A revision of the New World eximius lineage of Anelosimus (Araneae, Theridiidae) and a phylogenetic analysis using worldwide exemplars
Author
Agnarsson, Ingi
text
Zoological Journal of the Linnean Society
2006
2006-04-30
146
4
453
593
https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/j.1096-3642.2006.00213.x
journal article
10.1111/j.1096-3642.2006.00213.x
0024-4082
5426885
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.