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GENERAL NOTE: This republication is intended for free dissemination at no charge. Please attribute the source correctly. Please address all suggestions
and corrections to the author. This version and republication copyright©2006 by David Edwin Hill.
David E. Hill
213 Wild Horse Creek Drive, Simpsonville, South Carolina, 29680
platycryptus@yahoo.com
Original pagination is retained for reference in this republication of the article originally published as: Hill, D. E. 1978 The
behavior of Eris marginata. Peckhamia 1 (4): 63-70. All corrections, annotations, or additions are highlighted in [red]. The
original version of this paper was completed on May 20, 1975, in Corvallis, Oregon. Figure 7 has been added to include
some more recent observations of Eris in South Carolina. All of the other illustrations presented here were included in the
original (1975) version but were not included in the published Peckhamia version.
63
[Introduction]
page 1 of 10
Hill, D. E., 2006: Behavior of Eris marginata [RV1]
[Figure 2. Blackberry bush (Rubus) in Corvallis, Oregon. The stems and leaves of this plant are
inhabited by salticids of at least five genera, notably Eris and Pelegrina. The most conspicuous
species is Eris militaris, particularly in May when the courting males are common.]
[Figure 3. The author in the bush, looking for jumping spiders (Witham Hill, Corvallis, Oregon,
May 1975).]
Throughout the period of this study, individual spiders [The spiders were originally identified by descriptions
were observed continuously for varying periods of time provided by Peckham and Peckham (1909) and Kaston
ranging up to more than three hours. By remaining (1972, 1973). The distinctive sexually-dimorphic males
relatively still, the influence of the observer upon the and females (Figures 4, 5) are easily recognized in the
behavior of the spiders was minimized. Observations field.]
were facilitated by a combination of mild climate and a
complete lack of biting insects.
page 2 of 10
Hill, D. E., 2006: Behavior of Eris marginata [RV1]
Diurnal activity patterns Eris walk almost equally well on the underside of a leaf
or stem as they do on the top, although they use the upper
The life of E. marginata [E. militaris] is greatly regulated surfaces most often. In addition to the usual forward
by warmth and sunshine. The spiders generally travel on advance, Eris are quite capable of side-stepping and
the upper, illuminated surfaces of blackberry leaves and walking backward at variable speed. They often side-step
stems. They often move into the sunshine when feeding (lateral movement) quickly to a concealed position on the
upon prey. In the morning these spiders, emerging from underside of a stem or leaf when threatened by an
their shelters, flatten themselves along a leaf or stem in a observer moving overhead. Beyond the basic patterns of
normal orientation [in a plane, perpendicular] to the solar walking and other forms of movement, there are no real
radiation, perhaps to maximize the effect of the stereotyped sequences. Thus a jump may precede another
irradiation [to warm up]. When sunning in this manner, jump, or a run, or a series of slow turns. The sequence in
the appendages are held close to the body and the spider which each element of the locomotory repertoire is
moves little. The dorso-opisthosomal [dorso-telsomal] employed is largely determined by the environs. Thus, if
situation of the heart may greatly facilitate the rapid a spider can reach between two leaves, it may continue a
distribution of heat throughout the body of the spider. In walk. If not, it may jump, or turn to reach another object.
this regard it is significant that the opisthosoma [telsoma] Movement involves continuing interaction with
may be pivoted to bring the dorsum to face the sun more immediate possibilities [identified through both visual and
directly. Eris appear to be most active in the mid-day tactile (contact with silk or surface) senses, by these
heat. It is at this time that most active travel and spiders].
exploration takes place. It should be noted that the
diurnal activity pattern of this species may be quite The most commonly employed walking pattern may be
different in a different part of the United States. The termed an exploratory walk. In this walk the spider takes
Willamette Valley of Oregon cools quickly as evening a variable number of steps, then stops, then takes a few
approaches, and when compared to other areas, the mid- more steps, then stops, and so on. After several intervals
day heat is really rather mild (perhaps 20C). of this pattern, the spider may pause to survey the
environs systematically, pivoting the prosoma to orient
In the early evening, long before nightfall, Eris begin to the large anterior medial eyes (AME) in various
construct and occupy shelters between adjoining directions. Information received during this pause may
blackberry leaves. The spiders remain in these shelters lead the spider to alter its course completely (see below).
throughout the night, and emerge the next day only if the Periodically, during a pause or long stop, the omnipresent
weather is favorable. As the air cools in the evening, the dragline, a strand of strong silk continuously released by
spiders often may be seen sunning as in the morning. the moving spider, is secured to the substratum with an
adhesive attachment disk. Although this forward walk is
Locomotion characterized by steps, stops, and pauses, the actual
pattern is quite variable, as is the speed of the moving
Eris walk slowly about in their shelters in the evening, if spider.
they move at all, and are quite sluggish until they have
warmed up in the morning (unless threatened). The A fast moving spider running along a main stem may take
spiders are only active when the warmth and illumination many steps between stops, and may turn quickly to face
are favorable, and it is thus that these spiders, particularly each of several different directions only briefly during
the each infrequent pause. A slower spider may take only
one or two steps before pausing and surveying
64 systematically in all directions for a prolonged period of
time. Again, a fast moving spider may enter a slower,
vagabond males, engage in the greatest amount of travel exploratory phase of behavior at any time. Thus a male
in the middle of the day. Like other jumping spiders, Eris running along a stem may encounter an object of visual
employ at least five basic methods in their movement interest, such as a moving insect, and will subsequently
upon vegetation. Most of the time they are walking at move very slowly with frequent turns to survey the
various speeds (or running), with pretarsal foot pads in environs. Beyond basic feedback from proprioceptors
contact with the surface. They often jump to a sighted [either body position sensors or surface contact sensors],
target position. They may descend (rappel) on a dragline, complex visual stimuli are critical determinants of the
or climb a vertical dragline. Finally, like many other pattern of locomotion.
spiders, they can construct and climb on the underside of
a horizontal bridge of silk, or similarly use a thin plant
stem as a bridge.
page 3 of 10
Hill, D. E., 2006: Behavior of Eris marginata [RV1]
The actual stepping pattern consists of a loose alternation Eris may ascend either a newly-formed or a pre-existing
of legs I and III with the ipsilateral legs II and IV, and vertical dragline very quickly, utilizing a characteristic
contralateral legs I and III (see Land 1972). Pedipalps are vertical climbing behavior (found in many spiders)
often moved up and down, although this activity is not involving the handling of silk strands alternately by legs I
restricted to a period of forward locomotion, but also and II, with legs III and IV outstretched. During this
occurs during each pause. In what is perhaps an ascent, the silk line is wound into a ball which is
anthropomorphic sense, one might interpret this pedipalp immediately discarded as the spider attains its objective
flicker of salticids to be indicative of the state of general [Figure 6].
excitation of these animals. The legs I are frequently
raised in the air and flexed at the junction of femur and
patella in a sort of gesture. Infrequently the walk
becomes a relatively slow, continuous but jerky (with
many brief stops) advance. This pattern of jerky walk is
as distinctive as it is difficult to describe. [During this
staccato walk each step is actually broken into a sequence
of separate, rapid movements of the respective legs.]
Similar behavior has been observed in several other
salticids, including Phidippus [and Platycryptus].
65
The jump of Eris, like that of other [but not all] salticids
(Parry and Brown 1959), depends primarily upon force
exerted against the substratum by legs IV. Eris jumps
clumsily and inaccurately between grass blades which are
generally too weak to support this thrust. On blackberry
leaves, Eris often walk backwards to secure a foothold for
the jump. Each jump is preceded by visual examination
of the target object with the AME. The spider crouches
and flexes legs IV at the tibio-metatarsal joint, and raises
legs I and II (flexed at the femur-patella joint) to receive
the new surface. Just before each jump, the opisthosoma
[telsoma] pivots to bring the spinnerets into contact with
[or near] the substratum, and the spider releases an
adhesive mass of silk to form an attachment disk for the
dragline. When a jump is missed, the spider hangs by the
dragline and is thus able to recover to its former position. [Figure 6. Vertical descent and ascent with the dragline. The male at
A jump involves the exertion of a calculated thrust to left is stopping during a descent. The female at right is climbing a
project the spider upon a ballistic flight for the required dragline with the first two pairs of legs, with legs III and IV outstretched
distance, which may involve 1-20 cm of horizontal flight. (vertical climbing behavior).]
page 4 of 10
Hill, D. E., 2006: Behavior of Eris marginata [RV1]
16
17
A
walk and
15
pull line
14
13
walk and 7 12
pull line
10 11 climb under
6 line
3
8 9
climb under line
5
1
2 46
4
short jump
45
42
Eris militaris 44
near Brashier Pond
S. Greenville County 41 43
South Carolina walk and pull line 40
A-E: female NOV-8-2005 35 38 39
F: immature AUG-24-2005
33
37
31 climb under line
34 36
32
29
30
28
F
walk and pull line 27 climb under line
26
25
C
24
walk and
pull line 23
20 22
18 21
17
climb under line
19 D E
[Figure 7. Recent observations of Eris militaris in the southern part of Greenville County, South Carolina.]. A-B: Sequential (1-46) positions of an adult, or
near-adult, female (~6-7 mm) following and using a silk line to complete a course (~70 cm) across leaf litter (photomontage from 320x240x30fps
Quicktime® movie). This spider originally dropped a dragline and let it out into the air, but the dragline fell down to the ground pulling the spider down
with it. The spider alternately pulled on the silk as it walked with legs I outstretched (blue arrows), and climbed under the silk line where it was stretched
between leaves (white arrows). This is a good illustration of the ability of salticids to maintain a course even if interrupted. Turns in position (blue
rectangles) are attached to red rectangles indicating where they took place. Short green arrows indicate the direction faced by the spider at each position. 1,
After the line fell, the spider turned to look directly at the observer. 2-3, Advancing and pulling on the line. 4, Sudden turn to look at grass blades moving in
the wind. 5-8, Advancing and pulling. 8-10, Fast climb under the line. 10, Stopped and went behind a leaf to watch an ant. 11-13, Advancing and pulling.
13-15, Climb under line behind leaf. 16, Sudden turn. 17-18, Pulling. 19-24, Climb under line. 24-25, Walk. 25-26, Short jump. 26-28, Walk to tip of
leaf. 28-31, Climb under line. 31, Behind leaf, watching ant. 31-32, Walk. 33, Sudden turn. 34-36, Small turns, reaching for line. 36-40, Climb under
line. 40-42, Climb under line. 42-43, Upside-down jump. 43-46, Advancing and pulling. C-E: Close-up views of the spider shown in A-B. F: Immature
ballooning from the top of a shrub. The dragline is highlighted in white to improve its visibility in this picture. Eris of all ages bridge much more than do
many other salticids. Bridging behavior has never been observed in the related genus Phidippus. This spider would have been able to avoid the leaf-litter
(not its microhabitat of choice) if the bridge had not fallen to the ground after the spider started running it. Eris move very fast on these bridges.
page 5 of 10
Hill, D. E., 2006: Behavior of Eris marginata [RV1]
The spider then turns and pulls in the slack on the bridge At times the spiders consistently ascend as they climb or
with legs I, perhaps testing for attachment. Then the jump from leaf to leaf. Often they will survey a position
spider climbs quickly along the underside of this with the AME, and then employ an indirect route to attain
horizontal bridge, using all legs in what probably that sighted position, as necessitated by the arrangement
corresponds to the normal stepping pattern, to attain its of branches and twigs. Visual survey plays a primary
destination [Figures 7, 8]. It is of interest that a brief role in the determination of a course by Eris.
descent on the dragline invariably precedes the formation
of a bridge. It is likely that this behavior is necessary to Sensory activity
draw out the initial strands of silk to such a length as may
be captured, and subsequently drawn out, by the wind Eris constantly use the front eyes (AME, ALE) to survey
itself. As noted above, the descent which precedes the the environs during periods of activity, as noted above.
formation of a bridge is distinct from the rappel, in that During pauses, the spiders can employ a series of small
the former involves a rapid turn to ascend without pause. turns of the prosoma to survey the surroundings. They
may turn directly toward each of several nearby leaves in
succession, perhaps indicating that the lateral (ALE and
PLE) eyes may be capable of directing movement toward
a stationary object. [This behavior has also been
observed many times in Phidippus. This does not support
the oft-cited assumption that the lateral eyes of salticids
are used only for movement detection.] The AME and
ALE are brought to bear upon an object by turns of the
prosoma, which can be turned through a small angle
without stepping. In addition to lateral movement, the
front of the prosoma (face) may be raised to look up, or
lowered to look down. Eris will turn to face moving
objects (of sufficient size) at distances ranging from
several mm to more than 10 m [Figure 9].
[Figure 8. Bridging. At top an inverted female I ballooning silk with the wind.
Below the female climbs quickly under the resultant bridge.]
66
page 6 of 10
Hill, D. E., 2006: Behavior of Eris marginata [RV1]
It is well known that the AME of salticids are utilized for When the prey is held securely, after initial maceration or
the detailed evaluation of the immediate surroundings and mastication with fangs and chelicerae, Eris carry the prey
the determination of a course of movement through the to a feeding position. In the evening, spiders may carry
vegetation. These spiders are remarkably alert, and the the prey into the shelter to feed. At mid-day, a feeding
accuracy of each turn to face a stimulus is remarkable. spider may move into a sunny position with a good view,
Eris will observe an interesting object, such as a hunting such as the top of a leaf or stem. When feeding, the
wasp or a human, for a prolonged period of time without spider crouches against the substratum, holding the legs
moving. against the body. Fluids are rhythmically pumped in and
out of the prey. Eris occupy the better part of an hour
In addition to the visual sense, Eris display a great deal of feeding on a small
tactile sensitivity. The forelegs in particular are often
tapped against the surface. In addition the pedipalps are 67
frequently brought to touch the underlying plant. This
behavior may facilitate the role of chemosensory as well insect, such as a leafhopper, although the time of feeding
as mechanosensory setae. on even a particular prey item is quite variable.
Subsequently the remnants of the prey are dropped and
Predation on insects the spider grooms in place by rubbing the pedipalps in
loosely alternate fashion in front of [over the front of] the
Eris jump directly upon sighted prey, generally small chelicerae.
insects, which are encountered during a journey through
the vegetation. If the prey are detected at more that 1-2 Grooming
cm distance, the spider may creep slowly toward the prey
with pedipalps flickering up and down, to jump at a As mentioned above, the chelicerae are fastidiously
distance of 1-2 cm. These spiders will also hurtle groomed after each feeding. The chelicerae are also
themselves into the air in pursuit of flying insects. The rubbed against the pedipalps at other times. The
predatory jump is similar to the locomotory jump, apart pedipalps may on occasion be rubbed against each other
from the grasping behavior with chelicerae and legs I and as well, or used to groom the first leg. The hairy
II which is included in the former. The attack is preceded pedipalps function as the vertebrate eyelid or nictitating
by formation of an attachment disk [Figure 10]. membrane, to brush the vital AME clean. The lateral
surfaces of the legs and pedipalps may be brushed against
the substratum, or against adjacent appendages. At times,
any of the leg tips, including each pretarsus, may be
cleaned between the endites and chelicerae. This
grooming may facilitate the efficient adhesion of these
structures, perhaps as the preening of a bird contributes to
flight (see Hill 1977). Grooming occurs during a pause in
activity. It is commonly seen when the spider is resting
(so to speak) in a shelter or concealed position. Extensive
grooming follows feeding, but it may also occur after a
brief pause during a journey. A spider walking along a
stem may pause, turn in several directions, clear the tip of
a rear leg between the endites, and then continue the
walk.
page 7 of 10
Hill, D. E., 2006: Behavior of Eris marginata [RV1]
Intra-specific relations
68
[Figure 12. Cohabitation of male and female. The male (below) is adding silk to
the framework of the lower chamber. The female grooms in the sealed upper
chamber.]
page 8 of 10
Hill, D. E., 2006: Behavior of Eris marginata [RV1]
page 9 of 10
Hill, D. E., 2006: Behavior of Eris marginata [RV1]
Discussion REFERENCES:
The response of these small (5-7 mm) spiders to visual Enders, F. 1975 The influence of hunting manner on prey size,
particularly in spiders with long attack distances (Araneidae,
stimuli at distances of at least 10 meters is remarkable, as
Linyphiidae and Salticidae). American Naturalist 109: 737-763.
is the almost unerring accuracy of their jumping behavior
in three dimensions. [Griswold, C. E. 1987 A revision of the jumping spider genus
Habronattus F. O. P.-Cambridge (Araneae; Salticidae), with
Although E. marginata [E. militaris] is found in other phenetic and cladistic analyses. University of California
neighboring habitats at times (such as upon the heavy Publications, Entomology 104: i-ix, 1-344.]
horizontal bars of chain-link fences in the area), it is most
often encountered on the larger blackberry bushes. It is Hill, D. E. 1977 The pretarsus of salticid spiders. Zoological
Journal of the Linnean Society, London 60: 319-338.
seldom encountered with a sweep net in grassy areas that
are far removed from this habitat. Several aspects of the [Kaston, B. J. 1972 How to know the spiders. W. C. Brown
behavior of this species may help to explain this Company, Dubuque, Iowa: 1-289.]
distribution or microhabitat preference. First of all, the
horizontal leaves [or leaflets] of the blackberry are [Kaston, B. J. 1973 Four new species of Metaphidippus, with
adjoining and often overlapping at inner angles, thus notes on related jumping spiders (Araneae: Salticidae) from the
affording these spiders their typical shelter sites. Eris eastern and central United States. Transactions of the American
tend to move toward the elevated positions of blackberry Microscopical Society 92: 106-122.]
bushes relative to other plants in an open field. The
Land, M. F. 1972 Stepping movements made by jumping
bushes grow in open, unshaded, habitats which afford
spiders during turns mediated by the lateral eyes. Journal of
Eris the sunshine which they seem to enjoy. It could be Experimental Biology 57: 15-40.
said that the radiation requirements of the blackberry leaf
and this jumping spider coincide. The strength of the [Maddison, W. 1986 Distinguishing the jumping spiders Eris
stems and leaves of the blackberry is more than adequate militaris and Eris flava in North America. Psyche 93: 141-149.]
to support the jumps of this spider, which moves clumsily
through grasses. The same plant structure also facilitates Parry, D. A., and Brown, R. H. J. 1959 The jumping
the visual activities of Eris by providing relatively large mechanism of salticid spiders. Journal of Experimental Biology
vistas and extensive surfaces for movement, including the 36: 654-664.
avenues of large stems. With foot-pads, extraordinary
[Peckham, G. W., and Peckham, E. G. 1909 Revision of the
powers of vision for a creature of its size, and a complex Attidae of North America. Transactions of the Wisconsin
repertoire of behavior, Eris is well-suited for its role as a Academy of Science 16: 355-644.]
little tiger of the blackberry bush.
Richman, D. B. 1977 The relationship of epigamic display to
Among salticids, one might classify Eris marginata [Eris the systematics of jumping spiders (Araneae: Salticidae). Ph.D.
militaris] as a searcher, rather than a pursuer ([dichotomy Thesis, University of Florida: 1-162.
offered by] Enders 1975). Nonetheless one should
entertain the possibility that activity patterns in a
particular habitat may vary greatly from those observed
elsewhere, even for the same species, depending upon
relative availability of sedentary and actively moving
prey species.
page 10 of 10