tarantulas of texas - Natural Science Research Laboratory

TARANTULAS OF TEXAS
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THEIR MEDICAL IMPORTANCE,
' UD WORLD-WIDE BIBLIOGRAPHY TO
THE THERAPHOSIDAE (ARANEAE)
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R. G. Breene, D. A. Dean, J. C. Cokendolpher, and B. H. Rege
TARANTULAS OF TEXAS
THEIR MEDICAL IMPORTANCE,
AND WORLD-WIDE BIBLIOGRAPHY T O
THE THERAPHOSIDAE (ARANEAE)
R. G. ~reene',D. A. ~ e a nJ.~C.
, cokendolpher3, and B. H. ~ e ~ e r ~
First Printing, October 1996
Copyright O By The American Tarnntzila Society
PO Box 1617
Artesia, New Mexico 88211- 1617 USA
All rights reserved
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PO Box 1617, Artesia, New Mexico 8821 1-1617 USA
Department of Entomology, Texas A&M University, College Station, Texas 77843-2475 USA
2007 29th Street, Lubbock, Texas 7941 1 USA
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4616 South State Road 9, Greenfield, Indiana 46140 USA
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Copyright 0 1996 By the American Tarantula Society. All rights
reserved. Reproduction in whole or in part by any means or methods is prohibited unless authorized in writing by the copyright
holder.
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INTRODUCTION
Tarantulas are present and often conspicuous in nearly all areas of
Texas. The presence of large numbers of males is a frequent occurrence
on Texas roads during parts of spring, summer, and fall, inspiring numerous news media reports. Flooding occasionally causes mass movement
of tarantulas in some areas, attracting more media attention. Tarantulas
have become as associated with Texas as armadillos or the Texas homed
lizard. They have been gaining popularity steadily for decades in many
parts of the world as zoo and display animals, and as very low maintenance pets. Yet despite the popular attention, less is known about them than
just about any other animal with even a small fraction of their notoriety.
Little work has been completed on tarantulas for a number of reasons.
Although tarantulas strike terror into the hearts of the arachnophobic, they
are not medically significant compared to other groups of animals whose
researchers may compete for the same research dollar. For most species,
tarantula venom is not known to be harmful to humans, or in the species
where it is harmful, they do not generally deliver enough of it in the bite to
generate a large number of medical reports. They are not considered
useful as biological control agents of medically important pests, with the
possible exception of a potential to cause rodent mortality.
In agriculture, they are not suspected of being significant biological
control agents on pests of plants. Conversely, they are not known to
damage commercial crops or gardens. Consequently, there has been little
incentive to apply any agricultural resources toward their study.
Very few arachnologists have taken an active research interest in them
for a number of reasons. The long life cycle of tarantulas seriously complicates research efforts. Graduate students may run out of funding years
before the time required to adequately document the biology of a single
species in one location. Other potential researchers are repelled by their
notoriety with the general public. In some cases, araclmologists assume
that since they are so conspicuous, all work must have already been done
on them years ago.
This treatise concentrates on what is known about tarantulas of Texas,
discussing their biology, ecology and recorded geographic distribution.
The medical importance section applies to all species covering the entire
tarantula family. A bibliography to the family Theraphosidae is included.
BIOLOGY
Geographic Range and Habitat
The nearly 800 species of tarantulas belonging to the family Theraphosidae (Coddington & Levi 1991) are known from the warmer parts of
the world. In general, tarantulas can be divided into arboreal (living in
trees) or burrowing species (dwelling in the soil or under rocks). Treedwelling species inhabit more tropical areas. Burrowing species are found
throughout nearly all the geographic range of the theraphosid family. All
US tarantulas are thought to be burrowing species.
Burrowing species are split by some authors into two groups; opportunistic and obligate burrowers (Marshall 1996). Opportunistic burrowers
may take advantage of already existing habitats, modifying them for their
needs. These habitats include fallen trees, under rocks, firewood stacks,
rodent burrows or crevices in the ground. An example in Texas is
Aphonopelma anax (Charnberlin), a species that has been found extensively in South Texas modifying preexisting habitats, but is-likely capable
of constructing its own burrow, Obligate burrowers only rarely modifl
preexisting structures, excavating their burrows themselves according to
their needs. Marshall (1996) gives Aphonopelma chalcodes Chamberlin
(a more western species not found in Texas) as an example of an obligate
burrower.
With many species, the burrow entrance often appears as an undistinguished round opening in the ground. A thin film of silk may cover the
entrance at times, and more northern and western species plug the burrow
during the winter (Baerg 1958) or in times of extremely high temperatures. Individuals of certain southwestern species may also seal the
burrow in winter and during other specific periods of the year (Minch
1979a). In extreme South Texas, tarantulas are active and have been
collected during all months of the year (unpubl. data). Although no
evidence of burrow plugging has been observed in tarantulas from
southern Texas, the behavior is not ruled out. Small piles of excavated
soil can be seen occasionally near active burrows.
Baerg (1958) wrote that tarantulas can dig their own burrows in soil
using the fangs and manipulating the excavated soil with their pedipalps.
It has also been widely suspected that tarantulas take over abandoned
rodent dens. Young rodents (rats and mice), and grown mice are eaten by
many captive tarantulas. It is possible that active rodent burrows may be
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invaded and purged of rodents by larger tarantulas. The significance of
tarantulas as rodent predators in nature is unknown. Research of this land
is difficult at best and has yet to be attempted.
Many ground nesting animals have symbiotic relationships, or at least
tolerate other taxa of animals in their burrows, and tarantulas are no
exception. Cocrofi & Hambler (1989) reported a rnicrohylid frog in a
commensal relationship with the tarantula Xenesthis immanis (Ausserer)
in southeastern Peru. Harold A. Dundee, of Tulane University in New
Orleans reported a species of the narrow-mouthed frog genus Gastrophryne living in the burrows of Aphonopelma sp. in the US (pers. comm.
1991, 1995). Dundee noted that tarantula pedipalp contact with the slun
of the toad, and apparent recognition, prevented attack; suggesting a
chemical cue may be used to suspend predator behavior. He hypothesized
that tarantulas coexist with the toads in exchange for protection the toad
may provide against ants.
Baerg (1958) described the tarantula habitat in Arkansas as rocky
hillsides covered sparsely with vegetation, usually with a southern
exposure. In the College Station, Texas area, burrows fairly close together
(one meter or so apart) have been observed on level ground in St.
Augustine grass lawns in residential areas (unpubl. data). In South Texas,
the highest concentration of tarantula burrows has been found in residential St. Augustine grass lawns, golf courses, and mowed lots in small
coastal communities, also on non-sloping ground. An occasional active
tarantula burrow can be located in grazed (largely overgrazed) rangeland
and wildlife refuges in the South Texas area, but the highest tarantula
numbers appear in residential areas, as seen not only from the frequency
of burrow discovery, but also from the number of males observed in these
areas during mating season.
Tarantulas have been collected in the US from Arkansas, Oklahoma,
Colorado, Utah, New Mexico, Arizona, Nevada, California, and Texas
(Smith 1994). Baerg (1958) also listed parts of Missouri as within the
distribution boundaries of US tarantulas. Smith (1994) did not find any
material collected from Louisiana, Mississippi, Alabama, or Georgia.
Smith later related that he could not locate specimens from either Missouri
or Kansas (pers. cornm. 1995). Theraphosids have since been recorded
from Kansas (pers. cornm. H. Guarisco 1995) and Louisiana (pers.
comm. R. Moore 1996) (Rossman 1984). Roth (1993) mentioned a record
of a theraphosid found in Alabama.
Early in this century, a single female specimen was captured in the Dry
Tortugas Islands, Florida, and designated as the holotype for Brachypelma aureoceps (Charnberlin 1917). Smith (1994) believes it likely that
the specimen was accidentally imported on a ship, probably from Mexico,
and suggested the species be included and maintained in Brachypelma in
the hope that the species will eventually be located and identified in its
place of origin.
Tarantula Development and Longevity
The developmental stages of spiders have been named a wild variety of
terms by different workers over time. It sometimes appeared that researchers would coin a new word for a developmental stage if they didn't
admire any of the ones then available. We follow the standardization of
terms used to describe early development of spiders proposed by Downes
(1987).
The egg (embryo) hatches (eclosion, sheds its chorion, roughly the
"eggshell") into the postembryo instar while still inside the eggsac. The
postembryo in tarantulas resembles a large egg with a stubby-legged mite
glued by its posterior to the egg's surface. Although capable of moving its
appendages, the postembryo is not a mobile stage (at least in North
American species), and nutrients continue to be derived from internal yolk
located in its abnormally enlarged abdomen.
Still within the eggsac, the next shedding of the exoskeleton, or the first
true molt, begins the stage termed the first instar (Downes 1987). The
subsequent molt places the immature into the second instar, the next
brings the third instar, and so on. The total number of molts in tarantulas
will vary with the species, sex, and probably with the individual. Baerg
(1958) recorded a total of 22 molts for one Aphonopelma sp. individual.
The term "spiderling" is an arbitrary term, most often referring to the first
to third or fourth instars.
The stage preceding sexual maturity is called the penultimate instar,
and is followed by the adult, or ultimate instar. All male spiders and most
females, except for the more primitive mygalomorph spiders and a few of
the more advanced ones, do not molt again after reaching adulthood, with
rare exceptions. Tarantulas and some other spider females can survive the
ultimate instar and continue to molt, usually on a yearly basis. The first
true molt following the ultimate instar is termed the first postultimate
instar, the next the second postultimate instar, and so forth.
A number of inappropriate terms have frequently been applied to
various developmental stages of spiders, with nymph, and larva appearing
more often than others. A brief overview follows.
The spider egg (embryo) is a widely accepted term for the stage from
the time of egg deposition to the rupturing of the chorion at eclosion
(hatching) by Downes (1987), Holm (1940), and several others. An
embryonic event called reversion begins a process where many species of
spider embryos become somewhat elongate. A post-reversion embryo was
thought to be different enough to be its own stage, and was called a
prelarva (Vachon 1957) and a pullus (Canard 1984, Neet 1985). These
last two authors considered the embryo and the pullus as both falling
within the "primary period of spiders (the egg stage).
Terms of past usage for the postembryo instar have included quiescent
stage, or quiescent nymph (Ewing 1918), the incomplete stadia of the
postembryonal stadia (Holm 1940), a deutovum (Gertsch 1949), a larva
then prenymph of the larval period (Vachon 1957), a protonymph
(Jkzkquel 1960), the interchorional stage (Galiano 1973), the exchorionate
stage (Hydorn 1977), the endovitelline then exovitelline stages (Whitcomb
1978), the postpullus then prejuvenile of the juvenile period (Canard
1984), and finally, the postpullus then prelarva of the juvenile period
(Neet 1985).
Other terms used as equivalents for first instar have been the complete
stadia (Holm 1940), first nymph (Vachon 1957), first young of the
juvenile period (Canard 1984), and first larva of the juvenile period (Neet
1985).
The two major factors affecting the rate of spider development are
temperature and food availability. Baerg (1958) reported that 10 to 13
years with an approximate average of 16 to 17 molts were required from
egg to adult for the Aphonopelma sp. in his region of northwestern
Arkansas. In warmer areas, time to maturity is probably much shorter.
When maintained in the laboratory under elevated temperature regimes
with a continuous source of food, male Aphonopelma anax (Chamberlin)
can reach maturity in less than two years (unpubl. data), and females
probably in less than three. An average of about 1,000 days from egg to
maturity was reported for one of the world's largest tarantulas, the South
American Theraphosa blondi (Latreille) (Marshall & Uetz 1993). As with
so many other tarantula subjects, little is known about the field developmental times for most species.
The life expectancy of male tarantulas once the ultimate instar has been
reached is wildly variable. Some mature males of the tropical genus
Avicularia have approached or exceeded two years duration in captivity.
In South Texas, Aphonopelma anax males rarely survive more than two
or three months, regardless of the quality of laboratory care (unpubl.
data). Most reports gathered from a wide variety of tarantula species in
captivity relate a life expectancy for adult males of 6 to 18 months. For
North American burrowing species, male longevity is probably an artifact
of captivity and is not likely to be ecologically significant. Because of
natural enemies in the tarantula habitat, male life expectancy once he
abandons the burrow is probably more realistically measured in weeks or
days, or in some cases, perhaps hours.
The record life span for some Aphonopelma females approaches 30
years (Baerg 1958). Many North and Central American Aphonopelma
and Brachypelma females have been reported to live 10 years or more as
adults, but the bulk of the evidence is anecdotal from non-scientific
sources. The average life expectancy for wild female tarantulas endemic
to the US remains a mystery.
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Predation Ecology and Feeding
The significance and impact tarantulas may have upon ecosystems have
never been investigated. If viewed from an agricultural standpoint, tarantulas fit into the mobile, visually acute, or type MV predators along with
the bulk of the spiders and certain insects including mantids and hangingflies (Breene et al. 1993). Although the tarantulas themselves are not
likely to have acute vision, much of the prey they capture does, such as
crickets, grasshoppers, and prey capable of rapid escape behavior. Certain
type SE, or sessile external prey such as caterpillars found on the outside
of plants, are available to them when on the ground near burrows.
Prey species in the natural habitat have rarely been identified. Baerg
(1928) mentioned beetles and crickets as likely food sources. Smith
( 1994) reported beetle remains outside of burrows.
In captivity, tarantulas have been observed taking many living and
inanimate prey items. The list includes crickets, a wide variety of beetles,
cockroaches, frogs, toads, lizards, skinks, snakes, voles, birds of several
species, small fish, caterpillars, adult moths and butterflies, crayfish,
grasshoppers, katydids, termites, Drosophila and other flies (both adults
and maggots), newborn mice and rats, grown mice, hamburger, beef heart,
wasps, bees, and other tarantulas. This list is by no means complete.
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Spiders consume microscopic particulate matter that has essentially
been liquefied in one of two basic ways. In the first method, the fangs
located on the ends of the chelicerae are used to create small incisions in
the exoskeleton of the prey. Digestive enzymes are pumped out through
the mouth into the prey body and tissues are gradually liquefied and
sucked back out into the spider's mouth. The remaining husk of the insect
or other arthropod remains largely intact unless soft-bodied. This method
of feeding is typical of many web-weaving spiders including the cobweb
weavers (Theridiidae) and orbweavers (Araneidae).
In the second method, the spider tears the prey's body into pieces and
works them into a ball, or bolus. Digestive enzymes are poured onto the
ball and liquefied material is ingested. Some wolf spiders (Lycosidae) and
tarantulas use thls method, which simplifies the capture of multiple prey
items when available, instead of catching and securing each item singly.
Natural Enemies
Spider wasps (Hymenoptera: Pompilidae) may be significant natural
enemies of tarantulas in many areas of the US. Most species are slender
with dark, often blue or green metallic colored bodies and orangish,
yellowish, rusty or smoky colored wings, with a few having dark wings.
Many have a body about !h to % of an inch (about 20 mm), but some
southwestern species may stretch over an inch and a half (nearly 40 mm).
Their antennae are filament-like and distinctively coiled in many species.
Most of the 270 or so North American spider wasp species feed on
nectar as adults, but each wasp larva requires a spider to complete its
development into an adult. Adult spider wasps capture their victims after
a brief struggle by injecting a paralyzing venom through the stinger into
the spider's body, often around the base of a leg. Some species will abduct
and paralyze a spider first, then build a cell in the ground to accommodate
it. Others will build the cell first, and then go get the spider. Some smaller
spider wasp species may provision a cell with many spiders.
Once preparations are complete, the tarantula is positioned down in the
cell and a single egg is laid by the female wasp, usually on the abdomen of
the spider. The female then exits and seals the cell with debris. The egg
hatches and the maggot-like larva will burrow into the living spider where
it will feed until it attains enough mass to pupate, later emerging to the
surface and completing its life cycle. The spider dies at some point in the
wasp's feeding after a critical organ is damaged, but can potentially stay
alive, therefore maintaining a fresh food supply for the wasp, for weeks or
months.
Some species will avoid building a cell altogether. The female wasp
simply enters an existing tarantula burrow, stings the spider, lays an egg,
then departs, sealing the burrow behind her.
Some spider wasp species do not construct a burrow or cell, and
tarantulas are not the only spiders attacked by members of the Pompilidae
(Preston-Mafiarn & Preston-Mafham 1993). One African spider wasp
species was observed stinging and paralyzing a wolf spider, and once the
spider was immobilized, she laid an egg on the top (dorsal side) of its
abdomen and departed without returning. In this case, the venom lasted
only a short time, and the spider recovered after about 15 minutes. Once
the egg hatches, the larva burrows into the spider and develops internally.
Where the spider ends up once it eventually dies, and if that location will
be favorable for completion of the development of the young wasp, is
apparently an element left to chance by the wasp mother.
Spider wasp females lay only a few eggs during their brief lifetime. If
the paralyzed spider is rescued from the wasp by a human, in many cases
it will be able to recover from the effects of the venom and revert to
normal within two to six months.
Other tarantula natural enemies include predacious and entomophagous
birds and smaller vertebrate mammals large enough to handle them, some
species of ants, and man. Parasites include certain acrocerid and other
flies, mites, nematodes, and fungi. Although not a predator of tarantulas,
the masked chafer (Scarabaeidae) is reported to kill tarantulas when eaten
(Cokendolpher 1993).
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Seasonal History
Female and immature tarantulas are believed to remain in their burrows
permanently unless flooded, starved, or in some manner severely disturbed
into abandoning the burrow. After molting into an adult, males abscond
from their burrows to search for females. Until recently, it was thought
that all US tarantula species began the mating season in late summer after
most females had molted for the year. The mating season was thought to
last until mid November. Although Baerg (1958) and others had seen
males at different times of the year, most notably in May and June, they
suspected these were males that had somehow survived the winter and
were not particularly significant for mating purposes. Smith (1994)
thought that spring males might be a fail-safe device in case of an early
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hard winter. It is now known that there are at least two separate seasonal
strategies each used by different species (unpubl. data).
Fall ale Mating Strategy
The Aphonopelma sp. Baerg worked with (now thought to be an
undescribed species by Smith 1994), follows the fall male mating strategy.
Males molt into adulthood apparently in summer and fall, produce the
sperm web to charge the pedipalps, and begin to search for females from
August to November. The females retain the sperm in the spermathecae
over the winter and construct the eggsacs in June and July. Females begin
molting in late July and continue through the middle of September (Smith
1994).
Spring ale Mating Strategy
In Cameron County of South Texas, Aphonopelma anax (Chamberlin)
is the representative species practicing the spring male mating strategy.
Males molt into adulthood in April every year (although there are certain
to be some individuals molting earlier and later). They may spend a couple
of weeks to a month or more in their burrows before leaving to search for
females in May and June. The peak mating period appears to be late May,
early June as evidenced by the often large number of males appearing on
the roads largely at night, early, and late in the day. Females produce
eggsacs in June and July and molt in August and early September.
Although males have supposedly been observed in August and later in
the year from unsubstantiated reports from people living in South Texas,
we suspect those of being members of species other than A. anax in the
area, possibly Aphonopelma harlingenum (Chamberlin) or others. We
further suggest that the spring males observed by Baerg and others in
more northern areas were eiLherA. anax or other species that possibly also
use the spring male mating strategy. Only further research and collection
of specimens can provide definitive answers.
Other Strategies
In more arid regions in the southwestern US, the time of mating may be
partially or entirely dependent upon rainfall and temperature.
Courtship and Mating
After molting into an adult, the male builds a sheet-like sperm web and
secretes a drop of sperm onto it from his abdomen, which he then transfers
to his pair of pedipalps located anteriorly near his mouth (sperm induc-
tion). The pedipalps of the male spider function as the intromittent, or
secondary sexual organs. The pedipalps in females and irnmatures have
the general appearance of walking legs that have been reduced in size.
Male pedipalps appear thickened and more club-like. Other male characteristics include tibial spurs located ventrally on the first pair of legs
(leg I), and the overall length of the legs may often exceed that of the
female.
After the male mating preparation period, which may last from a few
days to several weeks, males abandons thier burrows to seek females.
These males are the tarantulas frequently observed crossing roads at night
and at dusk and dawn, sometimes in large numbers. Males (fitted with
radio transmitters) have been recorded traveling up to 1,750 meters in
relatively short periods of time (18 days) (Janowski-Bell 1995) while
searching for mates.
When a male locates a female's burrow, he taps the ground around the
entrance of the burrow, signaling the female that he is present and available. If the female is receptive, she responds by drumming the walls of her
burrow with her pedipalps and ascends to the entrance. The male may
retreat a few inches, perhaps to draw her out of the ground and away from
the burrow entrance. Once the pair touch, he taps her on her front legs
with his pedipalps until she raises them and opens her chelicerae, exposing
her fangs. The male hooks each female fang with a tibial spur and lifts,
balancing the female onto her hind legs. While in this position, he moves
his palps, continuously tapping, down her underside and inserts the end tip
(embolus) from one palpal bulb into the female genitalia, located toward
the anterior of the abdomen ventrally. The male usually repeats the
procedure with the other pedipalp. Afterward, he gently lowers and frees
the female from his mating hooks and backs away from the female,
sometimes rapidly.
After a successhl mating, the female builds an eggsac and deposits
anywhere from fewer than 100 to perhaps as many as 1,500 eggs. One
eggsac from a smaller-sized A. anax female yielded a total number of
1,392 eggs (unpubl. data). The average number of eggs laid will vary with
the species and the individual, and is probably reliant upon the nutritional
status of the female in previous months. Eggsac construction has been
observed on many occasions in the laboratory but not in the field, where it
apparently takes place in a circular, underground chamber probably fashioned for the purpose.
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In captivity, females may weave a saucer-like plate for egg deposition,
or they may entirely enclose themselves in a ball of silk. After the base of
silk is laid down, the yellowish eggs are laid and the female weaves more
layers of silk over the mass. She then gathers the eggsac together in an
irregular ball, which may have varying amounts of substrate incorporated
into it (vermiculite, peat moss, etc.) if available in the cage. In A. anax,
females have been observed on several occasions apparently incorporating
urticating hair into the external surfaces of the eggsac (unpubl. data).
Eggsacs from wild-caught females are more symmetrical in shape, and
appear to have a more intricate, compact construction.
Upon hatching, the spiderlings stay in the mother's burrow for a short
period of time before dispersing. Unlike most other spiders, tarantulas do
not disperse by ballooning, but instead by walking. Small tarantulas may
dig their own burrows initially and expand them as they grow. Otherwise,
they may occupy rodent or other burrows and convert them to their needs.
As mentioned previously, some of the larger irnmatures or females
disturbed from their burrows may take over active rodent burrows.
CLASSIFICATION
The major characteristics separating spiders from the remaining arachnids are the abdominal silk devices like the spinnerets, silk glands and
spigots, the modified pedipalpal tarsi serving as secondary sexual organs
or genitalia in the males, and the lack of segmentation on the abdomen
(except in some highly primitive species) (Coddington & Levi 1991).
Another key characteristic of spiders is the abdomen being narrowly joined to the cephalothorax by a petiole, the pedicel (Kaston 1978). The relationship of Araneae to the other arachnids is illustrated in Figure 1.
Spiders are broadly defined into two suborders. Suborder Mesothelae is
considered the most primitive and contains a single family with a few
dozen species. Suborder Opisthothelae includes infraorders Araneomorphae and Mygalomorphae. Tarantulas belong to the latter infraorder,
which includes 15 families and about 2,000 species. The most advanced
spider infraorder, Araneomorphae, currently contains 90 families and over
32,000 species. The most accepted cladistic hypothesis is illustrated in
Figure 2.
otLer Arachida
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'~'rigonotarbida
Schizornida
Uropygi
m
pedipalpi
L b l yPYgi
rOmO
~esothelae~i~histiidae
Aranea
~ o p i s t h o t h e l ~
Figure 1.--Cladistic hypothesis showing the relationships between the
Arachnids. Trigonotarbida is an extinct arachnid order. Adapted from
Coddington & Levi (199 1).
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r~typoidina-l
Abt
ro iaetidae
L~ornicephalae
Cytaucheniidae
astelloidin
Actinopodidae
Figure 2.--Cladistic hypothesis showing the relationships between the
Mygalomorphae families. Adapted from Raven (1985) and Coddington &
Levi (1991).
Although some non-theraphosid species within the Mygalomorphae
may look very similar to tarantulas, they do not share the theraphosid
characteristic of having only two claws with claw tufts present.
Tarantula Species
In Texas
Up to tlie publication by Smith (1994), most tarantulas from Texas
were either unidentifiable or reported as Aphonopelma hentzi (Girard) or
variations of that name; Dugesiella hentzi, Rhechostica hentzi, etc. Smith
expressed hope that his book would inspire more research on US tarantula
species. Numerous new species were described in his book using single
specimens. Large series of animals of both sexes will need to be collected
at the known localities of the species recognized by Smith and studied to
determine the variation that each species or population exhibits. While we
cannot currently verifjr his taxonomic judgments, they are the most recent
and therefore will remain valid until contradictory data is published. The
following discussion and records are based largely on Smith's publication.
Currently, two genera occur in North America, north of Mexico;
Aphonopelma Pocock, 1901 and Apnchepelma Smith, 1994 (one species
from Arizona). Genera synonymous with Aphonopelma include Chaunopelma, Clavopelma, Delopelma, Dugesiella, and Gosipelma (Raven
1985, Smith 1994). Raven (1985) determined Aphonopelma to be a junior
synonym of Rhechostica, but Aphonopelma was reinstated in a decision
by the International Commission of Zoological Nomenclature (199 1).
The generic names Delopelma, Dugesiella, and Rhechostica are occasionally still erroneously used in the scientific literature by a few
ecologists and physiologists, but they are frequently used in the popular
literature by people unacquainted with arthropod taxonomic methods.
Gertsch & Mulaik (1940) noted Tapinauchenius texensis Simon was
collected from Maverick County, near Eagle Pass, Texas; a border town
on the h o Grande River. In the original publication, Simon (1890) also
noted the species Tapinauchenius caerulescens Simon as being found in
Indian territory around Fort Sill, and probably not only in Oklahoma, but
in Texas as well. Smith (1994) notes that "Both Tapinauchenius
caerulescens Simon 1890 and Tapinauchenius texensis Simon 1890 are
believed to have been collected by George Marx - Washington DC - a
curator who was notorious for incorrectly labeling specimen jars. It is thus
hghly likely that both specimens hail, not from North America, but
Central America - if indeed they are Tapinauchenius." Smith (1994) also
points out that both type specimens are apparently lost, and they may have
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been based on immature specimens. Since no other specimens have been
collected as far as is known, the two species and hence the genus are not
included here. Aphonopelma marxi (Simon) and Aphonopelma mordax
(Ausserer) are also excluded, since there is not enough evidence to confirm that they actually occur in Texas.
Figure 3--Aphonopelma anax (Charnberlin) 2nd instars. While still getting
nutrients fiom their internal yolk sac, they can be kept together.
15
mo3yxapq waqyou pue sExaL waqnos lno@no.rq p ~ a ~ d s IClq!ssod
ap~
8 ~ a q a m' u o ~ a w 3-- ( u q ~ a q q 3 )XDUD muladouoydp.--~ den
~ p d pus
s ~
'mas
'yyxaaen
--
z
1
.sa!)unoD
(uourrs) a s u m ~tuladouoydv--~1
dtqq
Map 13--Aphonopelma waconum (Chamberlin) -- McLenna11County.
14--Aphonopelma hentzi (Girard) -- Smith (1994) places this species
present only in central and northern Oklahoma. Gertsch & Mulaik (1940)
recorded the species from Dallas, Potter, Starr, and Taylor Counties.
Additional records include Archer, Brown, Clay, Nacogdoches, Travis,
and Wichita. Others have broadly reported A. hentzi as residing in Texas,
Arkansas, Arizona, Kansas, Louisiana and Oklahoma. Male and female
specimens positively identified as A. hentzi have been collected from the
Shreveport, Louisiana area (pers. cornrn. R. West 1996). No range map.
15--Aphonopelmapseudoroseum (Strand) -- Listed only as collected from
an undisclosed location in Texas (Gertsch & Mulaik 1940), Smith (1994)
suggests the species be suspended. Rick C. West (pers. conun) suggests it
be retained pending new information. No range map.
Table 1. Texas Tarantula Species.
Species
Common Name
Location (County)
Texas tan tarantula
Aphonopelma anax (Chamberlin)
Cameron, Kleberg, possibly widespread
Aphonopelma armada (Chamberlin)
blackspot tarantula
Travis
Aphonopelma arnoldi Smith
none
Crosby
Aphonopelma breenei Smith
none
Cameron
Aphonopelma clarki Smith
none
Dallas
Aphonopelma gurleyi Smith
none
Cooke
Aphonopelma harlingenum (Chamberlin)
none
Cameron, Hidalgo
Texas brown tarantula
Aphonopelma hentzi (Girard)
Archer, Brown, Clay, Dallas, Nacogdoches, Potter, Starr, Taylor,
Travis, Wichita, widespread?
Texas blackspot tarantula
Aphonopelma heterops Chamberlin
Hidalgo
Aphonopelma hollyi Smith
none
Lubbock
Aphonopelma moderaturn (Chamberlin & Ivie) Rio Grande gold tarantula
Starr, Webb, Zapata
Aphonopelma pseudoroseum (Strand)
none
Texas
Aphonopelma steindachneri (Ausserer)
brownspot tarantula
Brewster, Pecos
Aphonopelma texense (Simon)
none
Starr
Aphonopelma waconum (Chamberlin)
none
McLennan
MEDICAL IMPORTANCE OF
TARANTULAS
The name "tarantula," in a broad sense, covered mygalomorphs of what
used to be called both the typical and atypical tarantulas in the past
(Kaston 1978). Currently, "tarantula" is reserved solely for members of
the family Theraphosidae (Breene 1995). This is important in a toxicological sense because a few of the non-theraphosid mygalomorph species
are some of the world's deadliest spiders. In Texas, the non-theraphosid
mygalomorphs are possibly more toxic to humans than tarantulas, but a
precise ranking has not been done. Non-theraphosid mygalomorphs are
uncommon in Texas and seldom come into contact with humans. The
following discus-sion will only deal with tarantulas. Because so many
non-native tarantulas are kept as pets, these will be considered as well as
the native species.
Because of the splitting of what was previously reported as A. hentzi, it
is difficult, perhaps impossible to assign a current species name to tarantulas recorded in the literature prior to Smith (1994). Unfortunately, much
of the toxicological and medical literature refers to tarantulas identified as
A. hentzi. Although somewhat confusing, we will refer to those tarantulas
as Aphonopelma hentzi (sensu lato). This indicates that we are referring
to the concept of this species in the broadest sense.
Tarantulas are medically important for both their bites and their
urticating hairs. Their bites can produce problems by the injection of
venoms and by the inoculation of disease-causing agents.
Urticaria
Many New World tarantulas have urticaria-inducing hairs on the back
of the abdomen (Cooke et al. 1972, 1973). These hairs are armed with
barbules (Fig. 1) and can be flicked towards an aggressor by the hind legs
of the tarantula. The flicking of these hairs results in a dorsal abdominal
bald spot. Itching and edema are problems associated with these hairs.
Contact may cause itching and weals that persist for weeks. Oral
antihistamines and topical corticosteroids are the mainstay of treatment
for urticaria (TOMES 1994). Severe reactions may require a week or two
of systemic corticosteroids. Ophthalmia nodosa (inflammation of the
conjunctiva, marked by the formation of a round swelling where each hair
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is embedded) has been reported after handling Brachypelma smithi (F. 0.
P.-Cambridge) (Stulting et al. 1985, Hered et al. 1988).
Tarantulas that are recorded as harmful to humans because of
urticating hairs are listed in Table 2. Cooke et al. (1972) suggested that
tarantulas that are aggressive and have potent venoms generally do not
have urticating hairs.
Bites
The biting apparatus of the tarantula consists of paired chelicerae. Each
chelicera has a broad base which houses the venom gland and terminates
in a large moveable fang. When the spider bites, it presses the sharp fangs
into the body of the victim and produces two separate punctures. Muscles
in the chelicerae can press the venom sacs, which forces the venom out the
small openings near the ends of the fangs. Tarantulas can control the
amount of venom (if any) that is injected. The amount of venom can also
be altered by the amount of time the fangs remain in contact with the
victim. This may account for the apparent lack of toxicity to humans of
bites from some of the larger tarantulas known to kill smaller vertebrates.
Possibly, the spider is able to determine that the larger human is not food
and not worth injecting with precious venom, especially when a bite with
centimeter long fangs will get an immediate withdrawal response.
Apparently, the age and sex of the spider can also affect the volume and
type of venom present. Likewise, the volume and potency of venoms can
be different between recently fed spiders and those not having fed (the
latter can involve more venom being injected).
In addition to the toxicity of the venoms, allergic responses are a
problem. It seems to be general knowledge that anyone allergic to insect
venoms could be allergic to spider venoms. We are unaware of any
scientific evidence to corroborate such a statement, other than that an
individual with an allergy is at greater risk of having a second or third
substance allergy. Venoms are complex and often quite different between
dissimilar taxa. It should not be assumed that a person allergic to honey
bee venom will be allergic to a specific tarantula venom. Until we know
that the venoms have specific allergens in common, we cannot assume that
they will cause similar reactions. It has been suggested (American Academy of Allergy) that immunotherapy for individuals with hymenopteran
sting allergies be specific (i.e., if you are allergic to honey bee venom,
your therapy should be with bee venom, not wasp or hornet venom).
More is known about the composition of the venom of A. hentzi (sensu
lato) than any other member of the family. While not known to be lethal to
humans, the venom of A. hentzi is lethal to both mice and insects
(Schanbacher et al. 1973a; and citations contained therein). The major
toxin in the venom is reported to be a necrotoxin, which causes necrosis of
heart muscle when injected into mice (Lee et al. 1974). Chan et al. (1975)
reported adenosine nucleotides from the venom and noted a synergistic
toxic effect on mice from at least one of these components with the necrotoxin. Other components in the venom include hyaluronidase (Schanbacher et al. 1973b), which is thought to function as a "spreading factor"
for the toxin; g-amino butyric acid, which could affect the transmission of
neural messages; and a small peptide, which might have toxic activity or
influence the toxic effects of the necrotoxin (Chan et al. 1975). The role of
the polyamines (especially sperrnine), which are also present in tarantula
venom, is uncertain (Cabbiness et al. 1980).
Tarantula venoms are complex and can cause several reactions besides
allergies. Extreme pain (requiring medical attention), necrotic lesions, and
death have been recorded. Gertsch (1949) stated that the bite of a common
tarantula of Central America, Sericopelma rubronitens (Ausserer), causes
symptoms in humans that persist for several hours. Biicherl (1971)
reported that deeper and larger wounds can result from bites to humans by
Acanthoscurria, Megaphobema, Xenesthis, Theraphosa, and some
Avicularia, Phormictopus, and Pamphobeteus. Southcott (1976) listed
two Australian species of Selenocosmia, bites from which have caused
severe effects to humans.
From the chapter by Bettini & Brignoli (1978), it is clear that many
tarantulas have venoms that can result in death or less severe symptoms
for smaller vertebrates. Records are fewer of bites on humans. A man
suffered muscular spasms and collapse following a bite by Selenocosmia.
Sericopelma was recorded as being considerably toxic to humans. A death
was attributed to a bite from a Phormictopus, and US Aphonopelma spp.
have produced bites resulting in localized tissue alterations, edema, pain,
and lynphadenitis in humans.
Russell & Gertsch (1982) reported Aphonopelma from California as
having been implicated in producing necrotic lesions in humans. These
authors also noted that not all cases attributed to spider bites were done so
correctly, and listed many diseases that have symptoms that are similar to
some spider bites. Generally with large tarantulas this point is not that
important, as it is difficult not to notice the spider when you are bitten.
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Russell (1987, 1988) listed the following genera of tarantulas from
which significant bites on humans are known: Selenocosmia, Pamphobeteus, and Aphonopelma. He also listed numerous other non-theraphosids implicated in human envenomations. Layzell (1989) reported the
results of a bite she received from a Pterinochilus murinus Pocock female, which included a night's stay in the local hospital.
No deaths were attributed to spider envenomations during 1989-1993 in
the US (Litovitz et al. 1990, 1991, 1992, 1993, 1994). The results of bites
reported during this five-year period are summarized in Table 3. For
comparative purposes, all envenomations reported to be by spiders are
included. Udortunately, the reports on the American Association of Poison Control Centers National Data Collection System did not have the
records for tarantulas separated from "other spiders" until 1989. Clearly,
tarantulas do not pose the potential health threats that recluse and widow
spiders do.
The puncture of human skin by tarantula fangs can result in the inoculation of disease-causing organisms that might have been on the intact
skin before the bite or that were present on the fangs. In addition to infection from the common skin-inhabiting bacteria and fungi, significant
infection involving other organisms is possible. Tetanus has been recorded in at least one case involving a spider (Lawrance 1825).
A single case of tularemia transmission by a spider bite was documented by Rowland & Grifliths (1988). The spider involved was not a
tarantula, but the authors suggested that tularemia should be kept in mind
when treating spider bites in areas where it is endemic. The symptoms
started one day after the bite with fever, chills, and localized inflammation. Right axillary adenopathy, severe myalgia, leukocytosis, and pulmonary signs then developed. The tularemia titers rose six-fold during this
time. The adult male patient recovered after treatment with antibiotics.
There are no first aid measures (such as tourniquets or venom extraction) of demonstrated value for tarantula bites. US tarantula venom is not
highly dangerous to humans, but they have large fangs and can produce a
paidid bite. Bites can be almost painless or produce a deep, throbbing
pain for an hour or so (Wong et al. 1987). An ice cube can be placed on
the bite for a short period of time to reduce the pain. The bite site should
be washed thoroughly with soap and water. Tetanus prophylactics should
be considered. Immobilization and elevation of the affected part may be
required (TOMES 1994). Treatment is symptomatic and supportive;
managing itching, pain, inflammation, and infection. Various antibiotics,
antihistamines, analgesics, and corticosteroids have been used (Wong et
al. 1987). For itching/inflammation, Russell (1 98 8) recommended a
corticosteroid-antihistamine-anestheticcream.
Table 2.--Tarantulas recorded as being harmful to humans (most species
of which are kept as pets in the US).
Taxa
~istribution
Source
Acanthoscuwia
Avicularia metallica Ausserer
Brachypelma albopilosum Valerio
South America
Cayenne
Costa Rica
Mexico
Lasiodora
Phrixotrichus (=Grammostola)
Psalmopoeus
Pseudotheraphosa apophysis Tinter
Theraphosa blondi (Latreille)
South America
South America
South America
South America
South America
Cooke et al. 1972,1973
Peters 1992
Peters 1992
Cooke et al. 1972, 1973
Hered et al. 1988, Stulting
et al. 1985, Peters 1992
Cooke et al. 1972, 1973
Cooke et al. 1972, 1973
Peters 1992
Portillo 1995
Peters 1992, Alroth 1995
With Urticating Hair
With Significant Bites
Acanthoscum'a
Aphonopelma chalcodes Charnberlin
Aphonopelma echina (Chamberlin)
Aphonopelma hentzi (Girard)
Aphonopelma sp.
Aphonopelma
Avicularia
Chaetopelma olivaceum (C. L. Koch)
Hapalopus sp.
Harpactirella lightfooti Percell
Heteroscodra maculata Pocock
Megaphobema
Pamphobeteus
Phormictopus cancerides (Latreille)
Phrixotrichus iheringi (Keyserling)
Poecilotheriafasciata (Latreille)
Poecilotheria ornata Pocock
Poecilotheria regalis Pocock
Pterinochilus
Pterinochilus murinus Pocock
Brazil
USA., Mexico
USA.
USA
New World
USA
Brazil
Syria, eastern Africa
South America
Africa
Africa
Brazil
Brazil
Bucherl 1971
Peters 1992
Peters 1992
Peters 1992
Peters 1992
Russell & Gertsch 1982
Bucherl 1971, Alroth 1995
Peters 1992
Peters 1992
Alroth 1995, Filmer 1991
Alroth 1995
Bucherl 1971
Biicherl 1971
Russell 1987, 1988
Brazil & West Indies Cooke et al. 1972
Biicherl 187 1, Alroth 1995
South America
Bettini & Brignoli 1978
Peters 1992
Brazil
Peters 1992
Schmidt 1989, Alroth 1995
Sri Lanka
Portillo 1995
Sri Lanka
Portillo 1995
India
Africa
Peters 1992, Schmidt 1989
Africa
Layzell 1989
R
Table 2.--Continued.
Selenocosmia crassipes (Koch)
Selenocosmia stirlingi Hogg
Sericopelma rubronitens (Ausserer)
Theraphosa
Xenesthis
Australia
Southcott 1976
Peters 1992
Australia, New Guinea
Southcott 1976
Peters 1992, Alroth 1995
Central America
Gertsch 1949
Central & South America
Bettini & Brignoli 1978
Peters 1992
Brazil
Bilcherl 1971
Brazil
Biicherl 197 1
Table 3.-- Annual average numbers of spider bites fkom the U.S.A. reported to
Poison Control Centers during 1989-1993 (Litovitz et al. 1990-1994). The
outcome of the bites were not always known and therefore the totals for the
columns labeled "Seriousness of Outcome of Bites" will not equal the total
number of bites.
Taxi
# Bites Treated in Health
Seriousness of
reported Care Facility
Outcome of Bites
None Minor Moderate Major Death
0
272
9
305 1,140
widow
2,404
813
spiders
recluse
spiders
1,453
8 13
other
spiders
560
113
55
11
tarantulas
1
3
32
1
0
0
CI
Figure 4.--Silhouette of urticating hair found on the abdomens of
tarantulas from Texas (Type I hair from Cooke et al. 1972).
CaU for Tarantulas
We urge anyone encountering tarantulas in Texas, or anywhere else in
the US, to send specimens, if possible (dead or alive) to Dr. R. G. Breene,
PO Box 1617, Artesia, New Mexico 8821 1. Live specimens can be packed semi-firmly in slightly damp paper towels inside a deli cup or margarine tub with a few small air holes. The cup can then be inserted into a
small box, surrounded with styrofoam peanuts or newspaper. Live specimens should be sent USPS priority mail.
Dead specimens ideally are "cured" in 55% or higher ethanol, methanol
or denatured alcohol (not isopropyl) for three weeks to a month. They can
then be placed in a self-sealing plastic bag with a few drops of alcohol,
placed into another bag, and sent in a small box parcel post.
The specimens will be forwarded to the appropriate taxonomists and
can greatly assist in a more accurate accumulation of knowledge of US
tarantulas.
We thank Dr. Norman V. Homer and Margaret E. Janowski-Bell
(Midwestern State University) for sending us their unpublished manuscript on tracking tarantulas with radio telemetry. We also thank Miep R.
O'Brien for extensive editing of this manuscript, Rick C. West and Dr.
Norman I. Platnick for valuable information in a number of areas.
Societies
Part of the following list of arachnid and tarantula societies, clubs and
publications is taken in part from an Internet page:
(http://members.aol.cornljccoke/society.html) entitled: Arachnological
Publications, Internet Discussion Groups / Databases, and Societies of the
World. Individuals with web searching capabilities should consult that
page for changes and updates.
ARACHNID (Internet listserver)
The arachnid mail list is a computer forum for the discussion of
spiders, scorpions, and other members of the class Arachnida. Of particular interest is the husbandry and breeding of tarantulas and scorpions
with emphasis on the needs of these animals when kept as pets. You may
31
subscribe to this list by sending a command (in the text part of the message) to: majordomo@bga.com. This command should read: subscribe
arachnid "your name" <your E-mail address> (the " " and < > are parts of
the command).
AMERICAN TARANTULA SOCIETY (ATS)
The American Tarantula Society, founded in 1991 (a non-profit organization, first published 1992), to promote the study and the dissemination
of information primarily concerning the arachmd infiaorder Mygalomorphae, especially Theraphosidae, but also to include other Araneae, and
other arachnid orders (exclusive of Acari). It is intended to maintain a
flow of information and cooperation between enthusiasts, students, teachers, and professional arachnologists. The Forum Magazine is published
six times per year ($15 yr US). Mygalomorph is published occasionally
when a sufficient amount of material has been accepted, and is not regularly scheduled for publication in any given year. The Forum Magazine
is written in a style easily understood by the hobbyist and contains articles, news items, advertisements, and exchange of materials. The
Mygalomorph is a scientific journal containing peer-reviewed articles,
written for scientists and advanced hobbyists. For further information
contact: ATS, PO Box 1617, Artesia, New Mexico 882 11 USA. Email:
7 1024.2662@compuserve.com - Phone (505) 748-2483. Additional information can also be found on the ATS home page at:
http:l/ www.cowboy.net/-spiderlATS .html
A society with this name was first started in 1978 and lasted until 1984.
That society produced the newsletter Tarantula Times. The newsletter, as
well as the society, had difficulty staying active. There were 21 issues of
the newsletter published from 1978 to 1984. The purpose of the American
Tarantula Society was to provide the opportunity for professionals and
laypersons to share their knowledge of the tarantula, to encourage the
study of the tarantula as it gains popularity as a pet, and to eliminate misunderstandings concerning the tarantula. A later goal of the society was to
have the pet industry breed their own supply of tarantulas, eliminating the
plunder of natural habitats and populations. Although that society is no
longer active, the problem of wild tarantula collecting for resale continues.
AMERICAN ARACHNOLOGICAL SOCIETY (AAS)
Scientific society, publishes Journal of Arachnology and newsletter. $30
yr, $20 students, $80 institutional ($90 outside USA) to: Dr. N. I. Platnick, AMNH, Central Park West at 79th St., New York, NY 10024 USA.
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BRITISH ARACHNOLOGICAL SOCIETY (BAS)
Scientific society, publishes the Bulletin of the British Arachnological
Society and a newsletter. USA £18 yr. UK currency or IPO to: S. H.
Hexter, 7 1 Havant Road, Walthamstow, London, E 17 3JE England.
THE BRITISH TARANTULA SOCIETY (BTS)
Originally founded as the British Tarantula Fellowship International in
1984, this group changed their name in 1986 to the British Tarantula
Society. l b s international group aims to further the study, keeping, and
breeding of tarantulas and scorpions and to educate the public that spiders
are not to be feared, but admired and studied. The quarterly Journal of the
British Tarantula Society was first published in 1986. This group holds an
Annual Show for members, where they have talks and discussions, and
members can voice their views and exhibit their tarantulas. For further
information concerning the society or membership contact the Secretary:
Mrs. Ann Webb, The British Tarantula Society, 81 Phillimore Place,
Radlett, Hertfordshire, WD7 8NJ, United Kingdom. Telephone: 0923 856
07 1. See also the BTS home page at:
http://www.wsu.edu:8080/-d0152871/BTS.html.
SONORAN ARTHROPOD STUDIES INSTITUTE (SASI)
SASI's goals are to increase understanding and appreciation of arthropods. Instar, Backyard Bugwatching, newsletter, $35 yr: SASI; PO Box
5624, Tucson, Arizona 85703 USA (520) 883-3945.
CENTRAL CALIFORNIA ARACHNID SOCIETY (CCAS)
Monthly meetings, special events, quarterly newsletter, lending library.
$10 yr. Jay Milles 4082 N. Benedict, Fresno CA 93722-4559 (209) 2251802, email carol~devine@csufresno.edu
INVERTEBRATA
Quarterly, covers many aspects of invertebrates, concentrating on successful care in captivity. $25 yr. Mascariiio, PO Box 2072 1., Los Angeles, CA 90006 ph. (213) 227-6566.
ROCKHAMPTON ARACHNOLOGICAL SOCIETY (RAS)
Quarterly Journal. $10 (Australian) plus mailing costs. Contact Doug
Wallace, 50 Naughton St., Wandal, Rockhampton 44700 Australia.
TARANTULA CLUB NEDERLAND (TCN)
This Dutch tarantula club is interested in captive breeding and care of
tarantulas. A newsletter was started in 1987. Interested individuals should
contact the group for additional information by writing: Mr. Rob. J.
Dumont, Tarantula Club Nederland, Waddenstraat 217, 2036 Le
Haarlem, The Netherlands.
Bibliog,aphy Of The TheraPhosidae
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mygale spider (Psalmopoeus cambridgii Poc.) Trans. Edinburg F.
Nat. Micr. Soc. 5: 402-406.
Adams, J. 1908. Further note on the
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Agnew, C. W., D. A. Dean & J. W.
Smith. 1985. Spiders collected from
peanuts and non-agricultural habitats in the Texas west cross-timbers.
Southwest. Nat. 30: 1-12.
Alberti, G., B. A. Afkelius & S. M. Lucas. 1986. Ultrastructure of spermatozoa and spermatogenesis in
bird spiders (Theraphosidae, Mygalomorphae, Araneae). J. Submicrosc. Cytol. 18: 739-753.
Anderson, J. F. & K. N. Prestwich.
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and above maximal aerobic capacity
in a theraphosid (tarantula) spider,
Brachypelma smithi (F. 0 . PickardCambridge). J. Comp. Physiol. 155:
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Andrews, J. S. 1930. The digestion of a
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Angersbach, D. 1975. Oxygen pressures in haemolymph and various
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Verh.
Dtsch. Zool. Ges. 1974: 277-280.
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Solifbgen, Palpigraden). Arch.
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Arrigoni, E. & A. Fanfani. 1974.
Caratterizzazione biologica del veneno della migale africana Phoneyusa
lesserti Dresco 1. Attivita "in vitro"
della muscolatura dell'apparato gastroenterico. I1 Fannaco, p. 4.
Arrigoni, E., A. Fanfani, A. Montani,
R. Scelsi & R. F. Villa. 1976. Effetti
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Univ. Pavia 4: 1-11.
Aruchami, M. & G. Sundara Rajulu.
1978. A study on the lipid component of the haemolymph of a spider
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Babu, K. S. 1969. Certain histological
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Poecilotheria. Am. Tarantula Soc.
Forum 5: 5-8.
Bustard, R. 1996. Poecilotheria care in
captivity. Am. Tarantula Soc. Forum
5: 44-48.
Charpentier, P. 1992. Heterscodra
maculata. J . British Tarantula Soc.
8(2): 11-19.
Charpentier, P. 1992. Heterscodra
maculata Pocock, 1899. Arachnides,
Bull. G. E. A. 15: 25-30.
Charpentier, P. 1992. The genus Avicularia. Exothermae l(1): 1-49.
Charpentier, P. 1992. Pterinochilus
murinus, morphology - biologybehaviour. Exothermae l(2): 1-72.
Cloudsley-Thompson, J. L. 1982. Tarantulas. Country Side 25: 26-29.
Cokendolpher, J. C. 1992. More tarantula bites. Am. Tarantula Soc. Forum 1: 56-59.
Dunlop, J. 1991. Venom research: the
story so far. J. British Tarantula Soc.
7(1): 18-19.
Dunlop, J. 1991. An African with hollow legs. J. British Tarantula Soc.
7(2) 8-10.
Dunlop, J. 1993. A functional explanation for posturing during tarantula
ingestion? Am. Tarantula Soc. Forum 2: 95-97.
m
m
~1
Dunlop, J. 1994. The mouthparts of tarantulas. Am. Tarantula Soc. Forum
3: 2-4.
Dunlop, J. 1995. Good models for a
poor species concept? Am. Tarantula Soc. Forum 4: 72-74.
Dunlop, J. 1996. Swimming in tarantulas. Am. Tarantula Soc. Forum 5:
79-8 1.
Eckardt, D. 1991. Mating tarantula
spiders: Always the new. J. British
Tarantula Soc. 7(2): 7-8.
Filmer, M. R. 1991. Southern Africa
spiders: An identification guide.
Struik Publ., Cape Town, South Africa 128 pp.
Gallon, R. 1991. Hybridslcaptive
breeding. J. British Tarantula Soc
7(2): 10-11.
Grimes, M. 1995. The proper way to
kill a king baboon. Am. Tarantula
Soc. Forum 4: 113-114.
Harp, J. 1992. On the status of a rare
tiny mygalomorph. Am. Tarantula
Soc. Forum 1: 64-66.
Harp, J. 1994. Passing tarantula water.
Am. Tarantula Soc. Forum 3: 91-92.
Harvey, D. A. D. 1989. The classification of mygalomorph spiders. part
11: Nemesiidae, Theraphosidae and
Paratropidae. J. British Tarantula
SOC.5(1): 21-26.
Harvey, D. A. D. 1991. The classification of mygalomorph spiders, part
111. J. British Tarantula Soc. 7(1):
14-16.
Harvey, D. A. D. 1991. The classification of mygalomorph spiders, part
IV. J. British Tarantula Soc. 7(2)
11-14.
Hurst, J. M. 1994. Imperial spiderlings. Am. Tarantula Soc. Forum 3:
74-76.
Hurst, J. M. 1995. To raise or not to
raise spiderlings. Am. Tarantula
Soc. Forum 4: 43-45.
Isler, Hans-Peter. 1991. Successful
breeding of bird-eating spiders in
captivity: Megaphobema robusta. J.
British Tarantula Soc. 7(1): 12-13.
Johnson-Delaney, C. 1991. Basic care
of tarantulas. J. Small Exotic Anim.
Med. l(1): 19.
Layzell, J. 1989. My first bite! J. British Tarantula Soc. 5(1): 15-16.
Layzell, J. 1991. Collecting tarantulas
in Ecuador. J. British Tarantula Soc.
7(2): 5-6.
Liczbinsky, B. 1994. Superman spider
snatches bus. Am. Tarantula Soc.
Forum 3: 155-156.
Marshall, S. D. 1995. Nomad pinktoes? Am. Tarantula Soc. Forum 4:
40-4 1.
Marshall, S. D. 1996. Old dog learns
new trick. Am. Tarantula Soc. Forum 5: 114-116.
Mason, T. 1993. Tarantula scavengers.
Am. Tarantula Soc. Forum 2: 74-75.
Miranda, L. 1994a. A bite in time
heals mine. Am. Tarantula Soc. Forum 3: 15-18.
Miranda, L. 1994. Killer tarantula police patrol. Am. Tarantula Soc. Forum 3: 92-93.
Moore, B. H. 1993. Pinktoe tarantulas.
Am. Tarantula Soc. Forum 2: 39-40.
Moore, B. H. 1994. Red rumped cannibals. Am. Tarantula Soc. Forum 3:
14-15.
O'Brien, M. R. 1996. On mice, men
and the Post Ofice (shipping tarantulas). Am. Tarantula Soc. Forum 5:
52-53, 56.
O'Brien, M. R. 1996. The Florida
anomaly. Am. Tarantula Soc. Forum
5: 83-85.
Peck, W. B. 1992. The tarantella. Am.
Tarantula Soc. Forum 1: 53-56.
Pkrez-Miles, F. & F. G. Costa. 1994.
Acanthoscurria atrox incorporates
urticating hair into its shedding mat.
Am. Tarantula Soc. Forum 3: 63-64.
Perez-Miles, F. & F. G. Costa. 1995.
Increased male activity during
stormy weather. Am. Tarantula Soc.
Forum 4: 153-154.
Portillo, R. 1995. Misidentification,
handling and envenomation. In, Invertebrates in captivity conf., Sonoran Arthropod Studies Inst., Tucson,
Arizona 186 pp.
Reger, B. H. 1994. Venomous hazards
down under. Am. Tarantula Soc.
Forum 3: 126,
Reger, B. H. 1994. Underwater Amazon adventure: death defLing diving
tarantula emulates submarine. Am.
Tarantula Soc. Forum 3 : 152-154.
Reger, B. H. 1995. Tarantula one-adays. Am. Tarantula Soc. Forum 4:
45-46.
Reger, B. H. 1995. Those perplexing
blonds. Am. Tarantula Soc. Forum
4: 85-86.
Schaefer, R. 1996. Bemerkungen zu
einigen gattungen der familie
Theraphosidae inklusive der beschreibung einer neuen art aus Paraguay: Tmesiphantes spinopalpus sp.
n. (Araneida: Theraphosidae). Arthropoda 4: 24-3 1.
Schmidt, G. E. W. 1990. Zur kenntnis
der gattung Mygafarachne (Araneida: Theraphosidae). Arachnol.
Anz. 9: 8-12.
Schmidt, G. 1991. Revision der gattung Megaphobema (Araneida:
Theraphosidae;
Theraphosidae).
Arachnol. Anz. 13/91: 11-13.
Schmidt, G. 1991. Eina neue Paraphysa art aus Equador (Araneida:
Theraphosidae;
Theraphosinae).
Arachnol. Anz. 10191: 8-12.
Schmidt, G. E. W. 1991. Eine neue
vogelspinnen aus Honduras Phormictopus atrichomatus sp. n. (Araneida: Theraphosidae: Theraphosinae). Arachnol. Anz., nr. 11: 7-10.
Schmidt, G. E. W. 1992. Das mannchen von Euathlus truculentus
Ausserer 1875 (Araneida: Theraphosidae: ~heraphosinae). Arachn.
Anz. 3: 9-13.
Schmidt, G. E. W. 1992. Brachypelma
auratum sp. n., die sogenannte
hochlandform von Brachypelma
smithi (Araneida: Theraphosidae:
Theraphosinae). Arachnol. Anz.
3(8): 9-14.
Schmidt, G. 1992. Brachypelma Simon
1890 oder Euathlus Ausserer 1875?
(Araneida: Theraphosidae; Theraphosidae). Arachnol. Anz. 1: 9-1 1.
Schmidt, G. E. W. & P. Klaas. 1993.
Eine neue Brachypelma-Spezies aus
Mexiko (Araneida: Theraphosidae:
Theraphosinae). Arachnol. Anz.
4(5): 7-9, 11-13.
Schmidt, G. E. W. & P. Klaas. 1994.
Eine neue Brachypelma species aus
Mexico Brachypelma boehmei sp. n.
(Araneida: Theraphosidae: Theraphosinae). Arachnol. Mag. 2(7): 715.
Schmidt, G. E. W. 1994. Avicularia
urticans sp. n. (Araneida: Theraphosidae: Aviculariinae), eine vogelspinnenart aus Peru. Arachnol.
Mag. 2(6): 1-7.
Schmidt, G. E. W. 1994. Grammostola
pulchripes ist ein synonym von G.
grossa. Arachnol. Mag. 2(7): 23.
Schmidt, G. E. W. 1994. Eine neue
Paraphysa-Art aus Brasilien (Araneida: Theraphosidae: Theraphosinae), Paraphysa horrida sp. n.
Arachnol. Mag. 2(12): 1-7.
Schmidt, G. E. W. 1994. Systematics
of Theraphosidae: some new results.
Am. Tarantula Soc. Forum 3: 98101.
n
n
-
n
Schmidt, G. E. W. & R. H. Krause.
1995. Eine neue art der Theraphosidae aus Vietnam: Selenopelma kovariki gen. et sp. n. (Araneida:
Theraphosidae: Selenocosmiinae).
Arthropoda 3 : 2 1-24.
Schmidt, G. E. W. 1995. Chromatopelma gen. n.; eine neue gattung der Theraphosidae (Arachnida:
Araneida: Theraphosidae: Theraphosinae). Arthropoda 3 : 25-26.
Schmidt, G. E. W. & V. von Wirth.
1996. Haplocosmia nepalensis gen.
et sp. n., die erste vogelspinne aus
Nepal (Araneida: Theraphosidae:
Selenocosmiinae). Arthropoda 4:
12-15.
Schmidt, G. E. W. & F. Kovarik. 1996.
Nesipelma insulare gen. and sp. n.
from the Nevis Island, Lesser Antilles (Arach-nida; Araneida: Theraphosidae). Arachnol. Mag. 4(6): 19.
Sherberger, F. 1993. Keeping tarantulas healthy: a quick guide for pet
stores. Am. Tarantula Soc. Forum 2:
3 1-33.
Sherberger, F. 1995. Sexing molts.
Am. Tarantula Soc. Forum 4: 181182.
Smith, A. M. 1988. Lyrognathus robustus, a new species of theraphosid
spider from Malaysia. J. British Tarantula Soc. 4(2): 15-19.
Smith, A. M. 1990. Taxonomic differences between the closely related
species Grammostola spatulata
Cambridge 1897 and Grammostola
cala Chamberlin 1917 from Chile.
J. British Tarantula Soc. 6(1): 1921.
Smith, A. M. 1991. A revision of the
genus Megaphobema Pocock, 190 1
(Araneida; Theraphosidae). J. British Tarantula Soc. 6(1): 14-19.
Smith, A. M. 1992. In defence of Raven's decision to make the genus
Brachypelma Simon 1891 a junior
synonym of Euathlus Ausserer 1895.
J. British Tarantula Soc. 7(3): 1419.
Smith, A. M. 1992. Redescription of
the Aviculariinae species Avicularia
minatrix Pocock 1903. J. British Tarantula Soc. 8(2): 22-26.
Smith, A. M. 1993. A new mygalomorph spider from Mexico (Brachypelma, Theraphosidae, Arachnida)
Brachypelma baumgarteni n. sp. J.
British Tarantula Soc. 8(4): 14-19.
Smith, A. M. 1993. Taxonomy focus.
J. British Tarantula Soc. 9(1): 1318.
Smith, A.M. 1993. How to keep tarantulas. Fitzgerald Publishing, London, 17pp.
Smith, A. M. 1994. A study of the genus Phormingochilus with a redescription of the species described by
Pocock (Araneae, Mygalomorphae,
family Theraphosidae, subfamily
Ornithoctinae). J. British Tarantula
SOC.9(4): 15-22.
Smith, R. 1992. Some thoughts on the
treatment of tarantula fungal infections. Am. Tarantula Soc. Forum 1:
30-32.
Stahnke, H. L. 1965. Tarantula regeneration. Turtox News 43: 236.
Stanton, B. 1993. Ornamentals: they're
not just for Christmas trees anymore. Am. Tarantula Soc. Forum 2:
60-6 1.
Stropes, D. 1994. The miracle spider,
Am. Tarantula Soc. Forum 3: 95-97.
Stropes, D. 1996. African beauties.
Am. Tarantula Soc. Forum 5: 4 1-42.
Sullivan, M. L. 1993. Tarantulas in
your refrigerator. Am. Tarantula
Soc. Forum 2: 105-106.
Tinter, A. 1991. Eine neue vogelspinne
aus Venezuela Pseudotheraphosa
apophysis n. gen. n. sp. (Araneae:
Theraphosidae:
Theraphosinae).
Arachnol. Anz., 16: 6-10.
von Wirth, V. 1991. Eine revision der
gattung Ornithoctonus Pocock, 1892
(Araneida: Theraphosidae: Ornithoctoninae). Arachnol. Anz. 12: 58.
von Wirth, V. 1991. Eine neue Vogelspinnenart aus Vietnam. Haplopelma schmidti sp. n. (Araneae:
Theraphosidae: Ornithoctoninae).
Arachnol. Anz. 18: 6-1 1.
West, R. C. 1984. An introduction to
the tarantula spiders of Trinidad,
Wisconsin. Living World 19831984: 54-58.
West, R. C. 1992. Not to everyone's
taste. Am. Tarantula Soc. Forum 1:
28-30.
West, R. C. 1993. Dealing with human
nature. Am. Tarantula Soc. Forum
2: 13.
West, R. C. 1993. Warning: do not eat
the tarantulas. Am. Tarantula Soc.
Forum 2: 3-4.
Williams, J. 1993. Adventures with
Australian mygalomorphs and other
creatures. Am. Tarantula Soc. Forum 2: 108-110.
Current popular Books on Tarantulas
Baxter, R. N. 1993. Keeping and
breeding tarantulas. Chudleigh Publishing, Essex, England. 89 pp. 32
plates.
de Vosjoli, P. 199 1. Arachnomania,
the general care and maintenance of
taran-tulas and scorpions. Advanced
Vivarium Systems, Lakeside, California. 80 pp.
Hancock, K. & J. Hancock. 1992.
Tarantulas: Keeping and breeding
arachnids in captivity. R & A
Publishing Limited, Somerset 147
PPMarshall, S. D. 1996. Tarantulas and
other arachnids. Barron's Hauppauge, New York 104 pp.
Reger, B. 1995. Tarantulas as a new
pet. T.F.H. Publications, Inc. Neptune City, New Jersey. 64 pp.
Schmidt, G. E. W. 1993. Vogelspinnen: vorkommen, lebensweise, haltung und zucht, mit bestimmungsschliissein fiir alle gattungen,
Vierte Auflage, Landbuch Verlag,
Hannover, 15 1 pp.
Schultz, S. A. 1984. The tarantula
keeper's guide book. New York,
Sterling, 1-128.
Smith, A. M. 1988. Tarantula Classification and Identification Guide.
Fitzgerald Publishing, London 178
PP.
Smith, A. M. 1990. Baboon spiders,
Trantuals of Africa and the Middle
East. Fitzgerald Publishing, London
142 pp.
Smith, A. M. 1994. Tarantula spiders,
tarantulas of the USA and Mexico.
Fitzgerald Publishing, London 196
PP.
Turbang, P. 1993. Guide des Mygales
klevees en terrarium. Delachaux et
Niestlk, Lausanne, Switzerland 157
PP
Webb, A. 1992. The proper care of tarantulas. T.H.F. Publications, Inc.
Neptune City, New Jersey. 288 pp.