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Lubin 1993
Lubin 1993
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Ecology, 74(7), 1993, pp. 1915-1928
? 1993 by the Ecological Society of America
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
INTRODUCTION
the habitat itself, and affect both current survival prob-
Habitat selection has received less empirical and the- ability and future prospects for survival and repro-
oretical attention than foraging decisions, although it duction. Consequently we lack a general, empirically
may be regarded as a branch of optimal foraging theory applicable framework or "currency" for assessing the
(Rosenzweig 1985, Orians 1991). One reason for this long-term costs and lifetime fitness benefits of move-
apparent neglect (Pulliam 1989) is the difficulty of re- ment to a new habitat. Short-term gains (e.g., food
lating habitat-selection decisions with particular fitness intake) are often used as measures of fitness, and a
consequences in natural settings. These difficulties arise direct relationship is assumed to exist between this
because decisions regarding movement to a new hab- readily measured variable and fitness. Nonetheless, this
itat are made over the same time scale as changes in assumption is rarely tested under field conditions
(Morse and Fritz 1987, Ellner and Real 1989).
I Manuscript received 9 March 1992; revised 20 July 1992;
Moves to new habitats are often associated with
changes in habitat requirements with age of the organ-
accepted 17 December 1992; final version received 19 Feb-
ruary 1993. ism. Such "ontogenetic niche shifts" (Werner and Gil-
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1916 YAEL LUBIN ET AL. Ecology, Vol. 74, No. 7
liam 1984) are caused by factors that relate to changing movement between resource patches, and type of food
body size or morphology, and may occur once or sev- consumed (Lima and Dill 1990). Predation during dis-
eral times within the life-span of an organism. Shachak persal is difficult to measure, and has not been ade-
and Brand (1983) coined the term "recurrent dispersal" quately studied in web-building spiders (Riechert and
to describe habitat shifts that occur occasionally Gillespie 1986). Web design may also act as a con-
throughout the lifetime of a sit-and-wait predator. They straint on dispersal: complex structures that are rich
suggested that recurrent dispersal enables normally in silk (e.g, sheet webs, three-dimensional-space webs,
sedentary predators to respond to changes in habitat funnel webs) are less frequently renewed than two-di-
quality. mensional orb webs, and may require more time and
Many species of web-building spiders relocate their energy to build (Tanaka 1989).
webs at intervals throughout their lifetimes (Janetos For L. revivensis many morphometric dimensions of
1982b). Their recurrent habitat shifts may be in re- the web are tightly correlated with body size, indicating
sponse to local variations in food supply (e.g., Gillespie that the web is modified continuously by the growing
1981, Olive 1982, Vollrath 1985), disturbance (Hodge spider (Lubin et al. 1991). Site features, however, could
1987b), the availability of suitable supporting struc- be changed only by relocating the web. Since web-site
tures for the web (Enders 1973, Hodge 1987a), or features were less strongly correlated with spider size
changes in microclimate of the web site (Riechert and than were web dimensions, we hypothesized that spi-
Tracy 1975, Biere and Uetz 1981). ders moved only infrequently. This pattern might re-
This paper reports an empirical study of recurrent sult from large energy costs or risks of moving, rela-
dispersal in a desert widow spider, Latrodectus reviven- tively small expected benefits to moving, or both.
sis Shulov (Theridiidae). L. revivensis builds its webs We examine here the spatial and temporal patterns
on scattered shrubs, which serve as foci for arthropod of web relocation in a population of L. revivensis, and
activity (potential prey) on sparsely vegetated slopes the causes, costs, and fitness consequences of shifting
in the Negev desert. After the initial dispersal of young habitat. The consequences of web relocation are as-
away from the egg sac, spiders shift to new web sites sessed on two time scales: the period of residence at
as they grow (Zilberberg 1988). the web site, and the lifetime of the individual. For
Habitat selection in L. revivensis is the choice of a individuals found at web sites of different quality, and
suitable shrub or type of shrub (species, size, location) for spiders at web sites with experimentally modified
in which to build a web. An important component of conditions, we compare the prey availability, growth,
the habitat-selection problem for a web-building spider survival, and lifetime reproductive success. We address
is to assess its current situation relative to the expected the questions: Why do spiders relocate their webs? What
outcome of web relocation. Prior information about do they gain or lose by doing so? What information is
the distribution of resources in the environment can available about the environment that could influence
increase an organism's expectation of growth, survival, movement decisions?
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 Natural history of L. revivensis
limited. The lack of information becomes more re- Latrodectus revivensis occurs in the central Negev
stricting in environments such as deserts, where the desert of Israel (Shulov 1948, Levy and Amitai 1983).
temporal or spatial distribution of critical resources The spiders have an annual life cycle: adult females
may vary greatly (Louw and Seely 1982). Vollrath (1985) are present from April to October, and produce egg
and Vollrath and Houston (1986) consider site tenacity sacs during May-September. The young emerge in the
in some web-building spiders to reflect, in part, this summer (June-October), or may overwinter inside the
lack of information about alternative web sites. Models egg sac to emerge in early spring. Thus, juveniles of
for habitat selection generally make unrealistic sim- various stages are present throughout most of the year,
plifying assumptions about the information available but adults are restricted to the summer months.
to an organism about alternative habitats. For example The web of L. revivensis consists of separate nest and
the "Ideal Free Distribution" model (Fretwell 1972), prey-capture components (Fig. 1; Szlep 1965, Lubin et
and subsequent extensions of that model (e.g., Rosen- al. 1991). The cone-shaped nest is built in a shrub, at
zweig 1981, Pimm et al. 1985, Sutherland and Parker about 2/3 shrub height, and is connected by bridge threads
1985), assume that individuals have perfect knowledge to a platform located at some distance from the shrub.
of all available habitats (but, see Bernstein et al. 1988, An array of threads stretches from the platform to the
1991). ground; these threads are sticky at their attachment to
High costs of moving to alternative web sites would the ground and function to trap terrestrial arthropods.
also favor infrequent moves (Bernstein et al. 1991). The nest is a complex structure. The lower, open end
Movements to new web sites may involve increased of the cone is composed of loosely woven silk; the
risk of predation. For active foragers, predation risk upper part is densely woven and is covered externally
influences decisions about time of activity, duration, with debris (plant parts, stones, snail shells, snail feces
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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 slope, from the wadi edge to a rock outcropping about
extends downward over the lower section of the nest. 50 m up the slope.
Web repairs, prey-capture activities and movements
to new sites occur at night. During the day, the spider Available web sites
generally remains concealed in the nest, which provides To determine if spiders select shrubs nonrandomly
shelter from predators and from direct insolation by species or height, we compared shrubs occupied by
(Konigswald et al. 1990). The main diurnal predators spiders with those available in the habitat. We sur-
on L. revivensis are birds, lizards, and mantids. Other veyed the perennial vegetation along linear transects,
spiders (e.g., Gnaphosidae) attack and kill L. revivensis each 570 m in length, at different distances from the
females in the nest and also may feed on juveniles or wadi bed. Point-quarter quadrats (Krebs 1989), situ-
eggs inside the egg sacs. ated at intervals of 5 m along each line, provided an
estimate of the relative abundance and distribution of
METHODS shrub heights of the different species along vertical and
horizontal gradients. In addition, an index of branch
Study site density was obtained for different shrubs by counting
The study was conducted in 1988-1990 on the rocky the number of branches that intercepted a string
slopes of a dry watercourse (wadi) in the Haluqim Ridge, stretched across the shrub at its maximum diameter
near Sede Boqer (30'50'N, 34046'E) in the Negev Des- and at the average nest height (2/3 shrub height; Lubin
ert highlands of Israel. The limestone slopes are sparse- et al. 199 1).
ly vegetated with dwarf shrubs (mainly Zygophyllum The shrubs used as web sites were surveyed in 1988,
dumosum, Artemisia herba-alba, Reaumuria negev- 1989, and 1990 and the data were analyzed separately
ensis, Hammada scoparia and Noaea mucronata; Ev- for November-April (cool season) and May-October
enari et al. 1982, Olsvig-Whittaker et al. 1983). These (hot season). The mean monthly temperature in the
shrubs, and some woody annuals, serve as web sites cool season was 13.40C (range of means: 9.8-1 8'C) and
for the widow spiders. in the hot season 23.40C (range 21.1-25.50C).
The study area included ;40 ha of both north-facing
and south-facing slopes of a small wadi. To reduce the Marked spiders
effects of habitat heterogeneity we restricted our ob- We estimated the costs and benefits of moving to a
servations to spiders found on the lower portion of the new web site by following marked spiders. In 1989 we
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1918 YAEL LUBIN ET AL. Ecology, Vol. 74, No. 7
marked all 65 juvenile and adult female spiders found when abandoned or when the spider died. Spiders that
on the north-facing slope with dots of enamel paint on molted were remeasured within a day or two of molting
their legs (tibiae) so that we could distinguish among and marked again (the paint marks are shed with the
individuals. (Males were not studied because they have exuviae). The number of times a spider molted at each
shorter life-spans than females and they do not con- web site was determined from the number of exuviae
struct webs as adults). The web sites were marked with attached to the nest wall.
numbered flags. We revisited the webs every 2 d and During the survey we found that marking sometimes
recorded the presence or absence of the spider, the influenced the tendency of recently molted spiders to
occurrence of males in or near the web, feeding activity, abandon web sites. Nearly half of all observed web
and egg-sac production. We used a small mirror, relocations (45% of 88 moves) occurred within 1 wk
mounted on a long handle, to observe the spiders inside of molting. We compared the tenure of web sites for
the nest without disturbing them. 11 newly marked and 11 unmarked (control) spiders
Web-site tenure was calculated as the average of the of the same size range (total length: 10-12 mm). Un-
maximum and minimum possible durations at a web marked spiders remained at web sites significantly lon-
site (sometimes several days elapsed between a spider's ger (median = 62 d, range: 1-169) than marked ones
disappearance from one web site and relocation at an- (median = 23 d, range: 5-99; Wilcoxon test, P = .02).
other). When a spider disappeared from the web, we Therefore, web-site tenure may be underestimated for
searched all shrubs within a radius of at least 50 m; if the 1989 survey data of marked spiders.
the spider was not found within 2 wk, it was presumed
dead. As shrub cover was sparse, all shrubs could be
searched exhaustively. Spider condition and reproductive success
Mortality could occur either in the web or during a The relative fitness of females residing at web sites
move. Mortality in the web was often obvious: dead of different quality was assessed during 1987-1990.
spiders were often found in the bottom of the nest; Two measures of relative fitness were used: body con-
gnaphosid spiders entered the nest through a small, dition and reproductive success.
round hole in the top of the nest and remained to To obtain measures of spider condition, the mass,
consume the nest occupant over several days; birds total body length, maximum abdomen width, and leg
(e.g., shrikes) made large, irregular holes in the side of length (tibia + patella of leg I or leg IV) were measured.
the nest, and lizards or snakes made large holes in the Within an instar the lengths of leg segments do not
bottom of the nest. change appreciably due to the hard cuticle (Vollrath
In cases where the cause of mortality was unknown, 1983). Body mass, abdomen width, and to some extent
we narrowed the uncertainty as follows. Since females body length do change with feeding (Vollrath 1987)
did not move once egg sacs were produced (Y. Lubin, and with reproductive condition. Body mass and ab-
personal observation), those disappearing while guard- domen width are tightly correlated (r2 = 0.98 in log-
ing egg sacs were categorized as "dead on web." Ju- log linear regression, N = 21), so abdomen width was
veniles and adults lacking egg sacs, that disappeared used as an alternative measure of body mass, which is
within 2 wk of a molt, with no evidence of predation, more difficult to measure in the field. An index of spider
were categorized as "died in moving." Some predation condition was initially based on the allometric rela-
mortality on the web may have been incorrectly di- tionship between leg length and average body length
agnosed as "died in moving." However, even if we for all spiders. The difference between a female's actual
assume that 50% of the disappearances diagnosed as body size, and the body size predicted from her leg
"died in moving" were actually due to predation, the length, was used as the index of condition. Two simpler
resulting error in our estimates of mortality due to indices of condition were also defined: (a) (body length*
other causes is <0.02, and our conclusions are unaf- abdomen width)'12/leg length, and (b) body length/ab-
fected (see Results: Mortality during web relocation). domen width. These indices were tightly correlated
We measured the spiders and their webs when first (pairwise r2s for linear regression all >0.8, for N = 50
found. We measured the spider's mass, total body unmarked adult females).
length, and the length of the tibia + patella combined Reproductive success was estimated by counting the
of one of the first pair of legs (henceforth, "tibpat"). number of egg sacs and eggs produced, decremented
Web measurements included total nest length, maxi- by the number eaten or parasitized. Most eggs hatched
mum nest diameter, and the lengths of the dense silk and the young emerged within 1 mo of laying. We
and debris sections of the nest cone. Web sites were removed hatched sacs and used a dissecting micro-
characterized by shrub species and the following mor- scope to count the eggshells remaining after hatching.
phometric parameters: height of the shrub, height of An unidentified wasp often parasitized the egg sacs and
the nest in the shrub, distance from the lower edge of could be recognized by the presence of pupal cases.
the nest to the capture platform, and height of the Other predators of eggs and newly emerged young, such
capture platform. If a spider moved, the new nest and as other spiders (Clubionidae and Gnaphosidae), left
web site were measured, and old nests were remeasured characteristic holes in the egg sac (easily distinguishable
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October 1993 HABITAT SELECTION IN A DESERT SPIDER 1919
from spiderling emergence holes), and were sometimes this case, because L. revivensis captures terrestrial ar-
found in the sac or nearby in the nest. thropods almost exclusively. We stress, however, that
the traps were used only to compare arthropod abun-
Shrub height andfitness dances among shrubs, not as a quantitative estimate
To examine correlations between shrub height and of actual prey capture rates.
measures of spider fitness, in May 1990 we observed The pitfall traps were plastic cups (depth and di-
50 unmarked spiders (48 adult females and two sub- ameter: 10 cm) sunk into the ground, one each on the
adults) that were not being used in other tests. We north, east, south, and west sides of shrubs not cur-
measured shrub height and nest height at the start of rently used as web sites. The cups were at approxi-
the observation period, and then spider size (body mately the distance from the shrub that the capture
length, abdomen width, and tibpat length of leg I), and web of L. revivensis would be found. At each of six
the number of egg sacs present in the nest were recorded sites (separated by at least 50 m) along the north-facing
at 2-wk intervals. We estimated survival time (mini- slope, traps were placed near one large Zygophyllum
mum tenure time, in days), the total number of eggs dumosum shrub (>50 cm height), one small Z. du-
produced, the number of egg sacs parasitized or at- mosum, and one Artemisia herba-alba (both <40 cm).
tacked by predators, and the total number and mass Thus, there were a total of 72 traps at 18 shrubs. The
of prey consumed during tenure of each web site. prey captured in each trap was recorded on each day
of 11 sessions consisting of 4-6 sequential sampling
Shrub-trimming experiment days each. Sessions were started roughly every 2 wk
To determine if the variation in shrub height had between 28 February and 20 August 1990. The taxon
long-term effects on survival and reproduction, we se- and body length class (5-mm length classes) of each
lected 32 spiders in Zygophyllum shrubs, measured the arthropod found in a trap were recorded. The lengths
web and web-site parameters listed above, and trimmed were converted to an estimate of prey mass using the
the branches of half of the shrubs, chosen at random, weight-length regression equation given above. In pre-
to the level of the top of the nest. Trimming was done liminary trials (eight sampling days in 1989) we found
during the day to minimize disturbance to the spiders. a significant correlation in prey capture among the four
All spiders were mature females, with the exception of traps at each shrub, so all analyses here are based on
two subadult females; eight had 1-2 egg sacs at the start the average prey capture per day for all traps at a shrub
of the experiment, the remaining spiders had none. We (number of prey per trap, or total mass of prey per
examined the webs at 2-wk intervals, noting the spi- trap).
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- Prey supplementation
mains, and estimated the number of eggs and the To test the effect of food supply on the tendency to
amount of predation or parasitism on them as de- shift web sites, we supplemented the natural prey of
scribed above. 25 unmarked spiders and compared web-site tenure
with that of 26 non-supplemented spiders. All spiders
Prey availability were juvenile females 5.5-8.0 mm in body length and
Prey availability at web sites was assessed indepen- were located in a 400-M2 area adjacent to the main
dently of the condition and fate of the spider occupying study area. Spiders were assigned randomly to the two
it. Prey remains were analyzed from nests of marked groups. Supplemented spiders received a mealworm
and unmarked spiders in 1989 (N = 65 nests) and from (Tenebrio monitor) every other day for 30 d. The webs
nests of 50 spiders used in 1990 to examine the rela- were examined every other day for presence or absence
tionship between shrub size and spider fitness (see of the occupant.
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 Data transformations
was abandoned was taken to the laboratory where we Data were transformed whenever necessary to achieve
counted the number of prey items and measured the an approximately normal distribution as required for
body length of each. The regression equation of Rogers parametric statistical tests. Frequently a logarithmic
et al. (1976), suitable for a range of arthropod taxa, transformation was adequate; if not we used the ex-
was used to estimate total body mass of prey (in mil- tended Box-Cox family of power transformations y =
ligrams) at each web from prey body length in milli- [(a + X)b - I /b for nonzero b, y = ln(a + x) for b =
metres: 0, with maximum likelihood estimates of a and b as-
suming a normal distribution of y. If the transformed
Mass = 0.0305 Length2-62.
data did not have a nearly normal distribution, as in-
We used pitfall traps to compare prey availability in dicated by nearly linear normal probability plots, we
the vicinity of shrubs. Pitfall traps were appropriate in used nonparametric statistical methods.
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1920 YAEL LUBIN ET AL. Ecology, Vol. 74, No. 7
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October 1993 HABITAT SELECTION IN A DESERT SPIDER 1921
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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.
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- Survival time alone accounted for roughly half the vari-
marked females in 1990 and monitored the effects of ance in fecundity (r2 = 0.60 for no. of sacs, r2 = 0.45
experimental manipulations of shrub height. For the for no. of eggs): females that survived longer at a web
latter, the range of shrub heights was 32-85 cm (mean site had more young.
= 59 cm, cv = 27%) and of nest heights 18-62 cm To remove the effects of spider size, we used the
(mean = 33 cm, cv = 19%). residuals from a regression of shrub height on spider
Fecundity (numbers of egg sacs and eggs) was sig- size (tibpat), instead of shrub height itself. This re-
nificantly correlated with shrub height, as were several defines a "large" shrub to be one that is larger than
measures of spider size (body length, abdomen width, average for the size of spider occupying it. These shrub
and tibpat [the length of tibia plus patella for one of height residuals also correlated with fecundity (Table
the first pairs of legs]) that correlate with fecundity 4). Moreover, adding shrub-height residuals to the re-
(Table 4). However it is difficult to separate the effects gression of the number of egg sacs on survival time,
of shrub height per se, from indirect effects due to significantly increased the amount of variance ex-
spider size and survival time at a web site, which cor- plained (r2 = 0.636, F45 = 6.95, P < .025). Thus, it
relate with both fecundity and shrub height (Table 4). 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.
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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 TABLE 5. Analysis of variance on the number and estimated
moved to new sites, based on the 1989 census of 65 marked biomass of prey captured in pitfall traps (using SAS pro-
spiders. N = number of observation dates. Probabilities of cedure GLM [SAS Institute 1989])*. ss= Type III sum of
survival were estimated per 2-d census interval. Overall squares. All main effects and significant interactions
probabilities of survival are estimated per census interval. (P < .05) are shown.
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. Ecology, Vol. 74, No. 7
there was a weak negative correlation (Spearman rank and unit variance. For both prey number and prey mass
correlation ra,=-0.28, P = .05) between shrub height there were no significant departures from the normal
and branch density: larger shrubs had lower branch distribution (Kolmogorov-Smirnov D = 0.13, P = .44),
density at the 2/3 shrub-height level. There was no cor- indicating an absence of positive or negative spatial
relation between shrub height and branch density in correlation among nearby shrubs, and no significant
Artemisia. Thus, differences in shrub architecture may autocorrelations (at a = .05).
explain moves from Artemisia to Zygophyllum, and We also found no evidence of temporal autocorre-
possibly moves from smaller to larger Zygophyllum lation over intervals longer than a single day. Analyzing
shrubs. each shrub separately, I/4 ? of the shrubs had a signif-
icant autocorrelation at a lag of 1 d (successive days,
Prey availability or last day of session N vs. first day of session N + 1).
We used pitfall traps to compare the availability of There were very few significant (P < .05) autocorre-
potential arthropod prey at Artemisia shrubs and at lations at higher lags up to 10 d (two sampling sessions),
large and small shrubs of Zygophyllum. To verify that well below the number expected by chance alone in
arthropods sampled by traps were representative of that many statistical tests. Analyzing all shrubs to-
prey taken by the spiders, we compared the distribution gether, only the autocorrelation between successive days
of arthropod taxa in the traps and in prey remains was statistically significant (r = 0.21, P < .05).
collected from 115 webs of L. revivensis in the study
area. The numbers of arthropods of the different taxa Prey supply and spider fitness
were positively correlated for the two sources (Ken- We looked for correlations between prey supply and
dall's T = 0.43, z = 2.65, P < .01, for 1951 prey items spider fitness by (1) testing the response to artificially
and 13 040 arthropods from webs and traps, respec- increased prey supply, and (2) examining correlations
tively). between spider fitness measures and the amount of prey
Arthropod captures in the traps were highly variable consumed, based on analyses of prey remains in the
over time (ANOVA, Table 5). The differences among nests.
sites and shrub types were also statistically significant, Prey supplementation.-There were no differences
but temporal variability (Session and Date effects) ac- between fed and unfed spiders in the tendency to aban-
counted for a much larger portion of the total vari- don web sites. Average web-site tenure was 9.4 ? 6.3
ability. d (mean ? 1 SD) for the fed group and 9.5 ? 7.1 d for
The deviations of individual shrubs from the habi- the control group. Between days 8 and 10 of the ex-
tat-wide overall trends (ANOVA main effects) were periment, there was a bout of unseasonably hot weath-
analyzed for temporal and spatial correlations. Any er, which may have caused the considerable mortality
such correlations could be important for movement that occurred then in both groups equally.
decisions, because they could allow spiders to predict Prey consumed and fitness measures. -Prey remains
prey availability at other sites "now" (spatial correla- in nests provided estimates of the total number and
tion) or at the same site "later" (temporal autocorre- mass of prey consumed during tenure at a web site,
lation). We report here only the results for the number and the rate of prey capture (number and mass of prey
of prey captured; the results for prey mass are nearly per day) for spiders of known web-site tenure (1989
identical but are less meaningful due to the additional census data).
errors introduced by estimating prey mass from the Total prey number and mass were correlated with
body sizes of prey items. leg length (tibpat, the length of tibia plus patella for
We tested for spatial correlations both across and one of the first pairs of legs) in the 1989 sample con-
within sampling days. For the former we used the pair- taining juveniles and adults, but not in the 1990 sample
wise cross-correlation coefficients between the time se- which was only adults (Table 6). Thus, as juveniles
ries for each shrub. The correlations were not signifi- grew and molted, prey consumption increased, but for
cantly higher for shrubs at the same site than for shrubs adults prey consumption was unrelated to leg length.
at different sites, indicating a lack of spatial correlation For a given leg length, however, the amount of prey
among nearby shrubs (r2 = 0.041 ? 0.046 [mean ? 1 consumed had strong positive correlations with body
SD], n = 18 same site; mean r2 = 0.033 + 0.047, n = size and fecundity.
135 between sites). Total prey accumulation (number and mass) was
To test for spatial correlations within sampling days, also correlated in both years with how long spiders
we computed for each sampling day the F statistic from remained at web sites (Table 6). However, for marked
a single-factor ANOVA with site as the treatment vari- spiders with known web-site tenure, the rate of prey
able. These F values were transformed (using the ap- capture (number per day) was negatively correlated
propriate F distribution) so that under the null hy- with tenure (the correlation with mass per day was also
pothesis of no spatial correlation the transformed values negative but not statistically significant). Negative cor-
would be independent normal variates with zero mean relations could result from (1) diminishing returns at
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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.
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1926 YAEL LUBIN ET AL. Ecology, Vol. 74, No. 7
site selection. Thus, while the energy cost of web build- visually hunting predators, and reduced the number of
ing may contribute to mortality during a move, it is supporting structures above the web. Nonetheless, nei-
unlikely to be the major cause of mortality. ther survival nor fecundity was reduced in the exper-
imental group.
Benefits of web relocation The lack of any significant effect of exposure on spi-
In spite of the large risk, virtually all L. revivensis der fitness could be explained if the experiments were
females moved at least once during their lifetimes after conducted during a period of cooler weather or when
initial dispersal from the maternal nest. The question the risk of predation was low. However, web relocation
is, then, what benefit is derived from recurrent dis- occurs throughout the summer, including the time pe-
persal movements? riod during which the manipulations were performed,
For many mobile organisms, the primary benefit to and temperatures were near average for that time of
moving to a new site is the expectation of improved year.
food supply at the new site relative to the current one. An alternative hypothesis is that web-site selection
In contrast to previous studies on web-building spiders in L. revivensis may be less precise than expected be-
(e.g., Vollrath 1985, Gillespie and Caraco 1987, but cause of the risks involved in moving and the low
see also Riechert and Tracy 1975), we showed this not probability of finding a better web site. The decision
to be the case for L. revivensis. Movement to new sites problem for a spider moving sequentially from shrub
in this species was apparently unrelated to prey avail- to shrub in search of a new web site is similar to the
ability. The type of shrub occupied (Artemesia, large mate choice problem as modeled by Real (1990), or
or small Zygophyllum) explained only a small amount the classical models of sequential job choice in eco-
of the variation in prey availability at web sites in nomics (Lipmann and McCall 1976). The optimal de-
comparison with the habitat-wide temporal trends that cision rule for such sequential choice problems is typ-
affected all web sites equally. The mean prey capture ically a "satisficing" rule: the item (shrub, mate, job
rate at the best shrub type was higher than that at the offer) chosen is the first encountered that meets or ex-
worst by only 10% for prey number, while the best and ceeds a minimum criterion (Lipmann and McCall
worst trapping sessions differed by 275%. Web relo- 1976). "Satisficing" criteria have been suggested to op-
cation to a different shrub type in order to improve erate in habitat selection decisions of other desert spi-
prey capture would seem to be an expensive gamble ders (Riechert 1985, Ward and Lubin 1993).
with little expected payoff. Furthermore, we found no In L. revivensis, the search cost is high due to the
spatial correlation in prey availability, so a spider's high risk of mortality during movement away from the
prey capture success at a particular web site does not web, thus the range of acceptable items will be broad,
provide any information about the quality of other and will include items that are far from optimal (Lip-
sites. The lack of temporal and spatial correlations also mann and McCall 1976, Real 1990). High mortality
implies that a spider has no reason to abandon a site during moves may also increase the amount of time
in response to a period of poor prey capture. an individual should remain in its current location,
The benefits of recurrent dispersal in L. revivensis tolerating poor or deteriorating conditions, before re-
appear to be associated with improved shrub architec- locating (Houston and McNamara 1986, Gilliam 1990).
ture. A likely benefit is the improvement of thermal Consequently much of the life-span would be spent
conditions within the nest during the transition from "making do" in suboptimal sites. Natural selection
the cool to hot season, and throughout the hot season. would then favor greater plasticity in habitat selection,
High daytime temperatures inside the nest force spi- either by increasing physiological tolerance, or by be-
ders to adopt thermoregulatory positions that expose havioral responses to reduce the impact of a poorer
them to visually orienting predators (e.g., birds and web site.
mantids) (Konigswald et al. 1990; Y. Lubin, unpub-
lished data). Spiders in larger shrubs, and in shrubs Conclusion: fitness measures for
with more open branch structure (e.g., Zygophyllum), habitat selection
will experience less extreme nest temperatures, because We have shown here that habitat selection is bene-
air temperature decreases with increasing height above ficial to the desert widow spider, L. revivensis. Spiders
the ground (Campbell 1977) and because nests in that moved to "good" web sites (taller shrubs) fared
sparsely branched shrubs will be more exposed to cool- better in the long run, as measured by their growth and
ing breezes than nests in dense shrubs. reproductive success. As food was not the direct cause
of success at a web site, indirect measures of fitness
Long-term consequences of habitat selection relating to food intake were not useful; the sole useful
The positive relationship observed between spider measure was fitness itself. Similar examples may
fitness and shrub height was tested experimentally by abound, whenever site-selection criteria are based on
trimming shrubs to nest height. This treatment in- qualities other than food availability.
creased the amount of solar radiation directly imping- Mortality may occur both on the web and during
ing upon the nest, exposed the nest from above to web relocation, but the probability of dying during a
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October 1993 HABITAT SELECTION IN A DESERT SPIDER 1927
move was 40% compared with <1% per day on the Gillespie, R. G. 1981. The quest for prey by the web-build-
web (Table 3). Current foraging theory generally treats ing spider, Amaurobius similis (Blackwell). Animal Behav-
iour 29:953-954.
mortality risk as a feature of the habitat (Gilliam 1990, Gillespie, R. G., and T. Caraco. 1987. Risk-sensitive foraging
Lima and Dill 1990). Here we have shown that the strategies of two spider populations. Ecology 68:887-899.
risk of mortality may be more important as a com- Gilliam, J. F. 1990. Hunting by the hunted: optimal prey
ponent of the move itself and will influence the decision selection by foragers under predation hazard. Pages 797-
whether to move at all, rather than which habitat to 820 in R. N. Hughes, editor. Behavioral mechanisms of
food selection. Springer-Verlag, New York, New York, USA.
choose. For L. revivensis, it appears that the main cost Hodge, M. A. 1987a. Macrohabitat selection by the orb
of moving is the decreased probability of survival, and weaving spider, Micrathena gracilis. Psyche 94:347-361.
there are no indirect measures (e.g., energy expended, 1987b. Factors influencing web site residence time
reduced food intake) that will substitute for survival. of the orb weaving spider, Micrathena gracilis. Psyche 94:
It remains paradoxical that experimental tests of the 363-371.
Houston, A. I., and J. M. McNamara. 1986. The influence
relationship between the preference for habitat features of mortality on the behaviour that maximizes reproductive
and fitness are inconclusive. "Satisficing" criteria for success in a patchy environment. Oikos 47:267-274.
web-site selection may partly explain the weak corre- Janetos, A. C. 1982a. Active foragers vs. sit-and-wait pred-
lation between fitness and prefered sites. Nonetheless, ators: a simple model. Journal of Theoretical Biology 95:
381-385.
the missing links between short-term advantages of 1982b. Foraging tactics of two guilds of web-spin-
habitat selection and dispersal, and long-term fitness ning spiders. Behavioral Ecology and Sociobiology 10:19-
consequences, bear further investigation. 27.
Konigswald, A., Y. Lubin, and D. Ward. 1990. The effec-
ACKNOWLEDGMENTS tiveness of the nest of a desert widow spider, Latrodectus
We are grateful to Helen Dallas, Arthur Dumosch, Iris Mus- revivensis, in predator deterrence. Psyche 97:75-90.
li, and Sonya Rosen who helped with various aspects of this Krebs, C. J. 1989. Ecological methodology. Harper & Row,
project, to Fiona Wierczk for Fig. 1, and to David Dickey for New York, New York, USA.
statistical advice. David Ward provided help and advice Levy, G., and P. Amitai. 1983. Review of the widow spider
throughout the study. We thank James Gilliam, George Hess, genus Latrodectus (Araneae: Theridiidae) in Israel. Zoo-
Leslie Real, and David Ward for critically reading the manu- logical Journal of the Linnean Society 77:39-63.
script. The study was supported by a U.S.-Israel Binational Lima, S. L., and L. M. Dill. 1990. Behavioral decisions made
Science Foundation grant #86-00092. This is contribution under the risk of predation: a review and prospectus. Ca-
Number 157 of the Mitrani Center for Desert Ecology. nadian Journal of Zoology 68:619-640.
Lippman, S. A., and J. J. McCall. 1976. The economics of
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