Rivet-like nest-building and agonistic behaviour of Thiania bhamoensis, an iridescent jumping spider (Araneae: Salticidae) from Singapore

9 Pages·2002·1.4 MB·English

by  Li, D

THE RAFFLES BULLETIN OF ZOOLOGY 200250(1): 143-151
@ National University of Singapore
RIVET-LIKE NEST-BUILDING AND AGONISTIC BEHAVIOUR
OF THIANIA BHAMOENSIS, AN IRIDESCENT JUMPING SPIDER
(ARANEAE: SALTICIDAE) FROM SINGAPORE
Daiqin Li, SeowH wa Yik and Wee Khee Seah
Department of Biological Sciences, National University of Singapore, 10 Kent Ridge Crescent,
Singapore 119260. Email: [email protected]
ABSTRAC7: -This is the first detailed description of nest-building and fighting behaviour of
Thiania bhamoensis Thorell, an iridescent jumping spider species, commonly known as fighting
spiders in South-east Asia. Its nest-spinning behaviour is unusual for a jumping spider. Thiania
bhamoensis built a silken nest by binding a pair of green leaves together with strong silken rivets.
Spiders rested, moulted, and oviposited in the narrow space between the bound leaves. Egg sacs are
usually flattened and densely woven for a salticids. However, the egg sac of 1: bhamoensis was
sealed to a leaf in the rivet-like nest but was not contiguous with the silken components (i.e., rivets)
of the nest. Pairwise fights between males were staged in the laboratory and videotaped. Twelve
major displays were identified, showing a large repertoire of agonistic displays of this species. The
behaviour of 1: bhamoensis is also discussed in relation to hypotheses concerning salticid evolution.
KEY WORDS. -Salticidae, Thiania bhamoensis, display, fighting spiders, rivet-like nest.
INTRODUCTION intraspecific interactions (Crane. 1949; Jackson &
Pollard,1 997). All these,t ogetherw ith their complex
A male's reproductives uccessg enerallyd ependso n its behaviour, make salticids well representativeso f
abilityt o acquireo r maintaina ccessto prospectivem ates. ethologicasl tudies( Richman& Jackson1, 992).
Competitionb etweenm ales for mates is common in
naturea nd conspecificm aleso ften communicated uring Salticidsh avea pair of largea nterior-mediaely es( AME),
the agonistic interactiont hrough threatd isplays.T hese known as the principale yes (Homann,1 928),a nd three
have been extensively studied for decades both pairs of smaller ('secondary') eyes responsible for
theoretically and experimentally. However, actual movementd etection( Land,1 971;1 974). The principal
knowledgea boutt he meaningo f agonisticd isplaysa nd eyesb, eingr esponsiblefo r acutev ision (Land, 1969ab, ),
the fighting decisionr ulesu nderlyingt heir occurrenceis enables alticidsp, rior to contactt,o discriminateb etween
poorly known (Mann et al., 2001). Male agonistic matesa nd rivals (Jackson& Blest, 1982;H arlande t al,
behaviouri s commoni n many specieso f spiders,w ith 1999;H arland& Jackson2 000ab, ). Acutev ision,m ade
best-known examples in jumping spiders from the possible by the salticid's principal eyes, createst he
Salticidae( Faber& Baylis, 1993; Taylor et al., 2000). potentialf or good assessmeonft the opponents(T aylore t
Salticids are one of the few groups of spidersw hose al.,2 000).A mong salticidsa, ll sex/agec lassess ometimes
major mode of locomotion is, as the name implies, performt hreatd isplaya ndr itualizedf ights,b ut this type
jumping. The Salticidaei s the largest (nearly 5000 of behaviouris mostp ronouncedin adultm ales( Crane,
describeds pecies)a nd diverse family (Coddington& 1949;J ackson1, 982).A lthoughs alticidsa rea ssumedto
Levi, 1991; Zabka, 1993), with memberso n every highly develops ucha ggressiveb ehaviourin conspecific
continent (except Antarctica) and on most oceanic males (e.g. Pechham& Peckham,1 889; Crane,1 949;
islands.T heses pidersa reo neo f the majora nimalg roups Forster,1 982; Jackson& Pollard, 1997),o nly a small
in which acutev isionh ase volved( e.g.,L and & Nilsson, fractiono f speciesin this larget ropical family has been
2002). Visual acuity andj umping ability have enabled subjected to detailed studies of contest behaviour.
salticidst o hunt their prey actively,i nsteado f relying on However,i nformation on the intraspecifici nteractions
silken snares. It is not surprising that their highly has been used to speculateth e evolution of salticids
developedv ision also plays an importantr ole in the (Jackson& Blest, 1982).T o evaluatet his, comparative
Received2 8 Nov 2001 Accepted 22 Feb 2002
143
Li et al.: Agonistic behaviouro f fighting spiders
data from a wide rangeo f speciesa re neededT. herea re To observei nteractionst,h e leaf was removedf rom the
73 describeds alticid speciesi n Singapore(D . X. Song, cage and clampedt o a stand and anotherm ale from
J.X. Zhang & D. Li, unpublishedd ata),b ut only three another cage was put on the leaf with the resident
species,E peusf lavobilineatus (Doleschall) (Jackson, male.T he initial displaysd efinedt he beginningo f the
1988), Phaeacius malayensis Wanless (Jackson & interaction; when one spiderd ecampeda nd the other
Hallas,1 986a)a nd Thorelliae nsifera( Thorell) (Jackson failed to follow defined the end, at which point the
& Whitehouse,1 989)h ave beens tudieds o far. spiders were removed. The male-male interactions
were videotaped for further descriptions. The
Thiania is an Oriental salticid genusc onsistingo f 17 descriptionsw ere basedo n 45 male-malei nteractions.
described but poorly known species. Thiania No individual male was usedm ore than once per day
bhamoensisT horell are commonly known as fighting or in the samep airing. All observationsw ere carried
spidersi n South-easAt sia becausem alesa re known to out between0 900 h and 1800h .
engage in fighting behavior both in nature and the
artificial situation (Rashid & Azirun, 1992). It is
known that a spider that wins battle after battle attains FIELD OBSERVATIONS
the statuso f "first king" until its performanced eclines
and is replacedb y anotherc hampion.F ighting spiders Nestsa nd nest-building behaviour
are normally kept in matchboxesw ith a piece of green
leaf and a spato f saliva to provide moisture. Thiania The nest-buildingb ehaviouro f 7: bhamoensiiss similar
bhamoensish as an iridescentg reen-bluec olouration, to that of Thianias p. from Malaysia (Jackson1, 9 86a).
males being more blue and femalesm ore green. It is In the laboratory,m aking a single rivet took c. 20-30
common belief that the darker the male is, the more min andi nvolveda variablen umbero f cycleso f making
aggressive it will be. Apart from a brief article silk attachmendt iscs to the leaf. Each cycle involved
describing male -male interactions in this jumping making three attachmentsI.n a cycle, the spider fIrst
spider( Rashid& Azirun, 1992),n o detaileds tudieso f stood betweent wo leaves,d orsal-side-upf,a cing into
male-male interactions have ever been conducted. the nest.T he spidert hens wepti ts abdomenfr om sidet o
Furthermore,i nformation on male-malei nteractionsi s side 2-5 mm for a few times,l aying strandso f silk on
needed for further investigation of factors that the substratuml,o weredt he abdomena, nd then moved
influence the outcomeso f male-malec ontests.I n the the spinneretsb acka nd forth fasteninga thread.A fter a
presents tudy,t he intraspecific display behaviouro f 7: pause( 0.5-1 s),t he spiderr aisedi ts abdomena nd,a t the
bhamoensis from Singapore is investigated. The samet ime, rotatedc . 1800b, ringing its spinneretso nto
aberrantn est spinning behavioura nd egg sacso f this the upper leaf surface( Fig. 1). The spiderm oved its
salticid are also describedi n this paper. spinnerets back and forth, making the second
attachmendt isc. Finally, the spiderm oved its abdomen
back to the bottom leaf, making the third attachment
MATERIALS AND METHODS disc closet o the first. Eachc ycle took up to 1 min, with
the spider gradually shifting its position for the next
Thiania bhamoensisw ere observed in nature and cycle. Therew as an interval of 0.5-3 min betweent wo
collected from Kent Ridge Park, SingaporeB otanic successivcey cles.A fter completiono f a rivet, the spider
Gardens and the National University of Singapore moved away for a certain distance,a nd either kept
campusin SingaporeT. hirty adultsm alesw erec ollected. inactive for a while or immediately startedb uilding
Spidersw ere individually housedi n plastic cylindrical anotherr ivet elsewhere.
containers( diameter:9 cm, height: 6.5 cm). They were
fed three adult Drosophila melanogastetrh reet imes a In nature, 7: bhamoensisw ere usually found between
week. Greenl eavesw ere provided for spiderst o build a horizontal pair of green leaves in a nest on most
theirn estsL. eavesr emainedg reeni n humidc agesf or up gardent rees and shrubsb ut were especially common
to two weeksa ndw erer eplaceda s theyd ried out. Water on Crinum asiaticum (spider lily) (Fig. 2). This plant
wasp rovidedv ia a cottonr oll thatp enetratedth rought he is native to tropical Asia and is often cultivated in
floor of the cage and was dipped into a water-filled gardens for aesthetic purposes because of their
containero utsidet he cage.C agesw ere cleanedo nce a stalkedw hite flowers. The leavesa re green,l ong and
week to remove any dead flies. Cardboardp artitions sessile with wide blades. Many leaves lay close
shieldedt he spidersf rom one anothers o thatt hey would together and 7: bhamoensisb uilt their nests in the
not have any prior experiencew ith conspecificsb efore spaceb etweent he leaves( Fig. 3). When leavesw ere
testing.L ighting (12:12L : D), temperature(2 5 :t 1°C), pulled apart,7 : bhamoensisa lmosta lwayse scapedb y
and humidity (70-80%)w erec ontrolled.L ightsc ameo n making sudden leaps and subsequentlyr emaining
at 0800 h and wento ff at 2000h . motionless,m aking it difficult to relocate.
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THE RAFFLES BULLETIN OF ZOOLOGY 2002
The nestc onsistedo f a spaceb etweent wo leavesa nd and pull the leavesa part (Fig. 3). Somen estsh ad 1-
the perimeter was about 4-10 silken rivets arranged 3 larger rivets. The large rivets were c. 12 x 5 mm
in a roughly circular or elliptical manner.T he rivets and had looser threads than the small rivets (Fig. 4).
were small or large. Most rivets were small and they The leaf surface within and near the nest was rarely
were c. 1 mm wide x 2-8 mm long. Its long axis was coated with silk threads so that wide, clear spaces
generally facing the interior of the nest. The small betweent he rivets (doors) allowed the spidert o enter
rivets were formed by many closely packed vertical and leave the nest. With many doors the spider was
strands of silk (2-6 mm long) connecting the two able to go in and out of the nest from almost any
leaves. Some effort was needed to break the rivets direction.
Figs. 1-6. (1) Thiania bhamoensiss pinninga rivet. Standingb etweentw o leaves( facing left), it lifts andr otatesa bdomento fasten
line to top leaf (lateral view); (2) Spider lily (Crinum asiaticum),a tropical plant on which Thiania bhamoensisb uild the nests
betweent wo leaves; (3) Thiania bhamoensisa t edge of nestp eering out (front view). Note the silken 'rivets' on both sides of the
spiders.L eavesh ave beenp artially lifted apart; (4) FemaleT hiania bhamoensissi tting aboveh er egg sac (front view). Note the
silken 'rivets' on the both left and right sides of the egg sac; (5) Male Thiania bhamoensiso n leaf in cryptic restp osture.F acing
up. Legs presseda gainstt he surfaceo f leaf; and (6) Male Thiania bhamoensis(d orsal view) walking on leaf.
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Li et al.: Agonistic behaviouro f fighting spiders
Like adultm alesa nd femalesj,u venilesa lso built rivet- that involved postures, which were static, with the
like nests.H owever,a dultsc ommonlys panl argern ests spiderh olding parts of its body in a particular stance
(20 x 30 mm to 30 x 50 rom) while juveniles built for a sustainedp eriod. Each elemento f behaviouri s
relativelys malln ests.I n naturej,u venilesb uilt their nests described below. When referring to the legs of the
betweenth e sames izedl eavesa s adults.N estsw ere not spider,l egs I are the most anterior pair; legs II, the
built in the absenceo f paired leaves.M ost times, only next most anterior pair, etc.
one individual was found in eachn est. Occasionallya,
malew asf ound cohabitingw ith a juvenile female. 1. Raised body. -Raised bodies were held higher
than normal above the substratum (Fig. 7). The
anteriorc ephalothoraxw as often held higher than the
Egg sacs posterior, but the abdomenw as held parallel to the
substratum.
In nature,a ll nestsw ere found containingo ne egg sac
each. The egg sac was c. 2-3 times larger than the 2. Abdomen flexed up or flexed down. -Thiania
spiderb ody length. It was white (opaque)in coloura nd bhamoensis often held its abdomen flexed up
roughly elliptical in shape.T he eggs were wrapped (deflected up c. 45° from the cephalothorax)o r flexed
severall ayers of lower and upper denselyw oven silk down (abdomen deflected down c. 45° from the
sheetso n the surfaceo f the lower leave inside the nest cephalothorax)w hile displaying with its legs I. Very
(Fig. 4). The egg sac was unusually flattened. The often, the cephalothoraxw as slightly raised when the
lower silk sheetw as generally larger than the upper abdomenw as lowered.
sheet.T he lower sheets ometimese xtendeda bout5 -8
mm out from the sideso f the upper sheet,b ut the upper 3. Arched-out legs. -Legs II, III and IV were
sheeta lwaysj oined the lower sheett ightly. stretchedo utwards,r aising up the body (Fig. 7).
4. Arched legs. -Only legs I were arched (Fig. 8).
NORMAL LOCOMOTION They were held roughly perpendiculart o the sides of
the body with the femur angled upward about 30°.
When at rest, 7: bhamoensisa dopted the "flattened The femur-patella and tibia-metatarsusj oints were
posture". When resting in the rivets betweenl eaves, flexed slightly down so that the tarsus contactedt he
the flattened body and legs were presseda gainstt he substratum. Legs were held about 120° apart.
surface of the leaf (Fig. 5), with legs I arched,l egs II Occasionally,l egs I were spreadm ore than 120° (but
angled forward, legs III arched-out and legs IV less than 180°)a part.
angled rearward. The palps angled ventro-mediately
so that they obscuredt he view of cheliceraef rom the 5. Hunched legs. -Only legs I were hunched by
front. being held to the sides of the cephalothoraxw ith the
femur angled upward about 50°. Legs were highly
Thiania bhamoensish ad two modes of locomotion -flexed at the femur-patellaj oints so that all segments
walking and running (Fig. 6). The spider normally distal to that joint pointed down and the tarsi
moved in a rapid stop-and-gog ait typical of style of contactedt he substratum.
locomotion, stepping rapidly a few centimeters,a nd
then standing for 0.5 -1 s, then stepping forward 6. Bent abdomen. -The abdomenw as frequently
again.T he movementw as rather rapid and agile, and twisted at the pedicela nd swivelled (c. 45°) to the left
the spider often alternated between walking and or right side of an individual's cephalothorax( Figs.
running, with walking predominating, although a 9,10,11& 12).
spider almost always ran when disturbed (e.g., if a
persona ttemptedt o pusha part leaves.T he spidera lso 7. Elevatedl egs. -Only legs I were elevated.T he two
often waved its palps, but not so much as it was most commonp ositions were:
standing.T here was little or no waving of legs when Position (1) -Arched legs I (see element 4) were
steppingo r during pausesb etweens teps. raised off the ground. They were held to the side of
the body about 120° or more but less than 180° apart
(Fig. 10).
ELEMENTS OF AGONISTIC BEHAVIOUR Position (2) -Legs I were held perpendiculart o the
side of the body, 180° apart and parallel to the
Interactions between 7: bhamoensism ales consist of substratum. All joints were fully extended, giving
displays that involved movementp atterns and those legs I a stiff appearance(F ig. 12).
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THE RAFFLES BUUETIN OF ZOOLOGY 2002
Fig. 7-14. (7) Thiania bhamoensisin a 'hunch' posture with arched out legs and raised body and cephalothorax( facing right).
Palps in nonnal posture; (8) Male Thiania bhamoensisp osturing with arched legs I. Palps in nonnal posture; (9) Thiania
bhamoensism ale posturing with archedl egs I elevatedi n Position 1. Abdomeni s bentt o the right of the cephalothoraxP. alps in
nonnal posture; (10) lWo Thiania bhamoensism ales sparing with each other, eachp osturing with legs I archeda nd elevatedi n
Position 1 and abdomenf lexed up and bentt o one side; (11)l Wo Thiania bhamoensism alesa pproachinge acho ther,w ith legs I
arched and abdomenf lexed up and bent to one side; (12) Samet wo Thiania bhamoensism ales (as in Fig. 11) initiating an
embrace.E acha pproachingt he other with legs I elevatedi n Position2 . Palps laterally erectinga nd almostt ouching; (13) Front
view of Thiania bhamoensism ale. Palpsh eld in nonnal posture: in front of the cheliceraea, rcheda nd hangingd own; and (14)
Front view of Thiania bhamoensism ale.P alpsi n nonnalp ostureb ut with the femoraa ngleds lightly upwards o thatt he palpsh ide
the cheliceraef rom view (c.f. Fig. 13).
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Li et al.: Agonistic behaviouro f fighting spiders
8. Jerky leg waving. -While stepping (forward or 15. Decamp. -To decamp,o ne spider ran or leapt
more often, to the side), males waved arched or away from the other.
elevatedl egs I (Position 1) by moving them up-then-
down (by a small amplitude) in a suddena nd rapid
motion. There was only one wave for each step the ORGANIZATION OF MALE-MALE
spider took, with legs I remaining arched in between AGONISTIC BEHAVIOUR
steps.
Whens pidersw ere not interactingw ith conspecificsa,
9. Normal palp posture. -The palps were held in male normally postured by standing or walking as it
front of the cephalothorax,a rcheda nd hanging down. held its stationary legs erect, hunched, or arched.
The palp femora extended ventro-laterally alongside Interactionsb egan when both males faced each other
the chelicerae with the rest of each palp angled from as far as 14 cm apart. Generally,t hey adoptedt he
ventro-medially so that the tips of the two tarsi were 'hunch' posturew herel egs I were huncheda nd legs II,
1-2 mm apart under the fangs (Fig. 13). Also, the III and IV were arched out, raising up the body. This
femora were usually angled slightly up such that the appearedto be a defensivea nd alerts tance.B oth males
rest of the palps obscuredt he view of the chelicerae approachedo ne anotherm aintaining this posture or
from the front (Fig. 14). sometimes one male remained stationary at this
posturea nd visually trackedi ts opponent.
10. Erect palp. -Palps were stiffly extendeda nd in
the lateral erect posture where they were held to the Next, sparring began. The legs I of one male was
sides of the chelicerae about 1200 away from each arched and the anterior cephalothoraxl owered from
other (Fig. 12). the 'hunch' position (Fig. 8). The other male usually
matchedt his display.A s they got closert o eacho ther,
11. Embrace. -The spiders approachede ach other legs I were elevatedi n position 2 (Figs. 9 & 10). Both
with legs arched or elevated in Position 1, moving malesu suallyh eld their abdomensjlexedu p/downa nd
them into Position 2 when they got closer. They bent to the sidew hile they sparredw ith eacho ther.T he
continued to advancea nd brought their laterally erect palps were held in their normal position.
palps into contactt o initiate an embrace( Fig. 12).
When both 'spiders approachedt,h ey usually stepped
12. Palpal pushing. -During embrace, with legs forward and backward or sometimest o the side and
extendedi n Position 2, palpal pushing occurred with tendedt o wave their legs I. Within about1 body length
laterally erectp alps. Each palp touchedt he palp of the of eacho ther (Fig. 11),t he legs I of both spidersw ere
facing spider then each spiderm oved or attemptedt o suddenlys traighteneda nd elevatedi n position2 . They
move forward. When pushing,l egs I of both spiders faced eacho ther with legsI in this positiona nd parallel
touched when they fluttered back and forth and to eacho ther (Fig. 12).
appearedt o be a blur. Also, during this period, the
anterior end of the cephalothoraxt ended to tilt up a They moved closert ogetherp, ausedm omentarilyf ace-
little with the bodies of both spidersf orming a 'tent' to-face and embracedt o engage in palp al pushing.
structure. With their bodies in suchc lose proximity, both males
occasionally started to grapple. Eventually, the
13. Grapple. -From embrace,b oth spiders pressed interactione nded when the dominantm ale causedi ts
their faces tightly together and moved their legs opponentt o break away and decamp.
forward over each other so that they interlocked.T he
legs I of one spidera ppearedto wrap aroundt he body
of the other spider in a bid to pin it down to the LEVELS OF CONTEST INTENSITY
substratum.
The following stagesw ere viewed as representinga
14. Step-to-the-sidea nd forward-and-backward. -sequence of increasing levels of intensity within
Spidersf aced but never ran directly into each other agonistic encounters based on consideration of
(charge).I nstead,w hen approachingt,h ey steppedt o proximity of males to each other and risk of injury.
the left or right or steppedf orward or backwarda few Transitionsf rom one staget o the next were viewed as
millimetres. When performing these behaviours,t he escalationsin encounterin tensity.
spideru suallyp ostured( Position 1) or wavedi ts legs I.
While steppingt o the side,t he spider's abdomenw as Levell. -The sparringd istanceb etweenth e two males
often bent away from the direction of stepping. was about four body lengths or more. Usually, one
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THE RAFFLES BULLETIN OF ZOOLOGY 2002
male approachedt he other that remained stationary putativelyp rimitive (Hallas & Jackson,1 986; Jackson
and they sparreda t that distance. & Hallas, 1986a, b). Thiania bhamoensis does
resembleL . viridis and the spartaeinesin making egg
Level2 . -The sparringd istanceb etweenth e two males sacs that are not contiguous with the silk of a nest.
was aboutt wo body lengths or less. In most casest,h is However, unlike L. viridis and the spartaeinest hat
proximity was due to the two malesa pproachinge ach make the more transparente gg sacs,1 : bhamoensis
other. spins the flat, denselyw oven egg sacs,d iffering from
most typical salticid egg sacs.T hesef indings suggest
Level 3. -Both males embracedt o engagei n palpal that the non-contiguousnesosf nest and egg sac silk
pushing. might be a primitive trait andt he typical enclosedn ests
might be an advanced trait of salticids. If this
Level4 . -Males were involved in a grapple with each hypothesisi s correct, the ancestorso f 1: bhamoensis
other. were salticids that madet ypical tubular nestsb ut also
kept the primitive trait in which the nest and egg sac
silk is not contiguous.T he spinning behaviouro f 1:
DISCUSSION bhamoensism ight modify in conjunction with its
becoming specialized at living between leaves
Nestsa nd eggs acs (Jackson, 1986a), representing a transition from
primitive to advanced salticids. However, the
Like the tubular nests spun by typical salticids, the phylogenyo f salticidsi s little known.
opened,r ivet-like nests of Thiania bhamoensism ay
servea s resting sites and as sites for moulting, mating Interestingly, like 1: bhamoensis,H olcolaetis and
and oviposition. However,t he nestso f 1: bhamoensis Euryattust,w o generao f more advanceds alticids,a lso
are morphologically different from the tubular nestso f build egg sacst hat are not contiguousw ith the silk of a
most salticids: 1: bhamoensism ade silken rivets and nest( Jackson1, 985,1 986a).F urthermore1, : bhamoensis
sandwiches itself between two leaves rather than resembleH olcolaetisa nd Euryattusi n spinningf lat and
enclosing itself in silk (also see Jackson, 1986a). denselyw ovene gg sacsa, nd noneo f theset hree genera
Furthermore,t ypical salticids spin their egg sacs by build the typical tubular nests (Jackson,1 986a).T his
covering a few layers of denses ilk that is continuous suggestsa possiblep hylogenetica ffinity of theset hree
with the enclosing silk of a tubular nest (Jackson, genera( Thiania,H olcolaetisa nd Euryattus).H owever,
1979) while 1: bhamoensis encloses its egg sac unlike 1: bhamoensisth at spins the egg sacs between
between two leaves in the rivet silk that is not leaves,H olcolaetisl ays the egg sacs on the trunks of
continuousw ith the silk of egg sac.W hy do the species trees and Euryattus in rolled up leaves.T his may be
from the genus Thiania (Jackson, 1986a)b uild this becauseo f differents electivep ressuresth e spidersf rom
aberrantf orm of nestsa nd egg sacs?O ne explanation these three genera may have evolved the spinning
has been proposed for this form of nest (Jackson, behaviourto uset he tree trunk or a leaf,i nsteado f silk,
1986a).T heseo pened,r ivet-like nestsm ay function as to preventt he nesta nd thus its egg sac is attachedt he
a significant anti-predatorm easure( Jackson, 1986a) trunk leaf insteadH. owever,n othingi s known aboutt he
becauset hese nests have many doors so that spiders selectivep ressureso n the spinning behaviouro f these
cane scapef rom almosta nyd irection. However,t o test threeg enera. In addition,t he phylogenetica ffinities of
hypothesisw e needt o show,f or example,t hat Thiania theset hree generaa re also not clear. Obviously,m uch
that spint he opened,r ivet-like nesth as higher survival morec omparativein formationo nt he biology of salticids
rate comparedt o salticids thatm aket he enclosedn ests. is needed.
This hypothesisi s now being investigated.
Other interesting questions also arise regarding the Agonistic behaviour
evolution of salticids. For example,i s the spinning
behaviour of 1: bhamoensisp rimitive among the In thoses alticidst hath aveb eens tudiedi n details,m ales
Salticidae or modified from typical tubular nestsb uilt have large and complex display repertoire (Jackson,
by majority of salticids? If the nest and egg sac 1977a, b, 1980, 1986b, 1989), comparable to the
spinning behaviouro f 1: bhamoensisis primitive, the maximumr epertoires izesr ecordedf or mammalsb, irds,
nests and/or egg sacs of 1: bhamoensiss hould be fish and sociali nsects( Moynihan,1 970;W ilson, 1975;
similar to those of primitive salticids. Lyssomanes Smith, 1977). In its male-male interactions, 1:
viridis and all spartaeine salticid genera (Portia, bhamoensisp erformed 12 'major displays' (according
Brettus, Cyrba, and Phaeacius)s tudied so far are to Moynihan's (1970) definition): Hunchedp osturing
149
Li et al.: Agonistic behaviour of fighting spiders
(1), posturing with legs I arched (2) or elevated in assistance of K. T. Ang for keeping the spiders alive
Positions 1 and 2 (3, 4), waving with legs I arched (5), with his ready supply of fruit flies and houseflies. We
bent abdomen (6), abdomen flexed up or down (7), thank H. K. Yip for his photographic assistance. Some
embracing (8), palpal pushing (9), grapple (10), photographs (Figs. 5 & 6) were kindly taken by Robert
stepping-to-the-side (11), stepping-forward-and- R. Jackson. Comments and suggestions from Naomi
backward (12). This is comparable to repertoire sizes Diplock, Wan She Lee, Lek Min Lim and one
that have been estimated for numerous other salticids anonymous reviewer greatly improved the quality of
(see Jackson, 1988). These crude estimates suggest that this manuscript.
complex display repertoires are widespread in
Salticidae.
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