; tibial apophysis variable ............................................... 123
35.229

35.230

146

Linyphiidae
bent embolus

123(122) Tailpiece of embolic division twisted into a spiral (T in Fig. 35.231) .......................................................................... 124

Tailpiece of embolic division not twisted into a spiral (T in Figs. 35.232-35.233) ..................................................... 125

124(123) Tibial apophysis with thick basoectal macroseta; tip of

embolus retroverted or pointing more or less toward proximal portion of palp ................ Disembolus, in part
Div. 24 species, 3 key out here Dist. widespread, more common in west Ref. Millidge 1981b

Tibial apophysis lacking thick basoectal macroseta; tip of embolus not retroverted, often terminates in wide coil at proximal portion of palp ............. Spirembolus, in part

Div. 43 species, 20 key out here Dist. western NA Ref. Millidge 1980b

35.232

T
125(123) Tibial apophysis appears to be two appressed projections

in dorsal view (Fig. 35.234) ..................... Epiceraticelus Tibial apophysis otherwise ........................................... 126

Div. 1 species: Epiceraticelus fluvialis CROSBY & BISHOP 1931 Dist. NY, OH Ref. Crosby & Bishop 1931

35.231

35.233

126(125) Tip of embolus initiates by an abrupt and acute bend (Fig.

35.232) .......................................... Ceratinopsis, in part Tip of embolus without such a bend (Fig. 35.233) ............ ....................................................... Grammonota, in part

Div. 24 species, 3 key out here Dist. widespread Ref. Bishop & Crosby 1930, Buckle et al. 2001, Paquin & Duprr 2003

Div. 26 species, 8 key out here Dist. widespread Refs. Bishop & Crosby 1933, Dondale 1959, Buckle et al. 2001, Paquin & Duprr 2003

127(121) Tibial apophysis definitely longer than length of rest of tibia (Fig. 35.235) .......................................................... 128

35.234

Tibial apophysis not longer than length of rest of tibia .... ........................................................................................ 130

35.235

128(127) Palpal femur distally wider than palpal bulb, tipped distoDiv. 1 species: Gonatium crassipalpum BRYANT 1933a Dist. AK, CAN, eastern USA Refs. Millidge 1981d, Paquin & Duprr 2003

dorsally with a sharp spine and numerous tubercles (Fig. 35.236) ............................................................ Gonatium
35.236

Palpal femur normal, unarmed distally ...................... 129

129(128) With two tibial apophyses (dorsal and ectal); embolus

recurved, tip pointing toward proximal end of palp; cymbium hollowed out on ectal side; paracymbium lacking (Fig. 35.237) ................................................ Sphecozone
Div. 1 species: Sphecozone magnipalpis MILLIDGE 1993d Dist. AZ Ref. Millidge 1993d Note this species may also key out at couplet 170

With one tibial apophysis; embolus tip points toward apical end of palp; cymbium and paracymbium normal (Fig. 35.235) .......................................... Silometopoides

Div. 1 species: Silometopoides pampia (CHAMBERLIN 1949) Dist. CAN Ref. Chamberlin 1949 as Minyriolus pampia

35.237

130(127) Embolus threadlike, running a teardrop-shaped course

across the palp (Fig. 35.238); eyes reduced in size; cephalic lobe sometimes densely clothed in short setae which are parted longitudinally (Fig. 35.239, Anacornia microps) . .......................................................................... Anacornia

35.238

Div. 2 species: Anacornia microps CHAMBERLIN & IVIE 1933a and Anacornia proceps CHAMBERLIN 1949 Dist. UT, WA Refs. Chamberlin & Ivie 1933a, Chamberlin 1949 Note both species may key out here

Eyes normal in size, and without this combination of characters ............................................................................ 131

35.239

Spiders of North America

Linyphiidae

147

131(130) Posterior declivity (PD) of cephalic lobe very steep, nearly

to more than vertical; clypeus (C) setose (Fig. 35.240); embolus short ................................................... Cheniseo
Div. 4 species Dist. AK, eastern NA Refs. Bishop & Crosby 1935c, Paquin & Duprr 2003

PD C

Posterior declivity of cephalic lobe much shallower than vertical; clypeus with few or no setae; embolus variable ... ........................................................................................ 132

35.240 35.241

132(131) Carapace almost trapezoidal in lateral view, with overhan-

ging clypeus, high peak, and long declivity (Fig. 35.241); cymbium with dark, setose projection dorsad of paracymbium (Fig. 35.242) ............................................... Eskovia
Div. 1 species: Eskovia exarmata (ESKOV 1989a) Dist. YT Ref. Eskov 1989a

Carapace not trapezoidal in lateral view; cymbium lacking setose process ................................................................ 133

133(132) Embolus long, whiplike, sinuous (Fig. 35.243); cephalic

lobe peaks at level of PME (Fig. 35.244); TmIV present ... ................................................................ Metopobactrus

Div. 1 species: Metopobactrus prominulus (O. PICKARD-CAMBRIDGE 1872b) Dist. CAN, n USA Refs. Bishop & Crosby 1935c [as Maso alticeps EMERTON 1909)], Roberts 1987, Paquin & Duprr 2003]

Embolus not sinuous; cephalic lobe peaks anterior or posterior to PME; TmIV variable ...................................... 134

35.242

35.243
134(133) Cymbium with basal outgrowth that fits over tibial apo-

physis (Fig. 35.245); embolus long, curved; TmIV present ............................................................ Procerocymbium

Div. 1 species: Procerocymbium dondalei MARUSIK & KOPONEN 2001c Dist. YT Ref. Marusik & Koponen 2001c

Cymbium normal; embolus short, thick, spinelike (Fig. 35.246); TmIV absent; sometimes with prominent stridulatory file on anterior of abdominal venter ........................ ...................................................................... Mecynargus
Div. 7 species Dist. AK, CAN Refs. Buckle et al. 2001, Paquin & Duprr 2003 Note Smodix reticulata (EMERTON 1915b), which probably belongs in Mecynargus, should key out here

35.244

No pits or cephalothoracic modification, 53 genera

135(53)

Eyes reduced in size or absent, usually cavernicolous ....... ........................................................................................ 136 Eyes normal, habitat variable ....................................... 139
35.245

136(135) Eyes absent ...................................................... Anthrobia

Div. 1 species: Anthrobia mammouthia TELLKAMPF 1844 Dist. AL, KY, TN, VA, WV Ref. Buckle et al. 2001 Note appeared as Anthrobia monmouthia in Tellkampf (1844), a lapsus calami. This genus includes two epigean species that will key out with Sciastes, and probably other troglobitic species in addition to Anthrobia mammouthia (J. Miller, pers. comm.)

35.246

Eyes reduced in size but not absent ............................. 137

137(136) Paracymbium with 2-3 conspicuous, very long setae (Fig.

35.247) ............................................... Islandiana, in part Paracymbium lacking very long setae ......................... 138

Div. 14 species, 3 key out here Dist. widespread Refs. Ivie 1965, Eskov 1987d

35.247

148

Linyphiidae

138(137) Tibial apophysis with one subapical point (Fig. 35.248);

paracymbium very thin (Fig. 35.248) ....... Caviphantes

Div. 1 species: Caviphantes saxetorum (HULL 1916) Dist. Palearctic, introduced to OR Ref. Roberts 1987

With multiple small tibial apophyses (Fig. 35.249); paracymbium normal .............................................. Phanetta

Div. 1 species: Phanetta subterranea (EMERTON 1875) Dist. e USA Ref. Millidge 1984a

139(135) Abdomen with sclerotized scutum or shield (Fig. 35.250);

reduced in some species to a small region at anteriormost portion of abdominal dorsum ..................................... 140 Without abdominal scutum ........................................ 141

35.249 35.248

140(139) Cheliceral fang sinuous (Fig. 35.251) ........... Ceratinella

Div. 14 species Dist. widespread except for southwest USA Refs. Crosby & Bishop 1925c, Chamberlin 1949, Paquin & Duprr 2003 Div. 41 species, 20 key out here Dist. widespread Refs. Crosby & Bishop 1925c, Buckle et al. 2001, Paquin & Duprr 2003

Cheliceral fang not sinuous ............ Ceraticelus, in part

141(139) With stridulating files on epigastric plates, activated by

sharp retrolateral angle of coxae IV (Fig. 35.252) ............. .......................................................................... Semljicola

35.250

Div. 4 species Dist. CAN, AK, CO Ref. Saaristo & Eskov 1996

Without stridulating ridge or retrolateral extensions of coxae IV ........................................................................ 142

142(141) Palpal patella with ventral apical apophysis (Fig. 35.253a35.253b) ........................................................................ 143

Palpal patella without ventral apical apophysis .......... 148

35.251

35.252

143(142) Patellar apophysis shorter to slightly longer than width of

rest of patella (Fig. 35.254); chelicerae with tooth (mastidion) on face ................................................................. 144 Patellar apophysis much longer than width of patella (Fig. 35.253b); chelicerae variable ........................................ 147

144(143) Cephalothorax orangish or reddish with black cephalic

region ................................................................. Tmeticus

Div. 1 species: Tmeticus ornatus (EMERTON 1914a) Dist. CAN, eastern USA Refs. Bishop & Crosby 1935c, Paquin & Duprr 2003

Cephalothorax tan, yellowish, or brownish; cephalic region not black or only black in interocular area ...... 145

35.253a

35.253b

145(144) Embolus longer than width of palp; tibial apophysis

longer than tibial length ........................ Gnathonarium

Div. 2 species Dist. AK, CAN Ref. Buckle et al. 2001

Embolic division without long embolus; tibial apophysis shorter than tibial length ............................................. 146

146(145) Body length less than 1.5 mm ............... Erigone, in part

Div. 56 species, 1 keys out here: Erigone praecursa CHAMBERLIN & IVIE 1939a Dist. NE Ref. Chamberlin & Ivie 1939a

Body length greater than 1.5 mm (usually greater than 2.0 mm) ................................................... Eperigone, in part

Div. 35 species, 2 key out here: Eperigone fradeorum (BERLAND 1932a) and Eperigone eschatologica (CROSBY in CHAMBERLIN 1924b) Dist. NA except NW Ref. Millidge 1987

35.254

Spiders of North America

Linyphiidae

149

147(143) Embolic division long, with multiple points, distinctively

shaped (Fig. 35.255); TmI at 0.80 or greater ...................... .................................................................. Ceratinopsidis

Div. 1 species: Ceratinopsidis formosa (BANKS 1892a) Dist. e USA Refs. Bishop & Crosby 1930

Embolic division otherwise; TmI at 0.60 or less ................ ................................................................. Erigone, in part

Div. 56 species, 54 key out here Dist. widespread Ref. Crosby & Bishop 1928a, Buckle et al. 2001, Paquin & Duprr 2003

148(142) Paracymbium with 2-3 conspicuous, very long setae (Fig.

35.256) ............................................... Islandiana, in part Paracymbium without very long setae ........................ 149

Div. 14 species, 10 key out here Dist. widespread Refs. Ivie 1965, Paquin & Duprr 2003

35.257 149(148) Palpus with thick, nearly straight, conductor-like sclerite

that is longer than width of palp. This sclerite extends well beyond the end of the palp (Fig. 35.257). Chelicerae robust, with a large tooth and hair tipped tubercles on its face ................................................. undescribed genus

35.255

Div. 2 species Dist. BC, MT, OR, WA Refs. Chamberlin 1949, Levi & Levi 1955 Note this udescribed genus includes Collinsia wilburi LEVI & LEVI 1955 and Spirembolus vasingtonus CHAMBERLIN 1949 a nomen nudum

Palpus lacking such a sclerite; chelicerae rarely robust or toothed .......................................................................... 150

150(149) Embolic division with conspicuous tailpiece that is elon-

gate (more than twice as long as wide), long (more than half as long as palp width), and oriented with its long axis parallel to the long axis of the palp (T in Figs. 35.258 35.260) ........................................................................... 151 Embolic division with tailpiece inconspicuous, absent, or otherwise than above (Figs. 35.271, 35.282, 35.284, 35.288-35.289, 35.292, 35.304, 35.306, 35.309-35.310) ..... ........................................................................................ 159

T
35.259

35.258

151(150) Tailpiece distinctly spiraled (T in Fig. 35.259) ............ 152

Tailpiece not spiraled (T in Figs. 35.258, 35.260) ....... 153

152(151) Embolic division with several short processes; dorsodistal

margin of palpal tibia often serrated; cymbium often with raised dorsal knob (Fig. 35.262) ................... Tachygyna

Div. 15 species Dist. Western NA Ref. Millidge 1984a

Embolus with a single point; palpal tibiae not serrated; tibial apophysis with a small secondary basal point; cymbium without raised dorsal knob ....................................... ........................................................ Spirembolus, in part
Div. 43 species, 3 key out here Dist. western NA Ref. Millidge 1980b

35.262 35.260

153(151) Dorsum of metatarsus I with row of large curved spines

(Fig. 35.263); TmI located at 0.50 to 0.60 ............ Scirites Setation of metatarsus I otherwise; TmI variable ....... 154

Div. 1 species: Scirites pectinatus (EMERTON 1911) Dist. CAN, n USA Ref. Bishop & Crosby 1938

35.263 154(153) Metatarsus I with two thick curved prolateral spines dis-

tally (Fig. 35.264); tibial apophysis with two points .......... ............................................................................ Wabasso

Div. 2 species: Wabasso cacuminatus MILLIDGE 1984a and Wabasso quaestio (CHAMBERLIN 1949) Dist. CAN, n USA Refs. Millidge 1984a, Paquin & Duprr 2003

35.264

Metatarsus I lacking two thick curved prolateral spines distally; tibial apophysis variable ................................. 155

150

Linyphiidae

155(154) Embolus looped back on itself at end of palp, forming

virtually a complete loop (Fig. 35.265); tarsal claws I-III strongly pectinate (Fig. 35.266) .......................................... ...................................................... Walckenaeria, in part
Div. 68 species, 30 key out here (the Walckenaeria acuminata group) Dist. widespread Refs. Millidge 1983, Paquin & Duprr 2003

Embolus not looped, but only curved or bent; tarsal claws not pectinate ................................................................. 156

35.266

156(155) Tibia with two apophyses, each longer than length of rest

of tibia, one dorsal and one ectal (Fig. 35.267) .................. ............................................................................. Coloncus Tibia with one apophysis, or with several short apophyses ........................................................................................ 157

Div. 5 species Dist. CAN, n USA Refs. Chamberlin & Ivie 1944, Chamberlin 1949, Paquin & Duprr 2003

35.265

35.267

157(156) Distal arm of paracymbium sharply pointed (Fig. 35.268);

apical end of embolic division divided into a sharp point and a long curving free part (Fig. 35.269) .......... Tutaibo
Div. 1 species: Tutaibo anglicanus (HENTZ 1850b) Dist. southeastern USA Ref. Bishop & Crosby 1930

35.268

Distal arm of paracymbium not pointed; apical end of embolic division a long curving free part ................... 158

158(157) Metatarsus IV with a trichobothrium ................................

...................................................... Grammonota, in part

Div. 26 species, 18 key out here Dist. widespread Refs. Dondale 1959, Buckle et al. 2001, Paquin & Duprr 2003

35.270 35.269

Metatarsus IV lacking a trichobothrium ............................ ........................... Ceratinopsis, in part and Styloctetor

Div. 24 species, 21 key out here + 2 species: Styloctetor purpurescens (KEYSERLING 1886b) and Styloctetor stativus (SIMON 1881g) Dist. widespread Refs. Bishop & Crosby 1930, Buckle et al. 2001, Paquin & Duprr 2003

159(150) Tibial apophysis definitely longer than length of rest of tibia (Figs. 35.270-35.272) ........................................... 161

Tibial apophysis not longer than length of rest of tibia .... ........................................................................................ 160
35.272

160(159) Tibial apophysis definitely shorter than width of nar-

rowest part of tibia, or lacking (Figs. 35.273-35.275) or as in Figure 35.376 ............................................................ 180 Tibial apophysis not shorter than width of narrowest part of tibia, but not as in Figure 35.376 ............................. 174

35.271 35.273

161(159) Palpal patella and palpal femur both distinctively longer

than cymbium (Fig. 35.270) ...................................... Tiso

Div. 2 species: Tiso aestivus (L. KOCH 1872b) and Tiso vagans (BLACKWALL 1834b) Dist. CAN, WA Refs. Locket & Millidge 1953, Roberts 1987

Cymbium as long or longer than palpal femur and/or patella ............................................................................ 162
35.274 35.275

162(161) Embolus less than half the length of cymbium .......... 163

Embolus longer than half the length of cymbium ..... 170

163(162) Distal portion of tibial apophysis strongly bent or even recurved (Fig. 35.277) .................................................. 164

Distal portion of tibial apophysis not strongly bent or recurved ........................................................................ 167
35.276 35.277

Spiders of North America

Linyphiidae

151

164(163) Cymbium somewhat hourglass-shaped in dorsal view;

embolic division appears bifid (Fig. 35.278) ...... Souidas

Div. 1 species: Souidas tibialis (EMERTON 1882) Dist. NH, QC (eastern North America) Refs. Crosby & Bishop 1936b, Paquin & Duprr 2003

Cymbium thicker in middle section in dorsal view; embolic division terminates in a single point ...................... 165

165(164) Tibial apophysis bent so tip points ectally (Fig. 35.279);

embolic division a sharp point ............................ Ivielum Tibial apophysis bent so tip points mesally ................ 166

Div. 1 species: Ivielum sibiricum ESKOV 1988b Dist. YT, Siberia Ref. Eskov 1988b

35.278 166(165) Embolic division with two processes, one rounded and

35.280

one sharp and hooked (Fig. 35.280) ..... Sciastes, in part

Div. 6 species, males are known for 5 of them, 1 species keys out here: Sciastes hastatus MILLIDGE 1984a Dist. YT, Siberia Ref. Millidge 1984a

35.279

Embolic division with one process, which is blunt apically but bearing a short sharp spine; tibial apophysis as in Fig. 35.281.................................................................. Zygottus
Div. 2 species: Zygottus corvallis CHAMBERLIN 1949 and Zygottus oregonus CHAMBERLIN 1949 Dist. BC, OR, WA Ref. Chamberlin 1949

167(163) Embolus somewhat recurved, so tip points toward proxi-

mal end of palp (Fig. 35.282); palpal tibia with small ventral lobe; TmI located at 0.70 or greater ............. Soulgas

Div. 1 species: Soulgas corticarius (EMERTON 1909) Dist. eastern NA Refs. Crosby & Bishop 1936b, Paquin & Duprr 2003

Embolus not recurved; palpal tibia lacking ventral lobe; TmI located at 0.60 or less ............................................ 168

35.281

168(167) Tibial apophysis shape distinctive (pistol shaped: Fig.

35.283); body uniformly pale in color; embolus long, curved .............................................................. Scylaceus Tibial apophysis a plain point or lobe; embolus terminating in a short, spinelike process; color variable ......... 169

35.282

169(168) Eye area black ................................... Masonetta, in part

Div. 1 species: Masonetta floridana (IVIE & BARROWS 1935) Dist. FL, GA Ref. Ivie & Barrows 1935 Note may also key out at couplet 176

Eye area concolorous with rest of cephalothorax ............... ................................................................. Sciastes, in part

Div. 6 species, males are known for 5 of them, 2 species key out here: Sciastes dubius (HACKMAN 1954) and Sciastes extremus HOLM 1967 Dist. WA, UT, CAN Refs. Millidge 1984a, Paquin & Duprr 2003

35.283

170(162) Embolus very sinuous, nearly forming a figure-8 shape

(Fig. 35.284); TmI located at 0.35 or less; tibial apophysis very long, thin, terminating in a sharp point ..................... .............................................................. Gnathonaroides

Div. 1 species: Gnathonaroides pedalis (EMERTON 1923) Dist. CAN and n USA Refs. Bishop & Crosby 1938, Paquin & Duprr 2003

Embolus not forming a figure-8 shape; TmI usually located at less than 0.35; tibial apophysis otherwise ......... 171

171(170) Embolus forms a wide proximal partial coil and termi-

nates in a small partial coil or hook (Fig. 35.285); tibial apophysis with long terminal and short basal projections, neither sharply pointed ................................ Sisis, in part
Div. 2 species, 1 keys out here: Sisis rotundus (EMERTON 1925a) Dist. AK, CAN Refs. Bishop & Crosby 1938, Paquin & Duprr 2003

35.284

Embolus coiled or not, but not forming large and small double coil .................................................................... 172

35.285

152

Linyphiidae

172(171) Embolus sinuous, S-shaped (Fig. 35.286); palpal tibia with

ventral lobe ................................................... Souessoula Embolus not S-Shaped, but coiled or recurved .......... 173

Div. 1 species: Souessoula parva (BANKS 1899b) Dist. eastern USA Ref. Millidge 1984a

173(172) Paracymbium absent; with two separate tibial apophyses,

dorsal and ectal; ectal margin of cymbium hollowed out; embolus recurved, so tip points toward proximal end of palp (Fig. 35.287) ........................... Sphecozone, in part
Div. 1 species: Sphecozone magnipalpis MILLIDGE 1993d Dist. AZ Ref. Millidge 1993d Note specimens may also key out at couplet 126

Paracymbium present; with one tibial apophysis with a single point; cymbium margins normal; embolus not recurved; embolus bifid ............ Typhochrestus, in part
Div. 3 species, 2 may key out here: Typhochrestus uintanus (CHAMBERLIN & IVIE 1939a) and Typhochrestus pygmaeus (SRENSEN 1898) Dist. CAN, AK, n USA Ref. Holm 1967

35.287 35.286

174(160) Free part of embolic division longer than width of cymbium (Figs. 35.288-35.289)........................................... 175

Free part of embolic division shorter than width of cymbium (Fig. 35.290) ........................................................ 176

175(174) Embolus makes complete coil around ventral face of palp

(Fig. 35.288); TmI located at 0.30-0.45 .............. Scironis

Div. 2 species: Scironis sima CHAMBERLIN 1949 and Scironis tarsalis (EMERTON 1911) Dist. CAN, AK, n USA Ref. Buckle et al. 2001

Embolus spirals at apical end of palp (Fig. 35.289); TmI beyond 0.45 ....................................... Sissicottus, in part

Div. 9 species Dist. widespread Refs. Miller 1999, Paquin & Duprr 2003 Note any of the species may key out here

35.288 176(174) Cymbium with basoectal groove that accepts the point of 35.289

the tibial apophysis (Fig. 35.290) ............................ Perro Cymbium without such a groove ................................ 177

Div. 1 species: Perro polaris (ESKOV 1986b) Dist. CAN, Siberia Ref. Eskov 1986b

177(176) Palp with large ventral conductor-like sclerite that points

ventroapically in ectal view (the posterior tooth of the embolic division) (Fig. 35.291); fourth metatarsus often with a trichobothrium ....................... Halorates, in part
Div. 16 species, 6 key out here Dist. widespread except southeastern USA Refs. Crosby & Bishop 1928a, Buckle et al. 2001, Paquin & Duprr 2003

Palp lacking such a conductor-like process; fourth metatarsus always lacking trichobothrium ......................... 178

35.290

178(177) Embolic division with multiple processes or points as

long or longer than the body of the embolic division (Fig. 35.292); sometimes with abdominal pattern and/or teeth on cheliceral face ............................... Eperigone, in part
Div. 35 species, 20 key out here Dist. widespread Refs. Millidge 1987, Paquin & Duprr 2003

Embolic division terminates as a single point or process .. ........................................................................................ 179

35.291

35.292

Spiders of North America

Linyphiidae

153

179(178) Tibial apophysis with multiple processes or with deep

notch producing appearance of two apophyses (Fig. 35.293); body uniformly pale; with tooth on cheliceral face ........................................................ Scyletria, in part
Div. 2 species, 1 key out here: Scyletria jona BISHOP & CROSBY 1938 Dist. CAN, eastern USA Ref. Bishop & Crosby 1938

Tibial apophysis a single point; eye area dark; cheliceral face lacking teeth ............................. Masonetta, in part

Div. 1 species: Masonetta floridana (IVIE & BARROWS 1935) Dist. GA, FL Ref. Ivie & Barrows 1935 Note may also key out at 166

180(160) Cymbium hourglass-shaped in dorsal view, narrowest in

middle (Fig. 35.294); embolus long, making more than one complete coil .................................................. Sisyrbe Cymbium not hourglass-shaped ................................. 181

35.293

35.294

Div. 1 species: Sisyrbe rustica (BANKS 1892a) Dist. NY Ref. Bishop & Crosby 1938

181(180) Tibia I somewhat swollen and curved (Fig. 35.295); with

small mastidion on cheliceral face; median row of four setae emerging from small tubercles between the posterior eye row and the thoracic groove ........................ Soucron
Div. 1 species: Soucron arenarium (EMERTON 1925a) Dist. CAN Ref. Crosby & Bishop 1936b, Paquin & Duprr 2003

35.295

Without the above combination of characters ........... 182

182(181) TmI located at 0.90 or greater; metatarsus IV usually with

trichobothrium ........................................................ Maso

Div. 4 species Dist. widespread Refs. Buckle et al. 2001, Paquin & Duprr 2003

TmI located at 0.75 or less; metatarsus IV usually lacking trichobothrium ............................................................. 183

183(182) Free part of embolus clearly longer than cymbium length ........................................................................................ 184

Free part of embolus clearly shorter than cymbium length ............................................................................ 186

184(183) Embolus makes 1 12 lateral loops (Fig. 35.296) .................

............................................................................. Lessertia

Div. 1 species: Lessertia dentichelis (SIMON 1884a) Dist. Palaearctic, introduced to BC Refs. Locket & Millidge 1953, Roberts 1987

35.296

Embolus sinuous, its path not coiling back on itself ......... ........................................................................................ 185

35.298

185(184) Embolus forms simple S-shape; with long, stout, bifid

conductor-like sclerite (part of the embolic division (Fig. 35.298); cymbium narrowed basally in dorsal view, with ectal groove; paracymbium normal (Fig. 35.299) ............. ........................................................................... Tunagyna
Div. 1 species: Tunagyna debilis (BANKS 1892a) Dist. AK, CAN, n USA Refs. Millidge 1984a, 1984b, Paquin & Duprr 2003

Embolus traces a complex, sinuous path (Fig. 35.300); lacking long, stout, bifid conductor-like sclerite; cymbium normal, without ectal groove; paracymbium very thin .... ............................................................................... Sitalcas
Div. 1 species: Sitalcas ruralis BISHOP & CROSBY 1938 Dist. NY Ref. Bishop & Crosby 1938

35.299

35.300

154

Linyphiidae

186(183) Cymbium with ectal groove which accepts ectal point

of tibial apophysis; cymbium narrowed basally in dorsal view (Fig. 35.301); palpal tibia with rounded apical notch, giving the appearance of two tibial apophyses .................. ................................................................ Scolopembolus
Div. 1 species: Scolopembolus littoralis (EMERTON 1913a) Dist. CT, NB Ref. Bishop & Crosby 1938

Cymbium lacking ectal groove or not narrowed basally; tibial apophysis usually otherwise ............................... 187

187(186) Cymbium of normal length, distinctly longer than wide .. ........................................................................................ 188

Cymbium rather short, almost as wide as long .......... 191

188(187) Embolic division with multiple apophyses ................. 189

Embolus terminates in a single point (although it may be bifid at apical end) ........................................................ 190

35.302

189(188) Palp with large ventral conductor-like sclerite that points

ventroapically in ectal view (posterior tooth of embolic division Fig. 35.302); never with abdominal pattern; rarely with teeth on cheliceral face ............... Halorates, in part

Div. 16 species, 10 key out here Dist. widespread except southeastern USA Refs. Crosby & Bishop & 1928a, Buckle et al. 2001, Paquin & Duprr 2003

Palp lacking such a conductor-like process (Fig. 35.303); sometimes with dorsal abdominal pattern; sometimes teeth on face of chelicerae ................. Eperigone, in part
Div. 35 species, 15 key out here Dist. widespread Refs. Millidge 1987, Paquin & Duprr 2003

190(188) TmI located at 0.30-0.35; embolus and embolic membrane

35.303

usually extend beyond end of cymbium (Fig. 35.304); embolus sometimes bifid apically ..................... Carorita
Div. 2 species: Carorita hiberna (BARROWS 1945) and Carorita limnaea CROSBY & BISHOP 1927) Dist. CAN, northern USA to CO Refs. Zujko-Miller 1999, Paquin & Duprr 2003

TmI located at 0.48 or greater; embolus usually does not extend beyond end of cymbium; embolus with a single point, ending in a fine hook-like extension of the embolus, but not bifid apically (palp as in Fig. 35.305) .................... .............................................................. Sisicottus, in part
Div. 9 species Dist. widespread except southeastern USA Refs. Miller 1999, Paquin & Duprr 2003 Note any of the species may key out here

35.304 191(187) Embolus and conductor (suprategular apophysis) point

35.305

at each other ; embolic division with long but blunt-ended free part and sharp accessory spine (Fig. 35.306) .............. .......................................................................... Vermontia
Div. 1 species: Vermontia thoracica (EMERTON 1913a) Dist. AK, CAN, northern USA Refs. Millidge 1984a, Paquin & Duprr 2003

Palpal configuration and embolic division otherwise ....... ........................................................................................ 192

192(191) Lacking a palpal tibial apophysis; cymbium often with

ectal lobe, or expanded/modified along ectal margin (Fig. 35.307) .................................................................. Eulaira
Div. 15 species Dist. widespread except for southeastern USA Refs. Chamberlin & Ivie 1945b, Buckle et al. 2001

With one or several apical palpal tibial apophyses; ectal margin of cymbium unmodified ................................. 193

35.306

35.307

Spiders of North America

Linyphiidae

155

193(192) With several short tibial apophyses; size greater than 1.2 mm ................................................................................ 194

Tibial apophysis a single point or process; size 1.2 mm or less ................................................................................. 196

194(193) Palpal tibia laterally compressed and dark in color, the

color contrasting with the palpal patella and femur (Fig. 35.308) .................................................. Scyletria, in part
Div. 2 species, 1 key out here: Scyletria inflata BISHOP & CROSBY 1938 Dist. CAN, eastern USA Refs. Bishop & Crosby 1938, Paquin & Duprr 2003

Palpal tibia concolorous with rest of palp, and not distinctly compressed laterally .......................................... 195

195(194) Embolus spirals or curls around embolic membrane at

end of palp (Fig. 35.309); distal arm of paracymbium curved and points toward proximal arm ........................... ..................................................................... Diplocentria
Div. 4 species Dist. CAN, n USA to CO Refs. Buckle et al. 2001, Paquin & Duprr 2003

35.308 35.309

Embolic division rounded posteriorly and not spiralled; distal arm of paracymbium not curved ............................. ................................................................ Sciastes, in part
Div. 6 species, males are known for 5 of them, 2 species key out here: Sciastes acuminatus (EMERTON 1913b) and Sciastes truncatus (EMERTON 1882) Dist. CAN, n USA Refs. Millidge 1984a, Paquin & Duprr 2003 Note Sciastes acuminatus probably belongs in Anthrobia. A new species of Anthrobia from eastern North America also keys out here (J. Miller, pers. comm.)

196(193) Embolic division with only one pointed process (Fig.

35.310) ............................................................ Annapolis

Div. 1 species: Annapolis mossi (MUMA 1945) Dist. MD, NB, NS Refs. Millidge 1984a, Paquin & Duprr 2003

Embolic division with more than one pointed process (Fig. 35.311) .................................................. Jacksonella

Div. 1 species: Jacksonella falconerii (JACKSON 1908) Dist. MN, palaearctic Refs. Locket & Millidge 1953, Roberts 1987

35.310

35.311

156

Linyphiidae

Female Linyphiinae
197(1)

Prolateral spine present on tibia I or, if prolateral spine on tibia I absent, then prolateral spine present on femur I .... ........................................................................................ 198 No prolateral spine on tibia I; no prolateral spine on femur I ........................................................................... 230

35.312

198(197) Some or all metatarsi with spines (check all legs) ...... 199

No metatarsi with spines .............................................. 218

199(198) Abdomen with conspicuous caudal tubercle (Fig. 35.312);

body red (fading to pale gray in alcohol) except for palps, eye region and caudal hump which are black (Fig. 35.312); epigynum as shown (Fig. 35.313) ...................... Florinda
Div. 1 species: Florinda coccinea (HENTZ 1850a) Dist. southeastern USA Ref. Blauvelt 1936

Abdomen without tubercle; often patterned dorsally; coloration brown or gray ............................................. 200

35.316

200(199) Carapace pale with median dark stripe bifurcating ante-

riorly; abdomen with pattern of narrow, dark chevrons on light background (Figs. 35.314-35.315) ...................... 201 Carapace and abdominal markings otherwise ............ 202
35.315

201(200) Dorsal stripe on carapace narrow (Fig. 35.315); epigynum

with triangular atrial opening and short dorsal scape (Fig. 35.316) ............................................................... Linyphia
Div. 1 introduced species: Linyphia triangularis (CLERCK 1757) Dist. ME Refs. Helsdingen 1969, Jennings et al. 2003

35.314

Dorsal stripe wider (Fig. 35.314); epigynum with scape usually extending well beyond epigastric furrow (Figs. 35.317-35.318), but short in Pityohyphantes limitaneus (EMERTON 1915a) (Fig. 35.319), atrium in form of shallow lateral depressions ............................... Pityohyphantes
Div. 16 species Dist. n and w NA Ref. Buckle et al. 2001

35.317

202(200) Epigynum with both dorsal and ventral scapes (Figs.

35.320-35.321) ........................... Bathyphantes, in part

Div. 26 species, 2 key out here: Bathyphantes keeni (EMERTON 1917a) and Bathyphantes reprobus (KULCZYNSKI 1916) Dist. widespread Ref. Ivie 1969

Epigynum with dorsal scape only, or with no scape ... 203

203(202) Metatarsi with ventral spines ....................................... 204

Metatarsi without ventral spines ................................. 210

35.318

35.319

204(203) Epigynum with an elongate scape extending well beyond epigastric groove ........................................................... 208

Epigynum otherwise .................................................... 205

Ventral scape Dorsal scape

35.320

35.321

Spiders of North America

Linyphiidae

157

205(204) Epigynum with mesal atrium and small, short dorsal scape (Figs. 35.322-35.323) .......................................... 206

Epigynum with broad mesal plate (Figs. 35.324-35.325) .. ........................................................................................ 207

206(205) Atrial opening tiny (Fig. 35.322) ............ Microlinyphia

Atrial opening large (Fig. 35.323) ....................... Neriene

Div. 9 species Dist. widespread Ref. Helsdingen 1969 Note some authors consider Neriene to be a synonym of Linyphia

207(205) Mesal plate of epigynum much wider than long, its sides

converging anteriorly (Fig. 35.324); carapace pale with black median and marginal bands ... Stemonyphantes

35.324

35.325

Div. 1 species: Stemonyphantes blauveltae GERTSCH 1951 Dist. n NA Ref. Helsdingen 1968

Mesal plate of epigynum with inverted T shape (Fig. 35.325); carapace uniform brown ...................................... ......................................................... Undescribed genus
Div. Linyphia catalina GERTSCH 1951, Linyphia rita GERTSCH 1951 and perhaps Linyphia tauphora CHAMBERLIN 1928a Dist. sw USA Ref. Gertsch 1951

35.326 208(204) Scape folded or otherwise modified at tip (Figs. 35.326-

35.327

35.327) ...................................... Lepthyphantes, in part Scape straight, unfolded (Figs. 35.329-35.331) ........... 209

Div. 29 species + several undescribed: a few of the larger species key out here Dist. widespread Refs. Zorsch 1937, Buckle et al. 2001

209(208) Scape relatively broad, with rounded tip (Fig. 35.328),

35.329

sometimes with a medial notch (Figs. 35.329-35.330) ...... ...................................................................... Allomengea

Div. 3 species Dist. n NA Ref. Helsdingen 1974

Scape long, with parallel sides and truncate tip (Fig. 35.331) ........................................................... Helophora
Div. 4 species Dist. n and w NA Ref. Buckle et al. 2001

35.328

35.330

210(203) AME larger than other eyes (Fig. 35.332); clypeus low,

about height of AME; anterior eye row strongly recurved; cheliceral teeth long; epigynum as shown (Fig. 35.333) .... ............................................................................ Tapinopa
Div. 2 species: Tapinopa bilineata BANKS 1893a, Tapinopa hentzi GERTSCH 1951 Dist. eastern NA Refs. Kaston 1981, Gertsch 1951

AME same size or smaller than other eyes (Fig. 35.334); clypeus height more than twice diameter of AME; anterior eye row nearly straight; cheliceral teeth short or long ....... ........................................................................................ 211

35.331

211(210) Chelicerae with 3-4 pairs of stout spines (Fig. 35.334);

cheliceral teeth long; trichobothrium present on metatarsus of leg IV; epigynum as shown (Fig. 35.335) ................. ......................................................................... Drapetisca

Div. Div. 2 species: Drapetisca alteranda CHAMBERLIN 1909a, Drapetisca oteroana GERTSCH 1951 Dist. n NA, NM Refs. Kaston 1981, Gertsch 1951

35.332

35.333

Chelicerae without stout spines; cheliceral teeth relatively short; no trichobothrium on metatarsus of leg IV ..... 212

35.334

35.335

158

Linyphiidae

212(211) Tibia I with ventral spines ............................................ 213

Tibia I with no ventral spines ...................................... 216

213(212) Tibia with 3-5 pairs of ventral spines (Fig. 35.336), epigy-

num as shown (Fig. 35.337); abdomen concolorous ......... .................................................................... Centromerita

Div. 1 introduced species: Centromerita bicolor (BLACKWALL 1833c) Dist. NF, NS, BC, WA Ref. Roberts 1987, Paquin & Duprr 2003

Tibia with 1 or 2 pairs of spines or a few scattered ones; epigynum otherwise; abdomen patterned or concolorous ....................................................................................... 214

214(213) Abdomen with dorsal folium (Figs. 35.338-35.339); epigy-

num as shown (Fig. 35.340) ............................. Estrandia Abdomen without folium, epigynum otherwise ........ 215

Div. 1 species: Estrandia grandaeva (KEYSERLING 1886b) Dist. n NA Ref. Blauvelt 1936, Paquin & Duprr 2003

35.339

35.340

35.338

215(214) Epigynum with short, broad, rigidly folded scape (Fig.

35.341) ...................................................... Oreophantes

Div. 1 species: Oreophantes recurvatus (EMERTON 1913a) Dist. n NA Refs. Helsdingen 1973b, 1981a, Paquin & Duprr 2003

Epigynal scape long, tip abruptly narrowed, usually curved or folded (Figs. 35.342-35.343) ................. Arcuphantes
Div. 7 species Dist. w NA Ref. Chamberlin & Ivie 1943

216(212) Legs short, femur I no longer than carapace; body conco-

lorous brown; basal segment of chelicera with anteriolateral row of short bristles (Fig. 35.344); epigynum as shown (Figs. 35.345-35.346) ................................. Centromerus
Div. 9 species Dist. e and n NA Ref. Helsdingen 1973c

35.341

35.343 35.342

Legs longer, femur I 114 to 2 times as long as carapace; abdomen frequently patterned; basal segment of chelicera without row of bristles ................................................. 217

217(216) Epigynum with short, straight scape, truncate posteriorly,

with a medial notch (Fig. 35.347) ................ Taranucnus

Div. 1 species: Taranucnus or nithes (BARROWS 1940) Dist. e NA from QC s to NC and TN Refs. Helsdingen 1973b, Paquin & Duprr 2003

35.344

Epigynum with longer, narrower scape without medial notch, usually curved or folded (Fig. 35.348-35.349) ........ .................................................... Lepthyphantes, in part
Div. 29 species + several undescribed: most species key out here Dist. widespread Refs. Zorsch 1937, Buckle et al. 2001

218(198) Prolateral spine on femur I .......................................... 219

35.345

35.346

No prolateral spine on femur I .................................... 223

35.347

35.348

35.349

Spiders of North America

Linyphiidae

159

219(218) Carapace with dark medial shield-shaped dark mark (Fig.

35.350); epigynal scape folded (Figs. 35.351-35.352); TmI 0.7 to 0.8; metatarsus IV with trichobothrium .................. ....................................................................... Poeciloneta
Div. 10 species Dist. n and w NA Ref. Buckle et al. 2001

Carapace without shield-shaped mark; scape straight or absent; TmI <0.5; metatarsus IV without trichobothrium ........................................................................................ 220

35.351

220(219) Epigynum with both dorsal and ventral scapes (Figs.

35.353-35.357); ventral scape may be very short (Fig. 35.357), or long (Fig. 35.355) but always with terminal pit ............................................... Bathyphantes, in part
Div. 26 species. 24 key out here Dist. widespread Ref. Ivie 1969

35.350 35.352

Epigynum without ventral scape ................................. 221

221(220) Dorsal scape relatively long, with terminal pit (Figs.

35.358-35.359) ............................................... Kaestneria

35.353 35.354 35.355

Div. 3 species Dist. n NA Ref. Ivie 196

Dorsal scape short (Fig. 35.360) or rudimentary (Fig. 35.361) .......................................................................... 222
35.356

222(221) Carapace and abdomen concolorous; legs relatively short,

with femur I about as long as carapace; epigynum as shown (Fig. 35.361)...................................... Porrhomma
Div. 9 species Dist. widespread Ref. Buckle et al. 2001

Carapace with dark cephalic region; abdomen patterned; legs relatively long, with femur I 1.5 times as long as carapace; epigynum as shown (Fig. 35.360) ............................. .................................................................... Pacifiphantes
Div. 1 species: Pacifiphantes magnificus (CHAMBERLIN & IVIE 1943) Dist. Pacific northwest Refs. Ivie 1969, Eskov & Marusik 1994 Note the male of Pacifiphantes magnificus is unknown. It is unclear whether Eskov & Marusik (1994) were correct in placing it in Pacifiphantes or Ivie (1969) in placing it as a close relative of Bathyphantes approximatus (O. PICKARD-CAMBRIDGE 1871a)

35.357

35.358 223(218) Epigynum without scape ............................................. 224 35.359

Epigynum with scape ................................................... 225

224(223) Epigynal plate ovoid, with anterior openings (Fig. 35.262);

dark unicolorous carapace and distinctive abdominal pattern with lateral light bars (Fig. 35.363) ....... Frontinella

Div. 2 species Dist. Frontinella pyramitela (WALCKENAER 1841), widespread and Frontinella huachuca GERTSCH & DAVIS 1946, AZ Refs. Blauvelt 1936, Gertsch & Davis 1946 Note Frontinella pyramitela is referred as Frontinella communis (HENTZ 1850a) in Platnick (2005)

35.361 35.360

Epigynal plate ovoid, with posterior openings (Fig. 35.364); carapace patterned; dorsum of abdomen dark with paired light spots (Fig. 35.365) ........... Graphomoa
Div. 1 species: Graphomoa theridioides CHAMBERLIN 1924a Dist. LA, TN Ref. Chamberlin 1924a Note according to Gustavo Hormiga (pers. comm.) Graphomoa theridioides is a synonym of Pocobletus coroniger SIMON 1894a and Graphomoa a synonym of Pocobletus SIMON 1894a

35.363

35.362

35.364 35.365

160

Linyphiidae

225(223) Epigynum as in Fig. 35.366..................... Jalapyphantes

Div. 1 species: Jalapyphantes cuernavaca GERTSCH & DAVIS 1946 Dist. AZ Ref. Gertsch & Davis 1946

Epigynum otherwise .................................................... 226

226(225) Abdomen patterned (Fig. 35.367); epigynal scape narrow,

straight or slightly bent ventrally (Figs. 35.368-35.369) .... ...................................................................... Linyphantes Abdomen concolorous; epigynal scape otherwise ...... 227

Div. 19 species Dist. Pacific northwest Ref. Chamberlin & Ivie 1942a

227(226) Epigynum scape rolled dorsally and with lateral lobes (Fig. 35.370-35.372) .............................................................. 228

35.368

Epigynum scape rigidly folded .................................... 229

228(227) Trichobothrium present on metatarsus IV; epigynum as

shown (Fig. 35.371) ........................................ Microneta

35.369

Div. 1 species: Microneta viaria (BLACKWALL 1841a) Dist. n and w NA Refs. Kaston 1981, Saaristo 1974a, Paquin & Duprr 2003

No trichobothrium on metatarsus IV; epigynum as shown (Figs. 35.372-35.373) ............................ Agyneta, in part

Div. 45 species and perhaps as many more undescribed, a few western species key out here Dist. widespread Refs. Buckle et al. 2001, Paquin & Duprr 2003 Note some authors treat Meioneta HULL 1920 as a distinct subgenus or genus. Agyneta sensu stricto has a trichobothrium on metatarsus IV while Meioneta does not

35.370 (expanded view) 229(227) Basal part of scape concave distally, terminal part of folded

35.371

scape does not extend beyond it (Fig. 35.374) ................... ............................................................................ Saaristoa

Div. 1 species: Saaristoa sammamish LEVI & LEVI 1955 Dist. w NA Ref. Levi & Levi 1955

Basal part of scape convex distally, terminal part often extends beyond it (Fig. 35.375) .... Oreonetides, in part
Div. 6 species + several undescribed, 3 described species key out here Dist. n and w NA Ref. Helsdingen 1981a

35.372

35.373

230(197) Epigynum scape rolled dorsally (Figs. 35.370, 35.37635.377) .......................................................................... 231

Scape not rolled ............................................................ 233

231(230) Epigynal scape narrow, tightly rolled, without lateral lobes

(Figs. 35.376, 35.378) .................................... Macrargus

35.374

Div. 1 species: Macrargus multesimus (O. PICKARD-CAMBRIDGE 1875c) Dist. n NA Ref. Kaston 1981, Paquin & Duprr 2003

Scape wider, more loosely rolled, with lateral lobes (Figs. 35.377, 35.379) .............................................................. 232

35.376

35.377

35.378

35.379

Spiders of North America

Linyphiidae

161

232(231) Spiracle forward 1/4 to 1/3 of way to epigastric furrow(Fig.

35.380); epigynum very similar to that of Agyneta (Fig. 35.381) ................................................... Tennesseellum

Spiracle in normal position at base of spinnerets (Fig. 35.382); epigynum as shown (Figs. 35.383-35.384) .......... ................................................................ Agyneta, in part
Div. 45 species and perhaps as many more undescribed, most key out here Dist. widespread Refs. Buckle et al. 2001, Paquin & Duprr 2003 Note some authors treat Meioneta HULL 1920 as a distinct subgenus or genus. Agyneta sensu stricto has a trichobothrium on metatarsus IV while Meioneta does not. Anibontes longipes CHAMBERLIN & IVIE 1944 will key out here, and perhaps the unknown female of Anibontes mimus (CHAMBERLIN 1924) as well

35.380

35.381

233(230) Anterior eyes approximately equally spaced; abdomen

patterned (Fig. 35.385); epigynum with short, broad scape notched medially (Figs. 35.386-35.387) ............ Wubana
Div. 7 species Dist. n and w NA Refs. Chamberlin & Ivie 1936b, Buckle et al. 2001, Paquin & Duprr 2003

35.383 35.382

AME much closer to each other than to ALE; abdomen concolorous; epigynum otherwise .............................. 234
35.384

234(233) Legs relatively long and slim, with femur I length 1 1/3

times that of carapace; leg spines short and thin, barely distinguishable from setae; eyes small, especially AME (Fig. 35.388); epigynum long and straight with bluntly rounded tip (Fig. 35.389) .............................. Oaphantes
Div. 1 species: Oaphantes pallidulus (BANKS 1904b) + 2 undescribed Dist. CA to BC Ref. Chamberlin & Ivie 1943

35.386

Legs shorter, with femur I no longer than carapace; leg spines of normal size; eyes larger; epigynum with folded scape, tip with stretcher usually visible (Figs. 35.39035.392) .......................................................................... 235

35.385

35.387 235(234) Body under 1.7 mm in length; stridulating ridges on

epigastric plates (Fig. 35.393); epigynal scape short (Fig. 35.390) ..................................................................... Maro
Div. 2 species: Maro nearcticus DONDALE & BUCKLE 2001 and Maro amplus DONDALE & BUCKLE 2001 Dist. n NA Refs. Dondale & Buckle 2001, Paquin & Duprr 2003

Body usually over 1.7 mm in length; no stridulating ridges on epigastric plates; epigynal scape longer (Figs. 35.39135.392) .......................................... Oreonetides, in part
Div. 6 species + several undescribed, 3 described species key out here Dist. n and w NA Ref. Helsdingen 1981a

35.388

35.389 35.390

35.391 35.392

35.393