p their medical importance, ud world-wide bibliography to m the

TARANTULAS OF TEXAS P THEIR MEDICAL IMPORTANCE,

' UD WORLD-WIDE BIBLIOGRAPHY TO

m THE THERAPHOSIDAE (ARANEAE)

n

R. G. Breene, D. A. Dean, J. C. Cokendolpher, and B. H. Rege

TARANTULAS OF TEXAS THEIR MEDICAL IMPORTANCE,

AND WORLD-WIDE BIBLIOGRAPHY TO THE THERAPHOSIDAE (ARANEAE)

R. G. ~reene' , D. A. ~ e a n ~ , J. C. cokendolpher3, and B. H. ~ e ~ e r ~

First Printing, October 1996 Copyright O By The American Tarnntzila Society

I

PO Box 1617 Artesia, New Mexico 882 1 1 - 16 17 USA

1 All rights reserved

1 PO Box 1617, Artesia, New Mexico 8821 1-1617 USA 2 Department of Entomology, Texas A&M University, College Sta- tion, Texas 77843-2475 USA

2007 29th Street, Lubbock, Texas 7941 1 USA 4 4616 South State Road 9, Greenfield, Indiana 46140 USA

Copyright 0 1996 By the American Tarantula Society. All rights I

reserved. Reproduction in whole or in part by any means or methods is prohibited unless authorized in writing by the copyright

R 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.

7

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

m 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 com- plicates 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 Thera- phosidae (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). Tree- dwelling 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 exten- sively 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 (1 996) 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 undis- tinguished 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 temp- eratures. 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

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 Gastro- phryne living in the burrows of Aphonopelma sp. in the US (pers. comm. 199 1, 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

m 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 Brachy- pelma aureoceps (Charnberlin 19 17). Smith (1 994) 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 CI

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 m 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 n 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.

Predation Ecology and Feeding The significance and impact tarantulas may have upon ecosystems have

never been investigated. If viewed from an agricultural standpoint, taran- n tulas fit into the mobile, visually acute, or type MV predators along with the bulk of the spiders and certain insects including mantids and hanging- flies (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

rn ( 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.

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, R

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 n (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 n 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. n 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. rn

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).

Seasonal History m

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 F

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) w thought that spring males might be a fail-safe device in case of an early

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 eiLher A. 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 avail- n able. 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

I 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 fash- ioned for the purpose.

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 join- ed 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 Araneomor- phae 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.

Figure 1 .--Cladistic hypothesis showing the relationships between the - Arachnids. Trigonotarbida is an extinct arachnid order. Adapted from Coddington & Levi (1 99 1).

12 n

o tLer Arachida

n

'~'rigonotarbida

S chizornida

pedipalpi

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L b l y PY gi

~ e s o t h e l a e ~ i ~ h i s t i i d a e

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Aranea

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ro iaetidae

L~ornicephalae

C ytaucheniidae

astelloidin

Actinopodidae

Figure 2.--Cladistic hypothesis showing the relationships between the Mygalomorphae families. Adapted from Raven (1 985) 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 n

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, 190 1 and Apnchepelma Smith, 1994 (one species from Arizona). Genera synonymous with Aphonopelma include Chauno- pelma, 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

n by the International Commission of Zoological Nomenclature (1 99 1).

The generic names Delopelma, Dugesiella, and Rhechostica are occasionally still erroneously used in the scientific literature by a few

n 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 rn

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 (1 994) also points out that both type specimens are apparently lost, and they may have

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 con- firm 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

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1 z .sa!)unoD

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Map 13 --Aphonopelma waconum (Chamberlin) -- McLenna11 County.

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--Aphonopelma pseudoroseum (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)

Aphonopelma anax (Chamberlin) Texas tan tarantula 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

Aphonopelma hentzi (Girard) Texas brown tarantula Archer, Brown, Clay, Dallas, Nacogdoches, Potter, Starr, Taylor, Travis, Wichita, widespread?

Aphonopelma heterops Chamberlin Texas blackspot tarantula 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

" 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.

n 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

r 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 Aca- demy 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 m

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 (Schan- bacher 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 n 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 n 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 n 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 m

Avicularia, Phormictopus, and Pamphobeteus. Southcott (1976) listed two Australian species of Selenocosmia, bites from which have caused severe effects to humans. n

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.

Russell (1987, 1988) listed the following genera of tarantulas from which significant bites on humans are known: Selenocosmia, Pampho- beteus, 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 Poi- son 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 docu- mented 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 pul- monary 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 extrac- tion) 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-anesthetic cream.

Table 2.--Tarantulas recorded as being harmful to humans (most species of which are kept as pets in the US). Taxa ~is t r ibut ion Source R

With Urticating Hair

Acanthoscuwia South America Cooke et al. 1972,1973 Avicularia metallica Ausserer Cayenne Brachypelma albopilosum Valerio Costa Rica

Mexico

Lasiodora South America Phrixotrichus (=Grammostola) South America Psalmopoeus South America Pseudotheraphosa apophysis Tinter South America Theraphosa blondi (Latreille) South America

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 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 Bucherl 1971 USA., Mexico Peters 1992 USA. Peters 1992 USA Peters 1992 New World Peters 1992 USA Russell & Gertsch 1982 Brazil Bucherl 197 1, Alroth 1995 Syria, eastern Africa Peters 1992 South America Peters 1992 Africa Alroth 1995, Filmer 199 1 Africa Alroth 1995 Brazil Bucherl 1971 Brazil Biicherl 1971

Russell 1987, 1988 Brazil & West Indies Cooke et al. 1972 South America Biicherl 1 87 1, Alroth 1 995

Bettini & Brignoli 1978 Peters 1992

Brazil Peters 1992 Sri Lanka Schmidt 1989, Alroth 1995 Sri Lanka Portillo 1995 India Portillo 1995 Africa Peters 1992, Schmidt 1989 Africa Layzell 1989

Table 2.--Continued. Selenocosmia crassipes (Koch) Australia Southcott 1976

Peters 1992 Selenocosmia stirlingi Hogg Australia, New Guinea

Southcott 1 976 Peters 1992, Alroth 1995

Sericopelma rubronitens (Ausserer) Central America Gertsch 1949 Central & South America

Bettini & Brignoli 1978 Peters 1992

Theraphosa Brazil Bilcherl 1971 Xenesthis Brazil Biicherl 1 97 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

spiders widow 2,404 813

recluse 1,453 8 13 spiders

None Minor Moderate Major Death 305 1,140 272 9 0

other 560 113 spiders

tarantulas 55 11 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 pack- ed semi-firmly in slightly damp paper towels inside a deli cup or margar- ine 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 par- ticular interest is the husbandry and breeding of tarantulas and scorpions with emphasis on the needs of these animals when kept as pets. You may

3 1

subscribe to this list by sending a command (in the text part of the mes- sage) to: [email protected]. 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 199 1 (a non-profit organ-

ization, first published 1992), to promote the study and the dissemination of information primarily concerning the arachmd infiaorder Mygalomor- phae, 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, teach- ers, 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 reg- ularly scheduled for publication in any given year. The Forum Magazine is written in a style easily understood by the hobbyist and contains art- n icles, 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 16 17, Artesia, New Mexico 882 1 1 USA. Email: 7 [email protected] - 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 mis- understandings 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. Plat- nick, AMNH, Central Park West at 79th St., New York, NY 10024 USA.

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 3 JE 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 85 6 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 arthro-

pods. 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) 225- 1 802, email [email protected]

INVERTEBRATA Quarterly, covers many aspects of invertebrates, concentrating on suc-

cessful care in captivity. $25 yr. Mascariiio, PO Box 2072 1 ., Los Ange- les, 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 Adams, J. 1907. Observations on a

mygale spider (Psalmopoeus cam- bridgii Poc.) Trans. Edinburg F. Nat. Micr. Soc. 5: 402-406.

Adams, J. 1908. Further note on the mygale spider Trans. Edinb. F. Nat. Micr. Soc. 6: 81-82.

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. Lu- cas. 1986. Ultrastructure of sper- matozoa and spermatogenesis in bird spiders (Theraphosidae, Myga- lomorphae, Araneae). J. Submi- crosc. Cytol. 18: 739-753.

Anderson, J. F. & K. N. Prestwich. 1985. The physiology of exercise at and above maximal aerobic capacity in a theraphosid (tarantula) spider, Brachypelma smithi (F. 0 . Pickard- Cambridge). J. Comp. Physiol. 155: 529-539.

Andrews, J. S. 1930. The digestion of a mouse by a tarantula. Proc. Indiana Acad. Sci. 39: 305.

Angersbach, D. 1975. Oxygen pres- sures in haemolymph and various tissues of the tarantula, Eurypelma helluo. J. Comp. Physiol. 98: 133- 145.

Angersbach, D. 1975. Die sauerstof- fiersorgung der gewebe zweier spinnen, Eurypelma helluo und Cupiennius salei: lokale p. 2. Mes- sungen in der pH molyrnphe und in verschiedenen muskeln. Verh. Dtsch. Zool. Ges. 1974: 277-280.

Angersbach, D. 1978. Oxygen transport in the blood of the tarantula Eu- rypelma calijiornicum: o2 and pH during rest, activity and recovery. J. Comp. Physiol. 123(2): 113-125.

Anonymous. 1777. Tarantula. Onom. Med. Comp. 7: 442-444

Araque, A., W. Ferreira, S. Lucas & W. Bruno. 1992. Glutamyc post synaptic block by Pamphobeteus spider venom in crayfish. Brain Res. 571: 109-1 14.

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Aruchami, M. & G. Sundara Rajulu. 1978. A study on the lipid compo- nent of the haemolymph of a spider Poecilotheria fasciata. Curr. Sci. (Bangalore) 47: 3 92.

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Ausserer, A. 1871. Beitrage zu kenntnis der Arachnidien familie der Territelariae Thorell (Mygalidae autor). Verhandl. K. K. Zool. Bot. - Gesell. Wien 2 1: 177-224.

Ausserer, A. 1875. Zweiter Beitrag zur Kenntnis der Arachniden-Familie der Territelariae Thorell (Mygalidae autor). Verhandl. K. K. Zool. Bot. Gesell. Wien 25: 125-206.

Babu, K. S. 1969. Certain histological and anatomical features of the central nervous system of a large Indian spider, Poecilotheria. Am. 2001. 9: 113-1 19.

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Baerg, W. J. 1922. Regarding the habits of tarantulas and the effects of their poison. Sci. Month. 14: 482- 489.

Baerg, W. J. 1922. The effect of the poison of tarantulas. J. Parasit. 8: 86-89.

Baerg, W. J. 1926. Regeneration of appendages in the tarantula; Euvy- pelnra cali$ornica Ausserer. Ann. Entmol. Soc. Am. 19: 512-513.

Baerg, W. J. 1928. The life cycle and mating habits of the male tarantula. Quart. Rev. Biol., 3: 109-1 16.

Baerg, W. J. 1929. Cocoon-making by the tarantula. Ann. Entomol. Soc. Am. 22: 161-164.

Baerg, W. J. 1929. Some poisonous arthropods of North and Central America. Congr. Int. Entomol. 4, Ithaca Trans. 2: 418-438.

Baerg, W. J. 1937. Laboratory care in tarantulas. In, Culture methods for Invertebrate Animals, Gal tsoff [Ed], Ithaca, pp. 243-244.

Baerg, W. J. 1938. Tarantula studies. J. New York Entomol. Soc. 46: 3 1-43.

Baerg, W. J. 1945. The black widow and the tarantula. Trans. Conn. Acad. Arts Sci. 36: 99-1 13.

Baerg, W. J. 1958. The tarantula. Univ. Kansas Press, Lawrence 88 PP-

Baerg, W. J. & W. B. Peck. 1970. A note on the longevity and molt cycle of two tropical theraphosids. Bull. British Arachnol. Soc. 1: 107-108.

Baird, R. L. 1932. A study of the large banana spider. Ohio J. Sci. 32: 32 1-

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Selected ~ i b l i o g r a ~ h ~ of ~ o b b y ~ r i e n t e d Material (Non-~efereed Publicatione)

Alroth, G. 1995. How dangerous are Brigida, R. A. 1993. Two blonds in the tarantulas in reality? J. British Ta- hand. Am. Tarantula Soc. Forum 2: rantula Soc. 1 l(1): 20-25. 99-101.

Anonymous. 1987. Keeping exotics: Brigida, R. A. 1994. Blond goliath egg Theraphosid or bird-eating spiders. gulpers. Am. Tarantula Soc. Forum Exotic Entomol. Group Newsl. 1987 3: 8-10. (Autumn) : 1 -3. Brigida, R. A. 1994. Togo starburst

Anonymous. 1989. Cambridge taran- glider pilots. Am. Tarantula Soc. tula project 1988. Camb. Exped. J. Forum 3: 160. 1988-1989: 32-35. Brigida, R. A. 1995. Arachnopulse.

Barbeua, J. T. & M. J. Blasczyk. 1994. Am. Tarantula Soc. Forum 4: 123, Tarantula care in captivity. Lore 125. 44(3). Bustard, R. 1996. Social behavior in

Breene, R. G., 111. 1993. Poisonous Poecilotheria. Am. Tarantula Soc. beetle kills tarantulas. Am. Taran- Forum 5: 5-8. tula Soc. Forum 2: 15-16. Bustard, R. 1996. Poecilotheria care in

Breene, R. G., 111. 1993. Food for captivity. Am. Tarantula Soc. Forum thought and tarantulas. Am. Taran- 5: 44-48. tula Soc. Forum 2: 21-22. Charpentier, P. 1992. Heterscodra

Breene, R. G., 111. 1993. Nick and the maculata. J . British Tarantula Soc. spider wasp. Am. Tarantula Soc. Fo- 8(2): 11-19. rum 2: 23-25. Charpentier, P. 1992. Heterscodra

Breene, R. G., 111. 1994. Nomenclatu- maculata Pocock, 1899. Arachnides, ral nemesis: Systematic headaches Bull. G. E. A. 15: 25-30. and taxing taxonomy. Am. Taran- Charpentier, P. 1992. The genus Avi- tula Soc. Forum 3: 3 1-37. cularia. Exothermae l(1): 1-49.

Breene, R. G., 111. 1994. Legalized Charpentier, P. 1992. Pterinochilus Latrodectus: Dangerous animals murinus, morphology - biology- laws in the USA. J. British Taran- behaviour. Exothermae l(2): 1-72. tula Soc. 9(3): 6-9. Cloudsley-Thompson, J. L. 1982. Ta-

Breene, R. G., 111. 1994. Rodent ran- rantulas. Country Side 25: 26-29. sackers? Pesticide mockers? Do Cokendolpher, J. C. 1992. More taran- m Texas tans prefer humans? Am. Ta- tula bites. Am. Tarantula Soc. Fo- rantula Soc. Forum 3: 102-104. rum 1: 56-59.

Breene, R. G., 111. 1995. Tarantula ER. Dunlop, J. 1991. Venom research: the Am. Tarantula Soc. Forum 4: 10-13. story so far. J. British Tarantula Soc.

Breene, R. G., 111. 1995. Beware the 7(1): 18-19. ides of March, and tarantula experts. Dunlop, J. 199 1. An African with hol- Am. Tarantula Soc. Forum 4: 108- low legs. J. British Tarantula Soc. m 111. 7(2) 8-10.

Breene, R. G., 111. 1995. Endangered Dunlop, J. 1993. A functional expla- species. Am. Tarantula Soc. Forum nation for posturing during tarantula 4: 184-185. ingestion? Am. Tarantula Soc. Fo- ~1

rum 2: 95-97.

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. Taran- tula 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 Af- rica 128 pp.

Gallon, R. 199 1. 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: 113-1 14.

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. 199 1. Successful breeding of bird-eating spiders in captivity: Megaphobema robusta. J. British Tarantula Soc. 7(1): 12-13.

Johnson-Delaney, C. 199 1. Basic care of tarantulas. J. Small Exotic Anim. Med. l(1): 19.

Layzell, J. 1989. My first bite! J. Brit- ish Tarantula Soc. 5(1): 15-16.

Layzell, J. 199 1. 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 pink- toes? Am. Tarantula Soc. Forum 4: 40-4 1.

Marshall, S. D. 1996. Old dog learns new trick. Am. Tarantula Soc. Fo- rum 5: 114-1 16.

Mason, T. 1993. Tarantula scavengers. Am. Tarantula Soc. Forum 2: 74-75.

Miranda, L. 1994a. A bite in time heals mine. Am. Tarantula Soc. Fo- rum 3: 15-18.

Miranda, L. 1994. Killer tarantula po- lice patrol. Am. Tarantula Soc. Fo- rum 3: 92-93.

Moore, B. H. 1993. Pinktoe tarantulas. Am. Tarantula Soc. Forum 2: 39-40.

Moore, B. H. 1994. Red rumped canni- bals. 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, In- vertebrates in captivity conf., Sono- ran 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 Ama- zon adventure: death defLing diving tarantula emulates submarine. Am. Tarantula Soc. Forum 3 : 152-154.

Reger, B. H. 1995. Tarantula one-a- days. 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 bes- chreibung einer neuen art aus Para- guay: Tmesiphantes spinopalpus sp. n. (Araneida: Theraphosidae). Ar- thropoda 4: 24-3 1.

Schmidt, G. E. W. 1990. Zur kenntnis der gattung Mygafarachne (Aran- eida: 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 Para- physa art aus Equador (Araneida: Theraphosidae; Theraphosinae). Arachnol. Anz. 1019 1: 8-12.

Schmidt, G. E. W. 1991. Eine neue vogelspinnen aus Honduras Phor- mictopus atrichomatus sp. n. (Aran- eida: Theraphosidae: Theraphosi- nae). Arachnol. Anz., nr. 11: 7-10.

Schmidt, G. E. W. 1992. Das man- n nchen von Euathlus truculentus Ausserer 1875 (Araneida: Thera- phosidae: ~heraphosinae). Arachn. Anz. 3: 9-13. n

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 n 1890 oder Euathlus Ausserer 1875? (Araneida: Theraphosidae; Thera- phosidae). 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: Thera- phosinae). Arachnol. Mag. 2(7): 7- 15.

Schmidt, G. E. W. 1994. Avicularia urticans sp. n. (Araneida: Thera- phosidae: Aviculariinae), eine vog- elspinnenart 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 (Aran- eida: Theraphosidae: Theraphosi- nae), 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: 98- 101.

Schmidt, G. E. W. & R. H. Krause. 1995. Eine neue art der Theraphosi- dae aus Vietnam: Selenopelma kov- ariki gen. et sp. n. (Araneida: Theraphosidae: Selenocosmiinae). Arthropoda 3 : 2 1-24.

Schmidt, G. E. W. 1995. Chro- matopelma gen. n.; eine neue gattung der Theraphosidae (Arachnida: Araneida: Theraphosidae: Thera- phosinae). 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 An- tilles (Arach-nida; Araneida: Thera- phosidae). Arachnol. Mag. 4(6): 1- 9.

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: 181- 182.

Smith, A. M. 1988. Lyrognathus ro- bustus, a new species of theraphosid spider from Malaysia. J. British Ta- rantula Soc. 4(2): 15-19.

Smith, A. M. 1990. Taxonomic differ- ences between the closely related species Grammostola spatulata Cambridge 1897 and Grammostola cala Chamberlin 1917 from Chile. J. British Tarantula Soc. 6(1): 19- 21.

Smith, A. M. 1991. A revision of the genus Megaphobema Pocock, 1 90 1 (Araneida; Theraphosidae). J. Brit- ish Tarantula Soc. 6(1): 14-19.

Smith, A. M. 1992. In defence of Ra- ven's decision to make the genus

Brachypelma Simon 1891 a junior synonym of Euathlus Ausserer 1895. J. British Tarantula Soc. 7(3): 14- 19.

Smith, A. M. 1992. Redescription of the Aviculariinae species Avicularia minatrix Pocock 1903. J. British Ta- rantula Soc. 8(2): 22-26.

Smith, A. M. 1993. A new mygalomorph spider from Mexico (Brachy- pelma, 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): 13- 18.

Smith, A.M. 1993. How to keep tarantulas. Fitzgerald Publishing, Lon- don, 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 infec- tions. 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 any- more. 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. 199 1. 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: Orni- thoctoninae). Arachnol. Anz. 12: 5- 8.

von Wirth, V. 1991. Eine neue Vogel- spinnenart aus Vietnam. Haplo- pelma 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 1983- 1984: 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. Fo- rum 2: 108-1 10.

Current popular Books on Tarantulas Baxter, R. N. 1993. Keeping and

breeding tarantulas. Chudleigh Pub- lishing, 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, Calif- ornia. 80 pp.

Hancock, K. & J. Hancock. 1992. Tarantulas: Keeping and breeding arachnids in captivity. R & A Publishing Limited, Somerset 147 PP-

Marshall, S. D. 1996. Tarantulas and other arachnids. Barron's Haup- pauge, New York 104 pp.

Reger, B. 1995. Tarantulas as a new pet. T.F.H. Publications, Inc. Nep- tune City, New Jersey. 64 pp.

Schmidt, G. E. W. 1993. Vogelspin- nen: vorkommen, lebensweise, hal- tung und zucht, mit bestimmung- sschliissein 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 Classifi- cation 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.

p their medical importance, ud world-wide bibliography to m the

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