Araneae

Related terms:

Genus, Hymenoptera, Arthropods, Fungi, Mites, Larvae, Lepidoptera, Coleoptera

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Arachnids☆
Paul A. Selden, in Reference Module in Life Sciences, 2017

Araneae
Araneae is the largest arachnid order (unless mites are considered as one), with
113 families, 4033 genera, and 46,499 species described to date. Spiders are dis-
tinguished from other arachnids by their silk-producing spinnerets at the end of
the abdomen and the prosomal poison glands exiting through their chelicerae
modified as fangs (Fig. 3). Spiders are among the very few animals that use silk
throughout their entire lives. A narrow stalk (the pedicel) joins the abdomen to
the prosoma, allowing great flexibility and precise orientation of the abdominal
spinnerets. Pedipalps are short and leg-like, except in adult males, where they
are modified for sperm transfer. Colors are predominantly dull tans, browns, and
blacks, but many common, conspicuous spiders are colorful, even iridescent. The
abdomen of most spiders shows no trace of ancestral segmentation, unlike that of
other arachnids. The Neotropical tarantula Theraphosa leblondi (Theraphosidae) is
the largest spider at about 10 cm in body length. The smallest spiders are the tiny
orb-weaving Symphytognathidae, adults of which are less than 1 mm long.
Fig. 3. Spider (Araneae: Araneidae: Argiope aurantia).

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Spiders – The Generalist Super Preda-

tors in Agro-Ecosystems
K. Samiayyan, in Integrated Pest Management, 2014

15.1 Introduction
The order Araneae ranks seventh in global diversity of animals, after the five
largest insect orders (Coleoptera, Hymenoptera, Lepidoptera, Diptera, Hemiptera)
and Acari among the arachnids (Parker, 1982) in terms of species described or
anticipated. Spiders are among the most diverse groups on Earth. Incidentally,
earliest records date spiders to some 300 million years ago, 150 million years
before flies began buzzing around (Ranjit Lal, 1995). Among these taxa, spiders
are exceptional for their complete dependence on predation as a trophic strategy.
In contrast, the diversity of insects and mites may result from their diversity in
dietary strategies – notably phytophagy and parasitism (Mitter et al., 1988). They
are of economic value to man because of their ability to suppress pest abundance in
agro-ecosystems. Faced with the need to reduce pesticide usage on the world’s crops
and optimize natural biological control, a full investigation of the means by which
spiders influence pest abundance is long overdue. Also, in recent years, there has
been a realization by ecologists that components of agro-ecosystems are tractable
to manipulation and that spiders are convenient model organisms. Consequently,
there are a growing number of investigations in which spiders in agro-ecosystems
are used as tools to gain fundamental insights into the role of generalist predators in
community and ecosystem function. There is a little rhyme of English origin, but of
uncertain age and derivation which says: ‘If you wish to live and thrive, Let a spider
run alive’.

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Molecular Assembly in Natural and En-

gineered Systems
Martin Humenik, ... Andrew Smith, in Progress in Molecular Biology and Transla-
tional Science, 2011

II Spiders
Spiders (Aranea) belong to the class Arachnida which includes the scorpions, ticks,
and mites, and are part of the Arthropod phylum. Spiders can basically be defined as
air-breathing Arthropods with two body parts, the cephalothorax and the abdomen.
They have eight legs attached to the cephalothorax, and chelicerae which are ven-
om-injecting fangs. Most spiders have eight simple eyes but some species have
more or fewer eyes. Spiders are able to ingest only liquids, as they do not possess
mandibles, and thus their venom liquidizes their prey prior to the spider ingesting
it. The most important feature that has captured the interest of materials scientists
is the fact that all spiders produce silk from spinnerets on their lower abdomen,
which vary between species from two to eight. These spinnerets have a selection of
spigots through which the silk is extruded from internal glands. The arrangement
and number of spigots vary between the species, sex, and age of the spider.3–6 It is
likely that the use of silk evolved once in spiders, and then over time has evolved into
multiple different forms to fulfill different applications or functions for the spider.

The Aranea order is the seventh most diverse order of all organisms and consists
of over 42,000 species in 109 families.7 All except one known species of spider8 are
predators and primarily feed on insects. The Aranea can be broken down into three
suborders, the Mesothelae being the most ancient, consisting of only one family. The
other two suborders are larger sister groups Mygalomorphae and Araneomorphae,
which diverged around 240 million years ago.9
The Mygalomorphae have downward-pointing chelicerae and incorporate the large
bodied spiders that include the spiders more popularly known as “tarantulas” (Ther-
aphosidae) and “funnel web spiders” (Hexathelidae). However, some species are only
a centimeter long and the smallest less than a millimeter.10 Some of the larger
Mygalomorphae prey on the young of larger animals utilizing their venomous bite to
kill or anesthetize them. This suborder of spiders tends to live in burrows or retreats
and use silk threads to wrap up their prey, for the production of egg cases, and to
extend their sensory range outside of their burrows. Thus they have not evolved any
obvious specialized applications for their silks that would generate highly engineered
threads.

The Araneomorphae is the largest suborder consisting of about 90 families, 2700

genera, and over 32,000 species. They have chelicerae that point diagonally down-
ward and cross over one another. Many spiders use silk just for egg-wrapping and
the lining of burrows, but do not use it for hunting. However, there are a number
of spiders that utilize prey-capture devices made of silk. The prey-capture devices
produced by Araneomorphae are varied, ranging from substrate sheets, aerial sheets,
orb webs, cob webs, and bola, all suited to different preys (Fig. 1). As the capture
of prey is an important driving force in the evolution of predators, some of these
spiders have evolved to have silk with properties that make it capable of stopping
and holding the prey even in flight (Fig. 2). As a consequence of the evolution of
prey-capture devices and hence of the properties of spider silk, the variety of silks
produced by Araneomorphae has expanded.11 This has allowed some spider species
to take a more passive role in hunting, just having to get to their captured prey and
kill it before it manages to escape from the web.

Fig. 1. Various web structures used by spiders, from the basic burrow arrangement
to sheets and then the more complex and familiar orb web. The three structures on
the right are developments from the orb web using the same materials. Black lines
represent MA silk, gray lines flagelliform silk covered in aggregate silk, and circles
represent aggregate silk added to MA silk in cob webs and bola.

Fig. 2. Association between web spinning and species diversity; the numbers repre-
sent species.Modified with permission from Ref. 11.

The prey-capture devices that have received the most attention are the orb webs
produced by some Araneomorphae of the Orbiculariae clade. The spiders of this
clade produce the largest range of silks with up to seven silk types in adult female
spiders.12 The orb web was an evolutionary move away from webs based on the
surrounding substrate toward an aerial frame that is more suited to capturing aerial
prey. The frame of the orb web, which is made of major ampullate (MA) silk, is
connected by mooring threads made from the same material to nearby structures.
The frame then supports radii, also made from MA silk, which are used initially as
connection points to lay down an auxiliary spiral made from minor ampullate (MI)
silk. This MI silk is later replaced by the capture spiral which has sticky properties for
capturing flying insects. The ancestral orb web weavers, namely, cribellate spiders,
relied on a capture spiral consisting of a dry pseudoflagelliform silk. Cribellate silk
is manually combed by special devices on the hind legs of the spiders and is placed
between two threads of the pseudoflagelliform silk.13 Cribellate silk is still seen in the
Deinopoidea super family of spiders. It forms a tufted surface on the capture spiral
with sufficient surface area for van der Waals forces to hold onto the prey.14,15

The ecribellate spiders lack the cribellate silk gland. Ecribellate spiders including the
Araneoidea, a sister group to the Deinopoidea, have replaced the pseudoflagelliform
and cribellate silk capture thread with a more elastic flagelliform silk coated with
an adhesive viscid glue (aggregate silk) that is laid down as the orb web is spun,
meaning that a similar sized web can be spun in one-sixth of the time of cribellate
webs.16
The orb web, whether produced by an ecribellate or a cribellate spider, is an impres-
sive piece of engineering with varying elasticities and strengths in the component
threads. In the case of orb webs made by Araneus sericatus, they are capable of
trapping objects with a kinetic energy of up to about 2 × 10− 4 J, sufficient to capture
a house fly, but, interestingly, not strong enough to capture larger insects. This is
most likely to prevent the spider from having to deal with a prey that can potentially
injure the spider while it struggles. The various structural components of a web are
all held under tension, with each component of the structure having different ranges
of tension, and this is balanced out throughout the web, even though there may be
varying numbers of radii on the top and bottom halves of the orb web.17 Spiders
do not need a specific orientation to spin orb webs, as it has been shown that it is
possible for spiders to spin orb webs in microgravity with little difference in structure.
The only major apparent difference is that in microgravity there was no difference
between the top and bottom half of an orb web. This is most likely due to the lack
of gravity supplying a reference point and its effect on the weight of the structure.18
Additionally, it has been shown that orb-weaving spiders can sense the amount of
the silk components that they have available. Spiders that are starved and prevented
from consuming their own webs, which is a common occurrence in orb weavers,
can ration out there remaining stocks to ensure that a complete web is spun but
with smaller radii and a decrease in spiral turns in the web as well as spacing of the
aggregate droplets that make the capture thread sticky.19

The species within the Orbiculariae clade account for roughly one-fourth of cur-
rent spider diversity; however, most of these species no longer spin orb webs but
have evolved to spin cob webs (Theridiidae) and sheet webs (Linyphiidae), or bola
(Mastophoreae), or abandoned using silk as a prey-capture technique altogether,
indicating that the orb web is just a stepping stone in the evolution of one method
of prey capture. The largest spider clade, the retrolateral tibial apophysis (RTA) clade,
mostly do not use silk for prey capture at all, highlighting the specialist nature
of the orb weavers. It appears that spiders have evolved toward improving and
simplifying the methods of prey capture to reduce the required energy expenditure
for feeding, with the later cob webs of the Theridiidae relying on less adhesive
viscid glue and some webs of Linyphiidae lacking sticky silk altogether, while the
bola of the Mastophoreae requiring a small silk fiber and just a droplet of sticky
silk at the end of the thread. However, these other structures are often used to
capture different categories of prey such as terrestrial prey, or specific species of
prey. For example, Mastophoreae tend to target specific moth species, but are still
opportunistic enough to try capturing nearby prey with their sticky bola. As such, it
seems that the development of the orb web resulted in the evolution and expansion
of the repertoire of spider silks and their properties, which led to the development
of new methods of prey capture that would not have been possible with the earlier
silks.
Researchers have concentrated their efforts on the orb-weaving spiders, specifically
the ecribellates, as their silk appears to have the best properties. Thus the main
species that are discussed here are those belonging to the Araneidae family with
most research concentrating on species from the genii Argiope, Araneus (specifically
Araneus diadematus), and Nephila (specifically Nephila clavipes). Additionally, much
research has been undertaken on the Lactrodectus genus, more commonly known as
“black widow spiders,” from the Theridiidae which spin cob webs but still utilize the
same range of silks as the orb weavers. Early research has concentrated on the MA
silk for two reasons. Firstly, this is the standard silk produced by spiders, which acts
as a lifeline or dragline, the more common name for this silk, and thus is easy to
collect. Secondly, this silk appears to be the strongest silk, as it is used for the frame
and radii of the orb web. Since this is the best characterized spider silk, we start by
describing it in detail.

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Spiders (Araneae) useful for pest limita-

tion and bioindication
Patrick Marc, ... Frédéric Ysne, in Invertebrate Biodiversity as Bioindicators of Sus-
tainable Landscapes, 1999

5.2.1 The basic concept

The relationship of Araneae species to vegetation type has been studied for a long
time (Allred, 1975). All arachnologists recognise the close correspondence between
the vegetation and the composition of the associated spider community. This corre-
spondence is related to the vegetation architecture or cover rather than its diversity.
Numerous workers have detailed these relationships and several reviews referring to
synecological studies have been published (Clausen, 1986; Uetz, 1991; Duffey, 1993;
Wise, 1993). These studies were performed in a wide range of biotopes in temperate,
tropical and desert climates including stony debris and litter, herbaceous, shrub
and tree layers. However, as suggested by Wise (1993), the mechanisms by which
the vegetation affects habitat selection can be complicated, and several variables,
including vegetation structure, microclimate, prey composition, and exchange with
adjacent areas are involved (Fig. 18). Vegetation structure (e.g. height, percentage
of cover) is probably the most important parameter involved in web site selection.
Availability of suitable structures for web or retreat attachments directly influences
the composition of web-building spider communities. Moreover, both vegetation
and leaf-litter structure (depth and composition of plant debris) determine spe-
cific microclimatic conditions in a given site. The specific microclimate acts as a
determinant factor in the composition of wandering and non-wandering spider
communities in relation to the ecological valence of each species (Tretzel, 1952).
More precisely, as suggested in previous studies on ground layer and shrub layer
spider communities (Canard, 1990; Ysnel et al., 1995), although a constancy in the
presence of functional groups for a given vegetation structure can be observed,
within a group, the specific dominance can change from year to year and from
place to place. This change in specific dominance probably depends on variations
in microclimatic conditions in relation to the ecological valence of species.

Fig. 18. Diagram showing the main factors that influence the composition of spider
communities and their intervention level.

Experiments involving artificial habitat manipulations have provided evidence that

spider communities quickly react to these variations in vegetation structure (Wise,
1993). As a rule, spiders are highly sensitive to habitat structure. Therefore, vari-
ations in spider communities can give an index of zoological changes associated
with natural or anthropic (human) changes in vegetation structure. The analysis of
spatio-temporal changes in the constitution of spider communities could offer an
approach for biodiagnosing the evolution or stability of different biotopes when
vegetation is affected by human activities. Due to the position of spiders in the
trophic scheme, other ecological information relating to other components of the
fauna (potential prey and predators of spiders) inhabiting the habitat could be
inferred from the spider community analysis.

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Spider Physiology and Behaviour

Shawn M. Wilder, in Advances in Insect Physiology, 2011

1.2 Why spiders?

Spiders (Arachnida, Araneae) represent one of the largest groups of obligate car-
nivores with almost 40,000 recorded species (Ubick et al., 2005) and are found in
nearly all terrestrial communities (Foelix, 1996; Riechert, 1974). In addition, spiders
can have a large influence on the structure and function of ecological communities
through their direct and indirect effects on herbivores and omnivores (Fagan et
al., 2002; Hurd and Eisenberg, 1990; Krivan and Schmitz, 2004; Moran and Hurd,
1998; Riechert, 1974; Schmitz, 1998; Schmitz, 2004; Van Hook, 1971). Even just the
presence or chemical cues of a spider can cause herbivores to shift their feeding
behaviour to reduce overall consumption of plants or change the types of plants con-
sumed, with resulting effects on plant community composition and nutrient cycling
(Hawlena and Schmitz, 2010; Schmitz, 2003). In addition to natural systems, spiders
may also have important effects on lower trophic levels in agricultural ecosystems
(Riechert and Lockley, 1984). Increased densities of spiders in agroecosystems can
result in herbivore suppression (Schmitz et al., 2000), reduced herbivory (Hlivko and
Rypstra, 2003) and increased crop yield (Riechert and Bishop, 1990; Snyder and Wise,
2001). The effects of spiders on natural and agricultural ecosystems depend upon
the abundance, habitat use and foraging activity of spiders (e.g. Riechert and Bishop,
1990; Schmitz, 1998; Settle et al., 1996), and hence, it is important to understand
the factors (e.g. nutrition) that influence spider foraging, growth and reproduction.

Interestingly, while the effects of spiders on community composition and ecosystem

functioning are due to the feeding behaviour of spiders, relatively little is known
about the nutritional ecology of spiders (e.g. what nutrients are required for optimal
growth, reproduction and survival and do spiders obtain the required nutrients in
nature?). The relative lack of research on spider nutrition could be due in part to
the widespread beliefs that spiders are polyphagous and indiscriminate predators
and that the nutritional composition of food is relatively unimportant for predators
because all prey are of relatively high quality, at least compared to herbivores
feeding on plants (Riechert and Harp, 1987). Yet recent studies are challenging these
long-held assumptions about the nutritional ecology of spiders and carnivores, in
general (Jensen et al., 2011a,b; Mayntz et al., 2005; Raubenheimer et al., 2007; Wilder
and Eubanks, 2010). Insect prey can vary in nutritional content and nutrients in prey
can have a large effect on spider performance (Banjo et al., 2006; Mayntz and Toft,
2001; Wilder et al., 2010). Dynamic feedbacks between spider nutrition and prey
nutrient content have the potential to cascade through food webs from the top down
(e.g. changes in cannibalism, intraguild predation and direct and indirect effects on
herbivore and plant communities; Elgar and Crespi, 1992; Hawlena and Schmitz,
2010; Polis and Holt, 1992; Polis et al., 1989) or the bottom up (e.g. changes in the
quantity, nutrient composition and distribution of prey carcasses and spider excreta;
Schmitz et al., 2010).

In addition to being interesting in their own right, spiders represent a model system
for studying carnivore nutrition in general. There are few groups of carnivorous
animals as diverse and widespread as spiders and much is known about the natural
history, physiology, ecology and evolution of many spiders (Barth, 1985; Foelix, 1996;
Nentwig, 1987; Ubick et al., 2005; Wise, 1993). Spiders are also ideal candidates
for laboratory and field research due to their sit-and-wait foraging habits, which
means that they require much less space and change locations less frequently
than other carnivore groups (Foelix, 1996; Uetz, 1992). In terms of evolutionary
research, there is wide variation among spiders species in many traits including
levels of activity (e.g. wandering vs. web building Blackledge et al., 2009), sexual size
dimorphism (Head, 1995; Hormiga et al., 2000), reproductive strategies (Fromhage
et al., 2005; Miller, 2007), diet specialization (Li and Jackson, 1996; Meehan et al.,
2009) and mimicry (Jackson, 1992; Nelson and Jackson, 2006). Many of these traits
have evolved independently multiple times, which facilitates the use of rigorous
evolutionary and comparative analyses (e.g. Grafen, 1989; Purvis and Rambaut,
1995). This combination of characteristics (e.g. natural history information, ease of
use in experiments, extraoral digestion and evolutionarily diverse traits) facilitates
integrative studies of spider nutrition that connect physiology, ecology and evolution
in both the laboratory and the field.

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Neuroparasitology and Tropical Neu-

rology
Oscar H. Del Brutto, in Handbook of Clinical Neurology, 2013

Taxonomy and distribution of venomous spiders

Spiders are arthropods of the class Arachnida, order Araneae. As occurs with snakes,
spiders have been considered as potentially lethal creatures since ancient civiliza-
tions, and the fear of these animals has been surrounded by myths and legends.
From the more than 40 000 described species of spiders, only a few are really
dangerous to humans (Vetter and Isbister, 2008). Venomous spiders with human
relevance are classified into seven families: Theridiidae, Sicariidae, Hexathelidae,
Ctenidae, Actinopodidae, Araneidae, and Oxyopidae.

Theridiidae (cobweb spiders, tangle-web spiders) are distributed all over the world
with the exception of Antarctica. This family has more than 100 genera, most of
which are harmless. However, the genus Latrodectus is highly venomous and one
of the few genera of spiders that can kill humans (Lucas, 1988; Garb et al., 2004).
Latrodectus spp. are called widow spiders because some females eat the male after
mating (Andrade, 1998). Only females are dangerous to humans due to the large
size of their venom glands and fangs. Widow spiders are most often black–hence the
common name “black widow”–and usually have a red hourglass or another colored
mark in their bodies (Fig. 28.4). One Latrodectus spp. (L. hasselti) deserves special
mention, as it is the most common venomous spider in Australia; it is often called
“redback spider” due to a distinctive red strip along the dorsal abdomen. Other
members of the family Theridiidae with clinical significance to humans, are species
of the genera Steatoda and Achaearanea (Isbister and Gray, 2003a).

Fig. 28.4. Widow spiders (Latrodectus spp.) are easily recognized by their black color
and the red hourglass or another colored mark on their bodies.(Downloaded from
http://en.wikipedia.org/wiki/File:Black_Widow_11-06.jpg on July 3, 2012.)

The family Sicariidae includes two genera: Loxosceles and Sicarius. Both are highly
venomous to humans. Loxosceles spp. (recluse spiders, violin spiders) are evenly
distributed over tropical and subtropical areas of the world (Lucas, 1988; Schenone,
2003; Vetter, 2005). Sicarius spp. (assassin spiders) live in desert areas of South
America and Africa (Newlands and Atkinson, 1988). Spiders of both genera can live
under extreme weather conditions and when deprived of food and water.

Hexathelidae (funnel-web spiders) are mainly distributed in rainforest areas of Aus-

tralia, New Zealand, and Asia (Isbister and Gray, 2004). Some other species inhabit
the Mediterranean region and South America. The common name of these spiders is
derived from their ability to construct funnel-shaped webs, where they wait for their
prey at the end of the funnel. Three genera of Hexathelidae have dangerous species:
Atrax, Hadronyche, and Macrothele. While the venom of these spiders is potent, only
a few cases of severe envenoming are reported every year (Isbister et al., 2005).

Ctenidae (wandering spiders) inhabit Central and South America. Only species of the
genus Phoenutria (armed spiders, banana spiders) may be dangerous to humans (Lu-
cas, 1988). Phoenutria spp. are large spiders with an aggressive behavior (Silva et al.,
2011). Other venomous spiders with clinical significance to humans include those
of the genus Missulena (family Actinopodidae); these are called “mouse spiders” and
are found in Australia and Chile (Isbister and Gray, 2004). Also, spiders of the family
Araneidae (Orb-weaver spiders) and Oxyopidae (Lynx spiders) may be dangerous to
humans, although reports of severe envenoming are scarce.
Other spider families have been blamed for being dangerous to humans. However,
the damage that they produce is minimal and mainly related to mechanical injury
of the skin and not to the injection of venom (Isbister and Gray, 2003b; Isbister
and Hirst, 2003; Isbister and Framenau, 2004). Among these are members of the
family Sparassidae (huntsman spiders), Salticidae (jumping spiders), Lamponidae
(white-tailed spiders), Lycosidae (wolf spiders), Desidae (black house spiders), Mi-
turgidae (yellow sac spiders), and Agelenidae (hobo spiders).

The term “tarantula” has been inaccurately used for centuries. Originally used to
name an old world spider of the family Lycosidae (Lycosa tarantula), unjustified fear
of this arthropod–that was even the origin of an Italian dance called tarantella,
which was supposed to cure the effects of its bite–caused a number of spiders from
different families to be referred to by that name. When Europeans conquered The
Americas and Africa, they called almost any large or hairy spider found in these
new lands a tarantula. Nowadays, the term should be restricted to members of the
family Theraphosidae, most of which are harmless to humans or produce only minor
medical problems that are not related to envenoming, but to an urticarian effect of
spider hairs when they come in contact with the eyes of the victims (Isbister et al.,
2003a).

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Amino Acids, Peptides and Proteins

Nicolas Andreotti, ... Jean-Marc Sabatier, in Comprehensive Natural Products II,
2010

5.10.5 Peptides from Spider Venoms

Figure 5. Venomous spiders. (a) Mygalomorphae-type chelicerae from Goliath
bird-eating spider (Theraphosa blondi); (b) Araneomorphae-type chelicerae from yel-
low sac spider (Cheiracanthium punctorium); (c) Venezuelan suntiger (Psalmopoeus
irminia); (d) black widow (Latrodectus hesperus); (e) red-knee tarantula (Brachypelma
smithi); (f ) mouse spider (Missulena bradleyi). Photos from (a) to (f ) by J. Smith (Cre-
ative Commons Attribution ShareAlike License), R. Altenkamp (Creative Commons
Attribution ShareAlike License), B. Smith (Creative Commons Attribution ShareAlike
License), Fir0002 (GNU free documentation license), B. Smith (Creative Commons
Attribution ShareAlike License) and Fir0002 (GNU free documentation license),
respectively.

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Nature of the Belowground Ecosystem

and Its Development during Pedogene-
sis
Richard John Haynes, in Advances in Agronomy, 2014

4.4 Macrofauna
Soil macrofauna have body widths >2 mm and include the orders Araneae, Opil-
iones, Scorpiones, Isopoda, Diplopoda, Chilopoda, Isoptera, Hymenoptera, and
Lumbricidae. The arachnids Araneae (spiders), Opiliones (harvestmen), and Scorpi-
ones (scorpions) are a group of important predators in soils, which feed on insects
such as beetles. Some Opiliones are also omnivores and scavengers. Chilopoda (cente-
pedes) are a group of important predators, which inhabit moist microhabitats. They
are thought feed on most soft-bodied fauna of reasonable size. Isopods (woodlice)
and Diploda (millipedes) generally feed on plant residues that have been partially
decomposed by fungi and bacteria.

Isoptera (termites), Hymenoptera (ants), and Lumbricidae (earthworms) are considered

as ecosystem engineers (Jouquet et al., 2006). Such organisms often change the
chemical, physical, and structural properties of their habitat with impacts on other
biota and ecosystem functions (Jones et al., 1994; Jouquet et al., 2006). These groups
move and mix large amounts of soil and plant litter. Termites exhibit a diversified
phylogeny and a broad range of social behaviors and feeding habits (Jeffery et al.,
2010). They are the most abundant macrofauna in many tropical ecosystems and
they live in colonies containing from 3000 to 250,000 individuals and have a complex
social structure (Whalen and Sampedro, 2010; Roisin and Korb, 2011). Termites
depend on obligate symbiosis with gut microflora to digest lignocellulosic materials.
The gut microflora varies from species to species and often includes a diverse
group of bacteria, archaea, and protists (Eggleton, 2006; Ohkuma and Brune, 2011).
Decaying wood and other plant residues are often the main food source of termites
although some feed on soil organic matter and are able to decompose humic
materials. While some termites construct large mounds, others instead construct
nests within the soil. Termites thoroughly mix soil minerals and organic residues,
add decomposed residues to the soil and release essential plant nutrients from
organic residues (Jouquet et al., 2011).

Ants also live in complex colony structures with typically 10,000 to 12,000 worker ants
(Whalen and Sampedro, 2010). They are often generalized predators, scavengers,
or opportunistic omnivores. Some species construct mounds aboveground while
others construct their nests within the soil. The latter mix significant amounts of
organic residues with soil mineral particles (promoting aggregation) while the soil
around mounds is generally enriched in organic matter (Folgarait, 1998; Frouz and
Jilkova, 2008).

Earthworms are hermaphroditic segmented worms that require moist conditions

for their activity. They are often the most important soil macrofauna in temperate
regions (forests, grasslands, agricultural fields) and tropical forests where their
numbers typically range from 50 to 2000 m−2 with 2–10 species often being present
at any one site (Lee, 1985; Dlamini and Haynes, 2004). They are often divided into
three main ecological classes: epigeic, anecic, and endogeic (Coleman et al., 2004).
Epigeic species live in the litter layer and they progressively fragment litter and
promote its decomposition. Anecic species feed and cast in the surface litter but have
permanent burrows and can live at depths of 1 m or more. They form continuous
macropores to depth and because they pull litter down their burrows, the walls
become lined with organic matter. Endogeic earthworms live in the top 15 cm of
soil and ingest soil with a preference for particulate organic matter. They continually
burrow and backfill their burrows and other voids with their casts. Turnover rates
of soil in temperate grasslands induced by earthworms commonly vary from 40 to
70 t ha−1 year−1 (Bouche, 1983).

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Spider Physiology and Behaviour

Roger D. Santer, Eileen A. Hebets, in Advances in Insect Physiology, 2011

1 Introduction
Our knowledge of arachnid physiology and behaviour varies tremendously across
orders. While much is now known with respect to the Acari and Araneae (e.g. see
Barth, 2002 and chapters in this volume), we know less about the Scorpiones, and
much less still about the Amblypygi, Schizomida, Thelyphonida, Ricinulei, Pseu-
doscorpiones, Opiliones, Palpigradi, and Solifugae (e.g. see Beccaloni, 2009). In
this chapter, our focus is on the Amblypygi, commonly known as whip spiders. We
hope to demonstrate that although many aspects of their sensory and behavioural
biology are shared with their better-known cousins, there is much about them
that is unique. Here, we hope to motivate neuroethological investigation of this
lesser-known arachnid order, and to share something of the appeal that drew us to
begin our own investigations.

The Amblypygi belong to the Tetrapulmonata, an arachnid clade that also includes
three other extant orders: Thelyphonida (whip scorpions), Schizomida (short-tailed
whip scorpions), and Araneae (spiders) (Dunlop, 2010; Shultz, 2007). As in spi-
ders, the body of whip spiders is divided into a prosoma (  cephalothorax) and
an opisthosoma (  abdomen), separated by a narrow waist called a pedicel. Whip
spiders also share with spiders a prosomal sucking stomach and some genital and
sperm characteristics (e.g. Dunlop, 2010). On this basis, some authors have grouped
the Amblypygi with the Araneae under the name Labellata, although this is not
supported by the most recent studies (see Dunlop, 2010; Shultz, 2007; Weygoldt,
2000). Whip spiders possess neither the spinnerets nor venom glands of spiders
(Weygoldt, 2000).
Phylogenetic studies most commonly group whip spiders with the Uropygi (com-
prising Thelyphonida and Schizomida) within the Pedipalpi clade (e.g. Dunlop, 2010;
Fahrein et al., 2009; Shultz, 2007). Within this clade, the pedipalps are raptorial.
Furthermore, only the rearmost three pairs of legs are used in walking and the first
pair has evolved into thin and elongated mechano- and chemosensory feelers. In
whip spiders, these ‘antenniform legs’ (sometimes called ‘whips’) are so long that
they can be 2.5 or more times the length of the walking legs (Fig. 1) and, in some
large neotropical species, can have a span of nearly 60 cm (Foelix and Hebets, 2001).
They are an incredible example of convergent evolution with the antennae of insects.
In contrast to whip scorpions, whip spiders have no flagellum, which is why they
are sometimes given the common name ‘tailless whip scorpions’ (Weygoldt, 2000).
Whip spiders differ from both whip scorpions and spiders in being dorsoventrally
flattened, which enables them to scuttle into narrow crevices, and by a reduction of
the patellae of the legs, which are the breaking points for autotomy (a property that
has pros and cons from the point of view of the experimentalist) (Weygoldt, 2000).

Fig. 1. The basic anatomy of a whip spider. The species shown is Phrynus marginemac-
ulatus, a species found in the south-eastern USA and on Caribbean islands. In this
species, the carapace of a large male has a width of approximately 8 mm. Note the
presence of spiny, raptorial pedipalps, and hugely elongated ‘antenniform’ forelegs
which are used as sensory structures and not for locomotion.

The Amblypygi are a small order. A recent count identified 158 recognised species,
divided into 17 genera and 5 families—a very small number in comparison to
the 42,055 species, 3821 genera, and 110 families of Araneae recognised at the
time of writing (Harvey, 2007; Platnick, 2011). The oldest fossils that can be firmly
identified as whip spiders come from the late Carboniferous period (Dunlop, 2010).
These fossils resemble the most basal extant species, Paracharon caecus, which has
short pedipalps that articulate up and down rather than side to side like the more
modern species (Dunlop, 2010; Weygoldt, 2000). This single extant species and those
identified from Carboniferous fossils were grouped into the suborder Palaeoam-
blypygi (Weygoldt, 1996, 2000). A second suborder, Euamblypygi, comprises the
more modern whip spiders, and within this, fossils from the early Cretaceous period
have been assigned to the family Phrynidae, and two to the living genus Phrynus
(Dunlop, 2010; Weygoldt, 1996, 2000).

Whip spiders generally inhabit the tropics and sub-tropics, with a few species found
in almost temperate zones, but none in regions that experience extreme cold or
snow (Weygoldt, 2000). Whip spiders are not normally found in deserts, but Damon
variegatus can be found in savannah-like habitats, hiding under rocks or in caves
(Weygoldt, 2000), and Phrynus neomexicanus can be found in the Sonoran desert of
the southern USA (Hebets, personal observation). Most whip spider species inhabit
rainforests (Weygoldt, 2000). Species smaller in size tend to be found within the
leaf litter, whereas the larger-bodied species (including neotropical Heterophrynus
and Phrynus species which will feature heavily in this review) are found on large
trees and rocky outcrops (Weygoldt, 2000). These larger species tend to prefer big
trees with buttress roots (Dias and Machado, 2006; Hebets, 2002), which means that
for large species like Heterophrynus longicornis, selective logging and general habitat
destruction are a serious issue (Dias and Machado, 2006). Most species appear to
be strictly nocturnal, spending the day hidden away deep within buttress roots, in
burrows, or under bark, and emerging to sit and wait for prey after dark (Hebets,
2002; Weygoldt, 2000). Our own experimental species, Phrynus marginemaculatus,
can be found under limestone rocks in the pine hammock of the south-eastern USA
and on Caribbean islands. Remarkably, this species can breathe under water by use
of a plastron (Hebets and Chapman, 2000b), an ability that no doubt helps it to
persist in these frequently inundated habitats.

Weygoldt (2000) reports that many rainforest species will readily inhabit caves. For
example, in Kenya, Damon diadema can be found both in coastal caves and the
surrounding forest (Weygoldt, 2000). However, there are also species that have
become adapted to life in caves and are found only in these habitats (e.g. Baptista
and Giupponi, 2002, 2003). These may lack medial or lateral eyes (or both), but their
physiology and behaviour are not yet well studied. Given that populations of some
species appear to be confined to caves, whereas others are found in nearby forests,
comparative studies could provide useful insights into physiological adaptations to
cave living.

In this review we are primarily concerned with the fascinating sensory and be-
havioural biology of these ancient arachnids. Before delving into this, however,
we must address two important issues. Firstly, there is some confusion regarding
species nomenclature. For example, much early physiological work was carried
out on ‘Admetus pumilio’, a no longer recognised species that is now divided into
Heterophrynus batesii and H. longicornis (Weygoldt, 1974; as described in Harvey,
2003). As such, the authors of some early physiological work state that their studies
may have been conducted on representatives of both species (e.g. Foelix and Troyer,
1980; Foelix et al., 1975), but the impression from subsequent work is that the likely
species identification is H. longicornis (e.g. Beck et al., 1977; Foelix and Hebets,
2001; Foelix et al., 2002; Igelmund, 1987). Here, we will follow the identifications
in the original publications unless a more accurate alternative is clearly indicated.
Secondly, a large volume of early literature on whip spiders was published in the
German language, and is sadly not easily accessible to many. In both cases we direct
readers to Peter Weygoldt's recent volume (Weygoldt, 2000) and, for coverage of
the German language work, to the review of Foelix and Hebets (2001). Our aims
in this chapter are to complement these works by providing an updated review of
whip spider sensory physiology and its role in behaviour, with the hope that we will
contribute to an emerging accessible picture of this order that will motivate future
research. Our review will broadly be divided into two sections: firstly, an account
of whip spider sensory biology (Section 2), followed by a description of the state of
knowledge on the roles of these adaptations in the guidance of behaviour (Section
3). We will finish by attempting to draw some general conclusions and highlighting
important areas for future study (Section 4).

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Phylum Arthropoda
D. Christopher Rogers, Ilse Bartsch, in Thorp and Covich's Freshwater Invertebrates
(Fourth Edition), 2019

Arachnida: Subclasses
Arachnids are a large group of mainly terrestrial organisms classified in 15 extant
orders (see Proctor et al., 2015). Representatives of four of these orders, Araneae
(true spiders) and three orders of the subclass Acari (mites)—Mesostigmata, Sar-
coptiformes, and Trombidiformes—are the only chelicerates habitually found in
and around freshwater habitats. Among these, members of only a few groups of
Sarcoptiformes and Trombidiformes are truly aquatic. Phylogenetic relationships of
arachnid orders are not clearly understood, and the classification is undergoing sub-
stantial revision based on interpretation of morphological, behavioral, and genetic
data. Here we follow the classification outlined by Proctor et al. (2015) in Volume I
of this series.

1 Body conspicuously divided into cephalothorax and abdomen, spinnerets

present, spiders ………………………… Araneae [p. 552]
1  Body less conspicuously divided into gnathosoma (capitulum) and idioso-
ma, spinnerets absent, mites………………………… Acari [p. 553]
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