COMPARATIVE STUDIES OF DICTYNA AND MALLOS (ARANEAE, DICTYNIDAE) III. PREY AND PREDATORY BEHAVIOR
PSYCHE

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Reprinted from PSYCHE, Vol. 83, No. 3-4, September-December, 1977
COMPARATIVE STUDIES OF DICTYNA AND MALLOS
(ARANEAE, DICTYNIDAE):
III. PREY AND PREDATORY BEHAVIOR
By Robert R. Jackson*
North Carolina Division of Mental Health Services
Research Section, P. O. Box 7532
Raleigh, N. C. 27611
Introduction
Although spiders are a major group of predaceous arthropods
(see Turnbull, 1973), the types of prey consumed in their natural
habitats are known for relatively few species. Some of the more
noteworthy studies have employed daw monitoring of webs of
araneids (Robinson and Robinson, 1970) and immunological tech-
niques with lycosids (Greenstone, 1978); however, very little infor-
mation is available for the dictynids. There is particular interest in
the diet of dictynids because different species in this family live
under a variety of types of social organization (Jackson, 1978).
Discussions of the prime movers in the evolution of social phenom-
ena frequently emphasize the type of prey taken by social predators
(Wilson, 1975). An important factor for some species (e.g., army
ants, canids, and killer whales) seems to be the ability of groups of
individuals acting together to handle relatively large and dangerous
prey. In order to evaluate the importance of this factor in the
evolution of social phenomena in spiders, we need information
concerning the diet and predatory behavior of species with differ-
ing types of social organization.
The species in this study belong to the closely related genera,
Mallos and Dictyna. These are small cribellate spiders (body length
usually 5 mm or less). Observations of actual feeding and other
behavior related to predation were made tn the western United
States of America in June and July, and in south-central Mexico
in September. Additional observations were made in the labora-
* Present address: Department of Zoology, University of Canterbury, Christchurch
1, New Zealand.
Manuscript received by the editor January 15, 1978.
267
Psyche
[September-December
268
tory. Also, arthropod carcasses in webs were collected and iden-
tified. Data are given as means ± S.D.
Most dictynid species are solitary, each individual generally liv-
ing alone in an individual web that does not touch other occupied
webs. Communal, territorial species (M. trivittatus Banks, D. al-
hopilosaFranganillo, D. calcarata Banks) live in web complexes,
consisting of web units connected to each other by silk. M. gregalis
Simon (communal, non-territorial) lives in communal webs not
subdivided into web units. Aggressive apd dîfcnnibalistic behavior
are virtually non-existent in this species, and indiquais routinely
feed in groups on the same prey. The other species are aggressive
and cannibalistic, and most often they feedj oh© tspMigr per prey.
In this paper basic information.’ CwBfewifng the feeding behavior
and diet of varied species Iwîl be presented, and a specific hypoth-
esis will be discussed: namely^ predation, on relatiwly large and
dangerous prey an important factor, in M gregalis? Other aspects
of the feeding behavt#$h£ .$& jgïegdlis havp; been reported eliswhere
(Burgess, 1975; Jackson, 1979a; Witt,
Data concerning M. gregalis were gathered in conjunction with
another ftudiy (Jackson, 1979a) îfj^hiçh the reader should ce&efbr
a description of laboratory method s^Large webs” were communal
webs built on plants in the laboratory, each probably containing
several hundred spiders (Jackson and Smith, 4579); and these were
not enclosed. “Small wefes^ (built by four spiders each) and “single-
female webs” were built inside plastic cafes. Data concerning where
the spider first grasped the fly came from all thro® types of webs;
data concerning size and composition of feeding groups came from
large webs only.
DIET
Diptera were the predominant prey upon which Dictyna and
Malios were observed feeding (Table 1), and these dominated the
collection of carcasses (Table 2). The data in Table 2 should be
viewed as a list of probable rather than certain prey of these species,
since some were possibly not fed upon by the dictynids. Two small
Diptera in webs of M. niveus and one small Diptera in a web of
D. iridentata were still filled with hemolymph. Probably these
were captured flies on which the spiders had not yet fed completely,
this species came from spending many hours observing a particular
Jackson — Dictyna and Mallos
269
Table 1. Number of instances of dictynids feeding on different types of prey
listed according to their estimated relative sizes (prey size/spider size). When more
than one individual fed on the same prey item ( .relative prey size
based on largest spider.
Species    Type of Prey    mailer    Number of Prey – Same – Larger size as than Spider Spider        Total
Dictyna    Diptera    #    i    1 \Iv’    3
.IlÉity^a çptplëta    |;1|p!ftera    0    …    i    3
Dictyna phylax    Diptera    ‘ iv:    ;§    o 1
Dictyna tiidêntata    Diptera    i        ‘ I    In
Mallos dugesi    Diptera            0
Mallos niveus    – Diptera                l-r
Mallos^ trivittatus        ■
&irpidopterab    . # ‘•            4
Conspecifie Spider    0    ■f    ‘ 0    9
sTi’puHdae: 14
Qjher38
My approach to the web may h^ve disturbed the spider, causing it
to depart from the prey. A living tipfl^ Shught in a M. trivittatus
web \9| be discussed later. All other carcasses in Table 2 were dry,
hollow, and almost |||tirelv intact, which is the usual condition of
prey of these spiders after feeding has occurred. Spiders inject en-
zymes into their prey, and digestion takes place primarily outside
the spider’s body. The spiders ingest the prey’s tissues in fluid form.
Unlike some other spiders, ®o noticeable mastication of the prey
occurs with dictynids. Since other species of spiders (sàlticids, tetrag-
nathids, etc.) frequently were found inside or near webs containing
dictynids, possibly some of the arthropod carcasses in Table 2 were
prey of these species, but most were probably prey of the dictynids.
Predation on conspecifics (cannibalism) is discussed elsewhere
(Jackson, 1979b).
Circadian Pattern of Feeding and Other activities
Many more data are available concerning M. trivittatus than for
the other species. Most of the observations of feeding (88%) for
[September-December
Table 2. Number of arthropod carcasses (“prey remains”) found in webs occu-
pied by dictynids. Listed according to their estimated relative sizes (prey size/spider
size). When more than one individual dictynid occupied the same web, relative prey
size based on largest spider. Unidentified Dictyna: sp. no. 1, Querecho Plains, New
Mexico, U.S.A.; sp. no. 2, Whiskey Mountain, Wyoming, U.S.A.; sp. no. 3, Lake
Chapala, Jalisco and Michoacan, Mexico.
Species    Type of Prey    Number of Prey Smaller Same Larger than Size as than Spider Spider Spider            Total
Dictyna albopilosa    Diptera    8    5    2    15
Franganillo
Dictyna annexa    Diptera    30    16    1    47
Gertsch & Chamberlin    Coleoptera    0    1    0    1
Dictyna bellans    Diptera    2    2    0    4
Chamberlin    Lepidoptera1    0    0    1    1
Dictyna calcarata    Diptera    55    16    4    75
Banks    Coleoptera    0    2    2    4
Homoptera2    2    0    0    2
Hymenoptera3    0    2    0    2
Lepidoptera    0    0    1    1
Dictyna coloradensis    Diptera    21    1    2    24
Chamberlin    Hemiptera    0    0    1    1
Dictyna compléta    Diptera    1    5    1    7
Chamberlin & Gertsch
Dictyna tridentata    Diptera    42    46    37    125
Bishop & Rudeman    Coleoptera    0    0    1    1
Hemiptera    0    0    1    1
Dictyna phylax    Diptera    9    3    0    12
Gertsch & Ivie
Dictyna sp. no. 1    Diptera    26    7    0    33
Hymenoptera4    0    0    1    1
Lepidotera1    0    0    1    1
Conspecific    1    0    0    1
Dictyna sp. no. 2    Diptera    0    4    4    8
Hymenoptera4    0    0    1    1
Dictyna sp. no. 3    Diptera    7    8    6    21
Homoptera2    1    0    0    I
Jackson — Dictyna and Mallos
271
Species    Type of Prey    Number of Prey Smaller Same Larger than Size as than Spider Spider Spider            Total
Mallos dugesi    Diptera    3    1    2    6
Becker
Mallos niveus    Diptera    57    38    18    113
O. P. Cambridge    Coleoptera    0    3    2    5
Homoptera2    0    1    0    1
Hymenoptera4    0    1    2    3
Orthoptera5    0    0    1    1
Thysanoptera    2    0    0    2
Salticid spider    0    0    1    1
Mallos trivittatus    Diptera6    163    20    38    221
Banks    Coleoptera    1    0    0    1
Homoptera2    3    0    0    3
Hymenoptera3    1    0    0    1
Lepid optera    0    11    5    16
Neur optera    1    0    0    1
Conspecific    2    3    0    5
‘Moth
2Aphid
3 Ant
“Wasp
’Grasshopper nymph
6Tipulidae: 33
Other Diptera: 188
web complex, located in a culvert through which a creek passed in
the Chiracahua Mountains of Arizona. This large web complex
was estimated to contain more than 10,000 individuals of trivit-
tatus (Jackson and Smith, 1979). Since initial observations sug-
gested that feeding occurred predominantly in the late afternoon
and early evening (see below), one hour was spent inside the culvert
on each of 12 evenings (5 in June; 7 in July); and records were kept
for all observed cases of feeding. Diptera and other insects in the
vicinity were especially active at this time of the day, and this was
generally true in other habitats of M. trivittatus and the other dic-
tynids.
[September-December
Table 3. Temporal pattern of activity of spiders in their natural habitats. Time
of day: early morning and early evening, within 2 hr before and after sunrise and
sunset, respectively. Duration of observation estimated. Walking: without spinning
and exclusive of intraspecific interactions. Intraspecific interactions described else-
where. (Jackson, 1979b). Dictyna phylax and Malios dugesi observed in day only.
Species    Time of Day    Duration of Observation (hr)    No of Spiders Feeding    No. of Spiders Walking    No. of Spiders Spinning    No. of Intraspecific Interactions
Dictyna calcarata    Early Morning    3    2    5    6    1
Day    5    0    0    0    0
Early Evening    2    1    0    0    0
Dictyna compléta    Early Morning    2    1    0    0    0
Day    4    2    0    0    0
Early Evening    2    0    0    1    0
Dictyna phylax    Day    6    2    0    0    0
Dictyna tridentata    Early Morning    6    1    2    0    2
Day    14    0    0    0    0
Early Evening    4    1    0    0    0
Mallos dugesi    Day    7    1    0    0    0
Mallos niveus    Early Morning    5    0    0    0    0
Day    14    2    0    0    0
Early Evening    5    0    0    2    1
Mdllos trivittatus    Early Morning    17    0    3    0    0
Day    34    3    0    0    0
Early Evening    19    53    18    3    9
Jackson — Dictyna and Mallos
273
With the exception of the evening observations in the culvert,
the amount of time spent observing webs was recorded only ap-
proximately. These estimates were used for the calculations in
Table 3. Based on these data, It seems that feeding and general
activity of the dictynids in this study occur predominantly in the
evening.
Initial Contact op Spider WfpfIPREY
Certain spiders, such ts some araneids and theridiids, wrap their
prey either before and/ftr after biting; however, this does not occur
in the Dictynidae. Tbeêe spiders seem to simply rush out and bite
the prey. If the prey is Violep|b| struggling, the spider may walk
or stand§Sl the vicinity” ufttil JtWrvjity subsides.
Bristowe (1958) reported that dictynids invariably grasp their
prey initially by 4 leg. The initiation of fefling was seen for one
Mi tmaeus £§$ five M.    In    .each    caite,    the    spider    initially
grasped a leg antenna of the prey. Of the spiders already feeding
when found, some were feeding on the head, thorax, or abdomen
of the prey (Fig. 1), although data were not recorded. M. gregalis,
M.    trivittatus,M.    and D. çalcarata were maintained and
fed in the laboralplii^ and it was noted that the spiders sometimes
initially grasped the prey by its head or body rather than by an
appendage. For M. gregmlis in fj§| laboratory, the location at
which the spider first grasped the» prey was recorded for 66 indi-
viduals: leg»’4H; head, 15%; abdomen, 14$$; thorax, 11%; wing,
9%; antenn^,    of    these    flies    were    active    when    contacted.
Once I saw an opilionid walk    web    unit    containing an
adult female M.    The spider rushed out of its nest and
grasped a leg of the opilionid with Its chelicerae. Immediately, the
spider released the opilionid and returned to its nest, suggestive of
opilionids being distasteful to dictynids (see Bristowe, 1941). Sev-
eral minutes later, the opilionid escaped from the web.
Extension Lines
Webs of M. trivittatus frequently contain long, heavy lines of
silk (extension lines) that extend to objects some distance from the
mesh (Jackson, 1978). Once in Utah I found an extension line
fastened at one end to a mesh, with a female M. trivittatus inside
the nest. On the other end, a tipulid fly was tethered by its thorax.
Psyche
[September-Decembe
Fig. 1. Adult female Mallos trivittatus (body length: 7 mm) at East Turkey
Creek (Chiracahua Mountains, Arizona) feeding on tipulid fly. Fly grasped at
ventral thorax.
The tipulid flew in circles continuously for 10 min while I observed,
after which I collected the fly and the spider. Of the set of M. tri-
vitattus observed feeding in nature, 9% were on extension lines at
the time; and 5% of the arthropod carcasses found in webs of M.
trivittatus were found on extension lines. M. gregalis webs also
have extension lines, and these spiders sometimes fed on flies caught
on extension lines.
1977]    Jackson    —    Dictyna    and Malios    275
Feeding Groups Size and Composition
The few cases in wMch more than one spider fed on the same
prey in species other than M.    are    described    elsewhere
(Jackson, 1979b). In the laboratory, the size and composition of
the group feeding on the fly was fecorded 15 min after it contacted
the web, and cases lf§ which no sliders were feeding at the end of
the 15 min are excluded. Group size was 4.8 8*1.96 spiders (range:
1-15;H= 38). In the cases in which a single spider fed on the fly,
three were females, two were immatutès, and none were males. One
of the immatures was a second instar; the other was almost adult size,
In^eJ&ès in^wEMr’iffoi^han one spider fed    were
three groups consisting of    femal^éà
females and itnrtatuTes but no males; 2, jéSi& ând immatures but
ife females; ari$    and    immatures.    casual
observations, single males feeding’tH flies and groups consisting’#’;
females and malyfe » immatur^.’W^^i^^feut groups of more
than onW lBf’;but no females or finmatwéfe?%ife%#f noticed.
Grotfps t# m<&e than ftrindividuals have beeif sèëÉ^^
Based on arthropod Wg^^wund ura|^Kljffid observations of
actual feeding in nature, Diptera seem to constitute the major prey
of the closely related species of D and in this study.
Billaudelle (1957), Bristowe (1958), and Wiehle (1953) commented
on dictynids ifeSfc.nn Diptera,’ ants, and lice. Unfortunately,
only limite# i^Mlation is available con^&ing the natural prey
of M.    gregaliSmÊÊcommunal,species. I was not
able to find this species when I was in Mexico. Diguet (1909a, b,
1915) and Burgess (1976 and personal communication) noted that
Diptera seem to be the primary prey of this species in nature, al-
though wasps are also fed upon. The Diptera seem to be predom-
inantly ones of body lengths of approximately 5 to 10 mm, such
as the “domestic fly” (presumably Musca domestica), tabanids, and
bot flies. Burgess collected a portion of a web in Mexico; and
when examined in the laboratory, it contained a great number of
carcasses, all of Diptera in the size range of 5 to 10 mm. In the
laboratory, M. gregalis has thrived for several years on a diet of
M.domestica almost exclusively. The natives of Michoacan have
Psyche
[September>December
276
given this species the name el mosqu During the rainy season,
they take portions of communal webs from trees and place these
in and around their homes, using them as fly traps (Berland, 1913;
Diguet, 1909a, b, 1915; Gertsch, 1949).
Burgess (1975) has demonstrated that vibrations within a fre-
quency range comparable to the wing beat frequency of Musca
domestica is the most effective stimulus for eliciting predatory be-
havior from M. gregalis. Furthermore, the web transmits vibra-
tions within this frequency range more readily than ones with other
frequency characteristics. It seems that the web has characteristics
that are particularly appropriate for the predominant prey species.
The vibration transmission properties of webs of species have
not been investigated jp&;
Some Diptera may be captured when they fly into Dictyna and
Mallos webs. However, it was noticed that many Diptera tend to
land on the stems and leaves of herbs and shrubs, on rock ledges,
and on other olÿjÿf on    dictyndds    tend to build their webs.
Perhaps the majority of Diptera are captuftd when they inadver> ■
tently use ê> web as a perch. Musm doMestiça were frequently
captured, Seemingly in this manner, on webs of M. gregalis in the
laboratory. These webs were kept in the open, on plain and other
objects. Id the laboratory. During routine feeding, house flies were
thrown in% Jtie comrai^li webs, but many inadvertently escaped
into the room beforehand. Frequently these were seen subsequently
landing on the webs and adhering to, the S§|§. Thrown flies would
seem more comparable to flying Digteta,; and there is no evidence
that the ratio of flies captured to ones that escaped differed for flies
landing on the web compared ter ones thrown into the web (Jack-
son, 1979a).
The extension lines in webs of M. gregalis and M. trivittatus
may have a function related to predation. Diptera may find them
to be particularly attractive perches and become trapped when they
land on them. Another cribellate species, Miagrammopes (Ulo-
boridae) has a single thread snare, and it reportedly captures Dip-
tera that use the thread as a perch (Akerman, 1932).
Dictynid webs have nests, which are tubular structures of more
densely woven silk; and the spiders tend to reside in their nests
when not active. Spiders in various families (e.g., Agelenidae, Eresi-
dae, Dysderidae) which have nests in their webs often transport prey
to the nest before feeding(see Bristowe, 1958; Krafft, 1971). Araneid
Jackson — Dictyna and Mallos
277
spiders tend to transport prey to the hub of the web before feeding
(Robinson and Olazarri, 1971), Although data were not collected,
it was noticed that arthropod carcasses tended to be concentrated
near the nests of the solitary and the communal, territorial species;
and many of the feeding dictynids were near their nests at the time.
These observations suggest that dictynids transport prey to their
nests, although actual transport has not been seen. Billaudeie
(1957) noted that D.    carries    prey from the periphery to
the center of the web.
Most dictynid webs tend be 2-dimensional;’i.é., most of the
silk of the web t& in a single plane. In contrast, the communal
webs of M. gregalis tend to be 3-dimensional; and the nests are in
the interior of the webs, beneath the surface sheet on which flies
are captured. Although flies were oeeasionally pulled into the in-
terior of Webs by spiders, in the vast majority of eases the prey was
fed upon at the capture site in communal web’s in the laboratory’.’ ■
Returning to fhrhypothesis proposed at the beginning of this
paper, ü tifte ‘g>r#ÿ MM. gregalis rebt&itely large and dangerous
compared that of other dictynids? Diptera are apparently the
primary prey of most species. Since Diptera such as muscids/ culi-
cidids, êtè.^ould n$$Pièfem elpfj^fpÿ! dangeroiïsïor dictynids, dif-
ferences in the danger associated with    preÿvwould not
seem important. Adult females of M.    the largest sex/ age
class, tend1 to weigh 4 to 21 mg, adult Mtt’sëa dfamestica tend to
weigh 10 to 20 mg (Witt, et al., 1978jÇfff prey of M. gregalis is in
this weight range, then prey tends to range from approximately
equal in size to individual spidersSib a few times larger. In the
solitary and in the iommunulg- territorial species, prey were often
smaller than the spiders. However, the difference in relative prey
size among species is not absolute. Many prey of solitary and
communal, territorial species were equal to or larger in size than
the spiders (see also Bristowe, 1958; Wiehle, 1953).
Since prey sizes overlap for different dictynids, we need quanti-
tative data from which variances can be calculated for relative prey
size. Data from the natural habitats of M. gregalis in Mexico are
especially needed. It will be tentatively concluded that M. gregalis
preys primarily on relatively large prey. However, the differences
in relative prey size do not seem dramatic. In a sense, the social
organization of M. gregalis seems very different from that of the
other dictynids, with great numbers of spiders living and feeding
Psyche
278
together in the same communal webs. If diet is a major factor in
the evolution of social phenomena in dictynid spiders, we might
expect the diet of M. gregalis to differ greatly from that of other
dictynids. Although differences in prey size seem to occur, perhaps
the most interesting finding in this study is that there is consider-
able overlap in prey sizes of different dictynids. We need to con-
sider the possibility that predation on relatively large and dangerous
prey is only one among other equally or more important factors
acting as prime movers In the evolution of social phenomena in the
Dictynidae and perhapiifor other groups as well.
^Hwary
Based on arthropod carcasses in    observations    of actual
feeding, Diptera seems to be,the maj    and    Mallos.
M. gregalis, a species that routinely feeds in grouggg may tend to
prey upoxfr^ëlafS^ll large mseV Compared to He, other species.
However, reltf    ■ p^tfep for species of all types of –
iVfpsial organization. apparent differences occur in the degree
to which prey dangerous. These observations ap? not to be
expected from the^^^th^sis thp the prime mo\»||f $£$ evol%
tion of social phenpjnena in spidffs^The a^|y of predators act-
ing as a group to handle relatively large and dangerous prey. Al-
though legs of flies «ar^frequently grasped    may
initially grasp almost any- part of the $)&4$ize of feeding groups
varies greatly, ranging from 1 to more than 20. Thejlg|$fjtaii:is
proposed thafe^py is captured    and    Maljjfgg    primarily
when flf^t use Webs as resting sites. Feeding and other activity
occur especial^’ ^ |be early evening and early morning.
For valuable discussions and comments on the manuscript, I
would like to thank P. N. Witt, M. C. \p|^k, £>. E. Smith, and J. W.
Burgess. Special thanks go 40 W. J. Gertsch for his assistance in
the identification of spiders. C. E. Griswold, P. S. Jackson, and
V. D. Roth are gratefully acknowledged for helping me locate
spiders in the field. The assistance of the Southwestern Research
Station of the American Museum of Natural History is gratefully
1977] Jackson — Dicîyna and Mallos
279
acknowledged. Thanks go to R. B. Daniels for typing the manu-
script. This work was supported in part by the North Carolina
Division of Mental Health Services, Research Section and by
N.S.F. grant number BMS 75-09915 to P. N. Witt.
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1957.    Zur Biologie der Mauerspinne Dictyna civica (H. Luc.) (Dictynidae,
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1941. The comity of spiders. Vol. II. ^London: Ray Society. 560 pp.
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