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The Tooth Cave pseudoscorpion ''( Tartarocreagris texana)'' is a small cave-dwelling
arachnid Arachnids are arthropods in the Class (biology), class Arachnida () of the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, opiliones, harvestmen, Solifugae, camel spiders, Amblypygi, wh ...
of the family
Neobisiidae Neobisiidae is a family of pseudoscorpions distributed throughout Africa, the Americas and Eurasia and consist of 748 species in 34 genera. Some species live in caves while some are surface-dwelling. Characteristics The body color ranges from re ...
. The species was originally assigned to the genus ''Microcreagris'', and then reassigned to ''Austalillocregris'' before arriving at its current genus, ''Tartarocreagris''. The Tooth Cave pseudoscorpion has a small geographic distribution, known to occur in only two caves in
Travis County Travis County is located in Central Texas. As of the 2020 census, the population was 1,290,188. It is the fifth-most populous county in Texas. Its county seat and most populous city is Austin, the state's capital. The county was established i ...
, Texas. Additionally, it is tentatively identified in two nearby caves in Texas, but more research is necessary to accurately define the range where the species lives. There is little known about the biology, life cycle, and life history of the Tooth Cave pseudoscorpion due to a lack of research and study. Because of this, the Tooth Cave pseudoscorpion is listed on the
IUCN Red List The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is an inventory of the global conservation status and extinction risk of biological ...
as data deficient. Under the
Endangered Species Act The Endangered Species Act of 1973 (ESA; 16 U.S.C. § 1531 et seq.) is the primary law in the United States for protecting and conserving imperiled species. Designed to protect critically imperiled species from extinction as a "consequence of e ...
, the Tooth Cave pseudoscorpion is listed as
endangered An endangered species is a species that is very likely to become extinct in the near future, either worldwide or in a particular political jurisdiction. Endangered species may be at risk due to factors such as habitat loss, poaching, inv ...
.Hammerson, G., Schweitzer, D., Master, L., Cordeiro, J., Tomanio, A., Oliver, L., & Nichols, J. (2008). "Ranking Species Occurrences: A Generic Approach and Decision Key". ''NatureServe''. https://www.natureserve.org/sites/default/files/eo_rank_specifications-generic_guidelines_and_decision_key_05.08.2020.pdf Retrieved 9 April 2022. Recovery plans for the species focus mostly on preserving the
karst Karst () is a topography formed from the dissolution of soluble carbonate rocks such as limestone and Dolomite (rock), dolomite. It is characterized by features like poljes above and drainage systems with sinkholes and caves underground. Ther ...
ecosystem that the pseudoscorpions live in, especially including lessening human impacts on the environment.


Description

Little information exists describing what Tooth Cave pseudoscorpions look like since they live in a few dark caves in Texas and are difficult to find.O'Donnell, L., Elliott, W. R., & Stanford, R. (1994). ''Recovery Plan for Endangered Karst Invertebrates in Travis and Williamson Counties, Texas'' https://ecos.fws.gov/docs/recovery_plan/940825.pdf Retrieved Apr 9, 2022. Some inferences are made about the Tooth Cave pseudoscorpions, however, through the commonalities of the
Neobisiidae Neobisiidae is a family of pseudoscorpions distributed throughout Africa, the Americas and Eurasia and consist of 748 species in 34 genera. Some species live in caves while some are surface-dwelling. Characteristics The body color ranges from re ...
family and other cave-adapted species. Tooth Cave pseudoscorpions fall under the category of
troglobites A troglobite (or, formally, troglobiont) is an animal species, or population of a species, strictly bound to underground habitats, such as caves. These are separate from species that mainly live in above-ground habitats but are also able to live u ...
, which are species that spend their entire lives in caves. Some distinct features of troglobites include attenuated
palps Pedipalps (commonly shortened to palps or palpi) are the secondary pair of forward appendages among chelicerates – a group of arthropods including spiders, scorpions, horseshoe crabs, and sea spiders. The pedipalps are lateral to the chelicera ...
and long, slender
appendage An appendage (or outgrowth) is an external body part or natural prolongation that protrudes from an organism's body such as an arm or a leg. Protrusions from single-celled bacteria and archaea are known as cell-surface appendages or surface app ...
s.Campbell, L. (2003). ''Endangered and Threatened Animals of Texas; Their Life History and Management''. Texas Parks and Wildlife. (pg 117-121) https://tpwd.texas.gov/publications/pwdpubs/media/pwd_bk_w7000_0013.pdf Retrieved Apr 9, 2022. Tooth Cave pseudoscorpions are large for cave-dwelling species in this area, measuring about 4 mm (0.15 in) in length. The species has no tail and no stinger, but has tiny pincers which it uses to capture small
arthropod Arthropods ( ) are invertebrates in the phylum Arthropoda. They possess an arthropod exoskeleton, exoskeleton with a cuticle made of chitin, often Mineralization (biology), mineralised with calcium carbonate, a body with differentiated (Metam ...
s.Endangered and Threatened Wildlife and Plants; Final Rule to Determine Five Texas Cave Invertebrates To Be Endangered Species. (1988). ''Federal Register'', ''53''(180), 51-55. https://www.govinfo.gov/content/pkg/FR-1988-09-16/pdf/FR-1988-09-16.pdf#page=51 Retrieved Apr 9, 2022. The Tooth Cave pseudoscorpion also has a set of small teeth on these tiny pincers that help in latching onto its prey. Pseudoscorpions, like arthropods, have a protein called
sclerotin Sclerotin is a component of the cuticle of various Arthropoda, most familiarly insects. It is formed by cross-linking members of particular classes of protein molecules, a biochemical process called sclerotization, a form of tanning in which ...
on their outer layer. This protein gives the specimen a hard
exoskeleton An exoskeleton () . is a skeleton that is on the exterior of an animal in the form of hardened integument, which both supports the body's shape and protects the internal organs, in contrast to an internal endoskeleton (e.g. human skeleton, that ...
, as well as a reddish color. Surface-adapted pseudoscorpions have more sclerotin than cave-adapted pseudoscorpions. Cave-adapted pseudoscorpions have reduced pigmentation and therefore less of a reddish color due to a lack of exposure to sunlight. Because Tooth Cave pseudoscorpions are cave-adapted, they have less rigid exoskeletons than surface-adapted pseudoscorpions. Tooth Cave pseudoscorpion palps are golden brown, and their body and legs are light tan. Additionally, troglobites have severe vision reduction. Tooth Cave pseudoscorpions show this characteristic through their lack of eyes. Since this species doesn't rely on eyesight, they must rely on other senses, such as sensory hairs to catch prey and navigate in the complete darkness of the caves.


Life history

There is limited information about the life history of the Tooth Cave pseudoscorpion. But, there is more information on development, reproduction, and dispersal available about the order pseudoscorpion.


Tooth Cave pseudoscorpion

The Tooth Cave pseudoscorpion lives in an environment below ground, called a subterranean environment. This species has a lifespan of more than one year, based on what scientists have seen in captivity. This type of environment does not expose Tooth Cave pseudoscorpions to the different seasons. Thus, the Tooth Cave pseudoscorpions do not have a seasonal reproduction cycle. It instead reproduces year-round. Individuals, once born, grow and develop until sexual maturity is reached. Sexual maturity is often a body length of 4.1 mm (0.161 in) for females and 3.96 mm (0.156 in) for males. The individual body parts have specific proportions once individuals reach adulthood. The top section of the exoskeleton (
carapace A carapace is a dorsal (upper) section of the exoskeleton or shell in a number of animal groups, including arthropods, such as crustaceans and arachnids, as well as vertebrates, such as turtles and tortoises. In turtles and tortoises, the unde ...
) is three times as thick as long; the femur is 1.5 times as long as the carapace; the pincer ( chela) is 2.55 times as long as the carapace; the mouth (
chelicerae The chelicerae () are the arthropod mouthparts, mouthparts of the subphylum Chelicerata, an arthropod group that includes arachnids, horseshoe crabs, and sea spiders. Commonly referred to as "jaws", chelicerae may be shaped as either articulated ...
) is two-thirds as long as the carapace. Further inspection of the female genitalia (
spermatheca The spermatheca (pronounced : spermathecae ), also called ''receptaculum seminis'' (: ''receptacula seminis''), is an organ of the female reproductive tract in insects, e.g. ants, bees, some molluscs, Oligochaeta worms and certain other in ...
) where the species intakes male sperm can distinguish Tooth Cave pseudoscorpions from other closely related species.


Pseudoscorpions in general

It must be noted that the information below is about the pseudoscorpion order as a whole, not just Tooth Cave pseudoscorpions, because there is no information available for the specific species. The information about the pseudoscorpion order is likely similar to what the information would be for the Tooth Cave pseudoscorpion. Pseudoscorpions do not take part in courtship practices or compete to find mates. To reproduce, males leave their sperm in sacs (
spermatophore A spermatophore, from Ancient Greek σπέρμα (''spérma''), meaning "seed", and -φόρος (''-phóros''), meaning "bearing", or sperm ampulla is a capsule or mass containing spermatozoa created by males of various animal species, especiall ...
s) in areas frequently visited by females. When a female sees a sperm sac, she intakes the sac into her
gonopore A gonopore, sometimes called a gonadopore, is a genital pore in many invertebrates. Hexapods, including insects, have a single common gonopore, except mayflies, which have a pair of gonopores. More specifically, in the unmodified female, it is ...
. Offspring grow inside the mother until she gives birth.. The first stage after birth is called the protonymph stage, in which the offspring typically stays close to the mother. Most pseudoscorpion species are mobile during this stage, but some are immobile and completely reliant on their mother for nutrients. The next two stages, deutonymph and tritonymph, occur as the pseudoscorpion develops and increases in body size. It sheds the exoskeleton to make room for the increasing size. Once they reach individual maximum body size and sexual maturity, pseudoscorpions reach what scientists term adulthood. Pseudoscorpions do not molt after reaching this stage and permanently leave mothers to live alone. Pseudoscorpions build silk chambers from galea, a type of tube extending from the body. These silk chambers are where pseudoscorpions molt during their various life stages (protonymph, deutonymph, and tritonymph stages). The pseudoscorpion order also uses these silk chambers to take shelter and make brood sacs for immobile young.


Ecology


Diet

Tooth Cave pseudoscorpions are food generalists and eat plants or other small animals available to them. The main diet of the Tooth Cave pseudoscorpion consists of invertivores, which are small insects, and other similar-sized arthropods, such as
mite Mites are small arachnids (eight-legged arthropods) of two large orders, the Acariformes and the Parasitiformes, which were historically grouped together in the subclass Acari. However, most recent genetic analyses do not recover the two as eac ...
s. Like other cave-dwelling species, they also rely on surface plants for nutrients. Surface plants in caves mostly consist of leaf mulch, plant roots, and organic debris. Tooth Cave pseudoscorpions also find nutrients in deposited organic matter, especially from bats and cave crickets. The cave cricket deposits feces and eggs in areas accessible to Tooth Cave pseudoscorpions. Tooth Cave pseudoscorpions also have been known to eat decaying bodies of cave crickets.


Behavior

There is currently no information on the mating behavior of Tooth Cave pseudoscorpions. However, for pseudoscorpions in general, there are no courtship behaviors associated with reproduction. Because the male leaves
sperm Sperm (: sperm or sperms) is the male reproductive Cell (biology), cell, or gamete, in anisogamous forms of sexual reproduction (forms in which there is a larger, female reproductive cell and a smaller, male one). Animals produce motile sperm ...
in sacs that a female later picks up with specialized organs, the two individuals do not meet each other, but produce
offspring In biology, offspring are the young creation of living organisms, produced either by sexual reproduction, sexual or asexual reproduction. Collective offspring may be known as a brood or progeny. This can refer to a set of simultaneous offspring ...
together. Tooth Cave pseudoscorpions are also known to
forage Forage is a plant material (mainly plant leaves and stems) eaten by grazing livestock. Historically, the term ''forage'' has meant only plants eaten by the animals directly as pasture, crop residue, or immature cereal crops, but it is also used m ...
for food at night, briefly leaving the cave to find nutrition in the surrounding plant and cricket populations near the cave.''Tooth Cave Pseudoscorpion (Tartarocreagris texana) 5-Year Review: Summary and Evaluation.'' U.S. Fish and Wildlife Service. https://ecos.fws.gov/docs/tess/species_nonpublish/2560.pdf Retrieved Apr 9, 2022.


Habitat

Tooth Cave pseudoscorpions need to live in small, dark caves that have acidic groundwater that dissolve
calcium carbonate Calcium carbonate is a chemical compound with the chemical formula . It is a common substance found in Rock (geology), rocks as the minerals calcite and aragonite, most notably in chalk and limestone, eggshells, gastropod shells, shellfish skel ...
. The acidic groundwater is crucial for the survival of the Tooth Cave pseudoscorpions because it creates various types of geographical features, like caves, sinkholes, fractures, and interconnections with other caves. These features act as a natural barrier to the outside world. This barrier ensures that the
humidity Humidity is the concentration of water vapor present in the air. Water vapor, the gaseous state of water, is generally invisible to the human eye. Humidity indicates the likelihood for precipitation (meteorology), precipitation, dew, or fog t ...
levels remain relatively high and the temperature remains fairly stable. This is an important factor for the temperature-sensitive Tooth Cave pseudoscorpion. Karst environments also ensure that there is proper drainage of water from the cave to further maintain the prime humidity level. Each type of cave that the Tooth Cave pseudoscorpion lives in is unique. Thus, individual populations have adaptations to their specific environment. However, the Tooth Cave pseudoscorpion cannot adapt to climate change as fast as it is increasing temperatures.


Range

The Tooth Cave pseudoscorpion is only confirmed in two caves: Tooth Cave and Amber Cave, in Travis County, Texas. The caves span the Balcones Canyonlands, ranging from eastern to southeastern
Edwards Plateau The Edwards Plateau is a geographic region forming the crossroads of Central, South and West Texas, United States. It is named in honor of Haden Edwards. It is bounded by the Balcones Fault to the south and east; the Llano Uplift and the Lla ...
. Generally keeping to the caves, the Tooth Cave pseudoscorpions have a range of approximately 1 km from the caves before the environment becomes unsuitable for them to live. The caves are currently in peril of falling in due to the urbanization of the local communities, so the populations may shrink in the near future.


Conservation


Population size

The Tooth Cave pseudoscorpion population size is currently ranked as “H,” or “historical data.” The current information about population sizes and distribution is lacking, with too little research to classify the population in a different way. This could be due to many factors such as imprecise locations, disturbances to the population, and unstable habitats. There is also a lack of researchers currently studying the species.


Past and current geographical distribution

There is little to no data on the past and current distribution of the Tooth Cave pseudoscorpion. Due to a lack of research and study of the species, there is also little known about where they live. All that is confirmed is that they live in Travis County, Texas, in 2 caves, including Tooth Cave and Amber Cave, and are suspected to live in other nearby caves. However, the extent of their range is unknown.


Major threats

The threats to the Tooth Cave pseudoscorpion are mostly indirect. The largest and most imminent threat is human activity that destroys their habitat and food sources. The species was first listed as endangered on September 16, 1988, due to restricted distribution and threats from
urban development Urban means "related to a city". In that sense, the term may refer to: * Urban area, geographical area distinct from rural areas * Urban culture, the culture of towns and cities Urban may also refer to: General * Urban (name), a list of peop ...
. This development caused stressors to their habitat, including destruction,
degradation Degradation may refer to: Science * Degradation (geology), lowering of a fluvial surface by erosion * Degradation (telecommunications), of an electronic signal * Biodegradation of organic substances by living organisms * Environmental degradation ...
, and fragmentation. Travis County, Texas experienced an increase in human population between 1980 and 2017, rising from 419,573 to 1,226,698. In addition, single and multi-family housing units increased by 394% over the 46-year period between 1970 and 2016. With these population increases, came significant urban development. The karst cave environments where Tooth Cave pseudoscorpions live are next to residential neighborhoods. This puts them at an increased risk for
habitat degradation Habitat destruction (also termed habitat loss or habitat reduction) occurs when a natural habitat is no longer able to support its native species. The organisms once living there have either moved elsewhere, or are dead, leading to a decrease ...
and pollution. Karst habitats are specifically threatened by urbanization due to the dangers of cave infilling, drainage pattern changes, chemical contamination, and disruption of surface plant and animal distribution. Cave habitats are also threatened by mining and quarrying practices that decrease karst environments. The Tooth Cave pseudoscorpion is also threatened by invasive predators and competitors, especially the
red imported fire ant ''Solenopsis invicta'', the fire ant, or red imported fire ant (RIFA), is a species of ant native to South America. A member of the genus ''Fire ant, Solenopsis'' in the subfamily Myrmicinae, it was Species description, described by Swiss ento ...
. This ant species arrived in Travis County in the 1970s, and has been recorded in over 140 caves. The ants prey upon the Tooth Cave pseudoscorpion, but they also compete for the same food sources, especially cave crickets. Declines in cave cricket populations are also linked to declines in similar karst
invertebrate Invertebrates are animals that neither develop nor retain a vertebral column (commonly known as a ''spine'' or ''backbone''), which evolved from the notochord. It is a paraphyletic grouping including all animals excluding the chordata, chordate s ...
s to the Tooth Cave pseudoscorpion. However, direct correlations between the cave cricket and the Tooth Cave pseudoscorpion have not been studied. The tawny crazy ant species was first documented in Travis County in 2002, and is potentially a threat to the Tooth Cave pseudoscorpion. This invasive species has the potential to disrupt native ecosystems in a similar way as the
red imported fire ant ''Solenopsis invicta'', the fire ant, or red imported fire ant (RIFA), is a species of ant native to South America. A member of the genus ''Fire ant, Solenopsis'' in the subfamily Myrmicinae, it was Species description, described by Swiss ento ...
.''Recovery Plan Amendments for 20 Southwest Species'' (2019). U.S. Fish and Wildlife Service. Retrieved Apr 9, 2022. The Tooth Cave pseudoscorpion is also threatened by natural factors. Its limited distribution and reliance on a specific ecosystem make it vulnerable to population declines and decrease the likelihood of establishing new colonies. They are also extremely small and have little to no ability to move large distances on the surface. The fragility of cave ecosystems also affects their ability to thrive. Additionally, there is little to no regulation in Texas that protects the species. The Texas Commission on Environment Quality (TCEQ) does not include protection for karst habitats and the City of Austin’s Environmental Criteria Manual is not sufficient to protect cave cricket foraging areas. It also does not apply to all karst invertebrates.


Listing under the ESA

The Tooth Cave pseudoscorpion was listed under the
ESA The European Space Agency (ESA) is a 23-member international organization devoted to space exploration. With its headquarters in Paris and a staff of around 2,547 people globally as of 2023, ESA was founded in 1975 in the context of European ...
as
endangered An endangered species is a species that is very likely to become extinct in the near future, either worldwide or in a particular political jurisdiction. Endangered species may be at risk due to factors such as habitat loss, poaching, inv ...
on September 16, 1988, with a recovery plan formed in 1994. The listing cited human urbanization as a leading cause of the endangerment of the Tooth Cave pseudoscorpion. It is not currently listed on the IUCN Red List, as there is not sufficient data for the IUCN to categorize it.


Five-year review

A five-year review initiated in 2009 classified the recovery priority for the species as 2C. A recovery priority number of 2C means there is a high degree of threat and a high recovery potential. The C indicates that the species’ recovery conflicts with economic activity such as water demands and development projects. The next review in 2018 determined that while there was a recovery plan in place with the goal of down-listing the species, there were no delisting criteria identified. The review suggested that research on similar karst invertebrates could be applied to the Tooth Cave pseudoscorpion to develop delisting criteria. Furthermore, the five-year review found that no down-listing criteria have been met since 2009. The review identified two potential cave sites that could begin to meet delisting criteria, but there was insufficient information to determine whether the sites qualified.


Species status assessment

Due to a lack of research and study of the Tooth Cave pseudoscorpion, there were no updates to the biology and habitat, abundance, population trends, demographic features, demographic trends, genetics, variation,
taxonomic classification In biology, taxonomy () is the scientific study of naming, defining ( circumscribing) and classifying groups of biological organisms based on shared characteristics. Organisms are grouped into taxa (singular: taxon), and these groups are given ...
, or changes in
nomenclature Nomenclature (, ) is a system of names or terms, or the rules for forming these terms in a particular field of arts or sciences. (The theoretical field studying nomenclature is sometimes referred to as ''onymology'' or ''taxonymy'' ). The principl ...
in the most recent species status assessment. The 2018 five-year review agrees with the 2009 review on the distribution of the species: five caves with records of the species from four karst fauna regions in Travis County, Texas. There is no data specific to the Tooth Cave pseudoscorpion, although research on troglodytic arachnids suggests that they may travel through a network of underground voids. A study documents similarities between Tooth Cave spider populations in caves in Travis County and the caves the Tooth Cave pseudoscorpion is known to inhabit. This study implies that the spiders move between caves over time through underground openings. The species status assessment assumes that if the Tooth Cave pseudoscorpion also lives in connected caves, its population may be capable of traveling. There is not enough information to estimate population sizes or sustainable reproductive rates of the Tooth Cave pseudoscorpion. The assessment used measurements of surface habitats to estimate Tooth Cave pseudoscorpion population information. Using this data, it was suggested that agriculture within 50 meters of an inhabited cave significantly reduced the species richness of cave organisms. This is due to chemical contamination, the loss of surrounding native vegetation, and urbanization. Development projects were also suggested to be more harmful to cave ecosystems than agricultural projects. The 2018 review also used 2016 aerial photographs of surrounding caves to assess habitat elements. It found that there were only two caves with high to moderate resiliency that the species is known to live in.


Recovery plan

The recovery plan for the Tooth Cave pseudoscorpion aims to down-list the species from endangered to
threatened A threatened species is any species (including animals, plants and fungi) which is vulnerable to extinction in the near future. Species that are threatened are sometimes characterised by the population dynamics measure of ''critical depensatio ...
. The plan hopes to succeed through further research, protecting and managing habitats, creating educational programs, and monitoring all active programs. It was originally approved on August 25, 1994, and developed by Lisa O'Donnell, Ruth Stanford, and William Elliot of the Austin Texas US Fish and Wildlife Service. An amendment to the recovery plan was approved on August 28, 2019. The plan lists the following criteria to down-list the Tooth Cave pseudoscorpion from endangered to threatened: # At least one high-quality protected karst
fauna Fauna (: faunae or faunas) is all of the animal life present in a particular region or time. The corresponding terms for plants and fungi are ''flora'' and '' funga'', respectively. Flora, fauna, funga and other forms of life are collectively ...
area per karst fauna region # At least three total medium- or high-quality protected karst fauna areas per karst fauna region # A minimum of six protected karst fauna areas range-wide # A minimum of three high-quality karst fauna areas range-wide # All karst fauna areas are medium or high quality In the fall of 1988, governmental agency representatives in Austin, Texas made the Balcones Canyonlands Conservation Plan (BCCP). This plan allowed taking endangered species outside preservation areas. The Tooth Cave pseudoscorpion, among six other endangered cave species from Travis County, was included in this plan. It requires all refugia to include land large enough that populations of each species are protected. It also accommodates industrial and utility development in the area while preserving habitats. Two types of corridors were created to contain large parts of the inhabited land, but also to be used for other purposes. Primary corridors are existing corridors that would receive the majority of new development. Secondary corridors are existing corridors that would not need development. Planned corridors are unconstructed corridors that would be created if a habitat became endangered. The Endangered and Threatened Animals of Texas publication in 2003 lists many preservation actions for karst invertebrates that apply to the Tooth Cave pseudoscorpion. It suggests that recovery efforts focus on researching the
diversity Diversity, diversify, or diverse may refer to: Business *Diversity (business), the inclusion of people of different identities (ethnicity, gender, age) in the workforce *Diversity marketing, marketing communication targeting diverse customers * ...
of animals and plants in cave habitats. This would also help to control the red imported fire ant population that threatens the Tooth Cave pseudoscorpion. In December 2002, the Williamson County Karst Foundation created a long-term method to conserve the three endangered invertebrates in its county, including any possible populations of the Tooth Cave pseudoscorpion. Other private landowners and governmental agencies were included in this plan to help with conservation efforts. The publication also provides management guidelines for karst invertebrates. These include preserving the known cave sites, avoiding altering surface drainage patterns, preserving native vegetation, preventing groundwater contamination, restricting human visitation, and controlling exotic species (especially fire ants).


References

{{DEFAULTSORT:Tartarocreagris Texana Endemic fauna of Texas Neobisiidae Travis County, Texas Cave arachnids Animals described in 1969 Taxonomy articles created by Polbot