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Biotropica. 2022;00:1–15.
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1wileyonlinelibrary.com/journal/btp
1 | INTRODUCTION
Global biodiversity is experiencing a major crisis (Barnosky
et al. Barnosky et al., 2011; Ceballos et al., 2015), where habitat
fragmentation and loss are drivers in the reduction of population
and extinction of species (Brook et al., 2008; Brooks et al., 2002;
Hoekstra et al., 2005). Regions with significant levels of biodiver-
sity, which are increasingly threatened, deserve special attention
Received: 13 August 2021
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Revised: 26 January 2022
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Accepted: 12 July 2022
DOI: 10.1111/btp.13151
ORIGINAL ARTICLE
Crickets as indicators of ecological succession in tropical
systems, New Caledonia
Jeremy Anso1,2 | Amandine Gasc1 | Edouard Bourguet1 |
Laure Desutter- Grandcolas2 | Hervé Jourdan1
This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium,
provided the original work is properly cited.
© 2022 The Authors. Biotropica published by Wiley Periodicals LLC on behalf of Association for Tropical Biology and Conservation.
1CNRS, IRD, IMBE, BPA5, Aix Marseille
Univ, Avignon Univ, Nouméa Cedex, New
Caledonia
2CNRS, UMPC, EPHE, UA , Institut
de Systématique, Evolution, Muséum
National d'Histoire Naturelle, Sorbonne
Universités, Paris Cedex 05, France
Correspondence
Amandine Gasc, CNRS, IRD, IMBE, BPA5,
Aix Marseille Univ, Avignon Univ, 98848
Nouméa Cedex, New Caledonia.
Email: amandine.gasc@ird.fr
Funding information
Action Transversale du Muséum (ATM):
Biodiversité actuelle et fossile. Crises,
stress, restauration set panchronisme
lemessage systématique, Grant/Award
Number: Barcode; Grand Observatoire du
Pacifique Sud: AAP GOPS 2013, Grant/
Award Number: Bioacoustique des grillons
de Nouvelle- Calédonie; Agence Nationale
de la Recherche (ANR) through the ERA-
Net BiodivERsA Project, Grant/Award
Number: FFII, JE 288/7- 1; Government
of New Caledonia, Grant/Award Number:
Bourse d’encouragement à la recherche
universit
Associate Editor: Eleanor Slade
Handling Editor: Alexander Smith
Abstract
Crickets (Ensifera, Grylloidea) are not commonly used as ecological indicators in con-
trary to other Orthoptera (e.g., grasshoppers and katydids). However, they are sensi-
tive to environmental changes and abundant in tropical regions. To evaluate whether
crickets are relevant bioindicators of tropical ecosystems, we investigated cricket as-
semblages along a tropical ecological gradient. We collected crickets during both day
and night in southern New Caledonia for three stages of ecological succession: open
shrubland, preforest, and forest. Simultaneously, we measured several environmental
variables, such as temperature and relative humidity, at each sampling site. Cricket
species assemblages showed a clear response to ecological succession. The highest
and lowest species richness and abundances of individuals were, respectively, found
in forest and shrubland, with species specialized in each ecological stage revealing
the conservation value of each of these stages. Similar results were found when con-
sidering only the part of cricket communities with the ability to acoustically commu-
nicate. This work is part of a larger research program about Neocaledonian crickets
and contributes to support the use of acoustic approaches to monitor tropical en-
vironments. In conclusion, these findings highlight the potential value of crickets as
an environmental indicator in tropical ecosystems. The results also contribute to the
discussion of the intrinsic conservational value of shrublands in New Caledonia and
similar ecotypes.
Abstract in French is available with online material.
KEYWORDS
bioindicator, community ecology, conservation, crickets, ecoacoustics, ecological succession,
taxonomic inventory, tropics
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ANSO et al.
for conservation and restoration (Dawson et al., 2014; Mittermeier
et al., 2011; Myers et al., 2000). Conser vation programs often rely on
biodiversity monitoring using diversity indices (Barlow et al., 2007).
Two of the most common diversity indices are species richness and
evenness (Ricotta, 2005). Such diversity indices are dependent upon
species inventories, which are time- consuming (Lawton et al., 1998),
expensive, and require highly skilled specialists (Levrel et al., 2010).
The United Nations has therefore strongly encouraged the scien-
tific community to develop biodiversity indicators for biodiversity
conservation (Strategic Plan for Biodiversity 2011– 2020, United
Nations, 2020, IPBES, 2019).
Bioindic ators are generally used to assess environmental change,
to monitor specific disturbances, or to estimate taxonomic diversity
(Gerlach et al., 2013; McGeoch et al., 2011). Such indicators serve
several applications, including assessing the condition of the envi-
ronment (Niemi & McDonald, 2004), following ecosystem resto-
ration programs (Alignan et al., 2014), or prioritizing the management
of habitats (Fartmann et al., 2012). The challenge in selecting taxa
as bioindicators in complex habitats, such as tropical forests, is that
they require sufficient knowledge about their distributions and nat-
ural fluctuations, with a fairly resolved taxonomic knowledge. This
is frequently absent in tropical environments (Moritz et al., 2001).
Among invertebrates, Orthoptera (spanning greater than 28,000
valid species; Cigliano et al., 2020) are widely used to assess ecolog-
ical or environmental changes (Gerlach et al., 2013). They are a key
resource for upper food web species, such as lizards, spiders, and
birds (Bor ges et al., 2013; Prete lli et al., 2014; Shea et al., 2009; Taylor
et al., 2012) and play an important role in recycling organic matter
(Prather et al., 2013). Related to ecological changes, Orthoptera spe-
cies richness and composition have been negatively associated with
mowing and grazing (Rada et al., 2014; Rambo & Faeth, 1999), and
they are regularly used to assess the success of restoration programs
(Alignan et al., 2014, 2018; Hugel, 2012; Rácz et al., 2013). Among
Orthoptera, grasshoppers and katydids are commonly use d to assess
environmental change and restoration in temperate regions where
they are well- diversified and can be easily identified, especially in
open habitats (Fartmann et al., 2012; Marini et al., 2009; Schirmel
et al., 2011). Grasshoppers were also found to be markers for forest
succession in Western Amazonia (Amédégnato, 1997; Amédégnato
& Descamps, 1980).
Crickets (Orthoptera, Grylloidea) are the other large group of
Orthoptera, with approximately 6000 species (Cigliano et al., 2020).
Crickets are studied as m odels in etholog y, neurobiology, physiology,
and bioaco ustics (G erhardt & Hub er, 2002; Kulkarni & E xtavour, 2019)
but receive less focus in ecology. Most ecological studies assessing
restoration or habitat management using Orthoptera assemblages
have not separated grasshoppers, katydids, and crickets. In partic-
ular, only a few have considered the cricket community alone (e.g.,
Hoffmann et al., 2002; Szinwelski et al., 2012). While underexplored,
crickets as potential bioindicators in the endangered tropics hold
significant potential because of their high level of diversification
and endemism in the tropics (Cigliano et al., 2020), abundance, and
local ecological specializations (Desutter- Grandcolas, 1995, 1997),
and use of recognizable acoustic signals (Anso, Jourdan, & Desutter-
Grandcolas, 2016; Diwakar & Balakrishnan, 2007) when using pas-
sive acoustics for monitoring.
The goal of the present study was to evaluate whether crick-
ets, at the species and community level, might be qualified as
relevant bioindicators of tropical ecosystems. The effect of vege-
tation and bioclimatic attributes along shrubland to forest gradient
on cricket assemblages was assessed. Research was performed in
New Caledonia, which was advantageous for two reasons. First, a
detailed description of the cricket species living in the area of in-
terest was needed (Anso, Jourdan, & Desutter- Grandcolas, 2016;
Desutter- Grandcolas et al., 2016), and second, New Caledonia is
representative of various tropical ecosystems. Crickets represent a
well- diversified part of the original and disharmonic insect fauna of
New Caledonia (Grandcolas et al., 2008), with 40 genera and more
than 180 species recorded today in the archipelago. A total of 19 of
the genera and more than 90% of the species are endemic to the
territor y (Desutter- Grandcolas et al., 2016). Guessestimate of the
total richness of the cricket fauna, based on the putative pattern of
species assemblage in New Caledonia (Anso, Jourdan, & Desutter-
Grandcolas, 2016), leads to the hypothesis that the actual number
of crickets in New Caledonia should be much more important, es-
pecially because of the high microendemism that characterizes this
fauna (Grandcolas et al., 2008; Nattier et al., 2011). Nevertheless,
the cricket fauna living in our research area is well described (Anso,
Jourdan, & Desutter- Grandcolas, 2016; Desutter- Grandcolas et al.,
Desutter- grandcolas et al., 2016). New Caledonia is additionally a
biodiversity hotspot (Myers et al., 2000), with a high endemism rate
(Grandcolas et al., 2008), and experiences multiple disturbances, in-
cluding species invasion, and habitat loss (Ibanez et al., 2019; Jaffré,
Bouchet, & Veillon, 1998; Jaffré, Rigault, & Dagostini, 1998; Pellens
& Grandcolas, 2009). Therefore, furthering ecological knowledge in
this context holds high value for local conservation plans by helping
to evaluate forest restoration programs, which are of high impor-
tance due to mining activity in New Caledonia (Losfeld et al., 2014).
In this study, a large taxonomic inventory was conducted during
both day and night to measure the overall abundance, species rich-
ness, composition, and structure of the cricket communities. Data
were collected in three ecological stages defined by a gradient of
microclimate and vegetation characteristics (forest, preforest, and
shrubland) in the south of the New Caledonian archipelago's main
island, on ultramafic soil. New Caledonian ecosystems are not nat-
urally fire- prone ecosystems (McCoy et al., 1999; Stevenson et al.,
2001), but fire is promoted by human populations since their arrival.
Ultramafic substrates are characterized by low nutriment and slow
vegetation growth (Isnard et al., 2016; Pillon et al., 2021), several
decades according to McCoy et al. (1999) and Pillon et al. (2021).
On these soil compartments, the succession is marked by the tran-
sition from open shrubland to arbustive shrubland to preforest to
forest stages. Each stage shows clear- cut vegetation formation in
terms of stem density, litter cover, and stratification (Jaffré, Rigault,
& Dagostini, 1998; McCoy et al., 1999). The context of succession on
ultramafic soils is similar to South Africa and Southwest Australia,
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ANSO et al.
with shrubland that resembles the South African fynbos or the
Southwest Australian kwongankaroo (Pillon et al., 2021).
Analyzing these data, the following questions were addressed:
(1) Are assemblages of crickets following the ecological succes-
sion? (2) Are any specific cricket species an indicator of the eco-
logical stages? And (3) is acoustic monitoring a suitable method for
the evaluation of the ecological stages? This research is part of a
larger research program conducted in New Caledonia focusing on
biodiversity monitoring using taxonomic and acoustic methods. The
results will thus be interpreted and discussed within the larger find-
ings of this program. In particular, a section of the discussion will be
dedicated to Passive Acoustic Monitoring (PAM) (Sugai et al., 2019)
for extracting information about the cricket community indicating
their tropical habitat.
2 | METHODS
2.1 | Study sites
Cricket communities were studied in the southern region of Grande
Terre, the main island of New Caledonia, an archipelago located in
the Southwest Pacific Ocean. Sites were selected along the same
geological ultramafic substrate at low altitude (231 ± 66 m; Table 1).
To study the ecological succession of cricket fauna, three vegetation
stages— forest, preforest, and shrubland— were considered. Four
sites were chosen within each stage for a total of twelve sites.
Forests on the island consist of trees with large stems, a deep
leaf litter entirely covering the ground, a closed canopy, and a max-
imum vegetation height of 30 m. The tree layer is composed of di-
verse, dominant families, such as Araucariaceae (Agathis sp. and
Araucaria sp.), Sapotaceae (Iteiluma sp., Planchonella sp., and Pichonia
sp.), Myrtaceae (Syzigium sp. and Eugeania sp.), and Proteaceae
(Kermadecia sp. and Macadamia sp.). Several unidentified species of
palm trees, pandanus, and ferns dominate the shrub and herbaceous
layers of the forest.
Preforest can be as dense as the forest but is composed of dif-
ferent plant species, characterized by a greater canopy openness
and lower diameter tree stems. As in the forest, the bare ground
is covered. The tree layer of preforest includes an assemblage of
Styphelia cymbulae, Hibbertia lucens, Alphitonia neaocaledonica, and
Gymnostoma deplancheanum, while the herbaceous layer is domi-
nated by Lepidosperma perteres.
Shrubland is characterized by a high bare ground expanse and an
absence of a continuous tree layer. More specifically, shrublands en-
compass small patches of vegetation with a thin leaf litter, separated
by bare ground. The maximum vegetation height is around 3 m.
Shrublands were dominated by shrubs but also include herbaceous
species such as Sannantha sp., Eugenia sp., Alphitonia neocaledonica,
Hibbertia lucens, and H. pancheri.
2.2 | Cricket sampling
Crickets were collected between November 2013 and April 2014. In
each of the twelve sites selected (see section 2.1), two squared par-
cels with 10 m sides were delimited. Sampling was performed under
clear sky meteorological conditions from hours 0900 to 1700 by
day, and 1900 to 0000 by night. On each parcel, crickets were col-
lected using the same collection method of ten 30- minute sessions,
with five total collections by day and five by night. Crickets were
located using both sight and song cues in the field only based on
human perceptions, which is the most appropriate sampling method
for crickets (Touroult et al., 2021). Another common method to
collect insects seems inefficient for crickets (Touroult et al., 2021)
such as (1) light trap that rarely attracts crickets, (2) pitfall that only
captures specimens foraging in the leaf litter, (3) fogging, a method
previously tried in New Caledonia leading to collecting mostly juve-
niles that could not be identified (LDG, pers. obs), or (4) interception
trap that captures only flying insects, very rarely crickets. For each
individual, activity (e.g., singing, eating, mating, resting) and micro-
habitat (e.g., trunk, leaf litter, vegetation, rocks, height from ground)
were noted before the capture. Collected specimens were kept
for identification, as most revealed new species to science (Anso,
Jourdan, & Desutter- Grandcolas, 2016). Specimens are deposited
in the Museum national d'Histoire naturelle de Paris with a refer-
ence collection in the Nouméa IRD center in New Caledonia. Species
identifications were performed in the laboratory by comparison with
the specimens in the deposited collections and using the classifica-
tion derived from the extensive molecular phylogeny of crickets
(Chintauan- Marquier et al., 2015). Based on previous taxonomic
descriptions including acoustic production (i.e., Anso, Jourdan, &
Desutter- Grandcolas, 2016; Desutter- grandcolas et al., 2016), we
were able to attribute to each species a presence or absence of
singing abilities. Of the 1030 individuals collected, 54 were dis-
carded from the analyses. These specimens were not identifiable at
the species level because of their destruction due to unfavorable
TABLE 1 Location and GPS information of the 12 sites. The
coordinate reference system used was WGS84
Location Habitat
Longitude
East
Latitude
South
Forêt Nord Shrubland 166.93501 22.32277
Grand Kaori Shrubland 166.89436 22.28460
Chute de la Madelaine Shrubland 166.85268 22.23568
Rivière Blanche Shrubland 166.70796 22.13625
Forêt Nord Preforest 166.93134 22.32259
Grand Kaori Preforest 166.89383 22.28000
Pépinière Preforest 166.96355 22.27103
Rivière Blanche Preforest 166.68033 22.15280
Grand Kaori Forest 166. 89674 22.28535
Mouirange Forest 166.68086 22.20416
Pic du Pin Forest 166.82715 22.24680
Rivière Blanche Forest 166.686 43 22.15142
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ANSO et al.
conservation conditions (56%) or from the absence of morphological
information for crickets in the juvenile development stage (44%). A
table of the cricket inventory (Table S1) and the list of species full
names (Table S3) are provided in the supplementary information.
2.3 | Environmental variables
During cricket sampling, environmental attributes were measured in
each parcel, including the percentage of bare ground, the percent-
age of three vegetation layers (herbaceous, shrub, and tree), the
number and diameter (dbh) of stems, the vegetation height, and the
plant species richness. Canopy openness was extracted from photos
taken with a 180° hemispherical (fisheye) lens in each parcel corner
and in its center. From the photos, canopy openness was then cal-
culated using the Gap Light Analyzer software (version 2.0, Simon
Fraser University, Cary Institute of Ecosystem Studies) by estimating
the percentage of light in the forest overstorey (Frazer et al., 1999).
Daily temperature (°C) and relative humidity (%) were recorded
every 30 minutes for 15 days at a 50- cm height above the ground
using waterproof thermo- hygrometer sensors (HOBO U23 Pro v2;
HOBOware software). A table of the environmental variables is
available in the Supplementary Information (Table S2).
2.4 | Statistical analysis
Two datasets were created for statistical analyses. The first included
the environmental attributes averaged per site, while the second
described the cricket assemblages, defined here as the number of
individuals per species averaged per site. All statistical analyses were
performed using R (software; R Core Team, 2019), with the type I
error set at 5%.
To evaluate differences between ecological stages a Kruskal–
Wallis test followed by the post hoc Conover test for multiple
comparisons with the Bonferroni correction was applied (Conover
& Iman, 1979). This test was run separately on species richness
(number of species), overall abundance (total number of individual
crickets), and environmental variables. This test was computed using
the function posthoc.kruskal.conover.test of the “PMCMR” R package
(Pohlert, 2014).
To visualize the differences in cricket assemblages along the eco-
logical succession, a Nonmetric Multidimensional Scaling (NMDS)
was performed on cricket assemblages across sampling sites
(Legendre & Legendre, 2012; Minchin, 1987). The Bray– Curtis dis-
tances of cricket assemblage were calculated between sites. NMDS
analysis with 1000 iterations and two dimensions was then per-
formed on each of these distance matrices. To explain each of the
four NMDS observations, the environmental variables were fitted
into ordination, and their significance was assessed with a permu-
tation test (n = 999 permutations). The percentage of variance in
cricket assemblages that can be explained by the ecological stages
was assessed with a permutation test (n = 999 permutations) applied
to the same Bray– Curtis distance matrix. This analysis was similarly
conducted on cricket assemblages and summarized in four ways: (1)
cricket species abundances, (2) stridulatory cricket species abun-
dances, (3) presence or absence of cricket species, and (4) presence
or absence of stridulator y cricket species. The function metaMDS
was used to perform the Bray– Curtis distance and the NMDS anal-
yses, while the permutation test was performed using the function
envfit for environmental data. Lastly, the permutation test applied
on the distance matrix of cricket assemblages was performed using
the function adonis. These three functions are from the “vegan” R
package (Oksanen et al., 2019). The function s.class from the “ade4”
R package was used for the graphical scatter diagram representation
of the ecological stages in the NMDS space (Dray & Dufour, 2007).
The beta diversity (β) of cricket assemblages measured as the
Sorensen dissimilarity was partitioned in the turnover (βSIM) and
the nestedness (βNES) components, being, respectively, the Simpson
dissimilarity and the nestedness- resultant fraction of Sorensen dis-
similarity. Abundances were first reduced to a simple presence or
absence of the cricket species at each site. Then, their overall beta
diversity was partitioned using the function beta.multi from the
“betapart” R package (Baselga et al., 2012; Baselga & Orme, 2012).
The same analysis was conducted on the presence or absence of the
cricket species for each ecological stage.
To evaluate the most relevant species between paired ecologi-
cal stages, a similarity percentage analysis (SIMPER) was performed
on both the abundances per cricket species and the presence or
absence of cricket species (Clarke, 1993; Warton et al., 2012). The
Bray– Curtis dissimilarity was decomposed in species contribution.
This analysis was performed using the function simper to the “vegan”
R package (Oksanen et al., 2019). The R code used to compile, manip-
ulate, and analyze the data is available as Appendix S4 and h t tps ://
github.com/agasc/ Anso- et- al- 2022- Biotr opica.
3 | RESULTS
3.1 | Cricket species along the succession
A total of 976 specimens, 30 species, and 16 genera were collected,
belonging to four families (Gryllidae, Phalangopsidae, Trigonidiidae,
and Mogoplistidae) and six subfamilies (Table 2). The most abundant
species were Bullita fusca (Gorochov, 1986) and Agnotecous azurensis
Desutter- Grandcolas, 2006 in both forest (40% and 31%, respec-
tively) and preforest stages (35% and 25%, respectively). In shrub-
lands, Koghiella flammea Anso & Desutter- Grandcolas, 2016 was the
most abundant species (88%). Species richness increased gradually
along the ecological succession, from the lowest number of species
in shrubland (7 species), followed by observed increases in preforest
(15 species) and forest (20 species) (Figure 1). Each ecological stage
was additionally found to harbor exclusive species. Forest demon-
strated the highest ratio of unique species (60% including 12 spe-
cies) followed by shrublands (43% including 3 species) and preforest
(14% including 2 species). Forest and preforest shared eight species,
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ANSO et al.
TABLE 2 List of cricket species from the taxonomic inventory in alphabetical order. The symbol a indicates the number of specimens
collected
Family Subfamily Species Habitat (a)Microhabitat Stridulatory Capture time (a)
Gryllidae Eneopterinae Agnotecous azurensis Fo (143) / Pr (137) Leaf litter yes Day (102) / Night
(178)
Gryllidae Eneopterinae Agnotecous clarus Fo (1) / Pr (34) Leaf litter yes Day (7) / Night
(28)
Gryllidae Eneopterinae Agnotecous meridionalisbFo (16) Leaf litter yes Day (9) / Night (7)
Gryllidae Eneopterinae Pixibinthus sonicus Pr (16) / Sh (5) Leaf litter yes Day (11) / Night
(10)
Gryllidae Gryllinae Notosciobia affinis
paranola c
Pr (1) Leaf litt er/
burrows
yes Night (1)
Gryllidae Gryllinae Notosciobia minoris Fo (1) / Pr (1) Leaf litt er/
burrows
yes Day (1) / Night (2)
Gryllidae Gryllinae Notosciobia sp1bFo (2) Leaf litt er/
burrows
yes Day (2)
Gryllidae Podoscirtinae Adenopterus crouensisdSh (1) Understory no Night (1)
Gryllidae Podoscirtinae Adenopterus meridionalisbFo (1) Understory no Nig ht (1)
Gryllidae Podoscirtinae Calscirtus amoabFo (1) Understory ?Night (1)
Gryllidae Podoscirtinae Calscirtus magnusbFo (1) Canopy yes Night (1)
Gryllidae Podoscirtinae Matuanus affinis mirabilisbFo (2) Tr unk yes Night (2)
Gryllidae Podoscirtinae Pixipterus punctulatusdSh (1) Understory no Night (1)
Mogoplistidae Mogoplistinae Mogoplistidae sp1 Pr (2) / Sh (1) Leaf litte r/tru nk yes Night (3)
Mogoplistidae Mogoplistinae Mogoplistidae sp2 Pr (4) / Sh (2) Leaf litte r/tru nk/
understory
yes Day (1) / Night (5)
Mogoplistidae Mogoplistinae Mogoplistidae sp3dSh (6) Lea f li tter/
understory
yes Day (2) /Night (4)
Mogoplistidae Mogoplistinae Mogoplistidae sp7bFo (9) Understory yes Day (1) / Night (8)
Phalangopsidae Paragryllinae Protathra nanabFo (1) Trunks yes Night (1)
Phalangopsidae Paragryllinae Caltathra balmessae Fo (2) / Pr (2) Trun ks/roots no Night (4)
Phalangopsidae Paragryllinae Caltathra meunieri Fo (13) / Pr (2) Tr un ks/roots no Day (1) / Night
(14)
Phalangopsidae Phaloriinae Pseudotrigonidium ana Fo (3) / Pr (5) Trunks/leaf litter no Day (1) / Night (7)
Phalangopsidae Phaloriinae Pseudotrigonidium
caledonicab
Fo (1) Lea f litter/
understory
yes Day (1)
Trigonidiidae Nemobiinae Kanakinemobius spbFo (3) / Pr (1) Understory no Night (4)
Trigonidiidae Nemobiinae Koghiella flammea Pr (3) / Sh (116) Leaf litter yes Day (59) /Night
(60)
Trigonidiidae Nemobiinae Koghiella nigris Fo (53) / Pr (8) Bare soil yes Day (23) /Night
(38)
Trigonidiidae Nemobiinae Bullita fusca Fo (182) / Pr (98) Leaf litter yes Day (128) / Night
(152)
Trigonidiidae Nemobiinae Bullita mouirangensisbFo (21) Leaf litter yes Day (12) / Night
(9)
Trigonidiidae Nemobiinae Bullita obscuracPr (72) Leaf litter yes Day (37) / Night
(35)
Trigonidiidae Nemobiinae Paniella bipunctatusbFo (1) Trun ks no Day (1)
Trigonidiidae Trigonidiinae sp Pr (1) Leaf litter ? Night (1)
Note: N/A indicates data not available.
aNumber of specimens collected.
bSpecies unique to forest.
cSpecies unique to preforest.
dSpecies unique to shrubland.
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ANSO et al.
while preforest and shrublands shared four. Forest and shrubland,
however, shared no species.
Collected species additionally provided several unique character-
istics. For instance, 70% of the cricket species collected were stridu-
latory species (Anso, Jourdan, & Desutter- Grandcolas, 2016). It was
also found that forests contained the lowest percentage of acoustic
species when compared to total species richness (65%), followed by
shrublands (71.4%) and preforest (78.6%) (Table 2). Of all species col-
lected, a total of eleven were collected only once during the entire
taxonomic inventory. During this inventory, a larger number of spe-
cies were collected at night (90%) than during the day (60%).
3.2 | Environmental succession
Environmental results demonstrated: (1) a clear difference in veg-
etation and climatic parameters between ecological stages, with a
higher variability between shrubland and forested sites, and (2) a
progression in the variable values that follow the ecological stage
succession (Table 3).
A large number of vegetation parameters separated open shrub-
lands from preforest and forest stages: a higher bare ground layer,
a lower vegetation height, a lower number and size of stems, and a
higher canopy openness. Conversely, vegetation parameters were
identical between preforest and forest, with the exception of vege-
tation height, which was higher in the forest. The herbaceous layer
and tree layer thus differed between forest and shrubland only.
Plant species richness did not significantly differ when compared
among ecological stages.
Several climatic parameters differed in the succession. Mean and
maximum temperature, along with daily variation, followed the eco-
logical gradient, (i.e., the highes t and lowest values for shrubland and
forests, respectively). Mean and minimum relative humidity differed
significantly between forest and the two other ecological stages,
with lowest and highest values for shrublands and forests, respec-
tively. Daily humidity variation followed the ecological gradient (i.e.,
highest in shrublands, followed by preforest and subsequently for-
est). The minimum temperature and the maximum humidity were
not significantly different between the successional stages.
3.3 | Alpha diversity of cricket assemblages
Species richness of crickets was not significantly different between
sites of differing ecological stages due to large standard deviation
values. This was indicative of a high variation of intra- site species
richness values. However, the overall abundance of crickets was sig-
nificantly lower in shrubland than preforest and forest (Table 3).
3.4 | Beta diversity of cricket assemblages
Each ecological stage showed clearly separated cricket assem-
blages (Figure 2) based on NMDS ordination (stress = 0.065, two
dimensions). Thirteen environmental variables significantly con-
tributed to the ordination model, with six at a high significance
level (p < = .01): maximum temperature, daily temperature vari-
ation, mean temperature, percentage of canopy openness, veg-
etation height, and percentage of bare ground (Figure 2a). Cricket
assemblages in shrubland were the most distinct, with a habi-
tat characterized by an open canopy, a high percentage of bare
ground coverage, and a large percentage of herbaceous and shrub
growth. Cricket assemblages in forest and preforest were similar,
with habitats characterized by a high canopy height, a high num-
ber of stems with a large circumference, and a high percentage of
trees. These ecological stages explained 52% of the total varia-
tion in cricket assemblages observed from the taxonomic inven-
tory (permutation test: R2 = 0.522; p = .004). Similar results were
obtained when considering the presence or absence of the cricket
species (55%; permutation test: R2 = 0.551; p = .001). Notably, the
daily humidity significantly contributed to the ordination model
(Figure 2b).
Focusing only on cricket species with a stridulatory apparatus
produced similar results (Figure 2c). The ecological stages explained
53% of the total variation of the cricket assemblages (permutation
test: R2 = 0.527, p = .006). NMDS showed a clear succession of
cricket assemblages along the ecological succession (stress = 0.034,
two dimensions). Eleven environmental variables significantly con-
tributed to the ordination model with a high significance for veg-
etation height and mean temperature (p < = .01). Considering the
presence or absence of the cricket species, the ecological stages
explained 60% of the total variation of the cricket assemblages (per-
mutation test: R2 = 0.6, p = .001). For that dataset, thirteen signifi-
cant variables were identified for the ordination model, with seven
of high significance (p < = .01): percentage of shrub layer, percentage
of bare ground, vegetation height, percentage of canopy openness,
maximum temperature, daily temperature variation, and daily hu-
midity variation (Figure 2d).
Based on cricket assemblage spatial turnover and nestedness,
results indicated that spatial turnover almost exclusively con-
tributed to the beta diversity observed (β = 0. 874, βSIM = 0 .807,
βNES = 0.067). The same analysis performed for the three eco-
logical stages produced similar results (β = 0.698, βSIM = 0. 567,
βNES = 0.131). This suggests that species are replaced by others
with low loss of species from other sites and ecological stages.
Results from Section 3.3 that demonstrate a nonsignificant dif-
ference in species richness between sites of different ecological
stages further support this idea.
3.5 | Relevant species by successional stage
SIMPER analysis demonstrated that specific cricket species were
keys in distinguishing communities in the succession (Table 4), several
of these species emitting specific calls making them a good candidate
for being bioindicators detected by acoustic sensors. Based on spe-
cies abundance in the assemblages, Agnotecous azurensis, Koghiella
flammea, Bullita fusca, and B. obscura Anso & Desutter- Grandcolas,
|
7
ANSO et al.
2016 contributed the most to the differences between shrubland
and preforest stages. These species live in leaf litter, with the abil-
ity to produce a species- specific calling song. K. flammea, B. fusca,
and B. obscura are active by day, while A. azurensis species is active
by night. Considering the differences between shrubland and forest
stages, three species were found to contribute the most: B. fusca,
A. azurensis, and K. flammea. Finally, B. fusca, A. azurensis, B. obscura,
and K. nig ris Anso & De sutter- Grandcolas, 2016 contribute d the most
to the differences between forest and preforest stages. K. nigris is a
small nocturnal Nemobiinae (Trigonidiidae) living strictly in forested
habitat, in a peculiar microhabitat (i.e., bareground).
Applying this analysis to the species presence or absence in
the assemblages resulted in a larger number of contributing spe-
cies (Table 5). The species contributing the most to the differences
between preforests and shrublands were A. clarus D es ut te r-
Grandcolas, 2006, and K. flammea. A. clarus is acoustically active by
night and col lected on thr ee preforest sit es versus once in fo rest sites.
Differences between forests and shrublands were mainly associated
with A. azurensis, Caltathra meurieri Anso & Desutter- Grandcolas,
2016, and K. flammea. C. meunieri is a nocturnal Phalangopsidae
living on large trunks or roots, without stridulatory apparatus (i.e.,
“mute species”) and found in every forest site but only at two prefor-
est sites. The two species A. clarus and C. meurieri were found to con-
tribute the most to the difference between preforests and forests.
4 | DISCUSSION
The results of this study highlight the central role of crickets in con-
servation biology and demonstrate that crickets could be relevant
bioindicators of the restoration process and integrity of the environ-
ment in tropical systems.
FIGURE 1 Cricket species assemblages along the ecological succession. * indicates stridulatory species.
8
|
ANSO et al.
4.1 | Cricket assemblages along
ecological succession
Cricket assemblages clearly responded to the different stages along
the environmental gradient, with several distinct species in each
stage. Overall cricket abundances were highest in forested habi-
tats (i.e., forest and preforest), suggesting that these habitats pro-
vide the best trade- off between food resources (Barberena- Arias &
Aide, 2003; Williams et al., 2008), shelter from predators (Brouwers &
Newton, 2009), and favorable moisture conditions. Oviposition sites
may also be more important and various in forested habitats than in
more open ones. This is especially important for litter- dwelling spe-
cies, explaining higher densities in closed habitats (Huber, 1989).
These results are similar to those obtained for forest- dwelling grass-
hoppers in Western Amazonia by Amédégnato and Descamps (1980).
Lower cricket densities in shrublands might be explained by smaller
optimal vegetation patches, but also warmer mean and high tem-
peratures, lower relative humidity, and limited food resources (Shik &
Kaspari, 2010). Moreover, in drier habitats, such as shrublands, cricket
density may be further constrained by higher predation rates in com-
bination with limited shelter (McCluney & Sabo, 2009, 2014).
Accumulated species richness similarly showed a pattern re-
lated to ecological succession, with the highest number of species
found in forested habitats (forest and preforest). This presumably re-
flected greater habitat complexity and microhabitat availability (Shik
& Kaspari, 2010). However, based on the number of species found
at each site, the differences were not significant between ecolog-
ical stages. This could mean that a larger number of sites is needed
to fully describe the cricket assemblages living in forests as compared
to preforests or shrublands. Although preforest and forest share a
large number of cricket species, the latter is characterized by a higher
percentage of unique species. This pattern is expected for rich forest
ecosystems (Rahman et al., 2012) and comparable with fruit- feeding
butterflies, birds, leaf litter amphibians, and the crickets documented
in Brazilian forests (Barlow et al., 2007; Szinwelski et al., 2012). While
habitat complexity and heterogeneity might explain higher species
richness in forests (Shik & Kaspari, 2010), they were surprisingly close
to preforests in their vegetation attributes; however, they were clearly
different in climatic conditions (i.e., temperature and relative humidity).
These observations highlight the importance of fine vegetation
attributes or microhabitats (e.g., dead trunks, woody debris, and gaps
around roots) along with microclimate condition (i.e., high moisture,
low variation of temperature and humidity) in explaining community
differences between these two advanced stages of forest regen-
eration, that are forest and preforest, in the southern part of New
Caledonia. Among the ten species exclusively found in the forest , eight
TABLE 3 Mean values (± standard deviation) of cricket assemblages, vegetation attributes, and climatic parameters in the three
ecological stages calculated from values averaged per sites. P represents the significance of the values after post hoc Kruskal– Wallis tests
according to Conover for pairwise multiple comparisons of the ranked data between stages
Shrubland Preforest Fores t P
Cricket assemblage
Species richness 3.5 ± 1.29 7 ± 2.71 7.5 ± 3.7 -
Overall abundance 16.5 ± 9.5 8 48.38 ± 15.19 5 7.13 ± 12.85 F o - S h * * , P r - S h *
Vegetation attributes
Herbaceous layer (%) 55.4 ± 24.1 23.8 ± 16.5 12.5 ± 13.4 F o - S h *
Shrub layer (%) 53.8 ± 13.8 31.2 ± 8.3 32.8 ± 12.1 -
Tree layer (%) 2.5 ± 2 37.2 ± 17. 5 35.3 ± 8.6 F o - S h *
Bare ground (%) 27. 5 ± 11. 5 0.6 ± 1.3 0 ± 0 Fo- Sh***, Pr- Sh***
Vegetation height (m) 2.25 ± 0.96 14 .9 ± 2.25 23.8 ± 4.33 Fo- Sh***, Fo- Pr**, Pr- Sh**
Number of stems (n°.) 0.5 ± 0.71 19 ± 11.6 15.9 ± 5.19 F o - S h * , P r - S h *
DBH of stems (n°.) 12.6 ± 14.8 56 ± 11.6 57. 5 ± 11. 5 F o - S h * , P r - S h *
Canopy openness (%) 90.6 ± 6.7 9.81 ± 3.15 7. 23 ± 0.75 F o - S h * * , P r - S h *
Plant species richness 1 7.4 ± 2.43 14.8 ± 2.33 18.1 ± 6.02 -
Climatic parameters
Mean temperature (°C) 21.5 ± 0.76 19.5 ± 0.53 18.6 ± 0.43 Fo- Sh***, Fo- Pr*, Pr- Sh*
Minimum temp. (°C) 16.2 ± 1.59 16.5 ± 0.97 16.6 ± 0.37 -
Maximum temp. (°C) 28.8 ± 2 24 ± 0.61 21.2 ± 0.71 Fo- Sh***, Fo- Pr**, Pr- Sh**
Daily temp. variation (°C) 12.6 ± 2.6 7.54 ± 1.26 4.65 ± 0.52 Fo- Sh***, Fo- Pr**, Pr- Sh**
Mean humidity (%) 78.5 ± 4.8 82.9 ± 1.5 88.1 ± 1.62 Fo- Sh**, Fo- Pr*
Minimum hum. (%) 47. 4 ± 8.91 59.1 ± 1.54 68.8 ± 2.61 Fo- Sh***, Fo- Pr*
Maximum hum. (%) 96.1 ± 2.45 97 ± 1.35 96.4 ± 1.28 -
Daily hum. variation (%) 48.8 ± 6 .51 3 7.9 ± 1.34 27.6 ± 1.39 Fo- Sh***, Fo- Pr**, Pr- Sh**
*p < .05; **p < .01; ***p < .001; Fo signifies Forest, Pr Preforest, and Sh Shrubland.
|
9
ANSO et al.
were found on vegetation strata, interestingly suggesting a close rela-
tionship to vegetation attributes (i.e., plant species or structure), and
likely further interrelated with the older age of forest habitat.
Some cricket species were previously described as potential in-
dicator of an ecological stage such as Pseudotrigonidium caledonica,
previously described as inhabiting forest understory and singing
on low vegetation with a unique and recognizable low- frequency
song (Desutter- Grandcolas et al. Desutter- grandcolas et al., 2016).
However, the statistical analyses presented in this paper did not
reveal Pseudotrigonidium caledonica (Otte, 1987) to be a significant
FIGURE 2 (a) NMDS based on cricket species abundances per site. (b) NMDS based on the presence or absence of cricket species per
site. (c) NMDS based on abundances of soniferous cricket species per site. (d) NMDS based on the presence and absence of soniferous
cricket species per site. Scatter diagrams (coefficient for the inertia ellipse size = 1.5) were drawn according to the habitat type. Significant
fitted environmental variables are shown with the following abbreviations: HER for herbaceous layer, SHR for shrub layer, TRE for tree layer,
BAR for bare ground, VH for vegetation height, DN for number of stems, DBH for number and diameter of stems, CO for canopy openness,
MT for mean temperature, HT for maximum temperature, DT for daily temperature variation, MH for mean humidity, LH for minimum
humidity, and DH for daily humidity variation.
Forest Shrubland
Preforest Bullita fusca 28% (28%) Agnotecous azurensis 28% (28%)
Agnotecous
azurensis
17% (45%) Koghiella flammea 22% (50%)
Bullita obscura 15% (60%) Bullita fusca 19% (69%)
Koghiella nigris 11% (71%) Bullita obscura 14% (83%)
Shrubland Bullita fusca 29% (29%)
Agnotecous
azurensis
26% (55%)
Koghiella flammea 19% (74%)
TABLE 4 Contribution percentage of
the most influential species for each pair
of habitats calculated based on species
abundances, with cumulative percentages
in parentheses. The most influential
species have with a minimum cumulative
percentage of 70% of the contributions
10
|
ANSO et al.
indicator of the forest. This is due to the fact that this species was
collected only once in our survey, which highlights an important bias
of our analyses towards rare species.
Previously, Desutter- Grandcolas (1997) have described the
cricket community in New Caledonia forested habitat for dif-
ferent soil types (volcano- sedimentary rocks) than those stud-
ied here (ultramafic soil). Their cricket community structure was
described as close to our forest community (in terms of species
richness and dominant species), with 23 species identified be-
longing to the same genera, but all different at the species level.
Taken together, the studies seem to confirm cricket genera
specialization for habitats of a given vegetation type and plant
structure (Anso, Jourdan, & Desutter- Grandcolas, 2016) and the
microendemism pattern of Orthoptera species in New Caledonia
(Grandcolas et al., 2008).
4.2 | Implications for conservation
The findings presented here provide strong evidence that cricket
species and their assemblages are sensitive indicators of ecologi-
cal succession in New Caledonia, with potential for applications
in conservation. There are several characteristics that make them
highly suitable as bioindicators for terrestrial monitoring in New
Caledonia. In combination with the bioindicator evaluation by
Gerlach et al. (2013), New Caledonian crickets are found in high den-
sities throughout the year, can be easily sampled by sight or hearing,
are sensitive to ecological stresses, and have a thorough taxonomic
understanding including various ranges of bioclimatic preferences
(Desutter- Grandcolas, 1997; Desutter- grandcolas et al., 2016; Otte
et al., 1987). Available data on assemblages of crickets in scientific
literature tend to show that crickets could be good bioindicators in
Forest Shrubland
Preforest Agnotecous clarus 9% (9%) Agnotecous clarus 11% (11%)
Caltathra meunieri 7% (16%) Koghiella flammea 10% (21%)
Pseudotrigonidium
ana
6% (22%) Agnotecous azurensis 8% (29%)
Koghiella nigris 6% (28%) Bullita fusca 8% (37%)
MOGOPLISTIDAE
sp2
5% (33%) MOGOPLISTIDAE
sp3
8% (45%)
Bullita fusca 6% (39%) Pseudotrigonidium
ana
8% (53%)
Bullita mouirangensis 4% (43%) MOGOPLISTIDAE
sp2
6% (59%)
Kanakinemobius sp 4% (47%) Pixibinthus sonicus 5% (64%)
Caltathra balmessae 3% (50%) Caltathra meunieri 6% (70%)
Notosciobia minoris 4% (54%)
Pixibinthus sonicus 4% (58%)
TRIGONIDIINAE sp 3% (61%)
Agnotecous azurensis 3% (64%)
Bullita obscura 4% (68%)
MOGOPLISTIDAE
sp1
3% (71%)
Shrubland Agnotecous azurensis 10% (10%)
Caltathra meunieri 10% (20%)
Koghiella flammea 11% (31%)
MOGOPLISTIDAE
sp3
7% (38%)
Bullita fusca 4% (44%)
MOGOPLISTIDAE
sp2
5% (49%)
Pixibinthus sonicus 4% (53%)
Pseudotrigonidium
ana
5% (58%)
Koghiella nigris 4% (62%)
Bullita mouirangensis 4% (66%)
Adenopterus
crouensis
4% (70%)
TABLE 5 Contribution percentage
of the most influential species for each
pair of habitats calculated based on
presence of the species, with cumulative
percentages in parentheses. The most
influential species have with a minimum
cumulative percentage of 70% of the
contributions
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11
ANSO et al.
other tropical areas, as they are most diverse in the tropics, espe-
cially in forested habitats. This reinforces the use of Orthoptera as
bioindicators (Gerlach et al., 2013).
All the ecological stages considered in this study are important
for conservation as each harbors a distinctive assemblage of cricket
species and their destruction would lead to irreversible loss of
unique biodiversity in New Caledonia. This provides valuable insight
for managers and decision- makers and further highlights the large
biodiversity of New Caledonian forests, with numerous endemic
genera and species.
Open shrublands are usually less studied for their fauna
(Chazeau, 1993) as they are often defined as the result of ecologi-
cal degradation by human activities. While this description may be
apt for particularly degraded areas, shrublands can be valuable for
conservation efforts. In the present study, open shrubland harbored
several distinct species and genera inhabiting leaf litter and shrubs.
Some species are particularly useful in capturing the impact of spe-
cific disturbances (i.e., fires, climatic oscillations) demonstrating the
different level of disturbances throughout the shrubland possibili-
ties. This was demonstrated with the discovery of the new mono-
typic genus Pixibinthus Robillard & Anso 2016 in open shrubland
and some preforest that provided useful information about fire re-
gimes and climatic fluctuations and their subsequent influences on
historic diversification processes in New Caledonia (Anso, Barrabé,
et al., 2016). While high species richness has long been established in
New Caledonian shrublands for plants, their value for animal conser-
vation is here highlighted. As gallery forests in the Neotropical re-
gion, shrublands are not only “forests- to- be” that serve as corridors
for forest species but host an original fauna of their own.
4.3 | Recommendations and perspectives for
passive acoustic monitoring
The sampling design is highly important when studying crickets.
Previous research on the response of cricket communities to vegeta-
tion succession has shown contrary or inconclusive results; however,
this could be due to a lack of taxonomic expertise, inadequate sam-
pling methodology, and/or the type of habitat (nontropical areas or
grasslands for example are usually poor in cricket species). For exam-
ple, while Araújo et al. (2015) found no relationship between habitat
degradation (from highly degraded to natural forested habitats) and
cricket abundances, the passive trapping system (i.e., pitfall trap)
used was not adequate to sample cricket communities (Gardiner
et al., 2005), potentially leading to bias in the results. Because nu-
merous cricket species live on vegetation, rocks, or tree trunks, with
restricted displacements on leaf litter (Desutter- Grandcolas, 1992,
2002; Jain & Balakrishnan, 2011), the low abundances of crickets ob-
tained by Araújo et al. (2015) (3– 5 individuals per habitat) cannot be
considered a sufficient evaluation of cricket communities. Similarly,
there are several other studies that were inconclusive for crickets
because of the low species richness recorded in nontropical steppes
or meadows (Fartmann et al., 2012; Marini et al., 2009).
When adequate sampling and taxonomic effort are present,
crickets are a powerful tool to evaluate the effect of succession on
insect communities in the tropics. The most comprehensive study
to date for cricket communities was conducted along a large for-
est regeneration gradient in Brazil (i.e., in a 300- year time period)
(Szinwelski et al., 2012). Positive correlations between species rich-
ness and forest age, canopy cover, and litter depth was a key find-
ing. Critically, Szinwelski et al. (2012) provided strong evidence of
cricket species succession and specificity along a large habitat chro-
nosequence, including open to closed habitats, with a comparable
turnover in species composition to our results in New Caledonia.
Similar results were obtained for a forest succession over 250 years
in Western Amazonia, where different cricket communities were
found in the forest plots according to their age of regeneration after
slash- and- burn cultivation (Desutter, 1990).
This work is part of a larger research program about
Neocaledonian crickets and contributes to highlight the potential
value of crickets as an environmental indicator of tropical ecosys-
tems. It further provides a unique opportunity for Passive Acoustic
Monitoring (PAM).
Firstly, New Caledonian cricket fauna comprises many acousti-
cally active species. In the present work, of the 30 species inven-
toried, 20 can produce sound, i.e., 67%. For comparison, among
the 23 species collected by Desutter- Grandcolas (1997), 14
(= 61%) can stridulate. Based on those two studies, bioacoustics
may capture up to two- thirds of cricket species in New Caledonia.
Secondly, calling songs are composed of a stereotyped repetition
of syllables (sensu Ragge & Reynolds, 1998) with a narrow carrier
frequency, and both the frequency spectrum and the call temporal
pattern allow for highly reliable species identifications (Diwakar &
Balakrishnan, 2007; Jain et al., 2014; Riede, 1997). Thirdly, cricket
calls are abundant and dominate the neocaledonian soundscapes,
particularly at night, whether it be in shrubland, preforest, or for-
est (Gasc et al., 2018). Thereby, Gasc et al. (2018) concluded that
only 6 days of passive acoustic recordings were necessary to de-
scribe the calling cricket community. Fourthly, calling cricket com-
munities reliable bioindicators of the surrounding environment
and their analyses can reveal major environmental perturbation.
In the present work, community analysis was performed only with
stridulating crickets revealing the same succession- based special-
ization as compared to the analyses using all species. As shown
by Gasc et al. (2018), the presence of the invasive ant Wasmannia
auropunctata resulted in a significant reduction in cricket species
and their acoustic activity.
In contrast with other methods, such as fogging or traditional
taxonomic inventories, the ecoacoustic approach is advantageous
as it is noninvasive, easy to implement, allow data collection over
large temporal and spatial scales, and in remote places (Sueur &
Farina, 2015). An acoustic approach for biodiversity monitoring is
meaningful in species- rich ecosystems, such as tropical rainforests
(Deichmann et al., 2018), where taxonomic impediments may be
overcome by the recognizable taxonomic unit of songs (Riede, 1998,
2018). Acoustic monitoring could not, however, be used to
12
|
ANSO et al.
characterize the sampled communities by their specific components
unless a preliminary taxonomic effort was conducted. When possi-
ble, taxonomic inventory to establish ground knowledge combined
with passive acoustic monitoring to sample natural environments
seems an effective strategy, and especially necessary in the trop-
ics where biodiversity still needs description. The research effort in
ecoacoustics conducted the last decade in New Caledonia within
the same research program, including the present work, brought
together sufficient taxonomic, ecoacoustic, and ecological knowl-
edge to confirm the value of crickets as bioindicators and support
the use of a passive acoustic approach to monitor ecosystems (Anso,
Jourdan, & Desutter- Grandcolas, 2016; Gasc et al., 2013, 2018).
Because crickets are globally distributed, particularly in endangered
biodiversity hotspots and in the tropics, they can be a key tool and
metric for conservation programs.
ACKNOWLEDGMENTS
We would like to thank Frédéric Rigault (IRD Noumea), Jean- Yves
Meunier (IRD Nouméa), Léa Colmas, and Alexia Omont (Master stu-
dents) who provided help with field work, habitat characterization
(FR) and mapping our sampled localities. We are also grateful to Eric
Vidal (head of IRD Nouméa) for his support, Alexandre Millon for
helpful comments on our statistical analysis, and Garett Pignotti
for providing thoughtful comments on the entire paper and English
edits. We also thank Thomas Ibanez who provided helpful insight
into community analysis. We thank the New Caledonian Direction
de l'Environnement de la Province Sud for providing permit col-
lection. This study was partly funded by a Ph.D. grant, “Bourse
d'encouragement à la recherche universitaire,” attributed to JA by
the Government of New Caledonia. Field and lab work were partly
supported by a funding attributed to HJ by Agence Nationale de
la Recherche (ANR) through the ERA- Net BiodivERsA Project
(FFII, JE 288/7- 1) and by a grant from the Grand Observatoire du
Pacifique Sud: AAP GOPS 2013 / “Bioacoustique des grillons de
Nouvelle- Calédonie.” The laboratory procedures were supported
by two Action Transversale du Muséum (ATM) research programs:
“Biodiversité actuelle et fossile. Crises, stress, restaurations et pan-
chronisme: le message systématique,” and “Barcode.” We would
like to thank the two anonymous reviewers for their constructive
comments.
CONFLICT OF INTEREST
The corresponding author confirms on behalf of all authors that
there have been no involvements that might raise the question of
bias in the work reported or in the conclusions, implications, or opin-
ions stated.
DATA AVAIL ABILI TY STATEMENT
The data and code that support the findings of this study are openly
available in the Dryad Digital Repository: https://doi.org/10.5061/
dryad.2280g b5w3 (Anso et al., 2022) and https://github.com/agasc/
Anso- et- al- 2022- Biotr opica.
ORCID
Amandine Gasc https://orcid.org/0000-0001-8369-4930
Laure Desutter- Grandcolas https://orcid.
org/0000-0002-7781-3451
Hervé Jourdan https://orcid.org/0000-0002-3756-4008
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SUPPORTING INFORMATION
Additional supporting information can be found online in the
Supporting Information section at the end of this article.
How to cite this article: Anso, J., Gasc, A., Bourguet, E.,
Desutter-Grandcolas, L., & Jourdan, H. (2022). Crickets as
indicators of ecological succession in tropical systems, New
Caledonia. Biotropica, 00, 1– 15. https://doi.org/10.1111/
btp.13151
