Environmental Entomology, XX(XX), 2020, 1–8

doi: 10.1093/ee/nvz167
Behavioral Ecology Research

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Saproxylic Cetoniidae (Coleoptera: Scarabaeoidea):
A ‘Females’ World’ or a Question of Dependence on
Deadwood?
Sandra Martínez-Pérez,1,3 Gerardo Sanchez-Rojas,2 Eduardo Galante,1 and
Estefanía Micó1

1
Centro Iberoamericano de la Biodiversidad CIBIO, Universidad de Alicante, Ctra de San Vicente del Raspeig s/n 03690 Alicante,
Spain, 2Laboratorio de Biología de la Conservación, Centro de Investigaciones Biológicas, Universidad Autónoma del Estado de
Hidalgo, Carretera Pachuca-Tulancingo km 4.5 s/n Colonia Carboneras Mineral de la Reforma, Hidalgo CP 42184, Pachuca, Mexico,
and 3Corresponding author, e-mail: sandra.martínez@ua.es

Subject Editor: William Morrison

Received 30 July 2019; Editorial decision 21 December 2019

Abstract
We explored the dependence of some Cetoniidae species on saproxylic environments and microhabitats in a
Mediterranean oak forest by analyzing species collected using different kinds of traps—log emergence, hollow
emergence, and interception traps—and the sex ratio of the species in each trap. Comparing the sex ratio of the
species collected via emergence versus interception was useful to unravel the degree of dependence on saproxylic
microhabitats. Among the species studied, Cetonia aurataeformis Curti, 1913 (Coleoptera: Cetoniidae) was the only
obligate tree hollow inhabitant. Special attention should thus be paid to the maintenance of tree hollows for the
species’ conservation in Mediterranean forests. A gradient of dependence on tree hollows was established from
the more dependent Protaetia (Potosia) cuprea (Fabricius, 1775) (Coleoptera: Cetoniidae) and Protaetia (Potosia)
opaca (Fabricius, 1787)  (Coleoptera: Cetoniidae)  to the less dependent Protaetia (Netocia) morio (Fabricius,
1781)  (Coleoptera: Cetoniidae). All the latter species can be considered facultatively dependent, to varying
degrees, on tree hollows. By contrast, the saproxylic affinity of Protaetia (Netocia) oblonga (Gory and Percheron,
1833) (Coleoptera: Cetoniidae), Tropinota squalida (Scopoli, 1783) (Coleoptera: Cetoniidae) and Oxythyrea funesta
(Poda, 1761)  (Coleoptera: Cetoniidae)  was doubtful. Generally, the sex ratio of the studied species was female-
biased. A possible explanation may be local male competition for females, suggesting the Cetoniinae is a female
world. However, the range of difference in the female-biased sex ratio among species suggests it is important to
explore other possible causes, such as differences in dispersal abilities.

Key words: Mediterranean oak forest, saproxylic environment, sex ratio, obligate tree hollow, facultative tree hollow

Saproxylic insect diversity is high in the Mediterranean region
(Ghaioule and Lieutier 2005, Baselga 2008, Micó et  al. 2013).
Coleoptera is widely represented among saproxylic species (Dajoz
and Álvarez 2001, Schlaghamersky 2003, Bouget et  al. 2008).
Among them, Cetoniidae (sensu Krikken 1984) are notable for the
functional importance of some of their species both as adults, acting
in some cases as pollinators, and as larvae, recycling plant matter
such as woody substrates (Micó et al. 2011, Sánchez-Galván et al.
2014, Sánchez et al. 2017).
Some species of European Cetoniidae seem to be invariably as-
sociated with woodlands, whereas others can be highly generalist
regarding their habitat (San Martín et al. 2001). Saproxylic spe-
cies can also differ with respect to the kind of woody microhabitat
that is exploited by larvae (e.g., logs, stumps, tree hollows) (Micó
et  al. 2011). In this way, many Trichiinae and Valginae larvae
have been found to feed mainly on rotten wood (Micó et  al.
2008), whereas most Cetoniinae (sensu Krikken 1984) are sap-
roxylophagous or saprophagous and commonly exploit tree hol-
lows or other woody substrates (Dubois 2009, Micó et al. 2011).
However, the extent to which the species depend on different
saproxylic microhabitats is poorly understood, and knowledge
regarding the facultative or obligate microhabitat preference of
one saproxylic species is typically extrapolated to the remaining
species of the same genus. Moreover, it is difficult to establish the
extent to which species classified as facultatively saproxylic are
reliant on deadwood environments. Knowledge of each species’
degree of dependence on different saproxylic environments could
help to detect the species most likely to be jeopardized by the

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2 Environmental Entomology, 2019, Vol. XX, No. XX
decline in woody substrates such as tree hollows (Ranius 2000,
Sebek et al. 2013, Micó 2018).
Different trapping methods commonly used for saproxylic spe-
cies can provide valuable information about the species’ habitat and
microhabitat requirements (Ranius and Jansson 2002, Hyvarinen
et al. 2006, Gouix and Brustel 2012, Quinto et al. 2013). Interception
traps, such as window traps (WT), do not allow differentiating all
the possible substrates that the species could be exploiting (i.e., cav-
ities, sap exudations, bark, dead branches, or deadwood in soil)
(Micó et al. 2013, Quinto et al. 2013, Micó 2018). In this way, WT
can reveal a habitat preference (e.g., a forest) but not a microhabitat
preference (Økland 1996). In contrast, emergence traps are con-
sidered to be a highly effective method for capturing species linked
to specific microhabitats. Such traps collect species shortly after their
emergence from immature stages and provide accurate informa-
tion about the species inhabiting these microhabitats (Quinto et al.
2013). Additionally, they are expected to provide real data on the
population sex ratio.
A bias in the real sex ratio in both emergence and intercep-
tion traps can also reveal different aspects of the species’ behavior.
A  biased sex ratio in interception traps can reveal a sex-biased
dispersal, with one sex emigrating more frequently and at in-
creased distances (Fontanillas et  al. 2004, Douadi et  al. 2007,
Gauffre et al. 2009); this approach can also be used to determine
oviposition preferences for tree microhabitats, when females are
captured more frequently through interception traps than through
emergence traps. A  bias in the real sex ratio in emergence traps
can also reveal differences in predation risk (Chiari et  al. 2013)
or interactions between relatives such as local mate competition
(Hamilton 1967).
Within this framework, our aim was to evaluate the depend-
ence of the Iberian Mediterranean cetonid species on saproxylic
habitats and microhabitats, estimating possible oviposition pref-
erences by comparing the species sex ratio obtained from both
emergence and interception traps. Our specific objectives in-
cluded: 1)  to assess the effectiveness of three sampling methods
for monitoring cetonid species: the WT, the hollow emergence trap
(HET) and the log emergence trap (LET); 2) to assess differences
in the species’ degrees of dependence on tree hollows; and 3)  to
infer the species behavior in saproxylic environments with respect
to its real sex ratio.
We hypothesized that tree hollows represent an important micro-
habitat for saproxylic species in Mediterranean regions because they
provide a stable abiotic environment and long-lasting resources
(Micó 2018). Moreover, as saproxylic cetonid females are expected
to fly near the trees to lay eggs in the tree hollows (Trizzino et al.
2014), a higher bias of females in WT compared to the real sex ratio
would predict the degree of a species’ dependence on tree hollows.
Materials and methods
Study Area
Data were obtained from surveys carried out in Mediterranean forests
located in 8 protected areas of the Iberian Peninsula: the biological
Reserve ‘Campanarios de Azaba’(Salamanca), Sierra de las Quilamas
Natural Area (Salamanca), El Rebollar Natural Area (Salamanca),
Las Batuecas-Sierra de Francia Natural Park (Salamanca), Sierra de
Bejar UNESCO Biosphere Reserve (Salamanca), Cabañeros National
Park (Ciudad Real), Sierra Espadán Natural Park (Castellón), and
Font Roja Natural Park (Alicante) (Table 1). All the studied areas are
mature forests dominated by Quercus species.
Beetle Sampling and Sex Identification
Adults of the Cetonid species were sampled using WT hanging
near tree trunks, HET and LET (272, 228, and 138 traps, respect-
ively). WT consisted of two transparent sheets (73  cm long and
42 cm wide) lying over a funnel and a collection container (Fig. 1)
(Bouget et al. 2008, Quinto et al. 2012); these traps were designed
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to capture active flying insects (Økland 1996) and are usually
referred to the sampling methods to study saproxylic assem-
blages (Økland 1996, Ranius and Jansson 2002). HET consisted
of a black acrylic mesh that completely sealed the tree hollow
and catcher pot attached to the mesh (Gouix and Brustel 2012,
Quinto et al. 2012) (Fig. 2). The LET was composed of a white
acrylic mesh encasing the log with a collecting bottle positioned
at the tip (modified from Owen 1989, Yee et  al. 2001) (Fig.  3).
Ethylene or propylene glycol was used as a preservative. Traps
were replaced once a month during one complete year in each
sampling site. Field studies were carried out from 2004 to 2018
(Table 1).
Males and females of each species were primarily distinguished
by morphological differences (Kašić-Lelo and Lelo 2007), and ne-
cessary corroborations were performed by extracting the external
genitalia using forceps.
Data Analysis
Data from all sampling sites were put together. The LET was fi-
nally excluded from the analysis due to the low number of cap-
tures (Results). A possible bias in sampling was dismissed because
both WT and HET were placed in all study sites. Moreover, the
species was not expected to present a different behavior depending
on the site because all the habitats are mature Quercus forests
in which all the microhabitats are present. In the same way, all
traps were active for a complete year on each site, preventing a
temporal bias.
Full inventory completeness was determined as the percentage
of observed species in relation to the number of species predicted by
the sample coverage estimator proposed by Chao and Jost (2012).
We utilized 100 replicate bootstrapping runs to estimate 95% con-
fidence intervals.
To evaluate differences in efficiency between each trap type, we
calculated sample-size-based rarefaction and extrapolation (R/E)
sampling curves (Chao and Jost 2012) for species richness (S) and
Hill number of order 1(1D)with INEXT software. 1D weighs each
species according to its frequency in the sample without favoring
either common or rare species (Jost 2006) and, therefore, can be in-
terpreted as the number of common (or typical) species in an assem-
blage. Significant differences were defined when curves were situated
outside the confidence limits of the other curves.
The difference in the abundance of individuals of each species
caught via both types of traps (WT and LET) was analyzed using
the Mann–Whitney U test. As multiple comparisons were made,
the alpha value was adjusted following the Bonferroni method
(Gotelli and Ellison 2004). The independence between species’ sex
and trap was calculated using 2  × 2 contingency tables, which
were statistically analyzed based on a likelihood ratio χ 2 test
(Zelen 1971). Where significant differences were found, standard-
ized residuals were analyzed to determine whether the observed
frequency differed significantly from what would be expected by
chance (Beasley and Schumacker 1995, Sharpe 2015). For each
species, the sexual proportion is expressed as female/males. All
calculations were performed using STATISTICA version 7.0
(Statsoft 2007).
Environmental Entomology, 2019, Vol. XX, No. XX 3

Table 1. Number of traps and dominant tree species in each studied area

Site Tree species sampled Number Number WT Number Coordinates Sampling year
HET LET

Campanarios Quercus rotundifolia Lam. 18 15 0 40° 29.769N 2010–2011

Quercus pyrenaica Willd. 10 12 0 6° 47.551W

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Quercus faginea Lam. 3 2 0
Quercus suber L. 0 1 0
Quilamas Q. pyrenaica 33 38 20 40° 35.642N 2012–2013
6° 03.201W 2014–2015
Rebollar Q. pyrenaica 18 39 39 40°21.10N 2014–2015
6°35.05W
Cabañeros Q. suber 9 37 0 39° 23.47N 2004–2005
Q. rotundifolia 32 18 0 4° 29.14W 2015–2016
Q. pyrenaica 22 14 0
Fraxinus angustifolia Vahl. 27 13 0
Q. faginea 8 8
Batuecas Q. rotundifolia 30 45 0 40° 27.291N 2012–2013
6° 08.088W
Espadán Q. suber 9 9 55 39° 52.00N 2015–2016
0°17.30O
Font Roja Q. rotundifolia 9 9 0 38°38.51N 2015–2016
0° 32.46W
Béjar Q. pyrenaica 0 12 24 40°25.26N 2017–2018
5°47.16O
Total 228 272 138

Campanarios (Biological Reserve Campanarios de Azaba), Quilamas (Quilamas Natural Area), Rebollar (The Rebollar Natural Área), Cabañeros (Cabañeros
National Park), Batuecas (Las Batuecas-Sierra de Francia Natural Park), Espadán (Sierra Espadán Natural Park), Font Roja (Font Roja Natural Park), and Bejar
(Sierra de Béjar UNESCO Biosphere Reserva).

Fig. 1. Window trap. Photo by D. Pérez.

Results
A total of 1,177 individuals belonging to 10 species and 5 genera of
Cetoniidae were collected. Only 2 species were collected via LET—
Oxythyrea funesta (Poda, 1761)  (Coleoptera: Cetoniidae)  and
Protaetia (Potosia) cuprea (Fabricius, 1775)  (Coleoptera:
Cetoniidae)—which represented 0.3% of the total number of indi-
viduals (Table 2); this type of trap was not very effective to monitor
Cetoniidae richness and diversity in the study areas and was thus
not included in the analyses. The sample coverage estimator showed
that our inventory completeness was 99.5% for WT and 99.6% for
HET. Cetoniidae diversity varied according to the WT and HET. WT
recorded the greatest richness and diversity (1D), followed by HET
(Fig. 4).
Fig. 2. Hollow emergence trap.
Species Abundance per Trap Type
The 3 most abundant species were Cetonia aurataeformis Curti
1913  (Coleoptera: Cetoniidae), P. (P.) cuprea and Protaetia
(Potosia) opaca (Fabricius, 1787)  (Coleoptera: Cetoniidae), repre-
senting 43%, 30% and 17% of individuals, respectively (Table 2).
Significant differences in the number of individuals between WT
and HET were found only for C.  aurataeformis (Mann–Whitney
Utests = 6053.5, P = 0.012), the latter being more frequent in HET
than WT (Fig. 5).
4 Environmental Entomology, 2019, Vol. XX, No. XX

Sex ratio

0
0

0
0
0
0
1
0
0
-

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LET

0
1
0
0
0
0
0
0
1
0
0
♀

0
0
3
0
2
0
0
0
0
1
0
Fig. 3. Log emergence trap.

Sex ratio

26.5
22.2

3.2
2.2
0.1
5

-
-
-
-
Species Sex Abundance per Trap Type
The number of females and males varied depending on the trap
type and species. Generally, the sex ratio was female-biased, with
the exception of Protaetia (Netocia) oblonga (Gory and Percheron,
1833)  (Coleoptera: Cetoniidae). Only C.  aurataeformis and P. (P.)

WT
cuprea showed sex-trap dependence (Table  3). The post hoc test

0
0
87
8
41
6
1
0
0
21
10
♂
showed that female C. aurataeformis were collected more than ex-
pected by chance in WT (z value = 2.97), whereas males were col-
lected more than expected by chance in HET (z value = 2.62), and
less than expected by chance in WT (z value = −4.74). In the case
Table 2. Species observed frequencies of females and males and sex ratio in different sampling methods

of females of P. (P.) cuprea, were collected more than expected by

2
1
536
133
5
1
6
46
1
210
131
♀

chance in WT (z value= 0.31), whereas males were collected more

than expected by chance in HET (z value= 1.65), and less than ex-
pected by chance in WT (z value = −1.27). Standardized residuals,
however, did not show significant values because the Z critical value
was not over ±1.96 (see MacDonald and Gardner 2000); moreover,
Sex ratio

Cramer’s V was 0.115, and therefore these residual values were not
1.8

3.5
3.5

9.8
significant. The greater a symmetries between male and female cap-
3
0
0
-

tures occurred in WT, where the most abundant species—C. auratae-

formis and P. (P.) cuprea—showed ratios of 22,2 ♀:1♂ and 26,5:1♂,
respectively (Table 2).
HET

0
0
154
130
0
0
4
2
0
12
6
♂

Discussion
This study provides a framework to assess saproxylic species’
degree of deadwood dependence using different kinds of traps
and the species sex ratio in each trap. The results demonstrated
that even congeneric species presented different degrees of de-
0
0
396
118
18
235
3
0
14
7
1
♀

pendence on deadwood. Although Cetoniidae populations were

generally female-biased, they were so to varying degrees, sug-
gesting possible differences in the species’ behavior and inter-
Sex ratio was calculated as females/males.

actions between relatives that could be conditioned by different

factors.
Our results showed that in the studied Mediterranean oak for-
Cetonia carthami aurataeformis

Protaetia (Eupotosia) mirifica

ests, deadwood on soil (logs) hosts less Cetonid species than other
Protaetia (Eupotosia) affinis

Protaetia (Netocia) oblonga

microhabitats such as tree hollows, WT and HET being more ef-

Protaetia (Potosia) cuprea
Protaetia (Netocia) morio

Protaetia (Potosia) opaca

fective at collecting and monitoring Cetoniidae species than LET.

WT is more effective at capturing this saproxylic family regarding
Tropinota squalida
Oxythyrea funesta

Trichius gallicus

both, species richness and abundance. Although tree hollows are

the predominant microhabitat for the development of the sap-
roxylic Cetoniinae species (sensu Krikken 1984), only two spe-
Species

cies—C. aurataeformis and Protaetia (Eupotosia) mirifica (Mulsant

Total

1842) (Coleoptera: Cetoniidae)—presented more individuals in HET

Environmental Entomology, 2019, Vol. XX, No. XX 5

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Fig. 5. Boxplots showing species abundance for each trap with the interquartile range boxes, median and outliers symbols. Name of species in bold indicates
significant differences in Mann–Whitney U test after Bonferroni correction (P < 0.025).

Fig. 4. Sample-size-based rarefaction and extrapolation (R/E) sampling curves for the Diversity metrics (S: species richness, 1D: Hill numbers of order 1) of each
sampling method. Shaded areas correspond to 95% confidence intervals.

Table 3. Pearson’s χ 2 and significance from the 2 × 2 contingency table test of independence were carried out on each species to test the
null hypothesis that their distributions in terms of sex were independent of trap type (P < 0.05)

Species Pearson’s χ 2 Df Prob. Level

Cetonia carthami aurataeformis 50.05 1 =0.0000

Oxythyrea funesta 0.56 1 0.7077a
Protaetia (Eupotosia) mirifica 1.6 1 0.5271a
Protaetia (Netocia) morio 0.03 1 0.8588
Protaetia (Netocia) oblonga 4.64 1 0.1736a
Protaetia(Potosia) cuprea 3.67 1 0.0311
Protaetia(Potosia) opaca 0.02 1 0.9005

a
Yate’s correction was applied to the χ 2 test when the expected frequency value was less than 5. Significant values in bold.

than in WT, whereas the remaining species were all more frequent
in WT than in HET (Table 2). This result indicates that both species
could be hollow-dependent. However, only in the case of C.  aura-
taeformis was this difference significant, despite P.(E.) mirifica being
considered an obligate saproxylic associated with tree hollows (Tassi
et  al. 2004, Micó et  al. 2010). The rarity of P.(E.) mirifica in all
of its range of distribution (Agoiz-Bustamante and Caselles 2009,
Micó et al. 2010) and the low number of specimens collected in this
study (24 specimens) may explain the lack of significance here. In
contrast, C. aurataeformis is common in the Mediterranean region
of the Iberian Peninsula (Micó and Galante 2002) (over 500 indi-
viduals were recorded in this study) and has been marked out as an
ecosystem engineer in Mediterranean oak tree hollows due to the
activity of its larvae in the wood mold substrate (Micó et al. 2015,
Sánchez et al. 2017).
The analysis of the sex ratio of each Cetonid species according
to each trap type helped to determine the species’ dependence on
saproxylic microhabitat but also the species’ behavior. It is expected
6 Environmental Entomology, 2019, Vol. XX, No. XX
that the sex ratio obtained from the emergence traps will provide a
good estimation of the real population sex ratio because such traps
collect the specimens that emerge after hatching—a single female of
Cetoniinae species can lay over approximately 40 eggs (Jones 1990).
Our data showed that, in all cases, the HET collected more females
than males; in fact, the sex-ratio values of the species ranged from
1.8 in C. aurataeformis to 9.8 in P. (P.) cuprea (Table 2). Many fac-
tors could cause a biased sex ratio in natural populations, such as
sex-biased predation, temperature influence or haplodiploid sex de-
termination. Acharya (1995) provided evidence of sex-biased pre-
dation on moths as a result of differences in the behavior of the
adults of each sex, and the same sex-biased predation was observed
in the beetle Melolontha melolontha (Linnaeus, 1758) (Coleoptera:
Melolonthidae) (Jones 1990). Other studies have also demonstrated
higher male mortality due to predation or parasitism on adults (e.g.,
Cade 1975, Burk 1982, Gwynne 1987, Sakaluk 1990, Rehfeldt
1992). However, there is no evidence that a selective predation of
larvae in insects could explain the biased sex ratio of the Cetonids
hatching in tree hollows. Moreover, examples of alterations in pro-
geny sex caused by biotic and abiotic agents such as temperature
have been previously reported in the case of some insects (Brown
and Chandra 1977, Nigro et  al. 2007). Empirical data under con-
trolled temperatures are required to test this hypothesis; this hy-
pothesis seems unlikely in our case; however, if we consider that
temperature and moisture are generally more constant and stable
in tree hollows compared to other environmental conditions (Park
and Auerbach 1954). In other cases, some species, such as Ambrosia
beetles (Xyleborini) (Coleoptera: Curculionidae) present a haplodip-
loid mating system of sex determination (Peer and Taborsky 2005).
In these mating systems, females store sperm and control the sex
of their offspring (Cole 1981, Suzuki et  al. 1984, van Dijken and
Waage 1987). Fertilized eggs develop into females (diploid), while
unfertilized eggs into males (haplodiploid) (Crozier 1975). However,
haplodiploid sex determination has not hitherto been found to exist
in the Cetonid species.
Cetonids seem to present a female-favored world, the causes of
which should be determined, particularly in terms of post-reproduc-
tive behavior. In general, a 1:1 sex ratio can readily be explained in
terms of selection acting on the individual (Fisher 1999), whereas
a deviation of this ratio can occur when: 1) brothers compete with
each other (i.e., local mate competition and local resource competi-
tion); and 2) individuals of one sex have an increased positive fitness
effect on relatives (local resource enhancement) (Hamilton 1967,
Clark 1978, Taylor 1981, Schwarz 1988, Frank 1998). In polygynous
mating systems such as the case of Cetonids beetles, males are there-
fore more likely to exhibit strong local mate competition (Perrin and
Mazalov 2000). The individuals are thus selected to produce a lower
proportion of the competitive sex (West et al. 2005). Consequently,
an increase in the number of females reduces the competition of the
males by the females (Davies et  al. 2012), and the degree of bias
would depend on the extent of local mate competition (Werren
1980). This could be the case of the studied Cetoniinae, where the
degree of bias between males and females varied greatly between
species, with the high sex ratio values of P. (P.) cuprea (9.8) being
particularly remarkable: a situation which could reflect an increased
level of competition in this species. A  high level of competition is
expected in dispersal-limited species (Davies et al. 2012). However,
no direct assessments of the dispersal of P. (P.) cuprea versus other
species with lower sex ratio values (such as C. aurataeformis (1.8))
have been performed to corroborate these hypotheses.
We detected that Cetonia aurataeformis can be highlighted
as a hollow-dependent species, not only because of its statistical
dependence on HET, but also due to the results of sex depend-
ence on WT (Table  3). In contrast, its congeneric Cetonia aurata
(Linnaeus, 1761)  (Coleoptera: Cetoniidae)  is considered a faculta-
tive tree hollow inhabitant because its larvae have also been found in
ant nests (Janssens 1960, Micó 2001) and compost (Paulian 1959).
As expected, C. aurataeformis females were captured more than ex-
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pected by random using WT, whereas males were captured more
than expected by chance in HET, but less than expected by chance
in WT. This result may reflect a post-reproductive behavior, where
males disperse on flowers, leaving the reproductive sites, whereas fe-
males usually remain near the tree hollows for oviposition (Trizzino
et al. 2014). Alternatively, the lower values of number of males in
WT (Table 2) could also be a consequence of an uneven predation
risk formales (Chiari 2011, Chiari et  al. 2013). However, the sub-
stantial deviation of the sex ratio, that ranged from 1.8 (natural sex
ratio) to 22.2 in WT could not be explained only by a biased preda-
tion risk to males. Moreover, the lack of males among the trapped
individuals in WT may reflect a difference in dispersal behavior be-
tween sexes. Local competition among same-sex relatives could also
provide an incentive for the adaptative adjustment of the dispersal
rate (Hamilton and May 1977, Frank 1986, Taylor 1988, Motro
1991). Consequently, females may be more abundant or more active
fliers (Watson 2003) or may have a distinct dispersal mode, and their
capturing via WT may hence be easier (Bouget et al. 2015). However,
the results of co-evolution of the sex ratio and sex-specific dispersal
are doubtful (Wild and Taylor 2004).
The species P. (P.) cuprea demonstrated sex dependence ac-
cording to the trap based on Chi-squared analyses. In contrast, the
Z critical value and Cramer’s V showed independence between sex
and trap (see Results). Thus, its obligate dependence with respect to
tree hollows is questionable. Despite this issue, the results illustrated
a clear trend indicating that the females were captured more in WT
than in HET. If we accept the hypothesis that a higher frequency of
females in WT is related to females’ behavior of searching for tree
hollows for possible oviposition locations, we ought to consider it to
be a saproxylic species. The same occurs with the other species be-
longing to the Potosia subgenus, P. (P) opaca. Moreover, both species
should be considered as facultative inhabitants of tree hollows be-
cause there were no significant differences between their frequencies
in HET and WT. According to the literature, both species have been
associated with tree hollows but also with the trunks of living palms
(Micó 2001, Tauzin 2007).
Both species belonging to the Netocia subgenus—Protaetia
(Netocia) morio (Fabricius, 1781) (Coleoptera: Cetoniidae) and P.
(N.) oblonga—were more abundant in WT than in HET (Table 2).
However, differences in that frequency were not found in either
case. The number of captures was low for both species, particularly
for P. (N) oblonga despite both species being frequently found in
the study area (Pers. Obs). The low number of individuals of P. (N.)
morio in HET revealed that tree hollows are poorly exploited by
this species, the larvae of which have been associated with wounded
parts of living trees (Schaefer 1958) and organic matter such as
manure (Micó 2001). In contrast, the captures of P. (N.) morio in
WT do not seem to be casual occurrences because this species is a
possible facultative saproxylic. However, the amount captured was
lower than that of the Potosia subgenus. On the other hand, the
captures of P. (N.) oblonga in our traps were still more haphazard
than in the case of P. (N.) morio. Moreover, the sex ratio of P. (N.)
oblonga in WT was male-biased, hence females do not seem to be
attracted by woody microhabitats. The literature has reported the
presence of this species in debris of vegetal matter accumulated in
ant nests and in rabbit latrines (Micó 2001). The same has been
Environmental Entomology, 2019, Vol. XX, No. XX
observed for Tropinota squalida (Scopoli, 1783)  (Coleoptera:
Cetoniidae) and O. funesta—both are abundant in clearing forests
of the Iberian Peninsula (Micó 2001), but few were collected in our
traps. The larval development of these species has been found in
decaying vegetative matter and manure (Balachowsky 1962, Micó
2001, Molina 2001). We, therefore, conclude that the saproxylic
nature of the P. (N.) oblonga, T.  squalida and O.  funesta species
is unlikely. Finally, only one individual of Trichius gallicus Dejean,
1821 (Coleoptera: Cetoniidae) was collected in our study; however,
contrary to the species mentioned above, the latter could be ex-
plained by the species’ rarity in the studied area. Its larval develop-
ment is associated with old trunks and different woody substrates
(Paulian 1959).
We also concluded that the analysis of the species’ sex ratio
based on emergence and interception traps helped to reveal the
degree of dependence on saproxylic microhabitats, such as tree
hollows, and demonstrated that microhabitat preferences of spe-
cies could not be extrapolated to other related species. Moreover,
the analysis of the Cetoniidae species population sex ratio re-
vealed possible local male competition for females, though to a
varying degree according to the species. Direct assessments of dif-
ferent species’ dispersal ability are needed to corroborate the hy-
pothesis of limited dispersal determining high levels of local mate
competition.
Acknowledgments
Financial support was provided by the ‘Ministerio de Economía, Industria
y Competitividad’ (CGL2016-78181-R). This research is part of Sandra
Martínez Pérez’s PhD studies granted by ’Ministerio de Economía, Industria y
Competitividad’ (BES-2017-080278).
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