Anelosimus (Agnarsson, 2004)

Agnarsson, Ingi, 2006, A revision of the New World eximius lineage of Anelosimus (Araneae, Theridiidae) and a phylogenetic analysis using worldwide exemplars, Zoological Journal of the Linnean Society 146 (4), pp. 453-593 : 462-470

ANELOSIMUS – CLADE 30

The results corroborate the monophyly of Anelosimus s.s. (excluding Kochiura and Selkirkiella , see Agnarsson, 2004), Figures 58 and 59 . Agnarsson (2004) found marginal support for Anelosimus , based on a single synapomorphy: ridges on the surface of the epigynal plate (4 -1, Figs 48A, B , 53B ). With more species in the analysis this synapomorphy supports the subsidiary Eb clade within Anelosimus (see Fig. 58 and below). Here Anelosimus synapomorphies are: an entire conductor (37 -1, Fig. 8C ), lack of pars stridens on prosoma

Characters Ste Eno Sel Ara Are Koa Kor Ach The koh sp1 lor rup big dud vit pul nel rab nig eth may sal ana chi pac dom jab dub exi oct juc ari bae pur stu gua ele tos pan ori fra tun

FEMALE GENITALIA

001. Epigynal ventral margin: 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0? 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 entire; with scape;

002. Epigynal ext.: cop. 0 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 openings not visible;

visible;

003. Epigynal depression: 0 – – – – – 1 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 narrow; wide;

004. Epigynal plate surface: 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 1 1 1 1 1 1 1 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 smooth; ridged;

005. Epigynal septum: absent; 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 present;

006. Copul. bursa ant. margin: 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 entire; medially acute;

007. Cop. duct to spermath: 1 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 posterior; lat./anterior;

008. Copul. duct rel. to 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 spermath.: apart;

encircling;

009. CD term. sclerot.: as 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0 1 1 1 1 1 1 1 other; distinctly sclerotized;

010. Sclerot.CD: ectal to the – – – – – – – – – – – – – – – – – – – – – – – 0 0 0 – – – – 0 0 0 0 0 1 1 spermathecae; caudal;

MALE GENITALIA

011. Tibial rim: slightly 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 asymmetric; scoop-

shaped;

012. Tibial prol.trichob.: => 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 two; one; none;

013. Cymbial ectal setae: 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 unmod.; curved towards bulb;

014. Cymbial dist. prom.: 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 entire; w/apoph.;

015. Cymbial mesal margin: 0 0 0 0 0 1 1 0 0 1 0 1 1 1 1 0 0 0 1 1 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 entire; incised; notched;

016. Cymbial tip 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 sclerotization: normal;

light;

017. Cymbial tip setae: 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 normal; thick and curved;

018. Cymbial sheath: absent; 0 0 0 0 0 1 1 0 0 0 0 1 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 present;

019. Lock placement: basal; 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 distal; central;

0 0 0

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1 1 1

1 1 1

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2 2 2

1 1 1

0 0 0

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0 0 0 0? 0

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1 1 1

1? 1

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1? 1

1 1 1

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2 2 2

1 1 1

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0 0 0

0 0 0 Characters Ste Eno Sel Ara Are Koa Kor Ach The koh sp1 lor rup big dud vit pul nel rab nig eth may sal ana chi pac dom jab dub exi oct juc ari bae pur stu gua ele tos pan ori fra tun

020. Lock mechanism: hook; 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 hood; Theridula ;

021. Cymbial hook orient.: 0 1 1 0 0 1 1 – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – downwards; upwards;

022. Cymbial hook: inside 0 1 1 0 0 0 0 – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – cymbium; ectal margin;

023. Cymbial hook tip: blunt; 0 0 1 0 0 0 0 – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – strongly tapering;

024. Cymbial hood size: – – – – – – – 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 narrow; broad;

025. Cymbial hood region: – – – – – – – 1 1 1 1 1 1 0 0 1 1 1 1 1 1 1 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 translucent; opaque;

026. Alveolus shape: suboval; 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0????? 0 0 0 0 0 0 0 0 0 0 0 0 0 0 mesal extension;

027. Tegulum ectal margin: 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 entire; protruded;

028. Tegular groove: absent; 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 present;

029. SDT SB I: separate; 0 – 0 0 0 0 0 0 0? – 0 0 0 0 1 1 0? 1 1?? 0 0? – 0 0 0 0 0 0 0 0 0 0 touching;

030. SDT post-SB II turn: 0 – 0 0 0 0 0 0 0? – 0 0 0 0 0 0 1 0 0 0 0 0 1 1? 1 1 1 1 1 1 1 1 1 1 1 absent; present;

031. SDT SB I & II segm. 0 – 0 0 0 0 0 0 0? – 0 0 1 1 1 1 1????? 1 1? – 1 1 1 1 1 1 1 1 1 1 alignment: diverg.;

parallel;

032. SDT SB I & II orient.: in 0 – 0 1 1 1 1 1 1? – 1 1 1 1 1 1 1????? 1 1? – 1 1 1 1 1 1 1 1 1 1 plane; out of plane;

033. SDT RSB I & II: absent; 0 0 0 1 1 0 0 0 0? – 0 0 0 0 0 0 0 0 0 0 0 0 0 0? 0 0 0 0 0 0 0 0 0 0 0 present;

034. SDT SB III: absent; 0 1 1 1 1 0 0 0 1? 0 0 0 0 0 0 0 0????? 0 0? – 0 0 0 0 0 0 0 0 0 0 present;

035. SDT SB IV: absent; – 0 0 1 1 – – – 0? – – – – – – – – – – – – – – –? – – – – – – – – – – – present;

036. C: simple; fan; Eno.; Arg.; 0 2 1 3 3 1 1 4 5 0 – 6 6 7 7 0 0 0 0 0 0 0 0 8 8 8 0 0 0 0 8 8 8 8 8 8 8 Ach.; The; rup.; tan.; cup;

037. C: with a groove for – 0 0 0 0 0 0 0 0 1 – 1 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 embolus; entire;

038. C surface: smooth; 0 1 1 0 0 0 0 1 0 0 – 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 heavily ridged;

039. C tip sclerotization: like 0 1 1 0 0 0 0 1 1 0 – 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 base; more than base;

040. Subconductor (SC): 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 absent; present;

041. SC pit upper wall: entire; – – – – – – – – – – – – – – – 1 1 0 1 1 1 0 0 0 0 0 0 0 0 1 1 1 1 1 1 0 0 forms a regular oval lip

1 1 1

– – –

– – –

– – –

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– – –

8 8 8

1 1 1

0 0 0

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1 1 1

0 0 0 1 1 1

– – –

– – –

– – –

0 0 0

0 0 0

0 0 0

0 0 0

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1? 1

1? 1

1? 1

0? 0

0? 0

–? –

8 8 8

1 1 1

0 0 0

0 0 0

1 1 1

0 0 0

042. SC narrowing at C base: – – – – – – – – – – – – – – – 2 2 2 2 2 2 0 0 0 0 0 1 1 1 2 1 1 1 1 1 0 0 abrupt; gradual; broad

043. E base-SC relation: E – – – – – – – – – – – – – – – 2 2 0 2 2 2 0 0 0 0 0 0 0 0 1 1 1 1 1 1 0 0 hooked in; E surpasses

SC

044. T ectally: <1/2 cymbial 0 0 0 0 0 0 0 0 0 0 1 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 cavity;> 1/2 cymbial cavity;

045. MA and sperm duct: not 1 1 1 1 1 0 0 – 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 in MA; inside MA;

046. MA-tegular connection: 1 1 1 1 1 1 1 – 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 broad; narrow;

047. MA form: unbranched; 0 0 0 0 0 0 0 – 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 two branches;

048. MA distal tip: entire; 1 1 1 1 1 1 1 – 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 hooded;

049. MA hood form: narrow, 0 0 1 0 0 1 1 – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – pit-like; scoop-shaped;

050. TTA form: entire; 0 1 1 0 0 1 1 – 0 0 – 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 grooved; excavate;

051. TTA: bulky; prong 0 0 0 0 0 0 0 – 0 0 – 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 shaped

052. TTA distal tip: entire; 0 0 0 1 1 0 0 – 2 0 – 0 0 2 2 0 0 0 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 Argyrodes ; hooked;

053. TTA hook distal branch: – – – – – – – – 0 – – – – 0 0 – – – 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 0 0 short; extending beyond;

054. TTA hook distal branch: – – – – – – – – 0 – – – – 0 0 – – – 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0 thick; thin, finger-like

055. TTA hook proximal – – – – – – – – 0 – – – – 0 0 – – – 1 1 1 2 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 branch: broad; bladelike

056. TTA surface 0 0 0 0 0 0 0 – 1 0 – 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 0 1 1 1 1 1 1 1 1 1 1 subterminally: smooth;

ridged;

057. TTA tip surface: smooth; 0 0 0 1 1 1 1 – 1 0 – 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 ridged;

058. E and TTA: loosely assoc.; 0 1 1 0 0 0 0 – 0 0 – 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 E enclosed in TTA ;

059. E tip surface: smooth; 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 denticulate;

060. E spiral curviture: gentle; 0 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0 whip like; corkscrew;

061. E tip: entire; bifid; 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1

062. E origin: ectoventral on 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 tegulum; retrolateral;

063. E surface: smooth; ridged; 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 1? 1 1 1 1 1 1 0 0 1 0 1 0 0 1 1 1 1 1 1 1

064. E shape: short to 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 moderate; extremely long;

065. E spiral: normal; thick, 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 much broader than tip;

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1 1 1 Characters Ste Eno Sel Ara Are Koa Kor Ach The koh sp1 lor rup big dud vit pul nel rab nig eth may sal ana chi pac dom jab dub exi oct juc ari bae pur stu gua ele tos pan ori fra tun

066. E distal rim: entire; 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 deeply grooved;

067. E terminus: abrupt; with 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 a distal apophysis;

068. E base: entire; lobed; 0 0 0 0 0 1 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

069. Embolus-dh connection 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 grooves: absent; present;

070. Embolus-dh grooves: – – – – – – – – – – – – – – – – – – – – – – – – – – 0 0 0 0 1 1 1 1 1 0 0 deep; short;

071. E spiral subterminally: 0 0 0 0 0 0 0 0 0 1 1 2 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 thin or filiform; thick;

rup.;

072. E spiral: entire; biparted; 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1? 1 1 1 1 1 0 1 1 1 1 2 1 1 1 1 1 1 1 ‘pars pendula’;

073. Eb orientation: towards E – – – – – – – – – – – – – – – 1? 0? 0 0 0 0 0 – 1 0 0 0 – 0 0 0 0 0 0 0 tip; towards tibia;

074. Eb: separates early from – – – – – – – – – – – – – – – 0 0 0 0 0 0 0 0 0 – 0 0 0 0 – 1 1 1 1 1 1 1 E; E and Eb tight;

075. Eb terminally: broad; – – – – – – – – – – – – – – – 0 0 0 0 0 0 0 0 0 – 0 0 0 0 – 0 0 0 0 0 1 1 narrow;

076. Eb distal portion, ectal – – – – – – – – – – – – – – – 0 0 0 0 0 0 0 0 0 – 0 0 0 0 – 1 1 0 0 0 0 0 margin: level; ridged;

077. Eb form, basally: flat, in – – – – – – – – – – – – – – – 0 0 0 0 0 0 0 0 0 – 0 0 0 0 – 1 1 1 0 0 0 0 ectal view; globose;

078. Eb form: separate – – – – – – – – – – – – – – – 0 0 0 1 1 1 0 0 0 – 0 0 0 0 – 0 0 0 0 0 0 0 apophysis; barely separate

079. E spiral tip:separated; Eb – – – – – – – – – – – – – – – 0 0 0? 0 0 0 0 0 – 0 2 2 2 0 1 1 1 1 1 1 1 and E tips juxtaposed

080. Eb snout: short, snug – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – 0 0 with E; long, separate;

081. Eb distal portion: entire; – – – – – – – – – – – – – – – 0 0 0? 0 0 0 0 0 – 0 1 1 1 – 2 2 2 0 0 0 0 with apophysis; ridged;

082. E tail: lobe simple; lobe – – – – – – – – – – – – – – – 0 0 0 0 0 0 0 0 0 – 0 0 0 0 – 1 1 1 0 0 0 0 bifurcate, snug with Eb

083. Extra tegular sclerite: 0 1 1 0 0 0 0 0 0? 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 absent; present;

SOMATIC MORPHOLOGY

084. AME (M): normal; on 0 0 0 1 1 0 0 0 0? 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 tubercle; eye region raised

085. AME size vs. ALE (M): 0 1 1 0 0 0 0 0 0? 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 subequal; smaller;

086. Chel. posterior margin: 1 0 0 0 0 0 0 1 1? 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 toothed; smooth;

087. Chel. post. tooth no.: – 2 0 2 2 0 0 – –? 0 0 0 0 0 0 0 0??? 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 four+ three; two; one;

0 0 0

0 0 0

1 1 1 1 1 1

0 0 0

1 1 1

1 1 1

0 0 0

1 1 1

1 1 1

0 0 0

0 0 0

0 0 0

1 1 1

1 0 0

0 0 0

0 0 0

0 0 0

0 0 0

0 0 0

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0 0 0 0 0 0

0 0 0

1 1 1 1 1 1

0 0 0

1 1 1

1 1 1

0 0 0

1 1 1

1 1 1

0 0 0

0 0 0

0 0 0

1 1 1

1 0 0

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0 0 0

0 0

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0 0 0

0 0 0

088. Chel. furrow: smooth; 0 0 0 1 1 0 0 0 0? 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 denticulate;

089. Carapace hairiness: 1 0 0 0 0 0 0 0 0? 0 0 0 0 0 0 0 0??? 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 sparse or patchy;

uniform;

090. Carap. pars stridens: 1 1 1 1 1 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 irregular; regular ridges;

091. Interocular area: flush 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 with clypeus; projecting;

092. Clypeus: concave or flat; 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 projected;

093. Ocular area setae (M): 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 sparse; in a dense field;

094. Labium-sternum 0 1 0 1 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 connection: seam; fused;

095. Sternocoxal tubercles: 1 1 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 present; absent;

096. Pedicel location: anterior; 0 0 1 0 0 0 0 1 0? 0 0 0 1 1 1 1 1 1 1 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 medial;

097. Abd. folium: spots or 0 1 1 0 0 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 blotches; central band;

098. Abdomen folium: 0 0 0 0 0 0 0 0 0 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 irregular; distinctly notched

099. Dorsal band: edge light; – 1 1 – – 0 0 – 1? 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 edge dark; Ameridion;

100. Abd. dot pigment: silver; 1 1 1 0 0 1 1 1 1? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 non-reflective, dull;

101. SPR form: weakly keeled; 1 0 0 1 1 1 1 0 0 1 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 strongly keeled;

102. SPR pick number: three 1 0 0 1 1 1 1 1 0 1 1 0 0 1 1 2 2 1 2 2 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 or less; four or more;

103. SPR insertion: flush with 1 1 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 abd. surface; on ridge;

104. SPR mesal pointing picks: 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 absent; present;

105. SPR mp-picks angle: 0 0 – 0 0 0 0 0 0 1 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 dorsal; perp. or ventral;

106. SPR: straight; distinctly 0 0 – 2 2 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 curved; argyrodine;

107. SPR dorsal pick 0 0 – 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 spacing:normal;

compressed;

108. SPR relative to pedicel: 1 0 1 0 0 1 1 1 0 1 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 lateral; dorsal;

109. SPR setae: separate; 0 0 0 0 0 0 0 0 0 1 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 tight;

110. Suprapedicillate 1 1 1 0 0 1 1 1 1? 1 1 1 1 1 1 1 1????? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 ventrolat. proprio.: abs.;

pres.;

0 0 0

1 1 1

0 0 0

0 0 0

0 0 0

0 0 0

0 0 0

1 1 1

0 0 0

1 1 1

1 1 1

0 0 0

1 1 1

1 1 1

1 1 1

0 0 0

1 1 1

1 1 1

1 1 1

1 1 1

1 1 1

1 1 1

1 1 1 0 0 0

1? 1

0 0 0

0 0 0

0 0 0

0 0 0

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1 1 1

1 1 1

1 1 1

1 1 1

1 1 1

1 1 1 Characters Ste Eno Sel Ara Are Koa Kor Ach The koh sp1 lor rup big dud vit pul nel rab nig eth may sal ana chi pac dom jab dub exi oct juc ari bae pur stu gua ele tos pan ori fra tun

111. Epiandrous fusule:in pair 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 of sockets; in a row;

112. Epiandrous fusule pair 2 0 2 3 3 1 0 1 1? 0 1 1 1 1 0 0 1? 0 0 1 1 1 1? 1 1 1 1 0 0 0 1 0 1 1 number:> ten; ≤ eight;

113. Colulus: present; absent; 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 invaginated;

114. Colulus size: large; small; 0 0 0 0 0 1 1 – – – – – – – – – – – – – – – – – – – – – – – – – – – – – –

115. Colular setae: present; 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 absent;

116. Colular setae number (F): 0 1 1 1 1 1 1 – – – 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 three or more; two;

117. Palpal claw dentition 0 0 1 1 1? 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 (female): dense; sparse;

118. Palpal tibial dorsal? 0 2 1 1 2 2 3 2 1 1 1 1 1 1 1 1? 1 1 1 3 3 1 1 1 1 1 1 1 1 1 1 1 0 0 1 trichobothria (female): 4;

3; 2; 5;

119. Femur I relative to II: 0 0 0 0 0 0 0 0 0? 1 0 0 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 subequal; robust, larger;

120. Leg IV rel. length (M): 1 1 1 0 0 0 0? 1? 0 0 0 0 0 0 0 1 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 third; second; longest;

121. Leg IV rel. length (F): 1 1 1 0 0 1 1 1 1? 0 1 1 1 1 1 1 1 1 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 third; second; longest;

122. Femur vs. metat.: metat. 1 0 1 1 1 1 1 0 1? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 longer; metat. shorter;

123. Metat. vs. tibia: metat.?????????? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 longer; metat. shorter;

124. Metatarsal ventral 0 0 1 1 1 1 1 0 0? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 macrosetae: normal;

thick;

125. Metatarsus vs. tibia?????????? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 length (male): longer;

shorter;

126. Metatarsal ventral 0 0 0 0 0 0 0 0 0? 0 0 0 0 0 1 1 1? 1 1 1 1 1 0 0 0 0 0 0 0 0 1 0 1 1 0 macrosetae: normal;

thick;

127. Tarsus IV comb: simple, 1 0 0 0 0 0 0 1 1? 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 straight; curved hooks;

128. Tarsal organ size: small 0 1 1 1 1 1 1 1 1? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 (normal); enlarged;

129. Tarsus IV central claw vs. 0 0 0 1 1 1 1 1 1? 1 1 1 1 1 1 1 1? 1 1?? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 lat. (M): short; longer;

130. Tarsus IV central claw 0 0 0 1 1 0 0 0 0? 0 0 0 0 0 0 0 0? 0 0?? 0 0 0 0 0 0 0 0 0 0 0 0 0 0 (F): normal; long; minute;

SPINNERETS

131. Spinneret insert.: normal; 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 abd. extend. beyond;

0 0 0

1 1 0

1 1 1

– – – 0 0 0

1 1 1

0 0 0

1 1 1

0 0 0

0 0 0

1 1 1

1 1 1

1 1 1

1 1 1

1 1 1

0 0 1

0 0 0

1 1 1

1 1 1

0 0 0

0 0 0 0 0 0

1? 1

1 1 1

– – – 0 0 0

1 1 1

0 0 0

0? 0

0? 0

0? 0

1? 1

1? 1

1? 1

1? 1

1? 1

1? 1

0 0 0

1 1 1

1 1 1

0 0

0 0 0

132. PLS FL spigot: = PLS CY; 0 0 1 – – 1 1 1 1 0 0 1 1 1 1 1 1?????? 0 0? 0 0 0 0 0 0 0 0 0 0 0 longer than PLS CY;

133. CY spigot bases: =<0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0? 0 0?? 0 0? 0 0 0 0 0 0 0 0 0 0 0 ampullates; huge;

134. CY shaft surface: smooth; 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0? 0 0?? 0 0? 0 0 0 0 0 0 0 0 0 0 0 grooved;

135. PLS AC spigot number: 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0? 0 0?? 0 0? 0 0 0 0 0 0 0 0 0 0 0 5+ four or less;

136. PLS FL spigot: present; 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0? 0 0 0? 0 0? 0 0 0 0 0 0 0 0 0 0 0 absent;

137. PLS posterior AG spigot 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1? 0 0?? 1 1? 1 1 1 1 1 1 1 1 1 1 1 shape: normal; flat;

138. PLS theridiid AG 0 0 – 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1? 1 1?? 1 1? 1 1 1 1 1 1 1 1 1 1 1 position: parallel; end to end;

139. PMS mAP spigot shaft: 1 1 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1? 1 1?? 1 1? 1 1 1 1 1 1 1 1 1 1 1 short;> CY shafts;

BEHAVIOUR

140. Web form: sheet; cobweb; 1 1 1 3 3 1 1 1 1?? 2 2?? 1????? 0 0 0 0? 0 0 0 0 0 0 0 0 0 0 0 mesh web; dry line-web;

141. Knock-down lines: 0 0 0 0 0 0 0 0 0?? 0 0?? 0????? 1 1 1 1? 1 1 1 1?? 1 1 1 1 1 absent; present;

142. Sticky silk in web: 0 0 1???? 0 0?? 1 1???????? 1 1???? 0 0 1? 1 1 1? 1 1 present; absent;

143. Egg sac: spherical to? 0 0 1 1?? 0 0?? 0 0?? 0??????? 0?? 0 0 0 0 0 0 0 0 0 0? lenticular; stalked;

144. Egg case: round; knob.;? 0 0 1 1?? 0 0?? 0 0?? 0??????? 0?? 0 0 0 0 0 0 0 0 0 0? rhomb.; elong.; spiky;

145. Web construction: 0 0 0 0 0 0 0 0 0? 1 1 1?????? 1 1 1 1 1?? 1 1 1 1?? 1 1 1 1 1 solitary; communal;

146. Mating thread: present; 0 0??? 0? 1 1???????????????????????????? absent;

147. Adult females per nest: 0 0 0 0 0 0 0 0 0? 0 1 1?? 0 0?? 0 0 0 0 0?? 1 1 1 1?? 0? 1? 1 one; multiple;

[148. Behavior: solitary; 0 0 0 0 0 0 0 0 0? 1 2 2?? 0 0?? 1 1 1 1 1?? 2 1 1 2?? 1 1 2? 2 subsocial; quasisocial;]

0 0 0

0 0 0

0 0 0

0 0 0

0 0 0

1 1 1

1 1 1

1 1 1

0 0 0

1??

1??

? 0?

? 0?

1??

???

0 0?

1 1? 0? 0

0? 0

0? 0

0? 0

0? 0

1? 1

1? 1

1? 1

0? 0

1??

1??

0??

0??

1? 1

???

1??

2? 1

Rows represent characters and columns taxa. The first state is ‘0’, second ‘1’ etc. ‘?’ represent missing data, ‘–’ inapplicable states. Refer to ‘Character descriptions and comments’ for further information on characters. Taxon abbreviations: Ste, Steatoda ; Eno, Enoplognatha ; Sel, Selkirkiella ; Ara, Argyrodes argyrodes ; Are, Argyrodes elevatus ; Koa, Kochiura aulica ; Kor, Kochiura rosea ; Ach, Achaearanea ; The, Theridion ; koh, A. kohi ; lor, A. lorenzo ; rup, A. rupununi ; big, A. biglebowski ; dud, A. dude ; vit, A. vittatus ; pul, A. pulchellus ; nel, A. nelsoni ; rab, A. rabus ; nig, A. nigrescens ; eth, A. ethicus ; may, A. may ; sal, A. sallee ; ana, A. analyticus ; chi, A. chickeringi ; pac, A. pacificus ; dom, A. domingo ; jab, A. jabaquara ; dub, A. dubiosus ; exi, A. eximius ; oct, A. octavius ; juc, A. jucundus ; ari, A. arizona ; bae, A. baeza ; pur, A. puravida ; stu, A. studiosus ; gua, A. guacamayos ; ele, A. elevatus ; tos, A. tosum ; pan, A. pantanal ; ori, A. oritoyacum ; fra, A. fraternus ; tun, A. tungurahua . Abbreviations sp. 1–7 refer to undescribed Anelosimus spp. numbered as in the text and figures.

(90 -0, Figs 14G , 15A ), three palpal tibial trichobothria (118 -1, Fig. 15G ), robust femur I (119 -1, Fig. 55G ) and communal (subsocial) behaviour (145 -1). The clade excluding Anelosimus sp. 1 (this undescribed species differs so much in genitalia from other species that it substantially changes the diagnosis of the Anelosimus and may, for the sake of clarity, later be placed in a new genus) is supported by: elongate (wide) epigynal plate (3 -1, Figs 19C, D , 44C, D, I, K ), incised cymbial mesal margin (15 -1, see e.g. Agnarsson, 2004, fig. 22A), terminally ridged theridiid tegular apophysis (57 -1, Fig. 55D ), lobed embolus tail (68 -1, Fig. 28B ), abdomen pattern (98 -1, Figs 27H–M , 49E ), and absence of sticky silk in web (142 -1, but see Discussion on character 142).

Figures

Figure 8. Anelosimus domingo, male palp from Peru. A, dorsal; B, apicomesal; C, mesal; D, ventral; E, apicoventral; F, ectal. Scale bars: 100 µm.

Figure 14. Anelosimus dubiosus. A, epigynum; B, epiandrous gland spigots; C, female posterior median and posterior lateral spinnerets; D, female prolateral cheliceral teeth; E, male stridulatory pick row; F, male prosoma dorsal; G, female prosoma dorsal. Scale bars: A,D,F,G, 100 µm; B, 20 µm; C, 10 µm; E, 50 µm.

Figure 15. Anelosimus dubiosus. A, male prosomal stridulatory ridges; B, female sternum; C, male prosoma; D, female prosoma; E, male fourth tarsal claws; F, female tip of fourth tarsus; G, female palpal tibia, as typical for Anelosimus with three dorsal trichobothria (arrows). Scale bars: A–D, 100 µm; E, 20 µm; F,G, 50 µm.

Figure 19. The ‘jucundus group’. A–E, Anelosimus arizona. A, male palp mesal, arrow points to the eb-ridge, larger in this species than in any other Anelosimus; B, same ventral; C, epigynum; D, internal epigynum dorsal; E, female habitus. F–I, Anelosimus jucundus. F, male palp mesal; G, same ventral; H, epigynum; I, internal epigynum dorsal. J–N, Anelosimus octavius. J, male palp expanded, mesal; K, male palp ventral; L, male palp expanded, ectal; M, epigynum; N, internal epigynum dorsal.

Figure 27. The ‘jucundus group’. A–M, Anelosimus baeza. A, male palp mesal, from Baeza, Ecuador; B, same, ventral view; C, male palp ventral, from Colombia; D, epigynum, from Baeza, Ecuador; E–G, epigynum, from Las Caucheras, Ecuador. E, ventral view; F, internal epigynum, ventral; G, internal epigynum dorsal; H, male habitus, from Baeza; I, male habitus, from Las Cuacheras; J, female habitus, from Las Caucheras; K, same, profile; L, ectal view of male abdomen, from Baeza; M, ectal view of male abdomen, from Las Caucheras. N–Q, Anelosimus puravida. N, male palp mesal; O, same, ventral; P, epigynum; Q, internal epigynum dorsal.

Figure 28. Anelosimus baeza from Baeza, male palp. A, mesal; B, ventral; C, subectal, note the ectal tegular process (arrow); D, ectal; E, details of conductor and embolus tip, ventral; F, same, mesal view. Scale bars: A–D, 100 µm; E,F, 50 µm.

Figure 44. The ‘studiosus group’. A–E, Anelosimus elegans. A, male palp mesal; B, same ventral; C, epigynum; D, internal epigynum dorsal; E, digital photograph of male holotype. F–O, Anelosimus guacamayos. F, male palp mesal; G, same ventral; H, male palp lacking distal E fork; I, epigynum; J, epigynum, female collected with ‘forkless’ male; K, internal epigynum dorsal; L, female habitus dorsal; M, male habitus dorsal; N, female habitus ectal; O, subadult male habitus ectal.

Figure 48. Anelosimus guacamayos. A,B, epigynum. A, ventral; B, subcaudal. C, epiandrous gland spigots; D, female palpal tibia, dorsal view showing the three trichobothria (arrows); E, female fourth tarsal claws, central claw removed – note strong accessory claws (arrows); F, female cephalic region. Scale bars: A,B,F, 100 µm; C,E, 20 µm; D, 50 µm.

Figure 49. The studiosus group. A–F, Anelosimus studiosus. A, mesal view of male palp; B, ventral view of male palp; C, epigynum; D, internal epigynum dorsal; E, male habitus dorsal; F, female abdomen dorsal. G–J, Anelosimus pantanal. G, male palp mesal; H, same ventral; I, epigynum; J, internal epigynum dorsal. K–N, Anelosimus fraternus. K, male palp (holotype) mesal; L, same ventral; M, epigynum; N, internal epigynum dorsal.

Figure 53. Anelosimus pantanal, female. A, epigynum ventral; B, epigynum, different specimen from same colony; C same, anterior view. D–F, spinnerets. D, all; E, anterior lateral spinneret, anterior is right; F, posterior lateral and posterior median spinnerets. G, fourth tibia, bearing six dorsal trichobothria (arrows); H, third tibia, bearing five dorsal trichobothria (arrows). Scale bars: A–D, 50 µm; E, 10 µm; F, 20 µm; G,H, 100 µm.

Figure 55. Characters I. A, Anelosimus ethicus epigynum, note scape (arrow); B, Anelosimus may epigynum, note anterior septum (arrow); C, Enoplognatha ovata male palp, note unmodified terminal portion of the theridiid tegular apophysis (arrow); D, Argyrodes argyrodes male palp, note simple terminal hook on the theridiid tegular apophysis (vertical arrow) with terminal ridges, and the unique Argyrodes conductor (horizontal arrow); E, Anelosimus sp. 1 (Australia), male palp with a ‘corkscrew’ embolus (arrow); F, Kochiura aulica, male palp, note cymbial marginal setae strongly bent towards the palpal bulb (arrows); G, Anelosimus ethicus male prosoma, note robust femur I (arrow) compared with femur IV.

Figure 58. A strict consensus of the two most parsimonious cladograms (L = 298, CI = 58, RI = 88). Numbers below branches indicate Bremer support, while those above branches are bootstrap values (above 50%); parsimony jackknife scores were nearly identical to the bootstrap scores and are not shown.

Figure 59. The preferred phylogeny with informal clade names and clade numbers in circles at the nodes.