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Annals of the Entomological Society of America, XX(X), 2020, 1–21

doi: 10.1093/aesa/saaa044
Review Review

A Review of Terrestrial and Canopy Malaise Traps


Michael J. Skvarla,1,4, Jonathan L. Larson,2 J. Ray Fisher,3 and Ashley P. G. Dowling3

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1
Department of Entomology, Penn State University, University Park, PA, 2Department of Entomology, University of Kentucky, Lexington,
KY, 3Department of Entomology and Plant Pathology, Fayetteville, AR, and 4Corresponding author, e-mail: mxs1578@psu.edu

Subject Editor: Gadi V. P. Reddy

Received 10 September 2020; Editorial decision 12 October 2020

Abstract
Malaise traps are important tools for the large-scale collection of arthropod taxa. Here, an extensive review
of the history and literature concerning Malaise and canopy traps is given. This review highlights how trap
design and placement can affect trap catch, which will help researchers to make more informed choices when
planning research endeavors. Additionally, terrestrial and canopy-style Malaise traps are compared to each
other and other types of arthropod traps.

Key words:  Malaise trap, insect collecting, review, intercept trap

“Since the time of Linnaeus the techniques of catching insects ‘rarefaction curve’, and combinations of those terms. Once rele-
has not improved very much.” vant literature was acquired, additional references were found by
– René Malaise 1937. scouring the citations. Steyskal (1981) and Achterberg (2009) pro-
vided extensive bibliographies on Malaise trap research and were
Malaise traps—which are large, tent-like structures made of fine
invaluable resources. The number of days between specific dates was
mesh netting—are one of the most widely used nonattractant, static
calculated using ConvertUnits (2020).
insect traps (Muirhead-Thomson 1991). They were first devel-
Herein, we use ‘Malaise trap’ to refer specifically to terrestrial
oped by René Malaise after he observed insects trapped inside of
Towne’s style Malaise traps (e.g., those traps set near, or in contact
a tent and provided the first passive method to survey and collect
with, the ground or over streams) and ‘canopy trap’ to refer to those
large numbers of flying insects. Malaise traps take advantage of the
traps suspended at considerable height above the ground, generally
fact that many insect species climb up after encountering a barrier.
in the forest canopy. While Malaise and canopy traps are based on
The traps primarily sample flying insects, especially Diptera and
similar designs and Malaise traps set in different environments (e.g.,
Hymenoptera, which are intercepted by the mesh walls, but also col-
field, forest, and over streams) may sample diversity as different as
lect numerous ground-dwelling species as well (Zilihona et al. 1998,
that sampled by Malaise and canopy traps, we make the distinction
Achterberg 2009).
between Malaise and canopy traps herein as such a distinction is
While Malaise traps are used extensively to survey insects and
made in the published literature. Other styles of Malaise traps (e.g.,
various short reviews have been written about them, there is no
Gressit Malaise traps) are referred to by their full names.
comprehensive review that cites much of the underlying literature.
Herein, we provide a review of Malaise traps, including the history
of their development, commonly collected taxa, factors that influ- History
ence the arthropods collected by the traps, and various uses and René Malaise (1892–1978) was inspired to invent a new type of in-
questions that have been investigated using Malaise traps. We also sect trap after watching insects fly into a tent and become trapped
compare Malaise traps to other arthropod survey methods and com- despite the open flaps. His design consisted of mesh fabric stretched
ment on potential problems concerning the use of Malaise traps. We over a wooden box frame open at one end with a collection cylinder
hope that this review will serve as a resource for anyone who uses at the top (Fig. 1; Malaise 1937). It would revolutionize the collec-
Malaise traps to collect insects. tion of flying insects.
Vecht (1939) was the first person to utilize Malaise’s design when
he successfully tested it in Burma and translated the design into
Material and Methods Dutch. However, it is unclear if his contribution had much impact
The first author conducted an extensive literature search on Google on the history of the trap design.
Scholar for terms such as ‘Malaise trap’, ‘canopy trap’, ‘SLAM trap’, Henry Townes (1913–1990) elaborated upon Malaise’s design
arthropod order and family names (e.g., ‘Diptera’ and ‘Tachinidae’), and presented a modified version of the trap at the 1959 Annual

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Fig. 3. Townes’ light-weight Malaise trap. After Townes (1972), used with
permission.

Others continued to experiment with the design of Townes’


Malaise trap in the hopes of maximizing various aspects of it.
Schroeder et  al. (1975) proposed a more durable design that used
a metal frame instead of wood and a bronze screen funnel instead
of plastic for use on wind-swept rangelands. Masner and Goulet
(1981) noticed some Hymenoptera would not readily climb up the
Fig. 1.  Malaise’s original trap. After Malaise (1937), used with permission. mesh into the collector, so designed a Malaise-type trap impregnated
with fast-acting insecticide and a collecting trough underneath.
Hutcheson (1991) suggested a modified collection jar to facilitate
easier servicing while Achterberg (2009) suggested angling the en-
trance of the collection jar at 45° instead of horizontally, as is the
case in commercial designs, in order to increase trap catch.
Additional modifications have been proposed so that Malaise-
type traps are more efficient at collecting specific taxa, including
Odonata (Glotzhober and Riggs 1998) and migrating Lepidoptera
(Walker and Lenczewki 1989), or answering specific questions, such
as periodicity by automatically segregating trap catch at certain time
intervals using a rotary device (Murchie et al. 2001).
Some designers have worked from Malaise’s concept rather than
simply modifying Townes’ trap design. For example, Gressitt and
Gressitt (1962) introduced a trap that consists of a large (7 m long
by 3.6 m high) sheet of fabric with collectors at either end that is sup-
ported between two poles or trees (Fig. 4). Walker (1978) modified
the Gressitt design so that insects from either side are collected sep-
arately and flight direction can be determined. Other designs include
Fig. 2. Large Malaise trap utilizing a prefabricated aluminum frame. After
large square traps only open to one side (Walker and Lenczewki
Marston (1965), used with permission.
1989) and small triangular traps (Fig. 13 in Achterberg 2009). Traps
with four fabric panes open to 360°, which were first proposed by
Meeting of the Entomological Society of America. After ‘many re- Nishida and Torii (1970) and modified by Yano et  al. (1975), are
quests for plans of the trap’, he published schematics for his design, commercially available as BugDorm SLAM (Sea, Land, and Air
though was worried the publication might ‘freeze the model at this Malaise) traps (MegaView Science Co., Ltd., Taiwan) (Fig. 5). SLAM
stage of development rather than encourage further experimentation traps are often used as canopy traps as they are easily raised into the
with it’ (Townes 1962). His fear may be correct in part: the most air and can be fitted with optional bottom collectors.
widely available and commonly used Malaise traps are only slightly Other modifications or redesigns have aimed to produce inex-
modified from Townes’ design. pensive, homebuilt traps, including designs proposed by Butler
Marston (1965) proposed improvements to Townes’ design, (1965, 1966), which consist of a modified bed net with a polythene
including the use of a prefabricated tubular aluminum frame instead collecting bag placed in the hole in the roof; Walker and Lenczewki
of using a wooden frame (Fig.  2). Móczár (1967) proposed add- (1989), which utilizes mosquito netting and monofilament shrimp
itional modifications to Marston’s to lighten Townes’ 13.5 pound nets; and Platt et  al. (1999), which uses wooden legs in a tripod
trap design. Townes (1972), after possibly reading of this improve- design. Will and Steplowski (2016) even presented designs for a 3D
ment, also designed a light-weight Malaise trap (Fig. 3). printed Malaise trap head.
Annals of the Entomological Society of America, 2020, Vol. XX, No. XX 3

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Fig. 4. Gressit and Gressit-style Malaise trap. After Gressitt and Gressitt
(1962), used with permission courtesy of the Bishop Museum.

Various methods have been described for using Malaise traps to


sample canopy arthropods. Some researchers have attached Malaise
traps to tall scaffolding or platforms constructed in the canopy
(Coulson et al. 1971, Crossley et al. 1973, Southwood et al. 1979).
Others tied Townes-style Malaise traps off to a wooden frame and
used ropes and pulleys in order to raise the structure into the canopy
(Hammond 1990, Faulds and Crabtree 1995, Basset et  al. 1997).
However, the advent of SLAM traps has made using Malaise-type Fig. 5.  BugDorm SLAM (sea, land, air Malaise) trap with bottom collector.

traps in the canopy much easier and is now the design used almost
exclusively in such studies. collector depending on whether the trap is set in the understory or
Malaise traps have been combined with other traps, including canopy (Vance et  al. 2007). However, the bottom collector may
light traps (Dufour 1980), window traps (Basset 1988; Fig. 6), and interfere with the collection of certain taxa (e.g., Cerambycidae), so
intercept traps, including colored pan traps (Fig.  7), in order to it may not be appropriate to always include one (DiGirollomo and
modify taxa collected or increase efficiency. The addition of yellow Dodds 2017).
pan traps beneath Malaise traps increases the trap effectiveness in Wet or dry killing agents may be used in the collecting head.
catching Diptera, Hymenoptera, Heteroptera, and Thysanoptera Both have advantages depending on the taxa targeted. Wet killing
(Darling and Packer 1988, Campos et al. 2000). Henry et al. (2018) agents—such as 70–90% ethanol or propylene glycol—also function
presented an ‘alpine Malaise trap’, which consists of two interlocking as a preservative, which is needed if traps are serviced on a weekly
clear plastic panes rather like a flight intercept trap, that can be com- or longer basis. Delicate specimens, especially Lepidoptera, may be
bined with a drop trap or colored sticky cards for use in windy, high- damaged by wet killing agents and unidentifiable beyond higher
altitude environments. taxonomic levels (e.g., family or genus). Dry killing agents—such
as naphthalene and insecticide-permeated strips—help alleviate this
Trap Setup and Implementation but require traps be serviced more often, potentially daily, as insects
Malaise traps can be erected in any location where flying insects are may damage themselves or other specimens before succumbing to
expected, including forests, open grassy areas and prairies, and wind- the killing agent, especially if an excess of specimens builds up in the
swept ridges, although care must be taken to secure them when high trap head. If interested in ectoparasites or phoretics present on flying
winds are expected. Trap location affects the taxa collected (Ozanne insects, dry killing agents have shown to be effective at killing the
2005). Insects often follow specific flight paths through vegetation ectoparasites on-host, often preserving the host–symbiont relation-
and a trap located along a flight path will catch more specimens than ship. Wet killing agents often result in detachment of the symbiont
one that is not (Matthews and Matthews 1983, Hutcheson 1990, and mixing of symbionts among individuals in the collection jar.
Southwood and Henderson 2000). Traps set in sunny, exposed areas
collect more insects than those in sheltered, shaded areas (Noyes 1989, Taxa Collected
Irvine and Woods 2007). Topography, wind, water, light, and other Diptera and Hymenoptera are generally the numerically dominant
abiotic conditions should also be taken into consideration (Gressitt taxa in Malaise traps, with Diptera often representing the majority
and Gressitt 1962, Richards and Windsor 2007). Additionally, some of specimens collected (Table 1). Because of this, Malaise traps are
researchers have suggested setting traps in a north-south orientation often used to generally survey diversity of diptera diversity. However,
with the trap head facing the sun’s zenith (Noyes 1989). they have also been used to specifically collect and survey many spe-
While environmental factors have been little studied, Matthews cific Dipteran taxa, including Tabanidae, Syrphidae, and Tachinidae
and Matthews (1969) reported temperature and precipitation had a (Table  2). Hymenoptera are generally the second most-collected
strong influence on trap catch, with the largest catches happening on taxa, though usually represent a much smaller percentage of the
hot, sunny days following rain. total catch than flies. Similar to Diptera, Malaise traps are often
The addition of a bottom collector to canopy traps can be im- used to survey general hymenopteran diversity but have also been
portant, as some taxa are preferentially caught in the top or bottom used to target specific taxa, including Braconidae, Ichneumonidae,
4 Annals of the Entomological Society of America, 2020, Vol. XX, No. XX

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Fig. 6.  Diagram of a combined Malaise/window trap. After Basset (1988), used with permission.

and Vespidae (Table 3). Besides Diptera and Hymenoptera, Malaise sequences of one of a few genes to identify the taxa in a sample
traps have been used to collect a variety of Arachnida, Odonata, (Aagaard et  al. 2016, Geiger et  al. 2016, Morinière et  al. 2016,
Coleoptera, Lepidoptera, and other insects (Table 4). Canopy traps Aagaard et al. 2017, Cancian de Araujo et al. 2017, Schmidt et al.
have been used to specifically collect Psocoptera, Thysanoptera, 2017, Cancian de Araujo et al. 2018, Watts et al. 2019).
Coleoptera, Diptera, Lepidoptera, Neuroptera, and Hymenoptera Malaise traps have been shown to be some of the most consistent
(Table 5). traps in terms of the composition of higher taxa collected, giving
Malaise traps run with wet collecting heads are generally con- credence to the confidence hymenopterists and diperists have that
sidered to be a poor choice when targeting Lepidoptera, as many they will invariably collect those taxa (Kitching et al. 2001). While
specimens lose their scales in liquid preservative (Walker and Crosby such consistency has not been studied in canopy traps, there is no
1988, Brown 2005, Krogmann and Holstein 2010). However, a reason to believe they do not sample similar groups irrespective of
number of studies have used Malaise trap-caught Lepidoptera for site locality.
genitalia studies (Schmidt 2014a,b, 2016, 2017; van Nieukerken
et al. 2016) and Schmidt et al. (2019) proposed a workflow for cur-
ating wet Lepidoptera that can be used to prepare pinned moths Application in Entomology Field Studies
and butterflies. Additionally, DNA work can still be conducted on Malaise traps are excellent tools for surveying biodiversity (e.g.,
wet lepidopterans, including metabarcoding of Malaise trap sam- Collections Unit 2017, Karlsson et al. 2020), especially when used
ples, which uses high-throughput DNA sequencing targeting short in conjunction with traps that collect nonoverlapping assemblages
Annals of the Entomological Society of America, 2020, Vol. XX, No. XX 5

2008, Rohr et al. 2009, Banks et al. 2010, Sellars and Hicks 2015,
Carey et  al. 2017, Wolton et  al. 2017, Pérez-Urbina et  al. 2018,
Westwood et al. 2018). Additionally, Malaise and canopy traps have
been used to investigate community differences in monospecific and
highly diverse tree canopies in agroforestry (Sperber et  al. 2004),
how arthropod communities change during plant succession (Hollier
and Belshaw 1992, Hutcheson 1999, Shlyakhtenok and Agunovish
2001, Nol et  al. 2006, Missa et  al. 2009, Rohr et  al. 2009), crop
stage (Silva et al. 2020), stand growth (Hutcheson and Jones 1999,
Luqman et  al. 2018), invasion by foreign plant species (Toft et  al.

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2001), or along environmental (Harris et al. 2000; Vas et al. 2001;
Lynch et al. 2002; Kato et al. 2004; Kitching et al. 2004; Karem et al.
2006; Hirao et al. 2008a, b; Carr 2010), latitudinal (Kitching et al.
2004, Veijalainen et  al. 2013), elevational (Kumar et  al. 2009, Le
Cesne et al. 2015, Musthafa and Abdullah 2019, Plant et al. 2020),
and farming intensity gradients (Volpato et  al. 2019). Differences
between arthropod communities associated with different tree
species (Basset et  al. 1996); vertical stratification within a habitat
(Roberts 1976a, Rose 1978, Hollier and Belshaw 1993, Asquith and
Kido 1994, Sárospataki and Markó 1995, Hammond et  al. 1997,
Fig. 7.  Bugdorm Townes-style Malaise trap with colored pan traps.
Preisser et al. 1998, Charles and Basset 2005, Grimbacher and Stork
2007, Sobek et al. 2009, Ulyshen 2011, Birtele and Hardersen 2012,
of arthropods, such as pitfall traps (e.g., Peck 1989, Benton 1995, Somavilla et  al. 2019); and attractiveness of flowers (Rohrig et  al.
Winchester and Ring 1996, Handler 2007, Missa et  al. 2009, 2008) have also been studied.
Oxbrough et  al. 2010, Skvarla 2015, Skvarla and Dowling 2017). When operated for long periods of time (e.g., weeks to years),
They can also be used to sample a specific subset of biodiversity, such Malaise and canopy traps can be used to investigate meteorological
as natural enemies or specific pests (e.g., Nishida and Torii 1970, variables affecting flight activity (Matthews and Matthews 1969,
Steinbauer et al. 2000, Toft and Changler 2004). McCravy (2018) Burnett and Hays 1974, Nyrop and Simmons 1986, Isard et al. 1999,
reviewed sampling and monitoring methods in agroecosystems, Briers and Cariss 2003, Witter et  al. 2012), diel activity (Rickleps
including the use of Malaise traps. 1975, Hammond 1990, Basset and Springate 1992, Springate and
While such biodiversity surveys have traditionally relied upon visu- Basset 1996, Spencer et  al. 1998, Isard et  al. 2000, Murchie et  al.
ally sorting and identifying specimens, more recently, metabarcoding 2001, Shlyakhtenok and Agunovish 2001), and seasonal or pheno-
has been used to quantify arthropod diversity and abundance in logical cycles (Evans and Owen 1965, Rickleps 1975, Denlinger 1980,
bulk collection samples, including Malaise traps (Gibson et al. 2014, O’Neill and Evans 1983, Wright et al. 1984, Elliott 1986, Hammond
Morinière et  al. 2018, Barsoum et  al. 2019, deWaard et  al. 2019, 1990, Hollier and Belshaw 1993, Dutra and Marinoni 1994, Ellis
Watts et al. 2019). A number of studies have examined how various and Simon Thomas 1994, Jackman and Nelson 1995, Toft and Beggs
issues affect metabarcoding results of Malaise trap samples, including 1995, Flint 1996, Tereshkin 1996, Spencer et al. 1998, Kaspari et al.
how field conditions affect DNA degradation in Malaise trap samples 2001, Shlyakhtenok and Agunovish 2001, Noguera et  al. 2002,
(Krehenwinkel et al. 2018) and quantifying the difference between Hicks et al. 2003, Sperber et al. 2004, Winterbourn 2005, Maleque
using preservative ethanol or homogenate (Marquina et  al. 2019). et al. 2006, Ulber and Nitzsche 2006, Eveleigh et al. 2007, Noriega
One study even screened Malaise trap samples for vertebrate DNA, et al. 2007, Pinheiro et al. 2002, Whitemore et al. 2008, Choi et al.
which presumably came into the samples via hematophagous insects 2010, Yuen and Dudgeon 2016).
such as mosquitoes (Lynggaard et al. 2019). Malaise and canopy traps can also be used to investigate insect
Malaise and canopy traps have be used to investigate the movement, such as movement within and between habitat patches
arthropod community associated with specific habitats, such as (Naranjo 1991, Spencer et al. 1999, Hossain et al. 2002, Briers et al.
specific tree species (Basset and Arthington 1992), tree fall gaps 2004, Gangurde 2007, Williams et al. 2007a, Macfadyen and Muller
(Horn et  al. 2005, Ozanne 2005, Nol et  al. 2006, Ulyshen et  al. 2013), including into agricultural areas (Dutcher et al. 1986, Dyer
2006, Richard and Windsor 2007, Hirao et al. 2008a, b), and dead and Landis 1997, Spencer et al. 1998, Isard et al. 1999, Irwin et al.
wood (Hutcheson and Jones 1999, Ozanne 2005). They have also 2000, Nicholls et al. 2001, Ulber and Nitzsche 2006, Samaranayake
been used to investigate differences between patches of similar and Costamagna 2019), flight patterns in relation to wind direction
habitat (Hutcheson and Jones 1999, Choi et al. 2010, Fraser et al. (Pruess and Pruess 1966, Isard et al. 1999), and movement during
2007, Fraser et  al. 2008), habitat patches under different treat- mate-seeking (Abbott 2006). They are also used to investigate move-
ments (burning: Cancelado and Yonke 1970, Campbell et al. 2007, ment and dispersal (Cookset and Wright 1987), especially of adult
McCracy and Baxa 2011, McCravy 2017, harvesting: Deans et al. aquatic insects (Buskirk 1975, Mendl and Müller 1979, Müller 1982,
2005, Newell and King 2009; herbicide treatment: McCravy and Jones and Resh 1988, Sode and Wiberg-Larsen 1993, Williams and
Berisford 2011; insecticide treatment: Dilling 2007, Dilling et  al. Williams 1993, Griffith et al. 1998, Briers et al. 2004, Peterson et al.
2007, Santos et  al. 2007; organic vs. conventional crop manage- 2004, Winterbourn 2005, Solem and Bongard 1987, Winterbourn
ment: Bleicher et al. 2006, Silva et al. 2020) and between different et  al. 2007). However, caution should be used as mark–recapture
habitat types (Coulson et al. 1971, Crossley et al. 1973, Greiler and studies have shown that instantaneous direction, which is indicated
Tcharntke 1993, Bomar 2001, Hicks et  al. 2003, Warriner et  al. by the side of the trap insects are collected on, may not always be
2004, Gittings et al. 2006, Tangmitcharoen et al. 2006, Cunningham a reliable way to determine overall direction of movement between
and Murray 2007, Rohr et al. 2007, Vance et al. 2007, Smith et al. habitats or along gradients (Macneale et al. 2004).
6 Annals of the Entomological Society of America, 2020, Vol. XX, No. XX

Table 1.  Examples of Malaise trapping studies and taxa collected

Diptera Hymenoptera Lepidoptera Coleoptera Hemiptera Plecoptera Collembola ‘Other’


Citation (%) (%) (%) (%) (%) (%) (%) (%)

Marston 1965 66.3 12.3 4.8 5.6 6.8 - 3.4 4.2


Geijskes 1968 58 19 14 4.6 2.3 - - 2.1
Matthews and Matthews 1969 52.2 16.9 7.1 3.1 9.4 8.3 1.7 11.3
Matthews and Matthews 1969 43.1 22.5 10 2 3.9 16.8 1.2 18.5
Matthews and Matthews 1969 14.7 1.2 0.8 0.3 0 76.9 0.7 83
Matthews and Matthews 1969 54.4 13 15.5 2.6 10.9 0.3 0.6 3.6

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Yano et al. 1975 69.7 4.6 4.6 0.8 13.6 0 3.2 6.7
Rose 1978 84.2 5.85 3.52 0.11 3.74 - - 2.58
Cooksey and Barton 1981 57 15 17 - 8 - - 3
Moeed and Meads 1987 84.2 4 1.8 2.3 - - 4.9 2.8
Basset 1988 47.8 8.1 3.7 10.3 19.5 - - 10.7
Basset and Arthington 1992 - - - - - - - -
Dutra and Marinoni 1994 85.1 3.2 5.7 1.7 2.5 <0.01 0.9 2
Dutra and Marinoni 1994 72 3.3 13.5 2.6 1.4 0 5.3 7
Campos et al. 2000 84.4 7.8 3.2 0.8 2.9 - - 0.9
Campos et al. 2000 64.3 10 1.3 7 10 - - 7.4
Campos et al. 2000 57.4 16.5 0.9 5.3 12.2 - - 7.7
Hughes et al. 2000 69.2 14.3 - - - - - 16.5
Kwok and Corlett 2002 80.6 3.9 5.3 6.3 - - - 3.9
Brown 2005 84 - - - - - - -
Brown 2005 81 - - - - - - -
Brown 2005 64 - - - - - - -
Horn et al. 2005 68.5 4.1 9.5 7.6 - - - 10.2
Horn et al. 2005 26.6 7.3 30.8 18.1 - - - 17.2
Karlsson et al. 2020 75 15 1.0 2.0 2.0 - 2.0 3.0

Citation Number of Duration per Number of Specimens/ Locality Trap type


specimens collected trap (days) traps trap/day

Marston 1965 2,927 7 1 418.14 Kansas, USA Malaise


Geijskes 1968 90,182 - - - Suriname Malaise
Matthews and Matthews 1969 23,722 91 1 260.68 New York, USA Malaise
Matthews and Matthews 1969 6,138 91 1 67.45 New York, USA Malaise
Matthews and Matthews 1969 7,008 91 1 77.01 New York, USA Malaise
Matthews and Matthews 1969 3,480 91 1 38.24 New York, USA Malaise
Yano et al. 1975 13,709 3.34 19 216.03 Thailand/China Malaise
Rose 1978 37,198 127 6 48.82 Malaysia Malaise/
canopy
Cooksey and Barton 1981 10,830 12 1 902.50 Arkansas, USA Malaise
Moeed and Meads 1987 45,965 365 1 125.93 New Zealand Malaise
Basset 1988 14,597 365 5 8.00 Queensland, Australia Composite
Basset and Arthington 1992 46,019 730 5 12.61 Queensland, Australia Composite
Dutra and Marinoni 1994 62,924 365 1 172.39 Parana, Brazil Malaise
Dutra and Marinoni 1994 38,868 365 1 106.49 Parana, Brazil Malaise
Campos et al. 2000 6,120 14 4 109.29 Minas Gerais, Brazil Malaise
Campos et al. 2000 2,436 14 4 43.50 Minas Gerais, Brazil Malaise
Campos et al. 2000 4,816 14 4 86.00 Minas Gerais, Brazil Malaise
Hughes et al. 2000 12,776 60 3 70.98 Colorado, USA Malaise
Kwok and Corlett 2002 53,897 940 4 14.33 Hong Kong, China Malaise
Brown 2005 4,646 5 1 929.20 Tambopata, Peru Malaise
Brown 2005 905 4 1 226.25 Tambopata, Peru Malaise
Brown 2005 1,064 3 1 354.67 Puntarenas, Costa Rica Malaise
Horn et al. 2005 - - 5 - South Carolina, USA Canopy
Horn et al. 2005 - - 5 - South Carolina, USA Canopy
Karlsson et al. 2020 20,000,000 54,268 73 368.5 Sweden Malaise

Malaise and canopy traps can be used to estimate the abundance Buskirk 1975, Kato et al. 2003, Kato et al. 2004; Odonata: Kirkton
of target pest species (Beggs et al. 1998), establish damage thresh- and Schultz 2001; Anura: Horn et  al. 2005; Chiroptera: Jong and
olds (Beggs and Rees 1999), or create an index of abundance, avail- Ahlen 1991, Fukui et al. 2006; Aves: Poulin et al. 1992, Rodenhouse
ability, and biomass of captured arthropods by themselves or as a and Holmes 1992, Duguay et al. 1997, Duguay et al. 2000, Johnson
prey base for insectivorous predators (Lynch et al. 2002; Araneae: and Sherry 2001, Kwok and Corlett 2002, Murakami and Nakano
Annals of the Entomological Society of America, 2020, Vol. XX, No. XX 7

Table 2.  Diptera families collected in Malaise traps Table 3.  Hymenoptera taxa collected in Malaise traps

Taxon Reference Taxon Reference

General Diptera Kitching et al. 2004, Roháček and Ševčik 2009, General Hymenoptera Darling and Packer 1988, Noyes 1989, Campos
Wolton et al. 2017 et al. 2000, Shlyakhtenok 2000, Sobek et al.
Agromyzidae Scheirs et al. 1997; Papp 2009, 2018 2009
Axiniidae Colless 1994 ‘Symphyta’ Holuša 2002, Brand et al. 2003, Harris 2006
Asilidae McCracy and Baxa 2011, McCravy 2017, Aculeata Volpato et al. 2019
Martín-Park et al. 2018 Apocrita Karem et al. 2006
Brachystomatidae Plant et al. 2020 Apoidea/Anthophila Bartholomew and Prowell 2005, Ngo et al.

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Calliphoridae Rosati and VanLaerhoven 2007 2013, Geroff et al. 2014, Sellars and Hicks
Cecidomyiidae Karlsson et al. 2020 2015, McCravy et al. 2016
Chironomidae Diserud et al. 2013, Karlsson et al. 2020 Aphelinidae Kissayi et al. 2017
Culicidae Graham 1969, Witter et al. 2012 Braconidae Shimbori and Shaw 2014, Shweta and
Dolochopodidae Pollet et al. 1989, Plant et al. 2020 Rajmohana 2018, Karlsson et al. 2020
Empididae Plant et al. 2020 Chalcidoidea Karlsson et al. 2020
Hybotidae Plant et al. 2020 Chrysididae Shlyakhtenok 2000, Strumia 2003
Mycetophilidae Økland 1994, Toft et al. 2001, Toft and Changler Crabronidae O’Neill and Evans 1983, Ghazi-Soltani et al.
2004, Jakovlev and Penttinen 2007, Papp 2018, 2010, Arbaei et al. 2015
Karlsson et al. 2020 Diapriidae Masner 1976a,b
‘Nematocera’ Salmela et al. 2007 Diaprioidea Karlsson et al. 2020
Odiniidae Parchami-Araghi et al. 2018 Embolemidae Amarante et al. 1999
Oestridae Capelle 1970, Cogley and Cogley 2000, Fleenor Encyrtidae Manuel Rodríguez et al. 2010, Silva et al. 2020
and Taber 2007, Witter et al. 2012 Eulophidae Silva et al. 2020
Phoridae Karlsson et al. 2020 Evaniidae Deans and Kawada 2008, Shweta and
Pipinculidae Skevington 2001 Rajmohana 2018
Psychodidae Quate 1999; Alexander 2000, 2001 Formicidae Collingwood 1981, Deyrup and Trager 1986,
Sarcophagidae Dahlem and Downes 1996, Whitemore et al. 2008 Longino and Colwell 1997, Delabie and Reis
Sciaridae Steffan 1972, Vilkamaa et al. 2007, Karlsson et al. 2000, Kaspari et al. 2001, Collingwood and
2020 van Harten 2005, Colby and Prowell 2006,
Sciomyzidae Carey et al. 2017 Framennau and Thomas 2008, Fisher et al.
Simuliidae Adler et al. 1983, Currie and Adler 2000, Witter 2009, Guerrero et al. 2010, Karlsson et al.
et al. 2012 2020
Stratiomyiidae Hauser 2008, Whitemore et al. 2008, Birtele and Ichneumonidae Noyes 1989, Bartlett et al. 1999, Sääksjärvi
Hardersen 2012 et al. 2004, Sperber et al. 2004, Sääksjärvi
Syrphidae Burgio and Sommaggio 2002, Thompson and et al. 2006, Ulber and Nitzsche 2006,
Zumbado 2002, Krčmar et al. 2005, Gittings Veijalainen et al. 2013, Gómez et al. 2018,
et al. 2006, Nol et al. 2006, Smith et al. 2008, Pérez-Urbina et al. 2018, Karlsson et al. 2020
Whitemore et al. 2008, Birtele and Hardersen Mutillidae Pitts et al. 2004, Pilgrim and Pitts 2006, Vieira
2012, Carey et al. 2017, Martín-Park et al. et al. 2017
2018 Mymaridae Vance et al. 2007, Shweta and Rajmohana
Tabanidae Roberts 1971, Anderson and Hoy 1972, Blume 2018, Silva et al. 2020
et al. 1972, Roberts 1976b, Hollander and Platygastridae Stevens and Austin 2007; Burks et al. 2013;
Wright 1980, Strickler and Walker 1993, Shweta and Rajmohana 2016, 2018; Manoj
Leprince et al. 1994, van Hennekeler et al. et al. 2017
2008, Martín-Park et al. 2018 Platygastroidea Karlsson et al. 2020
Tachinidae Cerretti et al. 2004, Inclan and Stireman III 2009, Plumariidae Penteado-Dias and Scatolini 2003, Silva et al.
Stireman et al. 2012 2020
Tephritidae Asquith and Kido 1994 Pompilidae Shlyakhtenok 2000
Tipulidae Dufour 1980, Toft and Beggs 1995, Peterson et al. Tenthredinidae Karlsson et al. 2020
2004 Trichogrammatidae Silva et al. 2020
Vespidae Archer 1990, Beggs et al. 1998, Beggs and Rees
1999, Shlyakhtenok 2000, Sackmann et al.
2001, Kumar et al. 2009, Aragão and Andena
2002, Iwata et al. 2003, Yuan and Dudgeon 2016, Law et al. 2018,
2016, Somavilla et al. 2019
Partidge et al. 2020). Collected taxa can also be associated with spe-
cific habitats and used as habitat indicators (Fraser et al. 2007).
Malaise traps can also be used to collect specific guilds of insects, Within a specific trap-style, the color and material of the trap net-
such as those attracted to corpses and potentially useful in forensic ting can affect the taxa collected. For example, coarse mesh is more
studies (De Jong 1994) or medically important species (Roberts effective in collecting Aculeata, fine mesh is more effective in collecting
1971, 1972; Alexander 2000). microhymenoptera, and both coarse and fine mesh are effective in col-
lecting Ichneumonoidea (Darling and Packer 1988). Black mesh in-
Factors Influencing Catch creases the overall number of specimens and species of Tabanidae and
Trap design significantly affects the taxa collected. Few studies have Culicidae that are collected (Roberts 1970, 1972; Hansen 1988) and
compared trap catch of different Malaise trap designs, but Matthews the addition of various colored fabric panels to commercially available
and Matthews (1983) found that Townes-style traps caught 10 times Malaise traps with black netting can influence and increase the number
as many specimens as Cornell-style Malaise traps. and diversity of pollinators collected (Campbell and Hanula 2007).
8 Annals of the Entomological Society of America, 2020, Vol. XX, No. XX

Table 4.  Arthropod taxa, excluding Diptera and Hymenoptera, col- Table 5.  Insect taxa collected in canopy traps
lected in Malaise traps.
Taxon Reference
Taxon Reference
Psocoptera Santos et al. 2007, Sokolova
Acari Zheng 1996; Behan-Pelletier and Winchester et al. 2010
1998; Clark 2004; Zheng 2002a, b; Ripka Thysanoptera Santos et al. 2007
and Szabó 2010; Skvarla et al. 2014 Coleoptera Tangmitcharoen et al. 2006,
Araeneae Wilkinson et al. 1980, Hauge and Midtgaard Hardersen et al. 2014
1986, Jennings and Hilburn 1988,  Cerambycidae Vance et al. 2003, Dodds et al. 2010,
Oxbrough et al. 2010, Vedel et al. 2011 DiGirollomo and Dodds 2017

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Coleoptera Hosking 1979, Hutcheson 1999, Hutcheson  Buprestidae
and Kimberley 1999, Harris et al. 2000,  Coccinellidae Santos et al. 2007
Toft et al. 2001, Kriska and Young 2002,  Curculionidae, Dodds et al. 2010, DiGirollomo
Grimbacher and Stork 2007, Stork et al. Scolytinae and Dodds 2017
2008, Ohsawa 2010, Musthafa and Diptera Tangmitcharoen et al. 2006
Abdullah 2019  Stratiomyidae Whitemore et al. 2008
 Buprestidae Bellamy 2000  Syrphidae Whitemore et al. 2008
 Carabidae Liebherr and Mahar 1979, Ulyshen et al. 2005,  Tachinidae Cerretti et al. 2004
Ulyshen et al. 2006, Cassola 2009, Meng  Sarcophagidae Whitemore et al. 2008
et al. 2012, Skvarla and Dowling 2017  Tephritidae Asquith and Kido 1994
 Cerambycidae Noguera et al. 2002; Warriner et al. 2002, Lepidoptera Tangmitcharoen et al. 2006
2004; Vance et al. 2003; Schiefer and Newell  Tortricidae Eveleigh et al. 2007
2010; Skvarla and Dowling 2017; Barimani  Zygaenidae Hoddle 2006
Varandi et al. 2018 Neuroptera Hollier and Belshaw 1993
 Elateridae Steiner 2000, Nol et al. 2006 Hymenoptera Tangmitcharoen et al. 2006
 Eucnemidae Hoffman et al. 2009
 Lampyridae Barrows et al. 2008
 Mordellidae Jackman and Nelson 1995 Disney et al. (1982) found that slightly altering the position of the
Collembola Fjellberg 1992 collecting container from the peak of a Malaise trap to just below
Ephemeroptera Peterson et al. 2004 the peak significantly reduced the overall number of certain Diptera
Hemiptera Cancelado and Yonke 1970, Hodkinson and species collected and somewhat reduced the number of species col-
Casson 1991, Le Cesne et al. 2015 lected. The age of individual Malaise traps has also been shown to
 Fulgoroidea Gonzon et al. 2006, Benton and McCreadie significantly alter trap catch, possibly the result of changes in color
2009
due to exposure to sunlight (Roberts 1975, Duarte et al. 2010).
Lepidoptera Owen 1969, Butler et al. 1999, Harris et al.
Species-specific factors of target taxa such as behavior, habitat
2004, Campbell 2007
 Gracillariidae Steinbauer et al. 2000 preference, and activity level influence trap catch. For instance,
 Sesiidae Steinbauer et al. 2000 while many species of Syrphidae are readily collected in Malaise
 Tortricidae Eveleigh et al. 2007 traps, some abundant species of this family avoid the trap al-
Mecoptera Byers 1973 together (Burgio and Sommaggio 2007). Malaise trap collections
Neuroptera Hollier and Belshaw 1992, 1993; Vas et al. of mosquitoes are biased toward Aedes Meigen, 1818 and Culex
2001; Abraham et al. 2003 Linnaeus, 1758 (Diptera: Culicidae) (Acuff 1976) while collections
Odonata Muzón and Spinelli 1995, Roble 1995, Flint of Agromyzidae are female-biased (Scheirs et al. 1997). When used
1996, Glotzhober and Riggs 1998 to collect spiders, Malaise traps sample a greater proportion of ar-
Opiliones Hicks et al. 2003
boreal, web-building species compared to terrestrial, active-hunting
Orthoptera Samways and Moore 1991, Quinn et al. 1993,
species (Jennings and Hilburn 1988, Oxbrough 2010).
Donnelly 1995, Johnson et al. 1995, Muzon
and Spinelli 1995, Bomar 2001
Plecoptera Peterson et al. 2004, Winterbourn 2005 Use of Baits With Malaise Traps
Pseudoscorpiones Aguiar and Buhrnheim 1998 The addition of various lures can increase the catch of specific taxa.
Thysanoptera Olsen and Midtgaard 1996 For example, the addition of carbon dioxide in the form of dry ice
Trichoptera Jones and Resh 1988, Sode and Wiberg-Larsen or compressed gas released over time increases the trap catch of
1993, Peterson et al. 2004, Winterbourn
hematophagous Diptera and mammalian ectoparasites (Easton et al.
2007, Winterbourn et al. 2007
1968, Smith et al. 1965, Geijskes 1968, Witter et al. 2012; Tabanidae:
Opiliones Hicks et al. 2003
Orthoptera Samways and Moore 1991, Quinn et al. 1993, Roberts 1971, Anderson and Hoy 1972, Blume et al. 1972, Roberts
Donnelly 1995, Johnson et al. 1995, Muzon 1976b, Hollander and Wright 1980, Strickler and Walker 1993,
and Spinelli 1995, Bomar 2001 Leprince et al. 1994, van Hennekeler et al. 2008, Culicidae: Breeland
Plecoptera Peterson et al. 2004, Winterbourn 2005 and Pickard 1965, Graham 1969; Oestridae: Capelle 1970, Wright
Pseudoscorpiones Aguiar and Buhrnheim 1998 et  al. 1984, Cogley and Cogley 2000, Fleenor and Taber 2007,
Thysanoptera Olsen and Midtgaard 1996 Witter et  al. 2012). Most studies have found octenol/1-octen-3-ol
Trichoptera Jones and Resh 1988, Sode and Wiberg-Larsen (French and Kline 1989, Schreck et  al. 1993, Krčmar et  al. 2005,
1993, Peterson et al. 2004, Winterbourn van Hennekeler et al. 2008, Krčmar et al. 2010), ammonia (Hribar
2007, Winterbourn et al. 2007
et al. 1992, Krčmar et al. 2010), acetone (Krčmar et al. 2010), lactic
Annals of the Entomological Society of America, 2020, Vol. XX, No. XX 9

acid (Krčmar et  al. 2010), and aged animal urine (Krčmar et  al. 2012, Barimani Varandi et  al. 2018). However, other studies that
2005, Krčmar et al. 2006, Krčmar et al. 2010, Tunnakundacha et al. used finer taxonomic units found that Malaise traps collect more
2017), as well as the addition of objects that reflect horizontally po- specimens of certain beetle families (e.g., Cleridae, Curculionidae,
larized light, such as large, round, black object like an inflated beach and Elateridae) or certain tribes and genera within a family (e.g.,
ball covered in black cloth (Catts 1970, Schreck et al. 1993, Blahó lebiine carabids) than light, window, and sticky, colored pan, or pit-
2009) increase the number of Tabanidae caught in Malaise and fall traps (Hosking 1979, Skvarla and Dowling 2017).
canopy traps, though some have not reported differences (Leprince Malaise and canopy traps collect fewer Stomoxys Geoffroy,
et  al. 1994). 1-octen-3-ol is also attractive to Culicidae (Nilssen 1762 (Diptera: Muscidae) and Tabanidae compared to Vavoua and
1998). 2,4-hexadlenyl butyrate and heptyl butyrate are highly po- Nzi traps, respectively (van Hennekeler et  al. 2008, Krčmar 2017,
tent, specific lures attractive to yellowjackets (Vespula Thomson, Tunnakundacha et al. 2017).

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1869 [Hymenoptera: Vespidae]) and Malaise traps baited with these Comparison of Malaise traps and colored pan traps, including
chemicals can be used to control yellowjacket populations over white, yellow, and blue pans, are mixed and depend on the taxon
small areas such as fruit orchards (Davis et  al. 1973). Methyl eu- being studied. For Hymenoptera, studies have found that pan
genol is attractive to some species of Hawaiian Drosophilidae and traps collect more diversity than Malaise traps (Campbell and
Muscidae (Asquith and Kido 1994) and rotting fruit has been used Hanula 2007, Alfred Daniel et al. 2018), pan traps collect more
to attract Odiniidae (Parchami-Araghi et al. 2018) specimens but are dominated by a few species while Malaise traps
Bouwer et  al. (2020) found that different traps (standard collect fewer specimens representing more species with a more
multiple-funnel, modified multiple-funnel, panel, and canopy even distribution of species (Wells and Decker 2006), or that pan
Malaise traps) had significantly different trapping efficacy for two and Malaise traps collect significantly different assemblages with
cerambycid species during field trials and that the traps significantly very little (12%) overlap (Finnamore et al. 2002). Depending
affected the structure of CO2 odor plumes downwind of the traps; on the locality, Malaise and canopy traps collect either a similar
they speculated that the trap effect on odor plumes may contribute species diversity or higher species diversity of pollinating bee-
to the differences in trapping efficacy, but were unable to conclu- tles compared to colored pan traps (Campbell and Hanula 2007,
sively reconcile the differences. Skvarla and Dowling 2017) and are more or less as efficient as
white pan traps depending on the family of Diptera considered
Terrestrial/Canopy Trap Comparison (Disney et al. 1982).
Within temperate forests, some studies have found that Malaise traps, When focused on lower taxa, different studies have found
when compared with canopy traps, catch more insect specimens and that Malaise and pan traps collect similar assemblages of bees
sample a higher diversity at both the family (Preisser et  al. 1998, (Bartholomew and Prowell 2005) or Malaise traps collect more bee
Rohr et al. 2007, Barkley 2009) and species level (Syrphidae: Birtele specimens and species (Geroff et al. 2014); the addition of colored
and Hardersen 2012; Tachinidae: Cerretti et al. 2004, Stireman et al. fabric to Malaise traps increases the number of pollinators collected
2012). However, other studies have found that Malaise and canopy (Campbell and Hanula 2007). Malaise traps compared to yellow
traps have similar observed species richness, although they capture pan traps collect more species and specimens of Platygastridae
significantly different species assemblages (Cerambycidae: Vance and Mutillidae (Shweta and Rajmohana 2016, Vieira et  al. 2017)
et al. 2003, Hardersen et al. 2014 Neuroptera: Hollier and Belshaw but fewer Diapriidae (Shweta and Rajmohana 2018). Within
1993), while others have found no difference in species composition Ichneumonidae, Malaise traps collect the same species assemblage
(Stork and Grimbacher 2006). Additionally, while observed species and more specimens of Pimplinae and Rhyssinae (Gómez et al. 2018)
richness is equivalent or higher in Malaise traps, expected species compared to Möricke/yellow pan traps while the opposite is true for
richness, which is based on various species richness estimators, may Orthocentrinae and Cryptini (Mazón and Bordera 2008, Aguiar and
be higher in canopy traps (Vance et al. 2003, Stireman et al. 2012). Santos 2010).
Finally, canopy and Malaise traps collect similar feeding-guild as- Within a species, sexes may be preferentially collected by either
semblages, at least when considering Coleoptera (Grimbacher and method: Malaise traps catch more male Ichneumonidae and female
Stork 2007). Agromyzidae while Möricke/yellow pan traps collect more female
When both styles of trap collect the same taxon, relative abun- Ichneumonidae and male Agromyzidae (Scheirs et al. 1997, Aguiar
dance per trap may vary significantly depending on the taxonomic and Santos 2010).
level analyzed (family: Barkley 2009; genus: Roberts 1976b; species: In tropical environments, Malaise traps and canopy fogging col-
Eveleigh et al. 2007). lect similar assemblages of Formicidae, which is significantly dif-
ferent than the assemblage sampled by processing leaf litter; thus,
Comparisons to Other Traps and Collecting Methods a combination of Malaise traps and leaf litter sampling is more
Aside from Maliase traps, there are a number of other trap designs efficient than either method alone (Longino and Colwell 1997).
entomologists can employ to survey capture insects, including pit- Furth et  al. (2003) found broad overlap between the flea beetle
fall traps (containers in the soil that capture arthropods as they (Chrysomelidae: Alticinae) taxa collected by Malaise traps and
wander across), sticky traps (a surface coated in glue or tanglefoot), canopy fogging in Costa Rica; they also found fogging to be more
glass-barrier traps (a glass container with an inverted funnel in the efficient than Malaise traps on a per sample basis but Malaise traps
bottom to allow entry but no escape), pan traps (open pans or bowls were more efficient on a per individual basis, so Malaise traps are
that may be different colors and contain soapy water or another more efficient over long time spans.
collecting agent), and several others. Some comparisons between Suction traps are more efficient than Malaise traps at col-
various trap types have been made, but it is not always clear which lecting Culicidae (Lothrop et  al. 2002), Neuropteroidea excluding
trap is preferable. Raphidioptera (Abraham et  al. 2003), and microhymenoptera
Malaise traps collect more specimens of Diptera, Hymenoptera, (Favret et  al. 2019). The comparison is more nuanced when com-
and Hemiptera, but fewer specimens of Coleoptera compared to paring brachyceran flies as ‘Malaise traps outperformed suction
glass-barrier, window, and sticky traps (Juillet 1963, Lamarre et al. traps in terms of measuring α diversity, but suction traps resolved a
10

Table 6.  Target taxa and trapping statistics from publications that reported accumulation or rarefaction curves for Malaise trap collections

Number of Fold increase


days needed to
Number of traps were Malaise Number of Spe- Total number of exhaustively survey
Taxon Area surveyed (ha) traps run trap days cies collected species estimated target taxon Reference

Anthophila 12 5 12 30 68 90 7 McCravy et al. 2016


Anthophila - 6 13 78 10 - - Sellars and Hicks 2015
Asilidae 680 12 161 1,932 26 30.5 3.9 McCravy 2017
Carabidae 4 5 266 1,330 15 64.5 10.5 Skvarla and Dowling 2017
Cerambycidae (LA) - 10 118 1,180 37 39–48 - Dodds et al. 2015
Cerambycidae (MA) - 10 119 1,190 39 40–56 - Dodds et al. 2015
Cerambycidae (NY) - 10 98 980 28 30–48 - Dodds et al. 2015
Cerambycidae 4 5 266 1,330 53 92 5.3 Skvarla and Dowling 2017
Coleoptera - 2 4 8 - - - Musthafa and Abdullah 2019
Curculionoidea 4 5 266 1,330 39 70 6.4 Skvarla and Dowling 2017
Diptera (trap 1; 49 families in- 4 1 365 365 590 1,071 - Borkent and Brown 2015, Brown
cluded in analyses) et al. 2018
Diptera (trap 2; 49 families in- 4 1 365 365 116 270 - Borkent and Brown 2015, Brown
cluded in analyses) et al. 2018
Encyrtidae - 6 60 360 52 55–73.8 - Manuel Rodríguez et al. 2010
Formicidae 1,500 16 395 6,320 - - - Longino and Colwell 1997
Fulgoroidea 211,415 22 1,249 27,478 55 57–81 Gonzon et al. 2006
Ichneumonidae 40,000 10 365 3,650 - - - Mazón and Bordera 2008
Ichneumonidae: Diplazontinae, 9.3 30 30 900 55 - - Fraser et al. 2008
Pimplinae, Diacritinae, and
Poemeniinae
Ichneumonidae: Pimplinae and 5600 27 105–363 5,550 88 101–114 - Sääksjärvi et al. 2004
Rhyssinae
Insecta 45,000,000 73 Various 54,268 4,000a 33,000 - Karlsson et al. 2020
Trichoptera 5 km stretch of 36 9 324 50 136–155 - Andersen and Kjaerandsen 2001
river
Vespidae: Polistinae 3069 3 365 1,095 20 - - Aragão and Andena 2016

A Malaise trap day is one Malaise trap run for one day, such that 10 Malaise trap days can be accomplished by one trap run for 10 d, two traps run for five days each, etc. Publications that reported traps were run for one
year that did not provide specific dates (e.g., June 2018–May 2019) were assumed to be 365 d.
a
After only 1% of trap catch was sorted and identified to species.
Annals of the Entomological Society of America, 2020, Vol. XX, No. XX

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Annals of the Entomological Society of America, 2020, Vol. XX, No. XX 11

higher degree of brachyceran community heterogeneity using β di- Finally, based on the handful of studies that have reported col-
versity’ (Favret et al. 2019). lection or rarefaction curves for Malaise trap collections, it is clear
Malaise traps are able to function in damp conditions so may be that it is extremely difficult to impossible to exhaustively sample
preferable to vacuum-sampling understory vegetation that is con- Malaise trap-collected arthropods in even a relatively small area
stantly wet (Noyes 1989). (Table 6). Those studies that calculated how much additional ef-
Malaise traps and sweep netting can differentially sample re- fort would be required to exhaustively sample their target taxa
lated genera; for example, within Tabanidae the majority of Tabanus reported a 3.9- to 10.5-fold increase in effort. How much effort
and Hybomitra are collected in Malaise traps while the majority of is required is determined by how large an area is being surveyed,
Chrysops are collected by netting (Tallamy et al. 1976, Strickler and how diverse the target taxa are, and how efficiently Malaise traps
Walker 1993). When targeting Ichneumonoidea, Malaise traps are collect them.

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more efficient per individual compared to hand rearing adults from
parasitized hosts (Bartlett 2000) but lack associated host data. Malaise
traps are less effective for Cerambycidae and social wasps compared Summary
to hand collecting (Noguera et al. 2002, 2007; Silveira 2002). Malaise traps revolutionized the collection of insects and other
Both pitfall and Malaise traps collect highly diverse arthropod arthropods and many iterations and refinements in design have been
assemblages but generally have little overlap in the species they proposed over the last 83 yr. Malaise trap catch is generally dom-
sample, so are complimentary techniques when trying to assess inated by Diptera and Hymenoptera, with actively flying species of
overall biodiversity with a limited number of techniques (Missa et al. other orders (e.g., lebiine carabids) also commonly represented. The
2009, Skvarla 2015; Araneae: Oxbrough et al. 2010; Carabidae and traps can be used for general collecting and biodiversity surveys as
Curculionoidea: Skvarla and Dowling 2017; Platygastridae: Manoj well as investigating insect movement, vertical stratification, and diel
et al. 2017). and seasonal patterns of abundance.
Considering more specific applications, social wasps are collected Many factors influence the taxa collected. Abiotic factors, such
more efficiently with watered-down honey bait than with Malaise as weather, season can variously affect the behavior of target taxa
traps (Noll and Gomes 2009) and when sampling pecan weevil and influence the species trapped. The trap itself can vary by design,
(Curculio caryae), Malaise traps situated in the first crotch of pecan orientation, and placement or the addition of various lures, all of
trees collected more beetles than cone emergence traps (Dutcher which will impact the resulting trapped species. Biotic factors that
et al. 1986). affect trap catch include the type and density of surrounding vegeta-
Traps that are left exposed (e.g., pitfall traps without a cover) or tion and species-specific behavior.
have the collection cup is at the bottom of the trap (e.g., bottom col- Whether Malaise traps are more or less efficient at collecting cer-
lector of SLAM traps, Lindgren funnel traps, and panel and intercept tain groups compared to other trap types is taxon-specific. Pitfall
traps) can be negatively influenced by rainfall, with excess water ei- traps and leaf litter extraction have the least overlap with Malaise
ther diluting the preservative or flooding the collection cup such that traps in taxa collected and are therefore good options to pair with
specimens are lost (M.J.S., personal observation). Malaise traps do Malaise traps when trying to maximize the arthropod diversity sam-
not experience such issues due to the design of the trap, so may be pled while minimizing effort.
more suitable in areas with heavy rainfall. Malaise traps are not without their drawbacks, however.
Commercially available traps are expensive and can be easily destroyed
by strong winds or large mammals like bears. And because they often
Limitations and Considerations collect large numbers of arthropods, the use of Malaise traps may raise
While many commercially available Malaise traps are well con- concerns over the capture and killing of nontarget by-catch.
structed, they are still ultimately made from fabric. Because of this, While Townes’ fear that Malaise trap development would stop
they are easily torn in high winds or by animals, such as moose that has not fully been realized, there is still potential to refine the trap
may view them as a threat or bears that seek out the ethanol that is design. Future efforts should focus on determining the differences
often used as a preservative. between various trap designs and improvements and the effects
Because Malaise trap samples are often extremely diverse taxo- they have on the taxa collected. Additionally, efforts should also
nomically and may consist of thousands or tens of thousands of in- be made to maximize the use and storage of Malaise trap by-catch,
dividual arthropods, it can be time consuming to sort trap samples as every trap represents a unique snapshot of the environment that
when specific taxa are targeted, especially when those taxa are better will never be precisely replicated in the future. Such efforts could
collected by other methods. Additionally, Malaise traps often gen- include collaborations between institutions that result in the storage
erate large amounts of by-catch, that is, arthropods that are collected and sharing of valuable materials that would otherwise be destroyed.
and killed but are not of interest to the study in question. While this Finally, Malaise traps have proven to be useful in controlling cer-
by-catch represents a unique snapshot of an area in space and time, tain pests when specific baits are used. This suggests the trap design
it is often discarded for a variety of reasons, including lack of space has potential for controlling other pests using the correct baits or
or funding to store the samples, lack of time or expertise to process circumstances.
and curate the by-catch, and lack of interest from the investigator
when the by-catch is not the taxon of interest (Buchholz et al. 2011,
Spears and Ramirez 2015). However, analyzing by-catch, or at least References Cited
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