Reproductive behavior of Homalonychus selenopoides (Araneae: Homalonychidae)

2011·8.4 MB·English

by  José Andrés Alvarado-Castro

2011.TheJournalofArachnology39:118-127
Reproductive behavior ofHomalonychus selenopoides (Araneae: Homalonychidae)
JoseAndresAlvarado-Castro'-^andMaria LuisaJimenez'-': ‘Laboratorio deAracnologiay Entomologia, Centrode
Investigaciones Biologicasdel Noroeste(CIBNOR), MarBermejo 195, Col. PlayaPalodeSanta Rita, La Paz, B.C.S.
23096, Mexico;^CentrodeEstudiosSuperioresdel EstadodeSonora(CESUES), UnidadAcademicaHermosillo, Ley
Federal delTrabajo s/n Esq. Israel Gonzalez, Col. Apolo, Hermosillo, Sonora 83100, Mexico
Abstract. HomalonychusselenopoidesMarx 1891 isendemictothecoastalplainsoftheSonoranDesertinthestateof
Sonora,MexicoandthesouthwesternUnitedStates.Althoughthespecieswasdescribedmorethanacenturyago,nothing
isknownaboutitsbehavior. WecollectedspidersinthesouthernSonoranDeserttostudytheirreproductivebehavior,
which we recorded with an infrared camera, mainly at night. Sperm induction was ofan indirect type; males wovea
triangularsperm web about 2 cm“ neartheground. Femalesand malesprepared threads ofsilk and sand. Courtship
behaviorwasintermediatebetweenlevelsIandII,andthecopulationpositionwasamodificationoftypeIII,wherethe
maletiedthefemale’slegswithsilkbeforemating.Sexualcannibalismmayoccurduringmating. Femalesbegantospin
theireggsacat~11 daysaftermatingandcompleteditin~15h,includingovipositioning.Theouterlayeroftheeggsac
containedsand,andthesacwassurroundedbyagarniturec cordsofsilkandsand,possiblytoprotecttheeggsfrom
desiccationandasabarriertoparasitesandpredators.
Keywords: Sperminduction,courtship,copulation,eggsacs
Homalonychus selenopoides Marx 1981 is endemic to concerned only with the systematics or phylogeny of
southwestern Arizona and small areas in southern Nevada homalonychids.
andCalifornia.InMexico,itoccupiesthecoastaldesertplains Here, we describe the reproductive behavior of H.
inthestateofSonoraand IslaTibtiron(Roth 1984;Crews& selenopoides under laboratory conditions, including sperm
Hedin2006). Despiteitsbroaddistribution,andmorethana induction, preparation of silk threads with adhering sand,
century after it was first described (Marx 1891), virtually courtshipandcopulation,andspinningoftheeggsac.
nothingis known about itsbehavior.Thisspeciesisincluded METHODS
in the family Homalonychidae, which is represented only by
the genus Homalonychus Marx 1891, including two species. We collected spiders in the bed and sloping sides ofthe I
Theotherspecies, H. theologusChamberlin 1924,inhabitsthe ephemeral stream El Macapul and surrounding area located .
Baja California peninsula, extreme southeastern California, northern of San Carlos, Sonora (27°59'00"N, lir02T6"W
athnadtsaroeutnhoetrncoNmemvoadnal.yHenocmoaulnotneyrcehdi(dVsetatreer&curCsoorkieanldoslpipdheerrs aofndth2e8S°i0e0r'r5a5E"lN,Agu1a1j1e°.03T'h0e5"cWl)i,maitnetihseveerxytrderym:ehsootutinhesrunmpmaerrt ^
2000); they are nocturnal and conspicuous. Adult males are andwarminwinter.Themeanannualtemperatureis22-24°C j
6u.s5u-a9l.l0yfmomu,ndainndfinaeduslatndfeomralseosilaarned7u.n0d-e1r2.r8ocmksm, waonodd,aorer wainndtetrhreaimnfeaallnisasnpnluital~r9a0in%falalndis~751-02%0,0remspme;ctisvuelmym(eIrNEaGnId |
debris.Typically,juvenilesandadultfemalescamouflagetheir 1999). Vegetation is desert scrubland with Bursera and
bodieswithfinesoilparticlesthatadheretothesetaeoftheir Jatropha predominating (INEGI 1984). Soils are weakly
integument, which allows the spider to blend in with the developed and shallow (< 25 cm), usually composed of
surrounding soil (Duncan et al. 2007). They are often found unconsolidated coarse-textured sand and fine gravel with
slightly buried in the sand with their legs extended (Roth rocky areas without soil or some soil found in depressions
1984). among the rocks (INEGI 2002). The stream bed is almost
Gertsch(1979)mentionedthatthefamilyHomalonychidae entirelysandandgravel.
was enigmatic because very little was known about it. Even We made 17 diurnal collections with 3^ participants
now, thereare few studiesavailable. Roth (1984)carried out between October 2007-April 2008. During this period, we
systematic studies of the family, Vetter & Cokendolpher captured 186adultand immaturespiders from understones,
(2000) described the egg sac and defensive posture of H. drycattledung, wood, bricks, orcardboard. Weplacedeach
theologus, and Dominguez & Jimenez (2005) reported on livespiderindividually inaplasticcontainerandtransported
sexual and cryptic behavior ofH. theologus. Crews & Hedin all ofthemto thelaboratoryin Hermosillo, Sonora, Mexico. '
(2006) explained the phylogenetic divergence of the two Male and female voucher specimens were preserved in 75%
species and Duncan et al. (2007) described the convergence ethanolanddepositedintheArachnologicaland Entomolog-
o'!HomalonychusandSicariusWalckenaer1847(Sicariidae)in icalCIBNORCollectionin La Paz. j
themorphologyoftheirsetaeforretainingsoilparticles.Other We maintained each live spider individually in a 500-ml
studies(Roth 1984;Griswoldetal. 1999;Milleretal.2010)are transparentplasticjarcontaining 1 cmsoilsubstratefromthe
collection siteand asmallcontainerofwetcotton forwater,
Specimens were initially fed crickets (Gryllidae) and cock- j
^Correspondingauthor. E-mail:[email protected] roaches(Blattellasp.),andlatermealwormlarvaeTenehriosp.
I
ALVARADO-CASTRO&JIMENEZ—REPRODUCTIVEBEHAVIOROFHOMALONYCHUS 119
(Dominguez & Jimenez 2005). We used mealworms because
theyareeasytocultivate. Thebreedingroom(3 X 3 m)was
kept at 18-28° C, under natural photoperiod, and 36-60%
relative humidity. We observed courtship and copulation in
this facility, but made observations ofsperm induction and
spinning of egg sacs in another small room. We recorded
spiderbehaviorwithan 8 mmdigitalcamcorderequipped to
recSopredrimnfrIanrduecdtiloignh.t—. We placed five males reared in the
laboratory and two field-collected males individually in
1750-mlclearplasticjars (13 cm diameter) with fine sand to
adepthof2.5cm.Weaddedasmallflatstoneforattachment
ofthespermweb,aswellasanarchedcardboardshelteranda
smallcontainerofwetcotton. From 14March-14April2008
from 20:00-08:00 h, we made momentary observations at
intervalsof20min usinganinfrared lightcamera. For these
specimens,theambienttemperaturewas 17.2-30.7°C,natural
phoMtaotpienrgiobde,haavnidor.2—0^F7r%omreJlaatniuvaerhyu-mMiadritcyh.2008, we formed Figure 1.—Homalonychus sdenopoides male during loading
25matepairingswitheightadultmalesand23 adultfemales ofsperm.
collected in the field (ageand reproductivestatus unknown).
Because we had few males that were very variable in their and3hadaflathorizontalroofatleast5X5cmataheightof
behavior,weusedmainlymalesthatwereactivelysearchingin 2.0-2.5 cmabove sand level. Weplaced five femalesin these
these trials; the other males were less active or fled from terraria,replacingthemevery4-5daysiftheyfailedtospinan
females.ThroughoutJuly2008,weformedanother20pairings egg sac. Observations lasted from 22 April-16 May 2008.
with14malesand12femalesrearedinthelaboratory,(virgins, Ambient temperature was 24.8-33.8° C, with natural photo-
ofknownage)plusonefemale fromthe field. In thesetrials, period,and 16-31%relativehumidity. Wedidnotobserveor
wemade these pairings at random, although the males were record the spinning of the egg sacs by females that had
alsovariablein behavior. Weformedadditional matingpairs copulatedinthelaboratoryinJuly2008;however,wenoticed
(one in October 2008 and 18 in July-August 2009) to see if thateachfemalehadproducedseveraleggsacs.
additional behavioral acts had been undetected during the RESULTS
ainniatliyalsesp.aiIrninalgls;thetsheesceaserse,suslotsmewfeermealensotandu,semdorienfrsetqautiesnttilcya,l Sperm induction.—Weobserved theentiresperminduction
males were used again to form new pairings. Observation processonce(02:38-03:00h),whenamalewoveaspermweb
schedules and laboratory conditions were as follows: in in 5.9 min, close to the sandy substrate; it was slanted and
January 2008, 14:30-18:00 h, 18-19° C, 50-60% relative attached to thecardboard shelterand to the wall ofthejar.
humidity;inFebruary2008, 15:00-20:00h,24-25°C,50-60% Themalestoodonthesubstrate,placedhisbodyontheweb,
relativehumidity(temperaturewasmaintainedwithanelectric and pressed against it twice. Infrared light failed to show
heater^inJuly2008,20:00-23:00h,24-28°C,36-55%relative spermdeposition.Subsequently,themalemovedapedipalpin
humidity.Weplacedindividualfemalesinglassterraria(20 X an arch-like motion from top to bottom on one edge ofthe
20 X 10 cm) containing a 2-cm substrate offine sand. We webtoloadthepedipalpwithsemen,rubbingtheventralpart
introduceda male20to 177minlater(median = 72min). If ofthecymbiumagainstthelowersurfaceoftheweb(Fig. 1)
the female was receptive, we filmed the behavior and withsoftmovements. Heraisedthispedipalptocarryoutthe
continued filming for 15 min aftercopulation. We separated same process with the other pedipalp. So, the semen was
individuals or changed their partners ifcopulation failed to deposited on the upper side of the web and it was then
occurwithin55min,orsooner,iftheytriedtoescape,orifan absorbed through to the underside. This stage took 7.8 min.
individual repeatedly ran from its partner or assumed a The male then climbed offthe web and rested on the sandy
defensive posturing of paired legs. When disturbed, these substrate.Theentireinductionprocesstook 16.5min.Wealso
spiders extend their first two pairs of legs together and observed the last 2 min ofsemen loadingofanothermaleat
forwardandthelasttwopairstogetherandbackward(Vetter 04:28 h,withapositionand processidentical totheonethat
& Cokendolpher 2000). In one trial in July 2008, we wehad observed in itsentirety. This male then rested on the
intErgogduscaecdctownsotrmuaclteiosn.s—imuWletanuesoeudsl2y0.capturedadult females, weTbhrfeore2l.a2bohr.atory-rearedmales(age6-8daysasadults)and
each of unknown reproductive status but with a large two field-collected males wove six sperm webs (one in
opisthosoma,toobserveeggsacspinning.Thesefemaleswere November 2007 and five in March-April 2008). Web
captured in the winter of 2008. We placed each female dimensions varied from 9 X 13 X 15 mm to 21 X 26 X
separatelyina 1750-mltransparentplasticjarcontaininga3- 28 mm. Webs were triangular, thin, and semi-transparent,
cm sand substrate and one ofthree types ofshelters: 1) an withoneorseverallayersofsilk(Fig. 2).Webshadtwostrips
arched piece ofcardboard; 2) flat stones glued togetherwith ofdensersheetsthatextendedfromthecentertooneedge;on
molding silicone; or 3) stones with a glass ceiling. Shelters 2 thisedge,themalearchedhispedipalpsduringinduction.The
120 THEJOURNALOFARACHNOLOGY
— —
Figure 2. SpermwebofHomalouychitsselenopoides. Figure 3. Thread of silk and sand spun by a Homalonychus
selenopoidesmale.
websweresetbetweenstoneorcardboardandthewallofthe
jar, inclined at anglesof40 70°, with a heightaboveground
bleevtewleaetnt1h2e-i2r4lmowme.stOnbeetmwaeelnew2o-v8emtmwoasnpdermatwethbesi,rahniogthheesrt cspoindseirdserecdopiunlaotuerdantawliycsei;s.tSheexsuealsebceohnadviomratwiansgsdivwiedreed innotto ®
wmeablse.woveoverapriorweb,andtwomaleswoverectangular (thGroenezasltaegzes1:98p9r;e-Dcoopmuilnatgiuoenz,&copJuilmaetnieonz,20a0n5d).poTshte-csoepquuleantcieosn
butWetheosbesewrevreednvoartiaotbisoenrsviendfdourrminagndcosniszteroufctoitohne.rTmawloemwaelbes,s osefcrebteihoanvioofraslilkaacntds saannddtthrraenasdist,ioanrefsruemqmueanrciizese,diinncFliug.di4n.g :
wofovtehetcroinatnagiunlearra~nd1 ctmh-ewmiedseh.spOetrhmerwmeablsesatwtoacvheedwetbostohenttohpe malPere-acdovpaulnacteidon:inDwuhraitngaphipseasreeadrcthotboefiandratnhedofmemamlaen,netrh,e j
sand that were 1-2 cm long, as short strips that went from exploring,walkingslowly,andgraduallyraisingandlowering s
“aggregates” ofsand and silk from the ground, stuck to the hisfirstpairoflegs.Themalecouldalsoapproachthefemale
wallofthejarorthecardboardshelters.Somespunelongated directly in a targeted manner when he apparently had
smiallkesshefiertsstw(o~ve1 sXmaltloer5.w5ecbms)beufpoorenusnadnedrytaakgignrgeglaatregse.rsOptehremr tidoemntaitfiinedgrhearn.geIdnf1r6omob0s.e1r-v3e9d.4cmoipunla(tmieodnisa,ns=ear1c1h.8tmiimne).prTihoer
webs. initial contact or touch between potential partners was with
Silk and sand threads.—In July 2008, five males placed in the tarsi ofthe forelegs. When the male reached a receptive i
glass terraria spun six threads of silk and sand in form of female, she became passive and he quickly and repeatedly
c“ocpourldast”io(nF,ig.a3n).dTharneoethtehrreadwsaswersepsupnunwbietfhooruetantdhetwospaifdteerr tthoeuctharesdiaofndhistafpopreeldeghserfoprr~oso1m-a3,s.oIpfitshtehofseommaal,eowrasleignsitwiailtlhy ij
participating in copulation. Males spun threads with their unreceptive,shecouldabruptlyretreatorwalkaway.Thenthe .
spinnerets, moving slowly with their legs close to their body maleinitiatedthecourtship.Femalesalsoinitiatedapproaches f
andconstantlytouchingthethreadwiththeirpedipalps.They or courtship; then the male could flee or begin tapping or
walkedveryclosetothefloor,weavingin thesametrack two begincourtship.Rejectionsinformofattacksagainstconsorts
oreven five times. Thespiders spun threads in4.7-18.3 min. wereobservedonlyinonepair; thefemaleattackedthemale ^•
Fourofthethreadsrangedfrom~8.0-17.3cm,withknobsor and laterthemaleattackedthefemale.
swellingsatoneorbothends.Twothreadswere1.9and2.4cm During courtship, the male drummed on the ground with
long,withonethickendandtheotherendbifurcated.Wedid his forelegs or with his first two pairs oflegs. Legs vibrated •
notobservereactionsoffemalestomalethreads, becausethe whentheywereincontactwiththeground.Theleftandright
males approached the female to mate before the females legs were extended and moved up and down quickly and
walkedonthethreads. InJuly2009,onefemalespunthreads alternately. Also, he drummed on the ground slowly and
with silk and sand prior to copulation. The female continu- gently with the pedipalps while moving forward or side-to-
ouslywovethesethreadswithherspinnerets,leavingagridof side. When a female initiated courtship, she approached the
threadsonthesand.Thethreadswereverythinintheformof maletotouchhim,thentooka“stalking”stancewhilemoving
a rosary, butwerevisiblebecause thesand grainsadhered to slowlyorswiftlywith oneormorequickapproaches. Ofthe
them. The male placed in this terrarium encountered the observed pairings, 50% included some period of male
female’s threads and immediately began spinning a thread courtship. In 25% of the 16 pairings, females approached
(coMrad)t.ingbehavior.—Weobserved 16successfulpairings,three afnrdomto<uch1e-d33m.a5lems.inWh(emendiitanocc=urr3e.d1,mmianl)eaconudrtsthheipfleamsatleed i
in January-March 2008 and 13 in July 2008. Two pairs of courtshiplastedonlyafewseconds.
ALVARADO-CASTRO&JIMENEZ—REPRODUCTIVEBEHAVIOROFHOMALONYCHUS 121
Copulation: Afteramaletouchedafemale,shebroughther After breakout, females rubbed their legs together to
legs toward her body, leaving the patellae almost touching remove the remnants ofthe bridal veil. 38% ofthe females
abovethecarapace;onlythetarsiandmetatarsiofthefourth duginthegroundatleastonetime,thenrubbedandwiggled
pair of legs were directed backward. The female remained thebackandbellyoftheirprosomaandopisthosoma,andlegs
passiveandmotionlessinaquiescentstate(Beckeretal.2005). inthesoil; sand particlesthenadhered to theirbodysurface.
Themaleclimbedontothebodyofthefemale,tappingherwith We did not observe this behavior in males. In all pairings,
thetarsi oftheforelegsandpedipalpsanywhereon the body males vibrated their opisthosoma after dismounting; they
andlegs.Thenthemaleclimbeduponesideorthebackofthe raised and lowered it with quick short movements. Also, the
female and settled on top ofthe female, facing the opposite malescleanedtheventralcymbiumofthepedipalps(presum-
direction.Duringmounting,themalecontinuouslytouchedthe ablycopulatorystructures)withtheirchelicerae.Theseactions
bodyofthefemale.Of16observedcopulations,insevenofthe occurredatleastonetimeineachmaleandtookplacewithina
mountings(44%),malesapproachedthefemalesfrontally;the few minutes aftercopulation. Males showed post-copulatory
othermountingsweremadefrombehindorfromoneside. courtshipin50%ofthecouplings.Wepresentthefullrangeof
While mounted, the male wove threads of silk in circles post-copulatoryactsandtheirsequencesin Fig.4.
aroundthelegsofthefemaleto formabroad ringtie, likea In January-March 2008, there were two cases where the
veil,coveringtheexposedsurfaceofthelegs,excepttarsiand maleswerecapturedandkilledbythefemaleswithinthefirst
metatarsiofthefourthpair.Themalealsoaddedsandtothe 7minofwaiting,withoutcourtshipormountingtakingplace.
silkon thesidesand bottomofthe female bodyas “counter Whenmaleswerekilled,theirbodycontentswereconsumedin
balances.” This web is known as the “bridal veil” (Bristowe the subsequent (undetermined) hours. In January 2008, we
1958; Dominguez & Jimenez 2005). While the male was observedoneeventofsexualcannibalismaftercopulation. In
weaving, hewas tappingthe female’s body and legs with his this case, after the last insertion ofthe pedipalp the female
forelegsandpedipalps.Thetyingwasrepeatedalternatelyand suddenlyextendedherlegs,broketheveilandquicklyreached
successively with insertions ofthe pedipalps (a tying always themaleasheattemptedtoescape;allthistookplaceinabout
precededinsertionofapedipalp). asecond. InOctober2008,therewasanothereventofsexual
During insertions of the pedipalps, the male placed the cannibalism, but thismalewascaughtduringmating. In this
quiescent female on her side, either right or left, moving to case,bothindividualswerelyingontheground,bellytobelly
thatsidewhilehewasembracingherwithhisfirstthreepairs in oppositedirections, when the femalegrabbed themale on
oflegsandrestingwiththefourthpaironthefloor.Themale’s theventralsideofhisopisthosoma.Thefemalebroketheveil,
left pedipalp was inserted into the genital opening of the brokefreeofthemale foramoment, andcaught him. These
femaleontheleftsidewhilethefemalewaslyingontheright maleswerealsoconsumedin thesubsequenthours.
sideorviceversa.Thepedipalpscouldbealternatelyinserted, In the 22 pairs that did not copulate in January-March
orapedipalpcouldbesequentiallyinserted. Duringinsertion 2008, we observed rejection by both males and females,
ofthe pedipalp, the male vibrated his legs II and IV on the immobility ofone or both partners, with or without legs in
samesideastheinsertedpedipalp.Inthe16observedpairings, paired position, and constant attempts to escape from the
the duration of copulation (mounting) ranged from 0.6- terrarium. Also, we observed that some males touched or
9.4 min (median = 1.9 min). The number of pedipalp stood on unreceptive females with their tarsi, but apparently
insertions per mating ranged from 2-12 (median = 2.5); of the females were notdetected. Ourwaitingtime to complete
85 individual insertions, 66% were done with the right thesetrialsrangedfrom—22-55 min.
pedipalpand 34%withtheleftpedipalp. Eggsacconstruction. EightfemalesthatcopulatedinJuly
Successful mating among pairs depended on the origin of 2008startedtospintheirfirsteggsacs9-13daysaftermating;
the females. Ofthe 25 pairs formed with the field-collected spinning was not filmed. Five females collected in the field
femalesinJanuary-March2008,thesuccessfulrateformating began spinning their egg sacs, but only four finished. We
was 12% because only three pairs mated; thus 88% of the recordedthespinningoftwoeggsacs from beginningtoend
femaleswereunreceptive.Onefemalecopulatedtwicewiththe and theothertwoafterthefirstphasehadstarted.
same male during the same session. On the other hand, the Thefemaleinitiatedtheeggsacconstructionbehaviorwhen
rateofsuccessofthe20pairsformedwithvirginlaboratory- sheexploredtheshelterroof;also,shecouldscratchthesandy
rearedfemalesinJuly2008was65%.Therewere 12ordinary substrate.Thenshestartedspinningtheeggsacbyweavinga
copulationsandonecaseinwhichafemalepresentedwithtwo silk sheet, thin and circular, on the roofofthe shelter. This
males,matedfirstwithone,thenminuteslatercopulatedtwice took54and69min.Thereafter,shewovethickdoublestrands
withtheother. Fiveof12virginfemalesreceivedasecondor ofsilkandsandintheshapeofcords.Whileshewasinverted
third partner after rejecting the previous male, but finally ontheceilingoftheshelter,shedroppedheropisthosomaand
100%ofthevirginfemaleswerereceptive.Theonlypairthat fourth pair oflegs grasping the shelter with her three other
includedafield-collectedfemaledidnotcopulate. pairsoflegs.Withherspinneretsincontactwiththesand,the
Post-copulation:Copulationfinishedwhen62.5%ofthemales femalesecretedsilk threadsandadded sand to thesein short
dismountedfromthefemalesandwithdrew,walkingawaywhile zig-zag strokes, leaving a cord behind her, which was also
they remained quiescent for a few seconds. Also, copulation foldedinazig-zagpattern. Afterwards, thefemaleraisedher
finished when 37.5% ofthe females were no longerquiescent, opisthosomaandthefourthpairoflegs,stayinginverted,and
extendedtheirlegsbreakingthebridalveil,andthemalesfled. attachingtotheceilingtheproximalendoftheextendedcord
Femalesusuallytooklessthan2stobreaktheveilandwalkor thatwasattachedtoherspinnerets.Thisprocesswasrepeated
run,althoughonefemaletook 16sandonetook 10min. withothercordstoformafirstoutercircleorringofthesand-
122 THEJOURNALOFARACHNOLOGY
FEMALE MALE
Figure 4.—Sexualbehavioralsequencesobservedin 18pairingsofHomcilonychusselenopoides.a)Pre-copulatorystage;b)Copulatorystage; i
c)Post-copulatorystage.Thenumbersadjacenttoarrowsrepresentthetotalnumberoftransitions.Sequencesthatoccurredoneortwotimesare
not included. Asterisks indicate the behavioral acts where a sequence began, and the numbers beside the asterisks indicatethe numberof 1
sequencesthatbeganintheseacts.
silk garniture ofthe future egg sac. During this process, the fromtheshelteratintervalstorestonthegroundortodigand l
sfepmuanlesilwkasstcreanntdrsalcloyncpeonstirtiicoanleldyiinnswiaderdth(iFsicgi.rc6l)e.(TFhige.g5a)rnaistsuhree accWuemulinafteerrseadndthattaktehneffreommaluensdelirnetdhethsehelitnetre.rior ofthe last '\|'
increasedprogressivelyinthickness,andtheinternalspacewas cordlayercircleoftheeggsacwithsilkbecausethetubewalls l
reducedtoincludethefemaleonly.Thefemaleloweredherself moved continuously, forming the inner layer oftheegg sac. Ui
—
ALVARADO-CASTRO&JIMENEZ—REPRODUCTIVE BEHAVIOROFHOMALONYCHUS 123
a
Figure 5. Homalonychusselenopoides female spinning the outer
ringofsilkcordsoftheeggsac.
The lower end of the tube was gradually withdrawn and
sealed, forming the completed egg sac. Afterward, females
wereimmobilefor5-6.5 h,withonlysporadicmovementsof
thetubularwall.Weinferredthatovipositionoccurredduring
thistime.Subsequently,femalesbrokethebottomsideoftheir
sacswiththeirfirsttwopairsoflegstoexit. Escapingrequired
28 s and 10.3 min for two females observed. Immediately
afterwards,each femaleembraced hereggsacandclosed the
exitrupturewith herspinnerets. Theothertwo females were
notobservedbecausetheywereontheoppositesideoftheegg
sacsfromwherewewerefilming. Ittook 14and 15.5 h from
the start ofweaving the silk sheet until the females emerged
fromthesac.
Thewholeeggsacconsistsoftwosections,athickexterior
garnitureofsand-silkcordsandtheeggsacinthecenter.The
wholestructureisshapedlikeashortcylinderandtheeggsac
—
Figure 7. Egg sacs of Honudonychus selenopoides. a) Egg sac
spun on awide, horizontal surface; b) Eggsacspun on a reduced,
slopingsurface.
can extrude from below, between the garniture of cords
(Fig. 7a). Six othercaptive females also spun egg sacs in the
laboratory. One female spun a flattened egg sac under an
inclined rock in a very narrow space (Fig. 7b). Later, this
femalespuntwootherflattenedeggsacsunderthesamerock.
Moreover,intheabsenceofashelter,fourunobservedfemales
deposited naked eggs directly on the sand surface and the
other female also deposited naked eggs on the woven cloth
thatcoveredthejar.
DISCUSSION
We observed all stages of reproductive behavior of H.
selenopoides. Mostreportsonspiderreproductionincludeonly
— somestages.Sperminductionhadnotbeenobservedbeforein
Figure 6. Full egg sac of Homalonychus selenopoides showing theHomalonychidae,andthefunctionofthebridalveilinH.
concentricarrangementofthesilkcords. selenopoidesstillremainsobscure.Apparently,addingsandto
124 THEJOURNALOFARACHNOLOGY
thesilkthreadsmadebymalesandfemalesandthegarniture Thepheromonesreleasedbyfemalesspidersasanattractant
ofcords ofsilk and sand surrounding the egg sacs spun by formalestoinducecourtshipareamplydocumented(Gaskett
females only occur in these spiders. We here discuss the 2007).However,inourstudy,onlyonevirginfemalespunsilk
functional role of these features and possible phylogenetic threads. It is possible that the small size of the terrarium
implicationsofthei—rsexual behavior. permitted pairs to meet more easily than in the field, so
Sperm induction. The horizontal, triangular shape ofthe spinning of silk threads by females (and males) was
sperm web matches what is commonly observed in spiders unnecessary, and these silk threads were by-passed in favor
(Foelix 1996). The square form is also common (Gertsch of direct contact between partners (Dondale & Hegdekar
1979). We found both web forms in different sizes, but the 1973). In the field, where these spiders are uncommon, silk
factors that determined the shape and size ofthe webs were threadscouldplaya—nimportantroleforlocatingmates.
notcleartous.AlthoughthespermwebofthesisterspeciesH. Mating behavior. In general, mating behavior of H.
theologiisistriangular,itsareaisonly2-Amm^(Dominguez& selenopoidesissimilartoH. theologus. Inbothspecies,males
Jimenez2005), much smaller than whatwe found among H. usually take the initiative and approach females; however,
selenopoides. Duration ofsperm induction is consistent with some H. selenopoides females made approaches and initial
observedbehaviorofmostspiders,whichrequirelessthanhalf contact to trigger the search ormalecourtship. Initiativeby
an hour to perform (Gertsch 1979). The filling ofpedipalps females for courtship was not observed in H. theologus
withspermcorrespondstotheindirectform(Foelix 1996)and (Dominguez & Jimenez 2005). Females starting courtship
is consistent with what is commonly reported for cursorial has also been observed in Lycosa spp. (Costa 1975; Rovner
spiders(Jackson&Macnab 1991).Thealternatingloadingof 1968). Although Homalonychus females are relatively seden-
pedipalpsissimilartoSchizocosacrassipes(Walckenaer 1837) tary(Crews&Hedin2006),itispossiblethat,intheirsexually
(Lycosidae),butdiffersinthatS. crassipesslowlyagitateseach receptive stage, they are more vagile. Active participationof
pedipalp after loading the sperm (T. H. Montgomery in bothsexesinsearchandcourtshipmayexplaintheirpresence
Gertsch 1979). Webs were not consumed by males, as in inpitfalltrapsinthecollectionarea. 15of17//. selenopoides
Sicarius(Levi 1967). specimenstrappedwereadultmales(47%)and adultfemales
Induction is a common phenomenon, but observing this (53%)(unpublisheddata).
behavior requires patience (Gertsch 1979). Reports of InH. selenopoides,mountingoccurredoneithersideofthe
induction vary from only descriptions of sperm webs female. Duringcopulation, themalesvibratedlegsII andIV,
(Dominguez & Jimenez 2005; Sierwald 1988), partial obser- incontrasttoH. theologus,wheremountingoccurredfrontally
vations of the induction process (Fraser 1987), single and males vibrated legs II and III during copulation
observation of the entire process (Levi 1967; Jackson & (Dominguez & Jimenez 2005). In both species, copulation
Macnab1991),andrepeatedobservationsoftheentireprocess could finish when the maleceased activity, dismounted from
(Rovner 1967; Stumpf 1990). When theprocesstakes several the female, and withdrew, but in H. selenopoides, there was
hours,itiseasiertoobserve,asinsomeTheraphosidae(Costa variation in theway toendcopulation. In this latterspecies,
&Perez-Miles2002).Themaleswestudiedwereverysensitive copulation also ends when the female suddenly spreads her
to light, sound, and vibration during sperm induction and if legs, breaksthenuptialveil, andthemalehasto flee.
disturbed, either ceased their activity or did not initiate it. CourtshipfallsbetweenlevelsIandIIdescribedbyPlatnick
Hence, we assume that successful observations ofinduction (1971), as in H. theologus (Dominguez & Jimenez 2005),
dependonitsduration(Costa 1975),sensitivityofthespecies Lycosidae, and Pisauridae. Evidently, the primary triggerof
to surrounding environmental events, and whether the courtship or mounting behavior in the male is the direct
induction is unpredictableor it occursimmediatelybefore or contact with the female, but we hypothesize that males can
aftSeirlkpsaenuddos-acnodputlharteaidosn.—orWceopuwlearteiosnu.rprisedtoobservemales talhseoredeitsecat acofnetmacatlesbeyxaphcehermoimcoanlestiinmutlhues.cWuteiclaessoufmevitrhgaitn
and females spinningthreads ofsilk and sand. We noted that females (Dondale & Hegdekar 1973). When males touched
immatureandadultspecimenshavetheirspinneretscontracted unreceptive and motionless field-collected females in some
intheopisthosomaand,likeothercursorialdesertspiders,donot pairs,theydidnotattemptmounting.Butinmostotherpairs,
createsecuritythreads. Hence,weassumethatreleasingthreads whenthemalestouchedvirginlaboratory-rearedfemales,they
whenmalesandfemalesaresearchingforpotentialmateshasa immediately attemptedmounting. Malespidersdetectphero-
role in sexual marking. The presence ofsex hormones in the mones by touching the females because they have tarsal
threadsispossiblebecausesilkisthemainhormonalsubstratein receptors involved in sexual recognition (Foelix 1996).
spiders; in other species both sexes emit and respond to Pheromones that attract or promote the courtship ofmales
pheromones (Gaskett 2007). Male silk can attract females inthefemalecuticlehavebeenreportedinatleast25speciesof
(Roland 1984) and promote the beginning ofcourtship (Ross spiders (Gaskett 2007). Pheromones in Homalonychus and
& Smith 1979). This function seems reasonable for H. theirroleinsexualbehaviordeservetobeinvestigated.
selenopoides, because it rarely occurs in the field (unpublished Homalonychus selenopoides take the “lycosid position of
data).Moreover,malesilkaffectscourtshipofconspecificmales copulation” (position III, Foelix 1996), similar to what is
(Ross&Smith1979;Ayyagari&Tietjen1987).Weobservedthat described for other wandering spiders, such as Lycosidae
a male walking on a thread produced by another male (Stratton et al. 1996), Pisauridae (Merret 1988), Agelenidae
immediately stopped and wove his own threadjust above the (Fraser 1987), Philodromidae, Clubionidae, Salticidae, and
previous one. There is no precedent in the literature for this Thomisidae (Foelix 1996). Basically, in this position, males
behaviororaboutspidersaddingsandtosilkthreads. mountfacingtheoppositedirectionfromthefemale,withthe
ALVARADO-CASTRO&JIMENEZ—REPRODUCTIVEBEHAVIOROFHOMALONYCHUS 125
ventralsurfaceofthemaleprosoma on thedorsal surfaceof Egg sac construction.—We have not found a precedent in
thefemaleopisthosoma.Inlycosids,malesleantowardseither anothergenusofspidersforgarnituresofsilkandsandcords
sideofthefemaletoinsertoneoranotheroftheirpedipalps. surrounding the egg sac as in Homalonychus. Although
In H. selenopoidesthis position is modified. The male places Sicariusattaches sand to thewall ofitseggsac(Levi & Levi
the quiescent female toward one side and then the other to 1969),itdoesnotmakeagarnitureofcords. BecauseSicarius
insertoneoranotherofhispedipalps,similartothereporton spp. inhabits deserts ofSouth America and southern Africa
Ancylometes hogotensis (Keyserling 1877) (Pisauridae) (Mer- (Platnick 2009), Dominguez & Jimenez (2005) suggest a
rett 1988) although in H. selenopoides the insertion of convergence between the two phylogenetically unrelated
pedipalps is not strictly alternating. Afterthis point, copula- genera as a response to harsh desert conditions. However,
tionisidenticaltothatofH. theologus(Dominguez&Jimenez therearedistinctdifferencesinthetimingandeggsacspinning
2005). process,form,andstructure,andthefactthatSicariusspp.use
The low frequency of sexual cannibalism observed is theirlegstoburytheireggsacswith sand.
consistent with theclaim that high frequencyofcannibalism The description ofthe egg sac of H. theologus (Vetter &.
isamythandnotcommonamongspiders(Foelix 1996).The Cokendolpher2000)isincompletebecauseit failstomention
two events ofsexual cannibalism here observed are the first thethickexteriorgarnitureofcords, although in a published
reportedfor Homalonychidae, becausethisbehaviorwasnot photographsomeofthemareapparent.Also,spinningofthe
observed in H. theologus (Dominguez & Jimenez 2005). For egg sac of H. theologus (Dominguez & Jimenez 2005) was
theothertwocasesofpredationuponmales,theseeventsdid made at an atypical site, the side wall ofthe container. We
not represent sexual cannibalism because there was neither infer that Homalonychus requires a shelterwith a horizontal
courtshipnorcopulation(Elgar 1992). roof for spinning typical cylindrical egg sacs with exterior
Regarding success in pairings, it is possible that H. garnitureofsilkcords.Wesuggestthatfurtherstudyisneeded
selenopoides females are monandrous. This would explain todefinethetypicalstructureandspinningprocessofeggsacs
the marked difference in the percentage of successful in H. theologus. We agree with Vetter & Cokendolpher’s
copulations between females collected in the field and the (2000)andDominguez&Jimenez’s(2005)hypothesisthatthe
virginfemalesobtainedinthelaboratory.Itislikelythatmost sandcoveringtheeggsacactsasaprotectionfrompredators
femalescollected in the field had already copulated since we andparasitesandamelioratestheintensedesertsummerheat,
alsocollected—adultmales. where temperatures can exceed 45° C. We suggest that the
Bridalveil. ThebridalveilisdefinedbyBristowe(1958)as cordgarniturehasthisfunct—ion,at least.
silkthreadsdepositedbymalesonfemalesduringcourtshipor Phylogenetic implications. Since the genus Homalonychus
copula.Althoughitoccursinspeciesofatleast 12families,the was described in 1891, it has remained in an uncertain
veil of H. selenopoides is only identical to H. theologus phylogenetic placement (Griswold et al. 1999). Historically,
(Dominguez&Jimenez2005). Accordingtothebriefdescrip- researchershavehypothesizedthatthereisarelationshipwith
tion of the veil of Thalassius spinosissimus (Karsch 1879) Pisauridae, Selenopidae, Zodariidae, Ctenoidea, and Pisaur-
(Pisauridae)(Sierwald1988),theshapeandwidthofthebundle oidea(Crews&Hedin2006).Proposalsbasedonmorphology,
appeartobesimilartothetwoHomalonychusspp.Theextent sexual behavior, and even on molecular analysis appear
oftyingisalsosimilartoA.hogotensis(Merrett1988),butinthe insufficienttodrawastablephylogenetichypothesis.
pisaurid,theveiliscomposedofanouterringatthedistalend Courtship and mating behaviors are considered important
oflegsI IIIandaninnerringatthelevelofthepatellae. characteristicsforreconstructingphylogeneticrelationshipsin
Several functional hypotheses have been proposed for the spiders (Platnick 1971; Bruce & Carico 1988; Stratton et al.
bridal veil (Ross& Smith 1979; Schmitt 1992; Dominguez& 1996).Basedonthematingposition,andoccurrenceandform
Jimenez 2005; Aisenberg et al. 2008). We cannot support or ofthebridalveil,Dominguez&Jimenez(2005)suggestthatH.
refutethesuggestionthattheveilinH. selenopoidesfunctions theologusisrelatedtoPisauridaeandcouldbeincludedinthe
asadeterrenttoothermalesduringcopulation. However,we superfamily Lycosoidea ofCoddington & Levi (1991). Based
doubtthattheveilinH. selenopoidesaidstoidentifythemale on morphologicalcharacters, Roth(1984)proposed retaining
as a consort because the veil is woven when the female is Homalonychidaeas aseparate family,criteria maintained by
receptiveandhasbecomequiescent,nordowebelievethatthe Coddington and Levi (1991). Griswold et al. (1999) lists
veilrestrainsthefemaletopreventherfromattackingthemale Homalonychidae and seven other families as groups whose
orinhibittheaggressivenessofthefemale,assuggestedforH. relationshipsin highertaxaareuncertain.
theologus(Dominguez&Jimenez2005).Weobservedfemales In a molecular survey, Miller et al. (2010) find Homalo-
that quickly broke free of the veil after copulation, ending nychidae are very closely related to Tengellidae, but the
their quiescence. The female that cannibalized her partner phylogenetic placement of both families was inconsistent.
immediately aftercopulation broke out and captured him in Penestomidae was very closely and consistently related to
about one second. Robinson & Robinson (1973) proposed Zodariidae,withallfourfamiliesincludedintheZodariioidea
that the main function of the bridal veil in all species that clade. The possible relationship of Homalonychus with
produceitistostimulatethefemale. Preston-Mafham(1999) zodariioids opens the possibility of finding homologies in
argued that courtship behavior in these species is very reproductive behavior; however, the sexual behavior of
rudimentary,butpheromonesintheveilmaycauseimportant Tengellidae and Zodariidae is too slightly known (Barrantes
physiologicalchangesinthefemaleepigynumtoprepareitfor 2008; Pekar & Krai 2001; Pekar et al. 2005) to make
insertion ofthe pedipalps. To fullydeterminethe role ofthe comparisons and afford a basis forconsidering relationships
bridalveilin Homalonychusrequiresfurtherinvestigation. with Homalonychidae.
126 THEJOURNALOFARACHNOLOGY
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