Anelosimus studiosus (Hentz, 1850)

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 : 499-512

THE STUDIOSUS GROUP

Diagnosis: Males of the studiosus group can be separated from the related jucundus group by a smaller, flat, embolic division b (e.g. Fig. 35H ), which is distally much narrower than in species of the jucundus group. The basal lobe of the embolus never surpasses the hood of the subconductor, but is rather hooked in it, or orientated towards it, unlike in the jucundus group. Epigyna are very similar among species, and to those of the jucundus group, but differ from the latter in the strongly sclerotized part of the CD being directly below, or ventral to the ectalmost margin of the spermathecae ( Fig. 49J ). The external epigyna in the studiosus group range from weakly to strongly ridged (e.g. Figs 44C, I , 49C, I, M ), whereas they are always strongly ridged in the jucundus group. Species of the studiosus group are generally smaller than those of the jucundus group, although the overlap is considerable. Palpal organs and epigyna are similarly smaller in the studiosus group than in jucundus the group.

Description: Males with a flat embolic division b that narrows gradually (or sometimes abruptly near the centre) towards tip, the E plus Eb covering only a portion of the tegulum (and other sclerites) ventrally. Tip of the Eb narrow compared with base. The lobe of the embolus usually indistinct not surpassing the hood of the SC. Epigyna are similar among species, but vary in having the epigynal plate weakly to strongly ridged. Strongly sclerotized part of the CD located directly below or ventral to the ectalmost margin of the spermathecae (e.g. Fig. 44D ).

Phylogenetics: The studiosus group ( A. studiosus sensu Levi, 1956, 1963 ) monophyly is supported by four unambiguous synapomorphies ( Fig. 60 ), the following two of which have perfect fit to the cladogram: sclerotized region of copulatory duct, mesal to ectal margin of spermathecae (10 -1, Fig. 44D ), and embolic division b terminally narrow and snout-like (75 -1, Fig. 44A ).

Composition: In his treatment of A. studiosus, Levi (1956: 419) discussed the extensive geographical variation he observed under the subheading ‘subspecies’. It seems clear now that his A. studiosus included numerous related species and here the following eight species are treated: Anelosimus studiosus , A. elegans , A. oritoyacu , A. pantanal , A. tungurahua , A. guacamayos , A. tosum and A. fraternus .

Distribution: From north-eastern USA to Argentina ( Figs 63C , 64C, E ). Most speciose in Ecuador, particularly at altitudes of 1000 m or above.

Natural history: Species of the studiosus group range from subsocial to social.

ANELOSIMUS TOSUM CHAMBERLIN, 1916

( FIGS 35A–F , 36– 38 , 64E )

Types: Female holotype from Peru, Huadquiña, vi.1911, Yale Peruvian Expedition, in MCZ, examined

Synonymies:

Theridion tosum Chamberlin, 1916: 229 , pl. 16, figs 1–4, ♂ ♀.

Anelosimus jucundus: Levi, 1956: 417 , synonymy here rejected.

Etymology: Chamberlin (1916) did not explain the species epithet.

Diagnosis: Anelosimus tosum has stark contrasting dark and white spots within the dorsal band on the abdomen ( Fig. 35E, F ). Males differ from most other species of the studiosus group in having a large Eb and a distinctly lobed E base entering or slightly surpassing the SC ( Fig. 35B ). It differs from the closely similar A. oritoyacu in having a broader and more rugose Eb tip and a smaller E tip fork ( Fig. 35B ). Females are difficult to separate from others in the A. studiosus group, but have unusually numerous ridges on the epigynal plate, especially towards the posterior end ( Fig. 37A ). Morphologically this species shows some characters intermediate to the studiosus and jucundus groups.

Male (IA40618): Total length 3.58. Prosoma 1.63 long, 1.25 wide, 1.07 high, brown, broad thicker bands around rim, and centre. Sternum 0.99 long, 0.91 wide, extending between coxae IV, dark brown. Abdomen 2.08 long, 1.90 wide, 1.98 high. Pattern as in other Anelosimus . Eyes subequal, about 0.08 in diameter. Clypeus height about 3.1 times AME diameter. Chelicerae with one large and two small prolateral teeth, 4–5 denticles retrolaterally. Leg I femur 2.41, patella 0.59, tibia 2.11, metatarsus 1.79, tarsus 0.81. Femur about 6 times longer than wide, metatarsus I about 14 times longer than wide. Leg formula 1423. Leg base colour yellowish, with distal tip of all segments darkened, tibia also with darker central bands, and femur I slightly darker than other segments. Tarsal organs distinctly distal (0.65–0.70) on tarsi I–II, central on III (0.50), slightly distal on IV (0.50–0.55). Four to five small trichobothria dorsally on all tibia, 5 on tibia I and III. Trichobothria on metatarsi I–III proximal (about 0.40–0. 50), absent on metatarsus IV. Two prolateral and one retrolateral trichobothria on palpal tibia. Palp as in Figures 35A, B , 36A–F .

Female (IA40618): Total length 5.20. Prosoma 2.28 long, 1.82 wide, 1.40 high, brown, broad thicker bands around rim, and centre. Sternum 1.30 long, 1.07 wide, extending between coxae IV, dark brown. Abdomen 3.25 long, 2.48 wide, 2.64 high. Pattern as in Figure 35E, F . Eyes subequal, about 0.10 in diameter. Clypeus height about 4.0 times AME diameter. Chelicerae with one large and two small prolateral teeth, 4–5 denticles retrolaterally. Leg I femur 2.28, patella 0.81, tibia 2.11, metatarsus 2.02, tarsus 0.98. Femur about 5 times longer than wide, metatarsus I about 13 times longer than wide. Leg formula 1423. Leg base colour yellowish with distal tip of all segments darkened, and a central band on femora and tibia, most distinct on legs I and II. Tarsal distal on tarsi I (0.70–0.75) and II (0.60–0.65), proximal on III (0.45–50) and IV (0.35–4.00). Six to seven small trichobothria dorsally on all tibia, 6–7 on tibia I, 6 on tibia III. Trichobothria on metatarsi I–III proximal (0.40–0.45), absent on metatarsus IV, distal (0.85) on palpal tarsus. Three dorsal trichobothria on palpal tibia. Epigynum as in Figures 35C, D , 37A .

Variation: Male total length 3.58–3.90, prosoma length from 1.5–1.69, first femur 2.21 – 2.60. Female total length 4.23–5.90, prosoma 1.85–2.28, first femur 2.21–2.60.

Additional material examined: BRAZIL. Rio de Janeiro, Teresopolis [22°24′0″S, 42°58′0″W], iii.1946, 900– 1000 m (H. Sick, AMNH), 1♀ [cf. IA40696] . COLOMBIA. Antioquia, San Vicente [6°17′0″N, 75°20′0″W], 2.i.1985 (M. A. Serna, NMNH), 1♀ [cf. IA40662] . Boyacá, Santuario de Fauna y Flora Iguaque, near margin of Laguna Iguaque (5°41′20″N, 73°26′7″W), 5–8.ii.1998, canopy fogging, 3450−3650 m (J. Coddington et al., NMNH), 1♂, 1♀ [IA111301]; 5♂ [IA1112]; 5–8.ii.1998, 2800 m (G. Hormiga et al., NMNH), 1♂, 1♀ [IA40529]; 1♂, 1♀ [IA40618]; 29 juv [cf. IA40630]; Near visitors centre (5°42′5.3″N, 73°27′20.1″W), 5–8.ii.1998, 2850− 3000 m (G. Hormiga et al., NMNH), 2♀ [IA40409]; 4♂, 17♀ [IA40531]; 3♂, 4juv [IA40631]; 2♀ [IA40633]; 1♂, 1♀ [IA40639]; 1♀ [IA40637]; 5.ii.1998, 1♂, 2♀, 2juv [IA40638]; 8.ii.1998, 2800 m (G. Hormiga, NMNH), 1♀ [IA40632] . Cundinamarca, La Calera, Cerro del Chocolatero , c. 5 km NE of Bogotá [4°42′0″N, 73°58′0″W], 31.i.1998, 3000 m (G. Hormiga et al., NMNH), 2♀, 15juv [IA40534]. Valle de Cauca, Atuncela [3°46′0″N, 76°42′0″W], 22.xi. 1969, 300 m ( MCZ), 1♀ [cf. IA40668] . ECUADOR, Chimborazo, c. 6 km NE of Chunchi on Panamerican Highway (2°15′48.96″S, 78°53′19″W), 10.vii.2004, 2380 m (I. Agnarsson et al., NMNH), ♂♂ ♀♀ [ IAV09 ] . Cañar, W of Suscal (2°28′1.2″S, 79°7′6.6″W), ♂♂ ♀♀ [ IAV10 ]. EL SALVA- DOR. San Salvador, Santa Tecla [13°40′0″N, 89°17′0″W], 15.x.1949 (J. B. Boursot, AMNH), 1♀ [cf. IA40216] . MEXICO. Michoacán, Pátzcuaro [19°30′0″N, 101°36′0″W], 12.vi.1941 (A. M. Davis, AMNH), 1♀, 1juv [cf. IA40517]; Tancítaro [19°19′0″N, 102°21′0″W], vi-vii.1941, c. 2000 m (H. Hoogstraal, MCZ), 1♂ [IA0222] . PERU. Cajamarca, Cuttervo [6°22′0″S, 78°48′0″W], 22.vi.1956, 2900 m ( V. Vegr., MCZ), 1♂, 8♀ [IA050201] . La Libertad, Pataz [7°43′0″S, 77°37′0″W], 26.iii.1988, 2000 m (D. Silva, MHNSM), 1♀, 8juv [IA40545]; Yalen [7°45′0″S, 77°33′0″W], 26.iii.1988 (D. Silva, MHNSM), 1♂, 2♀, 5juv [IA40551] .

Distribution: Found from Mexico to Peru ( Fig. 64E ), most records from 2000 m or above.

Taxonomic history: Chamberlin (1916) described this species based on a single female from Peru. Levi (1956: 417) considered all females with a strongly ridged epigynal plate to be A. jucundus and thus synonymized it with A. jucundus . It is clear now that species of both the jucundus and the studiosus group can have strongly ridged epigynal plates, and although A. tosum females resemble A. jucundus , the resemblance is closer to specimens from Colombia (also based on colour pattern), which based on the male clearly belong to the studiosus group. Examination of the type female of A. tosum also suggests that it belongs to the studiosus group, as the strongly sclerotized portions of the CD appear (without dissection) to be relatively close together, rather than extending ectal to the ectalmost rim of the spermathecae as is typical in the jucundus group. Nevertheless, it is notoriously difficult to identify unaccompanied females, and matching the type female to the description series from Colombia represents nothing more than a best guess.

Natural history: Information on the natural history of this species comes from field labels, and field notes of G. Hormiga (pers. comm.), who collected the description series, and my personal observations made in Ecuador in 2004. Most colonies encountered consisted only of juveniles, or an adult female with juveniles. This appears to be a typical subsocial species with single-mother nests where the mother dies before the young reach adulthood. Colonies were common in forest gaps, along edges (trails) and in secondary growth, but were rare in the forest understory. Heteropteran commensals ( Ranzovius ) and argyrodine kleptoparasites were seen in some nests.

ANELOSIMUS ORITOYACU SP. NOV.

( FIGS 35G–J , 39–41 , 63C , 66D )

Types: Male holotype, four male paratypes, and female paratype from Napo, Oritoyacu, 8.1 km S. of Baeza, Ecuador, 0°29.83′S, 77°52.43′W, 6.i.2002, L. Avilés, deposited in NMNH [ IA40626 ] .

Synonymy:

Anelosimus studiosus Levi, 1956 (in part); Levi 1963 (in part).

Etymology: The species epithet is a noun in apposition referring to the name of the type locality.

Diagnosis: Males can be diagnosed from all other Anelosimus of the studiosus group by the relatively large Eb and stout embolus fork ( Fig. 35H ); females are very difficult to separate from others, but may be identified by the epigynal plate being unusually high relative to its width ( Fig. 35I ).

Male (holotype): Total length 2.99. Prosoma 1.50 long, 1.17 wide, 0.91 high, brown, nearly covered with dark grey markings. Sternum 0.92 long, 0.81 wide, extending between coxae IV, brown nearly covered with dark grey markings. Abdomen 1.69 long, 1.24 wide, 1.16 high. Pattern similar to A. guacamayos . Eyes subequal, about 0.09 in diameter. Clypeus height about 3.4 times AME diameter. Chelicerae with one large and two small prolateral teeth, 4–5 denticles retrolaterally. Leg I femur 2.08, patella 0.65, tibia 1.79, metatarsus 1.50, tarsus 0.75. Femur about 7 times longer than wide, metatarsus I about 15 times longer than wide. Leg formula 1243. Legs base colour light brown, femur I, distal half of femur II, and tips of other femora, tibia, patella and metatarsus dark brown. Tarsal organs distal (0.50–0.55) on tarsi I–II, proximal (0.35–0.40) on III–IV. Four to six small trichobothria dorsally on all tibia, 5 on tibia I, 5 on tibia III. Trichobothria on metatarsi I–III proximal (0.40–0.45), absent on metatarsus IV. Two prolateral and one retrolateral trichobothria on palpal tibia. Palp as in Figures 35G, H , 39A–F , 40A–F .

Female (paratype): Total length 3.51. Prosoma 1.63 long, 1.27 wide, 1.04 high, brown with dark grey markings. Sternum 1.06 long, 0.86 wide, extending between coxae IV, brown with dark grey markings. Abdomen 2.02 long, 1.60 wide, 1.65 high. Pattern similar to A. guacamayos . Eyes subequal, about 0.10 in diameter. Clypeus height about 3.1 times AME diameter. Chelicerae with one large and two small prolateral teeth, 4–5 denticles retrolaterally. Leg I femur 1.95, patella 0.62, tibia 1.50, metatarsus 1.40, tarsus 0.72. Femur about 5 times longer than wide, metatarsus I about 11 times longer than wide. Leg formula 1423. Legs pale brown with distal tips of femora, tibia, patella and metatarsus dark brown. Tibia I also with a central band ventrally. Tarsal organs distal (0.55–0.60) on tarsus I, central (0.50) on II, proximal (0.40–0.45) on III–IV. Five to six small trichobothria dorsally on all tibia, 5 on tibia I, 5 on tibia III. Trichobothria on metatarsi I and III proximal (about 0.40– 0.45), central on II, absent on metatarsus IV. Three or four dorsal trichobothria on palpal tibia. Epigynum as in Figures 35I, J , 41A .

Variation: Male total length 2.80–3.05, prosoma 1.45–1.55, first femur 2.05–2.20. Female total length 3.35–3.70, prosoma 1.55–1.70, first femur 1.85–2.10. Female palpal tibia has three or four dorsal trichobothria; this variation can be asymmetric, i.e. differing between sides of the individual. Specimens from Jacala, Mexico, are large, 3.60 (male), 4.40 (female) but their identity is uncertain.

Additional material examined: ECUADOR. Chimborazo, 30 miles SW of Alausí [2°22′0″S, 79°4′0″W], 14.xi.1955, 2500 m ( CAS), 1♂ [IA40778]; 2♂, 2♀ [IA40783] . Napo, 3.9 km S of Baeza (0°28′0″S, 77°71′9″W), 13.i.2002, hand collected, c. 1500 m (L. Avilés, NMNH), 2♂ [ IALA29 ]; Oritoyacu 8.1 km S of Baeza (0°29’83′S, 77°52’43′W), 23.xi.2002, hand collected, c. 1500 m (P. Salazar & G. Iturralde, NMNH), 1♂, 1♀ [ IALA30 ] . Pichincha, Tandayapa [0°1′0″S, 78°46′0″W], xi.1984, 1700 m (G. Onore, MCZ), 1♂, 9♀ [cf. IA40764] . MEXICO. Hidalgo, Jacala [21°1′0″N, 99°12′0″W], 13.vi.1936, c. 1600 m (Davis, AMNH), 1♂, 2♀ [cf. IA40218] . Guerrero, Taxco , c. 1800 m, [18°34′0″N, 99°37′0″W], 15.viii.1943 (Bolivar, Pelaez & Osorio, AMNH), 1♂ [IA40202] . Morelos, Cuernavaca , c. 1500 m, [18°55′0″N, 99°13′0″W] (N. Banks, NMNH), 1♂ [IA40510] .

Distribution: Only known from Ecuador and Mexico ( Fig. 63C ). All collections made at altitudes around 1500 m or above.

Natural history: Anelosimus oritoyacu is social, with biased sex ratio. It seems to be mostly confined to the canopy, where it makes nests containing multiple adult males and females (L. Avilés, pers. comm.; my pers. obs.). As the sheet is placed in the canopy, the webs lack the aerial threads so typical of Anelosimus species , and rather resemble the webs of A. rupununi and A. lorenzo ( Fig. 66D ).

ANELOSIMUS TUNGURAHUA SP. NOV.

( FIGS 35K–Q , 42–43 , 64D )

Types: Male holotype and female paratype from Ecuador, Tungurahua, Baños, 1800 −2000 m, iii.1939, W. C. Macintyre ( MCZ), examined .

Synonymy:

Anelosimus studious: Levi, 1956: 419 (in part).

Etymology: The species epithet is a noun in apposition after the type locality.

Diagnosis: Males can be distinguished from most species by the shape of the Eb, which narrows abruptly near the mid region, similar to A. studiosus ( Fig. 35L ). It differs from A. studiosus in the Eb being much flatter, with a narrower and less rugose distal tip. Females may be separated from others by the large epigynal lip, and relatively small epigynal plate ( Fig. 35P ).

Male (holotype): Total length 2.60. Prosoma 1.30 long, 1.04 wide, 0.79 high, brown, with centre darker. Sternum 0.83 long, 0.71 wide, extending between coxae IV, dark brown. Abdomen 1.43 long, 1.16 wide, 1.12 high. Pattern as in Figure 35M . Eyes subequal, about 0.10 in diameter. Clypeus height about 2.1 times AME diameter. Chelicerae with one large and two small prolateral teeth, 4–5 denticles retrolaterally. Leg I femur 2.15, patella 0.55, tibia 1.82, metatarsus 1.66, tarsus 0.72. Femur about 7 times longer than wide, metatarsus I about 15 times longer than wide. Tarsus I with a ventral row of thickened setae. Leg formula 1243. Leg base colour yellowish-brown, distal tip of all segments darker, femur 1 darker than other femora. Tarsal organs distal (0.55) on tarsi I–II, proximal (0.40–0.45) on III–IV. Four to five small trichobothria dorsally on all tibia, five on tibia I and III. Trichobothria on metatarsi I–III proximal (0.35–0.45), absent on metatarsus IV. Two prolateral and one retrolateral trichobothria on palpal tibia. Palp as in Figures 35K, L , 42A–F .

Female (paratype): Total length 3.90. Prosoma 1.69 long, 1.20 wide, 0.99 high, yellowish-brown with centre and rim darker. Sternum 1.02 long, 0.89 wide, extending between coxae IV, dark brown. Abdomen 2.34 long, 1.90 wide, 2.03 high. Pattern as in Figure 35N, O . Eyes subequal, about 0.09 in diameter. Clypeus height about 3.0 times AME diameter. Chelicerae with one large and two small prolateral teeth, 4–5 denticles retrolaterally. Leg I femur 2.02, patella 0.52, tibia 1.79, metatarsus 1.69, tarsus 0.78. Femur about 6 times longer than wide, metatarsus I about 14 times longer than wide. Leg formula 1423. Leg base colour yellowish-brown, distal tip of all segments darker, femora and tibia also with central bands, especially prominent on leg I. Tarsal organs distal on tarsi I (0.55–0.60) and II (0.50–0.55), proximal on III (0.45–50) and IV (0.40–0.45), distal (0.85) on female palp. Four to seven small trichobothria dorsally on all tibia, 6–7 on tibia I, five on tibia III. Trichobothria on metatarsi I–III proximal (0.40–0.45, absent on metatarsus IV. Three, or sometimes four ( Fig. 43G ), dorsal trichobothria on female palpal tibia. Epigynum as in Figures 35P, Q , 43A .

Variation: Male total length 2.28–3.12 prosoma 1.00– 1.50, first femur 1.43–2.15. Female total length 3.19– 4.50 mm, prosoma 1.43–1.69, first femur 1.83–2.02.

Additional material examined: ECUADOR. Tungurahua, Baños [1°23′0″S, 78°25′0″W] , vii.1938 (W. C. Macintyre, MCZ), 17♂, 45♀, c. 1800 m, [IA050101] ; 24.iv.1939, 1800 m, 3♂ [IA050701] ; iii.1939, 1♂, 1♀ 10.iv.1939, 4♂, 8♀, 1juv [IA050301] ; iv.1939, 1850− 2000 m (W. C. Macintyre, MCZ), 1♂, 2♀ [IA050401] ; iii.1939, 5♂, 16♀, 5juv [IA052401]; vii-viii.1938, 2000 m (W. C. Macintyre, MCZ), 3♂, 1♀ [IA051101] ; 15–21.vi.1943 ( MCZ), 3♂, 21♀ [IA051301] .

Distribution: Only known from area of type locality ( Fig. 64D ), at altitudes of 1800−2000 m.

Natural history: Subsocial (L. Avilés, pers. comm.).

ANELOSIMUS ELEGANS NEW REPLACEMENT NAME

( FIGS 44A–E , 45 , 46 , 64E )

Types: Anelosimus elegans is a replacement name for Enoplognatha dubia Chamberlin, 1916 , 60: 233, pl. 17, fig. 3, a junior secondary homonym of Brattia (?) dubia Tullgren, 1910 (= Anelosimus dubius , see Miller, 2004). Female holotype of E. dubia came from Sorontoy, Peru (7000 ft.), in MCZ, examined. Not A. studiosus (Hentz) , contra Levi (1956: 419).

Synonymies:

Enoplognatha dubia Chamberlin, 1916 , preoccupied by Brattia (?) dubia Tullgren, 1910 (= A. dubius , see Miller, 2004).

Anelosimus studiosus: Levi (1956: 419 , in part), synonymy here rejected.

Etymology: The species epithet refers to their unusual and elegant posture, standing with legs in a plane parallel to the body, opened to the sides (L. Avilés, pers. comm.). The males of this species are also unusually shiny.

Diagnosis: Although differing in posture, colour intensity (shininess) and behaviour, this species is morphologically very similar to the sympatric A. guacamayos . The male is readily distinguished from most other Anelosimus by the gradually narrowing Eb ( Fig. 44A ), but is slightly less gradual than in A. guacamayos . Females are difficult to separate from others in the studiosus group, but have a relatively evenly broad epigynal plate ( Fig. 44C ), more gently curved than in, for example, A. studiosus . In A. elegans spermathecae are further apart than in other species of the studiosus group, so that the strongly sclerotized part of the CD is mesal to the spermathecae, instead of lying directly underneath them ( Fig. 44D ). Anelosimus elegans is apparently less social than A. guacamayos , having equal sex ratios and predominantly single female nests.

Male (IA40627): Total length 2.67. Prosoma 1.37 long, 1.04 wide, 0.86 high, dark shiny brown. Sternum 0.83 long, 0.71 wide, extending between coxae IV, dark brown. Abdomen 1.56 long, 1.16 wide, 1.07 high. Pattern as in Figure 44E . Eyes subequal, about 0.08 in diameter. Clypeus height about 3.0 times AME diameter. Chelicerae with one large and two small prolateral teeth, 4–5 denticles retrolaterally. Leg I femur 1.79, patella 0.59, tibia 1.59, metatarsus 1.40, tarsus 0.65. Femur about 6 times longer than wide, metatarsus I about 14 times longer than wide. Leg formula 1243. Leg formula 1243. Leg base colour orange, femora I and II darker brown and distal tip of femora III–IV and tibia darker. Tarsal organs slightly distal (0.50–0.55) on tarsi I, proximal (0.30–0.45) on II–IV. Four to six small trichobothria dorsally on all tibia, 5 on tibia I, 5 on tibia III. Trichobothria on metatarsi I–III proximal (about 0.35–0.40), absent on metatarsus IV. Two prolateral and one retrolateral trichobothria on palpal tibia. Palp as in Figures 44A, B , 45A–F .

Female (IA40627): Total length 3.77. Prosoma 1.89 long, 1.49 wide, 1.11 high, yellowish with sparse dusky grey markings, more concentrated in the cephalic region, and especially the thoracic groove. Sternum 1.16 long, 0.96 wide, extending between coxae IV, yellowish with dense dusky grey markings. Abdomen 2.02 long, 1.73 wide, 1.57 high. Pattern as in the male; see Figure 44E . Eyes subequal, about 0.09 in diameter. Clypeus height about 3.4 times AME diameter. Chelicerae with one large and two small prolateral teeth, 4–5 denticles retrolaterally. Leg I femur 2.28, patella 0.72, tibia 1.95, metatarsus 1.79, tarsus 0.75. Femur about 5 times longer than wide, metatarsus I about 12 times longer than wide. Leg formula 1423. Leg base colour yellowish, distal tip of femora, patella, tibia and metatarsi slightly darkened. Tarsal organs distal (0.60) on tarsus I, central (0.50) on II, proximal (0.30–0.40) on III–IV, most proximal on IV. Five to seven small trichobothria dorsally on all tibia, 5–6 on tibia I, 5–6 on tibia III. Trichobothria on metatarsi I–III proximal (about 0.30–0.45), absent on metatarsus IV. Three dorsal trichobothria on palpal tibia. Epigynum as in Figures 44C, D , 46A .

Variation: Male total length 2.47–2.67, prosoma 1.24– 1.37, femur 1.66–1.79. Female total length 3.77–4.03, prosoma 1.83–1.89, first femur 2.02–2.28. A female identified as A. cf. elegans from San Geronimo, Mexico, measured 5.85 in total length, prosoma 2.15, first femur 2.67.

Additional material examined: BRAZIL. Rio Grande do Sul, Pelotas [31°46′0″S, 52°19′0″W], 2.iii.1964, c. 0–5 m (C.M. Biezanko, MCZ), 1♀ [cf. IA053201] . COLOMBIA. Putumayo, Sibundoy [1°10′0″N, 76°53′0″W], viii.1963, 2200 m (M L. Bristol, MCZ), 1♂, 2♀ [IA051701] . ECUADOR. Morona Santiago, km 20 from Limón to Gualaceo , cloud forest (3°0′15.84″S, 78°30′50″W), 11.vii.2004, 2270 m (I. Agnarsson et al., NMNH), ♂♂ ♀♀ [ IAV12 ] . Napo, Cordillera Guacamayos, Cocodrilo (0°38′75″S, 77°47′45″W), 11.xii.2002 (L. Avilés, NMNH), 2♂, 1♀ [IA40563]; 4♂, 3♀ [IA40627] . Pichincha, near Mindo , roadside (0°1’35.79′S, 78°47’20.39′W), 25.vii.2004, c. 1400− 1800 m (I. Agnarsson and G. Iturralde, NMNH), ♀♀ [ IAV11 ] . MEXICO. Guerrero, Taxco (18°34′0″N, 99°37′0″W), viii.1978, c. 1800 m (P. Klass, NMNH), 1♀ [cf. IA0218] . Morelos, Cuernavaca [18°54′0″N, 99°13′0″W], viii.1996, c. 1500 m, hand collected (W. Maddison, MCP), 1♂ [ IALA13 ] . Oaxaca, San Geronimo [16°34′0″N, 95°6′0″W] ( AMNH), 1♀ [cf. IA40524] . PERU. Junín, Utcuyacu [11°40′0″S, 75°0′0″W], iii.1948, c. 3500 m (F. Woytkowski, AMNH), 1♂, 10♀ [cf. IA40695]; 15♂, 40♀, 50juv [IA40746] . Pasco, 15 km SE. of Oxapampa on Carretera Nueva a Villa Rica, Pampa (10°40′0″S, 75°18′0″W), 20.vi.1986 (D. Silva,), 4♂, 1♀ [IA1109] . Pasco, Oxapampa , 15 km SE of Oxapampa [10°40′0″S, 75°18′0″W], 20.vi.1986, 2000 m (D. Silva, MHNSM), 1♂, 4♀ [cf. IA40596]; 1♂, 4♀, 65juv [cf. IA40602]; 4♂, 2♀ [IA40605] .

Distribution: Occurs from Mexico to Peru ( Fig. 64E ). It is not certain that specimens from Mexico belong to this species; they are very large, but otherwise closely resemble specimens from other areas. Likewise the identity of the specimen from Brazil is in doubt, an unaccompanied female collected near sea level; all other records of A. elegans come from altitudes of 1300−3100 m.

Natural history: In Ecuador A. elegans is apparently subsocial, and has unbiased sex ratios (L. Avilés, pers. comm.; my pers. observ.). A sample from Peru contained c. 15♂, 40♀ and 50 juveniles, indicating that the level of sociality may differ between populations. The argyrodine Faiditus caudatus has been collected in the webs of A. elegans .

Taxonomic note: Based on subtle differences in morphology and different levels of social behaviour A. elegans and A. guacamayos are here treated as separate species. There is some evidence, however, that populations of A. elegans may be social (see Natural history) and further data are necessary to investigate if the observed variation is intra- or interspecific.

ANELOSIMUS GUACAMAYOS SP. NOV.

( FIGS 44F–O , 47 , 48 , 63C , 66B )

Types: Male holotype and female paratype from Ecuador, Cordillera Guacamayos, Cocodrilo , 0°38.75′S, 77°47.45′W, 14.xii.2002, P. Salazar, deposited in NMNH [ IA40624 ] .

Synonymy:

Anelosimus studiosus: Levi (1956: 419 , in part); Levi (1963: 36, in part).

Etymology: The species epithet is a noun in apposition based on the name of the type locality.

Diagnosis: Anelosimus guacamayos differs in behaviour from most species of the studiosus group, being social. Behaviour apart, separating this species from others in the studiosus complex is difficult. Males can be distinguished by the shape of the embolic apophysis, whose ectal edge is straighter than in other species ( Fig. 44F–H ). Some A. guacamayos males lack the fork at the embolus tip ( Fig. 44H ) reminiscent of A. fraternus from Haiti. I have not found a reliable way of separating the females morphologically from other species of the studiosus complex, except from A. elegans which has CD located more mesally in the internal epigynum.

Male (holotype): Total length 2.86. Prosoma 1.37 long, 1.09 wide, 0.84 high, brown, dusky grey markings most noticeable in centre, in streaks towards the rim and on rim. Sternum 0.86 long, 0.79 wide, extending between coxae IV, brown, dusky grey markings get denser closer to pedicel. Abdomen 1.63 long, 1.20 wide, 1.09 high, truncated in front. Pattern as in Figure 44M, O . Eyes subequal, about 0.10 in diameter. Clypeus height about 2.7 times AME diameter. Chelicerae with one large and two small prolateral teeth, 4– 5 denticles retrolaterally. Leg I femur 1.92, patella 0.49, tibia 1.69, metatarsus 1.56, tarsus 0.72. Femur about 9 times longer than wide, curved, metatarsus I about 17 times longer than wide. Leg formula 1243. Leg base colour brown, distal tip of tibia darker, coxae, trochanters and base of femora lighter than other parts. Tarsal organs central (0.50) on tarsus I, proximal (0.40–0.45) on II–IV. Five to seven small trichobothria dorsally on all tibia, five on tibia I, II and III. Trichobothria on metatarsi I–III proximal (0.35–0.40), absent on metatarsus IV. Two prolateral and one retrolateral trichobothria on palpal tibia. Palp as in Figures 44F, H , 47A–E .

Female (IALA34): Total length 4.05. Prosoma 1.80 long, 1.40 wide, 0.50 high, brown. Sternum 1.10 long, 0.90 wide, extending midway between coxae IV, brown. Abdomen 2.30 long, 1.65 wide, 1.70 high, pattern as in Figure 44L, N . Eyes subequal, about 0.10 in diameter. Clypeus height about 2.6 times AME diameter. Chelicerae with one large and two small prolateral teeth, 4–5 denticles retrolaterally. Leg I femur 2.30, patella 0.65, tibia 1.80, metatarsus 1.80, tarsus 0.90. Femur I about 7 times longer than wide, tibia I about 14 times longer than wide. Leg formula 1423. Leg base colour light brown, distal tip of femur and tibia dark grey, femora also with a central band. Tarsal organs usually slightly distal (0.55) on tarsi I and II, slightly proximal (0.45) on III and IV, distal (0.85) on female palp (but the positions vary slightly among specimens). Five to seven small trichobothria dorsally on all tibia, five on tibia III. Trichobothria on metatarsi I–III proximal (about 0.35), absent on metatarsus IV. Three dorsal trichobothria on female palpal tibia. Epigynum as in Figures 44I–K , 48A, B .

Variation: Male total length 2.34–2.95, prosoma 1.24– 1.45, femur I 1.80–1.95. Males in some collections lack the fork at the embolus tip (compare Fig. 44G to 44H), a variation among, but not within colonies. It is uncertain if this represents intraspecific variation or if ‘forkless’ males belong to a different species. While provisionally treated as conspecifics here, the more common forked condition is scored in the data matrix as all those from the type locality have the fork. Female total length 3.70–4.42, prosoma 1.80–1.89, first femur 2.28–2.30.

Additional material examined: ECUADOR. Napo, Cordillera Guacamayos, Cocodrilo , 18.8 km S of Cosanga river (0°38’75′S, 77°51′0″W), 29.xi.2002, hand collected, c. 1500 m (P. Salazar, NMNH), 1♂, 2♀ [ IALA31 ]; 11.i.2002 (L. Avilés, NMNH), 4♂, 2♀ [ IALA32 ]; 7.i.2002, 1♀ [ IALA33 ]; 15.5 km S. of Cosanga river [0°38’9′S, 77°47’4′W], 19.viii.1999, hand collected (L. Avilés, NMNH), 1♂, 1♀ [ IALA34 ]; 17.6 km S. of Cosanga river [0°38′9″S, 77°47′4″W], 19.viii.1999, hand collected (L. Avilés, NMNH), 1♂, 4♀ [ IALA35 ]; 16.65 km S of Cosanga (0°38’79′S, 77°47’45′W), 8.i.2002 (L. Avilés, NMNH), 6♂ [IA40556]; 17.6 km S of Cosanga, Cordillera de los Guacamayos (90°45′0″S, 77°51′0″W), 19.viii.1999, hand collected (L. Avilés, NMNH), 6♀ [ IALA02 ]; 1♂, 5juv [ IALA0301 ]; Cocodrilo (0°38′75″S, 77°47′45″W), 11.xii.2002 (L. Avilés, NMNH), 2♀ [IA40562]; NE of El Chaco, bridge at Rio Salado (0°12′9″S, 77°42′5″W), 19.vii.2004, 1290 m (I. Agnarsson et al., NMNH), [ IAV02 ], ♀♀ . Morona Santiago, km 6.7 from Limón Indanza (2°59′33″S, 78°26′3″W), 12.vii.2004, 1415 m (I. Agnarsson, NMNH), ♂♂ ♀♀ [ IAV01 ] .

Distribution: Known from Ecuador ( Fig. 63C ). This appears to be a mid-elevation species, found between 1200 and 2000 m.

Natural history: A. guacamayos is social with distinct primary sex ratio bias (L. Avilés, pers. comm.; my pers. obs.). It makes large, typical basket-shaped nests ( Fig. 66B ), and forms colonies of up to 1000 or more individuals, including numerous adult females and their clutches.

Most nests are found in open areas, typically in clearings, for example old landslides where often a cluster of nests may be found.

Taxonomic note: Here, specimens lacking the fork at the embolus tip ( Fig. 44H ) are treated as synonymous with the type. Apart from this detail, ‘forkless’ males otherwise strongly resemble the holotype. Females collected with ‘forkless’ males are not separable from females collected with the more common (‘normal’) males. Based on this evidence I presume that this difference represents variation within a species, but further studies should explore the possibility of these being two distinct species.

ANELOSIMUS STUDIOSUS ( HENTZ, 1850)

( FIGS 49A–F , 50 , 51 , 64C )

Types: Hentz’s types of Theridion studiosum , from Alabama, USA, have been lost ( Levi, 1956). Their identity is not problematic however; Hentz’s description is recognizable, and other specimens from Alabama are very similar to specimens from across the USA.

Synonymies:

Theridion studiosum Hentz, 1850 , 6: 274, pl. 9, fig. 5, ♀.

Theridion studiosum: Hentz 1875 , 145, pl. 16, fig. 5, ♀ Keyserling, 1884, 1: 20, pl. 1, fig. 7, ♂ ♀ (in part); Marx, 1890, 12: 520; 1892, 2: 156; Simon, 1894a, 1: 540; 1894b, 521; 1897, 862; Banks, 1902, 11: 272; Banks, 1903, 55: 340; 1904, 56: 125; 1906, 22: 187; Simon, 1903, 2: 989; Petrunkevitch, 1911, 29: 207; 1925, 27: 68; Comstock, 1912, 350, fig. 250, ♀ 1940, 365, fig. 350 ♀ Bishop and Crosby, 1926, 41: 183; Chickering, 1936, 55: 452; Fox, 1940, 53: 43; Mello- Leitão, 1941, 13: 250; 1946, 11: 36; Roewer, 1942, 1: 501; Muma, 1945, 38: 29.

Anelosimus studiosus : F. O. P.- Cambridge, 1902, 2: 395, pl. 37, figs 16–17, ♂ ♀ (probably A. studiosus , drawings difficult to recognize); Banks, 1910, 72: 20; Mello-Leitão, 1942, 2: 385; 1943, 37: 171; 1944, 3: 313; 1948, 100: 382; Gertsch, 1949, 167. Kaston, 1948, 20: 99, figs 178–181, ♂ ♀ Levi, 1956, 75: 407–422 (in part), figs 21, 23, possibly also figs 37– 39 ♂ ♀. Stejskal, 1976, 26: 344, fig. 4.2 ♀ (note that Stejskal’s photos are not recognizable); Kaston, 1981, 890; Breene et al., 1993, 56, fig. 20A–C ♂ ♀ Platnick, 2006; Agnarsson, 2004, figs 24(A–G), 25(A–F) ♂ ♀.

Anelosimus textrix, Chamberlin and Ivie, 1944 , 8(5): 37, probably an incorrect synonymy of Linyphia textrix Walckenaer, 1842 , 2: 281. L. textrix was illustrated in an unpublished manuscript ‘Spiders of Georgia’ by J. Thomas Abbot, cited in Walckenaer as ‘Abbot, 1792’ and is considered valid, see Levi, 1956: 419; Platnick, 2006; Mello-Leitão, 1945, 4: 215; Archer, 1946, 22: 54; 1950, 30: 22, pl. II, fig. 5– 6 ♂. Kaston, 1953, 166; Barnes, 1953, 23: 321.

Theridion magnificum Keyserling, 1884 , 2(1): 47–48, pl. 2, fig. 26, ♂ ♀.

Diagnosis: Males can be distinguished by the sharp constriction of the Eb centrally, and a relatively broad Eb tip ( Figs 49B , 50B ). Females are difficult to separate from others of the studiosus group, but most specimens differ from all other Anelosimus by having a more strongly procurved genital plate ( Fig. 49C ; note however that this feature is not universal, see under Variation).

Male (IA40656): Total length 2.65, prosoma 1.30 long, 0.90 wide, 0.75 high, brownish-orange with scattered grey flecks, narrow rim around the edge slightly darker. Sternum 0.75 long, 0.65 wide, extending between coxae IV, yellowish, with a darker rim around edge. Abdomen 1.40 long, 1.10 wide, 1.20 high, pattern as in Figure 49E . Eyes subequal, about 0.08 diameter. Clypeus height about 2.3 times AME diameter. Chelicera with one large and two small prolateral teeth, three or four denticles retrolaterally. Legs pale yellow, part of femur I and distal tip of tibia I slightly darker. Leg formula 1243. Leg I femur 1.80, patella 0.55, tibia 1.65, metatarsus 1.40, tarsus 0.65. Femur I about 7 times longer than wide, tibia I about 12 times longer than wide, thickening somewhat towards distal end. Four to five small trichobothria dorsally on all tibia, five on tibia I and III. Trichobothria on metatarsus I (0.35), II (0.40) and III (0.40) proximal, absent on metatarsus IV. Tarsal organs on tarsus I (0.45), II (0.40), III (0.35), and IV (0.40) proximal.Two prolateral and one retrolateral trichobothria on palpal tibia. Palp as in Figures 49A, B , 50A–E .

Female (IA40503): Total length 4.40, prosoma 1.60 long, 1.25 wide, 1.00 high, brownish-orange with grey flecks in cephalic area and narrow rim along edges. Sternum 0.90 long, 0.80 wide, extending between coxae IV, brownish-yellow, mostly covered by dusky grey dots, with a darker rim around edge. Abdomen 2.85 long, 2.45 wide, 3.00 high, pattern as in Figure 49F . Eyes subequal, about 0.10. Clypeus height about 2.9 times AME diameter. Chelicera with one large and two small prolateral teeth, three denticles retrolaterally. Legs pale yellow, part of femur I and distal tip of tibia I slightly darker. Leg formula 1423. Leg I femur 2.00, patella 0.60, tibia 1.70, metatarsus 1.55, tarsus 0.75. Femur I about 6 times longer than wide, tibia I about 9 times longer than wide. Four to seven small trichobothria dorsally on all tibia, 6–7 on tibia I, five on tibia III. Trichobothria on metatarsus I (0.40), II (0.40) and III (0.40) proximal, absent on metatarsus IV. Tarsal organ central (0.50) on leg I, proximal on legs II (0.45), III (0.40), and IV (0.45), distal (0.85) on palpal tarsus. Epigynum as in Figures 49C, D , 50H .

Variation: Male total length 2.00–3.50, prosoma 1.10–1.55, femur I 1.40–2.30. Female total length 3.50–4.70, prosoma 1.35–1.90, femur I 1.80–2.40. A male from Chapala, Mexico, measured 3.90 mm in total length, but the identity of the specimen is dubious. Like in other species of the studiosus complex the female epigynum of A. studiosus is highly variable. In the majority of specimens the epigynal plate is narrower and more strongly procurved than in any other species. However, in some specimens the plate may be broader and less procurved, then very similar to A. guacamayos (e.g. Fig. 44I ) and other species. The setae ventrally on metatarsus I are sometimes thickened in the males, sometimes not. Behaviour appears to be variable also, some populations prominently subsocial, others social.

Additional material examined: ARGENTINA. Córdoba, La Serrapita, Alta Garcia, Depto. Santa María [31°39′0″S, 64°25′0″W], 24.x.1967, c. 500 m (di Tada, MCZ), 1♂, 1♀ [cf. IA051001]. Santiago del Estero [27°47′0″S, 64°16′0″W], 2.iv.1965, c. 180 m (H. Levi, MCZ), 1♀ [cf. IA052701]. BRAZIL. Rio de Janeiro, Guanabara, Barra de Tijuca [23°0′0″S, 43°21′0″W], 16.iv.1965 (H. Levi, MCZ), 1♀ [cf. IA052801]; Petropolis [22°31′0″S, 43°11′0″W], 2–5.xi. 1945, 850 m (H. Sick, AMNH), 2♂, 3♀ [IA40303]. Rio Grande do Sul, Canela (29°21′0″S, 50°48′0″W), 2–8.i.1992, c. 600 m, hand collected (J.W. Thome, MCP), 1♀ [cf. MCP02]; Cordilheira, Cachoeira do Sul [30°2′0″S, 52°53′0″W], 30.xii.1993, c. 60 m (R. G. Buss, MCP), 1♀ [cf. IA40611]; Fazenda Souza, nr. Caxias [29°7′0″S, 51°1′0″W], 3–5.xi.1994, hand collected (A. A. Lise, MCP), 4♂ [cf. MCP01]; 18–21.xi.1993, 2♂, 4♀ [cf. IA40609]; Pelotas [31°46′0″S, 52°19′0″W], 2.iii.1964, c. 0–5 m (C. M. Biezanko, MCZ), 2♀ [IA050801]; São Francisco de Paula [29°26′0″S, 50°34′0″W], 21– 24.iii.1996, c. 600 m (A. A. Lise et al., MCP), 4♀ [cf. IA40607]; Viamão (30°5′0″S, 50°58′0″W), 22.xi.1995, hand collected (A. A. Lise, MCP), 2♀ [cf. MCP03]. COLOMBIA. Magdalena, Gaira [11°10′0″N, 74°13′0″W], xii.1975, 10 m (W. Eberhard, MCZ), 2♀ [cf. IA40667]. Meta, Pto. Lieras, Lomalinda (3°18′0″N, 73°22′0″W), 300 m (B. T. Carrol, MCZ), 2♀ [IA40761]; Villavicencio [4°9′0″N, 73°38′0″W], 12.xii. 1979, 470 m (M. Barreto, MCZ), 1♂ [IA40767]. Putumayo, Sibundoy [1°10′0″N, 76°53′0″W], viii.1963, 2200 m (M. L. Bristol, MCZ), 1♀ [IA053101]. Valle, above Piekindixii, 1972, 1800 m (MCZ), 1♀ [cf. IA052301]; Atuncala [3°46′0″N, 76°42′0″W], 17.xii.1969, c. 800 m (W. Eberhard, MCZ), 1♀ [cf. IA052101]; 1♂, 1♀ [IA051401]; 22.xi.1969, 1♀, 10juv [IA053001]; 23.xi.1969, 1♂ [IA051601]; Between Dagua and Loboguerrero [3°43′0″N, 76°40′0″W], 10.vii. 1970, 800 m (H. Triana, MCZ), 1♂, 1♀ [IA051201]; Río Calima [3°42′0″N, 76°33′0″W], v.1976, 1400 m (W. Eberhard, MCZ), 2♂, 3♀ [IA050601]; nr. Pance, P. N. N. Farallones de Cali, Res. Nat. Hato Viejo (3°20′53″N, 76°40′16.7″W), 12.ii.1998, 2300 m (G. Hormiga, NMNH), 6♂, 2♀ [IA40661]. Western Cordillera, Between Queremal and Buenaventura, 12.ii.1935 (H. F. Schwarz, AMNH), 1♀ [IA40702]. COSTA RICA. Alajuela, near Esparta [9°59′0″N, 84°40′0″W], xi.1981, c. 300 m (MCZ), 1♂ [IA40771]; Grecia [10°4′0″N, 84°18′0″W], 27.xi.1955 (B. Malkin, AMNH), 1♂ [IA40108]. Cartago, Turrialba [9°54′0″N, 83°41′0″W], 10–17.iv. 1944, 600 m (F. Schrader, AMNH), 1♀ [IA40112]; 23.vii-15.viii.1965 (A. M. Chickering, MCZ), 1♂ [IA40750]. Guanacaste, Comunidad [1033′0″N, 85°35′0″W], 19.ii.1967 (J. M. Nelson, MCZ), 1♀ [cf. IA0211]; Palo Verde, Bagaces [10°31′0″N, 85°15′0″W], 16– 22.i.1978 (W. Eberhard, MCZ), 1♂ [IA40765]. Heredia, Heredia, Universidad Nacional Autónoma (10°0′0″N, 84°7′0″W), 1–2.iv.1989 (J. Coddington, NMNH), 3♂, 3♀ [IA40559]. San José, San José [9°55′0″N, 84°4′0″W] (E. Schmidt, AMNH), 3♂ [IA40107]; Escazú [9°55′0″N, 84°8′0″W], 30.vii.1983,1300 m (H. Levi, MCZ), 1♀ [IA40772]; San Pedro de Montes de Oca [9°55′0″N, 84°2′0″W], iii.1983, 1000 m (W. Eberhard, MCZ), 1♀ [IA40756]; vii.1988 (W. Eberhard, NMNH), 6♀ [cf. IA40641]; 15 km N. of Puriscal [9°58′0″N, 84°19′0″W], vii. 1988, 600 m (W. Eberhard, NMNH), 2♀ [cf. IA40635]. CUBA. Ciudad de La Habana, Siboney, Oriente [22°4′0″N, 82°27′0″W], 26.vi.1955 (A. F. Archer, AMNH), 1♀ [IA40572]. ECUADOR. Pichincha, Calderón [0°5′0″S, 78°26′0″W], 5.vii.1989, c. 2500 m (L. Avilés, NMNH), 1♂ [IALA16]; 1♂ [IALA18]; area no censada [0°5′0″S, 78°26′0″W], 5.vii.1989, c. 2500 m (L. Avilés, NMNH), 3♂ [IALA17]; Ilalo [0°3′0″S, 78°32′0″W], viii.1999, hand collected (L. Avilés, MCP), 3♂, 1juv [IALA09]; Pululahua Crater [0°3′0″S, 78°32′0″W], viii.1999 (L. Avilés, NMNH), 3♂ [IALA15]; San Antonio del Tingo [00’6′S, 78°27′0″W], 1.vii.1989 (L. Avilés, NMNH), 1♂ [IALA19]; 1♂, 3juv [IALA20]. Tungurahua, Río Pastaza between Baños and Mapoto [1°25′0″N, 78°10′0″W], viii.1938 (W. C. Macintyre, MCZ), 1♂ [IA050901]; Río Pastaza near Mapoto [1°25′0″N, 78°10′0″W], 2.iv.1938, 1300 m (W. C. Macintyre, MCZ), 1♀ [cf. IA051901]. EL SALVADOR. San Salvador, San Salvador [13°42′0″N, 89°12′0″W], iii.1954 (J. B. Boursot, AMNH), 1♀ [IA40104]. GUATEMALA. Baja Verapaz, Los Ramones [15°0′0″N, 90°12′0″W], 25.vii.1947 (C. & P. Vaurie, AMNH), 1♂ [IA40105]. HAITI. l’Artibonite, Carrefour [19°24′0″N, 72°4′0″W], 23vii.1955 (A. F. Archer, AMNH), 1♀ [cf. IA40576]. HONDURAS. Atlandida, Lancetillal [14°54′0″N, 89°7′0″W], vii. 1929, 900 m (A. M. Chickering, MCZ), 1♀ [IA40751]. [no detailed locality] (Dyer, AMNH), 1♂ [IA40212]. JAMAICA. St. Andrew, [18°1′0″N, 76°54′0″W], xixii.1957 (A. M. Chickering, MCZ), 1♀ [IA40754]; Mona pasture [18°1′0″N, 76°54′0″W], 29.xi.1957 (MCZ), 3♂, 1♀, 15juv [IA40752]. St. Catherine, Spanish Town [17°59′0″N, 76°57′0″W], 15.viii.1974 (D. B. Jayasingh, MCZ), 1♀ [IA40753]; old harbor [17°58′0″N, 77°0′0″W], xi.1957 (A. M. Chickering, MCZ), 1♀ [IA40755]. Westmoreland, Negril [18°16′0″N, 78°20′0″W], 23–30.iii.1981, 90 m (H. & L. Levi, MCZ), 1♀ [IA40760]; Whitehouse [18°4′0″N, 77°59′0″W], 26.iii.1955 (A. M. Nadler, AMNH), 1♂ [IA40312]. MEXICO. Chiapas, Cintalapa [16°41′0″N, 93°42′0″W], 17.ix.1947 (H. Wagner, AMNH), 2♂ [IA40710]; Las Cruces Arriaga [16°17′0″N, 93°48′0″W], 18.ix.1947 (H. Wagner, AMNH), 2♂ [IA40678]; Ocosingo [16°54′0″N, 92°7′0″W], 25.vi. 1950, 900 m (C. & M. Goodnight and L. J. Stannard, AMNH), 1♂ [IA40304]; Río de las Flores [17°10′0″N, 91°16′0″W], 15.ix.1947 (H. Wagner, AMNH), 1♂ [IA40713]; 1♂ [IA40715]; 16.ix.1947, 1♀ [IA40210]; Tuxtla Gutiérrez [16°44′0″N, 93°6′0″W], 9.ix.1947 (H. Wagner, AMNH), 2♂, 2♀ [IA40221]. Distrito Federal, Coyoacan [19°20′0″N, 99°10′0″W], 28.vii.1947 (H. Wagner, AMNH), 2♂ [IA40729]. Guerrero, Acapulco, El Mirador Hotel [16°51′0″N, 99°54′0″W], 5.vi.1943 (F. H. Pough, AMNH), 1♀ [IA40725]; Taxco, km. 100, Rd. Taxco [18°45′0″N, 99°48′0″W], 10.viii.1946 (Goodnight, Bolivar & Bonet, AMNH), 1♀ [IA40204]. Hidalgo, 6 mi. N. of Jacala [21°5′0″N, 99°11′0″W], 23.vi.1955 (C. & P. Vaurie, AMNH), 1♂ [IA40688]; Chapulhuacán [21°9′0″N, 98°53′0″W], 20.v.1952 (M. Cazier et al., AMNH), 1♂ [IA40716]; Ixmiquilpan [20°29′0″N, 99°13′0″W], 15.viii.1947 (H. Wagner, AMNH), 1♂ [IA40119]; Jacala [21°1′0″N, 99°12′0″W], 13.vi.1936, c. 1600 m (A. M. Davis, AMNH), 1♂, 1♀ [IA40218]. Jalisco, Chapala [20°17′0″N, 103°11′0″W], 22vi.1941, c. 1500 m (A. M. Davis, AMNH), 1♂ [cf. IA40213]. Michoacán, Tzararacua Falls, 7 mi. from Uruapan [19°20′0″N, 102°4′0″W], 14.vi.1941 (A. M. & L. I. Davis, AMNH), 1♂, 2♀ [IA40203]. [♀ Michoacán], 7 mi. S. of Hidalgo [19°36′0″N, 100°34′0″W], 3.vii.1936 (L. I. Davis, AMNH), 1♀ [IA40699]. Morelos, Cuernavaca [18°55′0″N, 99°13′0″W], 3.vii.1941 (A. M. & L. I. Davis, AMNH), 1♀ [IA40679]; x.1944 (N. L. H. Krauss, AMNH), 1♂ [IA40717]; Oaxtepec [18°54′0″N, 98°57′0″W], 17v.1942 (AMNH), 1♀ [cf. IA40686]. Nayarit, 15 mi. W. of Tepic [21°29′0″N, 105°5′0″W], 25.vii.1954 (W. J. Gertsch, AMNH), 2♀ [IA40703]; Jesús María Cortés [21°43′0″N, 104°53′0″W], 25– 30.vii.1955 (B. Malkin, AMNH), 1♂ [IA40692]; Tepic [21°29′0″N, 104°53′0″W], 26.vii.1953 (P. & C. Vaurie, AMNH), 1♂ [IA40109]; 4.viii.1953, 1♂ [IA40120]; 2.viii.1947 (C. J. Goodnight, AMNH), 1♂, 1♀ [IA40217]; 2–7.viii.1947 (C. & M. Goodnight & B. Malkin, AMNH), 1♀ [IA40706]. Nuevo León, Horsetail Falls [25°21′0″N, 100°8′0″W], 11.vi.1936 (L. I. Davis, AMNH), 1♀ [IA40683]; Horsetail Falls, Cola de Caballo, San Juan R. Canyon [25°21′0″N, 100°8′0″W], 31.viii.1968 (J. E. Carico, NMNH), 1♀ [IA40401]; Linares [24°51′0″N, 99°33′0″W], 8.vii.1941 (L. I. Davis, AMNH), 1♂ [IA40215]. Oaxaca, Asunción, Nochixtlán [17°27′0″N, 97°17′0″W], 5.vii.1953, 2000 m (A. Robinson Jr., AMNH), 1♂ [IA40208]; Oaxaca [17°3′0″N, 96°43′0″W], 2.x.1946, 1550 m (H. Wagner, AMNH), 2♂, 1♀ [IA40727]. Puebla, Tecamalchalco [18°52′0″N, 97°43′0″W], 2.vii.1953, 2000 m (A. Robinson Jr., AMNH), 1♂ [IA40681]; Tehuacán [18°27′0″N, 97°23′0″W], 17–24.x.1944 (H. Wagner, AMNH), 2♀ [IA40205]. San Luis Potosí, Huichichuyán [21°30′0″N, 98°57′0″W], 19.v.1952 (AMNH), 1♂ [IA40707]; Nr. Ciudad del Maíz [22°24′0″N, 99°36′0″W], 19.viii.1947 (C. & M. Goodnight, AMNH), 1♀ [IA40207]; Tamazunchale [21°15′0″N, 98°47′0″W], 20.v.1952 (M. Cazier, W. Gertsch, & R. Schramme, AMNH), 2♂ [IA40201]; Valles [21°59′0″N, 99°0′0″W], vii.1959 (Steude, AMNH), 1♀ [IA40698]. Tamaulipas, 11 mi. N. of Victoria [23°54′0″N, 99°9′0″W], 22.v.1952 (M. Cazier et al., AMNH), 1♂ [IA40711]; 40 mi. S. of Linares [24°54′0″N, 98°14′0″W], 30.xi.1939 (A. M. & L. I. Davis, AMNH), 1♀ [IA40724]; Reynosa [26°4′0″N, 98°17′0″W], 2.v.1936 (J. Ruth, AMNH), 2♂, 1♀ [IA40712]. Veracruz, 14 mi. S. of Catemaco on Rt. 180 [18°13′0″N, 95°6′0″W], 23.vi. 1982, 400 m (F. Coyle, MCZ), 1♂ [IA051501]; Aroyac [19°3′0″N, 96°6′0″W], 12.xi.1941 (F. Bonet, AMNH), 1♀ [IA40685]; Papantla de Olarte [29°27′0″N, 97°19′0″W], 12.x.1947 (H. M. Wagner, AMNH), 1♂ [IA40209]; pass above Orizaba [18°50′0″N, 97°5′0″W], 29.vi.1944, 1950 m (L. I. Davis, AMNH), 1♀ [IA40310]; Tecolutla [20°29′0″N, 97°0′0″W], 13.x.1947 (H. M. Wagner, AMNH), 1♂ [IA40705]; Tlapacoyan [19°58′0″N, 97°12′0″W], 7– 8.vii.1946 (H. Wagner, AMNH), 1♀ [IA40206]. Taxco, viii.1978 (P. Klass, MCZ), 1♀ [IA40763]. [México ♀], Cerro Gordo [19°9′0″N, 100°7′0″W], 22.vi.1936 (AMNH), 1♀ [IA40219]. NETHERLANDS ANTI- LLES. Curacao, Curacao [12°4′0″N, 68°34′0″W], 22.xii.1962 (B. deJong, MCZ), 2♂, 2♀ [IA090101]. PANAMA. Bocas del Toro, Bocas del Toro, Corriente grande (9°18′0″N, 82°32′0″W), 13–17.iii.1980, hand collected (R. Ibanez, NMNH), 1♀ [IA0204]. Chiriqui, Boquete [8°46′0″N, 82°25′0″W], 4–11.viii.1954, 1100 m (A. M. Chickering, MCZ), 1♂, 1♀ [IA40747]; Renacilmento, 10 km W. of Volcán [9°13′0″N, 83°32′0″W], 10.viii.1983,1300 m (H. & L. Levi, MCZ), 1♀ [IA40773]. Panamá, Canal Zone, Barro Colorado Island [9°9′17″N, 79°50′53″W], 20.iv.1953 (A. M. Nadler, AMNH), 1♀ [IA40220]; v.1964 (A. M. Chickering, MCZ), 1♂, 1juv [IA40766]; 18–29.viii.1939, 1♂, 1♀ [IA060101]; 16.vi.-15.vii.1930, 3♀ [IA071001]; 3♀ [cf. IA40541]; Panama Canal Zone, Colon humid forest (9°21′0″N, 79°54′0″W), 2–14.vii.1979, canopy fogging, tree 4 (E. Broadhead et al., NMNH), 1♀ [IA40414]; tree 6 (E. Broadhead et al., NMNH), 2♂ [IA40416]; Panama City [8°57′0″N, 79°32′0″W], 15–30.vii.1979, canopy fogging (E. Broadhead et al., NMNH), 1♂ [IA40406]. TRINIDAD. Victoria, Gasparillo [10°19′0″N, 61°25′0″W], 4.xi.1944 (R. H. Montgomery, AMNH), 1♀ [cf. IA40704]. URUGUAY. Montevideo, Montevideo [34°54′0″S, 56°9′0″W], x-xi.2000 (F. Costa & C. Viera, NMNH), 1♂, 1♀ [IA40568]. USA. Alabama, Baldwin Co., Lagoon [30°21′0″N, 87°35′0″W], 12.x.1951 (A. F. Archer, AMNH), 1♂, 1♀ [IA40694]; Mobile Co., Dauphin Island [30°14′0″N, 88°6′0″W], 20.iv.1948 (A. F. Archer, AMNH), 2♀ [IA40574]; Tuscaloosa Co., Tuscaloosa [33°12′0″N, 87°34′0″W] (A. F. Archer, AMNH), 5♀, 9juv [IA40571]. Connecticut, Cobalt [41°33′0″N, 72°33′0″W], 2.viii.1939 (B. J. Kaston, NMNH), 1♂ [IA40652]. District of Colombia, Washington DC, Rock Creek Park (38°56′0″N, 77°2′0″W), 16.vii.1985, hand collected (J. Coddington, NMNH), 1♂ [IA1108]; (38°53′0″N, 77°1′0″W), 24.vi.1982, hand collected (J. Coddington, NMNH), 1♂ [IA1106]; National Arboritum [38°54′0″N, 76°58′0″W], 15.vi.1989 (J. Coddington, NMNH), 4♂, 2♀ [IA40503]. Florida, 5 mi. W. of Marianna [30°46′0″N, 85°19′0″W], 17.xi.1972 (A. Moreton, MCZ), 3♀ [IA042301]; 7 mi. E. of Apopka [28°40′0″N, 81°24′0″W], 20.viii.1944 (M. Ninenberg, AMNH), 1♀ [IA40118]; Alachua [29°47′0″N, 82°29′0″W], 10.v.1941 (H. K. Wallace, AMNH), 2♂, 3♀ [IA40106]; 28.iv.1937 (AMNH), 1♂ [IA40113]. Florida, Alachua Co. [29°42′0″N, 82°21′0″W], 8.iv.1938 (AMNH), 1♀ [IA40308]; Auburndale, Polk Co. [28°3′0″N, 81°47′0″W] (N. Banks, MCZ), 1♀ [IA041501]; Biscayne Bay [25°41′0″N, 80°9′0″W] (N. Banks, MCZ), 1♀ [IA041301]; Charlotte co., Punta Gorda [26°56′0″N, 82°3′0″W], 1–16.i.1946 (S. Rounds, AMNH), 1♀ [IA40690]; Clay Count’s Hammock, Alachua Co. [29°42′0″N, 82°21′0″W], 30.iii.1939 (H. Wallace, AMNH), 1♂ [IA40301]; Collier Co., Naples [26°8′0″N, 81°47′0″W], 18.i.1946 (S. Rounds, AMNH), 1♀ [IA40674]; Cox’s Hammock, Dade Co. [25°38′0″N, 80°30′0″W], 28.xii.1940 (Archer, AMNH), 1♀ [IA40102]; Dunedin [28°1′0″N, 82°47′0″W], 1924 (W. S. Blatchley, MCZ), 1♀ [IA40777]; Earmra SE. Isl. Everglades National Park, Dade Co. [25°14′0″N, 80°51′0″W], 28.i.1973 (A. Sheldon, MCZ), 1♂ [IA040701]; Dade co., Everglades, 28.xii.1950 (A. M. Nadler, AMNH), 1♂ [IA40693]; (in hammock area), [25°38′0″N, 80°24′0″W], 19.vi.1975 (L. Roth, MCZ), 1♂, 1♀ [IA042101]; Kendall [25°41′0″N, 80°19′0″W], 4.iii.1953 (A. M. Nadler, AMNH), 1♀ [IA40211]; 31.iii.1953 (A. M. Nadler, AMNH), 1♂, 1♀ [IA40684]; La Belle [26°45′0″N, 81°26′0″W], 25.xii.1952 (B. J. Kaston, NMNH), 2♀ [IA40654]; Lake Okeechobee [26°56′0″N, 80°47′0″W], 17.ii.1943 (W. Proctor & M. Cazier, AMNH), 1♀, 1juv [IA40111]; Lake Placid, Highlands Co. [27°17′0″N, 81°21′0″W], 25.ii.1976 (H. Levi, MCZ), 1♂, 1♀ [IA041401]; 3.ii.1943 (M. Cazier, AMNH), 1♀ [IA40114]; 25.i.1943, 1♀ [IA40115]; 1943, 1♀ [IA40101]; 3.ii.1943, 2♀ [IA40302]; 26.i.1943, 6♀ [IA40314]; Leesburg, Lake co. [28°48′0″N, 81°52′0″W], 1–11.iii.1954 (M. Statham, MCZ), 5♂, 1♀, 7juv [IA041901]; Matheson Hammock, 1/2 M south of Dade Co. [25°44′0″N, 80°18′0″W], 1952 (P. Porter, MCZ), 1♀ [IA041201]; Miakka River State Park, nr. Sarasota [27°20′0″N, 82°31′0″W], 6.iv.1936 (Gertsch, AMNH), 1♀ [IA40117]; Miami Beach [25°48′0″N, 80°7′0″W], vi.1944 (A. Bacon, AMNH), 1♀ [IA40116]; Naples [26°8′0″N, 81°47′0″W] (AMNH), 1♀ [IA40110]; Nassau Co. [30°36′0″N, 81°43′0″W], 28.iv.1935 (H. K. Wallace, AMNH), 1♀ [IA40676]; Orange Co., Orlando [28°33′0″N, 81°22′0″W], 11–14.xi.1946 (A. F. Archer, AMNH), 1♂, 6♀ [IA40575]; Orange Park, Trismen Estate [30°10′0″N, 81°43′0″W], 13.xi.1942 (MCZ), 1♀ [IA40775]; Orlando [28°33′0″N, 81°22′0″W], 15– 30.viii.1944 (M. Nirenberg, AMNH), 1♀ [IA40103]; Pine Crest off Tamiami Trail [25°44′0″N, 80°66′0″W], 1.iii.1936 (S. C. Bishop, AMNH), 1♀ [IA40313]; Royal Palm Hammock [25°59′0″N, 81°35′0″W], 21.i.1946 (S. Rounds, AMNH), 1♂, 3♀ [IA40675]; Sebastian [27°49′0″N, 80°29′0″W], 30.xi.1931 (G. Nelson, MCZ), 1♂ [IA041801]; Tampa, Mac Dill Field [27°57′0″N, 82°27′0″W], 15–19.iii.1943 (B. Malkin, AMNH), 3♂ [IA40311]; Tavernier [25°0′0″N, 80°31′0″W], 29.xi.1952 (A. M. Nadler, AMNH), 1♀ [IA40214]; Wewahitchka, Dead Lake [30°6′0″N, 85°12′0″W], 6.iv.1927 (MCZ), 1♂ [IA041001]; Winter Park [28°35′0″N, 81°21′0″W], 11.iv.1938 (W. J. Gertsch, NMNH), 1♂ [IA40656]; (N. Banks, MCZ), 3♂, 5♀ [IA042401]. Georgia, 5 mi. N. of Macon [32°54′0″N, 83°38′0″W], 15.vi.1939 (W. J. Gertsch, AMNH), 1♂ [IA40691]; Bar-M-Ranch, S. of Boston [30°45′0″N, 83°47′0″W], 25.vi.1978 (H., L. & F. Levi, MCZ), 1♀ [IA40776]; Gainesville [34°17′0″N, 83°49′0″W], 28.v.1943 (B. J. Kaston, AMNH), 2♂ [IA40309]; 8.vi.1940, 1♂ [IA40653]; Rabun Co., Talullah [34°43′0″N, 83°23′0″W], 20.viii.1961 (J. E. Carico, NMNH), 1♀ [IA40659]; St. Simons Island [31°8′0″N, 81°24′0″W], v.1911 (S. C. Bishop, AMNH), 1♂ [IA40307]. Kansas, Elk Lake City, Montgomery Co. [37°17′0″N, 95°54′0″W], 10.ix.1994 (H. Guarisco, MCZ), 10♂, 4♀, 14juv [IA040201]. Louisiana, Saint Tammany Co., Mandeville [30°21′0″N, 90°4′0″W] (R. V. Chamberlin, MCZ), 1♀ [IA40774]; Cheniere au Tigre, Vermilion Par [29°34′0″N, 92°12′0″W], 27.iv.1974 (D. A. Rossman, MCZ), 1♀ [IA052501]. Maryland (Drury, AMNH), 2♂ [IA40315]. Mississippi, Wilkinson Co., Centreville [31°5′0″N, 91°4′0″W], 1944 (A. F. Archer, AMNH), 1♂, 3♀ [IA40573]. North Carolina, Apex, Wake Co. [35°43′0″N, 78°51′0″W], 28.ix.2000 (I. Agnarsson, NMNH), 7juv [IA40403]; Ashville [35°35′0″N, 82°34′0″W], 8.vii.1933 (W. J. Gertsch, AMNH), 1♂, 1♀ [IA40682]; Carteret co. [34°49′0″N, 76°46′0″W], 29.vii.1950 (R. B. Barnes, AMNH), 50 juv. [IA40687]; Raleigh [35°47′0″N, 78°38′0″W], viii.1912 (C.S. Brimley, MCZ), 1♀ [IA040801]; (C. S. Brimley, NMNH), 1♀ [IA40501]; Schenck forest, Raleigh [35°47′0″N, 78°38′0″W], 20.ix.2000 (I. Agnarsson, NMNH), 15 juv [IA40402]; Swan Co., Oconolufte visitor centre [35°29′0″N, 83°19′0″W], 21.vii.1998 (F. Coyle & I. Agnarsson, NMNH), 2♀ [IA40657]; Sylva [35°22′0″N, 83°13′0″W], 16.iv.1938 (B. B. Fulton, AMNH), 5 juv [IA40306]; Wake Co., Apex [35°43′0″N, 78°51′0″W], 28.ix.2000 (J. Perry, NMNH), 21 juv [IA40660]. South Carolina, Charleston [32°47′0″N, 79°56′0″W], 15–30.vi.1943 (B. Malkin, AMNH), 1♀ [IA40680]; Horrey Co., Socastee, Maccaman River [33°41′0″N, 79°0′0″W], 3.vii.1961 (J. E. Carico, NMNH), 2♀ [IA40658]; McClellanville [33°5′0″N, 79°27′0″W], vii-viii.1945 (P. Vaurie, AMNH), 1♂ [IA40305]. Tennessee, Martel [35°48′0″N, 84°14′0″W], 8.vii.1950 (M. Cazier, NMNH), 1♀ [IA40677]. Texas, 3 mi. E. of Edinburg [26°18′0″N, 98°6′0″W], 12.iv.1937 (S. Mulaik, AMNH), 1♂, 1♀ [IA40721]; 7 mi. E. of Edinburg [26°17′0″N, 98°3′0″W], 14.x.1936 (S. M., AMNH), 2♀ [IA40316]; Beaumont [30°5′0″N, 94°7′0″W], iv-vi.1946 (E.D. Parmer, MCZ), 1♂, 1♀ [IA041601]; Beeville, Bee Co. [28°24′0″N, 97°44′0″W], 25.iv.1982 (D. Bickel, MCZ), 1♂ [IA040401]; Cameron Co. [41°26′0″N, 78°11′0″W], ix.1933 (S. Mulaik, NMNH), 1♂, 1♀, 8juv [IA40655]; i-iii.1936 (L. I. Davis, AMNH), 1♀ [IA40728]; Corpus Cristi [27°46′0″N, 97°24′0″W], 21.iii.1936 (AMNH), 1♂ [IA40697]; Cotulla [28°26′0″N, 99°13′0″W], 8.vii.1936 (L. I. Davis, AMNH), 3♀ [IA40719]; Dallas, Denton Co [32°46′0″N, 96°47′0″W], 11.x.1947 (S. E. Jones, MCZ), 1♀ [IA040901]; Edinburg [26°17′0″N, 98°9′0″W], 10.xii.1935 (Stockton, AMNH), 1♀ [IA40708]; 1.xii.1936 (S. Mulaik, AMNH), 1♀ [IA40720]; 10.x.1935 ([collector unknown], AMNH), 1♀ [IA40722]; 15.xi.1935 (J. L. Ledbetter, AMNH), 2♀ [IA40730]; Goose Island State Park, Aransas Co. [28°1′0″N, 97°2′0″W], 16.vi.1961 (A.R. Brady, MCZ), 3♂, 1♀ [IA042001]; Harlingen [26°11′0″N, 97°41′0″W] (AMNH), 1♂, 1♀ [IA40714]; 25.x.1936 (S. Mulaik, AMNH), 1♀ [IA40732]; Kingsville [27°30′0″N, 90°51′0″W], x.1934 (AMNH), 1♀ [IA40718]; 4.xi.1934 (S. Mulaik, AMNH), 4♀, 2juv [IA40731]; Liberty [30°3′0″N, 94°47′0″W], 12.vi.1937 (AMNH), 1♀ [IA40723]; Newton [30°50′0″N, 93°45′0″W], 13.viii.1938 (L. I. Davis, AMNH), 1♀ [IA40701]; S. of Pharr [21°11′0″N, 98°11′0″W], 5.iv.1936 (M. Welch, AMNH), 3♂, 1♀ [IA40689]; Stony Oaks, Denton Co. [33°13′0″N, 97°7′0″W], 11.vi.1944 (MCZ), 1♀ [IA041701]; Zapata Co. [27°1′0″N, 99°14′0″W], 10.iv.1936 (Welch, AMNH), 1♂ [IA40726]. Virginia, 1407 N. Garland, Fayetteville, Arlington [36°4′0″N, 94°9′0″W], 9.xi.1986 (Richard Leschen, MCZ), 1♀, 25juv [IA040101]; Dismal swamp W. of Lake Drummond, 7 mi. S of Suffolk, Nansemond Co. [36°38′0″N, 76°31′0″W], 18–19.v.1968 (E. Sabath, MCZ), 25♂, 9♀, 10juv [IA042201]; Great Falls [38°59′0″N, 77°17′0″W] (N. Banks, MCZ), 2♂ [IA041101]; Hampden-Sydney, Prince Edward Co. [37°14′0″N, 78°27′0″W], 20.vi.1982 (H. Levi, MCZ), 1♀ [IA040501]; Waynesboro, Augusta Co. [38°4′0″N, 78°53′0″W], 3.viii.1981, from webs (L. & H. Levi, MCZ), 11♀ [IA040601]. VENEZUELA. Mérida, Mérida, [8°36′0″N, 71°9′0″W] (Y. Lubin, MCZ), 3♀ [cf. IA030901]. Monagas, Jusepín [9°45′0″N, 63°31′0″W], xi.1974 (MCZ), 15♀ [IA052901]. Patos, 24.ix.1944 (R. H. Montgomery, AMNH), 1♀ [IA40745].

Distribution: USA to Argentina. Widespread and common over a large portion of the Americas from 39°N to 33°S and altitudes of 0–2500 m. In a range of habitats, but apparently not in lowland tropical rainforests.

Natural history: Extensive literature is available about the natural history and various aspects of the biology of Anelosimus studiosus (e.g. Simon, 1891; Brach, 1977; Buskirk, 1981; Avilés, 1987, 1993, 1999; Avilés & Maddison, 1991; Avilés & Gelsey, 1998; Furey, 1998; [note that Stejskal’s (1976) report from Venezuela cannot be specifically linked to A. studiosus ]). It is typically characterized as a predominantly subsocial species, with a single mother and offspring in each nest. Yet, some populations have up to several hundreds of individuals building large communal webs with extensive co-operation ( Furey, 1998). Apparently the behaviour of this species is thus highly variable. Brach (1977) discusses A. studiosus in Florida. He found that the majority of colonies consisted of a single female and its brood of the same or adjacent instar. The juveniles were fed by regurgitation at first but then started catching prey in the mother’s web. The mother usually disappeared by the time her juveniles reached the 5th instar. Eventually most of the offspring dispersed from their natal nest (often after mating there), but sometimes a single female or a female and a couple of juveniles (and males) stayed and started a new colony in the old nest. As they mature, the offspring are pursued out of the web by the mother, if still alive, but otherwise by the first female to mature. Males are always tolerated, but contribute little and can only handle tiny prey. The mother lays a single egg case at the time containing 31– 47 eggs. Individuals may wander around the web in search of small prey, but when a large prey item becomes entangled it is usually attacked first by the mother and then juveniles converge to feed on it. Juveniles were dependent on the mother and could not survive without her until after the third instar. Conspecific juveniles were never attacked (introduced or not) but juveniles of other theridiids were killed and eaten within a day of having been introduced. Introduced adult females were accepted by the brood but vigorously attacked by the mother. Brach (1977) attempted to create colonies by putting several adult females together, but these would show no co-operation and high aggression resulting in frequent cannibalism. Sometimes they killed prey together, but did not feed communally.

Furey (1998) studied A. studiosus in Tennessee, and as for Brach (1977) found that the majority of nests were single female plus offspring, but some nests had more than one female, with up to 29 adults in a nest. Although multi-female nests are fewer, contrary to Brach’s study, Furey (1998) found that 80% of A. studiosus females he encountered lived in groups. Females in solitary webs drove out other females whereas those in social webs accepted most of them and in general showed inter-attraction. He established multiple female nests with non-sibling females that lasted 3 years. In multi-female nests mothers did not discriminate between egg sacs (own and others).

Furey (1998) found female-biased sex ratio ranging between 4 and 6 females per male and concluded that the Tennesse populations are social.

Given that the studies were made on different populations, this may represent interspecific rather than intraspecific variation. However, I have not found any consistent morphological differences between specimens collected in subtropical versus northern temperate USA and all are thus treated as conspecific here.

Several mirid bugs of the genus Ranzovius have been reported living in the nests of A. studiosus , including R. contubernalis Henry , R. clavicornis (Knight) , R. fennahi Carvalho , and R. stysi ( Henry, 1984, 1999; Wheeler & McCaffrey, 1984).

Taxonomic note: Anelosimus studiosus as here circumscribed is both highly variable and very widespread, despite greatly limiting Levi’s (1956, 1963) concept of the species. The possibility that A. studiosus as currently delimited still represents a number of species certainly deserves further scrutiny. Conversely, it is also possible that the within-species variation is greater yet, and e.g. A. fraternus and A. tungurahua may represent further variation rather than biologically separate species. Behavioural and molecular work is urgently needed to compare both syn- and allopatric populations from the entire species range.

ANELOSIMUS PANTANAL SP. NOV.

( FIGS 49G–J , 52–54 , 63C )

Types: Male holotype and female paratype from Brazil, Pantanal, Mato Grosso do Sul, Cuiaba, Poconé ,

Rodovia Transpantaneira (hasta Pousada PIXAIM), Fazenda Araras, approximately 17°37′S, 57°28′W, 5.ix.1996, G. Hormiga and J. Coddington, deposited in MNR [IA40619] .

Etymology: The species epithet is a noun in apposition, referring to the type locality, the wonderful swamplands of Pantanal, Brazil.

Diagnosis: The males can be easily separated from other species of the studiosus group by the relatively small Eb and absence of a distal E fork ( Fig. 49H ). Females are difficult to separate from others of the studiosus group, but differ in having a small, weakly procurved, epigynal plate, the ectal margins barely extending beyond the ectal margin of the spermathecae ( Fig. 49J ).

Male (holotype): Total length 2.47. Prosoma 1.17 long, 0.87 wide, 0.66 high, yellowish, darker in centre and around rim. Sternum 0.71 long, 0.64 wide, extending between coxae IV, yellowish, darker around rim. Abdomen 1.30 long, 1.09 wide, 1.16 high. Pattern as in A. studiosus . AME slightly the largest, other eyes subequal, about 0.08 in diameter. Clypeus height about 2.1 times AME diameter. Chelicerae with one large and two small prolateral teeth, 4–5 denticles retrolaterally. Leg I femur 1.85, patella 0.42, tibia 1.69, metatarsus 1.53, tarsus 0.72. Femur about 8 times longer than wide, metatarsus I about 16 times longer than wide. Leg formula 1243. Leg base colour yellowish with distal tips of all segments slightly darker. Tarsal organs central distal (0.50) on tarsi I and II, proximal on III (0.35–0.40) and IV (0.40–0.45). Four to five small trichobothria dorsally on all tibia, five on tibia I, four on tibia III. Trichobothria on metatarsi I–III proximal (about 0.35–0.45), absent on metatarsus IV. Two prolateral and one retrolateral trichobothria on palpal tibia. Palp as in Figures 49G, H , 52A–F .

Female (paratype): Total length 3.58. Prosoma 1.37 long, 1.04 wide, 0.91 high, yellowish, darker in centre and around rim. Sternum 0.87 long, 0.78 wide, extending between coxae IV, yellowish, darker around rim. Abdomen 2.93 long, 1.73 wide, 1.82 high. Pattern as in A. studiosus . Eyes subequal, about 0.07 in diameter. Clypeus height about 2.1 times AME diameter. Chelicerae with one large and two small prolateral teeth, 4–5 denticles retrolaterally. Leg I femur 1.98, patella 0.52, tibia 1.72, metatarsus 1.63, tarsus 0.68. Femur about 7 times longer than wide, metatarsus I about 16 times longer than wide. Leg formula 1423. Leg base colour yellowish, distal tip of femora, patella, tibia and metatarsi slightly darkened. Tarsal organs distal (0.55) on tarsus I, slightly proximal (0.40–0.45) on II–IV. Five to seven small trichobothria dorsally on all tibia, 5 on tibia I and III. Trichobothria on metatarsi I–III proximal (about 0.40), absent on metatarsus IV. Three dorsal trichobothria on palpal tibia. Epigynum as in Figures 49I, J , 53A–C .

Variation: Male total length 2.35–2.55, prosoma 1.10–1.20, first femur 1.80–1.90, female total length 3.50–3.65, prosoma 1.35–1.45, first femur 1.95–2.10.

Additional material examined: BRAZIL. Mato Grosso do Sul, Pantanal, Poconé, Rodovia Transpantaneira (hasta Pousada PIXAIM), Fazenda Araras , [17°37′0″S, 57°28′0″W], 5.vii.1996, c. 150 m (G. Hormiga, MNR), 8♀ [IA40530]; Porto Cercado [17°25′0″S, 52°27′0″W], 2.viii.1992 (A. A. Lise & A. Braoul Jr, MCP), 1♀ [IA40612]; Pantanal [17°38′0″S, 57°29′0″W], 4– 10.viii.1992 (A. A. Lise & A. Braoul Jr, MCP), 1♀ [IA40614]; Fazienda Sta. Inés, Poconé [16°16′0″S, 56°37′0″W], 4–10.viii.1992 (A. A. Lise & A. Braoul Jr, MCP), 2♀ [IA40616] .

Distribution: Brazil, in lowland swamps of Mato Grosso ( Fig. 63C ).

Natural history: Field notes of J. Coddington (pers. comm.) indicate that A. pantanal is a typical subsocial species.

ANELOSIMUS FRATERNUS BRYANT, 1948

( FIG 49K–N , 63C )

Types: Male holotype from Haiti, Ennery [19°28′0″N, 72°29′0″W], 10.ix.1934, Darlington, in MCZ, examined [IA40768]. The specimen lacks the first pair of legs, except the femur on the left side, leg III is also missing from the left side .

Synonymies:

Anelosimus fraternus Bryant, 1948: 381–382 , figs 55, 57, ♂.

Anelosimus studiosus: Levi, 1956: 419 (in part) not Theridion studiosum Hentz , synonymy here rejected.

Etymology: Bryant did not explain the etymology; one may speculate that the species epithet refers to web sharing (fraternus = brotherhood), but the biology of this species is unknown.

Diagnosis: Anelosimus fraternus differs from most species of the studiosus group by having the embolus tip entire (lacking E-fork, Fig. 49L ). It differs from A. pantanal by a larger Eb, from fork-less specimens of A. guacamayos by a smaller, medially constricted Eb, and from both by having a more extensive row of abdominal stridulatory picks (about 14–16 on each side).

Male (holotype): Total length 2.28. Prosoma 1.17 long, 0.83 wide, 0.66 high, pale brown, darker in centre and around rim. Sternum 0.68 long, 0.59 wide, extending between coxae IV, pale brown with dusky markings. Abdomen 1.37 long, 1.07 wide, 1.07 high. Pattern as in A. studiosus . Eyes subequal, about 0.08 in diameter. Clypeus height about 2.3 times AME diameter. Chelicerae with one large and two small prolateral teeth, 4–5 denticles retrolaterally. Leg I femur 1.50, the type specimen is lacking other segments of legs I. When described ( Bryant 1948: 381) the legs were seemingly intact and she described the first pair of legs as ‘very long’. Femur about 7 times longer than wide. Leg formula 1423. Leg base colour yellowish with, femora slightly darker than other segments. Tarsal organs proximal (0.35–45) on tarsi II–IV. Three to five small trichobothria dorsally on tibia, 4 on tibia III. Trichobothria on metatarsi II–III proximal (about 0.40– 0.45), absent on metatarsus IV. Two prolateral and one retrolateral trichobothria on palpal tibia. Palp as in Figure 49K, L .

Female (IA40769): Total length 4.36. Cephalothorax 1.82 long, 1.22 wide, 1.11 high, brown, darker in centre and around rim. Sternum 0.99 long, 0.86 wide, extending between coxae IV, pale brown with dusky markings. Abdomen 2.73 long, 2.43wide, 2.56 high. Pattern as in A. studiosus . Eyes subequal in size about 0.08 in diameter. Clypeus height about 2.2 times AME diameter. Chelicerae with one large and two small prolateral teeth, 4–5 denticles retrolaterally. Leg I femur 1.95, patella 0.62, tibia 10.59, metatarsus 1.46, tarsus 0.68. Femur about 6 times longer than wide, metatarsus I about 9 times longer than wide. Leg formula 1243. Leg base colour yellowish with femora slightly darker than other segments. Tarsal organs distal (0.55–60) on tarsus I, proximal on tarsi II (0.45–50), and III–IV (0.40–0.45). Five small trichobothria dorsally on all tibia. Trichobothria on metatarsi I–III proximal (about 0.40–0.45), absent on metatarsus IV. Three dorsal trichobothria on palpal tibia. Epigynum as in Figure 49M, N .

Variation: Male only known from holotype. Female total length 4.30–4.50, prosoma 1.80–1.85, first femur 1.90–2.00.

Taxonomic note: Females collected on Hispaniola ( Isla Española, Haiti plus Dominican Republic) differ slightly from typical A. studiosus , and are here considered conspecific with the holotype male from Haiti. Due to the variability of A. studiosus , the limited specimen availability of A. fraternus , and absence of other than geographical evidence matching sexes, the validity and circumscription of this species certainly needs further scrutiny.

Additional material examined: HAITI. Ouest, Portau-Prince [18°32′0″N, 72°20′0″W] ( MCZ), 1♀ [cf. IA40932]; Carrefour [19°24′0″N, 72°4′0″W], 23.vii.1955 (A. F. Archer, AMNH), 1♀ [cf. IA40576] . DOMINICAN REPUBLIC. Sánchez Ramírez, Mina Pueblo Viejo, nr .

Hatillo [18°56′0″N, 70°15′0″W], 21.iii. 1984, 100 m (H. & L. Levi, MCZ), 1♀ [cf. IA40769] .

Distribution: Haiti and Dominican Republic ( Fig. 63C ), between c. 100 and 400 m.

Natural history: Unknown.

Figures

Figure 35. The ‘studiosus group’. A–F, Anelosimus tosum. A, male palp mesal; B, same ventral; C, epigynum; D, internal epigynum dorsal; E, female abdomen dorsal; F, same ectal. G–J, Anelosimus oritoyacu. G, male palp mesal; H, same ventral; I, epigynum; J, internal epigynum dorsal. K–Q, Anelosimus tungurahua. K, male palp mesal; L, same ventral; M, male habitus ectal; N, same, dorsal; O, female abdomen ectal; P, epigynum; Q, internal epigynum dorsal.

Figure 36. Anelosimus tosum, male palp. A, mesal view; B, submesal; C, ventral; D, ectal; E, apicoventral; F, details of embolus tip and conductor. Scale bars: 100 µm.

Figure 37. Anelosimus tosum. A, epigynum; B, epiandrous gland spigots; C, female prosoma dorsal; D, female stridulatory pick row; E, egg sac fibres; F, higher magnification of egg sac fibres. Scale bars: A,C–E, 100 µm; B,F, 10 µm.

Figure 38. Anelosimus tosum. A, female palpal claw. B–G, male. B, fourth tarsal claws; C, stridulatory pick row; D, anterior lateral spinneret; E, posterior median spinnerets, anterior view; F, cheliceral promarginal teeth; G, first tarsus ventral. Scale bars: A,C,F, 50 µm; B, 20 µm; D,E, 10 µm; G, 100 µm.

Figure 39. Anelosimus oritoyacu Ecuador, male palp. A, mesal view; B, submesal; C, ventral; D, ectal; E, details of distal sclerites, ventral; F, mesal view of distal sclerites. Scale bars: 100 µm.

Figure 40. Anelosimus oritoyacu Peru, male palp. A, dorsal view; B, ventral; C, caudoectal; D, apical; E, details of distal sclerites; F, close up of distal sclerites. Scale bars: A–E, 100 µm; F, 50 µm.

Figure 41. Anelosimus oritoyacu. A, epigynum; B, epiandrous gland spigots; C, male stridulatory pick row; D, female abdomen surface; E, female tarsus IV; F, female prosoma dorsal; G, male femur I. Scale bars: A,D,F,G, 100 µm; B, 20 µm; C,E, 50 µm.

Figure 42. Anelosimus tungurahua male palp. A, dorsal view; B, mesal; C, apical; D, ventral; E, ectal; F, details of embolus tip. Scale bars: A–E, 100 µm; F, 10 µm.

Figure 43. Anelosimus tungurahua. A, epigynum; B, epiandrous gland spigots; C, male stridulatory pick row; D, female prosoma dorsal; E, same, front; F, male spinnerets, anterior is left; G, female palpal tibia dorsal; H, male tarsus I claws. Scale bars: A,B,F, 50 µm; C–E, 100 µm; G,H, 20 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 45. Anelosimus elegans, male palp (B–D,F, specimen 1; A,E, specimen 2). A, dorsomesal; B, mesal; C, submesal; D, ventral; E, ventral view, note deeper and broader incision of the distal hematodocha (arrow) as compared with specimen 1; F, ectal. Scale bars: 100 µm.

Figure 46. Anelosimus elegans. A, epigynum; B, epiandrous gland spigots; C, female prosoma dorsal; D, male stridulatory pick row; E, male prosoma and leg I; F, male tarsus I claws. Scale bars: A,C–E, 100 µm; B, 50 µm; F, 10 µm.

Figure 47. Anelosimus guacamayos. A–E male palp. A, mesal; B, apical; C, ventral; D, ectocaudal, arrows indicate the tibial trichobothria; E, details of embolus tip and conductor. F, male stridulatory pick row, right side. Scale bars: A–D, 100 µm; E,F, 20 µm.

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 50. Anelosimus studiosus. A–E, male palp. A, mesal; B, ventral; C, ectal; D, details of embolus tip; E, tibia, note three trichobothria (arrows). F, male stridulatory pick row; G, epiandrous gland spigots; H, epigynum. Scale bars: A–C, 100 µm; E,H, 50 µm; D,F,G, 20 µm.

Figure 51. Anelosimus studiosus female. A, spinnerets; B, anterior lateral spinneret, anterior is right; C, posterior lateral, and posterior median spinnerets, right side; D, posterior median spinnerets, submesal view; E, colular setae (arrow), note absence of colulus; F, cheliceral teeth posterior view. Scale bars: A,E,F, 100 µm; B–D, 20 µm.

Figure 52. Anelosimus pantanal male palp. A, dorsal, note three tibial trichobothria (arrows); B, apicomesal; C, mesal; D, ventral; E, ventrocaudal; F, ectal. Scale bars: 100 µm.

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 54. Anelosimus pantanal. A, epiandrous gland spigots; B, male booklung cover; C, male abdominal apodeme; D, male stridulatory pick row; E, female stridulatory pick row; F, anterior lateral spinneret; G, posterior lateral spinneret, right side; H, posterior median spinnerets, anterior is right. Scale bars: A,F–H, 10 µm; B,C, 100 µm; D, 50 µm; E, 20 µm.

Figure 60. Character optimization on the preferred phylogeny, with clade numbers in circles at the nodes. Numbers above branches indicate characters changing on that node, filled circles indicate characters with perfect fit to the cladogram. All changes are shown; ambiguous character optimization was generally resolved in favour of parallel losses, over parallel gains of structures (see Methods).

Figure 63. A, Distribution of the Anelosimus lineage; B, distribution of A. domingo, A. dubiosus and A. jabaquara; C, distribution of A. oritoyacu, A. guacamayos, A. fraternus, A. pantanal and A. puravida.

Figure 64. A, distribution of A. analyticus, A. pacificus and A. chickeringi; B, distribution of A. octavius, A. jucundus and A. arizona; C, distribution of A. studiosus and A. eximius; D, distribution of A. baeza; E, distribution of A. tosum and A. elegans.

Figure 66. Anelosimus webs, all from Ecuador. A, A. studiosus; B, A. guacamayos; C, A. eximius; D, A. oritoyacu; E, A. domingo.