Myrmecia (ant)

Myrmecia is a genus of ants first established by Danish zoologist Johan Christian Fabricius in 1804. The genus is a member of the subfamily Myrmeciinae of the family Formicidae. Myrmecia is a large genus of ants, comprising at least 93 species that are found throughout Australia and its coastal islands, while a single species is only known from New Caledonia. One species has been introduced out of its natural distribution and was found in New Zealand in 1940, but the ant was last seen in 1981. These ants are commonly known as bull ants, bulldog ants or jack jumper ants, and are also associated with many other common names. They are characterized by their extreme aggressiveness, ferocity, and painful stings. Some species are known for the jumping behavior they exhibit when agitated.

Species of this genus are also characterized by their elongated mandibles and large compound eyes that provide excellent vision. They vary in colour and size, ranging from 8 to 40 millimetres (0.31 to 1.57 in). Although workers and queens are hard to distinguish from each other due to their similar appearance, males are identifiable by their perceptibly smaller mandibles. Almost all Myrmecia species are monomorphic, with little variation among workers of a given species. Some queens are ergatoid and have no wings, while others have either stubby or completely developed wings. Nests are mostly found in soil, but they can be found in rotten wood and under rocks. One species does not nest in the ground at all; its colonies can only be found in trees.

A queen will mate with one or more males, and during colony foundation she will hunt for food until the brood have fully developed. The life cycle of the ant from egg to adult takes several months. Myrmecia workers exhibit greater longevity in comparison to other ants, and workers are also able to reproduce with male ants. Myrmecia is one of the most primitive group of ants on earth, exhibiting differentiated behaviors from other ants. Workers are solitary hunters and do not lead other workers to food. Adults are omnivores that feed on sweet substances, but the larvae are carnivores that feed on captured prey. Very few predators eat these ants due to their sting, but their larvae are often consumed by blindsnakes and echidnas, and a number of parasites infect both adults and brood. Some species are also effective pollinators.

Myrmecia stings are very potent, and the venom from these ants is among the most toxic in the insect world. In Tasmania, 3% of the human population are allergic to the venom of M. pilosula and can suffer life-threatening anaphylactic reactions if stung. People prone to severe allergic reactions can be treated with allergen immunotherapy (desensitisation).

## Etymology and common names

The generic name Myrmecia derives from Greek word Myrmec- (+ -ia), meaning "ant". In Western Australia, the Indigenous Australians called these ants kallili or killal.

Ants of this genus are popularly known as bulldog ants, bull ants, or jack jumper ants due to their ferocity and the way they hang off their victims using their mandibles, and also due to the jumping behaviour displayed by some species. Other common names include "inch ants", "sergeant ants", and "soldier ants". The jack jumper ant and other members of the Myrmecia pilosula species group are commonly known as "black jumpers", "hopper ants", "jumper ants", "jumping ants", "jumping jacks", and "skipper ants".

## Taxonomy and evolution

Genetic evidence suggests that Myrmecia diverged from related groups about 100 million years ago (Mya). The subfamily Myrmeciinae, to which Myrmecia belongs, is believed to have been found in the fossil record of 110 Mya ago. However, one study suggests that the age of the most recent common ancestor for Myrmecia and Nothomyrmecia is 74 Mya, and the subfamily is possibly younger than previously thought. Ants of the extinct genus Archimyrmex may possibly be the ancestor of Myrmecia. In the Evans' vespoid scala, Myrmecia and other primitive ant genera such as Amblyopone and Nothomyrmecia exhibit behavior which is similar to a clade of soil-dwelling families of vespoid wasps. Four species groups form a paraphyletic assemblage while five species groups form a monophyletic assemblage. The following cladogram shows the phylogenetic relationships within Myrmecia:

### Classification

Myrmecia was first established by Danish zoologist Johan Christian Fabricius in his 1804 publication Systema Piezatorum, in which seven species from the genus Formica were placed into the genus along with the description of four new species. Myrmecia has been classified into numerous families and subfamilies; in 1858, British entomologist Frederick Smith placed it in the family Poneridae, subfamily Myrmicidae. It was placed in the subfamily Ponerinae by Austrian entomologist Gustav Mayr in 1862. This classification was short-lived as Mayr reclassified the genus into the subfamily Myrmicinae three years later. In 1877, Italian entomologist Carlo Emery classified the genus into the newly established subfamily Myrmeciidae, family Myrmicidae. Smith, who had originally established the Myrmicidae as a family in 1851, reclassified them as a subfamily in 1858. He again treated them as a family in 1871. Swiss myrmecologist Auguste Forel initially treated the Poneridae as a subfamily and classified Myrmecia as one of its constituent genera but later placed it in the Ponerinae. William H. Ashemad placed the genus in the subfamily Myrmeciinae in 1905, but it was later placed back in the Ponerinae in 1910 by American entomologist William Morton Wheeler. In 1954, Myrmecia was placed into the Myrmeciinae; this was the last time the genus was placed into a different ant subfamily.

In 1911, Emery classified the subgenera Myrmecia, Pristomyrmecia, and Promyrmecia, based on the shape of their mandibles. Wheeler established the subgenus Halmamyrmecia, and the ants placed in it were characterized by their jumping behavior. The taxon Wheeler described was not referred to in his later publications, and the genera Halmamyrmecia and Pristomyrmecia were synonymised by John Clark. At the same time, Clark reclassified the subgenus Promyrmecia as a full genus. He revised the whole subfamily Myrmeciinae in 1951, recognizing 118 species and subspecies in Myrmecia and Promyrmecia; five species groups were assigned to Myrmecia and eight species groups to Promyrmecia. This revision was rejected by entomologist William Brown due to the lack of morphological evidence that would make the two genera distinct from each other. Due to this, Brown classified Promyrmecia as a synonym of Myrmecia in 1953. Clark's revision was the last major taxonomic study on the genus before 1991, and only a single species was described in the intervening years. In 2015, four new Myrmecia ants were described by Robert Taylor, all exclusive to Australia. Currently, 94 species are described in the genus, but as many as 130 species may exist.

Under the present classification, Myrmecia is the only extant genus in the tribe Myrmeciini, subfamily Myrmeciinae. It is a member of the family Formicidae in the order Hymenoptera. The type species for the genus is M. gulosa, discovered by Joseph Banks in 1770 during his expedition with James Cook on HMS Endeavour. M. gulosa is among the earliest Australian insects to be described, and the specimen Banks collected is housed in the Joseph Banks Collection in the Natural History Museum, London. M. gulosa was described by Fabricius in 1775 under the name Formica gulosa and later designated as the type species of Myrmecia in 1840.

### Genetics

The number of chromosomes per individual varies from one to over 70 among the species in the genus. The genome of M. pilosula is contained on a single pair of chromosomes (males have just one chromosome, as they are haploid). This is the lowest number possible for any animal, and workers of this species are homologous. Like M. pilosula, M. croslandi also contains a single chromosome. While these ants only have a single chromosome, M. pyriformis contains 41 chromosomes, while M. brevinoda contains 42. The chromosome count for M. piliventris and M. fulvipes is two and 12, respectively. The genus Myrmecia retains many traits that are considered basal for all ants (i.e. workers foraging alone and relying on visual cues).

### Species groups

Myrmecia contains a total of nine species groups. Originally, seven species groups were established in 1911, but this was raised to 13 in 1951; Promyrmecia had a total of eight, while Myrmecia only had five. M. maxima does not appear to be in a species group, as no type specimen is available.

## Description

Myrmecia ants are easily noticeable due to their large mandibles, large compound eyes that provide excellent vision and a powerful sting that they use to kill prey. Each of their eyes contains 3,000 facets, making them the second largest in the ant world. Size varies widely, ranging from 8 to 40 mm (0.31 to 1.57 in) in length. The largest Myrmecia species is M. brevinoda, with workers measuring 37 mm (1.5 in); M. brevinoda workers are also the largest in the world. Almost all species are monomorphic, but M. brevinoda is the only known species where polymorphism exists. It is well known that two worker subcastes exist, but this does not distinguish them as two different polymorphic forms. This may be due to the lack of food during winter and they could be incipient colonies. The division of labour is based on the size of ant, rather than its age, with the larger workers foraging for food or keeping guard outside the nest, while the smaller workers tend to the brood.

Their coloration is variable; black combined with red and yellow is a common pattern, and many species have golden-colored pubescence (hair). Many other species are brightly colored which warns predators to avoid them. The formicine ant Camponotus bendigensis is similar in appearance to M. fulvipes, and data suggest C. bengdigensis is a batesian mimic of M. fulvipes. The number of malpighian tubules differs between castes; in M. dispar, males have 16 tubules, queens range from 23 to 26, and workers have 21 to 29.

Worker ants are usually the same size as each other, although this is not true for some species; worker ants of M. brevinoda, for example, vary in length from 13 to 37 mm (0.51 to 1.46 in). The mandibles of the workers are long with a number of teeth, and the clypeus is short. The antennae consist of 12 segments and the eyes are large and convex. Based on a study on the antennal sensory of M. pyriformis, the antennal sensilla are known to have eight types. Large ocelli are always present.

Queens are usually larger than the workers, but are similar in colour and body shape. The head, node, and postpetiole are broader in the queen, and the mandibles are shorter and also broad. Myrmecia queens are unique in that particular species either have fully winged queens, queens with poorly developed wings, or queens without any wings. For example, M. aberrans and M. esuriens queens are ergatoid, meaning that they are wingless. Completely excavated nests showed no evidence of any winged queen residing within them. Some species have queens which are subapterous, meaning they are either wingless or only have rudiments of wings; the queens can be well developed with or without these wing buds. M. nigrocincta and M. tarsata are "brachypterous", where queens have small and rudimentary wings which render the queen flightless. Dealated queens with developed wings and thoraces are considered rare. In some species, such as M. brevinoda and M. pilosula, three forms of queens exist, with the dealated queens being the most recognisable.

Males are easy to identify due to their perceptibly broad and smaller mandibles. Their antennae consist of 13 segments, and are almost the same length as the ants' bodies. Ergatandromorph (an ant that exhibits both male and worker characteristics) males are known; in 1985, a male M. gulosa was collected before it hatched from its cocoon, and it had a long but excessively curved left mandible while the other mandible was small. On the right side of its body, it was structurally male, but the left side appeared female. The head was also longer on the female side, its colour was darker, and the legs and prothorax were smaller on the male side. Male genitalia are retracted into a genital cavity that is located in the posterior end of the gaster. The sperm is structurally the same to other animal sperm, forming an oval head with a long tail.

Among the largest larvae examined were those of M. simillima, reaching lengths of 35 mm (1.4 in). The pupae are enclosed in dark cocoons.

## Distribution and habitat

Almost all species in the genus Myrmecia are found in Australia and its coastal islands. M. apicalis is the only species not native to Australia and is only found in the Isle of Pines, New Caledonia. Only one ant has ever established nests outside its native range; M. brevinoda was first discovered in New Zealand in 1940 and the ant was recorded in Devonport in Auckland in 1948, 1965 and 1981 where a single nest was destroyed. Sources suggest the ant was introduced to New Zealand through human activity; they were found inside a wooden crate brought from Australia. While no eradication attempt was made by the New Zealand government, the ant has not been found in the country since 1981 and is presumed to have been eradicated.

Ants of this genus prefer to inhabit grasslands, forests, heath, urban areas and woodland. Nests are found in Callitris forest, dry marri forest, Eucalyptus woodland and forests, mallee scrub, in paddocks, riparian woodland, and wet and dry sclerophyll forests. They also live in dry sandplains, and coastal plain. When a queen establishes a new colony, the nest is at first quite simple structurally. The nest gradually expands as the colony grows larger. Nests can be found in debris, decaying tree stumps, rotten logs, rocks, sand, and soil, and under stones. While most species nest underground, M. mjobergi is an arboreal nesting species found on epiphytic ferns of the genus Platycerium. Two types of nests have been described for this genus: a simple nest with a noticeable shaft inside, and a complex structure surrounded by a mound. Some species construct dome-shaped mounds containing a single entrance, but some nests have numerous holes that are constantly used and can extend several metres underground. Sometimes, these mounds can be 0.5 m (20 in) high. Workers decorate these nests with a variety of items, including charcoal, leaves, plant fragments, pebbles, and twigs. Some ants use the warmth by decorating their nests with dry materials that heat quickly, providing the nest with solar energy traps.

## Behaviour and ecology

### Foraging

The genus Myrmecia is among the most primitive of all known living ants, and ants of the genus are considered specialist predators. Unlike most ants, workers are solitary hunters, and do not lay pheromone trails; nor do they recruit others to food. Tandem running does not occur, and workers carrying other workers as a method of transportation is rare or awkwardly executed. Although Myrmecia is not known to lay pheromone trails to food, M. gulosa is capable of inducing territorial alarm using pheromones while M. pilosula can attack en masse, suggesting these ants can also induce alarm pheromones. M. gulosa induces territorial alarm behaviour using pheromones from three sources; an alerting substance from the rectal sac, a pheromone found in the Dufour's gland, and an attack pheromone from the mandibular gland. Despite Myrmecia ants being among the most primitive ants, they exhibit some behaviours considered "advanced"; adults will sometimes groom each other and the brood, and distinct nest odors exist for each colony.

Most species are diurnal, and forage on the ground or onto low vegetation in search of food, but a few are nocturnal and only forage at night. Most Myrmecia ants are active during the warmer months, and are dormant during winter. However, M. pyriformis is a nocturnal species that is active throughout the whole year. M. pyriformis also has a unique foraging schedule; 65% of individuals who went out to forage left the nest in 40–60 minutes, while 60% of workers would return to the nest in the same duration of time at dusk. Foraging workers rely on landmarks for navigation back home. If displaced a short distance, they will scan their surroundings, and then rapidly move in the direction of the nest. M. vindex ants carry dead nest-mates out of their nests and place them on refuse piles, a behaviour known as necrophoresis.

### Pollination

While pollination by ants is somewhat rare, several Myrmecia species have been observed pollinating flowers. For example, the orchid Leporella fimbriata is a myrmecophyte which can only be pollinated by the winged male ant M. urens. Pollination of this orchid usually occurs between April and June during warm afternoons, and may take several days until the short-lived males all die. The flower mimics M. urens queens, so the males move from flower to flower in an attempt to copulate with it. M. nigrocincta workers have been recorded visiting flowers of Eucalyptus regnans and Senna acclinis, and are considered a potential pollination vector for E. regnans trees. Although Senna acclinis is self-compatible, the inability of M. nigrocincta to appropriately release pollen would restrict its capacity to effect pollination. Foraging M. pilosula workers are regularly observed on the inflorescences of Prasophyllum alpinum (mostly pollinated by wasps of the family Ichneumonidae). Although pollinia are often seen in the ants' jaw, they have a habit of cleaning their mandibles on the leaves and stems of nectar-rich plants before moving on, preventing pollen exchange. Whether M. pilosula contributes to pollination is unknown.

### Diet

Despite their ferocity, adults are nectarivores, consuming honeydew (a sweet, sticky liquid found on leaves, deposited from various insects), nectar, and other sweet substances. The larvae, however, are carnivorous. After they reach a certain size, they are fed insects that foragers capture and kill. The workers also regurgitate food for other ants to consume. Young ants are rarely fed food regurgitated by adults. Adult workers prey on a variety of insects and arthropods, such as beetles, caterpillars, earwigs, Ithone fusca moths, Perga saw flies, and spiders. Other prey include invertebrates such as bees, cockroaches, crickets, wasps and other ants; in particular, workers prey on Orthocrema ants (a subgenus of Crematogaster) and Camponotus, although this is risky since these ants are able to call for help through chemical signals. Slaters, earthworms, scale insects, frogs, lizards, grass seeds, possum feces and kangaroo feces are also collected as food. Flies such as the housefly and blowfly are consumed. Some species, such as M. pilosula, will only attack small fly species and ignore larger ones. Nests of the social spider Delena cancerides are often invaded by M. pyriformis ants, and nests once housing these spiders are filled with debris such as twigs and leaves by the workers, rendering them useless. These "scorched earth" tactics prevent the spiders competing with the ants. M. gulosa attacks Christmas beetles, but workers later bury them.

Myrmecia is one of the very few genera where the workers lay trophic eggs, or infertile eggs laid as food for viable offspring. Workers laying trophic eggs have only been reported in two species; these species are M. forceps and M. gulosa. Depending on the species, colonies specialise in trophallaxis; queens and larvae eat eggs that are laid by worker individuals, but the workers do not feed on eggs. Neither adults nor larvae consume food during winter, but cannibalism among larvae is known to occur throughout the year. The larvae only cannibalise each other; this is most likely to happen when no dead insects are available.

### Predators, parasites and associations

Myrmecia ants deter many potential predators due to their sting. The blindsnake Ramphotyphlops nigrescens consumes the larvae and pupae of Myrmecia, while avoiding the potent sting of the adults, which it is vulnerable to. The short-beaked echidna (Tachyglossus aculeatus) also eats the eggs and larvae. Nymphs of the assassin bug species Ptilocnemus lemur lure these ants to themselves by trying to make the ant sting them, by waving its hind legs around to attract a potential prey item. Body remains of Myrmecia have been found in the stomach contents of the eastern yellow robin (Eopsaltria australis). The Australian magpie (Gymnorhina tibicen), the black currawong (Strepera versicolor), and the white-winged chough (Corcorax melanorhamphos) prey on these ants, but few are successfully taken.

The host association between Myrmecia and eucharitid wasps began several million years ago; M. forficata larvae are the host to Austeucharis myrmeciae, being the first recorded eucharitid parasitoid of an ant, and Austeucharis fasciiventris is a parasitoid to M. gulosa pupae. M. pilosula is affected by a gregarines parasite that changes an ant's colour from their typical black appearance to brown. This was discovered when brown workers were dissected and found to have gregarinasina spores, while black workers showed no spores. Another unidentified gregarine parasite is known to infect the larvae of M. pilosula and other Myrmecia species. This gregarine parasite also softens the ant's cuticle. Other parasites include Beauveria bassiana, Paecilomyces lilacinus, Chalcura affinis, Tricoryna wasps, and various mermithid nematodes.

M. hirsuta and M. inquilina are the only known species in this genus that are inquilines and live in other Myrmecia colonies. An M. inquilina queen has been found in an M. vindex colony. Myrmecia is a larval attendant to the butterfly Theclinesthes serpentata (saltbush blue), while some species, particularly M. nigrocincta, enslave other ant species, notably those in the genus Leptomyrmex. M. nigriceps ants are able to enter another colony of the same species without being attacked, as they may be unable to recognize alien conspecifics, nor do they try to distinguish nestmates from ants of another colony. Formicoxenus provancheri and M. brevinoda share a form of symbiotic relationship known as xenobiosis, where one species of ant will live with another and raise their young separately, with M. brevinoda being the host. Solenopsis may sometimes nest in Myrmecia colonies, as a single colony was found to have three or four Solenopsis nests inside. Lagria beetles and rove beetles in the genus Heterothops dwell inside colonies and skinks and frogs have also been found living unmolested within Myrmecia nests. Metacrinia nichollsi, for example, has been reported living inside M. regularis colonies.

### Life cycle

Like other ants, Myrmecia ants begin as an egg. If the egg is fertilised, the ant becomes a diploid female; if not, it becomes a haploid male. They develop through complete metamorphosis, meaning that they pass through larval and pupal stages before emerging as adults.

During the process of founding a colony, as many as four queens cooperate with each other to find a suitable nesting ground, but after the first generation of workers is born, they fight each other until one queen is left alive. However, occasional colonies are known to have as many as six queens coexisting peacefully in the presence of workers. A queen searches for a suitable nest site to establish her colony, and excavates a small chamber in the soil or under logs and rocks, where she takes care of her young. A queen also hunts for prey instead of staying in her nest, a behaviour known as claustral colony founding. Although queens do provide sufficient amounts of food to feed their larvae, the first workers are "nanitics" (or minims), smaller than the smallest workers encountered in older developed colonies. Several species do not have any worker caste, and solitary queens will raid a colony, kill the residing queen, and take over the colony. The first generation of workers may take a while to fully develop into adults; for example, M. forficata eggs take around 100 days to fully develop, while other species may take up to eight months.

Queens lay around eight eggs, but less than half of these eggs develop. Some species, such as M. simillima and M. gulosa, lay their eggs singly on the colony floor, while M. pilosula ants may lay eggs in a clump. These clumps have two to 30 eggs each with no larvae present. Certain Myrmecia species do not lay their eggs singly and form clumps of eggs, instead. The larvae are capable of crawling short distances without the assistance of adult workers, and workers will cover the larvae in dirt to help them spin into a cocoon. If cocoons are isolated from a colony, they are capable of shedding their skins before hatching, allowing themselves to advance to full pigmentation. Sometimes, a newborn can emerge from its pupa without the assistance of other ants. Once these ants are born, they are able to identify distinct tasks, a well known primitive trait. Myrmecia lifespans vary in each species, but their longevity is greater than many ant genera: M. nigrocincta and M. pilosula have a lifespan of one year, while M. nigriceps workers can live up to 2.2 years. The oldest recorded worker was a M. vindex, living up to 2.6 years. If a colony is deprived of workers, queens are able to revert to colony-founding behaviours until a sustainable workforce emerges. A colony may also emigrate to a new nesting spot altogether.

### Reproduction

Winged, virgin queens and males, known as alates, appear in colonies during January, before their nuptial flight. Twenty females or fewer are found in a single colony, while males are much more common. The nuptial flight begins at different times for each species; they have been recorded in mid-summer to autumn (January to early April), but there is one case of a nuptial flight occurring from May to July. Ideal conditions for nuptial flight are hot stormy days with windspeeds of 30 km/h (18 mi/h) and temperatures reaching 30 °C (86 °F), and elevations of 91 metres (300 ft). Nuptial flights are rarely recorded due to queens leaving their nest singly, although as many as four queens may leave the nest at the same time. Species are both polygynous and polyandrous, with queens mating with one to ten males. Polygynous and polyandrous societies can occur in a single nest, but particular species are either primarily polygynous or primarily polyandrous. For example, nearly 80% of tested M. pilosula colonies are polygynous while M. pyriformis colonies are mostly polyandrous. Nuptial flight takes place during the morning and can last until late afternoon. When the alates leave the nest, most species launch themselves into the air from trees and shrubs, although others launch themselves off the ground. Queens discharge a glandular secretion from the tergal gland, which males are strongly attracted to. As many as 1,000 alates will gather to mate. A queen was once found to have five or six males attempting to copulate with her. The queen is unable to bear the weight of the large number of males trying to mate with her, and will drop to the ground, with the ants dispersing later on. M. pulchra queens are ergatoid and cannot fly; the males meet the queen out in an open area away from the nest and mate, and these queens do not return to their nest after mating.

Both independent and dependent colony foundation can occur after mating. Isolation by distance (IBD) patterns have been recorded with M. pilosula queens, where nests that tend to be closer together were more genetically related to each other in comparison to other nests further away. Independent colony foundation is closely associated with queens which engage in nuptial flight in areas far from their home colony, showing that dependent colony foundation mostly occurs if they mate near their nest. In some cases, queens could seek adoption into alien colonies if there are no suitable areas to find a nest or independent colony foundation cannot be carried out. Other queens could try to return to their home nest after nuptial flight, but they may end up in another nest near the nest they originally came from. In multiple-queen societies, the egg-laying queens are generally unrelated to one another, but one study showed that it is possible for multiple queens in the same colony to be genetically related to each other. Depending on the species, the number of individuals present in a colony can range from 50 to over 2,200 individuals. A colony with less than 100 workers is not considered a mature colony. M. dispar colonies have around 15 to 329 ants, M. nigrocincta have over 1,000, M. pyriformis have from 200 to over 1,400 and M. gulosa have nearly 1,600. A colony can last for a number of years. Foraging behaviour among smaller workers which never usually leave the nest can be a sign of a colony's impending demise.

Workers are known to produce their own eggs, but these eggs are unfertilised and hatch into male ants. There is a chance of workers attacking a particular individual who has successfully produced male offspring due to a change in a workers cuticular hydrocarbon; cuticular hydrocarbons are believed to play a vital role in the regulation of reproduction. However, this is not always the case. Myrmecia is one of several ant genera which possess gamergate workers, where a female worker is able to reproduce with mature males when the colony is lacking a queen. Myrmecia workers are highly fertile and can successfully mate with males. A colony of M. pyriformis without a queen was collected in 1998 and kept in captivity, during which time the gamergates produced viable workers for three years. Ovarian dissections showed that three workers of this colony mated with males and produced female workers. Queens have bigger ovaries than the workers, with 44 ovarioles while workers have 8 to 14. Spermatheca is present in M. gulosa workers, based on eight dissected individuals showing a spermatheca structurally similar to those found in queens. These spermathecas did not have any sperm. Why the queen was not replaced is still unknown.

### Vision

While most ants have poor eyesight, Myrmecia ants have excellent vision. This trait is important to them, since Myrmecia primarily relies on visual cues for navigation. These ants are capable of discriminating the distance and size of objects moving nearly a metre away. Winged alates are only active during the day, as they can see better. Members of a colony have different eye structures due to each individual fulfilling different tasks, and nocturnal species have larger ommatidia in comparison to those that are active during the day. Facet lenses also vary in size; for example, the diurnal species M. croslandi has a smaller lens in comparison to M. nigriceps and M. pyriformis which have larger lenses. Myrmecia ants have three photoreceptors that can see UV light, meaning they are capable of seeing colours that humans cannot. Their vision is said to be better than some mammals, such as cats, dogs or wallabies. Despite their excellent vision, worker ants of this genus find it difficult to find their nests at night, due to the difficulty of finding the landmarks they use to navigate. They are thus more likely to return to their nests the following morning, walking slowly with long pauses.

### Sting

Myrmecia workers and queens possess a sting described as "sharp in pain with no burning." The pain may last for several minutes. In the Starr sting pain scale, a scale which compares the overall pain of hymenopteran stings on a four-point scale, Myrmecia stings were ranked from 2–3 in pain, described as "painful" or "sharply and seriously painful". Unlike in honeybees, the sting lacks barbs, and so the stinger is not left in the area the ant has stung, allowing the ants to sting repeatedly without any harm to themselves. The retractable sting is located in their abdomen, attached to a single venom gland connected by the venom sac, which is where the venom is accumulated. Exocrine glands are known in some species, which produce the venom compounds later used to inject into their victims. Examined workers of larger species have long and very potent stingers, with some stings measuring 6 millimetres (0.24 in).

## Interaction with humans

Myrmecia is one of the best-known genera of ants. Myrmecia ants usually display defensive behaviour only around their nests, and are more timid while foraging. However, most species are extremely aggressive towards intruders; a few, such as M. tarsata, are timid, and the workers retreat into their nest instead of pursuing the intruder. If a nest is disturbed, a large force of workers rapidly swarms out of their nest to attack and kill the intruder. Some species, particularly those of the M. nigrocincta and M. pilosula species groups, are capable of jumping several inches when they are agitated after their nest has been disturbed; jumper ants propel their jumps by a sudden extension of their middle and hind legs. M. pyriformis is considered the most dangerous ant in the world by the Guinness World Records. M. inquilina is the only species of this genus that is considered vulnerable by the IUCN, although the conservation status needs updating.

Fatalities associated with Myrmecia stings are well known, and have been attested to by multiple sources. In 1931 two adults and an infant girl from New South Wales died from ant stings, possibly from M. pilosula or M. pyriformis. Another fatality was reported in 1963 in Tasmania. Between 1980 and 2000, there were six recorded deaths, five in Tasmania and one in New South Wales. Four of these deaths were due to M. pilosula, while the remaining two died from a M. pyriformis sting. Half of the victims had known ant-sting allergies, but only one of the victims was carrying adrenaline before being stung. Most victims died within 20 minutes of being stung, but one of the victims died in just five minutes from a M. pyriformis sting. No death has been officially recorded since 2003, but M. pilosula may have been responsible for the death of a man from Bunbury in 2011. Prior to the establishment of a desensitisation program, Myrmecia stings caused one fatality every four years.

### Venom

Each Myrmecia species has different venom components, so people who are allergic to ants are advised to stay away from Myrmecia, especially from species they have never encountered before. Based on five species, the median lethal dose (LD<sub>50</sub>) is 0.18–0.35 mg/kg, making it among the most toxic venoms in the insect world. The toxicity of the venom may have evolved due to the intense predation by animals and birds during the day, since Myrmecia is primarily diurnal. In Tasmania, 2–3% of the human population is allergic to M. pilosula venom. In comparison, only 1.6% people are allergic to the venom of the western honeybee (Apis mellifera), and 0.6% to the venom of the European wasp (Vespula germanica). In a 2011 Australian ant-venom allergy study, the objective of which was to determine what native Australian ants were associated with ant sting anaphylaxis, 265 of the 376 participants in the study reacted to the sting of several Myrmecia species. Of these, the majority of patients (176) reacted to M. pilosula venom and to those of several other species. In Perth, M. gratiosa was responsible for most cases of anaphylaxis due to ant stings, while M nigriscapa and M. ludlowi were responsible for two cases. The green-head ant (Rhytidoponera metallica) was the only ant other than Myrmecia species to cause anaphylaxis in patients. Dogs are also at risk of death from Myrmecia ants; renal failure has been recorded in dogs experiencing mass envenomation, and one dog was euthanised due to its deteriorating health despite treatment. Sensitivity is persistent for many years. Pilosulin 3 has been identified as a major allergen in M. pilosula venom, while pilosulin 1 and pilosulin 4 are minor allergens.

### Sting treatment

The nature of treatment for a Myrmecia sting depends on the severity of stingose, and the use of antihistamine tablets are other methods to reduce the pain. Indigenous Australians use bush remedies to treat Myrmecia stings, such as rubbing the tips of bracken ferns onto the stung area. Carpobrotus glaucescens is also used to treat stung areas, using juices that are squeezed and rubbed onto the area, which quickly relieves the pain from the sting.

Emergency treatment is only needed if a person is showing signs of a severe allergic reaction. Prior to calling for help, stung persons should be laid down, and their legs elevated. An EpiPen or an Anapen is given to people at risk of anaphylaxis, to use in case they are stung. If someone experiences anaphylactic shock, adrenaline and intravenous infusions are required, and those who suffer cardiac arrest require resuscitation. Desensitisation (also called allergy immunotherapy) is offered to those who are susceptible to M. pilosula stings, and the program has shown effectiveness in preventing anaphylaxis. However, the standardisation of M. pilosula venom is not validated, and the program is poorly funded. The Royal Hobart Hospital and the Royal Adelaide Hospital are the only known hospitals to run desensitisation programs. During immunotherapy, patients are given an injection of venom under the skin. The first dose is small, but dosage gradually increases. This sort of immunotherapy is designed to change how the immune system reacts to increased doses of venom entering the body.

Before venom immunotherapy, whole-body extract immunotherapy was widely used due to its apparent effectiveness, and it was the only immunotherapy used for ant stings. However, fatal failures were reported, and this led to scientists to research for alternative methods of desensitisation. Before 1986 allergic reactions were not recorded and there was no study on Myrmecia sting venom; whole body extracts were later used on patients during the 1990s, but this was found to be ineffective and was subsequently withdrawn. In 2003, ant venom immunotherapy was shown to be safe and effective against Myrmecia venom.

### Prevention

Myrmecia ants are frequently encountered by humans, and avoiding them is difficult. Wearing closed footwear such as boots and shoes can reduce the risk of getting stung; these ants are capable of stinging through fabric, however. A risk of being stung while gardening also exists; most stings occur when someone is gardening and is unaware of the ants' presence. Eliminating nearby nests or moving to areas with low Myrmecia populations significantly decreases the chances of getting stung.

Due to their large mandibles, Myrmecia ants have been used as surgical sutures to close wounds. The ant is featured on a postage stamp and on an uncirculated coin which are part of the Things That Sting issue by Australia Post, and M. gulosa is the emblem for the Australian Entomological Society. Myrmecia famously appears in the philosopher Arthur Schopenhauer's major work, The World as Will and Representation, as a paradigmatic example of strife and constant destruction endemic to the "will to live".

> But the bulldog-ant of Australia affords us the most extraordinary example of this kind; for if it is cut in two, a battle begins between the head and the tail. The head seizes the tail in its teeth, and the tail defends itself bravely by stinging the head: the battle may last for half an hour, until they die or are dragged away by other ants. This contest takes place every time the experiment is tried."

Notable Australian poet Diane Fahey wrote a poem about Myrmecia, which is based on Schopenhauer's description, and a music piece written by German composer Karola Obermüller was named after the ant.

## See also

- Ant
- Bullet ant
- Fire ant
- List of ants of Australia

Genus of ants

Extant Early Cretaceous first appearances


Natasha Falle

Natasha Falle (born 1973) is a Canadian professor at Humber College in Toronto, Ontario, Canada, who was forcibly prostituted from the ages of 15 to 27 and now opposes prostitution in Canada. Falle grew up in a middle-class home and, when her parents divorced, her new single-parent home became unsafe, and Falle ran away from home. At the age of 15, Falle became involved in the sex industry in Calgary, Alberta.

Falle's pimp kept her falsely imprisoned and trafficked her across the country. He married her and tortured her, breaking several of her bones and burning her body. In order to cope with the trauma of prostitution and violence, Falle became dependent on cocaine and almost died. Eventually, she got out of prostitution and, with her mother's support, went through drug rehabilitation, finished high school, and eventually received a diploma in Wife Assault and Child Advocacy from George Brown College.

In 2001, Falle began counselling women in prostitution at Streetlight Support Services, and counselled more than 800 women in the subsequent decade, 97% of whom wrote on their intake surveys that they wanted to exit the sex industry. In order to make this statistic more widely known, Falle founded Sex Trade 101. She began offering training for police and partners with the Toronto Police Service's sex crimes unit. Falle was one of the main proponents of Member of Parliament (MP) Joy Smith's private member's bill, Bill C-268, which was passed in June 2010 as An Act to amend the Criminal Code (minimum sentence for offences involving trafficking of persons under the age of eighteen years), and she helped the Canadian government formulate their appeal of the decision of the Ontario Superior Court of Justice in Bedford v. Canada to strike down various prostitution laws. Falle advocates adopting a law in Canada analogous to Sweden's Sex Purchase Act, which would decriminalize the selling of sex and criminalize the purchasing of sex.

## Early life

Natasha Falle grew up in a middle-class home in Nova Scotia, a suburb of Toronto, Ontario, and a suburb of Calgary, Alberta. Her mother managed stores in the wedding industry and her father was a police officer with a vice squad, arresting drug dealers and pimps. While she was growing up, Falle had multiple family members with addictions. When she was a young teenager, Falle's parents divorced, and she subsequently lived in a single-parent home with her mother. They moved into an apartment in downtown Toronto. Falle's father did not pay her mother alimony and there was often no money for food, so Falle began stealing food to survive.

Falle had no role models. She began writing poetry about suicide and wearing black clothing. She acted out by stealing cars and using drugs recreationally. She started out with soft drugs and then moved on to using psychedelic mushrooms and LSD. Her associates in these criminal activities, who came from similarly dysfunctional backgrounds, provided her with a sense of belonging that she no longer found at home. Falle's mother had a series of boyfriends who abused her and made the house unsafe, so Falle ran away from home. She slept at friends' houses, on their couches and in tents in their backyards, concealing her presence from her friends' parents.

## Prostitution

### Initiation

After she turned 15, Falle had her first experience in the sex industry at a party in a bar in Chinatown, Calgary. A 25-year-old man at the party who had forged identity documents convinced her to sell sex for money and give him half of her earnings. Falle found this man well-dressed and attractive and did not think that he fit the stereotypical image of a pimp. She had nowhere to sleep that night and had no money for food, so she accepted his offer. The man to whom she sold sex that night had teeth that were rotten. They had sex on a soiled mattress in a poorly-ventilated restaurant attic, and she received \$100.

Falle convinced five of her friends who came from dysfunctional homes to join her in prostitution. Falle said that she and the other girls she knew her age who had gone into prostitution had felt as though that was their only remaining option. She said they all tried to stay free of pimps, organized crime, and drugs, but eventually succumbed. Falle later said, "We were prey for every paedophile, pervert, pimp and drug dealer that was out there." The man who had recruited Falle in Chinatown became her pimp. Adults began advertising Falle's sexual services in newspapers. Falle engaged in both indoor and outdoor prostitution, working as an escort, in strip clubs, and at massage parlours. At the legal establishments where she engaged in prostitution, escort drivers engaged in organized crime, selling guns, alcohol, cocaine, and stolen goods and pimping children in massage parlours.

### Abuse

Falle's pimp kept her imprisoned for the years that he prostituted her, and trafficked her across Canada to Edmonton, Vancouver, and Kelowna. Falle later said of herself and the other women in prostitution she knew, "I couldn't admit that I was not there by choice. We couldn't live in our own skin if we admitted that. We needed to believe that it was our choice." Falle's pimp told her he would marry her if she made enough money for him through prostitution. When she had made the amount of money her pimp required, she was 17 years old. The two got married that year.

Falle's pimp regularly beat her and she began to suffer from battered person syndrome. The worst beating she received was in a brothel where she and four other teenagers engaged in prostitution. Over the years that she was married to her pimp, she hoped he would change. She later said that she "wanted to love him, believe he'd never do it again." She tried to make him enough money through prostitution to convince him not to find another girl. He threatened her family, and conditioned her not to call the police. Fear of being labelled a "snitch" or "rat" prevented her from going to the police. None of the other women she knew in the sex industry were in healthy relationships. At her peak, Falle owned a Ford Mustang, bought her pimp a Mercedes-Benz, and lived in a four-bedroom penthouse apartment, but she continued to experience violence. She used her material possessions to elevate herself above the women around her who were dependent on cocaine; she dismissed them as "crackheads" and "crackhoes".

Some of the men who purchased sex with Falle were police officers, and she knew girls who used drugs with their drug counselors. Falle was brainwashed to believe that her only worth lay in prostitution, which she later said was why it took her so long to exit prostitution. While engaging in prostitution, Falle was frequently threatened by other women in the sex industry, she was verbally abused by her clients, she was stalked, and she was threatened with guns on multiple occasions, including by her pimp. She was also drugged several times, and was once sexually assaulted by an escort driver after having been drugged by him. At one point, she was kidnapped. She had to resort to pulling a knife on people at times when she was threatened with violence. Because of various instances of violence during her time in prostitution, Falle was injured in a variety of ways. Her pimp broke several of her bones and burned her body.

### Effects and aftermath

In order to cope with the psychological trauma of prostitution and violence, Falle became dependent on cocaine. Within two years, she was spending \$500 per night on cocaine. Her drug use resulted in delusions and caused her to become schizophrenic, and subsequently distrustful of those around her, fearing that people were coming to take away her drugs.

Eventually, the personal horror stories Falle heard from other women in her situation convinced her that she needed to get out of prostitution. Her substance dependence became so severe that she almost died. Her best friend was killed by her pimp. By this point, many of her friends in the sex industry had died, and she thought that she would be next. One night, Falle was hiding alone in a hotel room when she found a Bible placed there by Gideons International. She began reading the Psalms and later said, "For the first time I understood what God was saying to me. That night my life began to move in a new direction." On her 27th birthday, Falle left her husband and returned to her mother. Because of the delusions, Falle did not recognize her mother, thinking she had been replaced with a clone. Falle told her mother about being dependent on cocaine, entering prostitution, and marrying her pimp. Her mother accepted her back. It took several more years before Falle gave up hope that her husband would change.

Falle participated in a diversion program that allowed her to return to school. She said that she required a lot of support in order to fully exit prostitution, and found that support in her mother, her school, and Streetlight Support Services. Falle underwent a month of drug rehabilitation, which convinced her to become a counsellor. She then spent 90 days at the Toronto West Detention Centre and took seven months to finish high school. She went on to graduate from a college and a university. She received a diploma in Wife Assault and Child Advocacy from George Brown College and graduated with honours. To remove the burns her pimp had inflicted on her, she had laser surgery that cost thousands of dollars.

Falle uses the term "sex trade survivor" to describe herself. In 2012, Falle sought to bring a lawsuit against her former pimp to demonstrate to other women who are or have been forcibly prostituted that it is possible to oppose one's pimp. In 2013, she said that, while she has left much aggression and profanity behind since leaving the sex industry, some of the issues that she developed will stick with her until the end of her life. Jonathan Migneault of the Sudbury Star wrote that "Falle's story about her descent in and escape from prostitution is so horrific you almost don't believe the details." Sam Pazzano of the Toronto Sun wrote that, after 12 years of prostitution, Falle still has "attractive looks and [a] sharp mind."

## Activism

### Streetlight Support Services

In 2001, Falle began counselling women in prostitution at Streetlight Support Services. Once she received her diploma from George Brown College, she redeveloped the organization's Choices program, a four-week program that helps women leave prostitution. Between 1997 and 2005, Choices helped 325 women out of the sex trade. Women entered into the program through the criminal justice system after being arrested for offences related to prostitution. By 2011, Falle had conselled more than 800 women. Of these women, 97% wrote on their intake surveys that they wanted to exit the sex industry, and 95% wrote that they had been physically abused by either a pimp, a client, or another woman in the sex industry. Falle believed these statistics to be representative of women in prostitution in Canada because the women she counselled did not come to her by choice; they were required to undergo the counselling by court order. In 2010, Falle said that hundreds of the women she had met were controlled by pimps, either as sexual partners or as traffickers. She also said that women in prostitution tend to be moved around a lot because they tend to fetch higher prices when they are new to a region. One of the women she counselled through Streetlight Support Services had been kept in a closet when she was not providing sexual services to men.

### Sex Trade 101

Because Streetlight Support Services is administered by the office of the Attorney General of Ontario, Falle could not be politically active in her capacity as a counsellor with them. Falle said, "I couldn't be silent anymore, because the strong voices that we were hearing were the minority few saying [prostitution] is liberating, it's a job choice ... I couldn't stand that lie anymore." She said that this minority only want to continue in prostitution because "they've never had a healthy comparison." Falle founded and became the director of Sex Trade 101, a Toronto-based nonprofit organization dedicated to protecting the rights of women who have been in prostitution. She founded Sex Trade 101 because she saw women in the sex industry as victims rather than as criminals, and she believed that the Criminal Code depicted them as criminals. She founded the organization in order to make it more widely known that the vast majority of women in the sex industry want to leave. She met many women who had no family to support them in exiting the sex industry, and she decided to found an organization to provide that support. The organization helps prostitution victims exit the sex industry and improve their lives.

Sex Trade 101 calls itself "Toronto's only sex trade survivors and abolitionist organization" and is composed of 12 women who had escaped forced prostitution and who offer training for police in order to change perceptions about people involved in the sex trade. The organization is also involved in prevention work in high schools. The organization offers mentorship and peer support groups for women who are currently or were formerly in prostitution. It also partners with other organizations including the Servants Anonymous Society of Calgary and Sheatre, an Owen Sound-based interactive theatre company. In 2012, Falle and Bridget Perrier represented Sex Trade 101 in Owen Sound where they launched ReStart, a mentorship program to aid women and youth who are at risk of sexual victimization, to help people exit the sex trade, and to provide support once they have exited. At the workshop that launched this program, Falle said that some people in the sex industry engage in survival sex—trading sex for food, a place to sleep, alcohol, drugs, or the feeling that they are loved. She said that rural communities are common places for women and children to be groomed into prostitution. She identified Bruce—Grey—Owen Sound as particularly vulnerable because of the substance abuse in the area. By 2013, Sex Trade 101 had served more than 1,000 women. In May 2012, Falle spoke about human trafficking at high schools in Vancouver primarily because of the influence of a parent concerned about human trafficking in the area and its dangers to children.

### Police

Falle is a professor at Humber College in Toronto, where she teaches in the police foundations program, educating on the subject of social justice. One of her students, Brittany Swartzentruber, said that Falle's lectures would have a significant impact on Swartzentruber's career as a police officer, saying, "She was one of the best speakers I have ever heard." Falle partners with the Toronto Police Service's sex crimes unit, which developed the statistic that the average age for a girl to become involved in prostitution in Canada is 14. According to Falle's research, the average age is between 13 and 16. York Regional Police drugs and vice unit Detective Thai Truong said that all law enforcement personnel should hear stories like Falle's because "they remind us that they are not just prostitutes or escorts, but they are somebody's daughter, somebody's sister, somebody's wife."

### Bill C-268

In June 2010, Kildonan—St. Paul Member of Parliament (MP) Joy Smith's private member's bill, Bill C-268, was passed as An Act to amend the Criminal Code (minimum sentence for offences involving trafficking of persons under the age of eighteen years). Falle was one of the bill's five main proponents, the others being Timea Nagy, a woman who was trafficked from Hungary to Canada at the age of 20 and kept as a sex slave in a strip club; Tamara Cherry, a Toronto Sun journalist who writes about human trafficking in Canada; Brian McConaghy, a former RCMP officer who works with Ratanak International—another anti-human-trafficking organization; and Grand Chief of the Assembly of Manitoba Chiefs Ron Evans, who raised awareness about victims of the sex industry in Manitoba. Later that year, Falle said that the best way to fight human trafficking is to influence public opinion.

Later in 2010 in Winnipeg, Falle received an award at the first annual Honouring Heroes ceremony, which was organized by Joy Smith, who is also an anti-human-trafficking activist. Falle called Smith an angel, saying, "It's only been in the last few years since all those missing and murdered aboriginal women turned up dead did anybody care about us. So to have her speak out the way she is against [prostitution] is so empowering." Falle was one of five award recipients at the ceremony, the other four being the other primary proponents of Bill C-268.

### Bedford v. Canada

Falle took a sustained interest in Bedford v. Canada, a case that began in the fall of 2010 when laws against keeping a brothel, communicating in order to facilitate prostitution, and living off the avails of prostitution were struck down by the Ontario Superior Court of Justice as unconstitutional. When Falle first heard about this decision, she cried, and expressed surprise and disappointment. She said that she did not believe that the average Canadian realized the implications of striking down these laws, which she said were that "your next door neighbours can run a brothel right beside you. Your children could be exposed to condoms left on their driveway, johns propositioning them." She also expressed her concern that striking down anti-prostitution laws normalizes prostitution as a career option for children to consider. She also said that, without the laws, "pimps will be legitimate businesspeople [and] billboards advertising brothels could start appearing on roadsides." Ron Marzel, a lawyer attempting to have the anti-prostitution laws declared unconstitutional, said, "the reality is there are consenting adults who want to go into" prostitution. Falle was angered by this statement, and replied that 97% of women in the sex industry are not engaging in prostitution by choice, and that "the voices of the overwhelming majority of women who want to get out of prostitution are being drowned out by a vocal few."

The Government of Canada disagreed with Ontario's ruling and announced that the decision would be appealed. Falle helped the Crown formulate their case. That November, Falle was a panelist at a debate about the case. At the debate, which was hosted by the Queen's Law and Public Policy Club at Queen's University, Falle said that, whatever the outcome of the appeal, she was glad that the case was removing the taboos surrounding talking about prostitution. The following June, Falle and more than twelve others intervened in the case, arguing that the laws should be reinstated to protect women from pimps. As part of this anti-prostitution coalition, Falle said that all areas of the sex industry are unsafe, including escort agencies and strip clubs. Falle testified in the lower courts, saying that "the more they say the women are there by choice, the harder it is for us to convince police, social workers and everyone else that these people are vulnerable." She also said that striking down anti-prostitution laws sends a message to men that it is acceptable to harass women. Falle said that all the applicants in Bedford v. Canada—including Terri-Jean Bedford, the dominatrix seeking to have the anti-brothel law struck down—entered the sex industry as children. For this reason, Falle argued that neither of these women chose a life of prostitution as a consenting adult; that life had already been chosen for them by the time that they had reached the age of consent. Falle encouraged Canadians to take an interest in Bedford v. Canada because any Canadian girl can become a victim of prostitution.

Eventually, the laws against pimping and communicating provisions were determined to be constitutional and were retained. As of 2013, the constitutionality of the law against brothels was still under review. Falle said that brothels should not be legalized, saying, "I know firsthand and... from the disclosures of the many women that I've been able to counsel over the years that most violence happens behind closed doors." Falle said that if brothels were legalized, the police would have less legal ability to find women and children victimized by human trafficking. Bedford said that legalizing brothels would make prostitution safer for women because it would allow prostitution to take place indoors. Falle responded that most prostitution has already moved indoors, as the internet has made most street solicitation unnecessary, and argues that the move indoors has not made the women safer. She said that when she was in prostitution, she was even a little safer outdoors because of the added visibility to the public. In discussion of the case, Falle said, "I don't know anyone in the prostitution business who hasn’t ended up dead, in jail, or on drugs." She intended to bring a pimp stick to the Supreme Court of Canada in June 2013 as a visual aid in explaining how pimps often abuse the women they prostitute.

### Advocacy events

In October 2010, Falle picketed a courthouse in downtown Toronto in recognition of International Day of No Prostitution. She was joined by Trisha Baptie, Bridget Perrier, Katarina MacLeod, and Christine Barkhouse; all were former human trafficking victims and sex workers. At the protest, Falle said that "only 1% of prostitutes say they enjoy sex with johns and 97% say they want to get out."

In January 2011, Falle appeared at the Party for Freedom at York University in Toronto, which launched the Alliance Against Modern Slavery, a nonprofit organization seeking to combat human trafficking through partnerships, education, and research. Falle was joined by Glendene Grant, human trafficking victim Jessie Foster's mother; Kevin Bales, co-founder and president of Free the Slaves; Kate Todd, a singer-songwriter and actor; Janelle Belgrave of Samba Elégua Drummers and Peace Concept; Roger Cram of Hiram College; and Jeff Gunn, a guitarist. That September, Falle attended Toronto's second annual Freedom Walk, which was hosted by Stop Child Trafficking Now, Freedom Relay Canada, and Oakville's Free-Them. At this event aimed at raising awareness about human trafficking nationally and internationally, Falle was joined by other abolitionists including Tara Teng, who was Miss Canada at the time; Trisha Baptie, co-founder of EVE; Shae Invidiata, founder of Free-Them; Timea Nagy; Constable Lepa Jankovic; MP Joy Smith; MP Olivia Chow; and MP Terence Young.

In October 2012, Falle was the keynote speaker at a symposium on street prostitution and human trafficking, which took place at Croatian Hall in Greater Sudbury and was attended by approximately 100 people. Sergeant Corinne Fewster of the Greater Sudbury Police Service said that the manner in which Falle was aided in her transition out of prostitution is a good example for local social and health services to follow in helping other women exit the sex industry. The following month, Falle told her story at a lunch hosted by The Salvation Army in Vancouver. That year, Falle said, "Where there's high-track prostitutes, escorts, strippers and masseuses; there's pimp violence."

In March 2013, Falle spoke at the 57th Session of the United Nations Commission on the Status of Women, where she sat on a panel called "Survivors Speak: Prostitution and Sex Trafficking" that was sponsored by Coalition Against Trafficking in Women. That May, Falle spoke at a fundraiser for the Servants Anonymous Society of Calgary, an organization that supports sex workers in the city. It was her first time in Calgary since she had been part of the sex industry there. The fundraiser was called "Cry of the Streets: An Evening for Freedom" and raised money for Servants Anonymous Facilitates Exit, a women's shelter for those seeking to leave the sex industry. At the fundraiser, Falle advocated adopting a law in Canada analogous to Sweden's Sex Purchase Act, which decriminalized the selling of sex and criminalized pimping and the purchasing of sex. Falle argued in support of modelling prostitution law in Canada after Sweden's laws because she believes that prostitution cannot be regulated. At another time, she had said that "it's not Canada's laws that make prostitution unsafe, it's the johns who are raping and abusing the women and their pimps." Also in 2013, Falle said that the average age at which girls get involved in prostitution in Canada is 14, and that their emotional intelligence does not develop while in prostitution, preventing many of them from leaving. She also said that prostitution has grown significantly in Canada since she left the sex industry, and that most human trafficking victims in Canada were born and raised in Canada.

## See also

- List of kidnappings

Canadian academic


History of the Jews in Dęblin and Irena during World War II

Thousands of Jews lived in the towns of Dęblin and Irena [pl] in central Poland before World War II; Irena was the site of the Polish Air Force Academy from 1927. In September 1939, the town was captured during the German invasion of Poland and the persecution of Jews began with drafts into forced labor and the establishment of a Judenrat ("Jewish Council"). A ghetto was established in Irena in November 1940. It initially consisted of six streets and was an open ghetto (the Jews were not allowed to leave without permission, but non-Jews could enter). Many ghetto inhabitants worked on labor projects for Dęblin Fortress (a German Army base), the railway, and the Luftwaffe. Beginning in May 1941, Jews were sent to labor camps around Dęblin from the Opole and Warsaw ghettos. Conditions in the ghetto worsened in late 1941 due to increased German restrictions on ghetto inhabitants and epidemics of typhus and dysentery.

The first deportation was on 6 May 1942 and took around 2,500 Jews to Sobibór extermination camp. A week later, two thousand Jews arrived from Slovakia and hundreds more from nearby ghettos that had been liquidated. In October that same year, the Irena ghetto was liquidated; about 2,500 Jews were deported to Treblinka extermination camp while some 1,400 Jews were retained as inmates of forced-labor camps in the town. Unusually, the labor camp operated by the Luftwaffe—employing, at its peak, about a thousand Jews—was allowed to exist until 22 July 1944, less than a week before the area was captured by the Red Army. One of the last Jewish labor camps in the Lublin District, it enabled hundreds of Jews to survive the Holocaust. Some survivors who returned home were met with hostility, and several were murdered; all left by 1947.

## Background

Dęblin and Irena [pl] (Yiddish: מאדזשיץ, Modzhitz) are located 70 kilometers (40 mi) northwest of Lublin in Poland, at the confluence of the Vistula and Wieprz rivers and at an important junction on the Lublin–Warsaw rail line. Local Jews supported the January Uprising in 1863. By the end of the nineteenth century, the area was a center of Hasidic Judaism; most Jews followed the Modzitz dynasty, named after the town of Irena, and there were also Gur Hasidim. Modzitz Rebbe Yisrael Taub settled in the town in 1889. Other Jews were Bundists or Zionists. Many Jews made their living as peddlers, shopkeepers, or artisans (especially in leatherwork and metalwork) and later participated in the development of the town as a summer resort. In 1927, the civilian population of Dęblin and Irena was 4,860, including 3,060 Jews. Beginning in 1927, Dęblin housed the Polish Air Force Academy, whose airfield was one of the largest in Poland. In 1936 and 1937, there were some incidents in which non-Jewish merchants vandalized their Jewish competitors' shops.

## German invasion

During the German invasion of Poland which started World War II in Europe, the Luftwaffe bombed Dęblin between 2 and 7 September 1939, targeting the military airfield, Polish Army ammunition stores, and a nearby bridge over the Vistula. Most Polish soldiers left the town on 7 September; on 11 September the remaining Polish forces detonated the ammunition and withdrew. The German Army arrived on 12 or 20 September. Many residents, both Jews and other residents of the area, had fled to escape the aerial bombardment. In the following weeks, most returned at the urging of the Germans. The Jews found that their properties had been ransacked and plundered. They were conscripted into forced-labor units that cleared up the bomb damage and repaired some of the buildings, and the Jewish community was forced to pay a fine of 20,000 zlotys. During the German occupation, Dęblin and Irena became part of the Lublin District of the General Governorate occupation region. In late 1939, Jews were required to wear armbands with the Star of David, and some were conscripted for forced labor in Janiszów, Bełżec, and Pawłowice.

At the end of the year, the German occupiers forced Jews to form a Judenrat ("Jewish Council"), as an interface between the Jewish community and Nazi demands. The Judenrat was formed later than others in the region and had its offices on Bankowa Street. The first Judenrat officials worked to reduce the impact of German demands and warned residents of searches. According to the Encyclopedia of Camps and Ghettos, the first chair of the Judenrat, Leizer Teichman, was dismissed in early 1941; survivors reported that he and his secretary were deported to Wąwolnica with their families after trying to bribe policemen. The Judenrat was run temporarily by Kalman Fris, who resigned after a few months and was replaced by Vevel Shulman in August and then a Konin native named Drayfish in September 1941. According to Israeli historian David Silberklang, "Szymon Drabfisz" was the first chair and Teichman the second.

## Ghetto

A Nazi ghetto was established in Irena in November 1940 or early 1941, probably on the orders of Kurt Lenk, the region's new land commissioner. Some Jews were forced to move from their homes to the ghetto. The motive was probably to keep Jews away from the two nearby military installations, a Luftwaffe airfield and a German Army base, which were filled with troops in preparation for the 1941 invasion of the Soviet Union. The decree establishing the ghetto was not formally issued until 1942. Until 1942, Irena was an open ghetto (there was no physical fence, although the penalty for unauthorized departure was death) consisting of six streets. Its boundaries were initially Okólna Street, the Irenka River, Bankowa, and Staromiejska Streets. Staromiejska Street was removed from the ghetto in September 1942; its residents moved to Bankowa. Initially, non-Jews were allowed to enter and perhaps live in the ghetto, so many of the Jews were able to survive by trading material goods for food with them. A Polish-language secular school, run by Aida Milgroijm-Citronbojm, instructed 70 to 100 pupils. The Jewish Ghetto Police maintained order in the ghetto, and on the outside it was guarded by a Sonderdienst ("Special Services") unit consisting of local Volksdeutsche (ethnic Germans) and Ukrainian Auxiliary Police. The Sonderdienst unit was known for beating, humiliating, and killing ghetto inmates.

In late 1941, non-Jewish Poles were banned from entering the ghetto, resulting in waste piling up and typhus and dysentery epidemics. The sick were treated at a thirty-bed hospital run by Isaar Kawa, a doctor from Konin; with the help of non-Jewish doctors, medicines were imported from Warsaw. Simultaneously, new restrictions were imposed by the Germans: the use of stoves was banned, winter clothing confiscated, and fuel imports forbidden. More Jews began to leave the ghetto to obtain essentials, resulting in twenty young women being discovered outside the ghetto and killed. Because conditions were better in Lublin District than in Warsaw, some Jews fled to towns there including Irena; twenty Jews were executed for being unregistered refugees. The Judenrat's leadership was altered again as Drayfish was executed, accused of filing complaints with the Puławy County administration; he was replaced by the businessman Yisrael Weinberg.

In March 1941, 3,750 Jews lived in the ghetto, including 565 who were not from the area. These consisted mainly of Jews expelled from Puławy and the German-annexed Reichsgau Wartheland, as well as some Jews who had come to live with relatives to avoid being trapped in the Warsaw Ghetto. Overcrowding was severe; there were 7–15 people per room. More Jews arrived in May 1941: 1,000 from the ghettos at Warsaw, Częstochowa, and Opole, although they were not housed in the ghetto but at work sites in the area. On 13 and 14 May 1942, two transports totalling 2,042 Slovak Jews arrived from Prešov. Although they brought food with them, they were unused to the harsh conditions and many died of typhus due to the poor sanitation. More than 2,000 Jews were brought to Irena from nearby towns, having survived the liquidation of their communities because they had been selected to work. These included 300 people from Ryki, 300 from Gniewoszów and Zwoleń, and a group from Stężyca. Other Jews had entered the ghetto after escaping roundups in Baranów, Ryki, Gniewoszów, and Adamów (from early October). In August, 5,800 Jews were reported to be living in the ghetto, of whom only 1,800 were from Dęblin.

## Forced labor

Until late 1942, Jews earned wages as forced laborers, except those who were conscripted by the municipality for tasks such as street cleaning or snow clearing. Some Jews worked for German companies such as Schwartz and Hochtief, contracted by the Wehrmacht to do construction on the military bases in the town. The Ostbahn contracted Schultz to improve the railway between Lublin and Warsaw, a project which employed 300 Jews beginning in June 1941. Working conditions at this site were very harsh with Jews forced to work twelve hours each day. However, most Jews worked for the German Army or the Luftwaffe; women and children grew food for the military. Dęblin Fortress, which had been taken over by the German Army and where around 200 Jews from the ghetto worked, was the site of Stalag 307, a camp for Soviet prisoners of war, from 10 July 1941. An estimated 80,000 Soviet prisoners were imprisoned there; all died within a few months of starvation and disease. The Jews who worked there saw prisoners held in worse conditions than their own. The Jews from Dęblin and Irena tried to take the best jobs; 200 of the Slovak deportees ended up working for the municipality. An additional 200 of the Slovaks worked for the Schultz firm following an expansion. Survivors recalled that although German soldiers supervising the forced laborers tended to treat them relatively well, some ethnic Polish supervisors beat Jews, and the Ukrainian guards at the railway project were especially harsh, threatening prisoners with death.

In May 1941, in preparation for the invasion of the Soviet Union, the Luftwaffe commissioned the firm Autheried to improve the airfield. They recruited 200 Jewish workers in Opole, many of whom were Austrian Jews who had been deported from Vienna in February 1941 and wanted to escape the harsh conditions in Opole. These Jews, as well as another four hundred recruited in Irena and the Lipowa 7 camp in Lublin, and 500 ethnic Polish workers, worked twelve hours a day levelling the runway and building roads, walls, air-raid shelters, and earth fortifications. They also built a barbed-wire-enclosed complex adjacent to the runway for craftsmen to work in. Following the completion of the project in October, the workers were dispersed to various projects around the town, an action which may have been related to the typhus epidemic. Some of the Jews from Vienna stayed at the airfield, having obtained permits to work on gravel transportation and building fuel tanks; they were the nucleus for the later labor camp. After the first deportation from the Irena ghetto on 6 May 1942, additional Jews tried to secure permits, often obtained through bribery, to avoid deportation. By October 1942, the camp had 543 legal residents. At this point there were five forced-labor camps around Dęblin: the Luftwaffe camp, the fortress, the train station (300 Jews), Lipova Street (200 Jews and ethnic Poles), and another near the municipal boundary (300 Jews).

## Deportation, murder, and liquidation

### First deportation

The first deportation was on 6 May 1942, to clear space for the Slovak deportees who arrived a week later. A force composed of German and Ukrainian police and a few German SS and Ukrainian SS auxiliaries ordered Jews to assemble in the main square at 09:00. According to Yad Vashem, the Blue Police also participated. Probably because the Luftwaffe was seeking permission to recruit more laborers, the Jews had to wait for five hours. At 14:00, 1,000 Jews who already had work assignments were separated, and local employers began recruiting an additional 200–500 Jews. Forty or fifty people were killed trying to cross over to the working group. Around 2,300 to 2,500 Jews from Dęblin—mainly the elderly and families with small children—were marched to the Dęblin train station (some 2.5 km (1.6 mi) away) at 18:00 and sent to Sobibór extermination camp. The victims were beaten, and those who could not keep up were shot. Only late in the night were the Jews selected for labor allowed to return home. Every family was affected. After the deportation, the bodies of the dead were collected at the synagogue and removed from the ghetto in carts.

During the next two days, more than four thousand Jews from north Puławy County—200 from Bobrowniki (6 May), 1,800 from Ryki (7 May), 125 from Stężyca (7 May), 1,500 from Baranów (8 May) and 500 from Łysobyki (8 May)—were also marched to Dęblin and deported to Sobibór. This was half of a countywide extermination action in Puławy County—the first in a series of systematic, countywide deportations in the Lublin District as part of Operation Reinhard. The county's administrator, Alfred Brandt [de] ordered it shortly after Sobibór became operational; he was present in Dęblin during the deportation.

### Second deportation

The next deportation, which coincided with the liquidation of the two remaining ghettos in Puławy (Opole and Końskowola), took place on 15 October 1942; the previous day, all workers had been ordered to remain at their workplaces. Under the command of SS-Obersturmführer Grossman, gendarmes, Luftwaffe soldiers, and police auxiliaries rounded up the remaining Jewish population. Many Jews tried to avoid the roundup and enter the Luftwaffe camp, which at that time had 543 official residents. Some were turned away by the Jewish police there. Israeli historian Talia Farkash estimates that about 500, including 60–90 young children, managed to get in, mostly by paying bribes. Silberklang's estimate is that 1,000 Jews entered the camp, including 280 women and 100 children. The Jewish elder, Hermann Wenkert, later claimed that if he had allowed non-registered Jews entry, and they were discovered, everyone in the camp would have been killed. Instead, he bribed the Germans to obtain legal permits for an additional 400 Jews, which were mostly given to the wives and children of current residents.

Many of the Slovak Jews, not knowing what to expect, had lingered in the ghetto while packing their bags. About 215 to 500 Jews were shot and killed by the Ukrainians and Germans while the houses were being cleared. Between 2,000 and 2,500 people were deported to Treblinka extermination camp; mostly the Slovaks. Members of the Judenrat, Jewish police, and their families, about 100 in total, were retained to clean up the ghetto. Perhaps another 100 Jews were hiding without permission in the ghetto. On 28 October, the remaining Jews were sent to the Schultz labor camp, and the ghetto was officially liquidated. Either on that day or within the next few weeks, 1,000 people were removed from the labor camps, including the 200 workers at Dęblin Fortress and 200 to 500 of the Luftwaffe's workers. They were deported to Końskowola or Treblinka.

After this, Farkash estimates that there were about 1,400 Jews were still alive in the labor camps around Irena. Estimates of the number at the Luftwaffe camp range from 1,000 to 2,000, with another 300 at a railway camp near the passenger station, and 120 at another camp near the railway loading station. During May or July 1943, most of the remaining prisoners in the railway camps were deported to Poniatowa concentration camp via the transit ghetto in Końskowola; the rest were deported in late 1943. Hundreds of Jews were still alive in the remaining labor camps in Puławy County, but they were murdered during Operation Harvest Festival (2–3 November 1943). This operation also killed the prisoners of Poniatowa but did not affect the Jews at the Luftwaffe camp in Irena, who were unaware of it. Following Operation Harvest Festival, there were ten labor camps for Jews in the Lublin District with more than 10,000 Jews still alive.

## Luftwaffe camp

### Camp leader

The camp leader (German: Lagerälteste) at the Luftwaffe camp was Hermann Wenkert, a Jewish lawyer from Vienna who had been deported to Opole in 1941 and came to Irena with the volunteers. According to Wenkert's account, soon after his arrival he happened upon Eduard Bromofsky, an Austrian Luftwaffe officer with whom Wenkert had served during World War I. Bromofsky convinced the commander of the camp to promote Wenkert to his position. Even after Bromofsky was reassigned, the relationships that Wenkert had cultivated with the German authorities enabled him to retain his position. Wenkert populated the administration with other Viennese Jews, insisted on extra rations for himself and his family, and had his family transferred from Opole to Irena. Because of his close association with the German authorities and use of his position for personal benefit, some survivors considered him to be a collaborator. When it was possible, Wenkert negotiated with the authorities and paid bribes to avoid punishment; he also consulted respected prisoners in cases where the matter had not come to German attention. However, when he thought there was no other option and the survival of the camp was in the balance, Wenkert turned Jews accused of wrongdoing over to the Germans even though he knew that this would result in their deaths. In general, German-speaking Jews had a more positive impression of him than the local Jews, who spoke Yiddish. According to Silberklang, both views of Wenkert may be true: that he tried to secure benefits for himself personally while also working to keep the other Jewish prisoners alive.

### Conditions

Conditions were relatively good compared to other camps. Residents received food rations of about 1,700 kilocalories daily (enough to live), weekly baths, and medical care, and were allowed to practice their religion. The presence of 100 young children—whose existence was justified as increasing the productivity of their parents—was particularly unusual and not matched elsewhere in the Lublin District after Operation Harvest Festival. Aida Milgroijm-Citronbojm continued the Polish-language school that she had run in the ghetto. The SS, who did not know about the children's existence, periodically conducted surprise inspections of the camp, during which the children had to hide. According to Farkash, Wenkert's positive relationship with the camp command was partially responsible for the good conditions. Wenkert managed to get religious Jews exempted from working on Shabbat and allowed a chevra kadisha society to operate, burying the dead according to Jewish law. In 1943, about 80 Jews (including Wenkert) received kosher food for the week of Passover.

The camp had three German commanders, all at the rank of sergeant major: Kattengel (through March 1943), Dusy, and Rademacher (during the last two months). Although Kattengel was distrusted because he roamed the camp with a dog and whip, both Dusy and Rademacher were described positively by survivors. Dusy even arranged for one woman to receive treatment at a German hospital after she was badly injured. Wiszniewski, a local Volksdeutscher, was the lead foreman of the Jewish laborers and did not mistreat them. Autheried engineer Kozak brought food to the Jews he worked with. The Slovak Jews at the camp were the last major group to survive of the almost 40,000 Slovak Jews deported to the Lublin District in 1942. The Jews remaining in Prešov hired couriers (non-Jews from the Polish–Slovak border) to travel to the camp regularly until it was dissolved, carrying letters and bringing valuables and money. A committee was formed in the camp to distribute these among the Slovak Jews. Cases of theft by Polish Jews led to friction between the two communities.

Despite the relatively good conditions, some Jews tried to escape because they feared that the camp would be liquidated. The Luftwaffe camp command imposed collective punishment to deter escapees. Both theft and possessing foreign currency were punishable by hanging. Punishment could be excessive and arbitrary—nine Jews were executed by firing squad for causing an accidental fire—but less so than other Nazi camps. Reports of the Home Army from that time often described the local ethnic Polish population as hostile to "Jewish bands" which stole food and resources from Polish peasants; said reports, according to Zimmerman, therefore can be seen as reflecting "the eerie distance of mere observers". According to Farkash, in 1943, Wenkert allowed a group of Jewish partisans seeking refuge from a hostile unit of the Polish Home Army resistance group into the camp. Most Jews who tried to escape were captured, and others returned to the camp. Several members of the Kowalczyk peasant family were honored as Righteous Among the Nations for sheltering nine Jews who had escaped from the Łuków ghetto and the Dęblin camp on their farm in Rozłąki [pl], from late 1943 to mid-1944. Árpád Szabó, a Hungarian military doctor, was similarly recognized for smuggling a Slovak Jewish couple from the camp to Hungary.

### Liquidation

The camp was liquidated on 22 July 1944, the same day that the Red Army reached Lublin. At the time of the deportation, some 800 to 900 Jews were still alive, of whom 400 to 600 were from Dęblin or Irena, 40 from Vienna, and 70 to 80 from Slovakia. Two or three transports departed from Irena on 17 and 22 July. (Silberklang reports a third on 20 July). Since no one was willing to volunteer, the first transport carried about 200 people who had few connections in the camp, including single adults and fifteen children between three and six years of age. On the orders of camp security officer Georg Bartenschlager, the fifteen children were shot and killed upon their arrival in Częstochowa. The rest of the prisoners, including Wenkert and 33 young children, departed on the second transport with Rademacher, the German camp commander, who carried a letter of protection from the airfield commander. About fifty Jews escaped from the camp during the evacuation, but most were killed. The second transport arrived in Częstochowa on 25 July; the young children were kept separated for two and a half days but were not killed. The reason for this is not clear, but likely involved negotiations with Bartenschlager. In Częstochowa, the Jews worked at the four Hugo Schneider AG plants. Some were deported to Buchenwald and Bergen-Belsen in January 1945 while others were liberated by the Red Army. About half survived.

According to Farkash, the airfield camp in Irena is a "singular case" in the Lublin district and possibly in the entirety of occupied Poland. One unusual aspect of the camp was that it was run from start to finish by the Luftwaffe rather than the SS, although during 1943 and 1944, the SS tried to expand its role. According to Silberklang, being run by the Wehrmacht helped to ensure the camp's survival. However, other camps (such as three in Zamość county) were run by the Luftwaffe and were still liquidated before the end of 1943. Another reason was the importance of Jewish workers at Dęblin to the German war effort, which was continually emphasized by Wenkert in his dealings with the Germans. Overall, luck played a major role in the fortuitous combination of Jewish leadership, relatively friendly Germans, and the evacuation of the camp to Częstochowa rather than Auschwitz-Birkenau. Ninety-nine percent of the Jews from Lublin District were killed in the Holocaust, but hundreds from Dęblin and Irena survived the war.

## Aftermath

Dęblin was liberated by the 1st Polish Army by 27 July 1944. In 1945, Alfred Brandt was convicted and sentenced to death for his crimes by a Soviet military tribunal near Słupsk and executed by firing squad. Some of the Jewish survivors, who attempted to return home in January 1945, were told by the local Milicja Obywatelska police chief that it was illegal for them to settle in the town. In March 1945, survivors Łaja, Gitla and Fryda Luksemburg were murdered with the involvement of local police. Other Jews received threatening letters. Following the 1946 Kielce pogrom, a meeting in Dęblin was held to commemorate the victims. Locals interrupted the meeting with anti-semitic slogans. By 1947, no Jews remained in the town, which is still the case in the twenty-first century. As of 2011, the Dęblin Synagogue [pl] was in poor condition and being considered for demolition. A memorial plaque about the Irena ghetto was unveiled in Dęblin in 2015.

Ignaz Bubis, a former prisoner of the ghetto, was the chairman of the Central Council of Jews in Germany from 1992 to 1999.

Ghetto in Poland