253

THE USE OF BAITS AND PRESERVATIVES IN PITFALL TRAPS

By PENELOPE
GREENSLADE*
and P. J. M. GREENSLADE?
[Manuscript received June 17, 19711

Abstract
Pitfall traps containing baits and alcoholic preservatives were investigated, in the Solomon
Islands and in South Australia, as methods for sampling ants and members of other groups
that are active on the ground surface. Observations by previous workers that there may be
attraction to preservatives used in traps were extended and a method of testing for it is
suggested. Ants were not attracted to the preservatives used and when dealing with diverse
ant faunas simple pitfalls appear to be preferable to baited traps.

INTRODUCTION
Although pitfall traps have many disadvantages, they can be used in studies of
the occurrence and activity of invertebrates which move on the surface of the
ground (Greenslade 1964, Southwood 1966). If traps cannot be cleared frequently
they are often used with the addition of a killing agent and preservative. However,
Luff (1968) found that various Coleoptera were attracted to formalin used in this
way in England and van der Drift (1963) suggested that scolytid beetles were
probably attracted when alcohol was used in pitfalls in South America. Additional
cases of attraction to preservatives are described here and occurred while baits
and pitfall traps were being examined as quantitative or semiquantitative methods
for sampling ants and for assessing the composition of the soil surface fauna when
ants are dominant.
Observations were made in the Solomon Islands in 1964-1966, mainly in low-
land tropical rain-forest on Mt. Austen on Guadalcanal where the local ant fauna
exceeded 100 species. Further experiments were carried out in the Mediterranean
climate of the Adelaide district of South Australia at Glen Osmond (Aug.-Nov.
1968) and Belair (June, July 1969) in sheep-grazed pasture and savannah woodland;
here the number of ant species in a locality was about half that in the Solomon
Islands but they were still a dominant group on the ground surface.
RESULTS
Use of baits for sampling ants
Some ant species inhabit large, distinct nests which can be sampled destructively
while other species are restricted to soil and litter and are amenable to the same
sampling and extraction methods that are available for other soil animals. But
neither of these approaches can be applied to an important intermediate group
whose workers forage on the ground surface out of diffuse nests under stones or in
litter, or from small nests in the soil. The frequency and territories of such species
can be determined by distributing baits on the ground in their habitats (Brian,
Hibble and Kelley 1966) but this method has several disadvantages if there are
many species in the local ant fauna. First of all, baits are very selective and may not
attract species lacking behavioural flexibility ; in South Australia for example,
foraging workers of Myrmecia pyriformis F. Smith ignored baits of sugar in the
field although they will feed on it in laboratory cultures. Baits of sugar (sucrose)
or syrup (sucrose in water) do not attract exclusively predacious species which are
numerous in rain-forest and if baits of meat or dead insects are used they are often
monopolised by a dominant species; in the Solomons this was commonly Zridomyr-
mex cordatus F. Smith in forest and Solenopsis geminata (F.) or Paratrechina
longicornis (Latreille) in other habitats. The number and species of ants visiting
baits also vary with time of day and transient changes in weather. At Glen Osmond,
South Australian Museum, Adelaide, South Australia. Work in the Solomon Islands was sup-
ported by a grant from the Royal Society and this is gratefully acknowledged.
t Division of Soils, C.S.I.R.O., Glen Osmond, South Australia, 5064.

J . Ausf. ent. Soc., 1971, 10: 253-260.

254 PENELOPE GREENSLADE and P. J. M. GREENSLADE
TABLE1
CATCHES OF ANTS IN DIFFERENT BAITED TRAPS, GLEN OSMOND, SOUTH AUSTRALIA
Control
Species (Alcohol- Beer Beer-Syrup Syrup
glycerol)
Number of Individuals
Iridomyrmex spp. - 2 34 180
Chelaner sp. I 9 146 61 1
Pheidole spp. - 1 - 2
Camponotus spp. 1 - 5 3
Other spp. 2 - 5 4
Total 4 12 190 800
Number of spp. 3 4 I I

for example, two sets of 10 baits, A and B, consisting of paper soaked in a sugar-
water syru in petri dishes, were set out at 10 a.m. By noon it was noticed that baits
A, in shac re, had few ants but they were abundant at B, then in direct sunlight.
Later, when baits A were in sunlight, 450 ants belonging to four species were
recorded at them but only 15 ants representing two species were found at baits B,
which by this time, were shaded. Usually a trap is required to retain the catch and
reduce this source of variation. In trials at Glen Osmond, specimen tube traps
(mouth diameter 1.8 cm, 7.5 cm in depth) were partly sunk in the ground; in one
series of experiments baits of syrup, beer (which is attractive to some social Hymen-
optera (Thomas 1960)), and a beer-syrup mixture were compared with alcohol
(ethanol) containing a trace of glycerol. On this, as on other occasions traps were
4-3 filled with bait or preservative, After two days’ operation catches showed that
the baits ranked: syru > beer-syrup > beer > alcohol-glycerol in effectiveness
F
(Table 1). The level o efficiency obtained with syrup could be excessive for this
rate of trapping might seriously deplete a population if continued for a long period.
The alcohol-glycerol catches were attributed to ants exploring a new feature of
their environment and there was no indication of repellency since the few individuals
taken must have climbed into the traps.
In another experiment at Glen Osmond, baited and pitfall tra s were compared.
P
Twenty specimen-tube pitfalls containing a little alcohol-glycero were set out in a
line at 1 metre intervals, their mouths flush with the ground surface, and were flanked
by two arallel lines, each of 20 traps 1 metre away with their mouths projecting
K
above t e ground. In each outer line, beer-syrup alternated with alcohol-syrup;
these baits were selected as being less effective than syrup alone. In the trapping
period of a fortnight for which results are given in Table 2, traps were lifted once,
exchanging the two types of bait to counteract any position effects. As a few traps
were lost through trampling by sheep, the figures are corrected to 300 trapdays per
treatment. Both types of baited trap took more ants than the pitfalls, but the same
range and number of species were represented in all three treatments. The only
major difference was an apparent under-estimation of Camponotus species in the
pitfall traps but on the other hand these traps gave much larger catches of groups
other than ants. Some of these were probably simple pitfall captures while some were
attracted to the alcohol-glycerol. Here and in later tables attracted groups are
indicated on the basis of the internal evidence of each experiment modified where
necessary from results from many other traps. In these experiments baited traps
yielded little more information on ants than pitfalls, certainly not enough to justify
the extra trouble involved in using baits, especially as they tend to decompose, and
the additional variable (attraction) introduced to the results.
Preservatives in pitfall traps
In the Solomon Islands the attractiveness of a readily available preservative,
commercial methylated spirit, was examined in Jan. 1966 in forest on Mt. Austen.
Ten traps consisting of glass jars, mouth diameter 6 cm, with the lip level with the
 USE OF BAITS AND PRESERVATIVES IN PITFALL TRAPS 255

TABLE2
CATCHES IN PITFALL AND BAITED TRAPS, GLEN OSMOND, SOUTH AUSTRALIA
(EXCLUDING ADULT DIPTERA, MITES AND COLLEMBOLA)
~

Pitfalls Baits
Group -
Alcohol- Beer- Alcohol-
glycerol syrup syrup
Formicidae
Rhytidoponera spp. 3 25 8
Pheidole spp. 203 942 78
Chelaner sp. 323 2356 2521
Camponoius spp. 1 154 65
Other spp. 15 19 42
Total individuals (and spp.) 545( 12) 3496( 13) 27 14( 13)

Other groups
Coleoptera
Carabidae 1 - -
Staphylinidae, Aleocharinae' 4 3 -
,, , Omaliinae' - - 2
, oxytelus sp? 149 1 15
Hydrophilidaea 2 - -
Scarabaeidte, Onthophagus ~ p . ~ 24 - -
Anthicidae 11 - -
Ni tidulidae 3 - -
Others 154 I 1
Hymenoptera (other than Formicidae)' 1 - -
Diptera (larvae) 14 1 -
Heteroptera 16 - 4
Homoptera 25 4 5
Thysanopteraa 4 - 2
Araneida 28 2 4
Othersb 11 2 -

Total individuals, other groups 447 14 33

Groups attracted to alcohol-glycerol.
Lepidoptera (larvae), Isoptera, Opiliones and Chilopoda, all represented by less than 10 individuals.
TABLE3
CATCHES IN WATER AND METHYLATED SPIRIT TRAPS IN LOWLAND TROPICAL
RAIN-FOREST, GUADALCANAL, SOLOMON ISLANDS, DURING PERIOD 1 (see text)
Group Water Spirit
Coleoptera' 88 478
Formicidae 284 190
Dipteraa 14 58
Hemiptera' 4 41
Psocopteraa 1 -
Dermapteraa - I
Orthoptera' 6 25
Collembola 91 110
Araneida 9 19
Acan 10 13
Others 17 14
Totals 524 955
a Groups attracted to spirit.
Including Lepidoptera, Thysanura, Diplopoda, Chilopoda, Opiliones, Scorpionida, Thelyphonida,
Mollusca, earthworms and lizards, all represented by less than 10 individuals.

soil surface were placed in a line with alternate traps partly filled with spirit or water.
The traps were cleared at least twice every three days to reduce decomposition of the
catch in water. After a total of 45 trap days per treatment (period 1) the positions
of the two types of traps were exchanged and trapping was continued for a further
40 trap-days (period 2). Over both periods the total number of individuals trapped
was much greater in spirit (1582) than in water (673) although treatment differences
256 PENELOPE GREENSLADE and P. J. M. GREENSLADE

TABLE4
CATCHES OF COLEOPTERA IN SPIRIT TRAPS, SOLOMON ISLANDS, PERIOD 1
Group Number of Species Number of Individuals
Carabidae 1 1
Staphylinidae, Oxytelus spp. 2 101
, Aleocharinae 8 92
Other {taphylinidae 3 27
S taphylinoidea 5 21
Ni'iidulidae 5 68
Other Cucu'oidea 4 23
c u rculionidae 3 4
Scolytidae and Platypodidae 8 120
Other Coleoptera 6 21
Totals 45 478

CATCHES OF WIDE-MOUTHED JAR AND SPECIMEN-TUBE PITFALL TRAPS CONTAINING

ALCOHOL-GLYCEROL AS A PRESERVATIVE, GLEN OSMOND, SOUTH AUSTRALIA. A,
FIRST AND B. SECOND EXPERIMENT
Numbers of traps are shown in brackets.
A B
Group J~~~ Specimen J~~~ Specimen
Tubes Tubes
(3) (19) (4) (20)
Formicidae
Rhytidoponera spp. 1 - 2 -
Pheidole spp. 4 27 9 8
Chelaner sp. 13 23 7 20
Iridomyrmex spp. 21 9 22 22
Camponotus spp. 1 - 2
Other spp. 4 8 6 6
Total Formicidae 44 67 46 58
Other groups
Coleo tera
drabidae - 1 1 -
Staphylinidae, Aleocharinat a 24 6 9 8
, Oxytelus sp. 1 2 1 2
Nitiducdaea 3 - I I
Others 1 3 2
Hymenoptera (other than Formicidae) a 10 - 5 -
Dipteraa 586 84 Present, not recorded
Psocopteraa 1 - - -
Dermapteraa 21 2 13 3
Collembola
Poduromorph spp. Abundant, not recorded
Isotomidae 50 45
Entomobryidae 5 5 Present, not recorded
Smithuridae 63 68
Acari, Halotydeus and Penthaleus spp. 34 71
Acari, others 15 14 Present, not recorded
Othersb 2 6 5 3
:Attracted groups, remainder probable pitfall captures.
Hemiptera, Isoptera, Opiliones, Araneida, Chilopoda, Diplopoda, Mollusca, all represented by less
than 10 individuals.

varied from group to group (Table 3). In the Coleoptera, Oxytelus species (Staphyl-
inidae) and Scolytidae formed 90% of the catch in water and were associated with
decomposing, trapped Collembola. The only apparent pitfall captures in water
were a mere eight specimens which were not representative of the litter Coleoptera
known to be present on the site; this litter group included many paedarine,
 USE OF BAITS AND PRESERVATIVES IN PITFALL TRAPS 257

euasthetine and osoriine staphylinid species, Pselaphide, Scaphididae and others,

none of which were trapped. In contrast with this, large catches of Coleoptera were
made in spirit (Table 4) and most of the species had been collected in the area in logs
at an early stage of decay, at wounds on trees, in fungi or in fallen fruits. Many
other histerid, curculionid and scolytid species from these habitats were taken in
large numbers in other spirit and alcohol traps.
There were no treatment differences among the ants and although the most
abundantly trapped species, Zridomyrmex cordatus (263 individuals), was common in
two water traps, this proved to be a position effect. The next most numerous ant was
Rhytidoponera araneoides Le Gillou (49 individuals), a large nocturnal species, and
the remaining 162 ants belonged to 20 species. The traps were valuable in showing
that some of these, which, previously, were thought to be confined to soil and litter,
are in fact active on the surface, presumably at night. However, a very common
diurnal species, Paratrechina manni Donisthorpe, was poorly represented in the
catch and probably saw and avoided traps.
Among Collembola, as among the ants, there appeared to be no differences
between water and spirit catches. Although statistical significance might be claimed
for the increased catch in spirit compared with that in water (E = 91, X I = 4.13,
P < 0.05) the difference has no biological significance here. Altogether 20 Collem-
bola species were recognised including many Entomobryidae such as Paronella,
Lepiabcyrtus and Dicranocentrus species and large Sminthuridae which occur on
and near litter surface. Of the other groups trapped, Diptera and Hemiptera
(mainly an enicocephalid species) were consistently most numerous in spirit in this
and other series of traps in rain-forest in the Solomon Islands. In the Orthoptera,
higher catches were made in spirit than water for two of the three families present
on the site: Tetrigidae, water 3 individuals, spirit 2; Gryllidae and Gryllacrididae,
water 5, spirit 117. In other Solomon Islands traps, although not in these, Dermap-
tera were much more abundant in spirit and alcohol-glycerol traps than in water.
In further experiments with alcoholic preservatives in South Australia at Glen
Osmond, catches in wide-mouthed traps (mouth diameter 6 cm) were compared
with those in specimen tubes (diameter 1.8 cm). All the traps were partly filled with
alcohol-glycerol and sunk in the ground with the lip flush with the soil surface.
In Table 5A (first experiment) catches are divided between probable pitfall and
attracted captures. The species-composition of the catch was similar in both types
of trap, but the wide-mouthed jars took more individuals of attracted groups (all
of them winged insects) and the numbers were greater than could be accounted
for simply by differences in mouth diameter (Table 6). But the large-small trap ratio
of pitfall captures did correspond with that of trap diameters and this would
TABLE
6
ANALYSIS OF DATA IN TABLE 5 SECTION A, MEAN CATCHES PER TRAP, EXCLUDING
PODUROMORPH COLLEMBOLA
Catch Jars Specimen Tubes
All individuals 286.3 19.6
Attracted catch 215.3 4.9
Pitfall catch
Total 71 .O 14.6
Excluding Acari and all Collembola 15.3 4.0
Ants 14.7 3.5
Trap diameter (cm) 6 1.8
Trap area (cm’) 28.3 2.54

be expected since the catch depends on the amount of trap expased to an individual
moving horizontally. Possibly, entry to the larger jars is easier for flying insects, or
eddying wind currents give more rapid diffusion of the preservative from them. As
this experiment was operative only for a short period (two days) it was repeated
and results showed the same trends (Table 5, Section B).
In these Australian trials duromorph Collembola were often trapped in
g.
very large numbers and since t ey are difficult to sort accurately from large jars
258 PENELOPE GREENSLADE and P. J . M. GREENSLADE

0 1 2 3 4 5 6
TRAP DIAMETER (cm.)

FIG. 1.-Catches of Collembola in alcohol-glycerol pitfall traps differing in mouth diameter, Belair,
South Australia. Mean catch per trap It 2 SE. Solid line, expected catches if the total is distributed
according to the relative diameter of traps; broken line, expected catches if they were proportional to
the area of the mouths of traps.

they were not always counted. As in the Solomon Islands, the remaining Collembola
showed no apparent attraction to an alcoholic preservative (Table 5A), but there
were indications that some poduromorph species might be attracted. This was
investigated at Belair by comparing catches in traps of three diameters (Fig. 1).
Four traps of each size, all containing alcohol-glycerol, were set out in random
order in a 3 x 4 grid in an area 3 x 3 metres and were o erated for two days.
Catches, ranging from 12 to 149 Collembola per trap, were dpominated by poduro-
morph species of Hypogastrura. Their distribution in the different trap sizes showed
better agreement with expectation on the basis of captures resulting from attraction
than on the basis of a simple pitfall effect (Fig. 1) in that they appeared to vary as a
function of the area of the mouths of traps and were not proportional to their
diameters. Collembolan distributions are aggregated and, inevitably, the variances
of mean catches for each trap size are high. However, variances would continue to
rise even if catches were increased by extending the trapping period or using more
traps (Southwood 1966). Therefore in a final check at Belair, specimen tube traps
 USE OF BAITS AND PRESERVATIVES IN PITFALL TRAPS 259
were used to make a direct comparison of catches of Collembola in alcohol-
glycerol and water (Table 7). With totals of eight traps and 24 trap-days per treat-
ment, there was strong attraction of Hypogasrrura species to the alcohol-glycerol
traps and less difference between treatments for other groups of Collembola. Other
poduromorph species and forms with short appendages did not seem to be attracted
although species with longer appendages may have been; even so the numbers that
were trapped were low and attraction was certainly less than among Hypogastrura
species.
TABLE
I
CATCHES OF COLLEMBOLA IN WATER AND ALCOHOL-GLYCEROL PITFALLS, BELAIR,
SOUTH AUSTRALIA
~~ ~

Collembola Water Alcohol-glycerol

Hypogastrura speciesa 28 345
Others, long appendagesb 9 40
Others, shorter appendages' 10 5
~ -
Total 41 390
aAs determined from Womersley (1939), Hypogastrura manubrialis (Tullberg) and a few (c. 5 %)
bH. armata group.
Entomobryidae including Lepidocyrtus and Lepidoseira species ; Isotomidae, Isoromurus and Isotoma
spp. ; Sminthuridae, Parakafianna sp.
'Species of Tullbergia, Neanura and Folsomia and lsotomina sp., cf thermophila (Axelson) (one speci-
men), and three examples of an unidentified pseudachorutid.

DISCUSSION
Although pitfall traps have disadvantages they are not necessarily so great as
stated by Hayes (1970), and they have to be set against simplicity and ease of opera-
tion. It was a premise of the trials described here that only a marked improvement
over pitfall traps would justify the additional complications involved with baited
traps. We conclude that in diverse ant faunas any advantages OF baits are marginal
and pitfall traps are to be preferred. But baits may be worthwhile in studies of a
single, dominant species or in areas where the diversity of ants is low. When sampl-
ing ants, the use of alcoholic preservatives in pitfall traps does not appear to raise
problems of attraction. For many other groups this is not the case and there was some
correspondence between those attracted in the Solomon Islands and South Australia.
In both areas they included Diptera, Dermaptera and Coleoptera such as Oxyrelus
and Onthophagus species, Nitidulidae and omaliine Staphylinidae.
These trapping results show how the large ant populations found in both our
study areas may influence other groups inhabiting the ground surface; in turn this
affects the sampling methods needed in extensive surveys covering a number of
taxonomic groups. Carabidae, for instance, were notably infrequent in pitfall traps
in both areas in comparison with cool temperate climates where ants are less
numerous, and both Darlington (1943) and van der Drift (1963) have attributed the
scarcity of ground-living Carabidae in the tropics to the abundance of ants there.
It is also probable that ants limit the frequency of other large cursorial arthropods,
either by direct predation or interference or by preempting their niches. Large
species of invertebrates do occur in the ground-layer of lowland tropical rain-forest,
for example Orthoptera, Dermaptera, predacious Hemiptera and many Coleoptera,
but they are found mainly in temporary decomposition habitats which most ants
are poorly adapted to exploit on account of their dependence on permanent nest
sites. There is also a considerable ground fauna on the open floor'of lowland
tropical rain-forest apart from ants and micro-arthropods, but it consists to a large
extent of small species. These are capable of sheltering in refuges within the upper
soil and litter layer and, from the evidence of pitfall trap catches, few of them travel
extensively on the litter surface.
260 PENELOPE GREENSLADE and P. J. M. GREENSLADE

In the majority of the attracted species the response to preservatives will be for
feeding or oviposition and natural attractants are likely to be alcoholic fermentation
products of plant tissues containing rapidly decomposed polysaccharides. In the
Solomon Islands, the apparent attraction of Scolytidae and Oxytelus species to
animal decay in water traps reflects the wide range of materials to which sap- and
wood-feeding species and their predators respond. For example the palm weevil,
Rhyncophorus palmarum (L.) is recorded as being attracted to skatole (Hagley 1965),
although naturally this would be met only as a putrefaction product of animal
protein. The methylated spirit used in the Solomons contained pyridine which can
act as an attractant or repellent to insects (Dethier 1947) but its contribution to
catches, and that of glycerol in alcohol traps, are not known.
It might be expected that Hypogastmra species, especially H . manubrialis,
would be attracted to the preservatives we used, since this species forms dense
aggregations on carrion baits in the wetter months of year in South Australia.
Solomon Islands-Australian differencesin the trapping of Collembola are explained
by the absence of any Hypogastrura species in the former area where Hypo-
gastruridae as a whole are poorly represented (Greenslade 1969).
This account supports Luffs (1968) emphasis on the necessity for testing the
attractive (and possible repellent) effects of any preservative used in pitfall traps.
Large catches of flying insects such as Diptera always indicate attraction which is
also probable when species otherwise found at decaying fruits, sap and so on, are
taken in numbers. However, as Luff has shown for Carabidae, there may be attrac-
tion where it would not be expected. Testing for this may be difficult in cases where
catches in preservative cannot be compared with those in empty traps or water, for
exam le on account of predation in traps. But a possible method is provided by the
1
fact t at pitfall catches should be proportional to trap diameters, at least over the
range employed in these experiments, and attraction (and perhaps repulsion)
should be indicated by departures from a linear relationship. This also means that
in itfall trapping there will be no loss of efficiency if small traps are used instead
P
of arger ones (except for relatively large invertebrates) and the former are easier
to handle (Luff 1968) and allow optimum dispersion of sample points. But if baits
are used, larger traps should be more effective. In the Solomon Islands, wide-
a
mouthed itfall traps containing methylated spirit, with or without a carrion bait
suspende over them, were a very efficient collecting technique. Although they do
not provide quantitative data they do collect members of groups which move on
the litter surface and are not satisfactorily extracted in Tullgren funnels, for example,
Gryllidae and entomobryid and sminthurid Collembola with long appendages.
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