Diet composition and prey selectivity

of Colombian populations of a social

pseudoscorpion

L. F. Garcia, J. C. Gonzalez-Gomez,
J. C. Valenzuela-Rojas, E. Tizo-Pedroso
& M. Lacava

Insectes Sociaux
International Journal for the Study of
Social Arthropods

ISSN 0020-1812

Insect. Soc.
DOI 10.1007/s00040-016-0505-z

1 23
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 Author's personal copy
Insect. Soc.
DOI 10.1007/s00040-016-0505-z Insectes Sociaux
SHORT COMMUNICATION

Diet composition and prey selectivity of Colombian populations

of a social pseudoscorpion
L. F. Garcia1,4,5 • J. C. Gonzalez-Gomez2,3,5 • J. C. Valenzuela-Rojas2,5 •
E. Tizo-Pedroso5,6 • M. Lacava4,5

Received: 29 October 2015 / Revised: 21 July 2016 / Accepted: 19 August 2016

Ó International Union for the Study of Social Insects (IUSSI) 2016

Abstract Prey capture is a determinant aspect in arachnid potentially dangerous species which possess stingers or
sociality, since cooperative hunting allows the individuals behaved aggressively. When evaluating size, we found that
access to very large prey. Although this behavior has been P. nidificator is able to prey on arthropods up to eight times
extensively studied in some spiders, the knowledge about larger than itself. These results suggest that by hunting in
some minor groups such as pseudoscorpions is still scarce. groups, P. nidificator is able to capture not only larger prey,
The aim of this study was to analyze diet composition, prey but also prey which could be potentially dangerous for
selectivity and prey size in Colombian populations of the solitary individuals such as some ant species.
social pseudoscorpion Paratemnoides nidificator. We col-
lected prey captured by 112 colonies of P. nidificator as well Keywords Social behavior
Feeding habits
Foraging

as their potential prey. We found that hymenopterans Pseudoscorpiones
Atemnidae
(Formicidae) were the most consumed prey, while other
items found in the diet included dipterans, embiopterans,
hemipterans, mantids, neuropterans and spiders. When Introduction
comparing prey composition, we found a low similarity
between actual and potential prey. Paratemnoides nidifi- Sociality is an uncommon phenomenon among arachnids
cator showed a positive selectivity towards ant species (Avilés 1997). While degrees of sociality have been repor-
which lacked strong defensive mechanisms, while avoiding ted among different arachnids (Choe and Crespi 1997; Costa
2006), the most extreme cases of sociality occur among
spiders and pseudoscorpions, where examples of species
& E. Tizo-Pedroso that cooperate for prey capture and breeding can be found
tizopedroso@ueg.br (Avilés 1997; Ainsworth et al. 2002; Del-Claro and Tizo-
1
Pedroso 2009; Settepani et al. 2013; Yip and Rayor 2014).
Centro Universitario de la Región Este, Universidad de la
República, Simón del Pino 1132, Treinta y Tres, Uruguay
In spiders, food is a critical determinant of sociality, as
2
group number is regulated by prey availability and size
Universidad Surcolombiana, Semillero INVUSCO, Avenida
Pastrana Borrero-Carrera 1, Neiva, Huila, Colombia
(Whitehouse and Lubin 2005; Lubin and Bilde 2007;
3
Guevara et al. 2011). By hunting in groups, spiders are able
Grupo de Investigación en Zoologı́a, Universidad del Tolima,
Barrio Santa Helena Parte Alta, Ibagué, Tolima, Colombia
to capture prey which otherwise would be unavailable for
4
solitary individuals (Majer et al. 2015). This includes larger
Laboratorio de Ecologı́a del Comportamiento, Instituto de
Investigaciones Biológicas Clemente Estable, Avenida Italia
and more varied prey including those which may be
3318, Montevideo, Uruguay potentially harmful to the spiders, such as ants, wasps and
5 bees (Souza et al. 2007). A constant supply of food and the
Grupo de Investigación BEA, Corporación Huiltur, Carrera 5
No 8-67, Neiva, Huila, Colombia consumption of large prey are necessary for the mainte-
6 nance of a large group size (Giraldeau and Caraco 2000).
Universidade Estadual de Goiás, Câmpus Morrinhos,
Laboratório de Ecologia Comportamental de Aracnı́deos, For this reason, social spiders are predominantly found
Goiás, Brazil when prey richness is high or where insects several times

123
 Author's personal copy
larger than the spiders are frequent (Guevara and Aviles
2009; Majer et al. 2013a, b, 2015).
The Order Pseudoscorpiones is a diverse group of
Arachnids, with more than 3300 known species (Harvey
2011). Sociality in this group is poorly known compared to
spiders (Del-Claro and Tizo-Pedroso 2009). Previous stud-
ies on the diet and feeding behavior of social
pseudoscorpions have focused on the Northern and Central
American pseudoscorpion Paratemnoides elongatus (Banks
1895) (Atemnidae), and showed that this species can feed on
arthropods several times larger than adult individuals and
can capture these cooperatively (Brach 1978; Zeh and Zeh
1990). In South America, studies based on the ecology and
natural history of the cooperative species P. nidificator
(Balzan 1888), a pseudoscorpion commonly found under
the bark of living trees and which forms groups of up to 300
individuals, showed that Brazilian populations of this spe-
cies also feed on large ants and other large insects such as
stink bugs and beetles (Tizo-Pedroso and Del-Claro 2007).
Additional studies about prey capture under laboratory
conditions found that adults are mainly responsible for
providing food for the colony (Tizo-Pedroso and Del-Claro
2007, 2011). Despite this, there are no detailed studies
describing prey selection by these pseudoscorpions. Prey
selectivity is an important aspect, as it allows us to unveil
the trophic specialization and importance of different prey
types on the life history of an organism (Martin and
Wainwright 2011; Litsios et al. 2012). Considering the food
demands of pseudoscorpion colonies (Tizo-Pedroso and
Del-Claro 2007; Del-Claro and Tizo-Pedroso 2009), we
expect that the colonies preferentially capture large prey.
Therefore, the aim of this study was to determine the diet
composition and prey size selectivity of Colombian popu-
lations of the social pseudoscorpion, P. nidificator.
According to previous studies, social pseudoscorpions are
able to capture animals larger than themselves (Brach 1978;
Zeh and Zeh 1990; Hahn and Matthiesen 1993; Tizo-Pe-
droso and Del-Claro 2007), so we expected P. nidificator to
consume a range of large crawling arthropods such as ants
and bugs.
Methods
Prey composition and size
We analyzed the prey composition by evaluating the con-
sumed prey found associated with multiple colonies of P.
nidificator that were found in a cloud forest next to Neiva
city, Colombia. Since P. nidificator individuals are often
found associated with rough bark trees (Tizo-Pedroso and
Del-Claro 2007), we sampled colonies found between 0 and
3 m height in ten trees of Pseudosamanea guachapele
123
L. F. Garcı́a et al.
(Kunth) and 20 trees of Pithecellobium dulce (Roxb) Benth
(Fabaceae, Mimosoideae). These samplings were made by
four researchers between February and May 2014 over 20
non-consecutive nights, and were ended after completing 20
sampling hours for each collector. Since pseudoscorpions
are nocturnally active, all sampling was done between 20:00
and 22:00. We chose this time period because P. nidificator
individuals can be found outside their nests at this time.
During sampling, we collected only colonies which we
found to be consuming a prey. When we found pseu-
doscorpions feeding, the prey was removed and identified to
the lowest possible taxonomic level. Additionally, obser-
vations were made ad libitum when interactions between
pseudoscorpions and their prey were found in the field.
After collecting the prey, most of the pseudoscorpions hid
inside their shelters so it was not possible to collect all the
living individuals to measure them or determine how many
of them participated in prey capture. Nevertheless, in all
cases only adult pseudoscorpions were found capturing the
prey, therefore, we collected 30 adult individuals to estimate
their mean body size. The voucher specimens were depos-
ited in the entomological collection of South Colombian
University (Universidad Surcolombiana). Pseudoscorpions
feed on prey’s internal tissues leaving the exoskeleton
almost intact (Tizo-Pedroso and Del-Claro 2007). We,
therefore, photographed adult pseudoscorpions and
exoskeletons of consumed prey under a dissecting micro-
scope. The measurements of both prey and pseudoscorpions
(total body length) were made using the software imageJ
1.46r (Rasband 2015).
To identify the potential prey, we collected all arthropods
that we found crawling on trees, inside crevices and on tree
barks. Samplings were also made between 0 and 8 m in
height on the tree trunks during the same time lapse and with
the same effort per collector used to sample the actual prey.
Because some pseudoscorpions were found consuming
flying arthropods, we decided to sample all the vegetation
found 5 m around the trees using 500 sweep net passes per
sampling to collect flying insects, which climb up trees. The
collected individuals were placed in vials of ethanol (70 %).
Preserved individuals were placed in the entomological
collection of the Universidad Surcolombiana. We also used
the Morisita index to compare species composition between
actual and potential prey (Krebs 1999).
Prey selectivity
Prey selectivity was measured using a Manly index, which
evaluates selectivity as the ratio between proportion for
each item in the diet divided by the proportion for each item
on the environment (Manly et al. 2002). When the index
value is significantly larger than one it indicates a preference
for a certain prey item. In contrast, when the value is less
 Author's personal copy
Diet composition and prey selectivity of Colombian populations of a social pseudoscorpion

Table 1 Prey available in the environment and prey consumed by Paratemnoides nidificator
Prey Common name Actual Available Size for actual prey (mm)

Acromyrmex octospinosus Leaf-cutting ant 40 68 6.69 ± 0.29

Camponotus cf. brevis Carpenter ant 31 28 6.88 ± 0.28
Camponotus sp. Carpenter ant 18 74 5.89 ± 0.22
Camponotus cf. substitutus Carpenter ant 7 106 7.85 ± 1.36
Coreidae Leaf-footed bugs 4 25 12.03 ± 2.90
Pseudomyrmex sp. Elongate twig ant 4 1 6.93 ± 0.88
Crematogaster sp. Acrobat ant 3 40 6.88 ± 0.28
Ectatomma tuberculatum Selva ant 3 40 6.93 ± 1.09
Odontomachus sp.* Trap-jaw ant 2 18 2.87
Chrysopa sp.* Lacewing 2 6 7.27
Schizocosa sp.* Wolf spider 2 4 6.84
Mantoida sp.* Praying mantis 2 2 22.09
Embioptera* Foot spinner 2 1 8.94
Sciaridae* Dark-winged fungus gnat 2 1 3.03
Total 122 414
The sizes for actual prey are represented as the mean ± standard error. (* Indicates prey where only one individual could be measured)

than one, it indicates avoidance. Values near to one indicate

that the resource is used proportionally to its availability
(Desbiez et al. 2009). The statistical significance for this
index was analyzed using a Chi-square test with a sequential
Bonferroni correction for critical values (a = 0.003). Sta-
tistical analyses were performed with the software R for
Windows 3.1.1 (R Core Team 2015) and the package ade-
habitat (Calenge 2006).

Results

Diet composition and prey size

We recorded a total of 112 colonies consuming prey. All the
observed prey had been captured by adult pseudoscorpions
and were in good condition (i.e., slightly deteriorated) by
the time they were collected. Among the prey we found,
hymenopterans (family Formicidae) were the most abun-
dant followed by hemipterans, mantids, neuropterans and
spiders (Table 1). Since several morphospecies of ants were
recorded in comparison to the other groups, ants were
identified to a lower taxonomic level compared to the other
prey. The species composition was different between
available and consumed prey and exhibited a similarity of
3 % according to the Morisita index.
The rank sizes for consumed prey were most frequent
between 6 and 8 mm (Fig. 1). A gradual decrease in fre-
quency is observed as prey size increases. Paratemnoides
nidificator rarely captured prey bigger than 16 mm. The
mean size recorded for P. nidificator adults is
2.7 ± 0.2 mm, therefore, this suggests that these
Fig. 1 Histogram for the size of the prey captured by individuals of
Paratemnoides nidificator
pseudoscorpions are able to capture prey up to eight times
the size of an adult individual.
Prey selectivity
When analyzing the prey selection in P. nidificator, we
found that these show a marked selectivity for different
arthropods (v213 = 127.98, p \ 0.0001). After analyzing the
selectivity index (Wi), we found that when significant, both
positive and negative significant values were recorded in
ants (Fig. 2). Positive selectivity was registered in Acro-
myrmex octospinosus (Wi = 1.99, p \ 0.0001) and
Camponotus cf. brevis (Wi = 3.75, p \ 0.0001). In con-
trast, negative selectivity was observed in Camponotus cf.
substitutus (Wi = 0.22, p \ 0.0001), Crematogaster sp.

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 Author's personal copy
Fig. 2 Manly selectivity index
for the prey captured by
Paratemnoides nidificator.
Values above the horizontal line
indicate positive selectivity,
while values below it indicate
negative selectivity. Error bars
represent 95 % confidence
interval. (*Indicates prey which
are consumed in a significantly
different proportion when
compared to their availability)
(Wi = 0.25, p \ 0.0001) and Ectatomma tuberculatum
(Wi = 0.25, p \ 0.0001). No significant selectivity was
found for other arthropod groups such as Coreidae
(Wi = 0.56, p = 0.087), Mantoida sp. (Wi = 3.93,
p = 0.87), Chrysopa sp. (Wi = 1.13, p = 0.32), Embiop-
tera (Wi = 6.78, p = 0.22), Schizocosa sp. (Wi = 1.19,
p = 0.56) and Sciaridae (Wi = 6.78, p = 0.22). A similar
tendency was observed in some ant species such as Cam-
ponotus sp. (Wi = 0.86, p = 0.331), Odontomachus sp.
(Wi = 0.38, p = 0.018) and Pseudomyrmex sp.
(Wi = 13.57, p = 0.060).
Discussion
Our study provides the first quantitative description of prey
selectivity in pseudoscorpions under field conditions. In
spite of having a wide variety of available prey, pseu-
doscorpions showed a marked consumption for certain prey
groups, which is reflected by the low similarity between the
actual and the potential prey. The mantids and embiopterans
are reported here for the first time in the diet of P. nidifi-
cator, nevertheless, their consumption was low compared to
other prey such as ants and true bugs, which had been
previously recorded in Brazilian populations (Tizo-Pedroso
and Del-Claro 2007). The high consumption of ants and
bugs may be explained by the fact that these insects walk
actively on the tree bark, where pseudoscorpions build their
nests. In contrast, the low frequency of other prey items may
be because they were only temporarily present on the tree,
such as some flying insects, or because they do not move
actively on these, such as some spiders. These results agree
with previous findings by Tizo-Pedroso and Del-Claro
(2007), who report ants as the most important prey in
123
L. F. Garcı́a et al.
Brazilian populations of P. nidificator during the dry season.
We, however, surveyed for only a short period, so the diet
could change across seasons (Tizo-Pedroso and Del-Claro
2007).
Although there is limited information about prey size in
pseudoscorpions, especially for solitary species, most
studies report arthropods smaller or slightly larger than the
pseudoscorpions, including mites, collembolans and fruit
flies (Weygoldt 1969; Harvey 1986; Witt and Dill 1996).
Experiments of prey size preference show that solitary
pseudoscorpions prefer individuals which measure about
50 % of their own body length (Johnson and Wellington
1980), while social species like the Brazilian populations of
P. nidificator have been recorded consuming prey four
times larger than adult individuals (Tizo-Pedroso and Del-
Claro 2007). The P. nidificator Colombian populations
caught prey up to eight times their size, although the most
frequent prey captured were between approximately 1.8 and
3 times the body size of an adult pseudoscorpion.
Sociality allowed the pseudoscorpions to catch and use
prey several times larger than themselves, which otherwise
would be very difficult to capture. In North America, the
social pseudoscorpion P. elongatus is capable of capturing
large ants, beetles and millipedes (Brach 1978; Zeh and Zeh
1990). The capture of larger prey is a common trait in social
arachnids: for example, it has been shown that social spiders
are able to capture prey up to 10–20 times larger than an
adult individual (Nentwig and Wissel 1986). Consumption
of large prey by P. nidificator (prey larger than the pseu-
doscorpions) was confirmed by this study.
The selectivity values for the majority of prey were not
significant, suggesting that most of prey were consumed
according to their availability. Nevertheless, the positive
selectivity values recorded for some species such as A.
 Author's personal copy
Diet composition and prey selectivity of Colombian populations of a social pseudoscorpion
octospinosus (a leaf cutter ant) and Camponotus cf. brevis,
indicates that P. nidificator has a marked preference for
these prey. The consumption of leaf cutting ants was
reported for another pseudoscorpion species, Paratem-
noides elongatus (Zeh and Zeh 1990), but not for P.
nidificator. The preference for this ant species could be
explained by its lack of any defensive mechanisms, such as
stingers or secretions, rendering it more profitable.
Although we observed that Camponotus cf. brevis was able
to produce defensive secretions, it is smaller and less
aggressive than other Camponotus and other ant species
found in the zone, such as Ectatomma sp., Odontomachus
sp. or Pseudomyrmex sp., turning it into a more accessible
prey and explaining the positive selectivity recorded for this
species. Since the ant A. octospinosus consume a wide
variety of plants, including the fruits and shrubs and may
nest in the basement of some trees (Wetterer et al. 1998),
these might expose the ants more frequently to P. nidificator
favoring its consumption. A similar pattern is found for
Camponotus cf. brevis, which builds its nests in the tree bark
like other ants of this genus (Hansen and Klotz 2005).
The negative selectivity recorded for some ants might be
explained because they have strong defenses, or are par-
ticularly large. For example, the avoidance of the ant
Camponotus cf. substitutus may be due to the aggressive
behavior of this ant, since the authors observed it attacking
several insects. This species also exceeds the mean size for
prey captured by P. nidificator. A similar pattern occurs for
the ant Ectatomma tuberculatum, which not only possesses
a large size and a powerful stinger, but also is very
aggressive, being able to prey even on other ant species
(Tofolo et al. 2011). Although the ants Crematogaster sp.
are smaller than other species, these are very aggressive and
perform mass attacks, being a dominant species in trees
(Tschinkel 2002), which might turn them into a dangerous
prey for P. nidificator.
Although diet analysis alone cannot reveal the full
trophic strategy of an organism, it provides an initial insight
of its feeding ecology. Accordingly, our results indicate that
P. nidificator acts as an euryphagous predator, confirming
previous observations for this species. Besides that, it
selectively feeds on some ant species such as Camponotus
cf. brevis and A. octospinosus, suggesting that both species
are an important part of its diet. By foraging in groups, P.
nidificator is able to capture prey that are potentially dan-
gerous for solitary individuals.
Acknowledgments We thank Lorena Cardozo, Camilo Moncayo and
Elı́as Amorteguı́ for their help during the fieldwork. Stano Pekár, José
A. Hódar and Carmen Viera provided us with several suggestions and
literature to write this manuscript. We thank to Paola González
Vanegas, who kindly helped us with the insect identification. We also
thank to Antonio Agudelo who helped us to identify the mantid spe-
cies. Volker Manhert helped us with pseudoscorpion
identification. Sean Blamires, Karen Greenhalgh de Guzmán and
David Guzmán, greatly helped us with the English correction for the
manuscript. We are also indebted to two anonymous referees who
greatly improved this manuscript with their suggestions.
References
Ainsworth C, Slotow R, Crouch T, Lubin Y (2002) Lack of task
differentiation during prey capture in the group living spider
Stegodyphus mimosarum (Araneae, Eresidae). J Arachnol
30:39–46
Avilés L (1997) Causes and consequences of cooperation and
permanent-sociality in spiders. In: Choe JC, Crespi BJ (eds)
The evolution of social behavior in insects and arachnids.
Cambridge University Press, Cambridge, pp 476–498
Brach V (1978) Social behavior in the pseudoscorpion Paratemnus
elongatus (Pseudoscorpionida: Atemnidae). Insectes Soc 25:3–11
Calenge C (2006) The package adehabitat for the R software: a tool for
the analysis of space and habitat use by animals. Ecol Model
197:1516–1519
Choe JC, Crespi BJ (1997) The evolution of social behaviour in insects
and arachnids. Cambridge University Press, Cambridge
Costa JT (2006) The other insect societies. Belknap Press of Harvard
University Press, Cambridge
Del-Claro K, Tizo-Pedroso E (2009) Ecological and evolutionary
pathways of social behavior in Pseudoscorpions (Arachnida:
Pseudoscorpiones). Acta Ethol 12:13–22
Desbiez LAJ, Bodmer RE, Santos SA (2009) Wildlife habitat selection
and sustainable resources management in a Neotropical wetland.
Int J Biodivers Conserv 1:11–20
Giraldeau L-A, Caraco T (2000) Social foraging theory. Princeton
University Press, Chichester
Guevara J, Aviles L (2009) Elevational changes in the composition of
insects and other terrestrial arthropods at tropical latitudes: a
comparison of multiple sampling methods and social spider diets.
Insect Conserv Diver 2:142–152
Guevara J, Gonzaga MO, Vasconcellos-Neto J, Aviles L (2011)
Sociality and resource use: insights from a community of social
spiders in Brazil. Behav Ecol 22:630–638
Hahn NS, Matthiesen FA (1993) Notas biológicas sobre Paratemnus
minor (Pseudoscorpiones, Atemnidae). Rev Bras Biol
53:571–574
Hansen L, Klotz J (2005) Carpenter ants of the United States and
Canada. Comstock Publishing Associates, Ithaca
Harvey MS (1986) The systematics and biology of pseudoscorpions.
In: Austin AD, Heather NW (eds) Australian Arachnology.
Australian Entomological Society, Brisbane, pp 75–85
Harvey MS (2011) Pseudoscorpions of the World, Version 2.0,
Western Australian. http://www.museum.wa.gov.au/catalogues/
pseudoscorpions. Accessed 27 Oct 2015
Johnson DL, Wellington WG (1980) Predation of Apochthonius
minimus (Pseudoscorpionida, Chthoniidae) on Folsomia candida
(Collembola, Isotomidae).1. Predation rate and size selection. Res
Popul Ecol 22:339–352
Krebs CJ (1999) Ecological methodology, 2nd edn. Addison Wesley
Educational Publishers, Menlo Park
Litsios G, Pellissier LC, Forest F, Lexer C, Pearman PB, Zimmermann
NE, Salamin N (2012) Trophic specialization influences the rate
of environmental niche evolution in damselfishes (Pomacentri-
dae). Proc R Soc B-Biol Sci 279:3662–3669
Lubin Y, Bilde T (2007) The evolution of sociality in spiders. Adv
Study Behav 37:83–145
Majer M, Agnarsson I, Svenning JC, Bilde T (2013a) Social spiders of
the genus Anelosimus occur in wetter, more productive
123
 Author's personal copy
environments than non-social species. Naturwissenschaften
100:1031–1040
Majer M, Svenning JC, Bilde T (2013b) Habitat productivity
constrains the distribution of social spiders across continents:
case study of the genus Stegodyphus. Front Zool 10:1–10
Majer M, Svenning JC, Bilde T (2015) Habitat productivity predicts
the global distribution of social spiders. Front Ecol Evol 3:1–10
Manly BFJ, McDonald LL, Thomas DL, McDonald TL, Erickson WP
(2002) Resource selection by animals: statistical design and
analysis for field studies, 2nd edn. Kluwer Academic Publishers,
Dordrecht
Martin CH, Wainwright PC (2011) Trophic novelty is linked to
exceptional rates of morphological diversification in two adaptive
radiations of Cyprinodon pupfish. Evolution 65:2197–2212
Nentwig W, Wissel C (1986) A comparison of prey lengths among
spiders. Oecologia 68:595–600
R Core Team (2015) R: a language and environment for statistical
computing. R foundation for statistical computing, Vienna
Rasband WS (2015) ImageJ US National Institutes of Health,
Bethesda. http://imagej.nih.gov/ijA. Accessed 27 Oct 2015
Settepani V, Grinsted L, Granfeldt J, Jensen JL, Bilde T (2013) Prey
capture behaviour in the social spider Anelosimus eximius
(Araneae: Theridiidae): Responses to prey size and type. J Evol
Biol 26:51–62
Souza ALT, Gonzaga MO, Vasconcellos-Neto J (2007) Prey capture
behaviour in the social spider Anelosimus eximius (Araneae:
Theridiidae): Responses to prey size and type. Ethology
113:856–861
123
L. F. Garcı́a et al.
Tizo-Pedroso E, Del-Claro K (2007) Cooperation in the neotropical
pseudoscorpion, Paratemnoides nidificator (Balzan, 1888): feed-
ing and dispersal behavior. Insectes Soc 54:124–131
Tizo-Pedroso E, Del-Claro K (2011) Is there division of labor in
cooperative pseudoscorpions? An analysis of the behavioral
repertoire of a tropical species. Ethology 117:498–507
Tofolo VC, Giannotti E, Moleiro HR, Simoes MR (2011) Diet and
spatial pattern of foraging in Ectatomma opaciventre (Hy-
menoptera: Formicidae) in an anthropic area. Sociobiology
58:607–619
Tschinkel WR (2002) The natural history of the arboreal ant,
Crematogaster ashmeadi. J Insect Sci 2:1–16
Wetterer JK, Gruner DS, Lopez JE (1998) Foraging and nesting
ecology of Acromyrmex octospinosus (Hymenoptera: Formici-
dae) in a Costa Rican tropical dry forest. Fla Entomol 81:61–67
Weygoldt P (1969) The biology of pseudoscorpions. Harvard Univer-
sity Press, Cambridge
Whitehouse MEA, Lubin Y (2005) The functions of societies and the
evolution of group living: spider societies as a test case. Biol Rev
80:347–361
Witt DL, Dill LM (1996) Springtail postmolt vulnerability to
pseudoscorpion predation: Mechanisms and implications. J Insect
Behav 9:395–406
Yip EC, Rayor LS (2014) Maternal care and subsocial behaviour in
spiders. Biol Rev 89:427–449
Zeh JA, Zeh DW (1990) Cooperative foraging for large prey by
Paratemnus elongatus (Pseudoscorpionida, Atemnidae). J Arach-
nol 18:307–311