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The Southwestern Naturalist. 2022. 26(2):1-10

FOREIGN SPIDERS AND INSECTS IN ANELOSIMUS CF. ANALYTICUS (ARANEAE:

THERIDIIDAE) NETS IN SONORA DESERT, MÉXICO

JULIO JAVIER GASTÉLUM-REYES, MARÍA CRISTINA PEÑALBA-GARMENDIA, AGUSTÍN FU-

CASTILLO, NARCISO NAVARRO-GÓMEZ, REYNA AMANDA CASTILLO-GÁMEZ, AND ALF
ENRIQUE MELING-LÓPEZ*

Departamento de Investigación Científica y Tecnológica de la Universidad de Sonora

(DICTUS), Hermosillo, Sonora C.P. 83000.
*Correspondent: [email protected]

ABSTRACT-- Spiders are known to be aggressive, but there is a group that is social
and recently found several arthropod species, including spiders, in the spider Anelosimus
nests. We did monthly sampling of spider nests from March 2015 to September 2016 in an
area south of the Sonoran Desert where we observed several arthropod species within the
nests. In this work, we perform monthly sampling of spider nests from March 2015 to
September 2016 in southern part of Sonoran Desert. We analyzed the nests; we counted and
taxonomically identified the arthropod species, and calculated the Shannon-Wiener
Diversity Index and the Sorensen Similarity Index. In total, we analyzed 90 nests in that
period and identified 48 arthropod species, where two species were very abundant, one ant
specie and one spider specie, A. cf. analyticus. We recorded several species from some
spider families, and the Salticidae family had the higher richness. When the ambient
temperature decreased at the sampling site, we observed an increase in both richness and
abundance of arthropods within the nests

RESUMEN—Se conoce a las arañas por ser agresivas, pero existe un grupo que es
social y recientemente se encontró a varias especies de artrópodos incluyendo arañas en
nidos de la Anelosimus. En este trabajo realizamos muestreos mensuales de nidos de la
araña A. cf. analyticus desde marzo 2015 a septiembre 2016 en el sur del Desierto
Sonorense donde observamos varias especies de artrópodos dentro de los nidos. Contamos
e identificamos taxonómicamente las especies de artrópodos y calculamos el Índice de
Diversidad de Shannon-Wiener y el Índice de Similitud de Sorensen. En total analizamos
90 nidos e identificamos 48 especies de artrópodos, donde dos especies fueron muy
abundantes: una hormiga y la araña A. cf. analyticus. Registramos varias especies de
arañas, y la familia Salticidae tuvo la mayor riqueza. Cuando la temperatura ambiental
disminuía en el sitio de muestreo observamos un aumento en la riqueza y abundancia de
artrópodos dentro de estos nidos.

Spiders are aggressive and solitary (Riechert, 1982; Coddington and Levi, 1991;
Foelix, 1996) in contrast there social and subsocial species and one of most studied of these
group is the Genus Anelosimus (Brach, 1977; Furey, 1998; Avilés et al., 2001; Agnarsson
and Kuntner, 2005, Tapia and De Vries, 1980; Lubin and Bilde, 2007; Perkins et al., 2007).
The species is common in tropical and subtropical ecosystems (Agnarsson and Kuntner,
2005). In America, its range extend from East of tropic to Tennessee State (Deyrup et al.
2004, Perkins et al., 2007), and from West side to South America (Agnarsson and Kuntner,
2005). Although there are reports of this Genera in Arizona (Agnarsson, 2004). A. pacificus
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The Southwestern Naturalist. 2022. 26(2):1-10

in western México, Anelosimus arizona, a solitary spider in Sonora and Arizona, and A. cf
analyticus is endemic of southern California and northwestern of México (Levi, 1963). The
genus represented one of the most diverse group of the family Theridiidae with 2516 valid
species and 124 genera (World Spider Catalog, 2019). Most of the social spider came from
Theridiidae family (Agnarsson and Kuntner, 2005; Agnarsson, 2012) where there are a
matriarchal care and work distribution between individuals living on its own web, and
prolonging the group-living phase in subsocial spiders by supplementing colonies with food
(Krafft et al., 1986; Rypstra 1986; Ruttan, 1990; Gundermann et al., 1993; Schneider,
1995; Evans, 1998). Social and sub social behavior are neither common nor rare in spider
literature, but the information is most extent for solitary-living spider species (Agnarsson,
2002). But is common in scientific literature some reports indicating that exist spider
species living together for short time with other spider and insect species (Perkins et al.,
2007) and tolerate other spider species when a catastrophic episode is occurred (Fritzén,
2012). Agnarsson and Kuntner (2005) found two spider species in Africa, one of them of
Genus Anelosimus, where they are found living together too close (at least 1.0 mm
separated each other) in vegetation with a similar spider of the family Agelenidae. Recently
were described a group that tolerate other species of spiders and insects around and inside
the nest (Perkins et al., 2007) this model contrasting the conventional social-spider and sub-
social models also the solitary spider model.
Jones et al. (2007) and Deyrup et al. (2004) presented documentation of the mixed
strategy in both social and communal colony nests of A. studiosus that exhibit foreign
arthropods. The most extended explanation of social and sub-social behavior in spider
species is the cost-benefit hypothesis (Jones and Parker, 2000). It’s cheaper for a spider
group hunting a prey together than in solitary, also construct and repair its web, or take care
of eggs and spiderling (Jones and Parker, 2000; Vakanas and Krafft, 2001). One example
for beneficial foreign nest associates to spiders is beetles and lepidoptera larvae that clean
nests of waste material (Robinson 1977; Furey and Riechert 1989). Spiders that accept or
tolerates a group of foreign species inside its nest (Perkins et al., 2007) have not a well
explanation because the cost-benefit is to cooperative work in social species, but foreign
species can exhibit predator, kleptoparasites or facultative commensal behavior (Robinson,
1977; Furey and Riechert, 1989; Buskirk, 1981). In this paper, we described the annual
variation of arthropods diversity (spiders and insects) inside the Anelosimus cf analyticus
nests, from the northern most subtropical ecosystem closet to Sonoran Desert in México.
Previous observations suggest the spider tolerates both spider and insect foreign species
inside its nests and we focused in the abundance of associated arthropod species into the
nests for a year around. Our interest is to know if the foreign species are present all year
around or for a special season, and if there are certain species more abundant and or
frequency. We considered nests as an arena for mating and brood care, for foraging (Furey
and Riechert, 1989), and a place to be predated for ants or for another specialized spider-
predator. Some arthropod species are present in nests for shelter procurement and for
escape from predators (Robinson, 1977) or rest for short time, though foreign species might
find the same benefits and food also the same risks.

MATERIALS AND METHODS-- Study Site--Our study area is a mix ecosystem in

southern part of Sonora Desert and northern part of subtropical ecosystem. It is a thorny-
spiny shrubby vegetation (Martínez-Yrizar et al., 2010) located in 27°06’10’’N,
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109°29’13’’W. Temperature and rain (Figure 1) from the sampled area are more desert than
subtropical. The nests of the spider Anelosimus cf. analyticus are randomly distributed or
too close to each other, located inside of the thorn-spiny trees and shrubs where they are
protected from wind, rain, and predators, although a few nests are located on high end of
mesquite (specie from Prosopis) branches. The most exposed nests were not present all
year around, only those protected by plants. Most of the nest disappeared during rain and
wind time. Especially, the relative abundance of the nests is present close to agricultural
fence and roads. During the harvested or fumigate time most of nests disappear. Three-
dimension nests apparently are closed, but they are tie at various ends to plants by short
extended webs, the nests size ranged 64-500 cm3. We collected five-ten nests (90 nests in
total) of the A. cf. analyticus monthly from March 2015 to September 2016 (except in July)
and we had the opportunity to sample nests in January 2017. Each nest was placed inside of
the hermetic plastic bag and it were put in an ice container to be transported to laboratory
where all the nests were put together in freezer container to avoid possible predation. We
dissected each nest by hand and used stereo microscopy to pick all organisms through the
silk, also we separated the individual by group (spider, insect and others arthropods) and
taxonomically identified to genus or at least to family (Ubick et al., 2017; Borror and
White, 1998), and preserved in 95% ethanol for further genetic analysis. We also sexed all
spider individuals, we counted all moults from the nests. All nests were measured to know
if larger nests contain a more genus-rich or numerically abundant spider species, we define
genus richness as the number of heterospecific spider genera, which inhabit Anelosimus,
nests (Pruitt and Riechert, 2011), the same criteria we used for insect families.
Data analysis--All individuals were accounted for estimating number of the species
and its abundances, also to know if one or more foreign species are timing present. We used
the Sorensen Similarity Index (Ss = 2C/(a + b). C, means a common species; a, the specie
in sample or month a; b, species in sample or month b. The limit between differences and
not differences is 0.5) to determined similarities or differences between months (Magurran,
1988). We also used the Shannon-Weaver Diversity Index (H’) to determined variation in
species during the time sample period. We used analysis of variance to determine
differences between number of species, abundances and H’ during the sampled period, also
for three-month periods (Sokal and Rohlf, 1995; Walker y Lev 1969; Zar, 1984). We
consider three-month period as enough time that most species of spiders and insects
completed its life cycle as an adult. We used the Pearson correlation coefficient (Zar, 1984)
to determine the relation between nest size and abundance of spiders and insects,
temperature and rain. Statistical analyses were using Sigma Plot 11.0 and 95% of
confidence.

RESULTS--Temperature and rainfall showed a tendency to decrease values over time

(Figure 1), and all the biological variables to raise their values over time except the
abundance of insects (Figure 2) but there was no statistical relationship (P > 0.05). A total
of 1,062 individual arthropods was observed in the 90 collected nests, including 17
heterospecific spiders of 8 families (Table 1) and 31 insect species of 15 Orders (Table 2),
two of those spiders appeared only in January 2017. All the nests had arthropods, 78 of
them with spiders of A. cf. analyticus, 8 with only other spider species but not Anelosimus,
and 4 with no spiders but other arthropods, and a nest with spider ovisac. Most of the time
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there were more spiders than insects. The most abundant spider species was A. cf.
analyticus with 357 individuals (105 females, 41 males, and 211 juveniles), representing
more than 80% of all spiders. Neon sp. and Habronattus sp. were the second most abundant
also with females, males and juveniles (Table 3). Both spiders together with Misumena sp.
were the most frequent spiders but it was the only spider present during three consecutive
months. Coleoptera (5 species) and Hemiptera (8 species) were the most frequent Orders,
but the ants were the species more frequent and abundant and they were present for five and
four consecutive months. November 2015 was the most insect abundance with 316 and 6
insect species, three spider species (A. cf. analyticus, Mimetus sp and Neon sp.) with 34
individuals. December was the month with more insect species (10 spp) but 24 individuals,
three spider species (A. cf. analyticus, Misumena sp. and Pelegrina furcata).
Nest Size Analysis--The species was present during all around the samplings, many
times sharing the nests with other species of spiders and insects. There was no relation with
nest size and richness or abundance (P > 0.05). In November 2015, we collected a large-
size nest with more than 500 ants, four Anelosimus spiders and a Neon individual. In June
2016, we found a large-size nest with 25 Anelosimus spiders with two Thiodina spiders.
During May 2016 we found one medium-size nest with 57 ants, one Coleoptera sp 2, four
Neon, one Anelosimus spiders, and one ovisac; also, in the same date, we collected a
medium-size nest with eight arthropod species (27 individuals) and three spider species
(seven individuals) and five ovisacs. The spider moults were frequent during the study
period. In a medium-size nest collected in April 2016, we found two moults, one ovisac and
five spider species: one male of Anelosimus, and females of Neon, Synageles,
Chalcoscirtus, Peckhamia, all of them with only one individual, also the same nest had one
insect species with four individuals. Before rain during May 2015 and 2016, the ovisacs
were abundant. In a medium-size nest during July 2015, we found three moults from
Anelosimus, but we not found spiders only ants and wasps. In September 2016, we found
the medium-size nest with more Anelosimus individuals. In December 2015, we found a
medium-size nest with more insect species (10) and one Anelosimus spider, and in June
2016 one small-size single nests had 12 females and six males.
Three Months Analysis--From March 2015 to May 2015 we found 11 species, four
of those were spiders including A. cf. analyticus, but from December 2015 to February
2016 from 21 species, eight were spiders. Anelosimus was accompanied with a specie from
Family Tetragnathidae from March 2015 to June 2015, and until November 2015 with
Neon, other species were present but do not remain continuous over time (e.g., Synageles
sp.). From September 2015 to August 2016 we recorded Misumena sp., and from December
2015 to August 2016 we recorded Chalcoscirtus diminutus and Habronattus sp. Ants were
present from June 2015 to August to 2016.
Diversity (H’) Analysis-- There were statistical differences (P > 0.05) in both
abundances and species richness during the study period. The Similitude Index (Figure 3)
showed low values (<0.5) between two or three consecutive samplings in species richness.
The Shannon-Wiener Index was low (H’ = 0.71 vs H’Max = 1.68), but in April 2016 it was
high (H’ = 1.0 vs H’Max = 1.2). The analysis of three month showed statistic differences in
the H´, increasing from March 2015 to February 2016 and declined from March 2016 to
August 2016. The number of species increased from 11 (six insects and four spiders) in
March 2015 to 25 (18 insects and 7 spiders) in May 2016, then decrease to 19 (12 insects
and 7 spiders) in August 2016. The period with more spider was decembre-2015 February
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2016 with 21 (13 insects and eight spiders, five spider species from the anterior period).
Also, the H’ was higher (0.939, H’Max =1.32) in this period. There was no relationship
between size nest and abundance or number of species (P > 0.05). But there was significate
correlation with H’ and low temperature (r = -0.65, P < 0.05), also the species of both
insect and spider were increasing when temperature was decreasing from August 2015 to
May 2016 (r = 0.67, P < 0.05).

DISCUSSION--Social and sub-social behavior in Anelosimus species are common in

America tropical and subtropical ecosystems (Agnarsson and Kuntner, 2005; Agnarsson,
2006), but to north, e.g., California, Arizona and northwestern part of Mexico, it seems
Anelosimus lives as solitary species (Levi, 1963; Agnarsson, 2004). We categorized A. cf.
analyticus as a subsocial species most of the time, but it also showed solitary behavior
during May 2015 in a few nests, we did not observe other spider species nor spiderlings in
those nests but three insects of three species. A. pacificus show a similar behavior, the
spider is solitary almost all its lifespan with the juveniles passively receiving maternal care
before dispersal at an early instar (Agnarsson et al. 2010). We observed most of the time
several insects and spiders species inside of A. cf. analyticus’ nests. The Sorensen Index
always were low between two or three consecutive months, showed no similarities between
species scenarios, but the species richness showed a tendency to increasing through time
(Figure 2), for this reason the community inside the nests were changing each time but A.
cf. analyticus was always present and was the most abundant specie.
The species of both insect and spider were increasing when temperature was
decreasing, also abundances were increasing from August 2015 to May 2016 (r = 0.67, P <
0.05), it means that predatory spiders were on the rise as insect prey increased. Despite the
constant changes in richness and abundances, H’ was always low although there was a
slight increase (statistically not significant) but the general tendency was to decrease. There
was a lot of activity in the nests throughout the sampled study. Each three-month periods
the insect-spider groups were changing each other, Anelosimus was always present. We
found more than one species in each nest, we also recorded evidence of several spider
species used the nest for periods of two consecutive months, and for molting in presence of
Anelosimus and insects but only Misumena (Thomisidae) was found for three consecutive
months in the nests. Perkins et al. (2007) found same spider families we found, e.g.,
Salticidae and Tetragnathidae and others in nests from A. studiosus. Deyrup et al. (2004)
found several insect species inside the Anelosimus nests. Spiders and insects interacting in
the same arena could indicate they use the nests for different activities, e.g., protection,
predator scape or feeding. Most of the time we recorded 2-3 associated spider species in
each nest and in several cases together with insect species. Ants were present almost all the
time (5% of the total species but 75% of abundance), the H’ down in November was
because ants had high abundance. Studies of social spiders suggest that ant can reduce the
colony size, or it can extinguish a colony (Henschel 1998; Purcell and Avilés 2008) or
increasing ants’ abundance may exclude Anelosimus from its habitat by predation or
disturbance (Purcell and Avilés 2008). Insect and spider richness were increasing during
study period and this could represent advantage for A. cf. analyticus, e.g., jumping spider,
specialized in hunting ants, it could represent protection against ant predation for
Anelosimus in the studied nests, Purcell and Avilés (2008) suggest that jumping spiders
may not threaten the survival of Anelosimus spiders in the lowland rainforest. The spider
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group and ants could represent predators competing the niche. The other insects could use
the nest looking for feed, rest or escape safely, but the nest may represent a trap to die or a
safe place to live. It could indicate that spider’s species protect each other, protecting
themselves in specific places of the nest (Perkins et al. 2007). The diversity of species in
the nests give Anelosimus survival advantages. Foreign species could decrease the
predation mortality rate, and maybe for this reason Anelosimus is present all year around in
its own nests. Most of the time the richness of spiders increased one month after it fell. We
conclude that Anelosimus is a subsocial species due to the presence of some females, males
and juveniles in the same nest, and most of the time we found nests with more than one A.
cf. analyticus female spider. We categorized the other spiders as opportunistic species due
they were present inside the nest part of the time and their abundances were low.

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34 220

32 200

30 180

160
28
140
26
Temperature

120
24

Rain
100
22
80
20
60
18 40

16 20

14 0
M'2015 May Jul Sep Nov J'2016 Mar May
FIG. 1--Temperature (°C) and rainfall (mm) values from the sampled area during de
study. Peaks of maximum temperature and rainfall coincide in summer.
12 350
)

300
10

)
insect

insect
250
8

200
Species Richness (spider

6
Abundances (spider

150

4
100

2
50

0 0
M'2015 May Jul Sep Nov J'2016 Mar May
FIG. 2--Species richness and species abundances. Trends in richness and abundance
increase but insect abundance decreases
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1.4

1.2
)
H'max

1.0

0.8
Diversity Index (H'

0.6

0.4

0.2

0.0
M'2015 May Jul Sep Nov Jan'2016 Mar May
FIG. 3--Values of Diversity Index (H’) and maximum Diversity Index (H’Max).
During November the H’ was low due to the high abundance of ants.
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TABLE 1--Monthly and total spider abundances found inside nests of Anelosimus cf analyticus, during 2015-2016 period
and January 2017.
2015 2016 2017
Especies Mar Abr May Jun Jul Agt Sep Oct Nov Dic Jan Feb Mar Abr May Jun Agt Sep Jan Total
Anelosimus cf analyticus 11 7 10 37 4 20 2 14 30 28 14 8 9 10 19 49 3 59 23 357
Messua sp 1 1
Chalcoscirtus diminutus 1 1 2 1 5
Thiodina hespera 3 1 3 7
Misumena sp 1 1 1 1 2 1 7
Pelegrina furcata 1 2 3
Habronattus sp 4 1 7 1 1 14
Attidops cutleri 2 2
Peckhamia sp 2 1 1 1 5
Salticus sp 1 1
Synageles sp 1 2 2 1 3 9
Kibramoa sp 2 2
Scytodes sp 3 3
Mimetus sp 1 3 2 6
Neon sp 3 1 3 1 2 10 20
Sparassidae sp1 1 1
Tetragnathidae sp1 1 1 2
Monthly Abundance 13 11 10 41 5 22 7 19 34 30 21 14 11 18 31 61 4 62 30 444
Spider Richness 2 3 1 3 2 3 3 4 3 3 4 6 2 7 3 5 2 4 5
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TABLE 2--Spider names, females, males, juveniles, and number of nests where the
species were registered.
Spider Name Total Females Males Juveniles Nests
Anelosimus cf analyticus 357 105 41 211 78
Neon sp 20 14 4 2 12
Habronattus sp 14 8 2 4 12
Mimetus sp 6 3 1 2 4
Peckhamia sp 4 1 3 0 4
Synageles sp 9 7 0 2 5
Chalcoscirtus diminutus 5 4 1 0 4
Misumena sp 7 6 0 1 7
Thiodina hespera 7 6 0 1 6
Pelegrina furcata 3 3 0 0 3
Kibramoa sp 2 1 0 1 2
Scytodes sp 3 3 0 0 2
Tetragnathidae sp1 2 2 0 0 2
Sparassidae sp1 1 1 0 0 1
Attidops cutleri 2 0 0 2 1
Salticus sp 1 0 0 1 1
Messua sp 1 0 1 0 1
Total 444 164 53 227 ---
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The Southwestern Naturalist. 2022. 26(2):1-10

TABLE 3--Monthly and total insect abundances found inside nest of Anelosimus cf analyticus during 2015-2016 period.
Spp is the number of species from each Order.
2015 2016
Especies Mar Abr May Jun Jul Agt Sep Oct Nov Dic Jan Feb Mar Abr May Jun Agt Sep Total
Coleoptera (5 Spp) 1 2 1 2 2 1 1 3 1 3 2 2 3 24
Hemiptera (8 Spp) 1 3 1 1 6 10 4 7 6 2 2 5 3 51
Diptera sp 1 1 1 1 2 5
Diptera sp 2 1 1
Diptera sp 3 3 3
Formicidae sp 1 2 4 2 1 302 1 4 8 10 90 3 5 432
Formicidae sp 2 4 4
Psocoptera 2 1 3
Hymenoptera sp 1 2 1 2 1 4 10
Hymenoptera sp 2 1 3 4
Vespidae 3 3
Lepidoptera 2 2
Dermaptera 1 1
Neuroptera 3 5 8
Coccidae 3 6 4 4 17
Colembola 3 4 1 8
Cicadelidae 2 5 7
Orthoptera 3 1 6 1 11
Miridae 2 1 3
Monthly Abundance 2 5 3 7 8 11 5 14 316 24 12 4 23 31 105 10 6 11 598
Insect Richness 2 3 3 4 4 4 3 6 6 10 4 3 6 9 9 8 4 4 30
 TABLE 2--Spider names, females, males, juveniles, and number of nests where the
species were registered.
Spider Name Total Females Males Juveniles Nests
Anelosimus cf analyticus 357 105 41 211 78
Neon sp 20 14 4 2 12
Habronattus sp 14 8 2 4 12
Mimetus sp 6 3 1 2 4
Peckhamia sp 4 1 3 0 4
Synageles sp 9 7 0 2 5
Chalcoscirtus diminutus 5 4 1 0 4
Misumena sp 7 6 0 1 7
Thiodina hespera 7 6 0 1 6
Pelegrina furcata 3 3 0 0 3
Kibramoa sp 2 1 0 1 2
Scytodes sp 3 3 0 0 2
Tetragnathidae sp1 2 2 0 0 2
Sparassidae sp1 1 1 0 0 1
Attidops cutleri 2 0 0 2 1
Salticus sp 1 0 0 1 1
Messua sp 1 0 1 0 1
Total 444 164 53 227 ---