Daddy-long-leg giants: revision of the spider genus Artema Walckenaer, 1837 (Araneae, Pholcidae)
Author
Aharon, Shlomi
FB5EEA73-DD16-440C-B611-07F1C0C1300B
Blaustein Institutes for Desert Research, Ben-Gurion University of the Negev, Sede Boqer Campus, Midreshet Ben-Gurion, Israel. Alexander Koenig Research Museum of Zoology, Adenauerallee 160, 53113 Bonn, Germany. The Arachnid National Natural History Collection, The Hebrew University of Jerusalem, Edmond J. Safra Campus, Givat Ram, Jerusalem, Israel.
shlomi.aharon@gmail.com
Author
Huber, Bernhard A.
33607F65-19BF-4DC9-94FD-4BB88CED455F
Alexander Koenig Research Museum of Zoology, Adenauerallee 160, 53113 Bonn, Germany. & Email: b. huber @ zfmk. de & urn: lsid: zoobank. org: author: 33607 F 65 - 19 BF- 4 DC 9 - 94 FD- 4 BB 88 CED 455 F
b.huber@zfmk.de
Author
Gavish-Regev, Efrat
FC073F19-2202-4C89-8B43-CEA4CC5E2D50
The Arachnid National Natural History Collection, The Hebrew University of Jerusalem, Edmond J. Safra Campus, Givat Ram, Jerusalem, Israel. & urn: lsid: zoobank. org: author: FC 073 F 19 - 2202 - 4 C 89 - 8 B 43 - CEA 4 CC 5 E 2 D 50 & Corresponding author: efrat. gavish-regev @ mail. huji. ac. il
efrat.gavish-regev@mail.huji.ac.il
text
European Journal of Taxonomy
2017
2017-12-08
376
376
1
57
journal article
21948
10.5852/ejt.2017.376
4298f5cf-d85e-48fb-afe8-f12c99ad65b3
2118-9773
3838544
F67AE00B-28CF-48AF-89D6-31251B22BB7C
Artema nephilit
sp. nov.
urn:lsid:zoobank.org:act:
8C0CA94A-B5CF-4585-B51D-FF25711103E5
Figs 1
,
4–6
,
9–14
,
35–75
,
203
,
210
“
Artema mauriciana
” (misidentification) –
Bodenheimer 1937: 238
(“
Palestina
”); see Notes below.
“
Artema mauricia
” (misidentification) –
Dalmas 1920: 59
(Bodrum,
Turkey
).
Diagnosis
Males can be distinguished from all known congeners by their bulbal processes: process c (
Fig. 40
) projecting prolaterally, processes d and e absent (
Fig. 39
) (
A. magna
: process c robust, strongly curved prolaterally, process d distinct rounded projection on ventral side of bulb – see
Figs 159–160
;
A. doriae
and
A. transcaspica
: process d small, pointed towards ventrodistally) and by unique median projection on each male cheliceral process (
Figs 43–44
,
67
) (only
A. magna
with similar median projection but no modified hairs connect to main ridge as in
A. nephilit
sp. nov.
– see
Figs 163–164
). Females with semicircular epigynum (
Figs 45–50
); differing from
A. atlanta
by straight posterior epigynal margin; from
A. magna
by epigynal plate length to width ratio; from
A. bunkpurugu
by much less prominent anterior epigynal projections (AEP in
Fig. 48
) (cf.
Huber & Kwapong 2013
: figs 49, 53–54).
Etymology
The species epithet is derived from the feminine singular noun of the biblical name “Nephilim”, the giants who were seen by the twelve people sent by Moses to scout the Land of Canaan. It refers to the large size of the spider. Noun in apposition.
Figs 9–14.
Mating sequence of
Artema nephilit
sp. nov.
9
. Male (on the right) approaches female and touches female legs.
10
. Male lowers palps below body for final approach.
11
. Male searches for correct position on female, female abdomen is pushed backwards.
12
. Male rotates palps into final mating position (180° around longitudinal axis; note that the bulb is visible).
13
. Asymmetric insertion of palps (note that the male’s left palp is more deeply inserted than the right palp).
14
. Male cleans palps. Photos BAH.
Figs 15–24.
Artema atlanta
.
15
. Epigynum (MRAC 215041, Benin).
16–18
. Male left pedipalp (MRAC 120766, Sudan), prolateral, dorsal, and retrolateral views.
19–21
. Male left bulb and procursus, prolateral, dorsal, and retrolateral views, arrows in Fig. 20 show notches on distal margin of procursus.
22–24
. Male chelicerae (same specimen), lateral and frontal views. Scale lines: 15, 19–24 = 0.5 mm; 16–18 = 1 mm.
Material examined
Holotype
ISRAEL
:
♂
,
Northern District
,
Jordan Rift Valley
,
Mount Berniki
,
south Teverya
(
Tiberias
) (
32.777° N
,
35.541° E
),
in cave
, -
120 m
b.s.l.
,
16 Sep. 2013
,
B.A. Huber
,
S. Aharon
and
E. Gavish-Regev
leg. (
HUJ
Ara 16000).
Other material
ISRAEL
:
3 ♀♀
,
1 juv.
together with
holotype
, same data (
HUJ
Ara
15996–99);
1 ♂
,
1 ♀
, same data but deep in cave (
ZFMK
Ar
15218–19);
1 ♀
(
HUJ
Ara
16001),
1 juv.
(
HUJ
Ara
16002), same data;
1 ♀
,
HaTanur Waterfall
, near
Metula
(
33.269° N
,
35.581° E
),
15 Mar. 2011
,
L. Friedman
leg. (
CSZ
);
2 ♂♂
,
1 ♀
,
Oren Cave
,
Mount Karmel
(
32.714° N
,
34.975° E
), in cave,
70 m
a.s.l., about
3 km
from
Mediterranean Sea
,
17 Sep. 2013
,
B.A. Huber
,
E. Gavish-Regev
and
S. Aharon
leg. (
ZFMK
Ar
15225– 26);
1 ♂
,
1 ♀
,
1 juv.
,
En Perat
,
Nahal Perat
(
31.833° N
,
35.303° E
), in caves and a building,
260–300 m
a.s.l.,
15 Sep. 2013
,
B.A. Huber
,
S. Aharon
and
E. Gavish-Regev
(
ZFMK
Ar
15220–21);
1 ♀
,
1 juv.
, in pure ethanol, same data (
ZFMK
Isr
51);
1 ♀
, Ein ad-Duyuk, near Jericho (
31.895° N
,
35.421° E
),
25 May 1983
, P. Amitai leg. (
HUJ
Ara
16003);
8 juvs,
Qumeran Cave
(
31.741° N
,
35.459° E
),
7 Nov. 2012
,
L. Friedman
leg. (
SMNH
);
1 ♀
, in pure ethanol,
Nahal Dawid
,
En Gedi
(
31.470° N
,
35.390° E
), among rocks, -250 to -
300 m
b.s.l.,
10 Sep. 2013
,
B.A. Huber
,
S. Aharon
and
E. Gavish-Regev
leg. (
ZFMK
Isr
58);
1 ♀
, same data (
HUJ
Ara
16004);
1 ♂
,
1 ♀
,
En Gedi
(
31.470° N
,
35.390° E
),
20 Sep. 1975
, B. Feldman leg. (
HUJ
Ara
16005-6);
1 juv.
, same locality,
29 May 1968
, P. Amitai leg. (
HUJ
Ara
16007);
1 ♀
, in pure ethanol, Ma’ale Ya’ir (
31.215° N
,
35.359° E
), under large rocks, -
370 m
b.s.l.,
9 Sep. 2013
,
B.A. Huber
,
S. Aharon
and
E. Gavish-Regev
(
ZFMK
Isr
49);
2 ♀♀
, Nahal Temarim (
31.640° N
,
35.411° E
), under large rocks,
3 Sep. 2013
,
S. Aharon
leg. (
HUJ
Ara
16008-9);
2 ♀♀
,
1 juv.
, Sedom Cave (
31.087° N
,
35.395° E
), -
370 m
b.s.l.,
15 Aug. 1939
, A. Shulov leg. (
HUJ
Ara
16010-12);
1 ♀
, in pure ethanol, same locality,
10 Sep. 2013
,
B.A. Huber
,
S. Aharon
and
E. Gavish-Regev
(
ZFMK
Isr
47);
1 ♂
(
HUJ
Ara
16013),
1 ♀
(
ZFMK
Ar
15222), same data;
2 ♀♀
, same locality,
30 Oct. 1941
(
CSZ
);
3 juvs, Sedom Spring,
15 Aug. 1957
, H. Ginsburg leg. (
HUJ
Ara
16014–16);
3 ♀♀
, Arubotaim Cave (
31.101° N
,
35.390° E
), -
380 m
b.s.l.,
10 Sep. 2013
,
B.A. Huber
,
S. Aharon
and
E. Gavish-Regev
leg. (
ZFMK
Ar
15223);
1 ♀
, 2 juvs, in pure ethanol (
ZFMK
Isr
46),
1 juv.
(
HUJ
Ara
16017), same data;
1 ♀
, Malham Cave (
31.076° N
,
35.397° E
), -
370 m
b.s.l.,
10 Sep. 2013
,
B.A. Huber
,
S. Aharon
and E. Gavish-Regev leg. (
HUJ
Ara
16018);
1 ♀
(
ZFMK
Isr
44),
1 juv.
(HUJ 16038), in pure ethanol, same data;
1 ♀
, NW of ’En Tamar, Nahal Zin (
30.989° N
,
35.346° E
), in crevices, -
340 m
b.s.l.,
9 Sep. 2013
,
S. Aharon
,
E. Gavish-Regev
and
B.A. Huber
leg. (
HUJ
Ara
16019);
1 ♂
, molted to
adult
30 Oct. 2013
, NW of ’En Tamar, Nahal Zin (
30.991° N
,
35.347° E
), in tube near road tunnel, -
340 m
b.s.l.,
9 Sep. 2013
,
B.A. Huber
,
S. Aharon
and
E. Gavish-Regev
leg. (
ZFMK
Ar
15224);
1 ♀
,
Ramat Avedat
, near
Sede Boqer
,
Negev Desert
[~
30.84° N
,
34.77° E
],
Jan. 1987
,
V.
and
B. Roth
leg. (MCZ 34068);
1 ♂
,
1 ♀
, Kibbutz Samar (
29.831° N
,
35.022° E
), dumping ground,
100 m
a.s.l.,
11 Sep. 2013
,
S. Aharon
,
E. Gavish-Regev
and
B.A. Huber
leg. (
HUJ
Ara
16020–21);
1 ♀
, in pure ethanol, same locality,
19 Oct. 2013
, E. Gavish-Regev leg. (
HUJ
Ara
16022);
1 ♂
,
2 ♀♀
, in 80% ethanol, same data, in houses (
HUJ
Ara
16023–25);
1 ♂
,
1 ♀
, Eilat Field School buildings (
29.509° N
,
34.920° E
),
12 Oct. 2013
,
S. Aharon
and E. Gavish-Regev leg. (
HUJ
Ara
16026-27);
2 juvs, in pure ethanol, same data (
HUJ
Ara
16028–29).
Figs 25–29.
Artema atlanta
Walckenaer, 1837
, ZFMK Gui 82 and ZFMK Gui 111.
25–26
,
28
. Male ALS and PMS (arrows: one of the six small cylindrically-shaped spigots wider than others).
27
. Male left bulb, prolateral-distal view.
29
. Small teeth prolaterally on round end of process c. Scale lines: 25 = 0.06 mm; 26, 29 = 0.03 mm; 27 = 0.2 mm; 28 = 0.04 mm.
Figs 30–34.
Artema atlanta
Walckenaer, 1837
, ZFMK Ar 15217 (♀) and ZFMK Gui 82 (♂).
30
. Female stridulatory ridges.
31
. Absence of male stridulatory ridges.
32
. Male chelicerae with frontal row of modified (cone-shaped) hairs.
33
. Epigynal plate.
34
. Modified (cone-shaped) hair on chelicerae process. Scale lines: 30–31 = 0.1 mm; 32 = 0.2 mm; 33 = 0.6 mm; 34 = 0.01 mm.
Figs 35–44.
Artema nephilit
sp. nov.
35
. Male habitus (ZFMK Ar 15225, Israel, Mount Karmel, Oren Cave).
36–38
. Male left pedipalp (ZFMK Ar 15227, Jordan, Petra): prolateral, dorsal, and retrolateral views.
39–41
. Male left bulb and procursus: prolateral, dorsal, and retrolateral views.
42–44
. Male chelicerae (ZFMK Ar 15227, Jordan, Petra and HUJ Ara 16000, Israel, Mount Berniki): lateral and frontal views. Scale lines: 36–38, 42–43 = 1 mm; 39–41, 44 = 0.25 mm.
GREECE
:
1 ♀
,
Athens
,
Acropolis
[
37.97° N
,
23.72° E
], collection
Roewer
, date and collector not given (
SMF
RII
/3724).
TURKEY
:
1 ♂
, 2 penultimate
males
,
Bodrum
[
37.03° N
,
27.42° E
],
16 Sep. 1919
,
R. Varriale
leg. (
MSNG
);
1 ♀
,
Antalya Prov.
,
Kemer District
,
Çıralı
(
36.399° N
,
30.475° E
,
10 m
a.s.l.), in building near beach,
3 Aug. 2016
,
B.A. Huber
leg. (
ZFMK
, collected as
juvenile
, molted to
adult
in
Jul. 2017
, still alive as of
Nov. 2017
).
Figs 45–54.
Artema nephilit
sp. nov.
45–50
. Epigyna, ventral views, females from Berniki caves (45, 49: HUJ Ara 15996-7),’En Tamar (46: HUJ Ara 16019), Samar (47: HUJ Ara 16020), Petra (48: ZFMK Ar 15227) and HaTanur Waterfall (50: CSZ).
51–54
. Female abdomens, dorsal and lateral views, from Petra (51–52: ZFMK Ar 15227), and ’En Tamar (53–54: HUJ Ara 16019). Figs 45-47, 49-50, 53-54 Israel; 48, 51-52 Jordan. Scale lines: 45–50, 54 = 1 mm; 51–53 = 2 mm.
CYPRUS
:
1 ♂
,
Nicosia
[
35.16° N
,
33.36° E
], 1937,
F. Werner
leg. (
NHMW
19.551);
1 ♂
, 2 juvs,
Nicosia
, near
Agios Sozomenos
(
35.059° N
,
33.421° E
),
180 m
a.s.l., in cave,
5 Jul. 2016
,
M. Hadjiconstantis
leg. (
CMH
);
1 ♂
,
2 ♀♀
, same data but
20 Jan. 2016
(
CMH
);
3 ♀♀
,
Nicosia
,
Geri
(
35.102° N
,
33.417° E
),
150 m
a.s.l., in house,
15 Sep. 2016
,
M. Hadjiconstantis
leg. (
CMH
);
1 ♂
, 2 juvs,
Famagusta
,
Lisi
, (
35.106° N
,
33.680° E
),
60 m
a.s.l., in
Panagia Lysis
church,
15 Apr. 2016
,
M. Hadjiconstantis
leg. (
CMH
).
JORDAN
:
Aqaba Province
:
2 ♂♂
, Wadi Rām (
29.7405° N
,
35.4574° E
), among rocks,
830 m
a.s.l.,
12 Sep. 2013
,
B.A. Huber
leg. (
ZFMK
Ar
16161-62), collected as
juveniles
, molted to
adults
in
Dec. 2015
and
Sep. 2016
. –
Ma’an Province
:
2 ♂♂
,
2 ♀♀
, Petra (
30.324° N
,
35.447° E
),
900–950 m
a.s.l., rock-cut tombs, near theater,
13 Sep. 2013
,
B.A. Huber
leg. (
ZFMK
Ar
15227);
1 ♀
,
1 juv.
, in pure ethanol, same data (
ZFMK
Isr
36);
1 ♂
, same data but among rocks (
ZFMK
Ar
15228).
–
Karak Province
:
1 ♀
,
1 juv.
, Wadi Hasa (31.004°–
31.014° N
, 35.494
–35.506
°
E
), -330 to -
250 m
b.s.l., among rocks,
14 Sep. 2013
,
B.A. Huber
leg. (
ZFMK
Ar
15229–30);
1 ♀
, in pure ethanol, same data (
ZFMK
Isr
63). –
Madaba Province
:
1 ♂
,
1 juv.
, Wadi Mujib (
31.465° N
,
35.578° E
), -
380 m
b.s.l., among rocks,
14 Sep. 2013
,
B.A. Huber
leg.
ZFMK
(
Ar
15231).
Material assigned tentatively
(see Notes below)
YEMEN
:
1 ♀
, “
Yar Ju Amlah
” [=
Al
Ja’milah?,
17.11° N
,
43.56° E
], 1950,
M. Borri
and
M. Poggesi
leg. (
MZUF 128
).
UNITED ARAB EMIRATES
:
1 ♀
, Near Ra’s al-Khaimah (
25.46º N
,
55.96º E
), hand-collected,
8 Mar. 2005
, A. van Harten leg. (
ZFMK
Ar
15232).
Description
Male
(
holotype
)
MEASUREMENTS. Total body length 7.0, carapace width 3.1. Leg 1: 48.4 (11.9 + 1.4 + 13.9 + 18.3 + 2.9), tibia 2: 10.2, tibia 3: 8.0, tibia 4: 10.9; tibia 1 L/d: 35. Distance PME–PME 190 μm, diameter PME 190 μm, distance PME–ALE 90 μm, distance AME–AME 50 μm, diameter AME 170 μm.
COLOR. Carapace pale to light beige, with dark gray median band. Clypeus with wide light brown band (as in
Figs 4
,
9–14
,
35
). Legs pale yellow to light ochre with dark rings on femora subdistally, patellae + tibiae proximally, and tibiae subdistally, tips of femora and tibiae whitish; sternum pale, with narrow light brown margins. Abdomen light brown with white and dark dots forming stripes from dorsal to lateral side (as in
Figs 51–52
).
BODY. Ocular area slightly elevated. Carapace with median pit and distinctive posterior furrow. Clypeus unmodified. Sternum wider than long (2.3/1.5). Chelicerae as in
Figs 42–44
and as in
Figs 67–69
, with frontal row of ~20 modified (cone-shaped) hairs on each side that splits distally, with outer branch ending at tip of process, inner branch ending at tip of median projection; with indistinct small field of stridulatory ridges laterally (as in
Fig. 70
). Abdomen globose and high; gonopore with four epiandrous spigots.
PALPS. As in
Figs 36–38
. Coxa unmodified. Trochanter with short ventral projection. Femur with short, somewhat pointed retrolateral process proximally; ventral membranous area proximally bordered on both sides by sclerotized ridges; femur-patella hinges close together dorsally. Patella very short. Procursus with proximal dorsal process and weakly developed ventral pocket; with distal dorsal notch on prolateral margin (as in
Fig. 40
). Bulb with two processes close to membranous embolus (processes a and b, as in
Fig. 40
), process c (as in
Fig. 40
) strongly projecting prolaterally, and indistinct projection on ventral side of bulb (as in
Figs 39
,
64–65
). Area between processes b and c modified with granular surface (as in
Fig. 66
).
LEGS.Without spines;with long curved hairs, especially on tibiae and metatarsi; retrolateral trichobothrium on tibia 1 at 8%; prolateral trichobothrium present on all tibiae; pseudosegmentation not visible.
Male
(variation)
Tibia
1 in
14 other males: 9.2–20.6 (mean 14.4). Color pattern on abdomen varies from distinctive stripes consisting of black and white dots on pale to brown-ochre background to pale without any pattern (cf. females,
Figs 52–53
); leg color varies from light brown to ochre; ocular area usually light brown to ochre; median band on carapace sometimes reaches posterior margin of ocular area, forming short
Figs 55–66.
Artema nephilit
sp. nov.
, ZFMK Ar 15225-27 (♂) and HUJ Ara 15997 (♀).
55–56
. Combhairs on left male tarsus 4, prolateral views.
57
. Right male tarsus 2, showing absence of comb-hairs.
58–59
. Comb-hairs on left female tarsus 4, prolateral (-distal) views.
60
. Right female tarsus 2, showing absence of comb-hairs.
61
. Detail of male comb-hair on tarsus 4.
62
. Trichobothrium base, female palpal tibia.
63
. Male gonopore and epiandrous spigots.
64–65
. Male bulbal processes.
66
. Modified area between processes b and c on male genital bulb. Scale lines: 55, 58 = 0.08 mm; 56–57, 59, 63 = 0.1 mm; 60, 64–65 = 0.2 mm; 61–62 = 0.01 mm; 66 = 0.03 mm.
Figs 67–75.
Artema nephilit
sp. nov.
, ZFMK Ar 15225-27 (♂) and HUJ Ara 15997 (♀).
67–69
. Male chelicerae with frontal row of modified (cone-shaped) hairs that split distally.
70
. Male stridulatory ridges.
71
. Female stridulatory ridges.
72
. Epigynal plate.
73–74
. Female ALS (arrows: one of the six small piriform spigots wider than others).
75
. Female PMS. Scale lines: 67 = 0.2 mm; 68, 75 = 0.02 mm; 69 = 0.01 mm; 70 = 0.08 mm; 71 = 0.1 mm; 72 = 0.6 mm; 73–74 = 0.04 mm.
v-shaped mark. Cheliceral modified hairs vary slightly in numbers. Gonopore with variable number of epiandrous spigots (4–6) (
Fig. 63
).
Female
In general similar to male; tibia
1 in
26 females
: 8.0–16.5 (mean 12.5); several females with light brown carapace margins and subtle brown stripes on ocular area. Stridulatory files laterally on chelicerae more distinct than in males (
Fig. 71
). Epigynum almost twice as wide as long, semi-circular, posterior margin almost straight, gently undulating; epigynal plate consisting of two sclerotized lateral areas that appear swollen posteriorly, pale median area, and dark median sclerite anteriorly (variably large and variably distinct) (
Figs 45–50
); pair of projections anterior to epigynum (AEP) visible (
Fig. 48
) but not prominent as in
A. bunkpurugu
(cf.
Huber & Kwapong 2013
: figs 49, 53–54).
Natural history
The spiders were found both in natural habitats such as caves, deep crevices, and under boulders, as well as in man-made habitats such as buildings, under low culverts, and artificial caves. In caves, the spiders built irregular webs (not clearly dome-shaped webs) near the ground, under stones, in crevices, and on the cave ceiling. The spiders seemed to avoid the deeper, dark parts of the visited caves. In a large cave survey we conducted in
Israel
(
Aharon 2016
), we found
A. nephilit
sp. nov.
in caves especially along the
Jordan
Rift Valley, in a very hot and arid zone with low mean annual precipitation (
50–150 mm
). Only a few specimens were found outside the
Jordan
Rift Valley, in caves in areas with higher mean annual precipitation (
400–650 mm
) in a more mesic habitat. These findings suggest that the species is more abundant in xeric, hot and dry caves rather than in humid caves nearby. The spiders were always found on the east or south slopes that are dryer than the west and north slopes and thus seem to be adapted to arid conditions (
Aharon 2016
).
We examined more than 40 caves in
Israel
; 19 caves were inhabited by
A. nephilit
sp. nov.
, and most of these caves were dry. Some specimens were also found in hot, humid caves, but the species was never found in wet caves. In caves in southern
Israel
, where food abundance was low, we could not observe prey items. In a cave situated in the Golan Heights, in the north of
Israel
, where the annual precipitation is higher, we recorded remnants of isopods, tenebrionid beetles, small scorpions, and a spider. In the south of
Israel
, in a basement of the Eilat Field School, we observed remnants of adult males that had probably been killed by females (that were seen nearby), while in a cave in Nahal Perat, east of
Jerusalem
, we observed the predation of an adult male
A. nephilit
sp. nov.
by
Holocnemus pluchei
(Scopoli, 1763)
. Penultimate instar juveniles collected from caves in southern
Israel
(Arubatayim, Sedom, and Malcham) and reared independently in Bonn,
Germany
and Midreshet Ben-Gurion,
Israel
, molted to adult almost simultaneously after an unusually long period of 9–10 months (
September 2013
to
July 2014
) (
4 ♂♂
and
1 ♀
molted in Midreshet Ben-Gurion;
3 ♂♂
in Bonn). This simultaneous timing suggests that the long period is not an artifact caused by the food regime or differences from natural conditions, but rather reflects an intrinsic growth and maturation period.
A mating sequence was observed on
10 Jan. 2015
(in Bonn,
Germany
). Male from
Israel
, Sedom Cave; collected penultimate in
September 2013
; molted to adult in
July 2014
. Female from
Israel
, Mount Berenice; collected penultimate in
September 2013
; molted to adult in
September 2014
. Previous trials (with various males and females) performed in daylight had resulted in no male reaction when placed into the female’s web. Trials under red light resulted in male courtship, but the females, who had not recently fed, attacked the males. Two days after being fed with crickets, the female above finally accepted the male. Time specification shown as hr:min.
0:00 Male is gently transferred to female web.
0:
12 Male
approaches female, bounces slightly and touches silk lines with front legs; female directs herself towards male, vibrates slightly a few times.
0:
16 Male
touches female legs and approaches further (
Fig. 9
); female remains mostly still.
0:17 Final approach; male lowers palps below body (
Fig. 10
); searches for about 20 sec for correct position on female, apparently by searching for correct alignment of chelicerae relative to epigynum (
Fig. 11
); female abdomen is pushed backwards.
0:18 As soon as chelicerae are correctly placed (locked?) at epigynum, palps are rotated 180° around longitudinal axis (procursus moving outward and forward;
Fig. 12
); insertion of palps, start of copulation; female releases legs from silk, pulls all legs above body and is moved by male to perpendicular final mating position.
0:19–1:02 No major movements, only very slight jerks of male body, usually every 6–8 seconds, rarely with shorter (4 sec) or longer (10 sec) intervals; no rhythmic palpal movements; no female movements; no female stridulation; asymmetric insertion of palps, consistent over entire copulation (left palp more deeply inserted than right palp;
Fig. 13
).
1:02 Sudden separation, apparently initiated by male.
1:02–1:
12 Female
remains motionless; male cleans palps for about 10 min (
Fig. 14
), then also motionless.
1:18 End of observation.
About 24 weeks later, this female produced an egg-sac. After another two weeks (i.e., beginning of
August 2015
), about 40 spiderlings emerged; two weeks later again they molted for the first time. In
July 2017
, i.e., almost two years after eclosion, most remaining specimens (about 15) molted to adults.
Distribution
After the ubiquitous
Holocnemus pluchei
,
A. nephilit
sp. nov.
is the most common pholcid in the
Jordan
Rift Valley. It has been collected from underground shelters and caves from the Eilat Mountains (
Fig. 8
) in the south, to Nahal Ayun in the north. Specimens from outside the Rift Valley (
Israel
: Ramat Avedat near Sede Boqer; Oren Cave (
Fig. 7
), Mount Karmel;
Jordan
: Petra), as well as from
Cyprus
,
Greece
, and
Turkey
, indicate that the species is probably widely distributed in the area. Two females with very similar epigynum were collected from
Yemen
and the
United Arab Emirates
(
Fig. 1
).
Notes
We have not seen the material identified by
Bodenheimer (1937)
as
A. mauriciana
. However,
A. mauriciana
(=
A. atlanta
) does not seem to occur in
Israel
and there is no evidence that it ever existed there. We consider it likely that Bodenheimer was dealing with the common
A. nephilit
sp. nov.
rather than with the much rarer (i.e., in
Israel
)
A. doriae
.
The females from
Yemen
(“Yar Ju Amlah” = Al Ja’milah?) and from the
United Arab Emirates
have a very similar epigynum, but since no males from these localities are known, they are assigned tentatively.