Journal for Nature Conservation 61 (2021) 125985

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Journal for Nature Conservation

journal homepage: www.elsevier.com/locate/jnc

Environmental variables and distribution data support the preliminary

conservation status of a trapdoor spider in a transformed coastal ecosystem
Rodrigo M. Barahona-Segovia a, b, *, Rubén Montenegro c, Rodrigo González G. d
a
Departamento de Ciencias Biológicas y Biodiversidad, Universidad de Los Lagos, Av. Fuschlöcher 1305, Osorno, Chile
b
Centro de Estudios en Ecología Espacial y Medio Ambiente – Ecogeografía, Santiago, Chile
c
Museo Nacional de Historia Natural, Quinta Normal, Santiago, Chile
d
Tarántulas de Chile Citizen Science Program, Chile

A R T I C L E I N F O A B S T R A C T

Keywords: Mediterranean coastal ecosystems are major tourist destinations that support intensive real estate business de­
Actinopodidae velopments. However, these areas are also important as biodiversity hotspots. Coastal spiders have been poorly
Coquimbo region studied as indicators of habitat disturbance, especially those that depend on certain environmental variables.
Maximum temperature
Plesiolena bonneti is a trapdoor spider that exclusively inhabits the coastal areas of central Chile (i. e., the
Real estate business
Relative humidity
Coquimbo region, 29◦ 57′ S, 71◦ 20′ W), intensely affected by habitat loss due to urbanization associated with
Tourism tourist destinations. The main aim of this work was to establish, preliminary, the conservation status of P. bonneti
supported by different environmental variables that define the activity period and occurrence of P. bonneti males
and its distribution area. Trapdoor spiders are rare and difficult to observe in the field. and therefore, we used
datasets obtained from citizen science to establish the period of activity, and environmental variables (maximum
temperature, relative humidity, and month of observation) as predictors of its occurrence. Moreover, since it
lives in an area perturbed by urbanization, we propose a preliminary conservation status using the IUCN Red List.
We found that the species is observed significantly more in the austral autumn and that its occurrence (65 % of
variability through Akaike weight) depends on maximum temperatures of 17 ◦ C and relative humidity above 90
%. Its current distribution is mainly coastal, between the Atacama and Coquimbo regions. We propose that this
species should be considered a vulnerable species. The increased observation of male P. bonneti post-rains may be
driven by a physiological response to avoid water loss, which would coincide with the reproductive period. As
the largest number of records are observed in coastal urban areas, we propose two simple management measures:
the creation of nature reserves within the city, and an educational program for this spider.

1. Introduction
Coastal urban development in Mediterranean biomes has increased
exponentially over time (Barragán & de Andrés, 2015). These changes in
land use impact deeply on ecological processes and biodiversity, and are
a major threat to all Mediterranean coastal biomes (González,
Yáñez-Navea, & Muñoz, 2014; Kaniewski et al., 2013; Underwood,
Viers, Klausmeyer, Cox, & Shaw, 2009). Hotels, secondary housing, and
other buildings for tourism and recreational activities have promoted
coastal urbanization, and are main drivers in the native coastal
ecosystem transformation (Burak, Dogan, & Gazioglu, 2004; González &
Holtmann-Ahumada, 2017; Paskoff & Manriquez, 1999). In addition,
coastal ecosystem services and biodiversity in Mediterranean biomes
and how they respond to urbanization have been intensively researched
(Drius et al., 2019; Liquete, Piroddi, Macías, Druon, & Zulian, 2016).
Although spiders are perceived as biological indicators of coastal
change in the land, this role has rarely been assessed for this biome
(Bidegaray-Batista et al., 2017; Ghione, Simó, Aisenberg, & Costa, 2013;
Seer, ElBalti, Schrautzer, & Irmler, 2015). Spiders show high fidelity for
specific habitat resources or environmental conditions and have small
home ranges and distributions (Bidegaray-Batista et al., 2017; Seer et al.,
2015.) Over time, many authors in different ecosystems have associated
habitat complexity (e.g., vegetation diversity) as a main predictor of
richness or abundance of spider communities (Bowden & Buddle, 2010;
Carvalho et al., 2015; Jiménez-Valverde & Lobo, 2007; Rypstra, 1986).
However, some environmental variables are key to habitat-specialized

* Corresponding author at: Departamento de Ciencias Biológicas y Biodiversidad, Universidad de Los Lagos, Av. Fuschlöcher 1305, Osorno, Chile.
E-mail address: rbarahona13@gmail.com (R.M. Barahona-Segovia).

https://doi.org/10.1016/j.jnc.2021.125985
Received 18 January 2021; Received in revised form 20 February 2021; Accepted 24 February 2021
Available online 28 February 2021
1617-1381/© 2021 Elsevier GmbH. All rights reserved.
R.M. Barahona-Segovia et al.
spider groups, such as mygalomorph spiders (Campbell & Engelbrecht,
2018). This group possesses high surface-volume ratio and, therefore,
they are vulnerable to environmental changes (Nentwig, 2012). Canals
et al. (2011) and Mason, Tomlinson, Withers, and Main (2013) deter­
mined that higher temperatures and low relative humidity in the air
influence water loss via evaporation in mygalomorphs. Therefore, spe­
cific environmental conditions predict activity patterns and occurren­
ce/abundance of mygalomorphs and increase their vulnerability.
Trapdoor spiders (Actinopodidae) are mygalomorphs that inhabit
burrow nests whose outer opening is protected by an operculum artic­
ulate, offering protection against predators and microhabitat to main­
tain physiological conditions (Ferretti, Copperi, Schwerdt, & Pompozzi,
2014; Grismado and Goloboff ;, 2014; Goloboff, 1994; Main, 2001;
Shillington, 2002). Juvenile and mature males of mygalomorphs
disperse short distances and females are mainly sedentary (Main, 2001).
Many trapdoor spiders can live many time, reaching maturity after 6–7
years (Main, 2001). They are habitat-specialized and have small distri­
butional ranges (Bond, Beamer, Lamb, & Hedin, 2006; Bonte, Lens, &
Maelfait, 2006; Coyle & Icenogle, 1994; Decae, Mammola, Rizzo, &
Isaia, 2019; Rix, Wilson, & Harvey, 2019; Vincent, 1993; Yáñez &
Floater, 2000). Considering the high pressure on coastal Mediterranean
biomes by urbanization and other human activities, spiders with a
burrowing lifestyle, such as Actinopodidae, are vulnerable to habitat
loss and more prone to extinction (Bond et al., 2006; Ferretti et al., 2014;
Yáñez & Floater, 2000). Currently, 159 spider species (only 0.32 % from
the total,) from 49,136 species described (World Spider Catalog, 2021)
are threatened under the IUCN Red List criteria. It can also be noted on
this list that the population of 45 species is decreasing, and the popu­
lation trends for 112 species are unknown. Moreover, residential and
commercial development threat to 22 % (n = 35) of spiders assessed (i.
e., VU + EN + CR). Spider assessment for the IUCN Red List is encour­
aged, although spatial and abundance data for many species are scarce.
Cardoso, Erwin, Borges, and New (2011) propose filling distributional
gaps in arthropods using citizen science and biodiversity repositories.
However, only some conspicuous species can be recognized with this
method (Hart, Nesbit, & Goodenough, 2018; Jiménez-Valverde,
Peña-Aguilera, Barve, & Burguillo-Madrid, 2019; Wang, Casajus,
Buddle, Berteaux, & Larrivée, 2018) including mygalomorphs (Camp­
bell & Engelbrecht, 2018). Maes et al. (2015) reviewed the use of
opportunistic data taken from citizen science for the IUCN Red List
assessment and found that the distribution criterion (i.e., B1 or B2) was
most used for different taxonomic groups.
Coquimbo coastal ecosystems are placed in the northern limit of the
Mediterranean biome of Central Chile, with predominant semiarid
conditions. This region has experienced an increase in urbanization for
tourism, as well as other human activities, resulting in many socio-
environmental conflicts (Aguilera, Tapia, Gallardo, Núñez, &
Fig. 1. Male of Plesiolena bonneti Zapfe, 1961: (A) frontal view and (B) dorsal view. Photos: Diego Demangel and Victor Pastén.
2
Journal for Nature Conservation 61 (2021) 125985
Varas-Belemmi, 2020; Carranza et al., 2020; González &
Holtmann-Ahumada, 2017; González et al., 2014; Paskoff & Manriquez,
1999). Two genera of trapdoor spiders inhabit these ecosystems: Mis­
sulena and Plesiolena. Missulena is represented in Chile by M. tussulena
Goloboff (1994), while Plesiolena is an endemic genus, restricted to
Coquimbo region and is represented by two species: P. bonneti Zapfe
(1961; Fig. 1) and P. jorgelina Goloboff (1994). For P. bonneti and
P. jorgelina the only localities in which they are found are those
described by Goloboff (1994). Males of P. bonneti are easily recognizable
because their abdomen has two large red maculae separated by a wide
central black line (Goloboff, 1994). Natural history or ecology is
completely absent for both genera. Recently, after seasonal rains during
mid-autumn, citizens have reported that P. bonneti males disperse
through the landscape, especially in the remaining natural areas within
the Coquimbo-La Serena urban or semiurban lands. This behavior has
also been reported by Ferretti, Pompozzi, Copperi, Pérez-Miles, and
González (2012) and Goloboff and Platnick (1987) for other Argentinian
and Chilean mygalomorphs. In this work, we determine the baseline for
the activity period and environmental drivers that could predict occur­
rence/abundance, as well as distributional limits of P. bonneti, using
these data to propose a preliminary conservation status of this species in
a coastal area transformed by urbanization in the Coquimbo region. Our
aims are
(i) to assess the activity period of P. bonneti through time,
(ii) to match the occurrence of P. bonneti with environmental
predictors,
(iii) to identify and compare distributional limits of occupancy be­
tween different occurrence sources, and
(iv) to assess the conservation status of this trapdoor spider species
using the IUCN Red List criteria.
2. Material and methods
2.1. Study site
Plesiolena bonneti is endemic to the Coquimbo region (29◦ 54′ S,
71◦ 15′ W; Fig. 2A) and is found mainly in coastal scrubland habitats
(Goloboff, 1994; Fig. 2B). We concentrated our search effort in this re­
gion and neighboring areas. The coastal scrubland ecosystem of
Coquimbo is a Chilean Mediterranean hotspot characterized by low
daily and annual temperatures due to maritime fog (Squeo, Arancio, &
Novoa, 2005). In ancient times (from Pleistocene to last glaciation),
vegetation was from sclerophyllous forest, mixed with scrubland
(Gajardo, 1993). The Coquimbo region suffered M8.3 earthquake in
2015, generating a tsunami that affected specific areas of biological
interest and local cities (Chávez-Villavicencio, Tabilo-Valdivieso, &
R.M. Barahona-Segovia et al. Journal for Nature Conservation 61 (2021) 125985

Fig. 2. Distribution (alpha-hull model) based on observations of Plesiolena bonneti Zapfe, 1961 in (A) Atacama and Coquimbo regions, and (B) coastal Mediterranean
scrubland. Photo: Rubén Montenegro.

Jofré-Pérez, 2019; Contreras-López et al., 2017). Currently, endemic

Table 1
vegetation is composed of endangered trees, such as Myrcianthes
Social media groups surveyed for Plesiolena bonneti occurrences.
coquimbensis (Barneoud) Landrum & Grifo and Myrceugenia correifolia
(Hook. & Arn) O. Berg (García-Guzmán, Loayza, Carvajal, Letelier, & Group name Target N◦ Link
biodiversity volunteers
Squeo, 2012; Hechenleitner et al., 2005). Common endemic species are
Senna cummingii (Hook et Arn.) Irw. et Barneby and Echinopsis coquim­ Flora, Fauna & Many organisms +37,000 https://www.facebook.
Fungi de Chile com/groups/FloraFauna
bana (Molina) Friedich & Rowley as well as many herbaceous plant
FungiChile/
species, such as Solanum spp. and Alstroemeria spp. (Fig. 2B). The Insectos de Chile Insects and +31,000 https://www.facebook.
Coquimbo Region, to date, has experienced a great expansion of ur­ arachnids com/groups/7779145
banization due to tourism (Carranza et al., 2020). This has led to the 22238618/
appearance of luxury resorts, vacation’s houses, industries related to Comunidad Insects and +25,000 https://www.facebook.
Entomológica arachnids com/groups/15669076
recreational activities and deregulated regional planning (Aguilera
Chilena 06922926/
et al., 2020; Barahona-Segovia, Núñez-Hidalgo, González-Césped, & Arañas de Chile Only arachnids +25,000 https://www.facebook.
Rojas-Osorio, 2019; García-Guzmán et al., 2012; González & com/groups/aranasdech
Holtmann-Ahumada, 2017; González et al., 2014). These have been the ile/
Insectos y Insects and https://www.facebook.
main drivers of Chilean coastal transformation. +11,000
Arácnidos arachnids com/groups/1926137
chilenos 24217649/
Tarántulas de Chile Only https://www.facebook.
2.2. Data collection
+2600
mygalomorphs com/groups/2762069
72846798/
Plesiolena bonneti occurrences were obtained using citizen science
records from different social media and biodiversity repositories gath­
ered between 2014–2020. We used Facebook groups dedicated to collect proximity to urban centers were also corrected for their rarity, reported
opportunistic records of insect biodiversity or arachnids (Table 1). In by Zapfe (1961) and Goloboff (1994) and due to different field works on
addition, other records were added from biodiversity repositories, such its potential known distribution (see below). Thus, for each resulting
as iNaturalist or Ispot (supplementary material S1). Social media is used citizen science record we obtained for each observation: (1) the more
as a method to obtain species identification of different specimens accurately geographic location (or coordinates) as possible provided by
photographed in Chilean ecosystems and is supported by several tax­ volunteers. When the location provided is only the city (e.g., La Serena
onomists and parataxonomists (Barahona-Segovia, 2019; Jarić et al., or Coquimbo), we used the centroid of a triangle, using the points that
2020). We used the search engine of each Facebook group by using the surround the imprecise location as reference; (2) precise date of record
keywords “Plesiolena bonneti” AND “Plesionela bonneti” (misspelled (day/month/year) was consulted to each volunteer; (3) the original
name) OR “Plesiolena bonetti” (misspelled name) OR “Actinopodidae” photo, and (4) the number of spider individuals observed as a proxy for
OR “Migidae” OR “tarántula de lluvia” OR “araña de Lluvia.” The search abundance. We conducted follow-up contacts with citizen scientists to
by each Facebook group was conducted separately and by keywords obtain missing or reliable information, as well as natural history data,
using similar time (1 h per day, two days in the week, two times each when necessary, according to Devictor, Whittaker, and Beltrame (2010).
month). This standardized our data because it increased the probability We eliminated those records that were not responded. To ensure taxo­
of spatially and temporally randomizing the records. This avoids biases nomic identification, Rubén Montenegro (a Chilean Mygalomorph
related to the same volunteer taken photos, particular tourist destina­ taxonomist) validated the photos and assigned to the target species.
tions within the study area or to the summer periods. Potential biases in Records for the same specimen, but from different people, were
the frequency of this species in social media groups generated by excluded from the analysis (data records available in https://figshare.

3
R.M. Barahona-Segovia et al.
com/s/5888733c2d33a69b87e9; DOI: https://doi.org/10.6084/m9.
figshare.13697242). Environmental data for each opportunistic regis­
ter were recorded. We used as a possible environmental predictor
maximum, mean and minimum temperatures, as well as the relative
humidity associated with each citizen science record, given that
P. bonnetti males appear after the first rains in transitional coastal desert
areas. We obtained environmental variable values from meteorological
stations near occurrence records. Temperatures and relative humidity
were stored by day and month through years in www.meteored.cl.
2.3. Distribution and conservation assessment
We assessed the distribution with Extent of Occurrence (EOO) and
Area of Occupancy (AOO) using opportunistic records and then, we
complemented this information by conducting field samplings to detect
the spider species. First, we evaluated EOO using minimum convex
polygons, which is characterized for containing all occurrences inside a
polygon with internal angles not higher than 180◦ (IUCN, 2012). AOO is
the sum of all grid cells of 2 × 2 km and is similar to those used by the
International Union for Conservation of Nature (IUCN, 2012). We
selected three localities to sample: Chungungo (29◦ 26′ S, 71◦ 18′ W),
Huachalalume (29◦ 58′ S, 71◦ 13′ O) and Las Barrancas (29◦ 49′ S,
71◦ 18′ W). A local researcher manually searched burrows of P. bonneti,
between stones and plants through linear transects of at least
100 × 10 m until completing an area of 2000 to 5000 m2 of total
coverage per locality. Each sampling lasted between four and six hours.
To ensure taxonomical classification (realized by R. Montenegro), bur­
rows and spider individuals were collected and counted. Due to move­
ment restrictions between localities imposed by the Chilean government
due to the COVID-19 pandemic, Chungungo was replicated three times,
while the other locations were repeated once each. All records (both
opportunistic and fieldwork) were referenced and mapped using ArcGIS
10.4.1. To assess the preliminary conservation status under the IUCN
Red List criteria, we used the ConR package focused on B criterion
(Dauby et al., 2017). To assign conservation status, distribution can be
used for two pathways and thresholds: B1 using EOO values (Critically
endangered [CR] < 100 km2; Endangered [EN] < 5000 km2 and
Vulnerable [VU] < 20,000 km2), and/or B2 using AOO values (CR < 10
km2; EN < 500 km2 and VU < 2000 km2). In addition, to meet conser­
vation status for B1 and B2, two of the three conditions are needed: (a)
severely fragmented or few localities (CR = 1; EN ≤ 5 and VU ≤ 10); b)
continuous decreasing, or (c) extreme fluctuations. We calculated the
percentage of urbanization that currently covers the main population of
P. bonneti, located between Coquimbo and La Serena cities, to obtain an
estimation of habitat loss and fragmentation. We limited this area to the
North and South between localities of Punta Teatinos and El Panul,
respectively, while to the East it was limited by the Pan de Azúcar lo­
cality and by the Pacific Ocean to the West, resulting in a core area of
418.86 km2 for P. bonneti. To calculate its, we used a satellite Landsat
image 2020 and Chilean land use cadaster and vegetation information
from CONAF (2014) to identify natural and urban areas in this period.
2.4. Statistical analysis
We performed circular statistical analyses to know whether there are
temporal differences in occurrences among record sets. To calculate
statistical parameters, we converted observation dates of trapdoor spi­
ders with citizen science into angles (commonly 30◦ each). However,
since the records of the P. bonneti dates are not standardized, to avoid
this bias, we grouped the observations into three subsets of days per
month (at the beginning (1–10), the middle (11–20) and the end of
month (21–30/31)), which allowed us to standardize this effect, trans­
forming the records into angles of six degrees for each subset (in total 12
bars; see figure in results). To probe whether spider occurrence is uni­
formly distributed over the months, we used a Rayleigh test (z), where
the null hypothesis represented that the entire month had the same
4
Journal for Nature Conservation 61 (2021) 125985
occurrences and no seasonality pattern appeared. As the Rayleigh test
assumes a von Mises distribution, we probed temporary segregation of
P. bonneti (i.e., higher frequency of individuals in some months than
others) with a Watson U2, which was used to test the goodness-of-fit
between von Mises and the observed distribution. We performed all
circular analysis with Oriana 4.0 (Kovach Computing Services, htt
p://www.kovcomp.co.uk/oriana/, Anglesey, U.K.).
To compare the contribution of different data sources to AOO in the
distribution of P. bonneti, we used a chi-squared for given probabilities.
To assess whether the abundance observed is associated with environ­
mental predictors, we performed a General Lineal Model (GLM) anal­
ysis. First, we applied an autocorrelation analysis between temperature
variables (i.e., maximum, minimum and mean). We assessed multi­
collinearity via pairwise Pearson r correlation analysis with ggplot and
corrplot packages in R software (R Development Core Team, 2019). We
selected maximum temperature because this should affect mygalo­
morphs according to Canals et al. (2011). Later, we fitted the full model
using Poisson error distribution because our data are counts and we
applied the overdispersion test using AER package to the suitability of
this model. However, overdispersion was alpha = 4.23 and therefore, we
used a negative binomial distribution error as final model. To construct
candidate models, we used relative humidity (%) and maximum tem­
peratures (◦ C) for each occurrence recorded as main environmental
predictors, according to Canals et al. (2011) and Mason et al. (2013)
because these traits trigger high activity in other mygalomorphs. We
used the subsets of each occurrence as a covariate. We selected the most
adequate candidate models ranking through Akaike’s Information Cri­
terion with small-sample correction (AICc), where the ΔAICc (i.e., rep­
resents the difference between the better models with other candidates;
Martensen, Pimentel, & Metzger, 2008), and AICc weight (i.e., repre­
sents the weight of evidence in favor of the best model; Burnham &
Anderson, 2002). Subsequently, we assessed the goodness-of-fit of the
selected model by running a likelihood ratio test comparing it to the null
model. We performed the analysis in R software using MASS, Matrix,
nlme, MuMIn and averaging mode (R Development Core Team, 2019).
3. Results
3.1. Activity period
Overall, we obtained 37 opportunistic records: 25 from Facebook
groups, 10 from biodiversity repositories, and two from the literature;
however, only 30 presented all data required for validation. Plesiolena
bonneti occurrence record showed that 93.3 % (n = 28) were single
spiders. Two volunteers reported several spiders (i.e., 15–20). The main
activity period for Plesiolena bonneti was late May-June and therefore,
occurrences are not distributed uniformly throughout time (z = 21.002;
p < 0.0001). Moreover, the length mean vector = 0.88 and the proba­
bility of observing individuals of P. bonneti in their habitat was higher in
June (Watson U2 = 1.328; p = 0.005; Fig. 3).
3.2. Occurrences and environmental predictors
The negative binomial error distribution was the best fit for our
dataset with a log link function (AICc = 128.26, Akaike weight = >
0.997 compared to Poisson AICc = 183.49). GLM analysis revealed that
the most parsimonious models for observed abundance of P. bonneti
were influenced mainly by the interaction between maximum temper­
ature and relative humidity (Table 2; Table 3). The best-fitted model
accounts for 65 % of the observed data (Table 2). Furthermore, the
second fitted model accounts for 23 % of the observed data (Table 2) and
was represented by an additive model between maximum temperature
and relative humidity. For the best model, the mean maximum tem­
perature was 17.4 ± 0.62 ◦ C, whereas mean relative humidity was
91.02 ± 1.63 %. The abundance of burrows and adults of P. bonneti
detected in the fieldwork was 0.003 burrows/m2 and 0.001 individuals/
R.M. Barahona-Segovia et al. Journal for Nature Conservation 61 (2021) 125985

[VU] B1ab (iii) + B2ab (iii).

4. Discussion

4.1. Environmental predictors and spider activity

Most observations of P. bonneti males, occurred in the austral

autumn-winter, similar to other South American Actinopodidae spiders
(Ferretti, Pérez-Miles, & González, 2010; Pérez-Miles, Costa, & Gudynas,
1993). In this period, sexually active males leave their nests to locate
mates after seasonal rains and are consistent with Ferretti et al. (2010).
Our results demonstrated that observed males are associated with daily
maximum temperatures (mean = 17.5 ◦ C) interacting with high mean
relative humidity (91.02 %). These results can correspond to an
ecophysiological response produced by seasonal rains that generate
Fig. 3. distribution of occurrences of Plesiolena bonneti Zapfe, 1961 using cir­ optimal environmental conditions to avoid high water loss rates. In
cular analysis. Circular dashed lines and number on them represent the number relation to this, Canals et al. (2011) and Mason et al. (2013) found that
of individuals recorded into each angle. (*) represents significant activity water loss by evaporation in mygalomorph spiders depends on high
(p < 0.05). Photo: Diego Demangel. temperatures in ecosystems with high environmental fluctuations. This
represents an environmental limit for the observation of these spider
m2, respectively, at the Huachalalume locality. In the remaining loca­ species through time. Recently, de Oca, Pérez-Miles, and Clavijo-Baquet
tions, no burrows or individuals of P. bonneti were found. (2020) showed that the reproductive period of mygalomorphs is con­
strained by thermal preferences and that these triggers the mate period
in males. Ecosystems with stable environmental temperatures, as the
3.3. Distribution and conservation status
Coquimbo region (influenced by the sea), can favor mygalomorphs
because they can maintain optimal temperatures and maximum per­
We found statistically significant differences in the AOOs provided
formance, as observed in other spiders (Alfaro et al., 2013). Optimal
by literature (at Herradura bay and north of La Serena) and biodiversity
environmental conditions can activate high metabolic rates and
repositories (χ2 = 14.4, df = 1, p = 0.0001) being the latter higher than
dispersal mechanisms. However, it is necessary to perform laboratory
historic dataset. We also found this difference in data provided by
experiments, as physiological performance test (e. g., speed time or
biodiversity repositories and Facebook groups (χ2 = 12.96, df = 1,
rollover speed test), at different controlled temperatures, as well as
p = 0.0003,) being social media higher than iNaturalist + ISpot. Finally,
thermal preferences in the field to prove this hypothesis.
we obtained statistically significant differences between Facebook
The central zone of Chile has decreased its rainfall between 20 and
groups and literature (χ2 = 47.36, df = 1, p < 0.0001) being social media
45 % from 2010 to the present, generating a mega drought of great
higher than historic AOO. We expanded the current distributionthank to
longevity, unprecedented and driven in part by anthropic factors (Gar­
citizen records by 105 km to the Northwest, to Caleta Sarco locality,
reaud et al., 2020). In addition, the gradual decrease in rainfall, even in
Atacama Region and by 34.35 km to the South, Las Cardas locality.
Trapdoor spiders were recorded in coastal areas or in coastal ranges with
maritime fog influence. Convex minimum polygon estimates (full data, Table 3
no time restriction) were for 2,268.5 km2 (EOO without sea) and for Effect of each parameter on observed abundance of Plesiolena bonneti. Estimates
AOO was only 72 km2. Natural and urban areas in the core of EOO (i.e., of the covariate coefficients, the unconditional standard errors and 2.5 % and
Coquimbo-La Serena) represent currently 319.19 km2 and 99.67 km2, 97.5 % confidence intervals (CI) (lower and upper limits, respectively) for all
respectively, with a habitat loss of 23.79 % (Fig. 4A) over six years covariates used in the GLM to explain P. bonneti observed abundance. Parame­
(2014–2020). Therefore, we considered habitat loss generated by ur­ ters and values in bold represent the most important parameters in the analyses.
banization as a main threat to this spider species (Fig. 4C-D). Following Covariates Estimate Standard error 2.5 % 97.5 %
the IUCN Red List criteria, this trapdoor spider meets the threshold to intecept 25.4833 14.3301 − 2.5980 53.5747
classify as vulnerable species using subcriteria B1 (2,268.5 km2 < 5000 tmax − 0.8788 0.5393 − 1.9358 0.1781
km2) and B2 (72 km2 < 100 km2). Localities displayed by ConR show 10 hum − 0.2442 0.1489 − 0.5361 0.0476
hum*tmax 0.0104 0.0046 0.0013 0.0195
localities (or six sub-populations); however, urbanization affect inten­
mon 3.0259 3.8538 − 4.5272 10.5792
sively to seven localities (Fig. 4). For both pathways, habitat quality is hum*mon ¡0.0677 0.0281 ¡0.1228 ¡0.0126
decreasing due to urbanization, considering that a big urban area is mon*tmax − 0.0833 0.0837 − 0.0756 0.2524
placed on the core of EOO, habitat fragmentation is increasing. We
hum = relative humidity; mon = month; tmax = maximum temperature.
propose a preliminary conservation status, which should be Vulnerable

Table 2
Candidate model set of P. bonneti abundance based on maximum temperature (tmax) as the main explanation and relative humidity (hum). Months (mon) were used as
covariates. Generalized linear models with a negative binomial distribution ordered according to AICc, with associated degrees of freedom (DF), number of parameters
in the model (k), AICc, ΔAICc, Akaike model weights, theta and Likelihood Ratio stats (χ2).
Candidate models K loglik AICc ΔAICc Weight χ2 DF GoF Theta

tmax + hum + (tmax*hum) 5 − 56.06 124.62 0.00 0.65 21.51 26 0.71 1.48
tmax + hum 4 − 58.55 126.69 2.08 0.23 23.49 27 0.65 1.23
hum 3 − 61.65 130.04 5.41 0.04 25.67 28 0.59 1.00
mon + hum + (mon*hum) 5 − 59.08 130.65 6.03 0.03 24.21 26 0.56 1.21
tmax 3 − 62.30 131.52 6.89 0.03 26.66 28 0.53 0.98
mon + hum 3 − 61.54 132.68 8.06 0.01 25.66 27 0.53 1.00
mon + tmax 4 − 61.59 132.79 8.17 0.01 26.20 27 0.50 1.03
mon + tmax + (mon*tmax) 5 − 60.87 134.24 9.62 0.01 25.94 26 0.46 1.09

5
R.M. Barahona-Segovia et al.
Fig. 4. (A) Coquimbo-La Serena cities present urban (as proxy of habitat loss) and natural areas; (B) El Culebrón wetland as example of urban reserve inside the
Coquimbo city and (C-D) represent tourism impacts overlaid on the historical distribution of Plesiolena bonneti.
years where the El Niño-Southern Oscillation phenomenon is present,
affects water availability in the next year (Álvarez-Garreton, Boisier,
Garreaud, Seibert, & Vis, 2020; Cai et al., 2020). This climatic situation
can be relevant to species such as P. bonneti, which depends on the scarce
seasonal rains, as the length of these rains affects the dispersal period of
males, reducing the rate of encounters with females, and affecting
abundance in the long-term. In addition, an increase in mortality or a
decrease in offspring could occur by reducing the prey that are abundant
in a flowering desert.
4.2. Distribution
Mygalomorph spiders are highly sedentary with low vagility, limited
home range, and therefore, they are endemic to small areas (Pérez-Miles
& Perafán, 2017). These characteristics promote habitat specialization
and rarity, with small distributional ranges (Bond et al., 2006; Coyle &
Icenogle, 1994; Decae et al., 2019; Rix et al., 2019; Vincent, 1993; Yáñez
& Floater, 2000). Our study expands the northern and southern limits of
P. bonneti; however, this species has a low total EOO and AOO associated
with coastal ecosystems highly transformed by urbanization. Coquimbo
and La Serena cities, together, have a dense human population (+400,
000 people), which increases three-fold in summer. These cities are
placed at the core of the known distribution of P. bonneti and currently,
¼ of the original habitat of this trapdoor spider has been replaced for
human settlements. Many males have been recorded in these cities in
spaces that retain native vegetation. The probability of death of this
6
Journal for Nature Conservation 61 (2021) 125985
trapdoor spider could be high due to car or truck tires, the attack of
domestic animals (e.g., domestic dogs) or being crushed by people.
Therefore, this rare specialized-habitat species needs urgent measures of
protection as a red list priority and habitat management, even inside the
city. Although some invertebrate priority lists for threatened ecosystems
are available for Chile (Barahona-Segovia, 2019; MMA, 2020), P. bonneti
has not been included in any.
4.3. The citizen science importance in spider conservation
The study of the distributional limits, activity period, and environ­
mental variables that predict the occurrence of P. bonneti males is
shown, for the first time, thanks to citizen science. This method has been
used for Latrodectus, Argiope, Tegenaria, and Eratigena or mygalomorphs
to show important distributional or spatiotemporal occurrence gaps
(Campbell & Engelbrecht, 2018; Jiménez-Valverde et al., 2019; Hart
et al., 2018; Wang et al., 2018). The study of P. bonneti with citizen
science is possible because males have conspicuous reddish abdomen
with a wide central black line (Goloboff, 1994; Fig. 1). However,
because trapdoor spiders are specialized-habitat and scarce, they are
highly vulnerable to human activities (Bond et al., 2006; Bonte et al.,
2006; Engelbrecht, 2013). Urbanization for touristic purposes has
transformed large coastal areas of the Coquimbo region (Carranza et al.,
2020; García-Guzmán et al., 2012; González & Holtmann-Ahumada,
2017; González et al., 2014). Worldwide, this practice has modified
the distribution of other mygalomorphs, such as Apomastus and
R.M. Barahona-Segovia et al.
Macrothele in California and the Iberian Peninsula, respectively, and it is
the major threat to these species, to the extent of modifying their genetic
structure (Arnedo & Ferrández, 2007; Bond et al., 2006).
Here, we propose P. bonneti, preliminary, as a vulnerable species
because their small geographic distribution, both in B1 as B2. However,
much of our evidence is from citizen science and may be complemented
with more fieldwork to confirm distributional limits as presented in this
paper. Fieldwork showed low abundance of P. bonneti, which justifies
the use of citizen science to complement the study of this rare species.
However, replicability must be ensured to maintain a minimum quality
under citizen science following Jarić et al. (2020). Both B1 as B2 path­
ways are based on rapid urbanization of coastline as the main threat and
10 localities (or two sub-populations) could be distinguished. Moreover,
urbanization can rapidly affect several localities and increase the cate­
gory from current vulnerable to endanger if adequate local policies are
not taken. Habitat fragmentation could generate low immigration of
individuals and genetic structuration between localities on P. bonneti, as
it has been reported in other mygalomorphs (Arnedo & Ferrández, 2007;
Bond et al., 2006), thus generating population depletion over time. Our
study represents the first baseline about the ecology of this trapdoor
spider; however, systematic conservation efforts need other measures to
preserve this species in coastal ecosystems transformed by urbanization.
The contribution of citizen science is key to fill distributional gaps of
endemic flies (Barahona-Segovia, Pañinao-Monsálvez, & Barceló, 2018)
or modeling the invasion of exotic bumblebees or ladybeetles (Montalva,
Sepulveda, Vivallo, & Silva, 2017; Werenkraut, Baudino, & Roy, 2020).
Also, it can be used to determine the population size of migratory bird
species (Thomson, Vukasovic, & Estades, 2020) or record novel
predator-prey interactions (Barahona-Segovia & Pañinao-Monsálvez,
2020). Citizen science projects engage volunteers in the knowledge of
biodiversity and generate relevant scientific information for the
decision-makers to incorporate into a national policy for the conserva­
tion of nature. Also, the conservation status under the IUCN Red List
criteria uses commonly opportunistic records (from citizen science
projects) to meet the thresholds required to assign a threatened category
(Maes et al., 2015). This work demonstrated that citizen science records
are key to enlarge the distribution and to know environmental variables
that determine the occurrence of P. bonneti.
4.4. Habitat management and education for saving trapdoor spiders
As Coquimbo and La Serena have been placed in the core of P. bonneti
distribution, habitat management inside and outside cities is key. We
propose two main measures to protect this trapdoor spider: (1) generate
local nature sanctuaries or urban national reserves, for example, in the
El Culebrón wetland basin (Fig. 4B). This area is vulnerable to exotic
predators, such as feral dogs, recreational activities and deregulated
tourism (Bravo-Naranjo et al., 2019; Chávez-Villavicencio, Molina
Pérez, & Tabilo-Valdivieso, 2015). The establishment of this urban
natural area could protect not only P. bonneti as well as the endangered
coastal bee Caupolicana fulvicollis (Barahona-Segovia et al., unpublished
data) as well as other epigean fauna. (2) Environmental education:
spiders are among Chilean primary-school students’ most favorite
epigean arthropods (Pizarro-Araya, López-Cortés, Jewell, & Inostroza,
2014). Plesionela bonneti is a conspicuous and endangered spider and
school children can be taught about their vulnerability and role as a
predator of urban insect pests within the ecosystem. Biodiversity is the
most important priority for Chilean educational curricula. This is key to
successfully educate future citizens that will have to take political de­
cisions in terms of protection and conservation of our biodiversity
(Barahona-Segovia et al., 2019), in particular of the Chilean coastline.
5. Conclusions
We conclude that the use of citizen science was important to know
distributional limits, activity periods and basic environmental
7
Journal for Nature Conservation 61 (2021) 125985
associations in a rare and endemic trapdoor spider. This information can
be used as a baseline for new hypotheses in physiology or ecology, not
only in P. bonneti, but also for rare, habitat-specialized and endangered
species facing natural environment transformations, particularly in
coastal ecosystems highly impacted by tourism and urbanization.
Declaration of Competing Interest
The authors report no declarations of interest.
Acknowledgments
RMBS wants to thank Agencia Nacional de Investigación y Desarrollo
(currently ANID) for providing a postdoctoral scholarship [ANID-FON­
DECYT 3200817]. We also thank Laura Pañinao-Monsálvez for your
help in map creation and habitat loss calculations; to Diego Demangel,
Bojana Kuzmicic and Victor Pastén for the high-quality photos of
P. bonnetti and to all citizen volunteers who provided the records and
information about the trapdoor spider. Finally, we would like to thank
our reviewers, Adrian Paterson and Stefano Mammola for providing a
throatful review of our manuscript, and Hernán Madrid, Dana Ortega
and Peter D. Lewis English editing service.
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