Applied Soil Ecology 150 (2020) 103460

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Applied Soil Ecology

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Short communication

Food consumption of the cockroach species Blaptica dubia Serville T

(Blattodea: Blaberidae) using three leaf litter types in a microcosm design
⁎
Masoud M. Ardestania,b, , Vladimír Šustrb, František Hniličkac, Jan Frouza,b
a
Institute for Environmental Studies, Charles University in Prague, Benátská 2, Prague CZ-12801, Czech Republic
b
Institute of Soil Biology and SoWa Research Infrastructure, Biology Centre, Czech Academy of Sciences, Na Sádkách 7, České Budějovice CZ-37005, Czech Republic
c
Czech University of Life Sciences, Department of Botany and Plant Physiology, Kamýcká 129, Praha 6, Suchdol CZ-16500, Czech Republic

A R T I C LE I N FO A B S T R A C T

Keywords: This study investigated the effect of different plant litter types and developmental stages of the cockroach species
Assimilation efficiency Blaptica dubia on food consumption, animal growth, and excrement production. Three leaf litter species Alnus
Energy budget glutinosa, Acer platanoides, and Quercus robur were selected as model plants in a microcosm test in which plant
Excrement production litter was consumed by three different cockroach instar groups during five consecutive one-week intervals. At
Feeding activity
each interval, litter consumption, animal weight, and produced excrements were measured. The results showed
Organic matter decomposition
slightly higher consumption of A. glutinosa compared to other leaf litter species in all three age/size groups of
Soil ecosystem
Soil organisms animals. Animals gained weight with time, especially Group B (medium-sized class animals) feeding on Alnus.
The production of excrements was higher in Group C (big class animals) and increased in Alnus among litter
species. Foliar litter consumption and excrement production rates were highest in B and C animal groups and in
Alnus treatments; however, animal growth rates did not show a clear pattern, except for A. glutinosa. Measured
energy contents showed highest values for animals, but slightly higher for leaves compared to excrements. Our
results show that using energy budget parameters can help explain the relationship between litter type, cock-
roach sizes, and their consumption in this microcosm experiment.

1. Introduction
Soil organisms adapt themselves to the resource availability and
environmental characteristics in order to obtain most suitable com-
promise for their life requirements and to reach the best condition for
their survival and maintenance. The feeding activity of soil organisms,
especially decomposers, has been one of the main subjects that have
been noticed for many years (Lavelle et al., 1997; Hättenschwiler et al.,
2005; Frouz et al., 2015a, 2015b). These organisms play an important
role in the breakdown and development processes of organic material in
natural ecosystems (Wardle et al., 2004; Vos et al., 2011; Frouz, 2018).
In this way, they are important for two major reasons; plant litter is
major carbon input in soil, and its decomposition plays a principal role
in global carbon cycle (Lavelle et al., 1997; Frouz et al., 2015b; Frouz,
2018).
Previous research has shown that food quality and food availability
has strong impact on various physiological responses in soil organisms
(Cornwell et al., 2008; Makkonen et al., 2012). This brings an extra
selection pressures on the animals to show effective mechanisms of
regulating the amounts and balance of nutrients which they extract
from their environment (Simpson and Raubenheimer, 1993). They may
even compensate for nutrient imbalance by changing the rates of food
intake and by changing their food type to be able to quickly recover
from food deficiency (Chambers et al., 1995; Jones and Raubenheimer,
2001; Raubenheimer and Jones, 2006). Using energy budget experi-
ments will help researchers to find out more about the essential re-
quirements of the organisms for having maximum trade-off between
their energy investments in their natural habitats. In these experiments,
the consumption of litter by a specific herbivore species, the changes in
the body size (length, weight), and the produced excrements will be
monitored for a certain period of time. Energy budget parameters can
reveal more insights into the resource allocation in different physiolo-
gical life stages of the organisms and into the factors that affect these
processes.
The cockroach species Blaptica dubia Serville (Blattodea:
Blaberidae), originally from South America and named as “Dubia
cockroach”, has been one of the soil organisms found in tropical region
with high abundance in rain forests. They play an important role in
litter breakdown and decomposition processes. They feed on different
leaf litter species and listed in the opportunistic omnivorous species in

⁎
Corresponding author at: Institute for Environmental Studies, Charles University in Prague, Benátská 2, Prague CZ-12801, Czech Republic
E-mail addresses: mortazavi_m2000@yahoo.com, masoud.ardestani@bc.cas.cz (M.M. Ardestani).

https://doi.org/10.1016/j.apsoil.2019.103460
Received 23 September 2019; Received in revised form 27 November 2019; Accepted 28 November 2019
Available online 03 December 2019
0929-1393/ © 2019 Elsevier B.V. All rights reserved.
M.M. Ardestani, et al.
Table 1
The measured values for each instar of Blaptica dubia. The values are in mm and the 95% range of each parameter in 5 animals. For measuring each parameter
QuickPHOTO MICRO 2.3-Microscope software (Promicra, Czech Republic) was used. The animal groups were: A, small class animals (instar 1 to 3); B, medium-sized
class animals (instars 4 and 5); C, big class animals (instars 6 and 7).
N Instar 1 Instar 2 Instar 3
5 5 5 5
Pronotal width 4.30–5.42 6.33–7.35 7.51–9.44
Pronotal length 1.85–2.34 2.91–3.12 3.20–3.67
Head width 3.05–3.30 4.67–4.75 5.80–5.92
Total length 8.36–11.7 12.7–14.8 15.03–16.9
these areas (Schal et al., 1984; Mullins, 2015). In addition to their
abilities in litter consumption, they produce big excrements which are
easy to collect and examine for their physical structure and chemical
content. Consequently, they are among one of the important species for
producing organic matter in soil and soil formation. They have ap-
proximately long life stages with different developmental instars de-
pending on temperature. In laboratory, they are easy to rear in large
numbers and need minimum requirements for their maintenance.
There are several studies in the literature where the feeding activ-
ities of soil organisms have been studied (e.g., Mullins and Cochran,
1986; Hättenschwiler et al., 2005; Ristok et al., 2019). However, there
is a lack of physiological information on the selected cockroach species
B. dubia and their feeding behavior in simulated natural condition.
Moreover, there are little systematic studies exploring how age/size of
invertebrate decomposer development and food type affect consump-
tion and other characteristic of animal feeding. Up to now, there is no
information about energy budget parameters of cockroaches in the
literature. The cockroach B. dubia represents unique model to explore
this question. Based on above-mentioned facts about our selected test
species, the aim of the present study was to investigate the effect of
different plant litter types and developmental stages of B. dubia on their
leaf consumption and excrement production. We quantified the en-
ergetic expenditure of key physiological processes: food consumption,
growth (increase in animal fresh weight), and excrement production by
considering the effect of time, as an interactive parameter. We hy-
pothesize that B. dubia may consume alder leaf litter more than other
litter types based on its higher N content (Frouz et al., 2015b) and the
interaction of age/size of the animals and time influence litter con-
sumption.
2. Materials and methods
2.1. Plant litter
Three leaf litter species were used in the present study: Alnus glu-
tinosa (alder), Quercus robur (oak), and Acer platanoides (maple). Fresh
leaf litters were collected from a forest close to České Budějovice, Czech
Republic. They were air-dried in lab condition and stored for the ex-
periment. Then, the leaves were cut to small pieces (approximately
2 × 1 cm) for the bigger class animals (Groups 1 and 2, see below) and
to very fine pieces (approximately 0.2 cm) using a blender for the small
class animals (Group 3). The reason for cutting leaves to smaller pieces
is that these dry and fragile foliar litters are subjected to physical
change by insects and saprophytic microbial groups as well as natural
conditions (rain, wind, etc.). The air-dried leaves were then dried in an
oven at 60 °C for 24 h to have dry weight of leaves at the beginning of
each interval.
2.2. Test species
The cockroach species B. dubia Serville (Blattodea: Blaberidae) were
purchased from a local shop several years ago and reared under la-
boratory conditions with the temperature of 25 °C, relative humidity of
2
Applied Soil Ecology 150 (2020) 103460
Instar 4 Instar 5 Instar 6 Instar 7
5 5 5
11.5–12.4 14.4–14.6 15.3–16.9 18.8–19.3
3.70–4.18 5.46–5.92 6.24–6.64 9.13–9.28
6.70–6.80 7.20–7.35 8.55–8.75 10.1–10.2
18.0–19.3 23.1–26.8 28.2–33.8 34.8–40.1
60%, and normal day-night photoperiod. In big plastic containers
(38 × 25 × 15 cm), small wood pieces, leaf litter pieces, and egg-
shields were provided for them. Every two days, food (cabbage, fruits,
and cereal grains for bigger animals and dry wheat powder for smaller
animals) was added to the main culture containers. At the start of the
experiment, individuals were selected from the main stock, starved for
2 days, and then distributed in the experimental containers
(8.5 × 9 × 4.5 cm) with a lid. The boxes were aerated through holes in
the lid. The selection of animals for each instar classes was based on Wu
(2013) and Wu et al. (2013) with known developmental stages. These
stages were classified by their pronotal length and width, head size and
width, and body length. We prepared 3 animal age/size classes (groups)
for the experiment; Group A: small class animals (1st, 2nd, and 3rd
instars), Group B: medium-sized class animals (4th and 5th instars), and
Group C: big class animals (6th and 7th instars) (see Fig. S1, Supple-
mentary material). In each experimental plastic container, 6 (small
class), 5 (medium-sized class), and 4 (big class) animals were added.
Different parameters in each individual were measured using Quick-
PHOTO MICRO 2.3-Microscope software (Promicra, Czech Republic)
(Table 1). At the start of the experiment, the average fresh weight of
each animal in Groups A, B, and C was 0.10 g, 0.50 g, and 1.25 g, re-
spectively (n = 20).
2.3. Experimental design
Nine treatment variants including three leaf litter species were
combined with three animal sizes (3 × 3). Three replicates for each
litter species and animal size were considered. In each experimental
container, approximately 0.50 g of Q. robur, 0.60 g A. platanoides, and
1.30 g A. glutinosa oven-dried leaf litters were added. The amount of
leaf litter for each plant species was determined based on a pilot test in
which the feeding rate of each animal group was investigated
(Ardestani, unpublished). Based on these tests, we could find the exact
amount of each litter type for each animal species during each interval.
The dry leaves were sprayed with distilled water only at the beginning
of the test. The experimental container was kept in the big container
having water inside to moisturize the atmosphere of the box. This al-
lowed the leaves to be moistened and palatable for the animals during
each interval. Suitability of this method was proved in the pilot tests.
Five one-week intervals were conducted in the same way (5 weeks
in total). For each leaf litter species, additional three replicates were
considered as controls which were maintained at the same experimental
conditions and their fresh and dry weight were measured at the be-
ginning and at the end of each interval to account for microbial activ-
ities (degradation, see Table S1, Supplementary material; and detailed
method in Supplementary material). Every day, the produced excre-
ments from each replicate were collected and stored in separate con-
tainers to prevent microbial activity. Experimental containers were kept
in the climate chambers under 25 °C to simulate natural condition for
the cockroaches.
M.M. Ardestani, et al.
2.4. Plant litter, animal weight, and excrements
At the end of each interval, the remaining leaf litter was collected
from each replicate, oven-dried at 60 °C for 24 h and weighed as un-
consumed dry weight of leaves. The difference between the un-con-
sumed weight and the initial dry weight at the start of experiment was
used as consumed leaf litter. At the end of each interval, the animals
from each replicate were weighed as end fresh weight. Again, the dif-
ference between start and end values for the fresh weight of each re-
plicate was used for calculating the growth of animals during each in-
terval. At the end of each interval, the daily collected excrements from
each replicate were compiled, weighed (fresh weight) and after oven-
drying (at 60 °C for 24 h), dry weight of excrements was obtained.
Three additional replicates of oven-dried leaf litters (not added to the
experimental boxes) were prepared at the beginning of the experiment
for calorimetric analyses.
At the beginning of the test, ten individuals from each animal group
were taken directly from the main culture and fresh weight of them was
obtained. Then, they were frozen and dried at 60 °C for 24 h to measure
their dry weight. These sampled animals were used for calorimetric
analyses and calculation of energy budget parameters. In addition and
for the same purpose, some animals from the experimental replicates
were selected randomly after finishing the 5th interval. Then, fresh
weight of them was obtained and their dry weight was also measured
after freeze-drying. All dry materials were weighed using a micro bal-
ance with an accuracy of 0.001 g (KERN, EG 620-3NM, Germany). The
energetic content of leaf litter, excrements, and the animals at the start
and at the end of experiment was established by burning of the dry
materials in a micro-calorimeter (Ika-Calorimeter C700T, Hamburg,
Germany).
2.5. Data analysis
Full description of the method for obtaining energy budget para-
meters is shown in Supplementary material. Normality and homo-
geneity of data were tested using Shapiro-Wilk and Bartlett tests, re-
spectively. For comparing the difference in the consumed leaf litter,
fresh weight of animals, and excrements produced among all different
animal groups and different plant litter species eaten by those in-
dividuals at all intervals, consumption rates, and excrement rates,
three-way analysis of variance (ANOVA) was used. In addition, a two-
way ANOVA was also used to assess the difference in leaf consumption,
animal growth, and excrement production separately in each animal
group (for the effect of litter type) and its interaction with time.
Statistica 13.3 program package was used for statistical analyses.
3. Results
3.1. Litter consumption
Litter consumption was dependent on animal age/size groups
(F2,90 = 283, p < 0.001), litter type (F2,90 = 315, p < 0.001), and
the interaction of animal groups x leaf litter (F4,90 = 65.8, p < 0.001;
Table S2, Supplementary material). When comparing food type in each
animal group, highest litter consumption was found in A. glutinosa
treatment in Group C animals (Fig. 1 and Fig. S2, Supplementary ma-
terial; F2,30 = 174, p < 0.001). In Groups A and B, alder litter was also
more attractive than other leaf litter types (Fig. 1 and Fig. S2;
F2,30 = 31.5, p < 0.001 and F2,30 = 132, p < 0.001; respectively).
Leaf consumption was significantly changed with time in Group A
(F4,30 = 3.84, p < 0.05) and Group B (F4,30 = 3.13, p < 0.05). The
interaction of time x litter type was only observed in Group A animals
(F8,30 = 3.03, p < 0.05). Overall, the consumption of litter was re-
latively stable in time (Fig. S2).
3
Applied Soil Ecology 150 (2020) 103460
3.2. Animal weight
The animals' fresh weight was significantly changed with animal
class (F2,90 = 446, p < 0.001), interval (F4,90 = 2.65, p < 0.05),
litter type (F2,90 = 22.7, p < 0.001), and the interaction of animal
groups x litter (F4,90 = 8.74, p < 0.001). Fresh weight of animals was
quite constant in each animal group, with slightly higher values ob-
served in A. glutinosa treatment in Group C animals (Fig. 1 and Fig. S2;
F2,30 = 3.39, p < 0.05). The same trend was found in Group B animals
feeding on alder litter (F2,30 = 85.6, p < 0.001). In the latter case, the
effect of intervals (F4,90 = 8.40, p < 0.001) and the interaction of litter
type x time (F8,90 = 4.88, p < 0.001) was significant.
3.3. Excrements
The amount of produced excrements was significantly changed
among different animal groups (F2,90 = 256, p < 0.001), litter type
(F2,90 = 234, p < 0.001), intervals (F4,90 = 4.24, p < 0.01), and the
interactions of animal group x interval (F8,90 = 2.25, p < 0.05), an-
imal group x litter type (F4,90 = 52.1, p < 0.001), and interval x litter
type (F8,90 = 2.45, p < 0.05). Highest excrement production was
observed in alder treatments in Group C animals (Fig. 1 and Fig. S2;
F2,30 = 138, p < 0.001). In addition, both other animal groups pro-
duced higher excrements when feeding on A. glutinosa compared to
other litter types (F2,30 = 95.1, p < 0.001 for Group B and
F2,30 = 12.9, p < 0.001 for Group A). Only Group B animals showed
significant changes in excrement production in different intervals
(Fig. 1 and Fig. S2; F4,30 = 4.21, p < 0.01).
3.4. Energy budget parameters
Energy budget parameters were calculated for all animal groups,
litter types, and intervals; for example, Fig. S3 (Supplementary mate-
rial) shows these parameters for A. glutinosa in Group C animals.
Consumption, excrement production, assimilation and growth (pro-
duction) rates were lower at the first interval, and then slightly in-
creased for the first two parameters till the end of experiment. But, a
decrease was observed for assimilation and production rates at intervals
2–4. Then, they were increased at interval 5 (Table 2 and Table S3,
Supplementary material). In addition, assimilation efficiency and pro-
duction efficiency did not show a consistent pattern in different treat-
ments and varied in different intervals. Consumption rates and excre-
ment production rates were always lower for Group A compared to
other two animal groups at different intervals and among different leaf
litters and it followed: Group A < Group B < Group C. When com-
paring the values based on gram fresh weight of animals, Group B va-
lues were always higher than other groups (Table S3; Table S4, Sup-
plementary material). For different leaf species, the consumption rates
were usually: Alnus > Acer > Quercus (Table 2 and Table S3). Gen-
erally, animal growth rates were slightly higher for Group C animals
compared to other two groups and for alder compared to other two
litter types.
3.5. Calorimetric values
When considering the assimilation and excrement production based
on the energetic content of food (leaf litter), animals' dry weight, and
dry excrements, slight differences were found between different treat-
ments at different intervals (one-way ANOVA, p > 0.05; Table S5,
Supplementary material). Higher values were observed for the animals
compared to litter and excrements (p > 0.05). Slightly higher values
were also measured for leaf litter compared to excrements (p > 0.05).
Generally, no clear pattern was observed among leaf litter types and
animal groups (Table S5).
M.M. Ardestani, et al. Applied Soil Ecology 150 (2020) 103460

Fig. 1. The consumed leaf litter (a), animals' fresh weight (b), and produced
1 excrements (c) for three animal groups (A, B, and C) feeding on different leaf litter
(a)
Consumed leaves dry weight (g)

0.9 species during five intervals of the experiment with the cockroach Blaptica dubia
0.8 (Blattodea: Blaberidae). Each column represents the average of five intervals. The
animal groups were: A, small class animals (instar 1 to 3); B, medium-sized class
0.7 animals (instars 4 and 5); C, big class animals (instars 6 and 7). The plant species
0.6 were: Quercus robur (QU), Acer platanoides (AC), and Alnus glutinosa (AL). The
0.5 duration of each interval was one week. All values are based on total dry weight
(g) measured at the end of each interval. Three replicates for each treatment
0.4
(interval/animal/leaf litter) were used. Data represent mean ± standard error
0.3 (SE).
0.2
0.1
0
A-QU

A-AC

A-AL

B-QU

B-AC

B-AL

C-QU

C-AC

C-AL
6
(b)
5
Animal fresh weight (g)

0
A-QU

A-AC

A-AL

B-QU

B-AC

B-AL

C-QU

C-AC

C-AL

0.8
(c)
0.7
Excrements dry weight (g)

0.6
0.5
0.4
0.3
0.2
0.1
0
A-QU

A-AC

A-AL

B-QU

B-AC

B-AL

C-QU

C-AC

C-AL

4. Discussion The results showed time as an important factor when investigating

The results of the present study showed that A. glutinosa was more
consumed by the cockroach species B. dubia when comparing different
leaf litters for each animal group (Fig. 1 and Fig. S2). This is in
agreement with previous studies in which leaf litter consumption had
positively correlated with leaf N content in different soil organisms
(Warren and Zou, 2002; Loranger-Merciris et al., 2008). The C/N ratio
of 14.6 was measured for A. glutinosa compared to that of 58 for Q.
robur (Frouz et al., 2015b). In addition, higher aliphatics, phenolic
compounds, and slightly higher lignin; but lower polysaccharides were
measured for alder compared to oak leaf litter in the latter study (Table
S6, Supplementary material). The relationship between these chemical
compounds of different leaves and B. dubia feeding activity would help
to obtain clearer reasons attributed to lower or higher leaf litter con-
sumption in our future experiments. Moreover, palatability of leaf litter
depends on several factors; such as, litter nutritive value, toughness,
and levels of feeding-deterrents (Zimmer et al., 2002; David and Handa,
2010) which needs to be tested.
feeding of B. dubia. Cockroaches seemed to be adapted to this type of
leaf consumption (as the same leaf species was provided in each in-
terval) and the litter became more palatable and digestible even after
1–2 intervals (Fig. 1 and Fig. S2), especially in alder plants. Although
the results of the present study clearly showed that leaf consumption
would not be affected by this experimental design and time was not a
limiting factor for their feeding activities, longer duration experiments
might be needed to better simulate the natural litter consumption of
cockroaches and investigate the physiological mechanisms attributed to
leaf consumption.
The relationship between excrement production, animal fresh
weight, and leaf consumption was positively observed in our study. For
example, higher leaf consumption in A. glutinosa treatments resulted in
higher excrement production and higher body weight of the animals
(Fig. 1 and Fig. S2; Table 2). This suggests that animals allocated bigger
part of their feeding to their maintenance and growth, in our case
mostly to maintenance. Another part was excreted from the body. The
excrement production was higher in A. glutinosa treatments. But, this

4
M.M. Ardestani, et al. Applied Soil Ecology 150 (2020) 103460

Table 2
The consumption rate, animal growth rate, and excrement production rate (mg dry weight day−1 individual−1) for different animal groups and different leaf species
(panels A to C) in five intervals of the experiment Blaptica dubia (Blattodea: Blaberidae). The animal groups were: A, small class animals (instar 1 to 3); B, medium-
sized class animals (instars 4 and 5); C, big class animals (instars 6 and 7). The plant species were: Quercus rubor (QU), Acer platanoides (AC), and Alnus glutinosa (AL).
The duration of each interval was one week. Three replicates for each treatment (interval/animal/leaf litter) were used.
Parameter Time Alnus glutinosa Acer platanoides Quercus rubor

A B C A B C A B C

Litter consumption rate Interval 1 0.58 16.6 25.6 0.39 5.28 11.5 0.03 3.27 7.38
Interval 2 0.00 18.0 29.4 0.44 6.41 14.3 0.08 2.51 5.45
Interval 3 0.00 17.5 28.6 0.67 7.39 15.0 0.27 2.46 4.65
Interval 4 0.00 22.4 25.9 0.38 8.70 15.7 0.00 3.02 4.57
Interval 5 0.00 28.4 30.2 0.54 7.97 13.6 0.49 3.44 5.09
Animal growth rate Interval 1 0.00 2.46 9.87 0.00 2.06 0.77 0.00 0.59 1.97
Interval 2 0.00 2.45 3.58 0.00 0.43 0.00 0.00 0.40 0.00
Interval 3 0.00 2.18 0.50 0.27 0.42 1.54 0.07 1.26 0.00
Interval 4 0.17 2.49 1.75 0.04 0.00 2.78 0.00 0.33 2.35
Interval 5 0.50 2.74 2.84 0.21 0.61 1.77 0.00 0.00 0.00
Excrement production rate Interval 1 1.31 12.7 20.3 0.53 4.76 10.5 0.32 2.90 5.74
Interval 2 0.83 14.8 26.2 0.68 6.12 13.2 0.34 2.18 4.70
Interval 3 0.59 14.6 25.4 0.64 6.90 13.6 0.34 2.40 4.42
Interval 4 0.76 18.8 21.6 0.73 8.25 14.1 0.44 2.59 4.02
Interval 5 1.40 24.6 27.0 0.69 7.71 12.9 0.60 3.29 4.88

does not mean that the animals did not use the useful parts of litter.
They may only remove less digestible parts. However, this needs further
investigation to measure the characteristics of excrements at different
intervals. In addition, litter type significantly influenced leaf con-
sumption, animal fresh weight, and excrement production (Table 2 and
Table S3). The interaction between litter type and animal classes and
between litter type and time was also significant. These results confirm
that these factors are important when investigating leaf consumption in
different soil organisms.
Average leaf consumption rates of 20–30 mg dry weight day−1 in-
dividual−1 were obtained when the isopod Armadillidium vulgare fed by
two grass species Bromus erectus and Festuca rubra (Pey et al., 2019). In
the latter study, the values of 5–35 mg dry weight day−1 individual−1
were reported for the millipede G. marginata fed on the same leaf spe-
cies for 7 days. This is consistent with our calculated values for the
Groups B and C of B. dubia feeding on alder and maple leaf litter
(Table 2). Excrement production rates were similar between these two
studies. Assimilation rate of up to 5 mg dry weight day−1 individual−1
for the Group C animals feeding on alder litter were completely in
agreement with the reported values for both isopod and millipede
species (Pey et al., 2019). For other animal classes in the present study,
negative assimilation rates were also obtained which may be the result
of failing to properly consume the litter by cockroaches in this short-
term laboratory experiment.
Calculated assimilation efficiency of 10–21% for B. dubia fed on A.
glutinosa leaves was consistent with the reported values for millipedes
(Tajovský, 1992; David and Gillon, 2002). However, higher values
(with large variation) were calculated for some other soil organisms.
For example, the values of 46–76% was obtained for the larva of the
bibionid fly Bibio pomonae (Frouz and Šustr, 1996), or the values of
42–78% for the woodlouse Porcellio dilatatus (Sousa et al., 1998), those
of 14–53% for four millipede and one isopod species (Ardestani et al.,
2019), and the values of 41–87% for the woodlouse Porcellionides
pruinosus (Loureiro et al., 2006). The assimilation efficiency values
slightly decreased after the first interval to 10–15% and remained
constant till the end of the 5th interval. The reason was not clear since
the leaf consumption, excrement production, and animal weight did not
change from interval 1 to 5 for the A. glutinosa leaf litter. It might in-
dicate that the leaves were readily consumed, but not easily assimilated
in the body which needs further investigation. Assimilation efficiencies
of 10–21% obtained in the present study for the Group C animals
feeding on alder litter are in agreement with the efficiency of conver-
sion of ingested food values of 14–30% calculated for B. dubia feeding
on diets with different protein and fat content (Oonincx et al., 2015). It
should be noted that the efficiency of conversion of ingested food values
in the latter study was calculated as (weight gained/weight of ingested
food) × 100 on a dry matter basis.
In conclusion, the results of this study showed that the cockroach
species B. dubia preferred A. glutinosa among other leaf litter species.
The cockroach B. dubia can be used as a model species in other ex-
perimental setups due to its easy culturing, high instar production rate,
and easy maintenance in the laboratory.
Declaration of competing interest
The authors declare that there is no conflict of interest.
Acknowledgements
The work was supported by the Ministry of Education, Youth and
Sports of the Czech Republic-MEYS (projects LM2015075, EF16_013/
0001782). This work has also been supported by Charles University
Research Centre program No. 204069. We thank the reviewers for their
constructive comments on the manuscript.
Appendix A. Supplementary data
Supplementary data to this article can be found online at https://
doi.org/10.1016/j.apsoil.2019.103460.
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