Behavioral Ecology
doi:10.1093/beheco/arq040
Advance Access publication 13 April 2010
Using visual cues of microhabitat traits to find
home: the case study of a bromeliad-living
jumping spider (Salticidae)
Paula M. de Omenaa and Gustavo Q. Romerob
Pós-graduac
xão em Biologia Animal, Departamento de Zoologia e Botânica, Instituto de Biociências,
Letras e Ciências Exatas, Universidade Estadual Paulista, Rua Cristóvão Colombo 2265, CEP 15054-000,
São José do Rio Preto-SP, Brazil and bDepartamento de Zoologia e Botânica, Instituto de Biociências,
Letras e Ciências Exatas, Universidade Estadual Paulista, Rua Cristóvão Colombo 2265, CEP 15054-000,
São José do Rio Preto-SP, Brazil
a
There are many examples of predators having specialized microhabitat requirements, but the sensory mechanisms by which
predators detect, identify, and evaluate microhabitats are only poorly understood. The ability to use visual cues to select microhabitats was investigated using Psecas chapoda, a bromeliad-dwelling salticid spider. In this study, we manipulated real plants and
photos of plants to test whether P. chapoda uses plant architecture to select host plants and whether visual cues alone are sufficient
for them to select microhabitats. The use of photos on the experiment allowed us to exclude the potential influence of other
cues, such as color and odor, on host plant selection by the spider. Our results showed that P. chapoda selects their microhabitat by
evaluating architectural features of leaves and rosette of the host plants. Rosette-shaped plants (Agavaceae) were preferred over
other types of plant architecture. Spiders showed a preference for photographs of rosette-shaped plants having narrow and long
leaves, confirming that they can make these choices entirely on the basis of vision. These salticids can recognize and select
microhabitats bearing specific architectural features, which possibly reflects an adaptation to choose microhabitats that are
favorable to its survivorship. Key words: Bromeliaceae, host plant selection, plant architecture, visual cues, visual selection,
Salticidae. [Behav Ecol 21:690–695 (2010)]
icrohabitat selection is particularly important to increase
the survival rate and reproductive success of animals
inhabiting heterogeneous environments (Pianka 2000). Although spiders that live on plants are known for being selective
in their microhabitat and foraging sites, especially in relation
to the physical structure of plants (Gunnarsson 1996; Halaj
et al. 2000; Romero and Vasconcellos-Neto 2005a; Morse
2007), little is known about which sensory modality (e.g., visual,
olfactory, and tactile) they use to evaluate and select substrata.
Some studies have shown that spiders can select a substratum
based on color (Greco and Kevan 1994; Heiling et al. 2005),
odor (Aldrich and Barros 1995; Krell and Krämer 1998; Heiling
et al. 2004), texture (Morse 1988; Greco and Kevan 1994), or
the frequency of prey visits (Morse 2007 and references
therein). Most of the studies of substratum choice by spiders
relate to optimal foraging theory and measure how choice is
based on the amount of food resources that spiders can obtain
(e.g., Morse 2007 and references therein). However, many spider species spend their entire life cycle on specific substrata
and not only use them as foraging site but also as site for breeding, shelter for adults, and immature and as nursery for spiderlings (e.g., Cumming and Wesolowska 2004; Romero and
Vasconcellos-Neto 2005a; Romero 2006). To date, little is
known about microhabitat choices by these specialist spiders.
All jumping spiders have unique complex eyes and have
a spatial acuity, which is unparalleled in any other animals
M
Address correspondence to G.Q. Romero. E-mail: gq_romero
@yahoo.com.br.
Received 18 May 2009; revised 18 February 2010; accepted
23 February 2010.
The Author 2010. Published by Oxford University Press on behalf of
the International Society for Behavioral Ecology. All rights reserved.
For permissions, please e-mail: [email protected]
of comparable size (Blest et al. 1981; Williams and McIntyre
1980). The visual sense controls much of their activities, such
as orientation to prey capture (Hill 1979; Tarsitano and
Andrew 1999; Tarsitano et al. 2000; Li and Lim 2005;
Tarsitano 2006), courtship (Jackson 1977; Lim et al. 2007),
and antipredator and agonistic behavior (Taylor et al. 2001;
Elias et al. 2008). It has been suggested that such an acute
vision could have been essential for the evolution of these
behaviors in Salticidae (Jackson 1992; Jackson and Pollard
1996; Lim and Li 2006; Lim et al. 2007). Vision could also
be used in substrata choice by salticids and even contribute to
specialization and evolution for specific microhabitats, such as
bromeliads (Romero 2006). However, to date, there is no
study showing that jumping spiders use their complex visual
system to choose their host plants.
The ability to use visual cues in microhabitat selection was
investigated using Psecas chapoda, a bromeliad-dwelling salticid
spider that almost exclusively inhabits only 1 host species,
Bromelia balansae (Romero and Vasconcellos-Neto 2005a;
Romero 2006). The entire life cycle of P. chapoda, including
courtship behavior, mating, egg sac deposition, and population recruitment of the young occurs on this bromeliad
(Rossa-Feres et al. 2000; Romero and Vasconcellos-Neto
2005a, 2005b, 2005c). The spider uses the central concavity
of the B. balansae leaves as shelter and as site where females
deposit egg sacs (Rossa-Feres et al. 2000; Vieira and Romero
2008). The structure of the leaves is also utilized during courtship; females remain in the base of the rosette, whereas males
occupy the upper part of the leaves for their courtship displays (Rossa-Feres et al. 2000). Additionally, adult and immature spiders use the base of the rosette as a refuge from
predators (Romero and Vasconcellos-Neto 2005a; Omena
de Omena and Romero • Visual cues and host plant selection
691
and Romero 2008) and fire (Romero GQ, Omena PM,
unpublished data).
The evolution of P. chapoda specialization for this single host
species is still poorly understood and might be related to the
large availability of a substratum bearing a specific architecture (i.e., rosette of long and narrow leaves) that provides
numerous benefits to the spider (Romero 2006; Omena and
Romero 2008). P. chapoda appears to use certain architectural
features of microhabitat to select its hosts (Omena and
Romero 2008). When bromeliad species bearing a similar
(i.e., narrow and long leaves) or a different architecture
(i.e., wide and short leaves) from the host B. balansae was
experimentally introduced, P. chapoda colonized all the narrow long-leaved plants but not the wide and short-leaved ones
(Omena and Romero 2008). Although the existence of specific associations between spiders and plants depends on the
ability of spiders to find their hosts, to our knowledge, there
has been no experimental test of how jumping spiders actively
choose suitable microhabitats and which cues are used to find
their hosts.
In this study, we manipulated real plants and photographs of
plants to investigate if the bromeliad-dwelling jumping spider
P. chapoda selects for a specific microhabitat architecture and
whether visual cues of plant traits are sufficient for this spider
to select between microhabitats. This spider–plant relationship
was recently reported to be mutualistic, where spiders benefit
from occurring on the host and debris derived from their
biological activities improve plant nutrition (Romero et al.
2006). Therefore, information on host plant choice can help
to understand the evolution of this spider–plant relationship.
Specifically, we addressed the 3 following questions: 1) Does
P. chapoda recognize and select rosette-shaped plants? 2) does
this spider select rosettes bearing specific architectural features (i.e., long and narrow leaves)? and 3) can spiders use
visual information to select plants from photographs?
METHODS
General procedures
Spiders were collected near São José do Rio Preto city, São
Paulo State (Brazil). In the laboratory, they were maintained
individually with food ad libitum (Drosophila melanogaster)
and with a piece of moist cotton in dram vials (15-cm high,
7-cm diameter) for 1–2 days. The laboratory was set under
12-h illumination and the temperature ranged from 24 to
34 C. The experiments were carried out in an open grassland
space located on the experimental area of the Universidade
Estadual Paulista. Spiders were transferred individually from
the laboratory to the experimental area inside transparent
acrylic vials (5.5-cm high, 5.0-cm diameter). We used 30 males
and 30 females for each of the experiments.
Experiment I: selection of rosette architecture
To investigate if P. chapoda selects rosette-shaped plants over
others architectures, we used squared arenas containing 4
plant species, 1 rosette-shaped plant and 3 without rosette
architecture. The rosette-shaped plant used was Agave angustifolia (Agavaceae), an exotic plant, with long leaves, sharing
similar morphological traits with Bromeliaceae. The 3 non–
rosette-shaped plants were 1) Euterpe oleracea (Arecaceae),
which has long leaves, albeit without rosette formation;
2) Croton floribundus (Euphorbiaceae), which has long and wide
leaves; and 3) Delonix regia (Fabaceae), which has large leaves
but bears minute secondary leaflets (Figure 1A and Table 1).
Each arena (n ¼ 5) consisted of a square wooden board, with
sides of 0.5 m, surrounded by 4 plants, 1 individual of each
experimental plant species (i.e., Ag. angustifolia, E. oleracea,
Figure 1
Relationship between scores of factors 1 and 2 for the DA of rosetteshaped and non–rosette-shaped plants (A) and bromeliads (B).
C. fluribundus, and D. regia), which were placed in individual
pots (Figure S1a in Supplementary Appendix). We used the
pots’ edges to support the board, in a manner that the vegetative part of the plants was positioned above the boards, that
is, in the visual field of spiders, and the pot under the boards.
We conducted the experiment between 10:00 and 15:00 h
because P. chapoda lives in open area grasslands and withstands daily intense luminosity and high temperatures. At
the beginning of each day of the experiment, we arranged
the plants randomly by drawing at the vertices of the square
board and removed the existing fauna. For each replicate, we
placed the acrylic vial containing 1 spider in the center of the
arena, and then, we removed the lid from the vial to allow the
spider to visualize and freely select the substratum. We took
the observations from at least 3-m away to avoid human interference on the spiders’ behavior. A single spider was released in each trial; we considered that the spider had made
a choice if it jumped or climbed on a plant. Spiders were then
removed from the plant immediately. Between each trial, we
rotated the plants around the wooden board (i.e., in clockwise
and sometimes in anticlockwise direction) and cleaned the
board and plant with a flannel to remove the silk released
by the preceding spider.
Experiment II: selection of bromeliad traits
We also sought to test whether P. chapoda preferred rosetteshaped plants bearing specific architectural features, that is,
Behavioral Ecology
692
Table 1
Mean (61 standard error) of leaf or leaflet length (LL) and width (LW), LW:LL ratio, leaf or leaflet number (LN), distance between leaves or
leaflets (DL), number of plants (n), and rosette (R) (present [P] or absent [A]) of the plant species used in the experiments
Plants
LL (cm)
Bromelia balansae
Aechmea distichantha
Ae. blanchetiana
Ae. fasciata
Agave angustifolia
Euterpe oleracea
Croton loribundus
Delonix regia
70.45
50.75
40.20
28.7
28.7
30
12.15
1.17
6
6
6
6
6
6
6
6
LW (cm)
4.05
3.28
1.55
2.24
2.24
1.84
0.54
0.07
2.67
2.21
7.67
8.26
3.29
5.65
6.31
0.63
6
6
6
6
6
6
6
6
0.08
0.09
0.22
0.38
0.15
0.56
0.23
0.03
LN
18.7
31.1
15.6
16.6
30.3
6.3
15.1
2348.8
narrow and long leaves. For this purpose, we used 4 bromeliad
species (B. balansae, Aechmea distichantha, Ae. blanchetiana, and
Ae. fasciata) arranged in arenas as above. B. balansae and
Ae. distichantha have long and narrow leaves with margins covered with spines; however, the latter differs in its leaf base,
which has a lateral expansion that forms a phytotelmata
(Table 1). Ae. blanchetiana and Ae. fasciata also bear phytotelmata; the first has intermediate leaf length when compared
with B. balansae and Ae. fasciata, but the former has leaf width
as broad as Ae. fasciata. Ae. fasciata possesses the shortest leaves
of these bromeliads (Figure 1B and Table 1). We conducted
this experiment using the same procedures described for experiment I (Figure S1b in Supplementary Appendix).
Experiment III: visual choice for hosts using photographs
In this experiment, we aimed to test if visual cues from the
leaves of rosettes are sufficient for P. chapoda to select its host
plant or bromeliads that share similar architectural features
with B. balansae. For this, we used black-and-white photos of
the 4 bromeliad species used in the second experiment (i.e.,
B. balansae, Ae. distichantha, Ae. blanchetiana, and Ae. fasciata).
The photos allowed us to exclude the potential influence of
other cues, such as plant color and odor, on host plant selection by P. chapoda. The photographs of the bromeliad photos
were scaled to the actual size of real plants, which were presented on polystyrene boards (80 cm in width 3 60 cm in
height). We also removed the coloration to produce blackand-white images using Adobe Photoshop CS3, leaving only
the bromeliad image on a white background. We presented
the 4 photos on the edges of a square lumber board with sides
of 80 3 80 cm; the board was the arena’s base, and the photos
formed the 4 walls (Figures S1c and S1d in Supplementary
Appendix). Two observers recorded the spiders’ behavior
from ladders, which were 1.5-m high and 3-m away from the
arena to avoid interference. For each spider, we changed the
position of the photos (i.e., clockwise or anticlockwise), and
we removed the silk of the preceding spider from the photos
and arena with a flannel between trials. We recorded the latency for each spider to jump or climb on to a photograph
and also whether they landed on the bromeliad image or on
the white background.
Statistical analyses
We compared the number of choices made by spiders in
experiments I, II, and III using the G test for goodness of fit
(Sokal and Rohlf 1995). For the second and third experiments, pairwise comparisons were conducted using G test,
with required significance values to be adjusted using the
sequential Bonferroni procedure (Rice 1989). Because previous work has found that P. chapoda prefers rosette plants bearing long, but especially, narrow leaves than plants with other
architectures (Omena and Romero 2008), we grouped
LW:LL
6
6
6
6
6
6
6
6
1.32
2.46
1.36
1.38
3.17
0.39
0.58
221.55
0.038
0.045
0.175
0.310
0.121
0.20
0.524
0.54
6
6
6
6
6
6
6
6
0.001
0.003
0.008
0.038
0.006
0.025
0.118
0.015
DL (cm)
n
R
0
0
0
0
0
19.65 6 1.2
1.9 6 0.14
0.43 6 0.04
10
10
10
10
10
10
10
10
P
P
P
P
P
A
A
A
bromeliads with similar architecture into 2 categories to perform the pairwise comparisons: 1) plants that have narrow
leaves (i.e., B. balansae and Ae. distichantha) and 2) plants that
have broad leaves (i.e., Ae. blanchetiana and Ae. fasciata)
(Figure 1B). For these analyses, a values were adjusted to
0.025 by the Bonferroni correction. For the third experiment,
we compared the number of spiders landed on the bromelia
image and on the white background using the G test for
goodness of fit (Sokal and Rohlf 1995).
Plant characterization
We performed 2 discriminant analysis (DA): the first one to
characterize the rosette-shaped and non–rosette-shaped plants
and to determine what traits best discriminate between them
and the second one to characterize the bromeliad species. We
performed the analyses using some morphological parameters
of plants, such as leaf or leaflet length (LL), width (LW), LW:
LL ratio, and the number (LN) and distance between leaves or
leaflets (LD) (see Table I). Data were log10 transformed for
normalization and equalization of variances.
The analysis of rosette-shaped and non–rosette-shaped
plants classified about 96% of plant species (100% of B. balansae, 80% of Ae. distichantha, 100% of Ae. blanchetiana, 90% of
Ae. fasciata, 100% of Ag. angustifolia, 100% of E. oleracea, 100%
of C. floribundus, and 100% of D. regia) (Figure 1A). The most
important parameter that discriminated between plant groups
was the distance between the leaf axils and leaf width (F statistics: LD ¼ 699.88, LW ¼ 57.52, LW:LL ¼ 26.07, LN ¼ 32.12,
and LL ¼ 28.59). The architecture of plant species without
rosette shape (i.e., E. oleraceae, C. floribundus, and D. regia)
differed from those bearing rosette architecture (Figure 1).
Each one of those plant species formed a group distinct and
distant from those of rosette-shaped species (Figure 1).
Ag. angustifolia was similar to bromeliads, especially Ae. distichantha (Figure 1A). The analysis of bromeliads classified
about 95% of plant species (100% of B. balansae, 90% of Ae.
distichantha, 100% of Ae. blanchetiana, and 90% of Ae. fasciata)
(Figure 1B). The most important parameter that separated
groups of bromeliads was leaf width (F ¼ 95.91); the first
canonical variable (factor 1) better discriminated the species
(eigenvalue of factor 1 ¼ 37.36 and eigenvalue of factor 2 ¼
1.83). The bromeliad species were separated in 2 groups,
one of them contained B. balansae and Ae. distichantha, that
is, plants with narrow leaves, and the other Ae. blanchetiana
and Ae. fasciata, that is, bromeliads that have broad leaves
(Figure 1B and Table I).
RESULTS
Experiment I
Male and female P. chapoda exhibited a similar pattern of host
plant selection. Their distribution of choices was not at
de Omena and Romero • Visual cues and host plant selection
693
Figure 2
Percentage of active choices of non-bromeliad plants in experiment I
by P. chapoda. Males: n ¼ 25 and females: n ¼ 30.
random, with both sexes preferring Ag. angustifolia (females:
G ¼ 65.71, n ¼ 30, degrees of freedom [df] ¼ 3, P , 0.001 and
males: G ¼ 69.31, n ¼ 25, df ¼ 3, P , 0.001; Figure 2). Males
that did not select Ag. angustifolia (n ¼ 5) left the arena and
did not select any other available plant.
Experiment II
Females chose bromeliad species in a manner inconsistent with
random choice (G ¼ 15.8, n ¼ 30, df ¼ 3, P , 0.001; Figure 3).
A similar result was also found for males (G ¼ 21.58, n ¼ 30,
df ¼ 3, P , 0.001; Figure 3). Bromeliads bearing narrow leaves
were more frequently selected by both males (pairwise comparisons; G ¼ 18.03, n ¼ 26, 4, df ¼ 1, P , 0.001) and females
(pairwise comparisons; G ¼ 8.99, n ¼ 23, 7, df ¼ 1, P ¼ 0.003),
indicating preference for plants with long narrow leaves and
rosette formation in both sexes.
Experiment III
Similar nonrandom patterns of host plant selection were also
obtained using black-and-white photographs (females: G ¼
20.53, n ¼ 30, df ¼ 3, P , 0.001 and males: G ¼ 10.71, n ¼
30, df ¼ 3, P ¼ 0.013; Figure 4). Photographs of bromeliads
with narrow leaves were selected more frequently by females
(pairwise comparisons; G ¼ 18.03, n ¼ 26, 4, df ¼ 1, P ,
0.001) and males (pairwise comparisons; G ¼ 8.99, n ¼ 23,
7, df ¼ 1, P ¼ 0.003). Most of the females (87%) and males
(88%) that chose bromeliads with narrow leaves (i.e., B. balansae and Ae. distichantha) first selected the bromeliad image
and not the white background (females: G ¼ 13.71, n ¼ 26,
Figure 3
Percentage of active choices of bromeliads in experiment II by
P. chapoda. Significance level for pairwise comparison with the G test
is shown above the horizontal bars (ns ¼ nonsignificant; **P , 0.01).
Males: n ¼ 30 and females: n ¼ 30.
Figure 4
Percentage of active choices of bromeliad photos by females (A) and
males (B) of Psecas chapoda. Different bands in the bars indicate the
percentages of individuals that moved and firstly contacted with the
bromeliad image or with the white background. Bars representing
males and females are indicated by the symbols # and $, respectively.
Significance level for pairwise comparison is shown above the bars
(ns ¼ nonsignificant; **P , 0.01). Males: n ¼ 30 and females: n ¼ 30.
df ¼ 1, P , 0.001 and males: G ¼ 15.18, n ¼ 24, df ¼ 1, P ,
0.001; Figure 4 and Figures S1e and S1f in Supplementary
Appendix).
DISCUSSION
Our findings show that P. chapoda prefer plants that have the
architectural features of their host plant. The results from the
experiment in which we used photographs instead of real
plants demonstrate that the spider can make these choices
entirely on the basis of visual cues based on details of the
plant’s architecture. Previous studies have shown that salticids
respond realistically to video playbacks (Clark and Uetz 1990)
and to computer-generated animation (Harland and Jackson
2002; Nelson and Jackson 2006) and use visually based decisions in both mate and prey choice. However, here, we have
shown for the first time that a salticid can use visual cues of
plant architecture for microhabitat selection. Some authors
have reported that many jumping spiders can discriminate
colors (Nakamura and Yamashita 2000) and that they have
photopigment that is maximally sensitive to green in their
principal eyes (DeVoe 1975; Yamashita and Tateda 1976; Blest
et al. 1981; Nakamura and Yamashita 2000). Therefore, it is
possible that the color of bromeliads (i.e., variable shades of
green) could also play some role in microhabitat selection by
P. chapoda, which should be investigated in future studies.
The visual system of jumping spider has exceptionally high
spatial acuity (Land 1969a, 1969b; Blest et al. 1990; Clark and
Uetz 1990; Land and Fernald 1992), which may enable
P. chapoda to visually select its host plant. The host B. balansae
is a suitable site for this spider species (see Romero and
Vasconcellos-Neto 2005a; Romero 2006; Omena and Romero
2008), and it is extremely abundant with a conspicuous architecture that differs from others in the environment. This
makes the use of visual cues an effective way to locate it, which
may be of crucial importance when migrating among bromeliads to obtain food, mating opportunities, and shelter.
Because these spiders are likely to be more vulnerable to
predators when they are away from their host plants (Omena
PM, personal observation), an inability to quickly detect their
plants could significantly increase their mortality risk.
694
Both males and females detected and chose their bromeliad
host plant (B. balansae), as well as bromeliads that share similar traits with this host plant (e.g., Ae. distichantha), and nonbromeliad plants, which bear rosette architecture (i.e., Ag.
angustifolia). Romero and Vasconcellos-Neto (2005a) proposed that P. chapoda would be capable of recognizing and
evaluating the physical features of microhabitat. In this study,
we show that this species of salticid was able to distinguish
between specific architectural features of plants and that it
was selective for microhabitat architecture bearing rosette
shape and both narrow and long leaves, which is typical of
its preferred host.
In conclusion, P. chapoda can evaluate and distinguish physical structure of microhabitats and actively select its host
plants on the basis of shape. P. chapoda can select microhabitats based on visual cues of plant traits, including those available from black-and-white photos. The widespread availability
of a substratum bearing a conspicuous architecture (i.e., B. balansae) associated with the ability of P. chapoda in detecting this
substratum possibly favored the establishment of this spider–
plant association.
SUPPLEMENTARY MATERIAL
Supplementary material can be found at http://www.beheco
.oxfordjournals.org/.
FUNDING
A postgraduate fellowship from Fundacxão de Amparo à
Pesquisa do Estado de São Paulo (FAPESP; 06/59409-1 to
P.M.O.); Research grants from FAPESP (04/13658-5 and 05/
51421-0 to G.Q.R.).
The authors thank Dr Robert Jackson and one anonymous reviewer for
their valuable comments and suggestions on the first draft of the manuscript. The authors also thank Carlos E. N. Girardi, Diogo B. Provete,
Paulo E. Cardoso, Marcelo O Gonzaga, Michel V. Garey, Thiago
Gonc
xalves-Souza for advice and reviewing the manuscript; Fernando
B. Noll helped with discriminant analysis; and Gustavo C. Piccoli
and José C. Souza helped with the setup of the experiments and data
collection.
REFERENCES
Aldrich JD, Barros TM. 1995. Chemical attraction of male crab spiders
(Araneae, Thomisidae) and cleptoparasitic flies (Diptera, Milichiidae, and Chloropidae). J Arachnol. 23:212–214.
Blest AD, Hardie RC, McIntyre P, Williams DS. 1981. The spectral
sensitivities of identified receptors and the function of retinal tiering in the principal eyes of a jumping spider. J Comp Physiol A.
145:227–239.
Blest AD, O’Carrol DC, Carter M. 1990. Comparative ultrastructure of
layer I receptor mosaics in principal eyes of jumping spiders: the
evolution of regular arrays of light guides. Cell Tissue Res. 262:
445–460.
Clark DL, Uetz GW. 1990. Video image recognition by the jumping
spider, Maevia inclemens (Araneae: Salticidae). Anim Behav. 40:
884–890.
Cumming MS, Wesolowska W. 2004. Habitat separation in a speciesrich assemblage of jumping spiders (Araneae: Salticidae) in a suburban study site in Zimbabwe. J Zool. 262:1–10.
DeVoe RD. 1975. Ultraviolet and green receptors in principal eyes of
jumping spiders. J Gen Physiol. 66:193–207.
Elias DO, Kasumovic MM, Punzalan D, Andrade MCB, Mason AC.
2008. Assessment during aggressive contests between male jumping
spiders. Anim Behav. 67:901–910.
Greco CF, Kevan PG. 1994. Contrasting patch choosing by anthophilous ambush predators: vegetation and floral cues for decisions by
a crab spider (Misumena vatia) and males and females of an ambush
bug (Phymata americana). Can J Zool. 72:1583–1588.
Behavioral Ecology
Gunnarsson B. 1996. Bird predation and vegetation structure affecting spruce-living arthropods in a temperate forest. J Anim Ecol. 65:
389–397.
Halaj J, Ross DW, Moldenke AR. 2000. Importance of habitat structure
on the arthropod food-web in Douglas-fir canopies. Oikos. 90:
139–152.
Harland DP, Jackson RR. 2002. Influence of cues from anterior medial
eyes of virtual prey on Portia fimbriata, an araneophagic jumping
spider. J Exp Biol. 205:1861–1868.
Heiling AM, Cheng K, Herberstein ME. 2004. Exploitation of floral
signals by crab spiders (Thomisus spectabilis, Thomisidae). Behav
Ecol. 2:321–326.
Heiling AM, Chittka L, Cheng K, Herberstein ME. 2005. Colouration
in crab spider: substrate choice and prey attraction. J Exp Biol.
208:1785–1792.
Hill DE. 1979. Orientation by jumping spiders of the genus Phidippus
(Araneae: Salticidae) during the pursuit of prey. Behav Ecol Sociobiol. 5:301–322.
Jackson RR. 1977. An analysis of alternative mating tactics of the
jumping spider Phidippus johnsoni (Araneae, Salticidae). J Arachnol.
5:185–230.
Jackson RR. 1992. Eight-legged tricksters. Bioscience. 42:590–598.
Jackson RR, Pollard SD. 1996. Predatory behaviour of jumping spiders. Annu Rev Entomol. 41:287–308.
Krell FT, Krämer F. 1998. Chemical attraction of crab spiders (Araneae, Thomisidae) to a flower fragrance component. J Arachnol.
26:117–119.
Land MF. 1969a. Movements of the retinae of jumping spiders (Salticidae: Dendryphantinae) in response to visual stimuli. J Exp Biol.
51:471–493.
Land MF. 1969b. Structure of the retinae of the principal eye of jumping spiders (Salticidae: Dendryphantinae) in relation to visual
optics. J Exp Biol. 51:443–470.
Land MF, Fernald RD. 1992. The evolution of eyes. Annu Rev Neurosci. 15:1–29.
Li D, Lim MLM. 2005. Ultraviolet cues affect the foraging behaviour
of jumping spiders. Anim Behav. 70:771–776.
Lim MLM, Land MF, Li D. 2007. Sex-specific UV and fluorescence
signals in jumping spiders. Science. 315:481.
Lim MLM, Li D. 2006. Extreme ultraviolet sexual dimorphism
in jumping spiders (Araneae: Salticidae). Biol J Linn Soc. 89:
397–406.
Morse DH. 1988. Relationship between crab spiders Misumena vatia
nesting success and earlier patch-choice decisions. Ecology. 69:
1970–1973.
Morse DH. 2007. Predator upon a flower: life history and fitness in
a crab spider. Cambridge (UK): Harvard University Press.
Nakamura T, Yamashita S. 2000. Learning and discrimination of colored papers in jumping spiders (Araneae, Salticidae). J Comp Physiol A. 186:897–901.
Nelson XJ, Jackson RR. 2006. A predator from East Africa that chooses
malaria vectors as preferred prey. PLoS One. 1:e132. doi:10.1371/
journal.pone.0000132.
Omena PM, Romero GQ. 2008. Fine-scale microhabitat selection in
a bromeliad-dwelling jumping spider (Salticidae). Biol J Linn Soc.
94:653–662.
Pianka ER. 2000. Evolutionary ecology. 6th ed. San Francisco (CA):
Benjamin-Cummings, Addison-Wesley-Longman.
Rice WR. 1989. Analyzing tables of statistical test. Evolution. 43:
223–225.
Romero GQ. 2006. Geographic range, habitats and host plants of
bromeliad-living jumping spiders (Salticidae). Biotropica. 38:
522–530.
Romero GQ, Mazzafera P, Vasconcellos-Neto J, Trivelin PCO. 2006.
Bromeliad-living spiders improve host plant nutrition and growth.
Ecology. 87:803–808.
Romero GQ, Vasconcellos-Neto J. 2005a. The effects of plant structure
on the spatial and microspatial distribution of a bromeliad-living
jumping spider (Salticidae). J Anim Ecol. 74:12–21.
Romero GQ, Vasconcellos-Neto J. 2005b. Population dynamics, age,
structure and sex ratio of the bromeliad-dwelling jumping spider,
Psecas chapoda (Salticidae). J Nat Hist. 39:153–163.
Romero GQ, Vasconcellos-Neto J. 2005c. Spatial distribution and microhabitat preference of Psecas chapoda (Peckham and Peckham)
(Araneae, Salticidae). J Arachnol. 33:124–134.
de Omena and Romero • Visual cues and host plant selection
Rossa-Feres DdeC, Romero GQ, Gonc
xalves-de-Freitas E, Feres RJF.
2000. Reproductive behavior and seasonal occurrence of Psecas viridipurpureus (Salticidae, Araneae). Braz J Biol. 60:221–228.
Sokal RR, Rohlf FJ. 1995. Biometry: the principles and practice of statistics
in biological research. 3rd ed. New York: W.H. Freeman and Company.
Tarsitano MS. 2006. Route selection by a jumping spider (Portia labiata)
during the locomotory phase of a detour. Anim Behav. 72:1437–1442.
Tarsitano MS, Andrew R. 1999. Scanning and route selection in the
jumping spider Portia labiata. Anim Behav. 58:255–265.
Tarsitano MS, Jackson RR, Kirchner WH. 2000. Signals and signal
choices made by the araneophagic jumping spider Portia fimbriata
695
while hunting the orb-weaving spiders Zygiella x-notata and Zosis
geniculatus. Ethology. 106:595–615.
Taylor PW, Hasson O, Clark DL. 2001. Initiation and resolution of
jumping spider contests: roles for size, proximity, and early detection of rivals. Behav Ecol Sociobiol. 50:403–413.
Vieira C, Romero GQ. 2008. Maternal care in a neotropical jumping
spider (Salticidae). J Zool. 276:237–241.
Williams DS, McIntyre P. 1980. The principal eyes of a jumping spider
have a telephoto component. Nature. 288:578–580.
Yamashita S, Tateda H. 1976. Spectral sensitivities of jumping spider
eyes. J Comp Physiol A. 105:29–41.