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Revista de Biología Tropical, ISSN: 2215-2075 Vol. 69(4): 1224-1232, October-December 2021 (Published Nov. 25, 2021)

Trap rebuilding by Myrmeleon brasiliensis larvae (Neuroptera:

Myrmeleontidae) in response to flooding: the effect of body size

Vitor Gabriel de Oliveira Leite¹; https://orcid.org/0000-0003-4104-3093

Raquel Braga Aquino Florenciano¹; https://orcid.org/0000-0001-9019-5398

Edihanne Gamarra Arguelho¹; https://orcid.org/0000-0002-4549-6762

Tatiane do Nascimento Lima¹,²*; http://orcid.org/0000-0002-0656-1170

1. Universidade Federal de Mato Grosso do Sul, Campus de Aquidauana, Biodiversity Studies Laboratory, Rua Oscar

Trindade de Barros, Bairro da Serraria, Aquidauana, Mato Grosso do Sul, Brazil; vg454053@gmail.com, raquell.

aquino12@gmail.com, edihanne.arguelho@ufms.br

2. Universidade Federal de Mato Grosso do Sul, Programa de Pós-Graduação em Recursos Naturais, Faculdade de

Engenharias, Arquitetura e Urbanismo e Geografia, Cidade Universitária, Campo Grande, Mato Grosso do Sul, Brazil;

tatianenlima@gmail.com (*Correspondence)

Received 19-V-2021. Corrected 20-X-2021. Accepted 10-XI-2021.

ABSTRACT

Introduction: Immature forms of the antlion Myrmeleon brasiliensis (Neuroptera, Myrmeleontidae) build traps

in dry sandy soil to capture prey.

Objective: The aim of the present study was to investigate how the waterlogging of the soil due to rain affects

the trap rebuilding and relocation behavior of M. brasiliensis of different sizes.

Methods: The study was conducted between July and December 2019. Larvae M. brasiliensis were observed

and collected from a forest reserve in the municipality of Aquidauana in the state of Mato Grosso do Sul, Brazil.

Results: In the natural environment, most larvae rebuilt their traps in the same location seven days after the

simulation of rain, with a smaller diameter than that observed prior to the simulation of rain. In the labora-

tory, the movements of M. brasiliensis larvae and rebuilding of the traps after the waterlogging of the soil was

affected by body size. Larger larvae moved more and were more likely to rebuild their traps.

Conclusions: The saturation of the soil affects the foraging of M. brasiliensis larvae, which are impeded from

rebuilding their traps for a period. In situations of long periods of saturated soil, the mortality rate of the larvae

is high and rebuilding of the traps occurs after the soil dries out, but with a smaller trap size. These data suggest

that changes in the rainfall pattern can affect the population structure of M. brasiliensis larvae, with the selection

of larger individuals in situations of more severe rains. In this process, the smaller larvae are more affected, as

their foraging is impeded.

Key words: antlion; construction behaviour; foraging; Myrmeleontinae; rainfall.

de Oliveira Leite, V. G., Aquino Florenciano, R. B., Gamarra

Arguelho, E., & do Nascimento Lima, T. (2021). Trap

rebuilding by Myrmeleon brasiliensis larvae (Neuroptera:

Myrmeleontidae) in response to flooding: the effect of

body size. Revista de Biología Tropical, 69(4), 1224-1232.

https://doi.org/10.15517/rbt.v69i4.44478

ECOLOGY

Many organisms build traps to capture

prey. This foraging strategy evolved indepen-

dently in different groups, such as spiders and

insects (Alcock, 1972; Hansell, 2007; Ruxton

& Hansell, 2007). Sit-and-wait foraging behav-

ior involving trap building diminishes energy

that would otherwise be spent in the active

search for prey. However, energy is spent on

the building and maintenance of the trap (Botz

et al., 2003; Gotelli, 1993). Thus, environmen-

tal disturbances that cause the destruction of

the trap generate greater energy expenditure for

these organisms, which can affect their fitness

(Lima & Silva, 2017; Lucas, 1985).

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Immature forms of the family Myrmeleon-

tidae (Neuroptera order), which are known as

antlions, have trap-building behavior. Among

species of the sub-family Myrmeleontinae,

traps are built in a funnel shape in dry sandy

soil (Triplehorn & Johnson, 2005; Youthed &

Moran, 1969;). The three-dimensional archi-

tecture of the trap and the granulometry of the

soil cause prey items (arthropods moving along

the ground) to fall into the trap (Devetak et

al., 2020; Humeau et al., 2015;). Larger traps

enable the capture of both large and small prey

items. Upon falling into the trap, the prey is

directed to the bottom of the pit, where the

larva awaits to attack with its long falciform

mandibles (Heinrich & Heinrich, 1984; Nonato

& Lima, 2011).

Trap size is proportional to the body size

of the antlion larva and is positively related

to the successful capture of prey items (Dias

et al., 2006; Missirian et al., 2006). A larger

trap diameter increases the abundance and size

range of the prey, which is the main benefit of

increasing the capturing surface of the trap.

However, a larger pit requires greater energy

expenditure for maintenance and rebuilding

in cases of more severe damage (Burgess,

2009; Griffiths, 1985; Lima & Silva, 2017;

Lucas, 1985)

The destruction of antlion traps can occur

due to the interference of neighboring larvae,

which toss sand out of their funnels during

the building and maintenance of their traps

(Day & Zalucki, 2000; Simberloff, 1978). Trap

destruction can also occur due to environmen-

tal factors, such as wind and rain. In natural

environments, some species, such as Myrmel-

eon brasiliensis, Myrmeleon immaculatus and

Myrmeleon crudelis, build traps in locations

protected from the direct action of rain, such

as under plantlets or fallen tree trunks (Gotelli,

1993; Lima & Faria, 2007), whereas other spe-

cies, such as Cueta lineosa and Morter obscu-

rus, prefer open areas exposed to direct sunlight

and rain (Devetak et al., 2020; Griffiths, 1980).

Independently of the selection of the trap build-

ing location, all traps are destroyed in situations

of strong rains, which can affect development,

as the larvae are impeded from foraging until

the soil dries out again. In the Cerrado biome

(savanna) of Brazil, M. brasiliensis larvae were

found to wait approximately five days before

rebuilding their traps after rainfall (Freire &

Lima, 2019).

Larvae of Myrmeleon brasiliensis (Návas,

1914) pass through three instars before the

formation of the pupa for the emergence of the

adults. The traps built by these larvae range

from 9 to 35 mm in diameter (Missirian et al.,

2006). Given this variation in trap size, cost-

benefit ratios associated with the rebuilding

and relocation of a trap must vary as a func-

tion of body size and trap size. Larger larvae

have a greater energy supply, due to a greater

fat reserve. Thus, the hypothesis raised in this

work is that larger larvae may bear the costs

of relocating and rebuilding the traps (after

disturbing their traps) more efficiently than

smaller larvae. In this case, after a rain event,

the larger larvae would be the first to rebuild

their traps. The aim of the present study was

to investigate how the waterlogging of the soil

due to rain affects the building behavior and

relocation of the traps of M. brasiliensis larvae

of different sizes.

MATERIALS AND METHODS

This study was conducted in a Permanent

Protection Area (1 600 000 m²) located in the

municipality of Aquidauana in the state of

Mato Gross do Sul, Brazil (20°26’36” S &

55º40’02” W). Mean annual temperature and

rainfall in the area are approximately 26 °C and

1 250 mm, respectively. Rains are concentrated

between October and March and the monthly

precipitation index diminishes considerably

between May and September, reaching zero

(Coutinho, 2000). The present study was con-

ducted between July and December 2019.

In the reserve, ten quadrants measuring 40

x 40 cm were established with a metric tape

and string, separated by a distance of 1 m. In

each quadrant, the position of the antlion traps

was marked with wooden stakes (10 x 1 cm)

and the diameter of the traps was measured.

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The occurrence of rain was simulated with

the aid of a watering can, to observe the effect

of waterlogging on the rebuilding of the M.

brasiliensis traps. A total of 200 ml of water

were poured on each quadrant. This quantity of

water was chosen for being sufficient to destroy

the traps without forming a puddle. After

the rain simulation, the traps were observed

daily. The presence of traces of the larvae was

observed for the construction of new traps. In

this way, it was possible to follow the move-

ment of the M. brasiliensis larvae. Seven days

after the rain simulation, the rebuilding of the

traps was observed, and the diameters of the

rebuilt traps were measured. There was no

occurrence of natural rain during this period.

In the second part of the study, two experi-

ments were set up in the laboratory. For such,

M. brasiliensis larvae were collected from their

traps through visual searches and transported

to the laboratory. In the first experiment, 60

larvae were measured for body size (from the

head to the end of the abdomen, excluding the

mandible) and individually placed into plastic

recipients (20 cm length x 15 cm width x 10

cm height) containing 195 g of sand removed

from the collection site. After 24 h (time given

for all larvae to build traps), the diameter of

the traps was measured, and the location of

the larvae was marked. To evaluate the effects

of waterlogging, the occurrence of rain was

simulated. A total of 9 ml of water were poured

directly into the trap. Thus, the soil around the

trap remained dry, enabling the movement of

the larva. After seven days, observations were

made of the rebuilding of the traps, trap size

and whether the trap had been relocated within

the plastic recipient.

In a second experiment, another 60 larvae

were measured for body size (head to abdomen)

and placed into plastic recipients (20 cm length

x 15 cm width x 10 cm height) containing 195 g

of sand removed from the collection site. As

in the previous experiment, trap diameter was

measured after 24 h. Next, the effect of water-

logging of the soil after rain was observed. For

such, 30 ml was water were poured directly

onto the trap, forming a puddle so that the larva

became submerged. After seven days, the water

was drained, and the recipients were placed to

dry in a plant nursery. For 15 days, observa-

tions were made of the rebuilding of the traps,

trap size and mortality.

Al measurements were performed with

digital calipers (precision: 0.01 mm). Trap

size before and after the simulation of rain

in the field and in the laboratory, experiment

was compared using the paired t-test. Logistic

regression was performed to determine the

probability of the rebuilding of the trap after

the waterlogging of the soil in relation to body

size of the M. brasiliensis larvae as well as the

probability of the relocation of the trap in rela-

tion to body size.

RESULTS

An average of six M. brasiliensis larvae

(range: two to 11) were found in each quadrant

demarcated in the natural environment prior to

the rain simulation. After the rain simulation,

an average of four larvae were found per quad-

rant. The rebuilding rate at the same site was 61

%. The traps rebuilt after the rain were smaller

(d.f. = 36, T = 2.15, P = 0.03). Mean (± stan-

dard deviation) trap size was 26.11 (± 11.84)

mm prior to the rain simulation and 22.55 (±

10.50) mm after the rain simulation.

In the first laboratory experiment, 75 %

of the M. brasiliensis larvae rebuilt their traps

after the rain simulation, 11 % of which rebuilt

at the same site. Among those that relocated

their traps, the mean distance from the original

site was 55.35 mm. The probability of reloca-

tion increased as a function of body size (d.f. =

1, X² = 24.33, P = 0.01). The probability of trap

reconstruction also increased as a function of

larva size (d.f. = 1, X² = 20.35, P = 0.01) (Fig.

1A, Fig. 1B). Mean trap size was 25.83 ± 8.94

mm prior to rain simulation and 27.51 ± 7.54

mm after rain simulation. This difference was

non-significant (d.f. = 44, T = -1.43, P = 0.15).

In the second laboratory experiment, when

the M. brasiliensis larvae spent seven days in

inundated soil (all the sand was waterlogged),

67 % (considering a total of 60 larvae) did not

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rebuild their traps in the 15-day observation

period after the draining of the water and dry-

ing of the soil. Among those that rebuilt, a sig-

nificant reduction in trap size was found after

the waterlogging of the soil (d.f. = 19, T = 2.54,

P = 0.01). Mean trap size was 26.33 ± 12.58

mm before waterlogging and 21.88 ± 6.68 mm

after waterlogging.

The probability of reconstruction (d.f. = 1,

X ² = 21.61, P = 0.01) and relocation the trap

(d. f. 1 = X ² = 23.74, P = 0.01) after a period of

seven days in saturated soil increased as a func-

tion of body size (Fig. 2A, Fig. 2B). The mor-

tality rate at the end of the experiment (seven

days in saturated soil + 15 days of observation)

was 52 %. Some larvae were able to rebuild

their traps when the soil dried but died in the

subsequent days. Others died during the water-

logging period.

DISCUSSION

For trap-building insects, variables that

cause the destruction of the trap should be

considered during decisions regarding foraging

behavior (Gotelli, 1993; Lima & Lopes, 2016).

The selection of an adequate site for building a

trap is an important component of the foraging

behavior of sit-and-wait predators, especially

when movements are rare. The relocation and

the rebuilding of a new trap in the absence of

environmental disturbance rarely occur among

antlions, possibility due to the metabolic cost

associated with the rebuilding of the trap and

Fig. 1. A. Probability of displacement of M. brasiliensis larvae as function of body size after waterlogging of soil caused

by simulation of rainfall. B. Probability of reconstruction of traps by M. brasiliensis larvae as function of body size after

waterlogging of soil caused by simulation of rainfall (First Experiment).

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the risk of cannibalism (Lima, 2016; Lucas,

1985). In the present study, more than half (61

%) of the M. brasiliensis larvae in the natural

environment rebuilt their traps at the same site

seven days after the simulation of rain, reveal-

ing that the larvae remained in the same loca-

tion, waiting for the soil to dry.

Other studies have also demonstrated that

antlion larvae exhibit strong site fidelity for

the construction of their traps (Crowley &

Linton, 1999; Lucas, 1985). One of the rea-

sons for this behavior is the selection of a dry,

warm micro-habitat, as reported by Miler et

al. (2019) for antlions found the tropical forest

of Borneo. Moist soil hinders trap building by

agglutinating the soil particles and reducing

sliding movements between the grains, thereby

restricting the movement of the antlion larvae

(Devetak et al., 2005; Lucas, 1982). These

characteristics lead the trap-building larvae to

select dry locations.

In the natural environment, the determina-

tion of the location for the rebuilding of traps

implies choices regarding the availability of

prey items, intraspecific competition, and abi-

otic factors, such as the granulometry of the

soil (Devetak & Arnett, 2015; Farji-Brener,

2003; Scharf et al., 2008; Scharf & Ovadia,

2006). In the present study, the M. brasiliensis

larvae relocated the traps less in the natu-

ral environment compared to the laboratory.

This may have been due to the fact that the

larvae were alone in the plastic recipients in

the laboratory. As these larvae are cannibals

Fig. 2. A. Probability of displacement of traps by M. brasiliensis larvae as function of body size after waterlogging of soil

caused by simulation of rainfall. B. Probability of reconstruction of traps by M. brasiliensis larvae as function of body size

after waterlogging of soil caused by simulation of rainfall (Second Experiment).

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(Lima, 2016), avoiding relocation to sites close

to neighboring larvae could help ensure that the

larva does not become prey.

In the study conducted by Lima et al.

(2019), larger larvae were found more distant

from the clusters in which the oviposition of

the adult females occurred, suggesting that lar-

vae of a certain size perform movements with

greater frequency. According to Lucas (1985)

and Fisher (1989), larger larvae are found

more distant from their clusters in response

to temperature, as larger antlion larvae are

less capable of supporting warmer locations.

Studying Myrmeleon uniformis, Faria et al.

(1994) found that average trap size was sig-

nificantly larger in open areas than areas pro-

tected from direct sunlight, whereas Scharf et

al. (2008) found that Myrmeleon hyalinus in

desert environments selected shaded locations

and often moved to shaded areas when exposed

to the sun.

The laboratory experiment showed that

relocation for the rebuilding of new traps

occurs as a function of body size in M. brasil-

iensis, as larger larvae moved their traps more

than smaller larvae. The fact that the reloca-

tion of larvae is affected by body size may be

related to the greater energy storage capability

of larger individuals compared to smaller indi-

viduals. Lucas (1985) found that the metabolic

rates of Myrmeleon carolinus and Myrmeleon

crudelis were approximately 10-fold greater

during trap building than during rest. Bearing

this metabolic cost likely requires a certain

energy reserve, which is more difficult for

smaller larvae due to their lower prey capture

rate (Nonato & Lima, 2011). However, other

researchers found that small antlions relocated

with greater frequency than larger individuals

(e.g., Griffiths, 1993), which may suggest a

specific response for each species.

When the larvae were exposed to seven

days in waterlogged soil with no possibility of

relocating to areas with dry soil, the mortal-

ity rate was high (52 %). Moreover, the prob-

ability of rebuilding the traps after this period

was higher among M. brasiliensis larvae with

a larger body size. Thus, body size was an

important variable to enabling the rebuild-

ing of traps after the waterlogging of the soil.

However, the rebuilt traps were smaller than

those found prior to the disturbance. Varia-

tions in trap size occur in response to success

in capturing prey and the energy expenditure

caused by the disturbance to the trap (Jenkins,

1994; Hauber, 1999; Liang et al., 2010). Thus,

besides the high mortality rate, the foraging of

M. brasiliensis larvae can also be affected in

situations of long periods of waterlogged soil,

as the rebuilt trap is smaller and the success of

capturing prey items is positively related to trap

size (Nonato & Lima, 2011).

Trap maintenance is important to ensuring

antlion larvae access to food. The rebuilding

of traps after a disturbance generally occurs on

the same or following day (Boake et al., 1984;

Eltz, 1997). However, when rain is the disturb-

ing force, the larvae need to wait for the soil to

dry before rebuilding, which can take five days

in the Cerrado biome (Freire & Lima, 2019). In

the present study, trap rebuilding and relocation

was governed by the size of the larvae. Thus,

the waterlogging of the soil due to rainfall

affects the foraging of M. brasiliensis larvae,

which are impeded from rebuilding their traps

for a period. In situations of long periods of

waterlogged soil, the larvae have a high mortal-

ity rate; rebuilding of the traps occurs after the

soil dries out, but the traps are smaller. These

data suggest that changes in the rainfall pat-

tern can affect the population structure of M.

brasiliensis larvae, with the selection of larger

individuals in situations of more severe rains.

Ethical statement: the authors declare

that they all agree with this publication and

made significant contributions; that there is no

conflict of interest of any kind; and that we fol-

lowed all pertinent ethical and legal procedures

and requirements. All financial sources are

fully and clearly stated in the acknowledge-

ments section. A signed document has been

filed in the journal archives.

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ACKNOWLEDGMENTS

This research was developed with the

support of the Universidade Federal de Mato

Grosso do Sul – UFMS/MEC – Brazil and the

Universidade Estadual de Mato Grosso do Sul

(UEMS) - Aquidauana.

RESUMEN

Reconstrucción de trampas por larvas de

Myrmeleon brasiliensis (Neuroptera: Myrmeleontidae)

tras las inundaciones: efecto del tamaño corporal

Introducción: Los insectos inmaduros de hormiga león

de la especie Myrmeleon brasiliensis (Neuroptera, Myr-

meleontidae), construyen trampas en el suelo arenoso seco

para capturar a sus presas.

Objetivo: El objetivo de este trabajo fue analizar cómo la

inundación del suelo, como la lluvia, afecta al comporta-

miento de reconstrucción y de desplazamiento de las tram-

pas de M. brasiliensis según las diferentes tallas.

Métodos: El estudio fue conducido entre julio y diciembre

del 2019. Las larvas fueron observadas y recolectadas en

una Reserva Forestal en la ciudad de Aquidauana, Estado

de Mato Grosso del Sur, Brasil.

Resultados: Observamos que, en el ambiente natural,

siete días después de la lluvia simulada, la mayoría de las

larvas reconstruyeron a sus trampas en el mismo sitio y con

diámetro más pequeño que el observado antes de la lluvia

simulada. En el laboratorio fue observado que después de

la inundación del suelo, el desplazamiento de las larvas y

la reconstrucción de las trampas es afectado por el tamaño

de su cuerpo. Así, larvas de M. brasiliensis más grandes se

desplazan más y presentan una probabilidad más grande de

reconstruir a sus trampas.

Conclusión: De esa manera, la inundación del suelo afecta

el forrajeo de las larvas M. brasiliensis, que se quedan

impedidas por un tiempo durante la reconstrucción de

sus trampas. Además, en situaciones de largos periodos

de inundación del suelo, las larvas presentan una alta tasa

de mortalidad y la reconstrucción de sus trampas ocurre

después que se seca el suelo, pero esas trampas suelen

ser más pequeñas. Esos datos sugieren que es posible

que variaciones en el patrón de lluvia pueden afectar la

estructura poblacional de las larvas M. brasiliensis, pues en

situaciones de lluvias más fuertes, las larvas más grandes

suelen tener más éxito. En este proceso, las larvas más

pequeñas se ven más afectadas, ya que se impide su bús-

queda de alimento.

Palabras clave: hormiga-león; comportamiento de cons-

trucción; forrajeo; Myrmeleontinae; lluvia.

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