systematics of nearctic cybaeus (araneae�:�cybaeidae)

Systematics of Nearctic Cybaeus (Araneae : Cybaeidae)

Claudia R. CopleyA,C,D, Robb BennettB and Steve J. PerlmanC

ANatural History Section, Royal BC Museum, 675 Belleville, Victoria, BC V8W 9W2, Canada.BBC Ministry of Forests & Range, 7380 Puckle Road, Victoria, BC V8M 1W4, Canada.CDepartment of Biology, University of Victoria, PO Box 1700 STN CSC, Victoria, BC V8W 2Y2, Canada.DCorresponding author. Email: [email protected]

Abstract. Spiders in the genusCybaeusL.Koch (Araneae : Dictynoidea : Cybaeidae) are common forest-floor inhabitantsin western North America and Japan. Here we establish an initial phylogenetic framework for North American Cybaeus.Morphological details for eight proposed species groups are given, and these results, combined with molecular analyses ofone nuclear and onemitochondrial gene for six of the eight species groups, suggest that NorthAmericanCybaeus species arecontained in two broad clades, one Holarctic and one Nearctic (primarily Californian). The Holarctic clade contains thetetricus andangustiarum species groups,which containmostlywidely distributed species. TheCalifornian clade includes theadenes, aspenicolens, consocius, devius, septatus and tardatus species groups, all of which have very restricted ranges.ThegenusCybaeusand thePalaearctic speciesC. tetricus (C.L.Koch) (type speciesof thegenus) andC.angustiarumL.Kochare redescribed and illustrated. A key to species groups is provided. Nine new species endemic to the western Nearctic andincluded in the molecular analyses are described and illustrated: C. paralypropriapus Bennett, sp. nov. and C. wayneiBennett, sp. nov. (tetricus group); C. sanbruno Bennett, sp. nov. (adenes group); C. thermydrinos Bennett, sp. nov.(aspenicolens group);C. penedentatusBennett, sp. nov. andC. vulpinusBennett, sp. nov. (consocius group);C. chauliodousBennett, sp. nov. and C. somesbar Bennett, sp. nov. (septatus group); and C. gidneyi Bennett, sp. nov. (unplaced).

Additional keywords: arachnid, California, morphology, phylogenetics, species descriptions, spider, taxonomy.

Introduction

Members of the Cybaeidae (Araneae : Dictynoidea) are amongthemost common spiders encountered in forests ofwesternNorthAmerica and Japan.Despite this, little has beenpublishedon themexcept for some taxonomic, ontological and other work on theNorth American fauna (Bennett 1992, 2005, 2006), and newspecies descriptions and natural history information from Japan(Ihara 2004–2007; Ihara and Nojima 2004) and Korea (Kim andKim 2008). The family includes 12 genera and 162 species(Platnick 2009), with two-thirds of the species in the genusCybaeus L. Koch. They generally have an annual life cycleand are most common in moist woodlands, where they occurin litter and in association with cover objects on the forest floor.

Members of this family are Holarctic and primarily circum-Pacific indistribution,with twomajor centres of diversity in JapanandwesternNorthAmerica (see Platnick 2009).As inmany othergroups of organisms (Myers et al. 2000; Brooks et al. 2002;Lapointe and Rissler 2005; Bond and Hedin 2006), in NorthAmerica the greatest diversity of cybaeids is found in California,Oregon and Washington; only four species live east of theMississippi River (Platnick 2009). Western Nearctic cybaeidsare of conservation interest because many have highly restricteddistributions, primarily in California and southern Oregon, andthere is general concern for the unique biota of that area (Hobbsand Mooney 1998; Bond et al. 2006; Vandergast et al. 2007).

In fact, the majority of Cybaeus species have restricteddistributions (Chamberlin and Ivie 1932; Ihara 2004). Thismay be because, in contrast to many other spider taxa, cybaeidspiders shownopropensity to disperse by ballooning (R.Bennett,unpubl. data). Gene flow for cybaeid species is probably limitedand mediated primarily through mate-seeking, wandering males(Hendrixson andBond 2005; Crews andHedin 2006; Starrett andHedin 2007).

Cybaeids have few distinctive features other than genitaliccharacters, which are useful for the identification of generaand species (Chamberlin and Ivie 1932; Ihara 2004; Bennett2005, 2006). In part, because of its generalised morphology,the family has been understudied from a systematic andphylogenetic perspective. Definitive apomorphies and soliddiagnoses for the family and genus are lacking (Bennett 2005,2006): Cybaeus may be paraphyletic as presently construedand the phylogenetic affinities of cybaeids remain unclear.However, a recent molecular phylogenetic study (Spagnaand Gillespie 2008) has suggested a relationship of(Agelenidae + (Cybaeidae + Hahniidae)).

In his unpublished Ph.D. dissertation, Bennett (1991) madesignificant progress in the systematics of North Americancybaeids. He classified 61 North American species of Cybaeus(including 33 new species) into eight species groups using maleand female genitalic characteristics and revised several other

� CSIRO 2009 10.1071/IS09001 1445-5226/09/040367

CSIRO PUBLISHING

www.publish.csiro.au/journals/is Invertebrate Systematics, 2009, 23, 367–401

Nearctic cybaeid taxa. Here, we introduce the Cybaeus speciesgroups and the morphological characters that define them. Wealso present a preliminary molecular phylogenetic analysis ofCybaeus based on portions of two genes that we sequenced from20 Cybaeus species (including nine that are newly described inthis paper), representing six of the eight proposed species groups.In addition to the nine new species, two are redescribed to ensurecorrelation of our datasets with existing names. Our aim is toprovide a useful framework for understanding the diversity andevolution of Cybaeus.

Methods

Morphological examination

Preliminary examinations of spiders were carried out with thespecimens immersed in 75% ethanol under a Leitz Greenoughstereo dissecting microscope (maximum magnification 100�;Leitz, http://leitzmicroscopes.com/). Measurements of spiderparts were made using a micrometer reticle in one ocular lensof the microscope.

Vulvae (with epigyna attached) were excised from femaleabdomens with sharpened insect pins, temporarily cleared withclove oil in depression slides and examinedwith a Leitz Laborluxstereo compound microscope at magnifications of up to 400� or1000�. Initial hand sketches of epigyna, vulvae and some palpiwere made with the aid of a squared-grid reticle in the dissectingmicroscope or a drawing tube affixed to the compound scope.Most palpi were initially photographed in an S-570 Hitachiscanning electron microscope (Hitachi, www.hitachi.com), butbecause of problems obtaining good contrast for publication,most of the resultant SEMs of palpi were subsequently copied toink drawings. For scanning electron microscopy, male genitaliccharacters were prepared by specimen dehydration in absoluteethanol for several days followed by air drying, mounting onstandard SEMstubswithwhite glue and sputter coatingwith goldpalladium before examination and photography in the vacuumtube of the microscope.

Taxon sampling and choice of markersfor molecular phylogenetics

For the molecular analysis, we were able to obtain samplesfrom 20 Cybaeus species (including the nine that are newlydescribed in this paper) representing six of the eight speciesgroups (tetricus, angustiarum, adenes, consocius, aspenicolensand septatus) aswell as one unplaced species (Table 1).Cybaeotanana Chamberlin & Ivie was used as an outgroup because it is amember of the cybaeid family outside of the genus Cybaeus. Allspecimens were identified using keys and illustrations in Bennett(1991). Voucher specimens have been placed in the entomologycollection of the Royal BC Museum.

We sequenced portions of two genes, mitochondrialcytochrome c oxidase subunit I (COI) and nuclear histonesubunit 3 (H3). These genes often complement each other interms of resolution, with the more rapidly evolving COI offeringbetter resolution at shallow divergence. In animals, COI evolvesrapidly enough to distinguish between closely related taxa and,because there are readily available primers that can amplify thisgene across virtually all animal lineages, it has become themarkerof choice in animal DNA barcoding initiatives (Hebert et al.

2003). H3 has been used in several recent phylogenetic analysesof spiders (Arnedo et al. 2004; Maddison and Needham 2006;Spagna and Gillespie 2008).

DNA extraction, amplification and sequencing

DNA was extracted and purified from entire specimens, with theexception of genitalic vouchers (palps or epigynes), using theQiagen Dneasy Tissue Kit (QIAGEN, www.qiagen.com).Approximately 400 bases of the H3 gene were amplified withthe primers H3aF: 50-ATGGCTCGTACCAAGCAGACVGC-30

andH3bR: 50-ATATCCTTRGGCATRATRGTGAC-30 (Colganet al. 1998), and ~700 bases of COI were amplified with theprimers LCO1490: 50-GGTCAACAAATCATAAAGATATTGG-30 and HCO2198: 50-TAAACTTCAGGGTGACCAAAAAATCA-30 (Folmer et al. 1994) via PCR and were cleaned usinga Qiagen PCR Purification Kit. H3 genes were sequenced inboth directions directly from PCR products by Macrogen Corp.(Seoul, Korea). COI genes were first cloned (Stratagene cloningkit, www.stratagene.com) and then sequenced in both directionsat the University of Victoria sequencing facility. Chromatogramswere visualised and cleaned usingGeneious Pro (www.geneious.com). Sequences were aligned using both Geneious Pro andClustalW (Thompson et al. 1997) and compared with othersequences in GenBank using BLAST searches (Altschul et al.1990) in order to rule out any possible contamination. Allsequences are deposited in GenBank (see Table 1).

Phylogenetic analysis

Phylogenetic analysis was performed using maximum parsimonyandmaximumlikelihoodmethods, implemented inPAUP*4.0b10(Swofford 2003) and PhyML (Guindon and Gascuel 2003)respectively. H3 and COI datasets were analysed separately.All trees were rooted with Cybaeota nana. For maximumparsimony, heuristic searches were performed using 100random addition sequences and the tree-bisection-reconnection(TBR) branch-swapping algorithm.To assess confidence in nodes,500 bootstrap replicates were performed; these were full heuristicsearches as implemented with maximum parsimony, except that10 random addition sequences were used for each bootstrap. Formaximumlikelihood,wefirst usedModeltest (PosadaandCrandall1998) to determine which model of nucleotide evolution bestfit our data; this was found to be general time reversible(GTR) + invariable sites (I) + gamma distribution (G) for theCOI dataset and Kimura 2-parameter + I +G for H3. Maximumlikelihood was then implemented, with 100 bootstrap replicates toassess node support. TheMLtreewas visualised inNjplot (Perrièreand Gouy 1996). We used a one-tailed Shimodaira–Hasegawa(SH) test (Shimodaira and Hasegawa 1999; Goldman et al. 2000),implemented in PAUP*, in order to ask whether a tree constrainedsuch that the angustarium and tetricus species groups arereciprocally monophyletic is significantly less likely than ourbest H3 tree.

Results

Morphology

Primarily on the basis of genitalic characters, North AmericanCybaeus species can be classified into eight species groups.Among these species groups, there is also morphological

368 Invertebrate Systematics C. R. Copley et al.

Tab

le1.

Cyb

aeid

speciesused

inmolecular

phylogenetican

alysis

H3=nu

clearhiston

esubu

nit3

,COI=

cytochromecox

idasesubu

nitI

Species

Species

Geographicrange

Locality

Catalogue

number

GenBankAccession

grou

pH3

COI

Cybaeotana

naBritishColum

biato

Utah

toCalifornia

California;SierraCounty;

39.6000�N12

0.61

67� W

ENT008–00

8605

FJ263

766

FJ263

787

Cybaeus

tetricus

tetricus

CentralandeasternEurope

Slovenia;InnerCarniolaRegion;

45.7917�N14

.201

4�E

ENT008–00

8606

FJ263

770

Cybaeus

eutypu

stetricus

BritishColum

biato

Orego

nBritishColum

bia;Vancouv

erIsland

;50

.402

4�N12

8.06

48� W

ENT008–00

8607

FJ263

773

FJ263

791

Cybaeus

morosus

tetricus

Western

North

America

BritishColum

bia;HeakamieRiver;51

.034

9�N12

4.53

48� W

ENT008–00

8608

FJ263

775

FJ263

792

Cybaeus

multnom

atetricus

Oregonto

California

Oregon;

Douglas

County;

43.6472�N

123.88

33� W

ENT008–00

8609

FJ263

777

Cybaeus

paralyprop

riapus

tetricus

Orego

nOrego

n;Dou

glas

County;

43.647

2�N

123.88

33� W

ENT008–00

8610

FJ263

776

Cybaeus

waynei

tetricus

Oregonto

California

Oregon;

Douglas

County;

43.6472�N

123.83

33� W

ENT008–00

8611

FJ263

778

Cybaeus

angustiarum

angu

stiarum

Europe

Czech

Republic;Moravian-Silesian

Region;

49.5830�N18

.000

0�E

ENT008–00

8612

FJ263

771

FJ263

790

Cybaeus

giga

nteus

angu

stiarum

New

Yorksouthto

Georgia,

Alabama

North

Carolina;Buncombe

County;

35.3604�N82

.435

5�W

ENT008–00

8613

FJ263

769

FJ263

788

Cybaeus

patritu

san

gustiarum

Tennessee

toNorth

Carolina

Tennessee;BlountCounty;

35.6450�N

83.806

6�W

ENT008–00

8614

FJ263

768

FJ263

789

Cybaeus

reticulatus

angu

stiarum

Western

North

America

BritishColum

bia;Vancouv

erIsland

;48

.647

8�N12

4.59

55� W

ENT008–00

8615

FJ263

774

FJ263

793

Cybaeus

sign

ifer

angu

stiarum

Western

North

Americasouth

from

BritishColum

bia

BritishColum

bia;Vancouv

erIsland

;48

.454

4�N12

3.27

14� W

ENT008–00

8616

FJ263

772

FJ263

794

Cybaeus

shoshoneus

angu

stiarum

Idaho,

Washing

ton,

Orego

nandMontana

Washington;

Spokane

County;

47.866

6�N11

7.23

94� W

ENT008–00

8617

FJ263

767

Cybaeus

sanbruno

adenes

California

California;San

Mateo

County;

37.6943�N12

2.45

30� W

ENT008–00

8618

FJ263

780

FJ263

795

Cybaeus

thermydrino

saspenicolens

California

California;KernCounty;

35.7333�N11

8.56

66� W

ENT008–00

8619

FJ263

783

Cybaeus

hesper

consocius

California

California;San

Mateo

County;

37.2998�N12

2.26

95� W

ENT008–00

8620

FJ263

781

FJ263

796

Cybaeus

penedentatus

consocius

California

California;SantaCruzCounty;

36.5964�N12

2.04

24� W

ENT008–00

8621

FJ263

784

FJ263

797

Cybaeus

vulpinus

consocius

California

California;Calaveras

County;

38.3101N

120.7224W

ENT008–00

8622

FJ263

782

Cybaeus

chaulio

dous

septatus

Oregonto

California

California;Hum

boldtCounty;

40.9475�N12

3.64

59� W

ENT008–00

8623

FJ263

786

FJ263

799

Cybaeus

somesba

rseptatus

Oregonto

California

California;SiskiyouCounty;

41.1123�N12

3.12

86� W

ENT008–00

8624

FJ263

785

FJ263

798

Cybaeus

gidn

eyi

unplaced

Oregonto

California

California;Hum

boldtCounty;

41.4786�N12

4.00

00� W

ENT008–00

8625

FJ263

779

FJ263

800

Systematics of Nearctic Cybaeus (Araneae : Cybaeidae) Invertebrate Systematics 369

support for species falling into two major clades – one that isHolarctic in distribution, the other Nearctic and primarilyCalifornian. Below, we briefly summarise the characters thatdefine the species groups, aswell as the potential synapomorphiesthat unite some larger groupings. These hypotheses providea useful framework with which to interpret the results of ourmolecular phylogenies. Species groups and species descriptionsare discussed in greater detail in the taxonomic section.

In common with some other cybaeids (e.g. CybaeinaChamberlin & Ivie), males of most Cybaeus species possess aretrolateral patellar apophysis (RPA – Fig. 16). Usually the RPAis well developed and possesses a variable number of ‘peg setae’(Figs 20, 21). Secondarily, it may be greatly reduced (RPA –

Fig. 34) or vestigial and lacking peg setae.North American Cybaeus species fall into two major lineages

(Holarctic and Californian); this is reflected in whetherspermathecal heads are ‘in-line’ (angustiarum+ tetricusgroups) or reduced (aspenicolens+ septatus+ tardatus+devius+adenes+ consocius groups) respectively. Bennett (1991) used twosets of spermathecal head character states for inter-grouprelationships: size and location within the vulva. Plesiomorphicstates are spermathecal heads large (HS – Figs 4, 31, 44, 45) andlocated on distinct blind lobes (HS – Figs 12, 66, 95, 98).Apomorphic states are heads small (HS – Figs 12, 66, 95, 98)and located in-line with the rest of the spermathecal ducts(HS – Figs 31, 38, 44, 45).

The Holarctic clade consists of two species groups, althoughthe characters defining each are somewhat ambiguous. Thetetricus group is defined by the large spermathecal heads(plesiomorphy) occurring in line with the rest of thespermathecal ducts (apomorphy) (HS – Fig. 31). Although thelatter character is also possessed by members of the angustiarumgroup, in that group the spermathecal head morphology isintermediate between the ‘blind lobe’ and ‘in-line’morphologies (HS – Figs 4, 53). A single, posteriorly located,small epigynal atrium (AT – Fig. 51) may also represent anautapomorphy of the angustiarum species group.

The Californian clade consists of six species groups. Theadenes and consocius groups are united by the presence of a shortembolus. In Cybaeus males, the genital bulb of most speciespossesses a basally enlarged andmedially and distally elongated,thin embolus (E – Fig. 17). In some species (especially within theadenes and consocius groups), the medial and distal portions ofthe embolus are relatively short and thick (E –Fig. 54).Within thegenus, the long, thin embolus state is plesiomorphic and the short,thick state is apomorphic. The anteriorly located and usuallysingle epigynal atrium (AT – Fig. 7) and the well developed andusually distinctively looped copulatory ducts (CD –Fig. 8) definethe consocius species group, whereas the very short copulatoryducts broadly joined to the atrium (CD – Fig. 5) define the adenesspecies group.

Of the four remaining Californian species groups, three(aspenicolens+ septatus+ tardatus groups) are united by thepresence of medially convergent copulatory ducts (CD –

Figs 95, 98). The plesiomorphic condition is for the copulatoryducts to diverge laterally from the epigynal atrium. Theaspenicolens+ septatus groups are further united by thepresence of one or more anterior epigynal ridges or ‘wrinkles’(EW – Figs 64, 93, 99). The integument of the epigynum in

females of most species of Cybaeus is relatively featurelessanterior to the atrium; this state is plesiomorphic. Thefollowing apomorphies define these species groups. Thetardatus species group possesses longitudinally elongated,medially located copulatory ducts (CD – Fig. 10). Thecopulatory ducts and spermathecal stalks are greatly reducedand arranged nearly linearly in the aspenicolens species group(CD, SS – Fig. 6). Within the male patellar apophysis in theseptatus group, a single peg seta is isolated from and/or largerthan the remaining peg setae (PS – Figs 88–90). Finally, thedevius group copulatory duct morphology (long and convolutedwith proximal sections looped about or enveloping distalsections (CD – Fig. 9)) is unique among cybaeids.

H3

A portion of the H3 gene was sequenced for 20 Cybaeusspecies and the outgroup Cybaeota nana (Cybaeidae). Thedata matrix consisted of 378 bases, except for C. tetricus, forwhich 25 bases were removed from the ends because they weredifficult to read unambiguously. There were no gaps. Maximumparsimony and likelihood analyses resulted in similar topologiesin that there was no disagreement with respect to stronglysupported (i.e. bootstrap) groupings (Fig. 1). For maximumparsimony, 27 characters were parsimony informative andthere were 142 most parsimonious trees with a treelengthof 92 and a consistency index of 0.84. Shortest trees werefound in 100 out of 100 random addition sequences.Maximum likelihood yielded one most likely tree with a scoreof –lnL = 1087.88. Parameters in the ML model consisted of thetransition/transversion ratio = 4.0, proportion of invariablesites = 0.61 and the shape parameter = 0.83.

COI

A portion of COI was successfully amplified and sequenced for13 Cybaeus species and the outgroup Cybaeota nana (Fig. 2).

Cybaeota nana

C. signifer (angustiarum group)

C. somesbar (septatus group)C. chauliodous (septatus group)

C. penedentatus (consocius group)C. gidneyi (unplaced)

C. sanbruno (adenes group)C. hesper (consocius group)

C. vulpinus (consocius group)

C. thermydrinos (aspenicolens group)

C. tetricus (tetricus group)

C. multnoma (tetricus group)C. paralypropriapus (tetricus group)

C. waynei (tetricus group)

C. reticulatus (angustiarum group)

C. giganteus (angustiarum group)C. patritus (angustiarum group)

C. shoshoneus (angustiarum group)

C. angustiarum (angustiarum group)

C. eutypus (tetricus group)C. morosus (tetricus group)

0.01

9765

90

97

5674

67

8571

77

51

76

8852

99

97

79

83

5371

87

9954

54

50

Fig. 1. Cybaeus nuclear histone subunit 3 (H3) maximum likelihood tree,with Cybaeota nana as an outgroup. Numbers above and below the branchesindicate parsimony and likelihood bootstrap percent support respectively.(Only bootstraps greater than 50% are shown.)


Five species did not yield a COI sequence and the sequencesgenerated for C. vulpinus andC. thermydrinos had the hallmarksof numt pseudogenes (mitochondrial DNA that has beentransferred into the nucleus) (Bensasson et al. 2001). Thesesequences had high similarity to the other Cybaeus sequences(and other spider sequences in GenBank), but the C. vulpinussequence contained two deletions (one was out of frame) and17 unique amino-acid substitutions and the C. thermydrinossequence contained nine unique amino-acid substitutions. Thefinal data matrix consisted of 709 bases, except for C. gidneyi,C. eutypus and C. giganteus, for which 21, 68 and 72 bases,respectively, were removed from the ends because they weredifficult to read unambiguously. There were no gaps. Formaximum parsimony, 134 characters were parsimonyinformative and there were two most parsimonious trees with atreelength of 492 steps and a consistency index of 0.60. Shortesttrees were found in 75 out of 100 random addition sequences; thisincreased to 95 out of 100 when we held five trees at each stepinstead of one. Maximum likelihood yielded one most likely treewith a score of –lnL = 3102.42. Parameters in the ML modelconsisted of the rate matrix: A–C=0.00, A–G=19.46,A–T= 1.94, C–G=1.12, C–T= 4.27, G–T= 1.00, proportionof invariable sites = 0.56 and the shape parameter = 1.27.

For the most part, there is not much disagreement between themitochondrial COI and H3 topologies and both show support forthe two major divisions (Holarctic and Californian) suggested bymorphology. Within the Holarctic clades, the two genes differwith respect to the placement of C. angustarium and C. signifer.Although there is not enough signal or taxon sampling to testhypotheses of relationships within the Californian clade, our H3dataset enables us to ask whether the species groups in theHolarctic clade (tetricus and angustarium) are reciprocallymonophyletic. This is rejected because a tree constrained on a

topology of reciprocal monophyly is significantly different fromthe ML tree (–lnL = 1188.52, SH test: P< 0.0001).

Taxonomic descriptions

Here we provide a redescription of the genus Cybaeus, as well asa key to the species groups. Information about the distributionsand number of species in each of the species groups arealso provided. Because most easily accessed illustrations anddescriptive material of both C. tetricus and C. angustiarum areinsufficiently detailed for phylogenetic purposes, these relativelycommon European species are redescribed here. Cybaeustetricus is also the type species of the genus and was used inthis study; for this reason, it is presented first. In addition, thenine species of previously undescribed Cybaeus species thatwere used in the phylogenetic analysis are described.

The descriptions follow the format of Bennett (1988).The genus description of Cybaeus is detailed and descriptionsof individual species emphasise differences from the basicCybaeus plan. Measurements are presented in mm as‘sample range (mean� standard error)’; ‘N’= sample size,‘CL’ and ‘CW’ = carapace length and width and ‘SL’ and‘SW’= sternum length and width. Paratype specimens listed ineach new species description are those specimens examined byBennett during the original descriptive work (Bennett 1991);subsequently identified specimens are listed as ‘other materialexamined’ and are not considered to be paratypes. Museumcollection acronyms are AMNH=American Museum of

C. somesbar (septatus group)

C. chauliodous (septatus group)

C. penedentatus (consocius group)

C. gidneyi (unplaced)

C. sanbruno (adenes group)

C. hesper (consocius group)

C. angustiarum (angustiarum group)

C. eutypus (tetricus group)

C. morosus (tetricus group)

C. signifer (angustiarum group)

Cybaeota nana

C. reticulatus (angustiarum group)

C. giganteus (angustiarum group)

C. patritus (angustiarum group)

95

81

100

60

59

76

92

58

99

81

100

74

93

90

97

83 54

0.0576

Fig. 2. Cybaeus cytochrome c oxidase subunit I (COI)maximum likelihoodtree, with Cybaeota nana as an outgroup. Numbers above and below thebranches indicate parsimony and likelihood bootstrap percent supportrespectively. (Only bootstraps greater than 50% are shown.)

Fig. 3. Cybaeus multnoma, male habitus, Crescent City, CA, dorsal.Scale bar = 2.0mm.


Natural History (New York NY), BM (NH) =British Museum(NaturalHistory) (LondonEngland), CAS=CaliforniaAcademyofSciences (SanFranciscoCA),MCZ=MuseumofComparativeZoology (Harvard MA), MNHN=Muséum naturelle d’histoirenaturelle (Paris France) and RBCM=Royal British ColumbiaMuseum (Victoria BC). Genitalic abbreviations are explained inrelevant figure legends.

Family CYBAEIDAE

Genus Cybaeus L. Koch

Amaurobius (in part) C.L. Koch, 1839: 43, fig. 462.Cybaeus L. Koch, 1868: 46, figs 22–25. – Chamberlin & Ivie, 1932: 3;Roewer, 1954: 88;Bonnet, 1956: 1299;Roth&Brame, 1972: 19,figs6,10, 20, 27, 28; Brignoli, 1983: 484; Roth & Brown, 1986: 3, 12;Platnick, 1989: 406; Platnick, 2009.

Parauximus Chamberlin, 1919: 2, plate I, fig. 2.Namopsilus Chamberlin, 1919: 14, plate VI, fig. 1.Type species: Amaurobius tetricus C.L. Koch, 1839.

Diagnosis

A definitive diagnosis of Cybaeus is lacking and will remain sopending a revision of the entire genus. The combined presence oftwo or three pairs of ventral tibia I macrosetae (Fig. 14) (rarelyfour orfive pairsmay be present but of these the second and fourthpairs will be lateral to the others, not in line as in the generaCybaeina Chamberlin & Ivie and Cybaeota Chamberlin & Ivie)and, in the male, a retrolateral patellar apophysis with peg setae(Figs 16, 20, 21) diagnoses Cybaeus. The females are not easilydiagnosed but are usually characterised by large spermathecalbases (BS) and relatively convoluted vulval ducts includinga single conspicuous, complex secondary pore (SP) on eachspermatheca (Figs 5, 8, 12, 31) (see Bennett 1992 fordiscussion of spermathecal pores).

Description

Small- to large-sized spiders (Fig. 3), carapace lengths averaging1.94–5.0 (females); sexes usually subequal. Carapace darkly

4

6

5

7

8

Figs4–8. Cybaeus spp., females: 4,C.patritus,GrandfatherMountain,NC,vulva, ventral; 5,C.adenes,Marin County, CA, vulva, dorsal; 6,C. aspenicolens, Aspen Valley, CA, vulva, ventral; 7,C. consocius,MarinCounty, CA, epigynum, ventral; 8,C. hesper, Claremont, CA, vulva, dorsal. Scale bars = 0.10mm.AT= epigynal atrium, BS= base of spermatheca, CD= copulatory ducts, HS = head of spermatheca,SP = complex spermathecal pore, SS = stalk of spermatheca.


pigmented around eyes, pale yellow to dark reddish brownelsewhere, longer than wide, glabrous except for small numberof setae along midline and around eyes; dorsal groove short,longitudinal. Eight eyes (Figs13, 15) in two rows, occasionally allreduced or absent; in dorsal view both rows slightly recurved; infrontal view anterior row straight, posterior row procurved;anterior median eyes usually reduced (Fig. 13), about as largeas others inC. signifer (Fig. 15); anterior laterals largest; posterioreyes subequal and slightly smaller than anterior laterals; medianocular quadrangle widest posteriorly, ~1–1 2/3 times height ofclypeus. Three teeth on promargin of cheliceral fang furrow,retromargin with variable number of teeth and denticles. Bosspresent on retrolateral margin of each cheliceral base.

Sternum shield-shaped, extending posteriorly slightlybetween coxae IV, longer than wide, pale yellow to darkreddish brown, lightly clothed with fine setae. Labium short,wider than long. Serrula well developed.

Legs pale yellow to dark reddish brown, unmarked or at leastfemora banded (tarsus not usually marked), IV–I–II–III in order ofdecreasing length, no scopulae (except tibia I inmaleC. scopulatusChamberlin & Ivie) or other modified hairs, with three claws.Trochanters unnotched. All femora, patellae, tibiae andmetatarsi with macrosetae; usually two or three complete pairsof ventral tibia Imacrosetae (terminal pairmay be present, absent,or incomplete), pattern often 2(�1p)-1p-2-1p-2 (Fig. 14).Tarsi and metatarsi each with single row (palpal tibiae withtwo rows) of trichobothria increasing in length distally. Tarsalorgan typically araneomorph.

Abdomen pale, unmarked to dark grey with light heart markand row of well defined light chevrons dorsally (Fig. 3), lightlyclothed with fine setae. Spiracle just anterior to and as wide ascolulus; colulus a low swelling usually marked by two groupsof up to ten setae each. Anterior lateral spinnerets broad,bisegmented, separated by width of colulus, as long as orlonger than posterior lateral spinnerets, which are narrow,bisegmented, separated by width of anal tubercle; posteriormedian spinnerets small, contiguous, unisegmented, oftentriangular in cross-section; apical segments of anterior andposterior lateral spinnerets subequal, much shorter than basalsegments.

Anterior retrolateral patellar apophysis present (RPA –

Fig. 16) but variable: stout, digitiform, as long as width ofpatella (Fig. 49) to reduced to a small, inconspicuous hump(Fig. 34). Patellar apophysis apparently absent in someJapanese species (Yaginuma 1986). Few to many peg setaescattered about (Fig. 49), or in one or two groups on dorsaland/or lateral surfaces of patellar apophysis. Retrolateral tibialapophysis a carinate process usually slightly produced anteriorlyand extending posteriorly nearly entire length of tibia (Fig. 19), insome species reduced to 1/2 or less of length of tibia (Bennett2006: figs 34, 35).

Genital bulb simple. Petiole very small, on ventral retrolateralsurface of well developed basal haematodocha. Subtegulumnarrow, annular; tegulum broader, rounded with slight tegularridge anteroventrally, lacking conductor or median apophysis(T, TR – Figs 17, 18). Tegular apophysis (AA, PA – Figs 17, 18,

9 10

11 12

Figs 9–12. Cybaeus spp., females: 9, C. devius, Madera County, CA, vulva, dorsal;10, Cybaeus sp. (tardatus group), San Ysidro, CA, vulva, dorsal; 11, C. septatus, ShastaCounty,CA, vulva, ventral; 12, same, dorsal. Scale bars = 0.10mm.BS= baseof spermatheca,CD= copulatory ducts, HS= head of spermatheca, SP = complex spermathecal pore,SS = stalk of spermatheca.


13 15

14

Figs 13–15. Cybaeus spp., females: 13, C. reticulatus, Haida Gwaii, BC, eye group, anterior;14, same, left tibia I, prolateral, illustrating 2(+1p)-1p-2-1p-2 arrangement of tibial macrosetae;15, C. signifer, South Pender Island, BC, eye group, anterior. Scale bars = 0.20mm.

0.48 mm 0.30 mm 0.38 mm

75 µm75 µm136 µm

Figs 16–21. Cybaeus spp., males, left palpus: 16,C. gidneyi, Brookings, OR, ventral; 17,C eutypus, Nanaimo, BC, genitalbulb, ventral; 18, C. cascadius, Eugene, OR, genital bulb, ventral; 19, C. waynei, Forest Glen, CA, retrolateral tibialapophysis; 20, same, patellar apophysis, dorsal; 21, C. cascadius, Eugene, OR, tip of patellar apophysis, anterodorsal.AA= anterior portion of tegular apophysis, E = embolus, PA= posterior portion of tegular apophysis, RPA= retrolateralpatellar apophysis, RTA= retrolateral tibial apophysis, T = tegulum, TR= tegular ridge.


47, 48) prominent, retrolateral, attached to embolar base,supporting embolus; with anterior, occasionally speciesspecific, flange-like portion; medial trough; and usuallyspecies specific posterior portion. Embolus enlarged basally;long, narrow (E – Figs 17, 47, 48) to relatively short, thick(E – Figs 54, 55); usually smoothly curved clockwise (leftpalpus, ventral view). Receptaculum seminis (Figs 22–24)coiled through ~540� from fundus (F) to tip of ejaculatory duct(ED); base of fundus within subtegulum, heavily sclerotiseddistally, with relatively large lumen proximally, passingthrough middle haematodocha into tegulum; reservoir (R)usually slender but occasionally quite broad (Figs 23, 24), wellsclerotised, appressed to outer margin of tegulum, graduallynarrowing into long, thin ejaculatory duct in embolus.Epigynum very simple, lightly sclerotised; with atrium single(Figs 36, 51) or paired (Figs 93, 96, 106, 109), variously located;vulval components discernible to varying degree throughintegument. Copulatory ducts variable: separated (Figs 53,108, 111) or contiguous (Bennett 2006: figs 9, 11, 29) atatrium, usually relatively short (Figs 59, 60) or rarelyelongated, lightly or heavily sclerotised (see Bennett 2006).Heads of spermathecae with primary pores and of variableform: distinct lobes connected to spermathecal stalks/copulatory ducts by narrow duct (Bennett 2006; Figs 67, 99,109) to undifferentiated from rest of vulva and identifiableonly by presence of group of simple primary pores (Fig. 65).Stalks of spermathecae each with a single complex pore (Figs 5,12, 31) and of variable form: long, sinuous, gradually dilatingdistally (Bennett 2006: figs 46–49) to very short, undifferentiated

(Figs 59, 60, 72, 75, 78). Complex spermathecal pores usuallydorsolateral and near distal end of stalks. Spermathecalbases prominent; rounded or elongated with single,uncompartmentalised, large lumen (Figs 31, 72). Fertilisationducts short, inconspicuous; attached to distal end of stalks(Fig. 31) or to bases near to junction with stalks (Figs 59, 60).

Distribution

Holarctic with species described from Europe, China, Korea andespecially Japan and western North America. In North America,Cybaeus is mostly found in forested locales west of the westerncontinental divide; three species occur in the forested uplands ofthe eastern USA.

Etymology

Cybaeus is derived from a nominative masculine adjectivereferring to a cybaea, a type of ancient merchant ship (Bennett1991; Cameron 2005).

Key to Nearctic Cybaeus species groups (females only)

1. Spermathecal heads, stalks and bases forming three distinctly bulbousregions on each half of vulva (Figs 31, 38, 45, 53). Heads more(Figs 31, 38) or less (Figs 4, 53) continuous with rest of vulval ducting,occasionally on separate, distinct lobes (e.g. C. signifer Simon,C. reticulatus Simon) .........................................................................2

Spermathecal heads, stalks and bases otherwise; not forming threedistinctly bulbous regions on each half of vulva (Figs 5, 8, 60,75, 84, 101). Heads usually set somewhat apart from rest of vulvalducting................................................................................................3

22

23 24

Figs 22–24. Cybaeus spp., males, left genital bulb, receptaculum seminis: 22,C.multnoma,Klickitat County, WA, ventral; 23, C. perditus, Cape Alava Trail, WA, prolateral; 24, same,ventral. Scale bars = 0.10mm. AA= anterior portion of tegular apophysis, ED= ejaculatoryduct of receptaculum seminis, F = fundus of receptaculum seminis, PA= posterior portionof tegular apophysis, R = reservoir of receptaculum seminis.


2. Spermathecal heads continuous and in line with rest of vulval ducting(Figs 31, 38)...................................................................tetricus group

Spermathecal heads offset somewhat from rest of vulval ducting(Figs 4, 53)...........................................................angustiarum group

3. Vulval ducting reduced, sinuous with stalks and laterallyprojecting copulatory ducts very short (Figs 5, 60). Atrium single(Fig. 58) ......................................................................... adenes group

Not exactly as above; either vulval ducting not reduced and sinuousor copulatory ducts not short and/or projecting to lateral margins ofvulva (Figs 9, 10, 65, 71, 98). Atrium single (Figs 7, 71) or paired(Figs 93, 99).......................................................................................4

4. Copulatory ducts long, well developed (Figs 8–10, 72, 75, 78, 84).Epigynum usually without features other than atrium (Fig. 7)..........5

Copulatory ducts relatively short (Figs 6, 11, 12, 66, 95, 98, 101).Epigynum usually with anterior, transverse ridges or ‘wrinkles’(Figs 64, 93, 99).................................................................................7

5. Copulatory ducts relatively straight, longitudinally oriented (Fig. 10);vulva triangular ............................................................ tardatus group

Copulatory ducts convoluted, laterally located (Figs 8, 9, 72, 78, 84); vulvamore box-shaped, not triangular ........................................................6

6. Copulatory ducts with proximal sections partially enveloping or loopedabout more distal sections (Fig. 9). Atrium single.......... devius group

Copulatory ducts often distinctly looped but never with proximal sectionsenveloping or looped about more distal sections (Figs 8, 72). Atriumsingle (Figs 7, 71) or paired.......................................consocius group

7. Vulva very simple with copulatory ducts and spermathecal stalks muchreduced, nearly linear (Figs 6, 65) ........................aspenicolens group

Vulva more complex, copulatory ducts and spermathecal stalks not soreduced or linear (Figs 12, 95, 101)............................. septatus group

tetricus species group

The tetricus group is based on the Palaearctic type species of thegenus, C. tetricus (C.L. Koch) and includes nine described(including C. paralypropriapus, sp. nov. and C. waynei,

25 26

27 28

Figs 25–28. Cybaeus tetricus, male, left palpus: 25, Chamonix, France, bulb, ventral; 26, unknownlocale, tegular apophysis, retrolateral; 27, Chamonix, France, patella, dorsal; 28, unknown locale,tibia, retrolateral. Scale bars = 0.10mm. PA= posterior portion of tegular apophysis,RPA= retrolateral patellar apophysis, RTA= retrolateral tibial apophysis. Figures 25, 28 redrawnfrom Bennett (1991: figs 28, 30).


sp. nov., described below) and one undescribed western Nearcticspecies. As with the Nearctic angustiarum species, several of thetetricus species are very common and wide-ranging woodlandspiders. The Nearctic tetricus species are found primarily incoastal localities and occur from the middle of the AlaskanAleutian Islands south to Marin County, California and inlandas far as the Rocky Mountains of northern British Columbia andthe western slopes of the Cascade Range in Oregon.

Cybaeus tetricus (C.L. Koch)

(Figs 25–31)

Amaurobius tetricus C. L. Koch, 1839: 43.Cybaeus tetricus L. Koch, 1868: 50, figs 24, 25. – Chamberlin & Ivie,1932: 11,figs 22, 23;Roewer, 1954: 88;Bonnet, 1956: 1304;Heimer&Nentwig, 1991: 360, fig. 395; Jocqué & Dippenaar-Schoeman, 2006:80, fig. 17a–f; Platnick, 2009.

Material examined

Types. Six syntypes (2 <, 3 ,, one immature) from ‘Bavarian Alps’, inBM (NH), examined.

Other material examined. 2 <, 30 ,, no data (MNHN); France: 1 <,‘Gallia’, no date, (AMNH); Slovenia: 1 <, Veliki Otok, 2 km NW ofPostojna, 45.7917N 14.2014E, 13.vi.2008, coll. F. Stahlavski (RBCMcatalogue number ENT008–008606) (GenBank accession number:FJ263770); Switzerland: 2 <, 2 ,, Basel, no date, coll. Schenkel(AMNH); 1 <, Chamonix, 3.viii.1932, coll. Crosby (AMNH).

Diagnosis

The form of the posterior portion of the tegular apophysis(Figs 25, 26) diagnoses the male of C. tetricus. The female is

distinguished by the form of the vulva (Figs 30, 31) and atrium(Fig. 29).

Description

Abdomen patterned. Legs unbanded. Ventral tibia I macrosetaevariable: 2(�1p)-1p-2-1p(or 0)-1p(or 2).

Male

Patellar apophysis (Fig. 27) broad basally, relatively short,length ~2/3 width of patella; ~50 peg setae. Retrolateral tibialapophysis nearly as long as tibia (Fig. 28). Embolus long andnarrow. Posterior arm of tegular apophysis (Figs 25, 26) withbulbous base and ventrally directed, slender, uncinate tip.Variation (n= 8): CL 2.8–3.4 (3.1� 0.2); CW 1.95–2.50(2.22� 0.19); SL 1.33–1.60 (1.46� 0.11); SW 1.23–1.50(1.38� 0.10). Syntypes CL 3.1, 3.2; CW 2.15, 2.43; SL 1.40,1.58; SW 1.43, 1.45.

Female

Atrium single, large, nearly as long aswidth of vulva (Fig. 29).Vulva (Figs 30, 31) with spermathecal heads, stalks and basesevident as three distinct bulbous regions on each half of the vulvaand the heads continuous with the rest of the vulval components(not on distinct, separate lobes) (Fig. 31). Copulatory ducts(Figs 30, 31) widely separated at atrium but converging andcontiguous at midline of vulva. Variation (n = 11): CL 2.8–3.6(3.2� 0.2); CW 1.83–2.45 (2.18� 0.18); SL 1.33–1.68(1.49� 0.10); SW 1.17–1.50 (1.37� 0.11). Syntypes CL 3.3,3.4, 3.5; CW2.25, 2.35, 2.45; SL 1.53, 1.55, 1.60; SW1.43, 1.48,1.50.

29

30

31

Figs 29–31. Cybaeus tetricus, female, BavarianAlps: 29, epigynum, ventral; 30, vulva, ventral; 31,same,dorsal. Scale bars = 0.10mm.AT= epigynal atrium,BS= baseof spermatheca,CD= copulatoryducts, EG= epigastric groove, FD= fertilisation duct, HS= head of spermatheca, SP = complexspermathecal pore, SS = stalk of spermatheca.


Distribution

Central and eastern Europe (Bonnet 1956).

Cybaeus paralypropriapus Bennett, sp. nov.

(Figs 32–38, 113)

Material examined

Holotype. 1 <, USA: Oregon, ten miles south-south-west of Coos Bay,43.2667N 124.2833W, 17.vii.1968, coll. W. Ivie (AMNH).

Paratypes. USA: 2 <, 2 ,, Oregon, Coos County, Camp Myrtlewood,nr. Bridge, 28.vii.1954, coll. V.D. Roth (CAS); 1<, 2 ,, Oregon, CoosCounty, Camp Myrtlewood, nr. Bridge 27.vii./4.viii.1955, coll.V.D. Roth (CAS); 1 ,, Oregon, Coos County, Charleston, 10.vii.1951, coll. B. Malkin (AMNH); 2 ,, Oregon, Coos County,Charleston, 27.v.1952, coll. V.D. Roth (CAS); 8 ,, Oregon, Coos

County, Charleston, 30.ix.1959, coll. V.D. Roth (CAS); 1 ,, Oregon,Curry County, 20 mi. N Gold Beach, 15.ix.1971, coll. V.D. Roth(CAS); 1 ,, Oregon, Curry County, 1 mi. N Sixes, 30.ix.1959, coll.V.D. Roth (CAS); 3 ,, Oregon, Douglas County, 1 mi. S Reedsport,30.ix.1959, coll. V.D. Roth (CAS).

Othermaterial examined. USA:3,,Oregon,DouglasCounty,Elliot St.For. SR7500, off Hwy 38 beside Mill Ck., 43.6472N 123.8833W,23.ix.2003, coll. P. Paquin, D. Wytrykush (RBCM catalogue numberENT008–008610) (GenBank accession number: FJ263776).

Diagnosis

The male of C. paralypropriapus is diagnosed by the tiny, knob-like patellar apophysis (Fig. 34) and the form of the posterior armof the tegular apophysis (Figs 32, 33). The form of the atrium(Fig. 36) and vulva, especially the copulatory ducts (Figs 37, 38),distinguish the female.

32 33

3534

Figs 32–35. Cybaeus paralypropriapus, paratype male, near Bridge, OR, left palpus: 32, bulb,ventral; 33, tegular apophysis, retrolateral; 34, patella, dorsal; 35, tibia, retrolateral. Scalebars = 0.10mm. PA= posterior portion of tegular apophysis, RPA= retrolateral patellarapophysis, RTA= retrolateral tibial apophysis. Figures redrawn from Bennett (1991, figs321–324).


Description

Abdomen patterned. Legs banded (at least femora). Ventral tibia Imacrosetae: 2(�1p)-1p-2-1p-2.

Male

Very small, knob-like patellar apophysis (Fig. 34) onlongitudinal lateral swelling of patella; less than ten peg setaeclustered about tip. Retrolateral tibial apophysis nearly as long astibia (Fig. 35). Embolus relatively long, narrow. Posterior arm oftegular apophysis (Figs 32, 33) with long, narrow, sickle-shaped,prolaterally directed tip. Variation (n= 4): CL 4.2–5.3 (4.7);CW 3.1–4.0 (3.5); SL 1.90–2.30 (2.10); SW 1.70–2.00 (1.85).Holotype CL 5.0, CW 3.6, SL 2.25, SW 1.93.

Female

Single, small but conspicuous, transverse, medially locatedatrium (Fig. 36). Vulvawith spermathecal heads, stalks and basesevident as three distinct bulbous regions on each half of vulva;heads continuous with rest of vulval components (not on distinct,separate lobes) (Fig. 38). Copulatory ducts (Figs 37, 38) broad,proceeding directly anterior from edges of atrium, completely orpartly contiguous or separated entirely; diverging, narrowing andturning posteriorly at anterior edge of vulva. Spermathecal basesrelatively small. Variation (n= 19): CL 4.2–5.6 (5.0� 0.4); CW2.7–3.6 (3.3� 0.2); SL 1.85–2.38 (2.20� 0.14); SW 1.68–2.08(1.91� 0.11).

Distribution

Coastal regions of Coos, Curry and Douglas Counties in south-western Oregon (Fig. 113).

Etymology

The specific epithet is derived from the Greek adjectiveparalypros, meaning ‘rather poor’ and Priapos, the ancientGreek god of reproduction. It refers to the diminutive size ofthe male patellar apophysis.

Cybaeus waynei Bennett, sp. nov.

(Figs 19, 39–46, 112)

Material examined

Holotype. 1 ,, USA: Oregon, Jackson County, Ashland, Lithia Park31.vii.1959, coll. V.D. Roth, W.J. Gertsch (AMNH).

Paratypes. USA: 1 ,, California, Del Norte County, Crescent City,16.xi.1961, coll. W. Ivie, W.J. Gertsch (AMNH); 1 ,, California, DelNorte County, 3mi. EKlamath, 16.xi.1961, coll.W. Ivie,W.J. Gertsch(AMNH); 1 <, California, Humboldt County, 18 mi. S Klamath,13.viii.1953, coll. G.A. Marsh, R.O. Schuster (AMNH); 1 <, 1 ,,California, Mendocino County, 4 mi. S Rockport, 19.viii.1959, coll.V.D. Roth, W.J. Gertsch (AMNH); 4 <, 3 ,, California, MendocinoCounty, Rockport, 19.vii.1962 coll. V.D. Roth (CAS); 3 <, 19 ,,California, Trinity County, 2 mi. W Forest Glen, 21.viii.1959, coll.V.D. Roth, W.J. Gertsch (AMNH); 2 <, Oregon, Coos County,Charleston, vii.1950, coll. V.D. Roth (CAS); 1 <, Oregon, CurryCounty, 2 mi. N Gold Beach, 30.v.1948, coll. V.D. Roth, E. Brown

36

38

37

Figs 36–38. Cybaeus paralypropriapus, paratype female, Charleston, OR: 36, epigynum,ventral; 37, vulva, ventral; 38, same, dorsal. Scale bars = 0.10mm. AT= epigynal atrium,BS = base of spermatheca, CD= copulatory duct, HS = head of spermatheca, SS = stalk ofspermatheca.


(CAS); 1 <, Oregon, Curry County, 7 mi. E Brookings, Chetco R.,29.v.1952, coll. V.D. Roth (CAS); 1<, Oregon, Curry County, 8 mi. EGold Beach, Rogue R., 28.v.1952, coll. V.D. Roth (CAS); 1 <, 1 ,,Oregon, Curry County, Pistol River, 21.vi.1953, coll. B. Malkin(AMNH); 7 ,, Oregon, Curry County, Pistol River, 17.ix.1956, coll.B. Malkin (AMNH); 1 ,, Oregon, Douglas County, IslandCampground, 0.5 mi. S, 1 mi. E Steamboat, Hwy 138, 1200’,30.x.1971, coll. E.M. Benedict (MCZ); 1 <, Oregon, DouglasCounty, Susan Ck., E Glide, 23.vii.1962, coll. V.D. Roth (CAS);8 <, 27 ,, Oregon, Jackson County, Ashland, Lithia Park31.vii.1959, coll. V.D. Roth, W.J. Gertsch (AMNH); 2 <, 3 ,,Oregon, Jackson County, 25 mi. NE Ashland, 1.ix.1959, coll. W.J. Gertsch, V.D. Roth (AMNH); 1 <, Oregon, Jackson County, 8mi. S Divide Guard Sta., N Trail, 27.vii.1955, coll. V.D. Roth (CAS);1 <, 2 ,, Oregon, Klamath County, Crater Lake Nat. Pk., lakeshore to

79000, 7.ix.1941, coll. B. Malkin (AMNH); 1 ,, Oregon, KlamathCounty, Crater Lake Nat. Pk., 65000, 27.viii.1950 coll. B. Malkin(AMNH); 2 <, 2 ,, Oregon, Klamath County, Crater Lake Nat. Pk.,Lake Ck., no date, coll. D. Lowrie? (AMNH); 1 ,, Oregon, LaneCounty, Willamette Pass, 5000–5200’, 7.ix.1954, coll. B. Malkin(AMNH).

Othermaterial examined. USA:1,,Oregon,DouglasCounty,Elliot St.For. SR7500, off Hwy 38 beside Mill Ck., 43.6472N 123.8833W,23.ix.2003 coll. P. Paquin, D. Wytrykush (RBCM catalogue numberENT008–008611) (GenBank accession number: FJ263778).

Diagnosis

The form of the patellar apophysis (Figs 41, 42) and the posteriorarm of the tegular apophysis (Figs 39, 40) diagnose the male of

39 40

41 42

Figs39–42. Cybaeuswaynei, paratypemales, left palpus: 39, ForestGlen,CA,bulb, ventral;40, same, tegular apophysis, retrolateral; 41, same, patella, dorsal; 42, Pistol River, OR, tibiaand tip of patella, retrolateral. Scale bars = 0.10mm. PA= posterior portion of tegularapophysis, RPA= retrolateral patellar apophysis, RTA= retrolateral tibial apophysis.Figures redrawn from Bennett (1991: figs 26, 91–93).


C. waynei. The female is distinguished by the form of the atrium(Fig. 43) and vulva (Figs 44–46), especially the configuration ofthe copulatory ducts and the spermathecal stalks.

Description

Abdomen patterned. Legs banded (at least femora). Ventral tibia Imacrosetae: 2(�1p)-1p-2-1p-2.

Male

Patellar apophysis (Figs 41, 42) triangular, length ~2/3width of patella, 20 to 30 tightly grouped peg setae on dorsalsurface. Retrolateral tibial apophysis nearly as long as tibia(Figs 19, 42). Embolus relatively long, narrow. Posteriorarm of tegular apophysis (Figs 39, 40) with simple, elongate,prolaterally directed, acuminate tip. Variation (n= 25):CL 2.7–4.3 (3.5� 0.4); CW 1.95–3.1 (2.5� 0.3); SL 1.33–2.03(1.64� 0.17); SW 1.23–1.75 (1.46� 0.14).

Female

Atrium (Fig. 43) single, small, transverse, posteriorly locatedon epigynum. Vulva with spermathecal heads, stalks and basesevident as three distinct bulbous regions on each half of vulva;heads continuous with rest of vulval components (not on distinct,separate lobes) (Figs 44–46). Copulatory ducts (Figs 44, 46)often conspicuously visible through integument of unclearedepigyna; divergent at atrium, convergent and usuallycontiguous at heads of spermathecae (Figs 44, 45). Variation(n= 32): CL 2.9–4.8 (3.8� 0.6); CW 1.95–3.3 (2.6� 0.4);SL 1.35–2.18 (1.73� 0.24); SW 1.25–1.88 (1.52� 0.20).Holotype CL 3.4, CW 2.33, SL 1.63, SW 1.45.

Distribution and natural history

South-western Oregon from Douglas County south to northernMendocino County in north-western California (Fig. 112).Specimens have been collected from under rocks in wet forest;males have been collected from late May until early September.

Etymology

The specific epithet is a patronym honouring the late JohnnyWayne.Heprovided anexcellent solution to theminor problemofa name for this species when he died the day the original draft ofthis description was prepared (18 July 1990). A Canadiancomedian, Johnny Wayne was a master of the doubleentendre. His often twisted sense of humour appealed toseveral generations of Canadians of all ages.

angustiarum species group

The angustiarum group is based on the Palaearctic speciesC. angustiarum L. Koch and includes nine described and threeundescribed Nearctic species. Several are very common andwide-ranging western Nearctic woodland spiders. Threedescribed Cybaeus species are the only ones known fromeastern North America and range from north-westernNew York and adjacent Canada to north-eastern Alabama andnorthernGeorgia. All but one of the others occur inwesternNorthAmerica from Haida Gwaii (Queen Charlotte Islands) in BritishColumbia to the Rocky Mountains of Alberta south to northernCalifornia and western Montana. One species (C. reticulatusSimon) ranges from the tip of the Alaskan Aleutian Islands southto north-western California.

43 46

44 45

Figs 43–46. Cybaeus waynei, paratype females: 43, Ashland, OR, epigynum, ventral; 44,same, vulva, ventral; 45, same, dorsal; 46,Rockport,CA,vulva, ventral. Scale bars = 0.10mm.AT= epigynal atrium, BS= base of spermatheca, CD= copulatory duct, HS = head ofspermatheca, SS = stalk of spermatheca.


Cybaeus angustiarum L. Koch

(Figs 47–53)

Cybaeus angustiarum L. Koch, 1868: 47, figs 22, 23. – Dahl, 1931: 11,figs 16, 17; Roewer, 1954: 88; Bonnet, 1956: 1300; De Blauwe, 1973:12, figs 9–11; Platnick, 2009.

Cybaeus angustiorum: Heimer & Nentwig, 1991: 360, fig. 934.

Material examined

Types. 1 <, 1 ,, ‘from type series’ (no other data), in BM(NH),examined.

Other material examined. 1 <, 1 ,, no data, (MCZ); Czech Republic:5 ,, Beskydy Mountains, Libotin Wood, near Novy Jicin, BeskydyMts., 9.ix.2006, coll. J. Kral (RBCM); Poland: 1 ,, Czorsztyn,16.viii.1959 (AMNH).

Diagnosis

The large, thumb-like patellar apophysis (Fig. 49) diagnoses themale of C. angustiarum. The female is diagnosed by the form of

the epigynum (Fig. 51) and the copulatory ducts (Fig. 52) as wellas the general configuration of the vulva (Figs 52, 53).

Description

Abdomen patterned, legs unbanded.

Male

Patellar apophysis (Fig. 49) large, nearly as long as width ofpatella; anteriorly directed, with ~50 peg setae. Retrolateral tibialapophysis nearly as long as tibia (Fig. 50). Embolus relativelyshort and thin. Tegular apophysis (Figs 47, 48) relatively simple,posterior arm with short, stout, prolaterally directed acuminatetip. Variation: CL 3.8, 4.0; CW 2.5, 2.8; SL 1.78, 1.80; SW 1.53,1.58. Male from type series listed first.

Female

Epigynum (Fig. 51) with very small atrium; proximalsection of spermathecal heads usually visible as pair of

47 48

49 50

Figs 47–50. Cybaeus angustiarum, type series male, left palpus: 47, bulb, ventral; 48,tegular apophysis, retrolateral; 49, patella, dorsal; 50, tibia, retrolateral. Scale bars = 0.10mm.AA= anterior portion of tegular apophysis, E = embolus, PA= posterior portion of tegularapophysis, RPA= retrolateral patellar apophysis.


anterior transverse bars on epigynum. Copulatory ducts (Fig. 52)bowed, lightly sclerotised. Vulva (Figs 52, 53)with spermathecalheads, stalks and bases evident as three distinct bulbous regionson each half of the vulva; heads set somewhat apart from therest of the vulval components and relatively small (Fig. 53).Variation: CL 3.3, 3.4, 3.9; CW 2.25, 2.25, 2.7; SL 1.53, 1.55,1.75; SW 1.38, 1.43, 1.60. Female from type series listed second.

Distribution

Widespread in Europe (Bonnet 1956).

adenes species group

The adenes group, based on C. adenes Chamberlin & Ivie,consists of five described (including C. sanbruno, sp. nov.,described below) and four undescribed species with restrictedranges, occurringprimarily inwest central coastalCalifornia fromSonoma and Napa Counties south to northern Monterey County.One species occurs in the Sierra Nevada of Placer and EldoradoCounties in east central California.

Cybaeus sanbruno Bennett, sp. nov.

(Figs 54–60, 112)

Material examined

Holotype. 1 <, USA: California, San Mateo County, San BrunoMountain, XL Cave Canyon, 14.xii.1980 coll. D. Ubick (CAS).

Paratypes. USA: 1 ,, California, San Mateo County, San Bruno Mtn.,XL Cave Cñ, 27.vii./3.viii.1980, coll. D. Ubick (D. Ubick collection);1<, 4,, California, SanMateoCounty, SanBrunoMountain, XLCaveCanyon, 14.xii.1980, coll. D. Ubick (D. Ubick collection); 1 <, 1 ,,California, San Mateo County, San Bruno Mtn., XL Cave Cñ,30.xii.1980, coll. D. Ubick (D. Ubick collection); 1 ,, California,

San Mateo County, San Bruno Mtn., 20.i.1981, coll. D. Ubick(D. Ubick collection); 1 <, 2 ,, California, San Mateo County, SanBruno Mtn., XL Cave Cñ, 28.ix.1981, coll. D. Ubick (D. Ubickcollection); 1 <, California, San Mateo County, San Bruno Mtn.,XL Cave Cñ, 28.ix./16.xi.1981, coll. D. Ubick (D. Ubick collection).

Other material examined. USA: 1 ,, California, San Mateo County, Wboundary San Bruno Mtn State Park across from JFK School, ~100m,37.6943N 122.4530W, 12.iii.2006, coll. M. Hedin (RBCM cataloguenumber ENT008–008618) (GenBank accession numbers: FJ263780,FJ263795).

Diagnosis

The male of C. sanbruno is diagnosed by the form of the patellarapophysis (Figs 56, 57) and, in particular, the formof the posteriorarm of the tegular apophysis (Figs 54, 55). The female isdistinguished by the form of the atrium (Figs 58, 59) and thevulva (Figs 59, 60).

Description

Abdomen usually patterned, legs unbanded.

Male

Patellar apophysis (Figs 56, 57) prominent, digitiform, nearlyas long as width of patella, with 13 peg setae on anterodorsalsurface. Retrolateral tibial apophysis nearly as long as tibia, withadditional small carina dorsally (Fig. 57). Embolus relativelyshort, thick. Posterior arm of tegular apophysis (Figs 54, 55) bifiddistally, pincer-like with tips directed towards each other, smallbut prominent posteriorly directeddorsal ‘keel’.Variation (n= 5):CL 2.33–2.6 (2.4); CW 1.68–1.93 (1.80); SL 1.18–1.30 (1.22);SW 1.10–1.24 (1.15). Holotype CL 2.43, CW 1.80, SL 1.24,SW 1.14.

51

53

52

Figs 51–53. Cybaeus angustiarum, female: 51, Czorsztyn, Poland, epigynum, ventral; 52, typeseries, vulva, ventral; 53, same, dorsal. Scale bars = 0.10mm. AT= epigynal atrium, BS= base ofspermatheca, CD= copulatory duct, HS= head of spermatheca, SS = stalk of spermatheca.


Female

Epigynum (Fig. 58) with anteriorly located invertedU-shapedatrium, atrium widening posteriorly; posterior ends curvedanteriorly (inconspicuous but visible in cleared epigynum withcompoundmicroscope); height of atrium (from epigastric grooveto anterior margin) 1.75 times to twice width. Vulva (Figs 59, 60)simple, each half sinuous. Copulatory ducts very short, separatedat atrium but with conspicuous, contiguous (or nearly so),sclerotised matrices. Spermathecal heads simple, small, dorsallobes.Variation (n= 9): CL2.18–3.2 (2.5); CW1.50–2.13 (1.73);SL 1.11–1.48 (1.26); SW 1.04–1.40 (1.17).


Known only from San Bruno Mountain, San Mateo County,California (Fig. 112). Theholotype andparatypes collected on thesame date were in Ceanothus leaf litter. Pitfall traps (one in

chaparral) and hand collecting (under rocks and beating ferns)produced the other paratypes. The remaining specimenwas foundunder rocks in chaparral.

Etymology

The specific name is a noun in apposition taken from the typelocality.

aspenicolens species group

The aspenicolens group, based onC. aspenicolensChamberlin&Ivie, contains three described (including C. thermydrinos,sp. nov., described below) and two undescribed speciesentirely restricted to very small ranges on the western slopesof the central and southern Sierra Nevada in California.

54 55

56 57

Figs 54–57. Cybaeus sanbruno, paratype male, San Bruno Mtn, CA, left palpus: 54, bulb,ventral; 55, tegular apophysis, retrolateral; 56, patella and tibia, dorsal; 57, patellar apophysis andtibia, retrolateral. Scale bars = 0.10mm.E= embolus,K = tegular apophysis ‘keel’, PA= posteriorportion of tegular apophysis, RPA= retrolateral patellar apophysis, RTA= retrolateral tibialapophysis. Figures 54, 56, 57 redrawn from Bennett (1991: figs 207–209).


Cybaeus thermydrinos Bennett, sp. nov.

(Figs 61–66, 112)

Material examined

Holotype. 1 <, USA: California, Tulare County, 2.5 miles east ofCalifornia Hot Springs, matured 5.x.1959, coll. W.J. Gertsch,V.D. Roth (CAS).

Paratypes. USA: 1 ,, California, Tulare County, 2.5 miles east ofCalifornia Hot Springs, 5.x.1959, coll. W.J. Gertsch, V.D. Roth(CAS); 3 <, 4 ,, California, Tulare County, 10 mi. W Johnsondale,15.ix.1959, coll. W.J. Gertsch, V.D. Roth (AMNH); 2 <, California,Tulare County, 10 mi. W Johnsondale, matured 20.ix.1959, coll.W.J. Gertsch, V.D. Roth (CAS).

Other material examined. USA: 1<, 2 ,, California, Kern County, 2 miW of Greenhorn Summit, Hwy 155, ~50000, 35.7333N 118.5667W,09.xi.2003, coll. P. Paquin, D. Wytrykush (RBCM catalogue numberENT008–008619) (GenBank accession number: FJ263783).

Diagnosis

The male of C. thermydrinos is diagnosed by the form of thepatellar apophysis (Fig. 63) and the posterior arm of the tegularapophysis (Figs 61, 62). The female is distinguished by the formof the epigynum, atria, copulatory ducts and spermathecal stalks(Figs 64–66).

Description

Abdomen patterned, femora at least lightly banded.Ventral tibia Imacrosetae 2-1p-2-1p-0(or 1p). Most paratypes teneral.

Male

Patellar apophysis (Fig. 63) longer than wide, length equal towidth of patella, gradually tapering to dull point with 20–30 largepeg setae along dorsal and lateralmargins of distinct concavity on

anterior margin of apophysis, distal-most peg setae often insingle line. Retrolateral tibial apophysis nearly as long as tibia.Embolus relatively short, thick (Figs 61, 62). Posterior arm oftegular apophysis (Figs 61, 62) prolaterally directed with ventral‘keel’ and simple, acuminate tip posterodorsal to keel. Variation(n= 6): CL 2.5–2.8 (2.6); CW 1.78–2.10 (1.92); SL 1.21–1.33(1.26); SW 1.18–1.30 (1.23). Holotype 2.6, CW 1.90, SL 1.26,SW 1.22.

Female

Epigynumwith large transverse epigynal ‘wrinkle’ anteriorly;single (Fig. 64) or divided (Fig. 66) atrial ridgepartially obscuringand continuous with pair of inconspicuous, longitudinal,medially located atria. Vulva (Figs 65, 66) very simple withmuch reduced, nearly linear components. Copulatory ducts veryshort, exiting atria towards midline of vulva, heavily sclerotised,contiguous posteriorly. Heads of spermathecae on very smalldorsal lobes near midline of vulva, not much differentiated fromrest of spermathecae. Stalks of spermathecae very short, notcontiguous. Variation (n= 5): CL 2.30–2.6 (2.4); CW1.63–1.83 (1.70); SL 1.14–1.25 (1.20); SW 1.13–1.24 (1.17).

Distribution

Southern Tulare County, California on the western slopes of thesouthern Sierra Nevada (Fig. 112).

Etymology

The specific epithet is derived from the Greek for ‘hot spring’and refers to the type locality.

58

59 60

Figs 58–60. Cybaeus sanbruno, paratype female, San Bruno Mtn, CA: 58, epigynum,ventral; 59, vulva, ventral; 60, same, dorsal. Scale bars = 0.10mm. AT= epigynal atrium,BS= base of spermatheca, CD= copulatory duct, FD= fertilisation duct, HS= head ofspermatheca, SS = stalk of spermatheca.


consocius species group

The consocius group, based on C. consocius Chamberlin & Ivie,contains five described (including C. penedentatus, sp. nov. andC. vulpinus, sp. nov., described below) and four undescribedspecies. Nearly all of these species occur in California in smallranges on the coast from southernMendocinoCounty toMontereyCounty and in the Sierra Nevada from Plumas County south toTuolumne County. One species is restricted to the central coast ofOregon.

Cybaeus penedentatus Bennett, sp. nov.

(Figs 67–78, 113)

Material examined

Holotype. 1 <, USA: California, Santa Clara County, two miles northof Holy City, Highway 17, from Redwood duff, 6.xii.1966, coll.V.D. Roth, Ferguson (CAS).

Paratypes. USA: 1 ,, California, San Mateo County, 6 mi. SEHalf Moon Bay, 5.xii.1953, coll. V.D. Roth (CAS); 1 ,,California, San Mateo County, Pescadero Ck., SE Half MoonBay, 26.iv.1959, coll. R.O. Schuster (AMNH); 1 ,, California,Santa Clara County, two miles north of Holy City, Highway17, from redwood duff, 6.xii.1966, coll. V.D. Roth, Ferguson(CAS); 1 ,, California, Santa Clara County, Mt. Madonna,2.i.1954, coll. R. Burdick (CAS); 1 ,, California, Santa CruzCounty, Ben Lomond, 6.vii.1956, coll. V.D. Roth, W.J. Gertsch(AMNH); 2 ,, California, Santa Cruz County, Ben Lomond,6.vii.1956, coll. V.D. Roth, W.J. Gertsch (CAS); 1 ,, California,Santa Cruz County, Boulder Creek, 23.xii.1953, coll. V.D. Roth(CAS); 1 ,, California, Santa Cruz County, Brookdale [between BenLomond and Boulder Creek], iii.1913, coll. R.V. Chamberlin(AMNH); 1,, California, Santa Cruz County, Felton, 6.ii.1949,coll. P.H. Arnaud (CAS); 1 ,, California, Santa Cruz County,12 mi. S Los Gatos, 26.iii.1941, coll. S. Mulaik, D. Mulaik(AMNH); 1 ,, California, Santa Cruz County, Soquel, GroverGl., 2.viii.1980, coll. D. Ubick (D. Ubick collection).

61 62

63

Figs 61–63. Cybaeus thermydrinos, paratypemale, near Johnsondale,CA, left palpus: 61, bulb, ventral;62, tegular apophysis, retrolateral; 63, patella and tibia, dorsal. Scale bars = 0.10mm. E= embolus,K = tegular apophysis ‘keel’, PA= posterior portion of tegular apophysis, RPA= retrolateral patellarapophysis. Figures redrawn from Bennett (1991: figs 359–361).


Other material examined. USA: 1 ,, California, Santa Cruz County,Cave Gulch at Empire Grade Rd., 18.ii.1991, coll. D. Ubick (D. Ubickcollection); 2 ,, California, Santa Cruz County, Stump Cave, 8910,36.5964N 122.0424W, coll. P. Paquin, N. Dupérré, D. Ubick (RBCMcatalogue number ENT008–008621) (GenBank accession numbers:FJ263784, FJ263797).

Diagnosis

The form of the posterior arm of the tegular apophysis (Figs 67,68) diagnoses the male of C. penedentatus. The female isdistinguished by the form and location of the atrium (Figs 70,71, 73, 74, 76) and the form of the copulatory ducts (Figs 71, 73,74, 78).

Male

Patellar apophysis (Fig. 69) as long as width of patella, two(right patella) or six (left) peg setae. Retrolateral tibial apophysisnearly as long as tibia. Embolus relatively short, somewhatthickened. Posterior arm of tegular apophysis (Figs 67, 68)with rounded, posteriorly directed, basal projection; ventralmedial keel terminating in short, slightly dextrally twisted,acuminate tip. Variation: Holotype CL 2.6, CW 1.83, SL 1.26,SW 1.17.

Female

Anterior epigynal ‘wrinkles’ usually absent (Fig. 76), presentin some specimens (StumpCave, Ben Lomond (Fig. 70)). Atrium(Figs 70, 71, 73, 74, 76) small, transverse, medially located on

epigynum, of variable morphology. Copulatory ducts(Figs 71–75, 77, 78) well developed but somewhatmembranous and often difficult to discern, extendinganteriorly from atrium in close proximity or contiguously toanterior edge of vulva, then diverging and heading laterallythen posteriorly before merging with spermathecal stalks;medial and/or proximal sections often broad with ductsindistinct. Spermathecal heads on small, dorsal lobes(Figs 72, 75, 77). Variation (n= 12): CL 1.93–2.7 (2.3� 0.2);CW 1.30–1.85 (1.53� 0.16); SL 1.00–1.30 (1.16� 0.10);SW 0.96–1.25 (1.08� 0.09).


Coastal central California west of San Francisco Bay south toMonterey Bay (Fig. 113). Specimens have been collected underlogs and within a redwood forest.

Etymology

The specific epithet is from the Latin for ‘almost toothless’ andrefers to the small number of peg setae on the male patellarapophysis.

Cybaeus vulpinus Bennett, sp. nov.

(Figs 79–84, 112)

Material examined

Holotype. 1 <, USA: California, Tuolumne County, Fox Gulch, 1.5miles north of Columbia, 22000, 6.xii.1986, coll. D. Ubick (CAS).

64

65 66

Figs 64–66. Cybaeus thermydrinos, paratype female: 64, near Johnsondale, CA, epigynum,ventral; 65, same, vulva, ventral; 66, California Hot Springs, CA, vulva, ventral. Scalebars = 0.10mm. CD= copulatory duct, EW= epigynal ‘wrinkle’, HS= head of spermatheca,SS = stalk of spermatheca.


Paratypes. USA: 1 ,, California, Calaveras County, Calaveras BigTrees St. Pk., 13.vii.1981, coll. D. Ubick (D. Ubick collection); 1 ,,California, El Dorado County, 6 mi. E Camp Connell [?], 10.ix.1959,coll. W.J. Gertsch, V.D. Roth (AMNH).

Other material examined. USA: 1 ,, California, Calaveras County,Hwy. 49 0.2 mi S of Calaveras/Amador County line, N of MokelumneHill, ~210m, 38.3101N120.7224W,22.i.2005, coll.M.Hedin (RBCMcatalogue number ENT008–008622) (GenBank accession number:FJ263782).

Diagnosis

The form of the patellar apophysis (Fig. 81) and the posterior armof the tegular apophysis (Figs 79, 80, 82) diagnose the male ofC. vulpinus. The female is distinguished by the form and location

of the atrium (Fig. 83) and the form of the copulatory ducts(Figs 83, 84).

Description

The male and females were collected separately. They areconsidered to be conspecific because of the close proximity oftwoof the collection locales.Abdomenpatterned. Femora banded(very faintly in holotype). Ventral tibia I macrosetae 2-1p-2-1p-1p(or 2).

Male

Patellar apophysis (Fig. 81) short, length ~1/3width of patella,strongly directed anteriorly; 13 peg setae clustered mostly near

67 68

69

Figs67–69. Cybaeuspenedentatus, holotypemale,HolyCity,CA, left palpus: 67, bulb, ventral;68, tegular apophysis, retrolateral; 69, patella and tibia, dorsal. Scale bars = 0.10mm. K= tegularapophysis ‘keel’, PA= posterior portion of tegular apophysis, RPA= retrolateral patellarapophysis. Figures redrawn from Bennett (1991: figs 297–299).


tip. Retrolateral tibial apophysis nearly as long as tibia. Embolusrelatively short, somewhat thickened. Posterior arm of tegularapophysis (Figs 79, 80, 82) simple; tip short, curved, with slightdextral twist. Variation: Holotype CL 2.8, CW 2.00, SL 1.35,SW 1.30.

Female

The epigynum of this species has not been illustrated. Atrium(Fig. 83) broad, weakly arched, located medially on epigynum.Copulatory ducts (Figs 83, 84) well developed but lightlysclerotised and somewhat membranous, broadly contiguous atatrium, separate elsewhere; proximal sections anteriorly directedand convergent. Spermathecal heads on small, dorsal lobes.Stalks of spermathecae smoothly sinuous. Variation: CL 2.8,3.2; CW 1.88, 2.18; SL 1.33, 1.50; SW 1.22, 1.45. El Doradospecimen largest.


Calaveras,ElDoradoandTuolumneCounties in the central SierraNevadaof easternCalifornia (Fig. 112).One femalewas collectedin a north-facing mixed pine/oak woodland.

Etymology

The specific epithet is Latin for ‘of a fox’ and refers to the typelocality.

devius species group

Members of the devius group are rarely collected and mostare known from only one or two localities, all in the SierraNevada of California from the Lake Tahoe area south to

70 71

72 73

74 75

Figs70–75. Cybaeuspenedentatus, paratype females: 70,BenLomond,CA,epigynum,ventral;71, same, vulva, ventral; 72, same, dorsal; 73, Pescadero Creek, CA, vulva, ventral;74, Mt. Madonna, CA, vulva, ventral; 75, same, dorsal. Scale bars = 0.10mm. AT= epigynalatrium, BS= base of spermatheca, CD= copulatory duct, HS = head of spermatheca, SS = stalk ofspermatheca.


Tulare County. The group contains one described species,C. devius Chamberlin & Ivie; four are undescribed.

septatus species group

The septatus group contains one relatively common species,C. septatus Chamberlin & Ivie and four less common species,two of which are described below (C. chauliodous, sp. nov. andC. somesbar, sp. nov.); two remain undescribed. All haverelatively restricted ranges and occur from south-westernOregon south to Alameda and Sierra Counties in California.

Cybaeus chauliodous Bennett, sp. nov.

(Figs 85–98, 113)

Material examined

Holotype. 1 <, USA: California, Plumas County, south side of LakeAlmanor, 5.ix.1959 coll. V.D. Roth, W.J. Gertsch (AMNH).

Paratypes. USA: 1 ,, California, Plumas County, 9.2 mi. NE BucksLake, 4.ix.1988, coll. D. Ubick (D. Ubick collection); 2 <, 3 ,,California, Plumas County, from south side of Lake Almanor,5.ix.1959, coll. V.D. Roth, W.J. Gertsch (AMNH); 5 ,, California,SierraCounty, 2mi.NCalpine, 6.ix.1959, coll.W.JGertsch,V.D.Roth(AMNH); 3 ,, California, Sierra County, Sierra City, The Cups,6.ix.1959, coll. W.J Gertsch, V.D. Roth (AMNH); 1 <, Oregon,Jackson County, Ashland Summit, 14.x.1954, coll. V.D. Roth(CAS); 1 ,, Oregon, Jackson County, Beaver-Sulphur For. Camp,nr. Applegate R., 9.x.1964, coll. F. Beer (CAS).

Other material examined. USA: 1 ,, California, Humboldt County,Brannan Mtn. Rd., 6.7 mi W of Hwy 96, vicinity of Willow Ck.,40.9475N 123.6459W, ~230m, 15.iii.2006, coll. M. Hedin (RBCM);1 ,, California, Trinity County, Hwy 299, 40.4502N 123.1664W,20.ii.2003, coll. P. Paquin, D. Wytrykush (RBCM); 4 <, 2 ,,

Oregon, Jackson County, 12 mi SE of Ashland, Rd. 66, 42.3119N122.4972W, 7.xi.2003, coll. P. Paquin, D. Wytrykush (RBCMcatalogue number ENT008–008623) (GenBank accession numbers:FJ263786, FJ263799).

Diagnosis

The enlarged peg seta (Figs 87–90) and the posterior arm of thetegular apophysis (Figs 85, 86, 91, 92) distinguish the male ofC. chauliodous. The female is diagnosed by the form of theatrium (Figs 93, 96) and the configuration (Figs 94, 95, 97, 98) ofthe copulatory ducts and spermathecal stalks.

Description

Abdomen usually patterned, holotype abdomen nearlyconcolorous. Legs unbanded. Ventral tibia I macrosetaevariable, usually 2-1p-2-1p-1p.

Male

Patellar apophysis (Figs 88, 89) as long as or longer thanpatellar width, four or five peg setae with one much larger thanothers (Figs87–90).Retrolateral tibial apophysis nearly as longastibia (Fig. 87). Embolus relatively short, thin. Posterior arm oftegular apophysis with tip dextrally twisted, swollen, rounded,rather lip-like (Figs 85, 86, 91, 92). Variation (n= 4): CL1.85–2.38 (2.09); CW 1.30–1.78 (1.55); SL 0.98–1.22 (1.09);SW 0.90–1.14 (1.02). Holotype smallest specimen.

Female

Epigynum with one or more ‘wrinkles’ anteriorly (Figs 93,96). Pair of large, well separated atria (Figs 93, 96); visible in

76

77 78

Figs 76–78. Cybaeus penedentatus, paratype female, Brookdale, CA: 76, epigynum, ventral;77, vulva, ventral; 78, same, dorsal. Scale bars = 0.10mm.AT= epigynal atrium,CD= copulatoryduct, HS = head of spermatheca, SS = stalk of spermatheca.


79 80

81 82

Figs 79–82. Cybaeus vulpinus, holotype male, Columbia, CA, left palpus: 79, bulb, ventral;80, tegular apophysis, retrolateral; 81, patella and tibia, dorsal; 82, posterior portion of tegularapophysis, prolateral. Scale bars = 0.10mm. PA= posterior portion of tegular apophysis,RPA= retrolateral patellar apophysis. Figures redrawn from Bennett (1991: figs 282–284, 287).

83 84

Figs 83, 84. Cybaeus vulpinus, paratype female, Big Trees State Park, CA: 83, vulva,ventral; 84, same, dorsal. Scale bars = 0.10mm. AT= epigynal atrium, BS= base ofspermatheca, CD= copulatory duct, HS = head of spermatheca, SS = stalk of spermatheca.


dorsal view, not obscured by vulval ducts (Figs 95, 98).Copulatory ducts short, exiting atria towards vulval midline,contiguous anteriorly then proceeding laterally and diverging(Figs 95, 98). Spermathecal heads small, dorsolateral lobes.Spermathecal stalks (Figs 94, 95, 97, 98) very short, slightlysinuous, not contiguous. Variation (n= 13): CL 1.95–2.7(2.3� 0.3); CW 1.33–1.93 (1.60� 0.18); SL 1.01–1.30(1.14� 0.11); SW 0.96–1.28 (1.10� 0.11).

Note

The male from Ashland summit, Oregon (Figs 85–87, 89, 92)differs slightly from the Lake Almanor males (Fig. 88, 90, 91) inplacement of the enlarged peg seta (closer to the patellarapophysis tip in the Ashland summit male) and form of theposterior arm of the tegular apophysis (trough broader andshallower anterior to tip and anterior lip of tip narrower in the

Ashland summit male). Perhaps this specimen and the ApplegateRiver female discussed below are representatives of another newspecies. They are tentatively placed here pending collection of alarger series.

The Sierra County females (not illustrated) differ from theLake Almanor (Plumas County) specimens (Figs 93–95) inhaving somewhat elongated spermathecal bases (versusrounded in Lake Almanor females) and spermathecal headsextending laterally beyond the atrial openings (versus notextended laterally in Lake Almanor females). The singleApplegate River (Jackson County) female (Figs 96–98) hasrounded bases and laterally extended heads. The LakeAlmanor genitalic figures were drawn from a probably teneralfemale (lightly sclerotised, thin-walled vulva). This may explainthe differences noted here (see Bennett 2006 for a discussion ofontogeny and temporal variation in Cybaeus female genitaliccharacters).

85 86

87

Figs 85–87. Cybaeus chauliodous, paratype male, Ashland Summit, OR, left palpus: 85, bulb,ventral; 86, tegular apophysis, retrolateral; 87, tibia and tip of patellar apophysis, retrolateral.Scale bars = 0.10mm. PA= posterior portion of tegular apophysis, PS = enlarged peg seta,RTA= retrolateral tibial apophysis. Figure 85 redrawn from Bennett (1991: fig. 389).



Northern California and inland south-western Oregon (Fig. 113).In California, specimens have been collected from under rocks inPinus and Pinus/Quercus forests. Males have been collected inearly September and mid November.

Etymology

The specific name is a Greek compounded adjective meaning‘having a prominent tooth’ and refers to the enlarged peg setawhich is diagnostic for males of this species

Cybaeus somesbar Bennett, sp. nov.

(Figs 99–101, 113)

Material examined

Holotype. 1 ,, USA: California, Siskiyou County, 1 mile south ofSomesbar, 22.viii.1959, coll. V.D. Roth, W.J. Gertsch (AMNH).

Paratypes. USA: 1,, California, SiskiyouCounty, 0.5mi. NCecilville,18.vi.1989, coll. D. Ubick, (D. Ubick collection); 1 ,, Oregon,Josephine County, 0.3 mi. S Elk Creek, off Hwy 199, 16000, nodate, coll. E. Benedict (MCZ).

88 89

Figs 88, 89. Cybaeus chauliodous, paratype males, left palpus, tibia and patella, dorsal:88, Lake Almanor, CA; 89, Ashland Summit, OR. Scale bars = 0.10mm. PS= enlarged pegseta. Figures redrawn from Bennett (1991: figs 390, 391).

0.48 mm 75 µm 86 µm

Figs90–92. Cybaeus chauliodous, paratypemales, left palpus: 90,LakeAlmanor,CA,patellar apophysis, dorsal; 91, same,posterior portion of tegular apophysis, ventral; 92, Ashland summit, OR, same. PS = enlarged peg seta.


Other material examined. USA: 1 ,, California, Siskiyou County,Salmon River, vicinity of Matthews Ck. campground, ~17500,41.1123N 123.1286W, 10–12.vii.2005, coll. M. Hedin (RBCMcatalogue number ENT008–008624) (GenBank accession numbers:FJ263785, FJ263798).

Diagnosis

The male of C. somesbar is unknown. The female is diagnosedby the configuration of the atria (Fig. 99), copulatory ducts andspermathecal stalks (Figs 100, 101).

Description

Abdomen usually patterned. Legs unbanded. Ventral tiba Imacrosetae 2-1p-2-1p-1p(or 0, or 2).

Female

Epigynum with one or more ‘wrinkles’ anteriorly (Fig. 99).Pair of very small, crook-shaped, inconspicuous atria anteriorly

located on epigynum (Fig. 99, 100), in dorsal view atria obscuredor partially obscured by vulval ducting and not readily visible.Copulatory ducts short, much shorter than spermathecal stalks.Spermathecal heads small, dorsolateral lobes. Spermathecalstalks (Figs 100, 101) convoluted with three 180� bends,proceeding anteriorly from spermathecal heads then reversingdirection three times before entering bases. Variation: CL 2.13,2.23, 2.48; CW 1.40, 1.45, 1.65; SL 1.04, 1.14, 1.21; SW 1.01,1.03, 1.12. Holotype largest specimen.

Distribution

Klamath Mountains of north-western California and adjacentextreme south-western Oregon (Fig. 113). Apparently absentfrom the coast.

Etymology

The specific name is a noun in apposition taken from the typelocality.

93 94

95 96

97 98

Figs 93–98. Cybaeus chauliodous, paratype females: 93, Lake Almanor, CA, epigynum,ventral; 94, same, vulva, ventral; 95, same, dorsal; 96, Applegate River, OR, epigynum,ventral; 97, same, vulva, ventral; 98, same, dorsal. Scale bars = 0.10mm. AT= epigynalatrium, BS= base of spermatheca, CD= copulatory ducts, EW= epigynal ‘wrinkle’, HS = headof spermatheca, SS = stalk of spermatheca.


tardatus species group

Members of the tardatus group are rarely collected andapparently have very restricted ranges. One species,C. tardatus (Chamberlin), is described; four others areundescribed. All occur only in south-western California fromsouth-western Monterey County to San Diego County.

Unplaced species

Bennett (1991) was unable to place two described (includingC. gidneyi, sp. nov., described below) and one undescribedspecies of Nearctic Cybaeus into species groups. Based onspermathecal head morphology, he felt that these species fitsomewhere within the Californian clade. The molecularanalysis unequivocally places C. gidneyi in the Californianclade, but it is still unclear with which species group it is allied.

Cybaeus gidneyi Bennett, sp. nov.

(Figs 16, 102–111, 113)

Material examined

Holotype. 1 <, USA: Oregon, Curry County, 2 miles N Brookings,31.ix.1959 coll. V.D. Roth (CAS).

Paratypes. USA: 4,, California, Humboldt County,Ave. of theGiants,Ten-Taypo Tr., 14.ix.1971, coll. V.D. Roth (CAS); 1 ,, California,Humboldt County, Orick, 16.ix.1961, coll. W. Ivie, W.J. Gertsch(AMNH); 8 ,, Oregon, Curry County, 7 mi. E Brookings, Chetco R.,29.v.1953, coll. V.D. Roth (CAS); 9 <, 9 ,, Oregon, Curry County,2 mi. N Brookings, 31.ix.1959, coll. V.D. Roth (CAS); 2 ,, Oregon,Curry County, 5 mi. N Brookings, matured 2.x.1959, coll. V.D. Roth(CAS).

Other material examined. USA: 2 ,, California, Humboldt County,Redwood Nat. For., Bald Hill Rd. off Hwy 101, [nr. Orick], 41.4786N

124.0000W, 24.ix.2003, coll. P. Paquin, D. Wytrykush (RBCMcatalogue number ENT008–008625) (GenBank accession numbers:FJ263779, FJ263800).

Diagnosis

Cybaeus gidneyi is distinguished by the form of the patellar(Figs 16, 104) and tegular apophyses (Figs 16, 102, 103) inthe male and, in the female, by the form of the atria (Figs 106,109), the copulatory ducts (Figs 107, 108) and the spermathecalstalks (Figs 108, 111).

Description

Abdomen pale grey, unpatterned (one female specimenwith veryslight pattern), legs unbanded. Two or three complete pairs ofventral tibia I macrosetae, terminal pair complete or not; pattern2-0-2-1p-1p (or 2).

Male

Patellar apophysis (Figs 16, 104) long, slender; nearly as longas width of patella; 14–20 peg setae concentrated mostly aroundtip, some along posterior dorsal edge. Retrolateral tibialapophysis (Fig. 105) nearly as long as tibia. Embolus(Fig. 102) long, slender. Tegular apophysis (Figs 16, 102, 103)with slender base; anterior arm elongate, slender (length abouttwice width) with rounded tip; posterior arm with bulbous basenarrowing abruptly to anteroventrally directed, acuminate tipwith slight dextral twist. Variation (n= 10): CL 2.08–2.7(2.4� 0.2); CW 1.63–1.88 (1.75� 0.10); SL 1.13–1.30(1.21� 0.06); SW 1.09–1.21 (1.15� 0.05). Holotype CL 2.35,CW 1.65, SL 1.18, SW 1.12.

99

100 101

Figs 99–101. Cybaeus somesbar, holotype female, Somesbar, CA: 99, epigynum, ventral;100, vulva, ventral; 101, same, dorsal. Scale bars = 0.10mm. BS = base of spermatheca,CD= copulatory ducts, EW= ‘epigynal wrinkle’, HS = head of spermatheca, SS = stalk ofspermatheca.


Female

Epigynum (Figs 106, 109) with small pair of posteriorlylocated, curved atria. Copulatory ducts (Figs 107, 108, 111)opening from atria towards posterior midline of vulvathen turning anteriorly and expanding into large bulb-likestructures before turning posteriorly at anterior margin ofvulva and narrowing. Spermathecal heads (Figs 108, 111)small, dorsal lobes often at ends of long ducts. Spermathecalstalks relatively short, simple, proceeding anteriorly thenposteriorly before entering small, rounded spermathecal bases.Secondary pores dorsally at junction of stalks with bases in closeproximity to heads. Short fertilisation ducts exiting bases near tojunction with stalks. Variation (n= 22): CL 2.15–3.0 (2.6� 0.2);

CW 1.53–2.03 (1.73� 0.13); SL 1.08–1.43 (1.24� 0.08);SW 0.98–1.30 (1.14� 0.08).


Extreme south-western Oregon south to Humboldt County,California (Fig. 113). Specimens have been collected fromwithin rotting redwood (Sequoia sempervirens) logs.

Etymology

The specific epithet is a patronym honoring J. Dirk Gidney,a long-time friend who introduced RB to Oregon and otherinteresting places.

102 103

105104

Figs 102–105. Cybaeus gidneyi, paratype male, Brookings, OR, left palpus: 102, bulb, ventral;103, tegular apophysis, retrolateral; 104, patella and tibia, dorsal; 105, tibia and tip of patellarapophysis, retrolateral. Scale bars = 0.10mm. AA= anterior portion of tegular apophysis,E = embolus, PA= posterior portion of tegular apophysis, RPA= retrolateral patellar apophysis,RTA= retrolateral tibial apophysis. Figures redrawn from Bennett (1991: figs 306–308, 310).


Discussion

Both morphology and our preliminary molecularphylogenetic analyses support the existence of two majorlineages of North American Cybaeus: a Holarctic clade and aNearctic, primarily Californian, clade. Cybaeus data appear toreflect divergence patterns common to many taxa. Thesepatterns are attributed to the movement of continental landmasses and ancient oceans creating dispersal corridorsbetween regions that are no longer contiguous, as well asthe formation of the Rocky Mountains during the lateOligocene and continuing through the Miocene (25–5Mya), which caused major changes to moisture regimes(Sanmartin et al. 2001). Further discussion of these

divergence patterns in relation to each clade is includedbelow.

Holarctic clade

TheHolarctic assemblage of theangustiarum and tetricus speciesgroups is the most broadly distributed Cybaeus clade – withspecies that occur in Europe and eastern and western NorthAmerica. We did not examine east Asian specimens ofCybaeus species but recently published illustrations of femalegenitalic characters (Ihara 2005–2007; Kobayashi 2006) indicatethat at least some Japanese species can be placed within theangustiarum or tetricus groups (e.g. C. aokii Yaginuma,C. aquilonalis Yaginuma, C. enshu Kobayashi, C. gassan

106 107

109108

110 111

Figs 106–111. Cybaeus gidneyi, paratype female: 106, Brookings, OR, epigynum, ventral;107, same, vulva, ventral; 108, same, dorsal; 109,Orick,CA, epigynum,ventral; 110, same, vulva,ventral; 111, same, dorsal. Scale bars = 0.10mm. AT= epigynal atrium, CD= copulatory ducts,HS= head of spermatheca, SS = stalk of spermatheca.


Kobayashi,C. hatsushibai Ihara,C. jinsekiensis Ihara,C. kiiensisKobayashi, C. kokuraensis Ihara, C. kuramotoi Yaginuma,C. miyagiensis Ihara, C. nipponicus (Uyemura) andC. urabandai Ihara). Neither molecules nor morphology showstrong support for the individual monophyly of either theangustiarum or tetricus species groups.

In their study of 57Holarctic-distributed taxa (mostly insects),Sanmartin et al. (2001) found that trans-Atlantic dispersalsacross the Thulean land bridge that connected eastern NorthAmerica with western Europe during the early to mid-Tertiary(70–20Mya) accounted for a considerable number of theobserved distribution patterns. The vicariance of eastern NorthAmerica taxa from those of western North America wasintensified by the formation of the Rocky Mountains duringthe mid-to-late Tertiary (20–2 Mya) and concomitant rain-shadow effect (Hendrixson and Bond 2007). This pattern ismirrored in Cybaeus. Cybaeus patritus and C. giganteus aretwo of only three Cybaeus species that occur in eastern NorthAmerica. These species diverged from C. angustiarum~20million years ago (based on the commonly used estimateof 2.3%permillion years for arthropodmitochondrial divergence(Brower 1994; Masta 2000; Ayoub and Riechert 2004; Croucher

et al. 2004)); theyconsistently formawell supported clade andarephysically separated from other North American cybaeids by avast region of habitat unsuitable for these mesic-forest inhabitingspiders. These Appalachian species, as well as Cybaeusshoshoneus, found in north-eastern Washington, Idaho andMontana, form a cluster within the Holarctic lineage distinctfrom the remaining coastally distributed cybaeids.

It is thought that there were twomajor occurrences of forestedcorridors that enabled dispersal across the Beringian land bridge,first via a mesophytic forest during the early–mid Tertiary(65–20Mya) and then via a coniferous forest, during the lateTertiary (20–3 Mya). These periods of contiguous landmassesfacilitated faunal connections between western North Americaand eastern Europe in many taxa (Sanmartin et al. 2001) and thisis reflected in theH3 tree, with evidence of a Pacific coastal clade.The COI data show divergences between coastally distributedspecies that range from 20 to 10 Mya, so it is unclear whichof the particular Beringian connections was important toCybaeus dispersal. Some of the species in this coastal cladeare broadly co-distributed, such as C. morosus and C. eutypusand C. reticulatus. Of these species, C. eutypus has the moresoutherly distribution: central California north to the QueenCharlotte Islands. Cybaeus morosus and C. reticulatus rangefurther north (including the Queen Charlotte Islands), throughoutthe Aleutians, but do not extend south past San FranciscoBay. Cybaeus waynei and C. multnoma are found in Oregon

Fig. 112. Distributions of Cybaeus sanbruno (~), C. thermydrinos (&),C. vulpinus (¤) and C. waynei (*) in Oregon and California.

Fig. 113. Distributions of Cybaeus chauliodous ( ), C. gidneyi (!),C. paralypropriapus (~), C. penedentatus ($) and C. somesbar ( ) inOregon and California.


and northern California and C. paralypropriapus has a verylimited range along the coast of Oregon that is encompassedby all five other species.

AlthoughCybaeus signifer is foundwithin theHolarctic clade,as proposed by Bennett (1991), this coastally distributed species(ranging from the Queen Charlotte Islands in British Columbia,Canada south to Monterey County, California and east to theSierra Nevada) fails to cluster with the other species with similardistributions. Interestingly, this species is the only cybaeid withwell developed anteriormedian eyes (Chamberlin and Ivie 1932).As well, this species has markedly different behaviour frommostotherCybaeus, actively hunting at night on the trunks of trees andon the forestfloor (Bennett 2005).Most otherCybaeus spiders aresecretive, inhabit litter and coarse woody debris and are usuallyfound in association with insubstantial webs or, as is the case forsome Japaneses species, open-ended tube-like constructs.

Californian clade

The California Floristic Province is a biologically rich andendangered terrestrial ecoregion (Myers et al. 2000) andharbours an abundance of spider species and genetic diversity(Bond et al. 2001; Hedin 2001; Calsbeek et al. 2003; Bondet al. 2006; Crews and Hedin 2006; Starrett and Hedin 2007;Platnick and Ubick 2008). California is also a significantregion for Cybaeus species diversity, with 40 of the 63 NorthAmerican species represented by the species groups in theCalifornian clade, many of them with very limiteddistributions. Lapointe and Rissler (2005) analysed multipletaxa and found that divergence corresponded to the formationof the Sierra Nevada, Coast and Transverse mountain ranges~4–7million years before present (Calsbeek et al. 2003; Starrettand Hedin 2007) and resulting in high rates of endemism andspeciation. Despite this, estimates of species divergence dates forCybaeus are earlier than that, ranging from21 to 13Mya.Becausethese are rough estimates based on 2.3% per million years forarthropod mitochondrial divergence (Brower 1994), and spidermitochondrial evolution appears to be very different from that ofother arthropods (Masta 2000), there aremany reasons why theseestimates may be imprecise. The restricted distributions ofCybaeus in California may be ascribed to more localised andmore recent geological events including mountain building, sea-water incursions, plate tectonic activity and climatic shifts(Calsbeek et al. 2003; Jacobs et al. 2004; Lapointe and Rissler2005), as well as anthropogenic activity (Connor et al. 2002;Vandergast et al. 2008).

Although our limited molecular dataset does not allow us tosay much about relationships among species groups in theCalifornian clade, there does appear to be support in our COIdata for a close relationship between the adenes and consociusspecies groups. This support is also reflected in the position,number and shape of the female atrium (in addition to the shortembolus in the male). Cybaeus sanbruno, C. hesper andC. penendatus exhibit complete sympatry and extreme rangelimitation – inhabiting the Santa Cruz Mountains between SanFrancisco Bay and Monterey Bay. Cybaeus vulpinus, theremaining consocius species member, has a completelydisjunct range; it is found only in a small region of the SierraNevada Mountains.

Conclusion

In this paper, our goal was to provide a framework forunderstanding the evolution and diversity of the species-richbut understudied genus Cybaeus. To that end, we introducedthe eight proposed North American species groups, along withthe morphological characters that define them. We provided asystematic description of Cybaeus and the species groups andformally described nine new species. Finally, we documentedthe first molecular phylogenetic analysis of Cybaeus, using onemitochondrial and one nuclear gene. Both molecules andmorphology support the existence of two major clades, oneHolarctic and the other Nearctic, primarily Californian.

The Holarctic clade of Cybaeus might not consist of twomonophyletic species groups, but rather may be a single one thatrepresentsCybaeus sensu stricto, (the type speciesC. tetricus is amember of the Holarctic clade). The remaining Nearctic speciesgroups (the Californian clade) appear to represent speciesthat arose in North America and may warrant generic status inlight of their evolutionary history. The relationships among theseCalifornian taxa require further study, with molecular analysisof additional representatives of the various species groups.Complete resolution of the systematics of the genus requiresexamination of the numerous eastern Palearctic and Japanesespecies.

Acknowledgements

We thank the following museum curators and individuals for the loan ofspecimens used for the morphological component of this study: JacquelineHeurtault (Muséum national d’histoire naturelle), Paul Hillyard(British Museum (Natural History)), Herb Levi (Museum of ComparativeZoology), Norm Platnick (American Museum of Natural History), WojciechPulawski (California Academy of Sciences), Vince Roth (Portal AZ) andDarrell Ubick (San Francisco CA). Marshal Hedin (San Diego StateUniversity), Jiri Kral (Charles University), Pierre Paquin (Montreal QC)and Frantisek Stahlavski (Charles University) provided the critical freshCybaeus specimens without which the molecular work (and this paper)could not have been completed. Darren Copley kindly prepared finalversions of taxonomic plates and range maps. We are grateful for thesupport of the Royal BC Museum and the BC Ministry of Forests andRange. Funding was provided by an NSERC Discovery Grant to StevePerlman. Steve Perlman also acknowledges support from the CanadianInstitute for Advanced Research. Rob Cannings, Jason Bond andanonymous referees provided helpful comments on earlier versions of thismanuscript.

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Manuscript received 14 January 2009, accepted 13 August 2009


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