Web Relocation and Habitat Selection in Desert Widow Spider

Author(s): Yael Lubin, Stephen Ellner and Mandy Kotzman

Source: Ecology, Vol. 74, No. 7 (Oct., 1993), pp. 1915-1928
Published by: Ecological Society of America
Stable URL: http://www.jstor.org/stable/1940835
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Ecology, 74(7), 1993, pp. 1915-1928
? 1993 by the Ecological Society of America

WEB RELOCATION AND HABITAT SELECTION IN A

DESERT WIDOW SPIDER'

YAEL LUBIN
Mitrani Center for Desert Ecology, Jacob Blaustein Institute for Desert Research,
Ben Gurion University of the Negev, Sede Boqer 84990, Israel

STEPHEN ELLNER
Biomathematics Program, Department of Statistics, North Carolina State University,
Raleigh, North Carolina 27695, USA

MANDY KOTZMAN
Mitrani Center for Desert Ecology, Jacob Blaustein Institute for Desert Research,
Ben Gurion University of the Negev, Sede Boqer 84990, Israel

Abstract. We investigated web relocation and habitat selection in Latrodectus reviven-

sis (Theridiidae), a widow spider that constructs webs on scattered shrubs in the Negev
desert. We used repeated observations of individual spiders' habitat preferences, move-
ments to new web sites, growth, fecundity, and survival under natural and manipulated
habitat conditions to examine relationships between movement, habitat, and components
of fitness. Our main objectives were to determine the spatial and temporal patterns of
movement, and the causes, costs, and fitness consequences of shifting habitat.
Repeated censuses of individually marked females showed that web relocation in L.
revivensis is associated with spider growth. Spiders tended to move to larger shrubs after
one or two molts, and the size of web components and web-site features scaled to body
size. Moves occurred mainly in Spring (March-May), and most moves (74%) were by
juveniles. The main cost of web relocation was increased mortality: there was a 40% chance
of death during a move. A comparison of occupied shrubs with those available in the
habitat indicated strong site selection involving seasonally varying and age-dependent
preferences for particular shrub species, and a general preference for taller shrubs.
Prey availability at shrubs (as measured by pitfall traps, and by prey remains in nests)
varied spatially and was affected by shrub species and size. However these effects were
small compared to habitat-wide temporal variation in prey availability, suggesting that
web-site relocations would not result in substantially higher prey capture. Nonetheless,
movement to larger shrubs resulted in improved spider body condition and, ultimately,
greater reproductive success. The advantages of web-site relocation in this species may
relate to architectural features of shrubs that provide support for the web, and minimize
exposure to thermal extremes and to predators. However, experimental manipulation of
exposure by trimming shrubs did not decrease spider fitness. We suggest that web-site
selection in this species is less precise than expected because of the risk of dying during a
move.
Key words: desert spider; dispersal; habitat selection; Latrodectus revivensis; Negev desert; web
relocation.

INTRODUCTION
Habitat selection has received less empirical and the-
oretical attention than foraging decisions, although it
may be regarded as a branch of optimal foraging theory
(Rosenzweig 1985, Orians 1991). One reason for this
apparent neglect (Pulliam 1989) is the difficulty of re-
lating habitat-selection decisions with particular fitness
consequences in natural settings. These difficulties arise
because decisions regarding movement to a new hab-
itat are made over the same time scale as changes in
I Manuscript received 9 March 1992; revised 20 July 1992;
accepted 17 December 1992; final version received 19 Feb-
ruary 1993.
the habitat itself, and affect both current survival prob-
ability and future prospects for survival and repro-
duction. Consequently we lack a general, empirically
applicable framework or "currency" for assessing the
long-term costs and lifetime fitness benefits of move-
ment to a new habitat. Short-term gains (e.g., food
intake) are often used as measures of fitness, and a
direct relationship is assumed to exist between this
readily measured variable and fitness. Nonetheless, this
assumption is rarely tested under field conditions
(Morse and Fritz 1987, Ellner and Real 1989).
Moves to new habitats are often associated with
changes in habitat requirements with age of the organ-
ism. Such "ontogenetic niche shifts" (Werner and Gil-

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1916 YAEL LUBIN ET AL.
liam 1984) are caused by factors that relate to changing
body size or morphology, and may occur once or sev-
eral times within the life-span of an organism. Shachak
and Brand (1983) coined the term "recurrent dispersal"
to describe habitat shifts that occur occasionally
throughout the lifetime of a sit-and-wait predator. They
suggested that recurrent dispersal enables normally
sedentary predators to respond to changes in habitat
quality.
Many species of web-building spiders relocate their
webs at intervals throughout their lifetimes (Janetos
1982b). Their recurrent habitat shifts may be in re-
sponse to local variations in food supply (e.g., Gillespie
1981, Olive 1982, Vollrath 1985), disturbance (Hodge
1987b), the availability of suitable supporting struc-
tures for the web (Enders 1973, Hodge 1987a), or
changes in microclimate of the web site (Riechert and
Tracy 1975, Biere and Uetz 1981).
This paper reports an empirical study of recurrent
dispersal in a desert widow spider, Latrodectus reviven-
sis Shulov (Theridiidae). L. revivensis builds its webs
on scattered shrubs, which serve as foci for arthropod
activity (potential prey) on sparsely vegetated slopes
in the Negev desert. After the initial dispersal of young
away from the egg sac, spiders shift to new web sites
as they grow (Zilberberg 1988).
Habitat selection in L. revivensis is the choice of a
suitable shrub or type of shrub (species, size, location)
in which to build a web. An important component of
the habitat-selection problem for a web-building spider
is to assess its current situation relative to the expected
outcome of web relocation. Prior information about
the distribution of resources in the environment can
increase an organism's expectation of growth, survival,
and reproduction (Cohen 1967). For relatively sed-
entary web-building spiders, the ability to sample dif-
ferent habitats in order to obtain this information is
limited. The lack of information becomes more re-
stricting in environments such as deserts, where the
temporal or spatial distribution of critical resources
may vary greatly (Louw and Seely 1982). Vollrath (1985)
and Vollrath and Houston (1986) consider site tenacity
in some web-building spiders to reflect, in part, this
lack of information about alternative web sites. Models
for habitat selection generally make unrealistic sim-
plifying assumptions about the information available
to an organism about alternative habitats. For example
the "Ideal Free Distribution" model (Fretwell 1972),
and subsequent extensions of that model (e.g., Rosen-
zweig 1981, Pimm et al. 1985, Sutherland and Parker
1985), assume that individuals have perfect knowledge
of all available habitats (but, see Bernstein et al. 1988,
1991).
High costs of moving to alternative web sites would
also favor infrequent moves (Bernstein et al. 1991).
Movements to new web sites may involve increased
risk of predation. For active foragers, predation risk
influences decisions about time of activity, duration,
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Ecology, Vol. 74, No. 7
movement between resource patches, and type of food
consumed (Lima and Dill 1990). Predation during dis-
persal is difficult to measure, and has not been ade-
quately studied in web-building spiders (Riechert and
Gillespie 1986). Web design may also act as a con-
straint on dispersal: complex structures that are rich
in silk (e.g, sheet webs, three-dimensional-space webs,
funnel webs) are less frequently renewed than two-di-
mensional orb webs, and may require more time and
energy to build (Tanaka 1989).
For L. revivensis many morphometric dimensions of
the web are tightly correlated with body size, indicating
that the web is modified continuously by the growing
spider (Lubin et al. 1991). Site features, however, could
be changed only by relocating the web. Since web-site
features were less strongly correlated with spider size
than were web dimensions, we hypothesized that spi-
ders moved only infrequently. This pattern might re-
sult from large energy costs or risks of moving, rela-
tively small expected benefits to moving, or both.
We examine here the spatial and temporal patterns
of web relocation in a population of L. revivensis, and
the causes, costs, and fitness consequences of shifting
habitat. The consequences of web relocation are as-
sessed on two time scales: the period of residence at
the web site, and the lifetime of the individual. For
individuals found at web sites of different quality, and
for spiders at web sites with experimentally modified
conditions, we compare the prey availability, growth,
survival, and lifetime reproductive success. We address
the questions: Why do spiders relocate their webs? What
do they gain or lose by doing so? What information is
available about the environment that could influence
movement decisions?
Natural history of L. revivensis
Latrodectus revivensis occurs in the central Negev
desert of Israel (Shulov 1948, Levy and Amitai 1983).
The spiders have an annual life cycle: adult females
are present from April to October, and produce egg
sacs during May-September. The young emerge in the
summer (June-October), or may overwinter inside the
egg sac to emerge in early spring. Thus, juveniles of
various stages are present throughout most of the year,
but adults are restricted to the summer months.
The web of L. revivensis consists of separate nest and
prey-capture components (Fig. 1; Szlep 1965, Lubin et
al. 1991). The cone-shaped nest is built in a shrub, at
about 2/3 shrub height, and is connected by bridge threads
to a platform located at some distance from the shrub.
An array of threads stretches from the platform to the
ground; these threads are sticky at their attachment to
the ground and function to trap terrestrial arthropods.
The nest is a complex structure. The lower, open end
of the cone is composed of loosely woven silk; the
upper part is densely woven and is covered externally
with debris (plant parts, stones, snail shells, snail feces
October 1993 HABITAT SELECTION IN A DESERT SPIDER 1917

FIG. 1. The capture web (A) and conical nest (B) of Latrodectus revivensis. Capture web components: bridge threads (b),
platform (p), capture threads (t) with sticky portions (s) near the attachment to the ground. Nest components: upper part
made of dense silk (d) and covered partially with debris (db), nest opening (o).

and prey exoskeletons). The debris layer sometimes
extends downward over the lower section of the nest.
Web repairs, prey-capture activities and movements
to new sites occur at night. During the day, the spider
generally remains concealed in the nest, which provides
shelter from predators and from direct insolation
(Konigswald et al. 1990). The main diurnal predators
on L. revivensis are birds, lizards, and mantids. Other
spiders (e.g., Gnaphosidae) attack and kill L. revivensis
females in the nest and also may feed on juveniles or
eggs inside the egg sacs.
METHODS
Study site
The study was conducted in 1988-1990 on the rocky
slopes of a dry watercourse (wadi) in the Haluqim Ridge,
near Sede Boqer (30'50'N, 34046'E) in the Negev Des-
ert highlands of Israel. The limestone slopes are sparse-
ly vegetated with dwarf shrubs (mainly Zygophyllum
dumosum, Artemisia herba-alba, Reaumuria negev-
ensis, Hammada scoparia and Noaea mucronata; Ev-
enari et al. 1982, Olsvig-Whittaker et al. 1983). These
shrubs, and some woody annuals, serve as web sites
for the widow spiders.
The study area included ;40 ha of both north-facing
and south-facing slopes of a small wadi. To reduce the
effects of habitat heterogeneity we restricted our ob-
servations to spiders found on the lower portion of the
slope, from the wadi edge to a rock outcropping about
50 m up the slope.
Available web sites
To determine if spiders select shrubs nonrandomly
by species or height, we compared shrubs occupied by
spiders with those available in the habitat. We sur-
veyed the perennial vegetation along linear transects,
each 570 m in length, at different distances from the
wadi bed. Point-quarter quadrats (Krebs 1989), situ-
ated at intervals of 5 m along each line, provided an
estimate of the relative abundance and distribution of
shrub heights of the different species along vertical and
horizontal gradients. In addition, an index of branch
density was obtained for different shrubs by counting
the number of branches that intercepted a string
stretched across the shrub at its maximum diameter
and at the average nest height (2/3 shrub height; Lubin
et al. 199 1).
The shrubs used as web sites were surveyed in 1988,
1989, and 1990 and the data were analyzed separately
for November-April (cool season) and May-October
(hot season). The mean monthly temperature in the
cool season was 13.40C (range of means: 9.8-1 8'C) and
in the hot season 23.40C (range 21.1-25.50C).
Marked spiders
We estimated the costs and benefits of moving to a
new web site by following marked spiders. In 1989 we

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1918 YAEL LUBIN ET AL.
marked all 65 juvenile and adult female spiders found
on the north-facing slope with dots of enamel paint on
their legs (tibiae) so that we could distinguish among
individuals. (Males were not studied because they have
shorter life-spans than females and they do not con-
struct webs as adults). The web sites were marked with
numbered flags. We revisited the webs every 2 d and
recorded the presence or absence of the spider, the
occurrence of males in or near the web, feeding activity,
and egg-sac production. We used a small mirror,
mounted on a long handle, to observe the spiders inside
the nest without disturbing them.
Web-site tenure was calculated as the average of the
maximum and minimum possible durations at a web
site (sometimes several days elapsed between a spider's
disappearance from one web site and relocation at an-
other). When a spider disappeared from the web, we
searched all shrubs within a radius of at least 50 m; if
the spider was not found within 2 wk, it was presumed
dead. As shrub cover was sparse, all shrubs could be
searched exhaustively.
Mortality could occur either in the web or during a
move. Mortality in the web was often obvious: dead
spiders were often found in the bottom of the nest;
gnaphosid spiders entered the nest through a small,
round hole in the top of the nest and remained to
consume the nest occupant over several days; birds
(e.g., shrikes) made large, irregular holes in the side of
the nest, and lizards or snakes made large holes in the
bottom of the nest.
In cases where the cause of mortality was unknown,
we narrowed the uncertainty as follows. Since females
did not move once egg sacs were produced (Y. Lubin,
personal observation), those disappearing while guard-
ing egg sacs were categorized as "dead on web." Ju-
veniles and adults lacking egg sacs, that disappeared
within 2 wk of a molt, with no evidence of predation,
were categorized as "died in moving." Some predation
mortality on the web may have been incorrectly di-
agnosed as "died in moving." However, even if we
assume that 50% of the disappearances diagnosed as
"died in moving" were actually due to predation, the
resulting error in our estimates of mortality due to
other causes is <0.02, and our conclusions are unaf-
fected (see Results: Mortality during web relocation).
We measured the spiders and their webs when first
found. We measured the spider's mass, total body
length, and the length of the tibia + patella combined
of one of the first pair of legs (henceforth, "tibpat").
Web measurements included total nest length, maxi-
mum nest diameter, and the lengths of the dense silk
and debris sections of the nest cone. Web sites were
characterized by shrub species and the following mor-
phometric parameters: height of the shrub, height of
the nest in the shrub, distance from the lower edge of
the nest to the capture platform, and height of the
capture platform. If a spider moved, the new nest and
web site were measured, and old nests were remeasured
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Ecology, Vol. 74, No. 7
when abandoned or when the spider died. Spiders that
molted were remeasured within a day or two of molting
and marked again (the paint marks are shed with the
exuviae). The number of times a spider molted at each
web site was determined from the number of exuviae
attached to the nest wall.
During the survey we found that marking sometimes
influenced the tendency of recently molted spiders to
abandon web sites. Nearly half of all observed web
relocations (45% of 88 moves) occurred within 1 wk
of molting. We compared the tenure of web sites for
11 newly marked and 11 unmarked (control) spiders
of the same size range (total length: 10-12 mm). Un-
marked spiders remained at web sites significantly lon-
ger (median = 62 d, range: 1-169) than marked ones
(median = 23 d, range: 5-99; Wilcoxon test, P = .02).
Therefore, web-site tenure may be underestimated for
the 1989 survey data of marked spiders.
Spider condition and reproductive success
The relative fitness of females residing at web sites
of different quality was assessed during 1987-1990.
Two measures of relative fitness were used: body con-
dition and reproductive success.
To obtain measures of spider condition, the mass,
total body length, maximum abdomen width, and leg
length (tibia + patella of leg I or leg IV) were measured.
Within an instar the lengths of leg segments do not
change appreciably due to the hard cuticle (Vollrath
1983). Body mass, abdomen width, and to some extent
body length do change with feeding (Vollrath 1987)
and with reproductive condition. Body mass and ab-
domen width are tightly correlated (r2 = 0.98 in log-
log linear regression, N = 21), so abdomen width was
used as an alternative measure of body mass, which is
more difficult to measure in the field. An index of spider
condition was initially based on the allometric rela-
tionship between leg length and average body length
for all spiders. The difference between a female's actual
body size, and the body size predicted from her leg
length, was used as the index of condition. Two simpler
indices of condition were also defined: (a) (body length*
abdomen width)'12/leg length, and (b) body length/ab-
domen width. These indices were tightly correlated
(pairwise r2s for linear regression all >0.8, for N = 50
unmarked adult females).
Reproductive success was estimated by counting the
number of egg sacs and eggs produced, decremented
by the number eaten or parasitized. Most eggs hatched
and the young emerged within 1 mo of laying. We
removed hatched sacs and used a dissecting micro-
scope to count the eggshells remaining after hatching.
An unidentified wasp often parasitized the egg sacs and
could be recognized by the presence of pupal cases.
Other predators of eggs and newly emerged young, such
as other spiders (Clubionidae and Gnaphosidae), left
characteristic holes in the egg sac (easily distinguishable
October 1993 HABITAT SELECTION IN A DESERT SPIDER
from spiderling emergence holes), and were sometimes
found in the sac or nearby in the nest.
Shrub height andfitness
To examine correlations between shrub height and
measures of spider fitness, in May 1990 we observed
50 unmarked spiders (48 adult females and two sub-
adults) that were not being used in other tests. We
measured shrub height and nest height at the start of
the observation period, and then spider size (body
length, abdomen width, and tibpat length of leg I), and
the number of egg sacs present in the nest were recorded
at 2-wk intervals. We estimated survival time (mini-
mum tenure time, in days), the total number of eggs
produced, the number of egg sacs parasitized or at-
tacked by predators, and the total number and mass
of prey consumed during tenure of each web site.
Shrub-trimming experiment
To determine if the variation in shrub height had
long-term effects on survival and reproduction, we se-
lected 32 spiders in Zygophyllum shrubs, measured the
web and web-site parameters listed above, and trimmed
the branches of half of the shrubs, chosen at random,
to the level of the top of the nest. Trimming was done
during the day to minimize disturbance to the spiders.
All spiders were mature females, with the exception of
two subadult females; eight had 1-2 egg sacs at the start
of the experiment, the remaining spiders had none. We
examined the webs at 2-wk intervals, noting the spi-
der's presence, the number of egg sacs, and signs of
predation or parasitism. After each spider died or dis-
appeared, we collected its nest, examined the prey re-
mains, and estimated the number of eggs and the
amount of predation or parasitism on them as de-
scribed above.
Prey availability
Prey availability at web sites was assessed indepen-
dently of the condition and fate of the spider occupying
it. Prey remains were analyzed from nests of marked
and unmarked spiders in 1989 (N = 65 nests) and from
nests of 50 spiders used in 1990 to examine the rela-
tionship between shrub size and spider fitness (see
above). We estimated the relative amount of prey con-
sumed from the remains of arthropods (exoskeletons)
attached to the outer walls of the nest. Each nest that
was abandoned was taken to the laboratory where we
counted the number of prey items and measured the
body length of each. The regression equation of Rogers
et al. (1976), suitable for a range of arthropod taxa,
was used to estimate total body mass of prey (in mil-
ligrams) at each web from prey body length in milli-
metres:
Mass = 0.0305 Length2-62.
We used pitfall traps to compare prey availability in
the vicinity of shrubs. Pitfall traps were appropriate in
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1919
this case, because L. revivensis captures terrestrial ar-
thropods almost exclusively. We stress, however, that
the traps were used only to compare arthropod abun-
dances among shrubs, not as a quantitative estimate
of actual prey capture rates.
The pitfall traps were plastic cups (depth and di-
ameter: 10 cm) sunk into the ground, one each on the
north, east, south, and west sides of shrubs not cur-
rently used as web sites. The cups were at approxi-
mately the distance from the shrub that the capture
web of L. revivensis would be found. At each of six
sites (separated by at least 50 m) along the north-facing
slope, traps were placed near one large Zygophyllum
dumosum shrub (>50 cm height), one small Z. du-
mosum, and one Artemisia herba-alba (both <40 cm).
Thus, there were a total of 72 traps at 18 shrubs. The
prey captured in each trap was recorded on each day
of 11 sessions consisting of 4-6 sequential sampling
days each. Sessions were started roughly every 2 wk
between 28 February and 20 August 1990. The taxon
and body length class (5-mm length classes) of each
arthropod found in a trap were recorded. The lengths
were converted to an estimate of prey mass using the
weight-length regression equation given above. In pre-
liminary trials (eight sampling days in 1989) we found
a significant correlation in prey capture among the four
traps at each shrub, so all analyses here are based on
the average prey capture per day for all traps at a shrub
(number of prey per trap, or total mass of prey per
trap).
Prey supplementation
To test the effect of food supply on the tendency to
shift web sites, we supplemented the natural prey of
25 unmarked spiders and compared web-site tenure
with that of 26 non-supplemented spiders. All spiders
were juvenile females 5.5-8.0 mm in body length and
were located in a 400-M2 area adjacent to the main
study area. Spiders were assigned randomly to the two
groups. Supplemented spiders received a mealworm
(Tenebrio monitor) every other day for 30 d. The webs
were examined every other day for presence or absence
of the occupant.
Data transformations
Data were transformed whenever necessary to achieve
an approximately normal distribution as required for
parametric statistical tests. Frequently a logarithmic
transformation was adequate; if not we used the ex-
tended Box-Cox family of power transformations y =
[(a + X)b - I /b for nonzero b, y = ln(a + x) for b =
0, with maximum likelihood estimates of a and b as-
suming a normal distribution of y. If the transformed
data did not have a nearly normal distribution, as in-
dicated by nearly linear normal probability plots, we
used nonparametric statistical methods.
1920 YAEL LUBIN ET AL.
80-
Z60-
0 the comparison was restricted to available shrubs larger
W 40- . 15 cm for Artemisia and Zygophyllum, 20 cm for Ham-
201
0 in the lower part of the hillside, where Zygophyllum
ZYGOPHYLLUM HAMMADA OTHER
ARTEMESIA NOAEA
FIG. 2. Distribution of resident juveniles (U) and adults
(M) in shrubs of different species (cool and hot seasons com-
bined) in relation to the frequency of occurrence of the shrubs
in the study area (El). Total numbers of shrubs sampled =
1668, resident juvenile spiders = 282, resident adult spiders
= 194.
RESULTS
Habitat selection: distribution of
occupied web sites
The distribution of spiders in the four main shrub
species was significantly different from expected based
on the occurrence of these shrubs on the hillside (Fig.
2) (X2 = 799.02, df = 3, P < .001, for juveniles and
adults combined). Zygophyllum and Hammada were
used more often, and Artemisia less often, than ex-
pected from their relative abundances on the slope.
During the cool season, spiders occupied predomi-
nantlyArtemisia, moving into Zygophyllum at the start
of the hot season (Fig. 3). This corresponds to the tran-
sition period in the population from juveniles to adults.
A smaller peak in the number of webs in Artemisia
occurred in midsummer (June-July), when newly
emerged young began dispersing.
Both spider age (juvenile vs. adult) and season (hot
vs. cold) influenced the distribution of webs among the
different shrub species, and the two factors are not
independent (for all years combined, X2 = 111.39, df
= 1, P < .001). To determine the effects of spider age
and season, we analyzed 1988 and 1989 data combined
and 1990 data separately, as there were significant dif-
ferences between years in the distribution of webs (X2
= 25.97, df= 4, P < .001); years 1988 and 1989 did
not differ significantly (X2 = 5.26, df = 4, P = .2) and
could be combined. There were significant differences
in shrub use by juveniles and adults in all years (1988-
1989: x2 = 17.306, df= 4, P= .002; 1990: x2 = 69.4,
df = 4, P < .001). There were also significant differ-
ences between cool and hot seasons in the use of dif-
ferent shrub species (1988-1989: x2 = 22.68, df = 4,
P < .001; 1990: X2 = 15.89, df = 4, P = .003).
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Ecology, Vol. 74, No. 7
Juveniles and adults in both seasons were found in
taller shrubs than expected (t tests with unequal vari-
ances, P < .001 for Artemisia, Zygophyllum, Ham-
mada, and Noaea). Similar results were obtained when
than the minimum size used by juvenile females during
the entire period of observation. The cut-off sizes were
mada, and 33 cm for Noaea. In most cases, spiders
built their webs more often in larger shrubs than ex-
pected, indicating that the preference for taller shrubs
is not simply avoidance of shrubs below some mini-
mum size.
Webs of Lactrodectus revivensis were concentrated
shrubs were most abundant. The 20 m wide strip of
vegetation bordering the edge of the wadi contained
88.2% of all Zygophyllum shrubs in the vegetation sur-
vey (5-m and 15-m transect lines). In the cool season
47.8% of all webs were found in this 20 m wide strip,
and 71.3% of all webs in the hot season.
Although the spiders were nonrandomly distributed
on the slope in a way that parallels the distribution of
certain shrubs, the choice of shrubs on a local scale
could still be random. We therefore tested for habitat
selection on the local scale by comparing occupied
shrubs with their nearest neighbors. Occupied shrubs
were larger on average than their nearest neighbors
(mean shrub heights = 43 cm and 29 cm, respectively;
t tests with unequal variances, P < .001), and larger
on average than their nearest neighbors of the same
species (mean = 37 cm, P = .04). Thus, on the scale
of neighboring shrubs, spiders built webs in larger-
than-average bushes.
Habitat selection: individual movements
Marked spiders in the 1989 census moved to new
web sites at a rate of 6.7% per census, or 3.35% per
100
Cl)
z 80 ,
0
> 60-
LU
IL
0CO/4''
0
0 Jan Feb Mar Apr May June
_:
July Aug
MONTHS
Zygophyllum-- Artemisia --- Hammada Noaea
FIG. 3. Monthly changes in the frequencyof occurrence
of spiders in four differentshrub species. Total number of
spiders = 447 (adults and juveniles combined).
October 1993 HABITAT SELECTION IN A DESERT SPIDER 1921

day. Juveniles moved more frequently than adults: web 0.8

relocation of juveniles was 9.3% per day, and 74% of A
0.8- a Juveniles
all moves were by juveniles. To determine web-site
tenure for the marked spiders, we used only those in-
dividuals for which the dates of arrival at the web-site
COO
1III
0.6 Adults
and departure or death were known (to an accuracy of Cd)
? 2 d). Average web-site tenure for juveniles was 17.9
? 11 d (mean ? 1 SD, N = 18) and for adults 44.1 + 0.4
129.0 d (N = 24). Most moves to new web sites oc-
U04
curred in March and April, tapering off in May and
June (Fig. 4A). Distances moved were generally short
(Fig. 4B). S0.2-
Most marked spiders in 1989 (71%) molted once
before moving, 20.3% molted twice, and 8.7% moved
without first undergoing a molt (N = 69 nests examined
for the presence of exuviae). We also examined nests 0-
Feb Mar Apr May June' July
of unmarked spiders in 1987 (N = 53) and in 1989 (N
= 22). There was no significant difference between the 25X
two years; overall, 48% of the nests had one molt and B
9.3% had two molts, while 42.7% had none.
The habitat shifts suggested by the population dis- 20 U Juveniles
tribution patterns of L. revivensis were confirmed by
web relocations of marked spiders. Shrub height and LIAdults
nest height increased significantly with moves to new 15-
web sites (Table 1), and spiders shifted more frequently
from Artemisia and from other shrubs to Zygophyllum ~Z10-
than the reverse (X2 = 10.08, df = 4, P < .05). Of 50
shrub transitions, 52% were into Zygophyllum and 26% z
into Artemisia. Further evidence for nonrandom shrub 5-
selection comes from comparing web-relocation and
nearest-neighbor distances. Marked spiders moved
considerably further between web sites (mean ? SD:
6.4 ? 7.0 m) than would be necessary if they only 00-5 10 15 20 25 30 35 40
moved to the nearest larger shrub (1.8 ? 1.4 m) or to Distance between websites (m)
the nearest shrub of the same species (1.1 ? 0.8 m). FIG. 4. Moves by markedspidersin 1989. (A) Movement
rate (total number of moves per total numberof spidersob-
Moreover, spiders that moved greater distances tended served).(B) Frequencydistributionof distancesbetweennew
to obtain larger shrubs (Kendall's r = 0.20, P = .03, N and old web sites on successfulmoves forjuvenilesand adults
= 48 moves). separately.
To determine how web dimensions changed between
successive web sites, we compared the last measure- ability of surviving a 2-d interval between censuses
ments made at the new and old web sites (Table 1). was 59% for spiders that abandoned their web sites (n
The height of the capture web and the length and max- = 88 moves), vs. 98% for spiders that remained at their
imum diameter of the nest increased significantly, as web sites (n = 1193 census dates of 65 spiders). Ju-
did all measures of spider size. Thus, as spiders grow veniles and adults were equally successful in reaching
they move into taller shrubs, in which they construct new sites (60.0% of 65 moves of juvenile spiders vs.
larger nests placed higher in the shrub. Spiders also 56.5% of 23 adult moves, G = 0.08, P > .1). Moves
grow and modify their webs during tenure at a web early in the season were more successful than later
site. There were significant increases in spider size and moves for all spiders: 71 % survived in March (n = 21
in all nest variables during the spiders' tenure of a web moves), 622%in April (n = 53), 31 % in May (n = 13),
site: spiders molted, increased in size, and increased and none in June (n = 1). Displacement by other in-
the size of the nest concomitantly (Table 2). The dis- dividuals of L. revivensis was probably not a major
tance from the nest to the capture web decreased sig- cause of movement or of mortality on the web. We
nificantly with tenure, as a result of the spider enlarging observed the displacement of one female by another
its nest in the direction of the capture web. on only two occasions (out of 88 web-relocation events).

Mortality during web relocation Shrub height and spiderfitness

Web relocation entails a high risk of death (Table Web relocations tended to result in spiders occu-
3). Overall, in the census of marked spiders the prob- pying taller shrubs. We examined correlations between

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1922 YAEL LUBIN ET AL. Ecology, Vol. 74, No. 7

TABLE 1. Changes in measurements of marked spiders and nests at successive web sites measured in two different ways: (a)
differences between the last measurements made at successive web sites and (b) the proportion of measurements showing
an increase from one web site to the next. N = Number of web-site relocations.

Change between web sites

Variable N a) Mean ? 1 SD b) Proportion of increases
Spider
Body length (mm) 27 2.13 ? 1.39* 0.96*
Tibpatt (mm) 41 0.96 ? 0.93* 0.59NS

Nest
Nest length (cm) 29 35.6 ? 28.8 * 0.93*
Dense-silk length (cm) 29 -1.9 ? 10.7 NS 0.48NS
Debris length (cm) 29 2.9 ? 13.3 NS 0.45NS
Max. nest diameter (cm) 29 10.9 ? 10.5 * 0.90*
Capture web
Distance (cm) 5 -7.0 ? 12.8 NS 0.2 INS
Height (cm) 4 4.0 ? 4.0 * 1.00*
Web site
Nest height (cm) 41 10.5 ? 11.2 * 0.83*
Shrub height (cm) 41 9.8 ? 21.3 * 0.7 1*
* Ho is rejected at the 5% level (two-tailed tests), for null hypotheses (a) mean = 0 (t test) and (b) (proportion of increases) 0.5
(z test). NS = not significant.
t Leg length, i.e., the length of the tibia + patella for one of the first pairs of legs.

shrub height and measures of spider fitness for 50 un-
marked females in 1990 and monitored the effects of
experimental manipulations of shrub height. For the
latter, the range of shrub heights was 32-85 cm (mean
= 59 cm, cv = 27%) and of nest heights 18-62
(mean = 33 cm, cv = 19%).
Fecundity (numbers of egg sacs and eggs) was sig-
nificantly correlated with shrub height, as were several
measures of spider size (body length, abdomen width,
and tibpat [the length of tibia plus patella for one of
the first pairs of legs]) that correlate with fecundity
(Table 4). However it is difficult to separate the effects
of shrub height per se, from indirect effects due to
spider size and survival time at a web site, which cor-
relate with both fecundity and shrub height (Table 4).
Survival time alone accounted for roughly half the vari-
ance in fecundity (r2 = 0.60 for no. of sacs, r2 = 0.45
for no. of eggs): females that survived longer at a web
site had more young.
cm To remove the effects of spider size, we used the
residuals from a regression of shrub height on spider
size (tibpat), instead of shrub height itself. This re-
defines a "large" shrub to be one that is larger than
average for the size of spider occupying it. These shrub
height residuals also correlated with fecundity (Table
4). Moreover, adding shrub-height residuals to the re-
gression of the number of egg sacs on survival time,
significantly increased the amount of variance ex-
plained (r2 = 0.636, F45 = 6.95, P < .025). Thus, it
appears that fecundity is influenced by shrub height in

TABLE 2. Changes in measurements of marked spiders, nests, and web sites during tenure of a web site, measured in two
different ways: (a) differences between last and first measurements taken at a web site, and (b) the proportion of measurements
showing an increase during tenure at the web site. N number of web sites measured. Hypothesis tests as in Table 1.

Change at web site

Variable N a) Mean ? 1 SD b) Proportion of increases
Spider
Body length (mm) 35 1.66 ? 1.48* 1.00*
Tibpatt (mm) 93 1.01 ? 0.97* 0.6 1*
Nest
Nest length (cm) 33 37.2 ? 30.3 * 0.88*
Dense-silk length (cm) 33 8.3 ? 7.4 * 0.85*
Debris length (cm) 33 15.1 + 14.8 * 0.45*
Max. nest diameter (cm) 32 5.6 ? 11.0 * 0.72*
Capture web
Distance (cm) 11 -3.6 ? 4.4 * -0.9*
Height (cm) 11 0.2 ? 4 .0 NS 0.36NS
* See first footnote of Table 1.
t Leg length (tibia + patella for one of the first pairs of legs).

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October 1993 HABITAT SELECTION IN A DESERT SPIDER 1923

TABLE 3. The fates of spiders that remained at web sites or
moved to new sites, based on the 1989 census of 65 marked
spiders. N = number of observation dates. Probabilities of
survival were estimated per 2-d census interval. Overall
probabilities of survival are estimated per census interval.
TABLE 5. Analysis of variance on the number and estimated
biomass of prey captured in pitfall traps (using SAS pro-
cedure GLM [SAS Institute 1989])*. ss= Type III sum of
squares. All main effects and significant interactions
(P < .05) are shown.

Juveniles Adults All spiders Source of

Fate (N= 689) (N= 579) (N= 1277) variation Ss df MS F P
Stayed and lived 0.895 0.931 0.911 A) Number of prey
Stayed and died 0.011 0.031 0.020 Session .846 10 .0846 10.75 <.001
Moved and lived 0.056 0.021 0.041 Date .321 42 .0076 3.69 <.001
Moved and died 0.037 0.017 0.028 Site .025 5 .0051 2.50 .030
Probability of surviving Shrub type .018 2 .0091 4.47 .012
On the web 0.987 0.968 0.978 Session x Site .176 44 .0040 1.98 <.00 1
Move to a new site 0.600 0.545 0.586 Model 1.397 103 .0136 6.69 <.001
Error 1.481 730 .0021
B) Estimated prey biomass
Session 6.758 10 .631 3.07 <.001
a way that is independent of the relationships among Date 9.358 42 .223 2.81 <.001
shrub height, spider size, and survival. Site 1.283 5 .257 3.24 .007
Shrub type .460 2 .229 2.88 .057
To determine if the variation in shrub height had long- Model 17.75 59 .301 3.79 <.001
term effects on survival and reproduction, we selected Error 61.356 774 .0793
32 spiders in Zygophyllum shrubs, measured the various * The data were power-transformed prior to analysis to
web and web-site parameters, and trimmed the branches achieve approximately normal distributions; however the
of half of the shrubs to the level of the top of the nest. maximum likelihood power transformations were practically
equivalent to a log transformation, y = log(1 + x) (r2 0.92
This manipulation in effect placed the spiders in shrubs for prey number, r2 = 0.98 for prey mass, between the power
that were too small for them, and in addition placed and log-transformed values of the data analyzed).
their nests at the wrong location in the shrub, i.e., at the
top, instead of the typical location near 2/3 of the shrub's
height (Lubin et al. 1991). sition of the nest in the shrub did not significantly affect
We found no difference in survival of spiders in the survival or reproduction in adult females.
control and trimmed groups, nor were there significant
differences in either the number of egg sacs produced Shrub structure
or the total number of eggs laid (mean ? SD: trimmed, Differences in shrub architecture, as well as height,
2.8 ? 1.9 egg sacs, 728 + 450 eggs; control, 2.3 + 1.6 may explain moves of L. revivensis to new web sites,
egg sacs, 574 + 360 eggs; P > .05). Changes over the if larger spiders require more space between branches
duration of the experiment in nest size and in the rel- to accommodate their larger nests within the shrub.
ative amounts of dense silk and debris did not differ We therefore compared the density of branches in Ar-
in the two groups (P > .05 for all web variables). The temisia and Zygophyllum at 2/3 shrub height (the av-
percentage of egg sacs preyed upon, parasitized, or con- erage height of nests in shrubs; Lubin et al. 1991). The
taining dry eggs was higher for spiders in the experi- branch density was significantly greater in Artemisia
mental group than in the control group (41.6% vs. (0.42 ? 0.09 branches/cm, [mean ? 1 SD] N = 20)
24.8%, respectively), but this difference was not sig- than in Zygophyllum (0.07 ? 0.04 branches/cm, N =
nificant (P > .05; t test on arcsine-transformed pro- 50), even when the difference in shrub height was taken
portions). Thus, changing the shrub height and the po- into account (ANCOVA, P < .001). For Zygophyllum,

TABLE 4. Correlations of survival time (number of days at a web site) and fecundity (number of egg sacs and number of
eggs) with shrub height (shrub ht.), spider size (tibpat), abdomen width (abd. width), body length, and condition ([body
length abdomen width]0.5/tibpat). Data are from unmarked females (1990, N 50 spiders).

Shrub ht.
Measure Shrub ht. residuals No. sacs No. eggs Survival
No. sacs 0.39* 0.38* ...
No. eggs 0.36* 0.33* 0.92*
Survival 0.18 0.26t 0.77* 0.67*
Tibpatt 0.28* -0.04 0.05 0.12 -0.18
Abd. width 0.24t 0.14 0.55* 0.63* 0.30*
Body length 0.25t 0.11 0.47* 0.57* 0.26t
Condition -0.01 0.17 0.48* 0.5 1* 0.44*
* P < .05 (two-tailed)
t .05 < P < .1 (two-tailed); all others NS. (Pearson's correlation coefficients, r, for untransformed variables).
t Leg length (tibia + patella for one of the first pairs of legs).

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1924 YAEL LUBIN ET AL.
there was a weak negative correlation (Spearman rank
correlation ra,=-0.28, P = .05) between shrub height
and branch density: larger shrubs had lower branch
density at the 2/3 shrub-height level. There was no cor-
relation between shrub height and branch density in
Artemisia. Thus, differences in shrub architecture may
explain moves from Artemisia to Zygophyllum, and
possibly moves from smaller to larger Zygophyllum
shrubs.
Prey availability
We used pitfall traps to compare the availability of
potential arthropod prey at Artemisia shrubs and at
large and small shrubs of Zygophyllum. To verify that
arthropods sampled by traps were representative of
prey taken by the spiders, we compared the distribution
of arthropod taxa in the traps and in prey remains
collected from 115 webs of L. revivensis in the study
area. The numbers of arthropods of the different taxa
were positively correlated for the two sources (Ken-
dall's T = 0.43, z = 2.65, P < .01, for 1951 prey items
and 13 040 arthropods from webs and traps, respec-
tively).
Arthropod captures in the traps were highly variable
over time (ANOVA, Table 5). The differences among
sites and shrub types were also statistically significant,
but temporal variability (Session and Date effects) ac-
counted for a much larger portion of the total vari-
ability.
The deviations of individual shrubs from the habi-
tat-wide overall trends (ANOVA main effects) were
analyzed for temporal and spatial correlations. Any
such correlations could be important for movement
decisions, because they could allow spiders to predict
prey availability at other sites "now" (spatial correla-
tion) or at the same site "later" (temporal autocorre-
lation). We report here only the results for the number
of prey captured; the results for prey mass are nearly
identical but are less meaningful due to the additional
errors introduced by estimating prey mass from the
body sizes of prey items.
We tested for spatial correlations both across and
within sampling days. For the former we used the pair-
wise cross-correlation coefficients between the time se-
ries for each shrub. The correlations were not signifi-
cantly higher for shrubs at the same site than for shrubs
at different sites, indicating a lack of spatial correlation
among nearby shrubs (r2 = 0.041 ? 0.046 [mean ? 1
SD], n = 18 same site; mean r2 = 0.033 + 0.047, n =
135 between sites).
To test for spatial correlations within sampling days,
we computed for each sampling day the F statistic from
a single-factor ANOVA with site as the treatment vari-
able. These F values were transformed (using the ap-
propriate F distribution) so that under the null hy-
pothesis of no spatial correlation the transformed values
would be independent normal variates with zero mean
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Ecology, Vol. 74, No. 7
and unit variance. For both prey number and prey mass
there were no significant departures from the normal
distribution (Kolmogorov-Smirnov D = 0.13, P = .44),
indicating an absence of positive or negative spatial
correlation among nearby shrubs, and no significant
autocorrelations (at a = .05).
We also found no evidence of temporal autocorre-
lation over intervals longer than a single day. Analyzing
each shrub separately, I/4 ? of the shrubs had a signif-
icant autocorrelation at a lag of 1 d (successive days,
or last day of session N vs. first day of session N + 1).
There were very few significant (P < .05) autocorre-
lations at higher lags up to 10 d (two sampling sessions),
well below the number expected by chance alone in
that many statistical tests. Analyzing all shrubs to-
gether, only the autocorrelation between successive days
was statistically significant (r = 0.21, P < .05).
Prey supply and spider fitness
We looked for correlations between prey supply and
spider fitness by (1) testing the response to artificially
increased prey supply, and (2) examining correlations
between spider fitness measures and the amount of prey
consumed, based on analyses of prey remains in the
nests.
Prey supplementation.-There were no differences
between fed and unfed spiders in the tendency to aban-
don web sites. Average web-site tenure was 9.4 ? 6.3
d (mean ? 1 SD) for the fed group and 9.5 ? 7.1 d for
the control group. Between days 8 and 10 of the ex-
periment, there was a bout of unseasonably hot weath-
er, which may have caused the considerable mortality
that occurred then in both groups equally.
Prey consumed and fitness measures. -Prey remains
in nests provided estimates of the total number and
mass of prey consumed during tenure at a web site,
and the rate of prey capture (number and mass of prey
per day) for spiders of known web-site tenure (1989
census data).
Total prey number and mass were correlated with
leg length (tibpat, the length of tibia plus patella for
one of the first pairs of legs) in the 1989 sample con-
taining juveniles and adults, but not in the 1990 sample
which was only adults (Table 6). Thus, as juveniles
grew and molted, prey consumption increased, but for
adults prey consumption was unrelated to leg length.
For a given leg length, however, the amount of prey
consumed had strong positive correlations with body
size and fecundity.
Total prey accumulation (number and mass) was
also correlated in both years with how long spiders
remained at web sites (Table 6). However, for marked
spiders with known web-site tenure, the rate of prey
capture (number per day) was negatively correlated
with tenure (the correlation with mass per day was also
negative but not statistically significant). Negative cor-
relations could result from (1) diminishing returns at
October 1993 HABITAT SELECTION IN A DESERT SPIDER 1925

TABLE 6. Correlation coefficients (r) of prey numbers and cies, which resulted in a nonrandom distribution of
mass (totals per web and prey per web per day) with body spiders in the study area. We found that the correlation
size (tibpat), shrub height, tenure or survival time at a web
site, and fecundity. Data are from marked juvenile and between spider size and shrub height, previously ob-
adult females (1989 census, N = 65 spiders) and unmarked served on a random sample of webs (Lubin et al. 1991),
adult females (1990, N = 50 spiders); all variables were log- held also for transitions between successive sites for
transformed. Significance levels indicated are as in Table 4. individually marked spiders. Spiders grew, abandoned
No. prey Mass No./day Mass/day
their webs, and moved to new shrubs that were gen-
erally larger than the ones occupied previously. In large
Juveniles and adults (1989) shrubs they built larger nests further from the ground,
Tibpatt 0.22* 0.63* 0.08 0.47*
Body length 0.12 0.60* 0.01 0.50* and with capture webs located at greater distances from
Shrub height -0.02 0.20t -0.03 0.30t the nests.
Tenure 0.65* 0.44* -0.45* -0.17 Larger webs (and larger nests) clearly require larger
No. sacs 0.30* 0.29* 0.06 0.16 supporting structures. Enders (1975) showed that the
Adults (1990) orb weaver, Argiope aurantia (Araneidae), moved up
Tibpatt 0.02 0.00 in the vegetation as it matured, where it built larger
Body length 0.25t 0.40*
Abdomen width 0.29* 0.48* webs in the larger gaps available there. Other studies,
Shrub height 0.19 0.21 however, have mostly ignored growth as a major factor
Survival 0.59* 0.59* in determining web relocation in spiders, concentrating
No. sacs 0.61* 0.82*
0.62* 0.79*
instead on changing food supply or physical factors
No. eggs
(reviewed by Riechert and Gillespie 1986). In L. re-
: Leg length (tibia + patella for one of the first pairs of legs).
vivensis the changing structural requirements of grow-
ing spiders may explain some, but not all, of web-site
selection behavior. Most individuals moved after each
web sites due to prey depletion or (2) spiders receiving molt, thus providing the positive correlations between
more food grew faster and molted sooner and therefore spider size, nest size, and shrub size (Lubin et al. 1991).
abandoned their web sites earlier. However, some spiders moved after two molts, and up
For adults there was only a weak positive correlation to 43% moved without undergoing a previous molt.
between shrub height and the total prey numbers and Furthermore, the seasonal differences observed in shrub
mass (Table 6). The relationship between shrub height preferences of these spiders cannot be explained strictly
and total prey consumption may simply reflect the fact on the basis of growth requirements. Large costs of
that shrub height and prey consumption are both pos- moving, or benefits unrelated to growth, were sus-
itively correlated with body length. pected to influence movement decisions in this species.

DISCUSSION Costs of web relocation

Habitat selection is assumed to have a direct and
strong impact on fitness (Orians 1991). For relatively
sedentary animals such as sit-and-wait predators or
animals that defend a fixed territory, good sites will
provide long-term benefits in terms of survival, growth,
and reproduction. Few studies, however, assess the
benefits and costs of habitat selection in terms of long-
term fitness consequences (Morse and Fritz 1987, Ward
and Lubin, in press). We focused here on web-site se-
lection and web-relocation movements in a desert spider
and asked: What does the spider gain by moving? What
are the costs? And, how do the benefits and costs of
movement in the short term relate to spider fitness in
the long term? The desert widow spider, Latrodectus
revivensis, seemed ideally suited for this purpose: it
shifts to new web sites at intervals throughout its life-
time and, owing to its large size and the openness of
the habitat it occupies, it could be marked and ob-
served over most of its annual life cycle.
Habitat selection, growth, and movement
Latrodectus revivensis showed a strong preference for
larger-than-average shrubs and for certain shrub spe-
Tanaka (1989) considered the energy required to build
a new web to be the major cost of web relocation for
a sheet-web spider (Agelena limbata, Agelenidae) found
in mesic habitats in Japan. Janetos (1 982a, b) suggested
that spiders with complex, energy-rich webs should
move less frequently, due to energy constraints, than
those with less complex webs. The movement rates for
different species tend to support this idea: they range
from 10-45%/d for orb and sheet-web builders with
relatively sparse webs (Eberhard 1971, Enders 1975,
Janetos 1982b) to 3.35%/d in L. revivensis and 0.3%/d
in A. limbata (Tanaka 1989), species with complex,
energy-rich webs.
In L. revivensis mortality was a major cost of web
relocation. On average, 40% of spiders abandoning a
web site were unsuccessful in reaching a new site. The
overall mortality rate in this study was 2.4% daily, 58%
of which occurred during moves to new web sites (Ta-
ble 3). The high mortality rate occurring during moves
suggests that predation may be the major risk of shift-
ing to a new web site in this species. Predation risk,
notoriously difficult to measure under natural condi-
tions, may be underestimated in other studies of web-

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1926 YAEL LUBIN ET AL.
site selection. Thus, while the energy cost of web build-
ing may contribute to mortality during a move, it is
unlikely to be the major cause of mortality.
Benefits of web relocation
In spite of the large risk, virtually all L. revivensis
females moved at least once during their lifetimes after
initial dispersal from the maternal nest. The question
is, then, what benefit is derived from recurrent dis-
persal movements?
For many mobile organisms, the primary benefit to
moving to a new site is the expectation of improved
food supply at the new site relative to the current one.
In contrast to previous studies on web-building spiders
(e.g., Vollrath 1985, Gillespie and Caraco 1987, but
see also Riechert and Tracy 1975), we showed this not
to be the case for L. revivensis. Movement to new sites
in this species was apparently unrelated to prey avail-
ability. The type of shrub occupied (Artemesia, large
or small Zygophyllum) explained only a small amount
of the variation in prey availability at web sites in
comparison with the habitat-wide temporal trends that
affected all web sites equally. The mean prey capture
rate at the best shrub type was higher than that at the
worst by only 10% for prey number, while the best and
worst trapping sessions differed by 275%. Web relo-
cation to a different shrub type in order to improve
prey capture would seem to be an expensive gamble
with little expected payoff. Furthermore, we found no
spatial correlation in prey availability, so a spider's
prey capture success at a particular web site does not
provide any information about the quality of other
sites. The lack of temporal and spatial correlations also
implies that a spider has no reason to abandon a site
in response to a period of poor prey capture.
The benefits of recurrent dispersal in L. revivensis
appear to be associated with improved shrub architec-
ture. A likely benefit is the improvement of thermal
conditions within the nest during the transition from
the cool to hot season, and throughout the hot season.
High daytime temperatures inside the nest force spi-
ders to adopt thermoregulatory positions that expose
them to visually orienting predators (e.g., birds and
mantids) (Konigswald et al. 1990; Y. Lubin, unpub-
lished data). Spiders in larger shrubs, and in shrubs
with more open branch structure (e.g., Zygophyllum),
will experience less extreme nest temperatures, because
air temperature decreases with increasing height above
the ground (Campbell 1977) and because nests in
sparsely branched shrubs will be more exposed to cool-
ing breezes than nests in dense shrubs.
Long-term consequences of habitat selection
The positive relationship observed between spider
fitness and shrub height was tested experimentally by
trimming shrubs to nest height. This treatment in-
creased the amount of solar radiation directly imping-
ing upon the nest, exposed the nest from above to
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Ecology, Vol. 74, No. 7
visually hunting predators, and reduced the number of
supporting structures above the web. Nonetheless, nei-
ther survival nor fecundity was reduced in the exper-
imental group.
The lack of any significant effect of exposure on spi-
der fitness could be explained if the experiments were
conducted during a period of cooler weather or when
the risk of predation was low. However, web relocation
occurs throughout the summer, including the time pe-
riod during which the manipulations were performed,
and temperatures were near average for that time of
year.
An alternative hypothesis is that web-site selection
in L. revivensis may be less precise than expected be-
cause of the risks involved in moving and the low
probability of finding a better web site. The decision
problem for a spider moving sequentially from shrub
to shrub in search of a new web site is similar to the
mate choice problem as modeled by Real (1990), or
the classical models of sequential job choice in eco-
nomics (Lipmann and McCall 1976). The optimal de-
cision rule for such sequential choice problems is typ-
ically a "satisficing" rule: the item (shrub, mate, job
offer) chosen is the first encountered that meets or ex-
ceeds a minimum criterion (Lipmann and McCall
1976). "Satisficing" criteria have been suggested to op-
erate in habitat selection decisions of other desert spi-
ders (Riechert 1985, Ward and Lubin 1993).
In L. revivensis, the search cost is high due to the
high risk of mortality during movement away from the
web, thus the range of acceptable items will be broad,
and will include items that are far from optimal (Lip-
mann and McCall 1976, Real 1990). High mortality
during moves may also increase the amount of time
an individual should remain in its current location,
tolerating poor or deteriorating conditions, before re-
locating (Houston and McNamara 1986, Gilliam 1990).
Consequently much of the life-span would be spent
"making do" in suboptimal sites. Natural selection
would then favor greater plasticity in habitat selection,
either by increasing physiological tolerance, or by be-
havioral responses to reduce the impact of a poorer
web site.
Conclusion: fitness measures for
habitat selection
We have shown here that habitat selection is bene-
ficial to the desert widow spider, L. revivensis. Spiders
that moved to "good" web sites (taller shrubs) fared
better in the long run, as measured by their growth and
reproductive success. As food was not the direct cause
of success at a web site, indirect measures of fitness
relating to food intake were not useful; the sole useful
measure was fitness itself. Similar examples may
abound, whenever site-selection criteria are based on
qualities other than food availability.
Mortality may occur both on the web and during
web relocation, but the probability of dying during a
October 1993 HABITAT SELECTION IN A DESERT SPIDER
move was 40% compared with <1% per day on the
web (Table 3). Current foraging theory generally treats
mortality risk as a feature of the habitat (Gilliam 1990,
Lima and Dill 1990). Here we have shown that the
risk of mortality may be more important as a com-
ponent of the move itself and will influence the decision
whether to move at all, rather than which habitat to
choose. For L. revivensis, it appears that the main cost
of moving is the decreased probability of survival, and
there are no indirect measures (e.g., energy expended,
reduced food intake) that will substitute for survival.
It remains paradoxical that experimental tests of the
relationship between the preference for habitat features
and fitness are inconclusive. "Satisficing" criteria for
web-site selection may partly explain the weak corre-
lation between fitness and prefered sites. Nonetheless,
the missing links between short-term advantages of
habitat selection and dispersal, and long-term fitness
consequences, bear further investigation.
ACKNOWLEDGMENTS
We are grateful to Helen Dallas, Arthur Dumosch, Iris Mus-
li, and Sonya Rosen who helped with various aspects of this
project, to Fiona Wierczk for Fig. 1, and to David Dickey for
statistical advice. David Ward provided help and advice
throughout the study. We thank James Gilliam, George Hess,
Leslie Real, and David Ward for critically reading the manu-
script. The study was supported by a U.S.-Israel Binational
Science Foundation grant #86-00092. This is contribution
Number 157 of the Mitrani Center for Desert Ecology.
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