Arthropods associated with a dioecious bromeliad, Catopsis minimiflora Matuda (Bromeliaceae), in a shade coffee plantation (Coffea arabica L.) in the southeast of Mexico

Abstract

Shade coffee plantations are considered reservoirs of local flora and fauna. Epiphytic bromeliads are an important component of flora that inhabit not only shade trees but also coffee bushes in southeast of Mexico. At the same time, in these plants inhabit a diversity of arthropods poorly documented. We chose Catopsis minimiflora as the studied species because this bromeliad is abundant in coffee plantations and has a specialized reproductive system (dioecy). We counted the number of individuals of C. minimiflora growing over coffee bushes and shade trees and collected 58 bromeliads in two seasons (dry and rainy). We registered 2,048 arthropods (including 21 orders and 71 families) inhabited these plants. Based on hill numbers, no significant difference was found in richness between seasons; however, species dominance was higher in the rainy season. We estimated 27,215.5 arthropods/ha in the dry season and 31,227 arthropods/ha in the rainy season inhabited C. minimiflora that grow over coffee bushes. This arthropod community associated with C. minimiflora could provide ecosystem services such as pollination or depredation in a coffee agroecosystem. Epiphyte removal could have a negative effect on the abundance of this plant species, and in turn, it may have an impact on arthropods associated with them.

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Arthropods associated with a dioecious bromeliad, Catopsis minimiora
Matuda (Bromeliaceae), in a shade coffee plantation (Coffea arabica L.) in the
southeast of Mexico
Diego A. Jiménez-Garza
El Colegio de la Frontera Sur
Lislie Solís-Montero
El Colegio de la Frontera Sur https://orcid.org/0000-0002-5793-3376
Eduardo R. Chamé-Vázquez
El Colegio de la Frontera Sur
Nancy Martínez-Correa
Orquidiario y Jardin Botánico Comitán
Research Article
Keywords: arthropod diversity, epiphytic bromeliad, southeast Mexico, shade coffee plantation
Posted Date: April 29th, 2024
DOI: https://doi.org/10.21203/rs.3.rs-4092425/v1
License:   This work is licensed under a Creative Commons Attribution 4.0 International License.  Read Full License

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Abstract
Shade coffee plantations are considered reservoirs of local ora and fauna. Epiphytic bromeliads are an important component of ora that inhabit not only
shade trees but also coffee bushes in southeast of Mexico. At the same time, in these plants inhabit a diversity of arthropods poorly documented. We chose
Catopsis minimiora
as the studied species because this bromeliad is abundant in coffee plantations and has a specialized reproductive system (dioecy). We
counted the number of individuals of
C. minimiora
growing over coffee bushes and shade trees and collected 58 bromeliads in two seasons (dry and rainy).
We registered 2,048 arthropods (including 21 orders and 71 families) inhabited these plants. Based on hill numbers, no signicant difference was found in
richness between seasons; however, species dominance was higher in the rainy season. We estimated 27,215.5 arthropods/ha in the dry season and 31,227
arthropods/ha in the rainy season inhabited
C. minimiora
that grow over coffee bushes. This arthropod community associated with
C. minimiora
could
provide ecosystem services such as pollination or depredation in a coffee agroecosystem. Epiphyte removal could have a negative effect on the abundance of
this plant species, and in turn, it may have an impact on arthropods associated with them.
Introduction
The agroecosystems, mainly coffee plantations, are reservoirs of biodiversity (Hietz 2005; Hylander and Nemomissa 2008; Philpott et al. 2008; Solís-Montero
et al. 2019). Coffee plantations are mostly established in montane tropical rainforests, ecosystems with a high level of endemism that supply environmental
services such as water and nutrient recycling (Agudelo et al. 2012; Clark et al. 2014). Despite their biological and environmental importance of rainforests,
where coffee beans are grown, are suffering of deforestation. Forest fragmentation and slow recovery from perturbation are also associated to high
deforestation rates (Toledo-Aceves et al. 2012; Yepes-Quintero et al. 2011). Epiphytic orchids (Orchidaceae) and bromeliads (Bromeliaceae) found in
rainforests and coffee plantations are considered important to biodiversity (Hietz 2005). These epiphytes contribute to increasing canopy complexity and
provide resources to fauna that inhabit these ecosystems (García-Franco and Toledo-Aceves 2008).
Epiphytic bromeliads present crassulacean acid metabolism photosynthesis (CAM) (Crayn et al. 2004; Givnish et al. 2011; Givnish et al. 2014). Bromeliad
leaves form a cavity called tank, leaves have trichomes that absorb water and minerals (Crayn et al. 2004; Frank and Lounibos 2009; Quezada and Gianoli
2011; Katayama et al. 2020; Popp et al. 2003). Water and nutrients resulting from detritus decomposition and fecal matter of animals, or phytotelma, are
stored in bromeliad tanks (Marino et al. 2011; Wittman 2000). This makes bromeliads capable to maintain trophic nets that involve different organisms
including arthropods (Brouard et al. 2012; Cogliatti-Carvalho et al. 2010).
Arthropods, in particular insects, account for 90% of species found in the tropics (Nielsen 2011; Pimentel et al. 1992; Urbaneja et al. 2005) and may play
different roles in agroecosystems such as coffee plantations (Armbrech and Perfecto 2001). When the population abundance of these insects
increases, they are regarded as pests that damage the productivity of some crops (Guzmán-Mendoza et al. 2016). Yet, an increase in arthropod diversity does
not always represent an increment in pests (Armbrech and Perfecto 2001). For example, some insects can recycle nutrients through the decomposition of
other dead organisms to incorporate nutrients into the soil (Liria-Salazar 2006; Mavárez-Cardoza et al. 2005; Vergara-Pineda et al. 2020). Other insects
transport pollen grains from anthers to the stigma of plants to reach ovules and fertilize them (Armijo et al. 2020; Sheldon and Nadkarni 2015; Stefanescu et
al. 2018). And, other entomophagous insects help to control pests in different crops (Aguilar-Astudillo et al. 2019).
In coffee plantations located in Mexico is common to remove the epiphytes that grow over shade trees and coffee bushes. The rst option is known as
“destenche” and the second option as “desmusgue” (Cruz-Angón and Greenberg 2005; García-González et al. 2017; Solís- Montero et al. 2019). The
“desmusgue” usually takes place during the rainy season in Soconusco region (García-González et al. 2017; Solís-Montero et al. 2019). This agricultural
practice negatively affects the diversity of epiphytes that inhabit the coffee plantations (Solís-Montero et al. 2019). However, the diversity of arthropods
associated with abundant epiphytic bromeliads remains unknown.
This study aims to know the arthropod diversity associated with a dioecious bromeliad (
Catopsis minimiora
Matuda), which inhabits coffee plantations in
the Soconusco region, and the possible impact of removing epiphyte on arthropod diversity. We sought to answer the following questions: (a) Which is the
richness and abundance of arthropods associated with
C. minimiora
?; (b) What is the relation between morphologic characteristics of
C. minimiora
, and the
richness and abundance of arthropods?; (c) Which is the sex ratio and abundance of
C. minimiora
?; (d) Which is the possible impact of
desmusgue
over
population of
C. minimiora
and arthropods associated to this bromeliad?
Materials and methods
Studysite:Thisresearchwasconductedinashadecoffeeplantation(
Coffeaarabica
L.)locatedinBenito Juárez El Plan, Cacahoatán Municipality, Chiapas,
Mexico (15° 05’ 21.779’’ N, 92° 08’50.766’’W,1441m asl).Thissitewasselectedbecause“desmusgue”isnotpracticedhere. Four 10 x 30 m transects (300
m2 each transect) were established for a total of 1,200 m2. In eachsampling, we recorded temperature and humidity using a Data Logger, Elitech, RC-51H.
Thesamplingswereconductedduringthedryandrainyseasons(MarchandJuneof2022,CONAGUA 2022).
Studied species:
Catopsis minimiora
Matuda (1975) is a dioecious bromeliad of the Tillandsioideae subfamily. It is distributed across the southeast of
Mexico (Chiapas) and Guatemala at an altitude range from 1,200 to 1,800 m asl (Espejo-Serna et al. 2017). The Bromeliaceae family presents two types of
reproduction: sexual reproduction through seeds, and asexual reproduction through stolons (Mondragón et al. 2011). Asexual reproduction of
Catopsis
genera,
produces axillar or basal new plants from the mother plant to form tillers with several rosettes (Mondragón et al. 2011). In this study, we considered this tiller
as an individual. The individuals grow as epiphytic plants over shade trees or coffee bushes, namely phorophytes (Granados-Sánchez et al. 2004).
Catopsis
minimiora
female plants have larger owers and produce longer inorescences, yet they have fewer owers compared with male plants (Martínez-Correa
2019).

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Density and the sex ratio of
C. minimiora
:The density was obtained by counting the numberofindividualsof
C.minimiora
pertransect. We
countedyoungindividuals(over 15 cm in length) and adult individuals growing over shade trees and coffee bushes. To determine the sex ratio,
weonlyconsideredadult individuals presenting oralbuds, owers,fruits, orremainsofthem.
Bromeliad sampling:Only adults of
C. minimiora
growing on coffee bushes were collected ineach sampled transect; we tried to collect 16 bromeliads per
transect (eight males and eightfemales). But, in transects three and four the number of samples required was not reached; thus,we used
C.
minimiora
outside the transects. Transects one and two were sampled during the dryseason,whiletransects three andfourweresampled during therainy
season.
Before collecting
C. minimiora
individuals, we measured bromeliads height
consideringcoffeebushesasthebase;eachplantwasmeasuredfromthesoiltoitsgrowthonphorophyteusing a ex meter. We also measured the volume
(using a graduated cylinder of 500 ml) and thetemperature of the phytotelma water between 9 am and 11 am (using a TAYLOR thermometer).The water was
placed in a plastic container and the plants were stored in hermetic plastic bags
asfastaspossibletoavoidthearthropodsescape.Thesampleswerestoredinafridgeat approximately 6 °C in  Laboratorio de Artrópodos, Polinizadores,
Plagas y Vectores (APPYV) at El Colegiodela FronteraSur(ECOSUR), Tapachulacampus.
Sampleprocessing:Everyoneof
C.minimiora
wasdissectedover awhiteplastictrayfollowingDiaz (2021) methodology with some modications. We took
the following morphologic variables per individual: number of rosettes; plant length from the basal (i.e., the part where leaves join androots emerged) to the
apical part (i.e., longest leaf); diameter of each rosette (i.e., the
distancebetweentheapicalpartofthetwooppositeleaves);numberofleaves;andpresenceofreproductive structures(owers orfruits).
To collect arthropods, we dissected each individual and washed each rosette with tap water
overPetridishes.Finally,weobservedthesedishesbyusingastereoscopicmicroscope(Zeiss,Modelo Stereo Discovery. V20) and the arthropods were
separated into vials with 70% alcohol and stored atapproximately 6°C until their taxonomic identication.
Taxonomical identication:Weuseddifferentresourcesforarthropodtaxonomicalidentications such as manuals, voucher specimens’ comparison and
websites searching.For example, we identied at family level insects collected using the identication manual Introduction to the Study of Insects (Triplehorn
and Johnson 2005). To identify larvae, we used the identication manual Introduction to the Aquatic Insects of North America (Merritt et al. 2008). The spiders
were identied using the identication manual Spiders of North America (Ubick et al. 2017). The          specimens were deposited in Colección de Insectos
asociados a plantas cultivadas en la Frontera Sur (ECO-TAP-E) and Colección de Arácnidos del Sureste de Mexico (ECOTAAR) in ECOSUR, Tapachula campus.
Data analyses:For everyone of
C. minimiora,
we obtained the abundance (number of individuals), richness (number of families), and diversity (number of
Hill; Jost 2006) of arthropods. The diversity and rarefaction/extrapolation curves were obtained with iNEXT package version 3.0.0 (Hsieh et al. 2016). We used
a Principal Component Analysis (PCA) to analyze the morphological variables of
C. minimiora
(number of rosettes, length, diameter, and volume of
bromeliads). The volume wascalculated by following Nielsen’s (2011) methodology and using the following formula: V=(πr2h)/3. The scores of PC1
(interpreted as size) were used as a response variable; sex and season were the explicative variables in the Analysis of Variance (ANOVA). We used two
separated Generalized Linear Models (GLM); the rst GLM was used to analyze the effect of bromeliad size (PC1) on the diversity of arthropods (Hill numbers)
t with Gaussian; the second GLM was used to analyze the effect of season on the diversity of arthropods t with Gaussian. In these models, the Hill number
was the dependent variable; scores of PC1 and season were the independent variables. The sex ratio was analyzed using GLM t with binomial error (Wilson
and Hardy 2002). Finally, the density of each season was calculated by the following formula: Density = the number of
C.minimiora
/sampledarea(m2). To
extrapolate the abundance of individuals of
C.minimiora
per hectare, we multiplied the density for a hectare area (10,000 m2). Arthropod abundance was
extrapolated with the following formula: (Density of
C. minimiora
per ha) (Number of arthropods)/Number of
C.minimiora
. All analyses were performed
with R program version 4.2.1 (R Core Team 2020).
Results
Richness and abundance of arthropods:We counted 2,048 individuals that inhabited 58
C.minimiora
bromeliads,whichbelongto21ordersand71families(TableS1).TheordersPseudoscorpionidaandAcaricouldnotbeidentiedatthefamily
Entomobryomorpha, Hymenoptera, and Diptera, which account for 78% of the total of theindividuals. Entomobryidae and Formicidae were the most abundant
families, with 29% and 25%ofthetotal abundance, respectively.
Accordingtoseasonality,ahigherabundanceofarthropodswasregisteredduringtherainyseason(1,137individuals)comparedwiththedryseason;however,
didnotsignicantlydiffer between seasons(q0; Fig. 1a). Entomobryomorpha was the most abundant order during the dry season (53%), whereas
Hymenoptera and Diptera were the most abundant orders (45% and 28%, respectively) during the rainy season. However, both seasons presented a similar
number of families (53 the rainy seasonand 54 in the dry season). The order with the highest number of families was Araneae (17),followedby
Coleoptera(14), Diptera(9)andHemiptera(8).
Of the total of individuals collected, 25% were larvae, including three orders of holometabolousinsects (Diptera, Coleoptera, and Lepidoptera). Diptera was the
most abundant order with 487individuals from seven families. The Ceratopogonidae family was the most abundant order in thedry season, whereas
Limoniidae was the most abundant one in the rainy season. As for the
ordersColeopteraandLepidoptera,27andtwoindividualswereregistered,respectively.Duringtherainyseason,larvaeweremoreabundantthantheywereint
showed that the diversity of order 0 (observed richness) and order 1 did not present differences betweenseasons (Fig. 1a and 1b). However, order 2 diversity
showed differences between seasons, as the rainyseason showed a greaterdiversity offamilies (q2;Fig. 1c).

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Bromeliad morphology and its relationship with arthropod diversity.The rst and second components of PCA explained the 80% of variance found in
C.
minimiora
. The PC1explained 58% of the variance; all vectors were positive with similar values. This component was interpreted as bromeliad size (Table 1).
Bromeliad size was similar between males and females(
F1,55= 0.607, P= 0.439
) in both the rainy and dry seasons. But the size of bromeliad
individualsvariedbetweenseasons(
F1,55=8.847,P=0.004
). The individuals of
C. minimiora
sampledinthe dry season were larger in comparison to those
individuals sampled in the rainy season (Table 2).The GLM showed that bromeliad size (PC1) affected the richness of families (q0; Table 3). This is, the rainy
season favored the presence of dominant families (Fig. 1c), but it did not affect the plant size (Table 2).
Sex ratio:The sex ratio between male and female plants did not differ according to phorophytetype (
z = 0.635, P = 0.326
), but it differed among transects (
z =
0.206, P = 0.012;
Fig 2a) and betweenseasons (
z = 0.0631, P = 0.0002
). During the dry season, female plants were more abundant
thanmaleplants.Incontrast,maleplantsweremoreabundantduringtherainyseason (Fig.2b).
Density and extrapolation:The transect 1 recorded the highest population density of bromeliads in coffee shrubs and shade trees in dry season. In raining
season, the transect 3 recorded the highest density of bromeliads and arthropods (Table 4).
Discussion
The composition of arthropods associated to
C. minimiora
is like that of other epiphytic bromeliads (Alvarado and Barreno 2010; Arzivenko-Gesing 2008;
Leandro et al. 2014; Ospina-Bautista et al. 2004). We found that Entomobryomorpha was the most abundant order during the dry season; only one family was
registered (Entomobryidae). In contrast, during the rainy season, Hymenoptera was the most abundant order, being Formicidae the most abundant family
(Table S1). Regarding the richness of families, Araneae was the order with the highest value in both seasons. This result aligns with previous ndings that
attribute such richness to spiders’ anity for tridimensional structures such as bromeliad rosettes (Arzivenko-Gesing 2008; Campos-Serrano et al. 2017;
Wittman 2000).
Bromeliads interact with arthropods, such as ants, and spiders, which are sheltered in rosettes for food and reproduction (Bermúdez-Monge and Barrios 2011;
Leandro et al. 2014; Wittman 2000). Also, these arthropods defend bromeliads from herbivory attacks, something similar is thought to occur with coffee
bushes. It has been reported that ants play a signicant role in pest control of coffee crops, such as
Hypothenemus hampei
Ferrari (coffee berry borer), a pest
that damages coffee fruits (Jaramillo et al. 2006; Perfecto and Vandermeer 2006). On its part, spiders are generalist depredators; they consume a great variety
of arthropods including other spiders (Birkhofer and Wolters 2012; Pekár and Toft 2015). Spiders are of great importance, since they contribute to maintaining
low insect populations, which keeps agroecosystems regulated (Riechert and Lawrence 1997; Sarma et al. 2013).
As in other studies, we found that the major abundance and richness of aquatic larvae were classied in Diptera order (Castro 2018; Diaz 2021; Mosquera-
Murillo et al. 2016). The larvae included in this order were successful to inhabit the phytotelma of bromeliads due to their morphological adaptations
(spiracles and siphons), alimentary traits (i.e., feeding of type collecting and lter), reproductive and dispersion capacities (Bermúdez-Monge and Barrios 2011;
Herrera 2003; Mosquera-Murillo et al. 2016; Ospina-Bautista et al. 2004). The families with more abundance in our study were Ceratopogonidae,
Chironomidae, Limoniidae, Syrphidae, and Culicidae. It is important to point that some of these families are considered crops pollinators of cacao (Cazorla-
Perfetti 2014; Kaufmann 1975; Montero-Cedeño et al. 2019), avocado (Castañeda- Vildózola et al. 1999) and mango (Sánchez et al. 2018). However, these
crops do not grow at these altitudes, but they can be considered as possible pollinating agents of the wild plants that inhabit these agroecosystems.
Diaz (2021) found that bromeliad length determined arthropod diversity. Other authors have concluded that the whole structure of bromeliad is of importance
to arthropod diversity (Herrera 2003; Jocque and Field 2014; Nielsen 2011; Ospina-Bautista et al. 2004). We found that bromeliad size affected the richness of
families but did not affect the dominance of families, which was affected by the season (Table3). The rainy season favors the water stored in tanks that later
become a phytotelma (i.e., rich medium), in which larvae of insects, mainly of Diptera develop (Bermúdez-Monge and Barrios 2011; Mosquera-Murillo et al.
2016; Ospina-Bautista et al. 2004).
The phenology of
C. minimiora
has been previously reported: being the owering period from July to January and the fruiting period in December (Martínez-
Correa 2019). In March, a month corresponding to the dry season sampling, we found fresh and dry inorescences of
C. minimiora
. A higher percentage of
female plants (60.8%) over male plants was registered in the dry season. This result coincides with
Catopsis compacta
Mez life cycle, which disperses its
seeds from March to April (Escobedo-Sarti and Mondragón 2016). In contrast, in June (the rainy season), we found that oral buttons and male plants were
more abundant (86.3%). These ndings suggest that
C. minimiora
presents an annual owering of seven months (Martínez-Correa 2019) similar to other
members of the Tillandsioideae subfamily (Gentry 1974).
The different sex ratios found during the owering period are like those of other dioecious owering plants, which produce more male than female plants
(Barrett and Hough 2013; Munguia- Rosas et al. 2011). Male plants invest more resources to produce more owers, and in turn, more pollen grains (Escobedo-
Sarti and Mondragón 2016; Field et al. 2013) by this way increase the probability to pollinate female plants. Forrest (2014) mentions that male plants ower
earlier than female plants since it takes longer to accumulate resources needed at the productive stage.
Although it has been demonstrated that epiphyte removal increases owering and fruit production of coffee bushes (Solís-Montero et al. 2019; Toledo-Aceves
et al. 2012), this practice endangers ora and may affect arthropod diversity. Our estimations suggest that the removal of
C. minimiora
of a coffee plantation
could affect 54,420 arthropods/ha, mainly larvae due to phytotelmata loss. These arthropods could provide different services such as pest control (e.g.,
spiders) and pollination to some plants around coffee plantations (e.g., Ceratopogonidae and Syrphidae).
In conclusion, the shade coffee plantations in Soconusco provide with shelter not only to epiphytes with specialized reproductive systems, such as
C.
minimiora
, but also to arthropods associated to this bromeliad. These arthropods provide different ecosystem services in coffee plantations. Thus, the

Page 5/10
“desmusgue” practice could have a negative effect on the diversity of epiphytes, and in turn, it may has an impact on arthropods associated with them.
Declarations
Competing interests and funding: The authors declare no competing interest. The National Council of Humanities, Sciences and Technologies (CONAHCyT,
acronym in Spanish) granted DJG a scholarship to carry out postgraduate studies (CVU 1102196).
Acknowledgment
To Guillermo Ibarra Núñez and Hector Montaño Moreno for their guidance and support for the taxonomic identication of spiders. To Fernando García
Crisóstomo, for the logistical support in the eld and laboratory. To Martín Roberto Domínguez Fuentes and Erick Antonio Chacón Hartleven, for their advice
and support in the taxonomic identication of the Blattodea group and Diptera larvae, respectively. To Nelson Perez for his permission to sample his coffee
plantation.
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Tables
Table 1.Eigenvectors of two principal components (PC1 and PC2) of PCA of the morphology of
Catopsisminimiora
.
Morphologicaltraits PC1 PC2
Lengthofindividual 0.412 0.594
Diameterofindividual 0.561 -0.487
Numberofrosettes 0.389 0.558
Volumeofindividual 0.603 -0.312

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 
Table 2.Mean ± standard error of variables measured in
Catopsis minimiora.
Variable Transect1 Transect2 Transect3 Transect4 Dryseason Rainyseason
Heightat
phorophyte(m)

1.48±0.18

1.36±0.14

1.31±0.11

1.65±0.08

1.43±0.11

1.48±0.07
Length(cm) 26.05±3.39 23.15±1.14 20.28±0.92 19.60±1.91 24.8±1.97 19.94±1.04
Diameter(cm) 27.79±2.03 33.83±1.86 25.53±1.61 23.67±2.42 30.41±1.49 25.53±1.47
Volume(cm3)5 525±880 7 364±993 4 232±552 4 238±1983 6 322±669 4 235±1010
Numberof 4.75±0.68 3.84±0.58 2.78±0.50 2.07±0.33 4.34±0.45 2.42±0.30
rosettes      
Numberof 20.5±1.04 24.28±1.51 24.12±1.38 20.93±1.62 17.78±0.88 23.03±1.06
leaves      
Volumeof
phytotelma(ml)
0 46.54±5.54 53±11.90 58.38±19.26 46.54±1.24 55.59±1.65
Temperatureof
phytotelma(ºC)
0 19.42±0.12 20.95±0.23 21.66±0.28 19.42±0.08 21.29±0.19

Table 3.Summary of statistics of two generalized linear models (GLM) to analyze the size of
Catopsis minimiora
and season sampled on the arthropod
diversity (the number of Hill). The values in parentheses are the standard errors of the estimate.
Variable Estimate(SE)
Teststatistic(z) P
PC1   
q0 0.918(0.837) 0.026 0.038*
q1 0.405(0.267) 0.134 0.390
q2 0.277(0.220) 0.213 0.638
Season   
q0 -1.354(1.194) 0.226 0.256
q1 -1.578(0.815) 0.058 0.053
q2 -1.485(0.673) 0.031 0.027*

Table 4.Extrapolation of abundance of
Catopsis minimiora
and the number of arthropods per phorophyte type and season.

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Transect Plants/transect
(300m2)
Plantdensity(ind/m2)Plants/ha Arthropods
/plant±S.E.
Arthropods/ha
T1 Bush 33 0.11 1,100 29.17±1.94 32,087
T1 Tree 52 0.17 1,700  35,989
T2 Bush 21 0.07 700 31.92±2.19 22,344
T2 Tree 5 0.01 100  22,344
T3 Bush 30 0.1 1,000 54.42±12.7 54,420
T3 Tree 21 0.07 700  38,094
T4 Bush 10 0.03 300 26.78±1.47 8,034
T4 Tree 13 0.04 400  10,712
   Season  
Season Plants/
season(600m2)
Plantdensity(ind/m2)Plants/ha Arthropods
/Plant±S.E.
Arthropods/ha
Dry  111 0.185 1,850 30.36(0.52) 56,166
Rainy  74 0.123 1,233.33 40.60(2.21) 50,073.19
Figures
Figure 1

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Rarefaction/extrapolation curves based on sampling size with intervals of condence of 95% (shadow areas): a) Richness family (q0), b) exponential of
Shannon (q1), and c) inverse of Simpson (q2). Orange circle (rainy season) and blue triangle (dry season).

Figure 2
a) Sexual ratio of
Catopsis minimiora
per transect and b) per season in a shade coffee plantation. Grey (female) and dark grey (male).
Supplementary Files
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SuplementaryTable1.xlsx