Table Of Content2006. The Journal of Arachnology 34:176-185
COPULATORY BEHAVIOR AND WEB OF
INDICOBLEMMA LANNAIANUM FROM THAILAND
(ARACHNIDA, ARANEAE, TETRABLEMMIDAE)
Matthias Burger: Natural History Museum, Department of Invertebrates, Bernastrasse
15, CH“3005 Bern, Switzerland. E-mail: burgermatthias@hotmaiLcom
Alain Jacob: University of Bern, Department of Conservation Biology, Baltzerstrasse
6, CH-3012 Bern, Switzerland and Natural History Museum, Department of
Invertebrates, Bernastrasse 15, CH-3005 Bern, Switzerland.
Christian Kropf: Natural History Museum, Department of Invertebrates, Bernastrasse
15, CH-3005 Bern, Switzerland.
ABSTRACT. The present study reports for the first time on the behavior prior to, during and after the
copulation of a member of the haplogyne spider family, Tetrablemmidae and describes the web of this
species. Prior to copulation, male and female of Indieoblemma lannaianum from Thailand sometimes
avoided each other or the female scared the male away, apparently by vigorous vibrations of her body.
When first copulations were initiated, they lasted from 1.21 to 3.8 h with an average of 2.25 ± 0.71 h {n
— 17). Some females accepted a second male for mating 3-9 days after first mating. There was no
significant difference between the duration of first and second copulations but significantly more trials
were needed to induce the second copulations. In the copulatory position, the male was inverted and faced
in the same direction as the female. He seized the female’s opisthosoma with apophyses on his chelicerae
which fit into grooves on a female’s ventral plate in this way building a locking mechanism during
copulation. The pedipalps were inserted alternately. The web of I. lannaianum consisted of a longish
narrow sheet, which was made ofmany short threads forming a zigzag pattern and additional long oblique
threads overdrawing the sheet and functioning as signal threads.
Keywords: Haplogynae, copulation, locking mechanism, courtship
Spiders show an impressive array of vari- studies on the copulatory behavior of many
ous copulatory patterns and positions (e. g., haplogynes are still missing. This is especially
Gerhardt 1933; von Helversen 1976; Foelix true for members of the family Tetrablemmi-
1996; Huber & Eberhard 1997). Although dae as their behavior has not yet been ob-
there have been numerous studies on the mat- served in detail.
ing of a variety of spiders, the copulatory be- Tetrablemmids are armored spiders which
havior ofmany species still remains unknown. mainly live as soil-dwellers in the litterhabitat
Bristowe and Gerhardt described the mating of tropical rain forests (Shear 1978; Lehtinen
behavior of several entelegyne species, and of 1981; Burger 2005). They show a character-
haplogynes including members ofthe families istic pattern of abdominal sclerotization, and
Scytodidae, Pholcidae, Segestriidae, Dysderi- the carapace or the chelicerae of males are
dae and Oonopidae (Bristowe 1929-1931; often strongly modified (Shear 1978; Lehtinen
Gerhardt 1926-1930, 1933). In addition to 1981; Schwendinger 1989, 1994), The family
Gerhardt’s comprehensive work (reviewed by Tetrablemmidae is systematically placed as
Huber 1998a), the most detailed descriptions sister group of the Dysderoidea (Coddington
&
ofreproductive biology and copulatory mech- Levi 1991; Platnick et al. 1991).
anisms in haplogynes are given for the family The diminutive size (body length less than
Pholcidae (Uhl 1993a, 1993b, 1998; Uhl et al. 2 mm) ofmost tetrablemmids and the factthat
1995; Huber 1994, 1995, 1997, 1998b, 2002; many species are hard to find make behavioral
Huber & Eberhard 1997; Yoward 1998; Seng- observations difficult. In the present study we
let 2001; Schafer & Uhl 2002). However, investigate the mating behavior of a tetra-
176
BURGER ET AL.—COPULATION AND WEB OF INDICOBLEMMA LANNAIANUM 177
blemmid spider and compare the copulation day). When a female copulated for the second
duration of first and second copulations. We time no more males were offered. If a female
also describe avoidance behaviors of males did not accept a second male within 14 days,
and females, female aggressive behavior prior the procedure was stopped and the female was
to copulation, and we provide details of the considered as single mated.
web of this species. Only the six females that had mated twice
METHODS were included in the statistical comparison of
first and second copulations. To compare first
Specimens ofthe tetrablemmidIndicoblem- and second copulations, two-tailed Wilcoxon
ma lannaianum Burger 2005 were collected in Sign-Rank-tests for paired data were applied.
the primary evergreen hill forest of Doi Suth- We tested for a difference in copulation du-
ep, 1600 m elevation, near Chiang Mai ration and for a difference in the number of
(18°48' N, 98°59' E) in northern Thailand, trials needed until successful first and second
from 11-21 July 2003 by sieving. Types are copulations took place. We used nonparamet-
deposited in the Geneva Natural History Mu- ric statistics because most ofourgroups to test
seum, Switzerland and the Natural History differed significantly from a normal distribu-
Museum of Bern, Switzerland. Indicoblemma tion [duration of second copulations (Shapiro
W
lannaianum inhabits the middle humid leaflit- Wilk W-test; = 0.775621, P < 0.0351);
ter layer alongside little streams. Sixty-four number of trials for the first (W — 0.639893,
females and forty-four males were caught and P < 0.0014) and second copulations (W =
kept singly in little snap cap glass jars (3 cm 0.763674, P < 0.0270)]. Averages are given
diameter and 5 cm height) with ground gyp- ± standard deviation.
sum, which was moistened every day by one
RESULTS
or two drops of water so that the air humidity —
was almost saturated. The spiders were fed Pre-copulatory behavior. When a male
Collembola (Folsomia Candida Willem). All (Figs. 1, 2) was placed in the female’s jar
spiders were mature when collected and thus (Figs. 3, 4), both spiders usually walked
their mating history is unknown. around and appeared to meet each other ac-
For the mating behavior studies, the spiders cidentally. A male who came into contact with
were used one or two days after they were the threads of a female’s web appeared to
collected. Copulations were observed with a commence searching for the female. The fe-
binocular microscope (Wild M3; 6.4X, 16X, male then reacted by turning towards him. No
40X) and partially photographed using a dig- male was ever seen filling his pedipalps with
ital camera (Canon Power Shot G2). All sixty- sperm prior to copulation. Sometimes males
four females and forty-four males were used and females stood immobile at a distance of
mm
for the mating observations. For each pairing, about 3-5 facing each other for 8-10
the male was carefully removed fromhis glass minutes before contacting (Fig. 5). When the
jar with a paint-brush and placed into the fe- spiders met frontally, the male either grasped
male’s jar. The spiders were left together for the female directly and took the copulatory
30-60 min. Ifno copulation was initiated dur- position (Figs. 6-10) or he pushed her back
ing that time, the male was put back in his and upwards with his front legs. In the latter
own glassjar and another male was offered to case both of them palpated each other with
the female after a recovery period of 30 min. their front legs for a few seconds before the
The first palpal insertion by the male was tak- male took the copulatory position by going
en as the beginning of copulation. The end of backwards along the ventrum of the female
copulation was defined as the moment when (Fig. 11). When the male approached the fe-
the spiders physically separated. Post-copula- male from the side or from behind hejumped
tory behavior was observed for 15 minutes in at her and gripped her back with his chelic-
each case. erae. If the female kept running around, the
Eleven females which had copulated once male grasped her opisthosoma with his legs
were given the opportunity to mate a second (Fig. 12). Before mating, the male crept under
time. One day after a female had copulated in her from the side and took the copulatory po-
the lab for the first time, different males were sition (Figs. 6-10). —
offered to her one after another (1-3 males per Locking mechanism. When the male was
178 THE JOURNAL OF ARACHNOLOGY
—
Figures 1-8. Indicoblemma lannaianum, male and female. 1. Male, dorsal view; 2. Male in web,
anterior dorsal view, arrow pointing to signal thread; 3, 4. Female, dorsal view; 5. Male (left side) facing
the female prior to copulation; 6. Male (left side) and female in copula, arrow pointing to embolus of
male pedipalp; 7. Male (left side) and female in copula; 8. Male (right side) and female in copula.
BURGER ET AL.—COPULATION AND WEB OF INDICOBLEMMA LANNAIANUM 179
—
Figures 9—12. Indicoblemma lannaianum prior and during copulation, 9. Male (left side) and female
in copula; 10. Male (prosoma on the left side, ventral view) grasping the female (ventral plates in optical
cut, dorsal view) with apophyses on his chelicerae during copulation (asterisks); 11. Male (left side) and
female palpating each other with the front legs prior to copulation before the male moves into the copu-
latory position (arrow); 12, Male (left side) grasping the female’s opisthosoma prior to copulation before
creeping under her (arrow) and taking the copulatory position. Scale bars = 0.5 mm (9, 11, 12), 0.2 mm
10
( ).
180 THE JOURNAL OF ARACHNOLOGY
4.0
r c 18- •firstcopulation
3.5 • i 16- osecond copulation
Q.
ICQ 3.0' • 8 12-
10
2.5' • C0
• 8 8-
S3 2.0H t 6
Q. 4H
8 1.5' o o
13 2 o 15
— .
1.0 firstcopulations 0 '13 '^ 4I 5I 6I
female-no.
—
Figures 13-17. Copulation durations, trials for successful copulations and web ofIndicohlemma Ian-
naianum. 13. Copulation duration offirst copulations (n = 17); 14. Copulation duration offirstand second
copulations (n = 6; numbers beneath box-plots indicate female-no. corresponding to Fig. 15); 15. Number
of trials needed for each female to induce successful first and second copulation; 16, 17. Web. Abbrevi-
ations: sh = horizontal sheet, th = oblique signal threads.
in the copulatory position, he only used male or the male (or both) sometimes turned
apophyses on his chelicerae to grasp grooves around and walked away without copulating,
on a ventral plate ofthe female’s opisthosoma (iii) After having palpated each other with the
(asterisks in Fig. 10). The tips ofhis legs were front legs (once or several times in quick suc-
not used to hold the female’s opisthosoma cession), the spiders sometimes walked away
(Figs. 6-9). In this position the male often from each other. —
bent and stretched his legs in quick succes- Female aggressive behavior, We some-
sion. — times observed that when a male approached
Avoidance behavior. We observed three a female (or had faced her for some time or
types of behavior that appeared to be avoid- had already come into contact with her) she
ance behavior: (i) Sometimes, when put to- seemed to scare him away by vigorous vibra-
gether, the two spiders ran quickly in different tions of her body, especially the front legs.
directions, (ii) After facing each other and Such apparent female aggressive behaviorwas
staying motionless for several minutes, the fe- observed 18 times.
— —
BURGER ET AL.—COPULATION AND WEB OF INDICOBLEMMA LANNAIANUM 181
—
Copulatory behavior. Seventeen out of pated each other with the front legs for a few
64 females accepted a male for copulation seconds before they separated again (see
(first copulations). Forty-one trials were nec- “avoidance behavior” above). Sometimes the
essary to induce successful first copulations. female scared away the male (see “female ag-
Those first copulations lasted from 1.21 to 3,8 gressive behavior” a—bove).
h (median = 2.06; lower quartile =1,7; upper Uncertain cases. In some pairings, after
quartile = 2.74; Fig. 13). Eleven of the 17 the male had seized the female’s opisthosoma
females that had copulated once were tested with his chelicerae (or had already taken the
to see ifthey would accept a second mate. Six copulatory position), the female ran until the
ofthem accepted a second male forcopulation male loosened his hold and let her go. Some-
3-9 days after first copulation (average 4.3 ± times the female pushed her legs against the
2.3). The second copulations lasted from 0.36 male’s legs and she seemed trying to knock
to 5.75 h (median = 1.53; lower quartile = the male off. Such male-female interactions
0.48; upper quartile = 2.89; Fig. 14). There were observed five times and lasted from 1.5-
was no significant difference between the du- 10.62 minutes (average 5.33 ± 3.42). These
=
ration of first and second copulations (T uncertain cases were not counted as copula-
-4.5; df = 5; P < 0.44; Fig. 14). tions and consequently not included for the
For the females that mated twice, eight tri- analyses shown above.
als were necessary to induce successful first Web characterization and function.
copulations whereas 37 trials totally were nec- The web inhabited by /. lannaianum consisted
essary to induce successful second copula- of a longish narrow horizontal sheet which
tions (Fig. 15). The numbers of trials needed was made of many short threads forming a
to induce successful first and second copula- zigzag pattern (sh in Figs. 16, 17). The sheet
tions significantly differed (T = 10.5; df = 5; should be attached along dry leaves and small
P < 0.031; Fig. 15). branches on the ground. Long oblique addi-
In the copulatory position (Figs. 6-10) the tional threads functioning as signal threads
male was inverted and faced in the same di- overlaid the sheet and were partly connected
rection as the female. His legs did not touch with it (th in Figs. 16, 17). The spiders often
the female’s opisthosoma. The male only used stayed in contact with these threads (arrow in
his chelicerae to grasp the female (see “lock- Fig. 2, Fig. 16). They immediately reacted to
ing mechanism” above). After copulation had a prey touching the threads and successfully
begun, all his legs stayed motionless and captured it. No particular retreat for the spider
slightly bent. The male rested on the patellae was observed.
ofhis legs III and IV and on the anterior dor-
DISCUSSION
sal part of his opisthosoma.
During copulation both spiders remained Pre-copulatory behavior. The main rea-
calm and almost motionless. From time to sons for a male to court before copulation are
time the female could take a few steps and probably to identify himself as a mate of the
slightly change her position. The palps were same species, to avoid being mistaken for
inserted alternately (arrow in Fig. 6, Fig. 10). prey or to stimulate the female and convince
Towards the end of copulation, some females her of his quality (Eberhard 1985, 1996; Foe-
started to run around or tried to knock offthe lix 1996; Huber 1997; Huber & Eberhard
male using their legs. In all cases, the male 1997; Bartos 1998). In many haplogyne spi-
sprang away from the female when copulation ders the male courtship behavior prior to cop-
was finished by turning a somers—ault. ulation is restricted to abdominal vibrations or
Post-copulatory behavior. After their simple leg and palp movements (Bristowe
separation, the spiders mostly walked away 1929; Gerhardt 1929; Dabelow 1958; Uhl et
from each other. In some cases, they remained al. 1995; Huber 1994, 1995, 1998b, 2002;
close together and showed intense self-groom- Huber & Eberhard 1997; Bartos 1998; Senglet
ing. The male often ran his pedipalps through 2001). In pholcids, males often keep on mov-
his chelicerae. When the spiders met a second ing their pedipalps during copulation (copu-
time by walking around after copulation, they latory courtship) (Gerhard 1927; Uhl et al.
either both ran quickly in different directions 1995; Huber 1994, 1995, 1998b, 2002; Huber
and kept out of each other’s way, or they pal- & Eberhard 1997; Schafer & Uhl 2002). Other
182 THE JOURNAL OF ARACHNOLOGY
forms of male courtship behavior are tapping shorter copulations because of the simplicity
or jerking the female’s web (Bartos 1998), of their copulatory apparatus. However, stud-
cutting threads of the female’s web (Uhl et al. ies have shown that the copulatory organs of
1995; Bartos 1998; Senglet 2001), or spread- some haplogynes are in fact quite complex
ing the chelicerae (Jackson & Pollard 1982). (e. g., Uhl 2000; Burger et al. 2003), and little
In exceptional cases male pholcids even per- is known about the copulation duration of
form gustatorial courtship (Huber 1997). many haplogynes. A few of the haplogyne
Males of/. lannaianum showed several be- spiders investigated so far copulate longer
haviors that could have some sort of pre-cop- than one hour (Gerhardt 1927-1929, 1933;
ulatory courtship function. Some males pal- Uhl 1993a; Bartos 1998; Senglet 2001; Schaf-
pated the female and pushed herbackwards or er & Uhl 2002). The longest copulation
the male bent and stretched his legs in quick known for a haplogyne spider was longerthan
succession before copulation started. The 5 hours in the oonopid Silhouettella loricatula
grasping of the female’s opisthosoma by che- (Roewer 1942) (sub Dysderina loricata)
liceral apophyses could also have a courtship (Bristowe 1930).
function (Huber 1999). In 5% of the cases, It seems obvious that the function of a pro-
distinctive female aggressive behavior was longed copulation in general cannot be ex-
observed: the male may have been ready to plained by prolonged sperm transfer only
&
copulate, but the female prevented any further (Eberhard 1985; Suter Parkhill 1990), Dur-
interaction by scaring him away. This behav- ing long copulations, apparent risks are ac-
ior is a striking indication for pre-copulatory cepted, which should favor brief mating (e, g.
female choice as those females often mated increased danger from predators or interrup-
:
with other males afterwards. Similar female tion by another male or by the female before
aggressive behavior in haplogynes was ob- sperm transfer is completed; Eberhard 1996). j
served in certain pholcids prior to copulation Copulation duration may correlate with the
(Huber 1994, 1995; Huber & Eberhard 1997; amount of transferred sperm (Engqvist & !
Bartos 1998) or after copulation (Bartos 1998; Sauer 2003) and/or with fertilization success
Senglet 2001). The fact, that significantly (Andres & Rivera 2000) but Schafer & Uhl j
more trials were needed for successful copu- (2002) showed that longer copulations do not
lation if females had already mated (Fig. 15) indicate higher fertilization rates in their
could indicate that females become choosier study. The prolonged copulation duration in /.
with increasing copulation number as sug- lannaianum could be explained by sperm
gested by Schafer & Uhl (2002) for Pholcus competition hypotheses such as sperm prece- i
&
phalangioides Fuesslin, 1775. Nevertheless dence (Suter Parkhill 1990; Elgar 1998)
,
copulations also seemed to take place without and/or by hypotheses of sexual selection by
pronounced pre-copulatory courtship as some- cryptic female choice (the male could initiate ‘
times males just grasped the females force- processes in the female during copulation
fully with their chelicerae and took the cop- which increase his chances of siring her off-
ulatory position directly.— spring; Eberhard 1985, 1996). The prolonged '
Locking mechanism. A cheliceral lock- copulation could also serve as mate guarding.
ing mechanism (by apophyses or modified By guarding the female, a male can restrict
:
cheliceral hairs) during copulation in haplo- access to females from other males (Sillen-
gyes was reported for certain scytodids (Da- Tullberg 1981 Wynn & Vahed 2004) and con-
;
below 1958) and pholcids (Huber 1994, 1995, sequently guard and protect his own trans-
1998b, 2002; Uhl et al. 1995; Senglet 2001). ferred ejaculate (Schofi &. Taborsky 2002).
Lehtinen (1981) proposed that some tetra- These are all hypotheses yet to be tested but
blemmids use apophyses on their chelicerae to it appears that /. lannaianum could be a good
grasp corresponding grooves on a ventral species for testing these ideas.
plate of the female during copulation. The The second copulation offemale no. 4 (Fig.
present study confirms this suggestion for the 14) seemed to last disproportionately long
first time by live observat—ions. when compared to the other second copula-
Copulation duration. Copulation dura- tions, Excluding it from the analysis would
tion is quite variable among different spiders. result in a trend towards a decreased duration i
Elgar (1995) suggested that haplogynes have of the second copulations. Schafer & Uhl I
BURGER ET AL.—COPULATION AND WEB OF INDICOBLEMMA LANNAIANUM 183
(2002) suggested that shorter second copula- script. AJ. thanks the Natural History
tions resulted from a stronger conflict of in- Museum of Bern and the Swiss National Sci-
terest between the sexes over paternity in sec- ence Foundation for financial support.
ond matings.
LITERATURE CITED
Copulatory position.-^-The evolution of
&
the mating positions in spiders was discussed Andres, J.A. A.C. Rivera. 2000. Copulation du-
by von Helversen (1976). The most plesio- ration and fertilization success in adamselfly: an
morphic copulatory position is the one taken example of cryptic female choice? Animal Be-
haviour 59:695-=703.
by most mygalomorphs and haplogyees like
Bartos, M. 1998. Quantitative analyses of male
Oonops (Gerhardt 1930), Segestria or Filis-
courtship behaviour in Pholcus phalangioides
tata (Gerhardt 1928, 1929): the male ap- (Fuesslin, 1775) (Araneae, Pholcidae). In: Seiden
proaches the female frontally, pushes herbody P.A., editor. Proceedings of the 17* European
back and upwards and inserts his pedipalps Colloquium of Arachnology, Edinburgh 1997.
simultaneously or alternately (von Helversen British Arachnological Society, Burnham Beech-
1976). Among different spider groups, a es. p 171-176.
change from the plesiomorphic to a derived Bristowe, W.S. 1929. The mating habits ofspiders,
copulatory position took place coevergeetly: with special reference to the problems surround-
the male rests on the ventral side ofthe female ing sex dimorphism. Proceedings ofthe Zoolog-
and both partners face in the same direction. BriisctaolwSeo,ciWe.tSy.o1f93L0o.ndAonsu1p9p2le9m:e3n0t9a-r3y58n.ote on the
This position is seen in the oonopids SilhoueF matinghabits ofspiders. Proceedings oftheZoo-
tella (Bristowe 1930) and Xestaspis (Gerhardt logical Society of London 1930:395-413.
1933). Bristowe, W.S. 1931. The mating habits ofspiders:
Indieoblemma lannaianum takes the de- a second supplement, with the description of a
rived copulatory position. The behavior be- new thomisid from Krakatau. Proceedings ofthe
fore some copulations could be a remnant of Zoological Society of London 1931:1401-1412.
the one shown by spiders that take the pie- Burger, M. 2005. The spider genus Indieoblemma
siomorphic position: the male pushes the fe- Bourne with description of a new species (Ara-
male back and they palpate each other with neae: Tetrablemmidae). Bulletin of the British
Arachnological Society 13:97-111.
the front legs for a few seconds (Fig. 11) be- Burger, M., W. Nentwig & C. Kropf. 2003. Com-
fore he goes backwards along the veetrum of
plex genital structures indicate cryptic female
the female and takes the derived copulatory choice in a haplogyne spider (Arachnida, Ara-
position (Figs. 6-10). neae, Oonopidae, Gamasomorphinae). Journal of
Web characterization and function,— Morphology 255:80-93.
The web ofL lannaianum resembles the only Coddington, J.A. & H.W. Levi. 1991. Systematics
web known so far of a tetrablemmine spider. and evolution of spiders (Araneae). Annual Re-
In BrignoUella vulgaris Lehtieen 1981, it is a view of Ecology and Systematics 22:565-592.
small dense sheet attached along the surface Dabelow, S. 1958. Zur Biologic der Leimschleu-
of large dry leaves (Lehtieen 1981). Schwee- derspiene Scytodes thoracica (Latreille). Zoolo-
dinger (1989) described the web constructed gische Jahrbiicher Systematik 86:85-126.
Eberhard, W.G. 1985. Sexual selection and animal
by Perania nasuta Schwendinger 1989, a
genitalia. Harvard University Press, Cambridge,
member of the subfamily Pacullinae. It is a Massachusetts.
loose sheetweb in which the spider hangs up- Eberhard, W.G. 1996. Sexual selection by cryptic
side down at eight. female choice. Princeton University Press,
ACKNOWLEDGMENTS Princeton, New Jersey.
Elgar, M.A. 1995. The duration of copulation in
We are most grateful to Dr. Peter Schwee- spiders: comparative patterns. Records of the
dinger from the Geneva Natural History Mu- Western Australian Museum 52:1-11.
seum, who discovered the species and gave us Elgar, M.A. 1998. Sperm competition and sexual
selection in spiders and other arachnids. Pp.
important information about the type locality 307—339. In Birkhead, TR. & A.P. Mpller, ed-
in Thailand. The tap to Thailand was financed itors. Sperm Competition and Sexual Selection.
by a travel grant of the Swiss Academy of London, Academic Press.
Sciences (SAS) which is greatly acknowl- Engqvist, L. & K.P. Sauer. 2003. Determinants of
edged. We sincerely thank Prof. Dr. ClausWe- sperm transfer in the scorpionfly Panorpa cog-
dekind for helpful comments on the manu- nata: male variation, female condition and cop-
184 THE JOURNAL OF ARACHNOLOGY
ulation duration. Journal of Evolutionary Biolo- globosus (Araneae, Pholcidae). CanadianJournal
gy 16:1196-1204. of Zoology 74:905-918.
Foelix, R.F. 1996. Biology of Spiders. Second edi- Jackson, R.R, & S.D. Pollard. 1982. Thebiology of
tion. Oxford University Press, Oxford. Dysdera crocata (Araneae, Dysderidae): Intra-
Gerhardt, U. 1926. Weitere Untersuchungen zur specific interactions. Journal ofZoology London
Biologic der Spinnen. Zeitschrift fiir Morpholo- 198:197-214.
gic und Okologie der Tiere 6:1-77. Lehtinen, P.T. 1981. Spiders ofthe Oriental-Austra-
Gerhardt, U. 1927. Neue biologische Untersuchun- lian region. III. Tetrablemmidae, with aworldre-
gen an einheimischen und auslandischen Spin- vision. Acta Zoologica Fennica 162:1-151.
nen. Zeitschrift fiir Morphologic und Okologie Platnick, N.L, J.A. Coddington, R.R. Forster&C.E.
der Tiere 8:96-186. Griswold. 1991. Spinneret morphology and the
Gerhardt, U. 1928. Biologische Studien an grie- phytogeny of haplogyne spiders (Araneae, Ara-
chischen, corsischen und deutschen Spinnen. neomorphae). American Museum Novitates
ZeitschriftfurMorphologic und Okologie derTi- 3016:1-73.
ere 10:576-675. Schafer, M.A. & G. Uhl. 2002. Determinants ofpa-
Gerhardt, U. 1929. Zur vergleichenden Sexualbiol- ternity success in the spiderPholcusphalangioi-
ogie primitiver Spinnen, insbesondere der Te- des (Pholcidae: Araneae): the role of male and
trapneumonen. Zeitschrift fiir Morphologic und female mating behaviour. Behavioral Ecology
and Sociobiology 51:368-377.
Okologie der Tiere 14:699-764.
Gerhardt, U. 1930. Biologische Untersuchungen an Schofl, G. & M. Taborsky. 2002. Prolongedtandem
sudfranzosischen Spinnen. Zeitschrift fiir Mor- formation in firebugs {Pyrrhocoris apterus)
phologic und Okologie der Tiere 19:184-227. serves mate-guarding. Behavioral Ecology and
Sociobiology 52:426-433.
Gerhardt, U. 1933. Neue Untersuchungen zur Sex-
ualbiologie der Spinnen, insbesondere an Arten Schwendinger, P.J. 1989. On three new armoured
der Mittelmeerlander und der Tropen. Zeitschrift spiders (Araneae: Tetrablemmidae, Pacullinae)
from Indonesia and Thailand. Revue Suisse de
fiir Morphologic und Okologie der Tiere 27:1-
Zoologie 96:571-582.
75.
Huber, B.A. 1994. Genital morphology, copulatory Schwendinger, P.J. 1994. Four new Perania (Ara- ji
neae: Tetrablemmidae, Pacullinae)fromThailand
mechanism and reproductive biology in Psilo- ,
and Malaysia. Revue Suisse de Zoologie 101:
chorus simoni (Borland, 1911) (Pholcidae; Ara- 447-464. |
neae). Netherlands Journal ofZoology44:85-99. >
Senglet, A. 2001. Copulatory mechanisms in Ho-
Huber, B.A. 1995. Copulatory mechanism in Hol-
plopholcus, Stygopholcus (revalidated), Pholcus,
ocnemus pluchei and Pholcus opilionoides, with I
Spermophora and Spermophorides (Araneae,
notes on male cheliceral apophyses and stridu- Pholcidae), with additional faunistic and taxo- j
latory organs in Pholcidae (Araneae). ActaZool- nomic data. Mitteilungen der Schweizerischen '|
ogica Stockholm 76:291-300.
Entomologischen Gesellschaft 74:43-67.
Huber, B.A. 1997. Evidence for gustatorial court-
Shear, W.A. 1978. Taxonomic notes on the armored
ship in a haplogyne spider (Hedypsilusculicinus: spiders of the families Tetrablemmidae and Pa- j
Pholcidae: Araneae). Netherlands Journal ofZo- cullidae. American Museum Novitates 2650:1-
ology 47:95-98.
46.
Huber, B.A. 1998a. Spider reproductive behavior: a Sillen-Tullberg, B. 1981. Prolonged copulation: a i
review ofGerhardt’s workfrom 1911-1933, with male “postcopulatory” strategy in a promiscu- !I
implications for sexual selection. Bulletin ofthe ous species, Lygaeus equestris (Heteroptera: Ly-
British Arachnological Society 11:81-91. gaeidae). Behavioral Ecology and Sociobiology
Huber, B.A. 1998b. Genital mechanics in someneo- 9:283-289. j’
tropical pholcid spiders (Araneae: Pholcidae), Suter, R.B. & V.S. Parkhill 1990. Fitness conse-
with implications for systematics. Journal ofZo- quences ofprolonged copulation in the bowl and j
ology London 244:587-599. doily spider. Behavioral Ecology and Sociobiol- ;
Huber, B.A. 1999. Sexual selection in pholcid spi- ogy 26:369-373.
Ij
ders (Araneae, Pholcidae): artful chelicerae and Uhl, G. 1993a. Mating behaviour and female sperm
forceful genitalia. Journal of Arachnology 27: storage in Pholcusphalangioides (Fuesslin) (Ar- |
135-141. aneae). Memoirs ofthe Queensland Museum 33:
Huber, B.A. 2002. Functional morphology of the 667-674.
genitalia in the spider Spermophora senoculata Uhl, G. 1993b. Sperm storageandrepeatedeggpro-
(Pholcida, Araneae). ZoologischerAnzeiger241: duction in female Pholcus phalangioides Fues-
105-1 16. slin (Araneae). Bulletin de la Societe Neuchate-
Huber, B.A. & W.G. Eberhard. 1997. Courtship, loise des Sciences Naturelles 116:245-252.
copulation, and genital mechanics inPhysocyclus Uhl, G. 1998. Mating behavior in the cellar spider,
j
BURGER ET AL.—COPULATION AND WEB OF INDICOBLEMMA LANNAIANUM 185
Pholcus phalangioides, indicates sperm mixing. Wynn, H. & K. Vahed. 2004. Male Gryllus bima-
Animal Behavior 56:1155-1159. culatus guard females to delay them from mating
Uhl, G. 2000. Two distinctly different sperm stor- with rival males and to obtain repeated copula-
age organs in female Dysdera erythrina (Ara- tions. Journal of Insect Behavior 17:53-66.
neae: Dysderidae). Arthropod Structure & De- Yoward, P.J. 1998. Sperm competition in Pholcus
velopment 29:163-169. phala—ngioides (Fuesslin, 1775) (Araneae, Pholci-
Uhl, G., B.A. Huber & W. Rose. 1995. Male ped- dae) shorter copulations gain higher paternity re-
ipalp morphology and copulatory mechanism in ward than first copulations. In: Selden P.A., editor.
Pholcus phalangioides (Fuesslin, 1775) (Ara- Proceedings of the 17* European Colloquium of
neae, Pholcidae). Bulletin of the British Arach- Arachnology, Edinburgh 1997. BritishArachnolog-
nological Society 10:1-9. ical Society, Burnham Beeches, p 167-170.
von Helversen, O. 1976. Gedanken zur Evolution
der Paarungsstellung bei den Spinnen (Arachni- Manuscript received 27 August 2004, revised 23
da: Araneae). Entomologica germanica 3:13-28. February 2005.
