Table Of Content2002. The Journal of Arachnology 30:181-188
WEB—
SHARING A ON THE RELATION OF SOCIALITY AND
KLEPTOPARASITISM IN THERIDHD SPIDERS
(THERIDHDAE, ARANEAE)
Ingi Agnarsson: Department of Biological Sciences, George Washington University,
2023 G Street NW, Washington, D.C. 20052, USA & Smithsonian Institution,
National Museum of Natural History, Department of Entomology, 10^*^ Street and
Constitution Avenue NW, Washington D.C. 20560-0105, USA.
ABSTRACT. Sociality and kleptoparasitism occur commonly in theridiid spiders. In both behaviors a
number of conspecifics occupy a single web; gregariousness entails tolerance. Sociality has evolved several
times in theridiids, but kleptoparasitism seems to have arisen only once. All four or more instances of
sociality in theridiids are concentrated within a clade of relatively distal theridiids. This distribution of
sociality suggests common cause, i.e. the presence of some characteristics that may facilitate the evolution
of social behavior. The monophyletic genus Argyrodes, many of which are kleptoparasitic, is sister to the
clade containing all social theridiids. Sociality and kleptoparasitism may thus be phylogenetically related
in theridiid spiders; behaviors that facilitated the evolution of sociality could also have facilitated klep-
toparasitism. Both may have their roots in maternal care.
Keywords: Argyrodes, kleptoparasitism, maternal care, social behavior
Permanent sociality, or quasi-sociality, is tism by solitary spiders: the eresids
known in only about 20 of the more than Stegodyphus africanus (Blackwall 1866) and
&
37,000 described species of spiders, but based S. sabulosus Tullgren 1910 (Wickler Seibt
on the current classification, this represents at 1988), the salticids Simaetha paetula (Key-
least 12-16 independent origins of sociality serling 1882) (Jackson 1985) and several spe-
&
(Aviles 1997). Living in a group is atypical cies of Portia Karsch 1878 (Jackson Blest
spider behavior; spiders are famously solitary. 1982), the sparassid genus Olios Walckenaer
In most species even conspecifics are only tol- 1837 (Jackson 1987) [note: Elgar (1993) pre-
erated while mating (although aggression even sumably misread Jackson’s paper and indi-
here is common) and as very young juveniles. cates that these occur in groups], the symphy-
&
Living in a group therefore requires overcom- tognathid Curimagua bayano Forster
ing this kind of innate aggression. Many au- Platnick 1977 (Vollrath 1978), the uloborid
thors have pointed out that sociality in spiders Philoponella republicana (Simon 1891)
is not randomly distributed but rather concen- (Struhsaker 1969), and probably the oonopid
trated in a few lineages (Shear 1970; Burgess Oonops pulcher Templeton 1835 (Bristowe
1978; Krafft 1979; Buskirk 1981; D’Andrea 1958). In some cases, however, many conspe-
&
1987; Kraus Kraus 1988, 1990; Aviles cific kleptoparasites (or even a number of spe-
1997). Presumably such lineages may exhibit cies of spiders and non-spider arthropods
traits that facilitate sociality, or, in other (Eberhard et al. 1993)) occupy the same host
words, predispose the spiders to group living. web; examples are Sofanapis antillanca Plat-
& &
Spider kleptoparasitism, the occupation of nick Forster 1989 (Anapidae) (Ramirez
a heterospecific web to steal prey or silk, oc- Platnick 1999), Archaeodictyna ulova Gris-
&
curs in several families (Anapidae, Dictyni- wold Meikle-Griswold 1987 (Dictynidae)
&
dae, Eresidae, Sparassidae, Mysmenidae, (Griswold Meikle-Griswold 1987), Isela
Oonopidae, Salticidae, Symphytognathidae, okuncana Griswold 1985 (Mysmenidae) and
Theridiidae and Uloboridae) (Struhsaker many other mysmenids (Griswold 1985; Coy-
& &
1969; Wickler Seibt 1988; Elgar 1993; Ra- le Meigs 1989) in addition to Argyrodes
&
mirez Platnick 1999). Most of these in- Simon 1864 (Theridiidae) (e.g. Vollrath
stances represent opportunistic kleptoparasi- 1987)). These animals interact frequently and
181
THE JOURNAL OF ARACHNOLOGY
182
from maternal care; third, I discuss how the
notion of web sharing implies a clear distinc-
tion between territorial and non-territorial so-
ciality.
Although no detailed phylogeny of the fam-
ily Theridiidae has ever been published, Ag-
narsson et al. (2001), and Arnedo et al. (2001)
presented a preliminary theridiid phylogeny of
74 taxa at the XI Congress of Arachnology in
Badplaas, South Africa, based on morpholog-
ical and molecular data. This phylogenetic
analysis will be published elsewhere, but Fig.
2 summarizes the clades relevant to this ar-
gument, based on a combined analysis of the
morphological and molecular data. This ar-
rangement has been consistently found in pre-
vious analyses. This phylogeny is included
here to illustrate the argument; this admittedly
speculative hypothesis does not require this
—
Figure 1 . Communal feeding of several individ- particular cladistic structure to be valid.
uals (arrows) of two Argyrodes species with their All social theridiids occur in a relatively
host, Nephila clavipes. distal part of the cladogram, Anelosimus plus
the ‘Tost colulus clade” (Fig. 2). Both the cur-
rent classification and the provisional phylog-
may even feed together (Fig. 1, see also Rob- eny suggest at least four origins of sociality
&
inson Robinson 1973). As in sociality, such in this clade, within the genera Theridion Wal-
gregariousness and communal feeding must ckenaer 1805, Achaearanea Strand 1929 and
require diminished agonism. Anelosimus. The predominance of solitary
Theridiids are behaviorally diverse, ranging Theridion and Achaearanea species makes in-
from solitary web-less hunters (e.g. species of dependent origin in these lineages very prob-
the genus Dipoena Thorell 1869 (Levi 1953)), able. Cladistic analysis further suggests a dual
to species in which thousands of individuals origin of sociality in Anelosimus (Fig. 2). Al-
cooperate to build webs several cubic meters though not all Argyrodes are kleptoparasites,
in size e.g., Anelosimus eximius (Keyserling the behavior seems to have arisen once in the
1884). Most social spider species are theri- common ancestor of the whole lineage (see
diids (Aviles 1997) as are the equally famous Whitehouse et al. this volume). Argyrodes is
kleptoparasites, Argyrodes. These are the most sister to the Anelosimus plus lost colulus
conspicuous spider kleptoparasites, found clade. Thus all hve instances of group living,
worldwide in the webs of Nephila Leach or sharing a web, seem to be juxtaposed in
1815, and numerous other spiders. Although theridiid phylogeny. Does this distribution re-
both sociality and most instances of klepto- quire explanation, or is it simply coincidental?
parasitism typically involve web sharing (i.e., The definitive answer to this question must
the presence of more than one individual of await the finished phylogenetic product. Mad-
—
the same species in a single web excluding, dison (1990) proposed a “concentrated chang-
of course, mating pairs) the two behaviors es test” to calculate the probability of obtain-
X
have hitherto been considered entirely unre- ing, by chance alone, independent events in
lated (but see Whitehouse & Jackson 1993). a subclade of size F in a phylogeny of Z ter-
Recent phylogenetic research suggests other- minals. Given the preliminary nature of both
wise. In this paper three main points are made: the phylogenetic and natural history data cur-
hrst, I point out that the two types of behav- rently available, a calculation based on such
iors appear phylogenetically juxtaposed in data certainly cannot be definitive. However,
theridiid spiders; second I suggest that this even a marginally significant result at this
may be due to a fundamental similarity be- point would support the notion that something
tween the two, namely web sharing, stemming unusual has occurred in this branch of theri-
AGNARSSON—SOCIALITY AND KLEPTOPARASITISM 183
Outgroups
Outgroups
Other theridiids (e.g.
Dipoena)
Other theridiids (e.g,
Robertus)
Argyrodes
Anelosimus
“Lost colulus clade’
—
Figure 2. Major theridiid clades ex Agnarsson et al. (2001) and Arnedo et al. (2001). Triangles rep-
resent subsidiary lineages, bold branches are group-living taxa, dotted branches unknown behavior. Web
sharing (sociality plus kleptoparasitism) is notably clumped in the distal part of the cladogram suggesting
common
a cause.
diids, and therefore justifies some conjecture Many authors have argued that features re-
as to its genesis. In fact, the test says that so- quired for maternal care can result in post-
ciality (four occurrences in the Anelosimus juvenile tolerance and sociality if siblings
plus lost colulus clade) is significantly con- continue to cohabit after leaving the egg sac
centrated, a fact that in and by itself calls for (Shear 1970; Kullmann 1972; Burgess 1978;
explanation. In addition, the juxtaposition of Krafft 1979; Cangialosi & Uetz 1987, Aviles
sociality plus kleptoparasitism (five occur- 1997). It should be pointed out here that ma-
rences in the Argyrodes plus Anelosimus plus ternal care is itself a vaguely defined phenom-
lost colulus clade) is significant. Of course, if enon that may require more than one behav-
the number of independent instances were to
ioral mechanism (Krafft 1982). The most
decrease in the final analysis the distribution
obvious form of tolerance, and the one that
might be attributed to chance. Likewise, aug-
authors presumably have in mind when they
menting the overall size of the cladogram
would increase significance. As an increase in speak of the precursor to social behavior, is
tolerance between peers. “Peer tolerance”
the size of the theridiid cladogram is much
more certain than the discovery of novel in- among juvenile spiders is very widespread
phylogenetically; most newly emerged spider-
stances of sociality or kleptoparasitism, it
seems safe to conclude that their cladistic jux- lings “tolerate” each other at least briefly
taposition demands explanation if, as is ar- (Krafft 1982). “Offspring tolerance” requires
gued belov^, sociality and kleptoparasitism are the inhibition of predation on smaller individ-
fundamentally similar. I argue that kleptopar- uals, or at least on the egg sac. Finally, small
asitism and sociality are both more likely to spiders quite generally flee their webs if
evolve in the presence of a trait that reduces threatened by a large animal. Maternal care
agonism. may additionally require inhibition of the
THE JOURNAL OF ARACHNOLOGY
184
flight response, or “maternal tolerance” on Argyrodes has other characteristics of coop-
the part of the juveniles. erative living. Many Argyrodes moving on the
&
Kraus Kraus (1990) suggested that neo- host web will produce vibratory signals from
tenic retention ofJuvenile tolerance “obvious- numerous directions, which may confuse the
ly” explained sociality in the eresid genus Ste- host and can be considered a form of coop-
godyphus. For example, they explained the eration. Such effects have been identified as
smaller size of social Stegodyphus species in benefits of sociality (Allee 1931).
comparison to their solitary congenerics as As Aviles (1997) pointed out, the hypoth-
neoteny. In a similar manner Whitehouse esis that sociality evolved from maternal care
(1986) suggested that kleptoparasitism might predicts that sociality should be concentrated
have arisen through neoteny as an extension in lineages already exhibiting maternal care
of a fundamental “feeding with host” re- and that maternal care should precede social-
sponse of juveniles. Elgar (1993) showed that ity phylogenetically. Although more phylo-
obligatory kleptoparasitic (and thus group liv- genetic and natural history information is re-
ing) Argyrodes are smaller than their free liv- quired for a strong test, current evidence does
ing/opportunistic kleptoparasitic congenerics,
suggest that sociality in theridiid spiders is in-
&
but did not claim neoteny. Both Kraus
deed concentrated in a lineage where maternal
Kraus (1990) and Elgar (1993) treated taxa as care is common. Maternal care may be nec-
statistically independent units, which of
essary, but it certainly is not sufficient because
course inflates sample size and the potential it occurs in many solitary spider lineages (e.g.
for Type I errors, due to the failure to account Araneidae (Patel & Nigram 1991), Lycosidae
for shared history (Ridley 1983; Eelsenstein
(Eason 1964; Eujii 1979), Oxyopidae (Randall
1985). Thus the trends on which they based &
1977; Willey Adler 1989), Thomisidae
their conclusions should be re-analyzed. &
(Castanho Oliveira 1997; Evans 1998), and
In this view, sociality is homologous to the Uloboridae (Patel & Bradoo 1981) to name
juvenile tolerance exhibited in cases of ma-
but a few. Aviles (1997) pointed out that all
ternal care. Offspring tolerance (minimally as
but one of the social spiders belong to the in-
egg sac guarding), maternal tolerance and peer
fraorder Araneomorphae, but this is not sur-
tolerance seem all necessary prerequisites for
prising as Araneomorphae comprises the vast
maternal care but the prolongation ofjuvenile
peer tolerance may alone explain sociality, as majority of all spiders. However, she and oth-
ers have also pointed out that most social spi-
tolerant juveniles become tolerant adults. &
ders build webs (Shear 1970; Krafft 1979
Could kleptoparasitism also be modified
1982; Buskirk 1981; D’Andrea 1987), while
maternal care? The latter, from the juvenile’s
point of view, also means living and feeding about half of all spider families (and the ma-
in the web of a much bigger spider (the moth- jority of spider species) do not build prey cap-
er) who at some point becomes a potential ture webs. The importance of webs may lie
predator (Whitehouse 1986). If sociality is the with the silk itself, e.g. Krafft (1982) likened
prolongation of the tolerance required for ma- spider silk to the social pheromones of insects
because the vibratory information allows
ternal care, kleptoparasitism can be viewed as
communication from a distance. Krafft (1979)
co-opting or the exaptive application of ju-
venile tolerance in a novel context in which further suggested that sociality is relatively
the much larger host is no longer a conspecific more common in species that build three-di-
relative but an entirely different species. As in mensional webs than in orb weavers. Krafft,
maternal care, the kleptoparasites (both adults Buskirk, and D’Andrea attributed the relative
and juveniles) occur in groups that feed to- absence of sociality in orb weavers to the dif-
gether with the host. Interestingly, Argyrodes ficulty of cooperatively building and using an
&
kleptoparasites exhibit a wide range of “tol- orb-web (see also Cangialosi Uetz 1987).
erance” behaviors, from obligatory kleptopar- (Note: from the perspectives of Argyrodes
asitism in which adults tolerate the juveniles kleptoparasites, Nephila host orbs with their
and vice versa to facultative kleptoparasites extensive barrier webs are effectively three di-
that abandon their eggsacs and presumably are mensional). Theridiid web architecture varies
opportunistically cannibalistic (Larcher & greatly; some theridiids do not even make
Wise 1985). In addition, kleptoparasitism in webs. Theridiid sociality, however, occurs
AGNARSSON—SOCIALITY AND KLEPTOPARASITISM 185
only among lineages where three dimensional ternal care will be common and widespread
webs are prevalent. within this particular clade of theridiid spi-
But is the concurrence of maternal care and ders; at present such data are lacking for most
three dimensional webs sufficient to explain theridiid species. Finally the ideas presented
why members of this particular subset of ther- here may also apply to cases of social web
idiids are so prone to web sharing? Other spi- sharing in unrelated spider lineages and can
ders beside this clade of theridiids have three be tested there.
dimensional webs and maternal care but lack The argument followed in this paper sug-
further web sharing. Either web sharing sim- gests that spider “sociality” that consists of
ply has not evolved in these groups despite sharing a web may be fundamentally different
propitious conditions, or other unknown fac- from spider “sociality” that consists of tightly
tors are involved, for example peer tolerance. aggregated individual webs. “Web sharing”
Peer tolerance typically entails suppression of means two or more conspecifics in a single
both hetero- and conspecific aggression. In a web. Whether permanent or periodic, sociality
shared web, vibrations caused by conspecifics and communal kleptoparasitism are web shar-
and even quite unrelated spider species walk- ing, and all might have arisen from maternal
ing around are generally ignored (Krafft care (often labeled the “maternal care route”
1982), although struggling prey, of course, to sociality). In contrast, territorial “sociali-
trigger aggressive responses. Direct touch, ty,” the tight concentrations of interconnected
however, typically triggers aggression towards webs, differs distinctly. Known mainly from
heterospecific spiders only. In Agelena con- orb weavers such as Cyclosa Menge 1866,
sociata Denis 1965 a pheromone on the integ- Cyrtophora Simon 1864, Nephila, and Mete-
ument seems responsible for the inhibition of peira FO.R-Cambridge 1903 (Burgess 1978;
&
mutual biting (Krafft 1975). Very little is Krafft 1982; Aviles 1997; Burgess Witt
& &
known about spider pheromones and their 1976; Burgess Uetz 1982; Cangialosi
possible role in social communication (see Ti- Uetz 1987), in these cases each individual
etjen & Rovner 1982, for review). Mutual maintains its territory (i.e., its own web) and
feeding leads to exchange of digestive fluids communal feeding does not occur. Although
that may contain pheromones (Krafft 1969). such aggregations have been described as pos-
Kullmann (1972) suggested that contact che- sibly “a colonial social organization interme-
moreceptors in Stegodyphus sarasinorum diate between the solitary behavior typical of
might function to receive colony pheromones. most spiders and the communal behavior of
Such pheromones may be laid down on webs; the “social” spiders” (Cangialosi & Uetz
recently sex pheromones have been isolated 1987 p. 237), it is not likely to be intermediate
&
from spider webs (Schulz Toft 1993). The between solitarity and web-sharing sociality
study of such tolerance and communication because it never involves web sharing. Terri-
mechanisms may cast additional light on pat- torial sociality in the same genus may also be
terns of social and kleptoparasitic web-shar- periodic or permanent. Metepeira labrynthea
ing, and the relationship between the two. (Hentz 1847) aggregates facultatively around
The ideas presented in this paper can be resources, (called “fortuitous aggregations”
&
tested in several ways. First, the current phy- by Buskirk Uetz 1982), but M. spinipes
logeny juxtaposes kleptoparasitism and soci- EO.R-Cambridge 1903 forms permanent ter-
ality; if the phylogeny changes in such a way ritorial colonies. Thus territorial “sociality”
that the two are no longer adjacent, behavioral rather, as suggested by Burgess (1978), rep-
homology is falsified, although maternal care resents communal living that may arise
might still coincide with each independently. through a different evolutionary pathway, that
Second, if maternal care does not evolve at could be termed the “web aggregations
the node subtending kleptoparasitism and so- route.”
ciality in theridiids, the hypothesis of common To conclude, I suggest that sociality and
cause is suspect. Third, the hypothesis would kleptoparasitism in theridiids can both be
be strongly corroborated if an increase in the viewed as forms of web-sharing social behav-
length of time juveniles spend in their natal ior. The origin of both may have its roots in
web evolves at the kleptoparasitism-sociality maternal care, via the retention of juvenile
node. Fourth, this hypothesis predicts that ma- peer tolerance, sharing a three dimensional
186 THE JOURNAL OF ARACHNOLOGY
web. Web-sharing sociality and territorial so- strategies in spiders. Pp. 317-351. In Spider
ciality seem unrelated. communication, mechanisms and ecological sig-
&
nificance (Witt, P.N. J.S. Rovner, eds.) Prince-
ACKNOWLEDGMENTS
ton University Press, Princeton,
&
I would like to thank Jonathan A. Codding- Burgess, J.W. P.N. Witt. 1976. Spider webs: de-
ton for many fruitful discussions on the sub- sign and engineering. Interdisciplinary Science
Reviews 1:322-355.
ject and his numerous suggestions, which
Buskirk, R.E. 1981. Sociality in the Arachnida. Pp.
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