Nothomyrmecia

Nothomyrmecia
N. macrops workers
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Formicidae
Subfamily: Myrmeciinae
Tribe: Prionomyrmecini
Genus: Nothomyrmecia
Clark, 1934
Species: N. macrops
Binomial name
Nothomyrmecia macrops
Clark, 1934

Nothomyrmecia, also known as the dinosaur ant or dawn ant, is a rare genus of ants consisting of a single species, Nothomyrmecia macrops. A member of the subfamily Myrmeciinae, two workers were presumably collected in 1931 near the Russell Range from Israelite Bay in Western Australia, first described by Australian entomologist John S. Clark in 1934. The ant lives in South Australia, nesting in old-growth mallee woodland and Eucalyptus woodland. The ant only remained in literature and from two specimens until a group of entomologists rediscovered it 46 years later, 1,300 kilometres (810 mi) away from the original site. Due to the species' restricted distribution and potential threats to its survival, the IUCN lists the ant as Critically Endangered.

Some entomologists have suggested a relationship to the Baltic Eocene fossil ant genus Prionomyrmex based on morphological similarities, but this interpretation is not widely accepted by the entomological community. The ant's status as a so-called 'living fossil', owing to its primitive body structure, has stimulated studies on its morphology, behaviour, ecology, and chromosomes.

The average length of this species is 9.7–11 mm (0.38–0.43 in), and polymorphism does not occur among the castes. A queen will mate with one or more males, and during colony foundation she will hunt for food until the brood have fully developed. Queens are univoltine and only produce a single generation of ants annually. Two queens may found a colony together, but only one will remain once the first generation of workers arrives. Mature colonies are very small, with only 50 to 100 individuals in each nest. Workers are strictly nocturnal and only forage during cold nights, collecting arthropod prey and sweet substances such as honeydew from Hemiptera insects. They are solitary foragers and rely on their vision to navigate, and no evidence suggests that they use chemicals to communicate. Potential predatory threats are restricted to other ants.

Taxonomy

Nothomyrmecia was originally discovered by Amy Crocker[lower-alpha 1] in December 1931, from two syntype workers collected near the Russell Range from Israelite Bay in Western Australia.[2][3] However, entomologist Robert W. Taylor expressed doubt about the original discovery site of the ant, stating the specimens were probably collected from the western end of the Great Australian Bight, south from Balladonia.[3] It is unknown where Crocker collected the insects she sent to Australian entomologist John S. Clark, but the locations Clark provided had no Nothomyrmecia colonies present when the areas were surveyed. Despite searches, a 2012 report discussing the possible abundance of Nothomyrmecia in Western Australia did not confirm any sighting of the ant between Balladonia and the Western Australian coastal regions.[4] After Australian and American collectors had failed to find Nothomyrmecia for decades, Taylor and his party of entomologists from Canberra found a solitary worker in 1977 at Poochera, southeast of Ceduna, 1,300 kilometres (810 mi) from the site of the 1931 discovery.[3] Currently, Poochera is probably the only town in the world with ant-based tourism, where they have pictures stenciled on the streets.[5] Entomologists dub the ant as the "Holy Grail" of myrmecology after 46 years of searching for it.[6][7]

N. macrops worker specimen collected by Taylor

John S. Clark described Nothomyrmecia in 1934 as a new genus of Myrmeciinae.[2] He did so because the two syntypes bore no resemblance to any ants he knew of, although the extinct genus Prionomyrmex shared similar morphological characteristics with Nothomyrmecia. Clark notes that the head and mandibles of Nothomyrmecia and Prionomyrmex are somewhat similar, but the two can be distinguished by the appearance of the node (a segment between the mesosoma and gaster).[2] The type material is currently housed in the National Museum of Victoria in Melbourne. In 1944, British myrmecologist Horace Donisthorpe moved Nothomyrmecia to the subfamily Ponerinae, suggesting that all bulldog ants are Ponerine.[8] In 1951, Clark proposed the new ant subfamily Nothomyrmeciinae for his Nothomyrmecia, based on morphological differences with the other Myrmeciinae.[9] With this proposal, Clark assigned Nothomyrmecia to the newly established tribe Nothomyrmeciini. Its distant relationship with modern ants was confirmed after its rediscovery, and its placement within the Formicidae was universally accepted by scientists until 2000.[3] In 1985, American entomologists George Wheeler and Jeanette Wheeler placed Nothomyrmecia back into Myrmeciinae, but a 1988 publication transferred it back into its own subfamily.[10][11] The single waist node and other features such as a non-tubulated fourth abdominal segment led scientists to believe that Nothomyrmecia should be separate from Myrmecia and retain Clark's original proposal. This proposal would classify the ant into its own subfamily, despite many familiar morphological characteristics between the two genera.[12][13]

In 2000, Cesare Baroni Urbani described a new Baltic fossil Prionomyrmex species. After examining specimens of Nothomyrmecia, Baroni Urbani stated that his new species and N. macrops belonged to the same genus, and the name Prionomyrmex would replace the name Nothomyrmecia and the subfamily Nothomyrmeciinae must be called Prionomyrmeciinae.[14] Baroni Urbani states that Prionomyrmex is a paraphyletic relative to Nothomyrmecia, and the two genera were so morphologically similar that Nothomyrmecia must be synonymised. In 2003, Dlussky & Perfilieva separated Nothomyrmecia from Prionomyrmex on the basis of the fusion of an abdominal segment.[15] In the same year Ward & Brady reached the same conclusion as Dlussky & Perfileeva and provided strong support for the monophyly of Prionomyrmex, and, in addition, transferred both taxa as distinct genera in the older subfamily Myrmeciinae.[15][16] Later, Dlussky also refers only to Ward & Brady's classification.[16][17] In 2005 an 2008, Baroni Urbani suggested additional evidence in favour of his former interpretation as opposed to that of Ward & Brady's arguments.[18][19] This view, however, is not supported in subsequent relevant papers, which continue to use the classification of Ward & Brady, rejecting that of Baroni Urbani.[17][20][21][22]

The ant is commonly known as the dinosaur ant, dawn ant, or living fossil ant because of its primitive body structure.[20][23][24] Its specific epithet, macrops ("big eyes"), is derived from the Greek words makros "long, large" and ops "eyes".[24][25][26] The generic name Nothomyrmecia means "false bulldog ant".[24]

Nothomyrmecia and a Ponerinae ant, Platythyrea tricuspidata, share the highest number of chromosomes within Hymenoptera, with a diploid chromosome number 92–94.[27] Studies show that all Hymenoptera insects with a 2n above 52 are ants. Nothomyrmecia ants have dot-like acrocentric chromosomes, but metacentric chromosomes are sometimes present.[3]

Genetic evidence suggests that the age of the most recent common ancestor for Nothomyrmecia and Myrmecia is 74 million years old, giving a likely origin in the Cretaceous.[16] There are two hypotheses of the internal phylogeny of Nothomyrmecia: The subfamily Formicinae is more closely related to Nothomyrmecia than it is to Myrmecia, evolving from Nothomyrmecia-like ancestors. However, Nothomyrmecia and Aneuretinae may have shared a common ancestor; the two most likely separated from each other, and the first Formicinae ants evolved from the Aneuretinae instead. Currently, scientists agree that Nothomyrmecia most likely evolved from ancestors to the Ponerinae.[28] In the Evans' vespoid scale, Nothomyrmecia and other primitive ant genera such as Amblyopone and Myrmecia exhibit behaviour similar to a clade of soil-dwelling families of vespoid wasps.[29] The following cladogram generated by Archibald and colleagues shows the possible phylogenetic position of Nothomyrmecia among some ants of the subfamily Myrmeciinae. They suggest that Nothomyrmecia may be closely related to extinct Myrmeciinae ants such as Avitomyrmex, Macabeemyrma, Prionomyrmex and Ypresiomyrma.[30]

Label view of Nothomyrmecia collected by Taylor
Myrmeciinae

Archimyrmex

Myrmecia


Prionomyrmex janzeni

Prionomyrmex longiceps

Macabeemyrma

Avitomyrmex

Ypresiomyrma

Nothomyrmecia





Description

Closeup view of a worker's head

Nothomyrmecia is a medium-sized ant, measuring 9.7–11 mm (0.38–0.43 in), and polymorphism does not occur among any caste.[2][31] The mandibles, clypeus, antennae and legs are pale yellow. The hair is yellow, erect and reasonably long and common, but on the antennae and legs it is suberect (standing almost in an erect position) and shorter. The pubescence is white and abundant. While it shows similar characteristics with Myrmecia, Nothomyrmecia somewhat resembles Oecophylla weaver ants. Workers are nocturnal but navigate by vision, relying on large compound eyes.[32] Mandibles are less specialised than Myrmecia and Prionomyrmex, elongated and triangular. These mandibles are shorter than the head with 10 to 12 teeth. The head is longer than its width and broader at the back. It is broadest around the eyes with convex sides and a concave occipital border. The antennal scapes extend beyond the occipital border, and the second segment of the funiculus is slightly longer than the first, third and fourth segment. The node, pronotum, epinotum and thorax are longer than broad, and the mesonotum is just as long. The first segment of the gaster is broader than long by a third and broader at the back than the front with strongly convex sides.[2]

There is a retractable stinger with the furculae (attachment sites for the muscles which help protrude the stinger) present at the back of the abdomen.[3] A sting bulb gland is only known in Nothomyrmecia and Myrmecia, an exocrine gland with an unknown purpose. The location of the gland is between the Dufour's gland and the ducts of venom,[33] and there are more derived characters of the sting apparatus in Nothomyrmecia than Myrmecia.[34] It is the only known species of ant that contains a sting and a waist that has no postpetiole between the first and second gastral segments.[24]

Nothomyrmecia macrops male specimen

Queens look similar to workers, but several morphological features distinguish the two castes from each other. Although their body structure is similar to that of workers, queens are usually larger. Ocelli are highly developed, but the eyes on the queen are not enlarged. The structure of the pterothorax (the wing-bearing area of the thorax) is consistent with other reproductive ants, although it does not occupy as much of its mesosomal bulk. The queens are described as brachypterous ants: their wings are reduced to the point that they are not functional.[3][35] Their wings are noticeably rudimentary and stubby, barely overlapping the first gastral segment. Males resemble those of Myrmecia, but Nothomyrmecia males bear a single waist node. Exhibiting a primitive venational complement, the wings on a male are not stubby like a queen's; rather they are long and fully developed. They have a jugal anal lobe (a portion of the hindwing), a feature found in many primitive ants, but the basal hamuli and apical set present are only known in Nothomyrmecia ants. Most male specimens collected have two tibial spurs; the first is a long calcar and the second is short and thick. In both the queen and worker caste, males are also known to have a ventral stridulatory organ.[3]

In all castes, these ants have six maxillary and four labial palpae, a highly primitive feature. The females of this species have a 12 segmented antenna, whereas males have 13 segments. Other features include paired calcariae found on both the hind and middle tibiae, and the claws have a median tooth. The unspecialised nature of the cuticle is similar to Pseudomyrmex, a member of the subfamily Pseudomyrmecinae. Many of the features known in Nothomyrmecia are found in Ponerinae and Pseudomyrmecinae.[3]

The eggs of Nothomyrmecia are similar to that of Myrmecia, appearing as non-adhesive and subspherical. The larvae bear a primitive body structure with no specialised tubercles, sharing similar characteristics with the subfamily Ponerinae. However, the sensilla are more abundant on the mouthparts. The larvae are characterised into three stages: very young, young, and mature, measuring 2.8 mm (0.1 in), 6.3 mm (0.2 in) and 11 mm (0.4 in), respectively.[36] The cocoons have thin walls and produce meconium (a metabolic waste product expelled through the anal opening after an insect emerges from its pupal stage).[3] The cuticular hydrocarbons have internally branched alkenes, a feature rarely found in ants and most insects.[37]

In general, the body structure of all Nothomyrmecia castes exhibits the primitive nature of the species.[3] Notable derived features include vestigal ocelli on workers, brachypterous queens, and the mesoscutal structure on males. The morphology of the abdomen, mandibles, gonoforceps (an area apart of the external male genitalia) and the basal hamuli shows it is more primitive than Myrmecia. The structure of the abdominal region can separate it from other Myrmeciinae relatives (the fourth abdominal segment of Myrmecia is tubulate, whereas Nothomyrmecia has a non-tubulated abdominal segment). The appearance of the fourth abdominal segment is consistent with almost all aculeate insects, and possibly Sphecomyrma.[3]

Distribution and habitat

Reported occurrences of Nothomyrmecia according to the Atlas of Living Australia

Nothomyrmecia is present in the cool regions of South Australia and possibly Western Australia in mallee woodland and old-growth areas populated with various Eucalyptus species, including Eucalyptus brachycalyx, Eucalyptus gracilis and Eucalyptus oleosa.[38][39] It is unknown how widespread Nothomyrmecia is, although the ant may be more abundant if it is heavily associated with mallee woodland. Recently, however, Nothomyrmecia colonies were located in 18 areas along the Eyre Peninsula, a stretch of 400 kilometres (248 miles).[38][40] Nothomyrmecia ants colonise areas at elevations of 85 metres (278.9 ft).[31] Nests are found in soil and degraded limestone soil with Callitris trees present.[39][41] Colony construction only occurs when the soil is moist.[3] Nest holes are difficult to identify, as they are only 4–6 mm (0.16–0.24 in) wide holes under shallow leaf litter with no mounds or deposited soil present, though guards are regularly seen. Nest galleries are 4–5 mm (0.16–0.20 in) in diameter that descend further into the ground at about 60°, ending up to subelliptical and horizontal chambers 3–5 mm (0.12–0.20 in) wide.[3] These chambers are typically 18 to 43 centimetres (7.1 to 16.9 in) below the earth's surface.

Behaviour and ecology

Workers are nectarivores and can be found foraging on top of Eucalyptus trees, where they search for food and prey.[42] They search for prey in piles of leaves, killing small arthropods including Drosophila flies, microlepidopterans and spiderlings. Prey items are usually 4 millimetres (0.2 in) or less, and workers grab them using their mandibles and forelongs, stinging them in the process.[12][43] Workers also feed on sweet substances such as honey coated on trees and tend to Hemiptera insects for honeydew; one worker alone may feed on these sources for 30 minutes.[3][44] Despite being nectarivores, workers are known to drink hemolymph from the insects they capture, and a queen in a captive colony was observed consuming a fly.[3][31] Captured prey items are given to larvae, which are carnivorous.[3] Pupae may be given to larvae if a colony has a shortage of food. Workers lay trophic eggs used to feed the larvae, but sometimes the adults, including the queen and other sexual reproductive ants, consume them. Trophallaxis occurs among adults, the alates and larvae, with the workers exuding anal droplets to their nestmates.[3]

Workers are strictly nocturnal ants only active during cold nights; they do not emerge from their nests during the day.[31][40] Workers are most active at temperatures of 5–10 °C (41–50 °F), and they are more difficult to locate on warmer nights. The ability to withstand such cold temperatures suggests that Nothomyrmecia encounters very few competitors, including dominant diurnal ants that are sometimes found foraging during warm nights.[43] Unless a forager has captured prey, workers stay on trees for the remainder of the night until dawn, possibly relying on sunlight to navigate back to their nest.[43] There is no evidence that they use chemicals to communicate; instead, workers rely on visual cues to navigate around. The ants are solitary foragers.[43]

Nothomyrmecia workers from other colonies are not antagonistic towards each other, so they can forage on a single tree, although they attack if an outsider tries to enter a colony.[43] Ants such as Camponotus and Iridomyrmex may pose a threat to foragers or to a colony if they try to enter; Nothomyrmecia workers counter this by secreting alarm pheromones from the mandibular gland and Dufour's gland.[43] Foraging workers also engage in alternative methods to protect themselves from predators. Adopting a posture by opening the jaws in a threatening stance or deliberately falling onto the ground and remaining motionless until the threat subsides are two known methods.[3] With that said, Nothomyrmecia is a timid and shy species that retreats if exposed.

Two queens showing their brachypterous wings, as well as a worker and several cocoons

Nuptial flight does not occur in Nothomyrmecia. Instead they engage in long-range dispersal which presumably begins by late summer or autumn, with alates emerging around March and April, but sometimes a colony may be overwintered. These alates, born around January, are usually young when they begin to mate. Queens are seen around vegetation trying to flutter their wings, a behaviour seen in some brachypterous Myrmecia queens.[3][43] Due to the brachypterous nature of the queens, it is likely that the alates mate near their parent nest and release sex pheromones, or instead climb on vegetation far away from their nests and attract fully winged males.[43][45][46] Nothomyrmecia is a polyandrous species, in which queens mate with one or more males (an average of around 1.37) in a single mating season.[47] After mating, colonies can be found by one or more queens, making them pleometrotic. However, a colony with two queens reduces to a single queen when the nest is mature, forming colonies that are monogynous.[48][49] The queens will contest for dominance, but the subordinate queen is later expelled by workers who drag her outside the nest.[43] An existing nest with no queen may adopt a foraging queen looking for an area to begin her colony, as well as workers. Queens are semi-claustral and can be seen among workers searching for food, meaning that during the founding of the new colony, the queen has to forage so that she has sufficient food to raise her brood. Sometimes a queen will leave her nest at night with the sole purpose of finding food or water for herself.[3]

Eggs are not seen in nests from April to September, and foraging activity greatly reduces and nests are sealed to avoid overwintering. In one tested nest, eggs laid by late December hatched by mid-February, although pupation did not occur until March. However, Nothomyrmecia is univoltine and only produces a single generation of eggs per season, and it sometimes may take as many as 12 months for an egg to develop into an adult. Adults are either defined as juveniles or post-juveniles: juveniles are too young (perhaps several months old) to have experienced overwintering whereas post-juveniles have. The pupae are generally overwintered and begin to hatch by the time a new generation of eggs is laid.[31] Workers may be capable of laying reproductive eggs, although it is not known if these eggs give rise to males and fertile females.[50] It is also unknown whether workers are gamergates, but some nests possibly have inseminated workers that act as supplementary reproductives.[43] Eggs are scattered among the nest, whereas the larvae and pupae are set apart from each other in groups. The larvae are capable of crawling around the nest, swelling up dramatically and later buried in the ground to spin their cocoons. Small non-aggressive workers that act as nurses provide assistance for newborns to hatch from their cocoons. At maturity, a nest may only contain 50 to 100 adults.[3][42][51] In some nests, colony founding can occur within a colony itself: when a queen dies, the colony may be taken over by one of her daughters, or it may adopt a newly mated queen, restricting reproduction among workers; this method of founding extends the lifespan of the colony.[52][53] Nothomyrmecia can recognise brood of their own by licking larvae, which also enables them to identify an alien conspecific that is transferring its colony odour to the larvae.[12]

Reasons of brachyptery among queens are possibly due to population structure or ecological requirements within their geographical range. Brachyptery may have only evolved in this species quite recently, as the wings would have disappeared if they had no function for dispersal. Another plausible scenario is that brachyptery only forms in drought-stressed colonies; this case has occurred in several Monomorium colonies found throughout semiarid regions of Australia. However, there is insufficient evidence to confirm such a scenario.[3]

Workers and queens groom each other, especially around the posterior area of the mesosoma but not near the abdominal region. When an adult carries another nestmate or colonies are migrating, it takes on a pupal position until the ant releases its nestmate. Both queens and workers may be dragged by others, and males are seen riding on queens without trying to copulate. Waste material such as dead nestmates, cocoon shells, and food remnants are disposed of far away from the nest.[3]

Conservation

Before its rediscovery, entomologists feared that Nothomyrmecia had become extinct. Notable biologists such as E. O. Wilson and William Brown, Jr., made attempts to search for the ant, but neither was successful.[54] After its rediscovery in 1977, the International Union for Conservation of Nature listed Nothomyrmecia as Critically Endangered in 1996, stating that only a few small colonies were known,[1] and it is a protected species under the Western Australian Wildlife Conservation Act 1950.[38] However, the Threatened Species Scientific Committee states that the species is ineligible for listing under the Environment Protection and Biodiversity Conservation Act 1999. No evidence confirms that populations are declining with colonies naturally depauperate (lacking in numbers), and their distribution is potentially extensive due to the ants' habitat preference, living in old growth mallee woodland; it is possible that it could be widespread in areas with mallee vegetation across southern Australia.[38][55] With this said, it is unknown how widespread it is, and scientists are unsure if any threats are impacting the species.

Although threats are not known for sure, suspected anthropogenic threats that can significantly impact Nothomyrmecia include habitat destruction and fragmentation by railway lines, roads and wheat fields.[38][56] In the Ceduna area, local populations of the ant were almost eliminated after the area was bulldozed and burned during the installation of an underground telephone line, although nearby sites had a reasonable population, even larger than the destroyed site. Colonies may not survive tree-clearing, as they depend on overhead canopies to navigate.[56] Bushfires are another major threat to the survival of Nothomyrmecia, potentially destroying valuable food sources, including the trees they forage on, and reducing the population of a colony. While these ants may have recovered from previous bushfires, larger, more frequent fires may devastate the population. However, Nothomyrmecia ants can be safe from fires if they remain inside their nests.[23][56] Nothomyrmecia is also dependent on cold temperatures to forage and collect food, so climate change is a threat to its survival. An increase in the temperature will prevent workers from foraging, and very few areas would be suitable for the species to live in. An increase in sea temperature would also threaten Nothomyrmecia.

Measures needed to ensure the survival of Nothomyrmecia include fighting climate change, conducting surveys, and maintaining known populations through habitat protection.[23][24] Conservationists suggest that its remaining habitat should be fully protected from degradation, and further improvement is needed, although most known populations are found outside protected areas in vegetation alongside roads. Local councils should also be aware of the conservation status of Nothomyrmecia, so future land use can be decided more appropriately at a local level. Not all colonies are found in unprotected areas, as some have been found in the Lake Gilles Conservation Park and the Chadinga Conservation Reserve. Other colonies have been found on private property, where the owner of the property intends to conserve them. More research is needed to know the true extent of the ant's geographical distribution.[24]

Notes

  1. ↑ Referred to as Miss. A. E. Baesjou in Clark's article.

References

  1. 1 2 Social Insects Specialist Group (1996). "Nothomyrmecia macrops". IUCN Red List of Threatened Species (IUCN) 1996: e.T14849A4465171. doi:10.2305/IUCN.UK.1996.RLTS.T14849A4465171.en.
  2. 1 2 3 4 5 Clark, J.S. (1934). "Notes on Australian ants, with descriptions of new species and a new genus" (PDF). Memoirs of the National Museum of Victoria 8: 5–20. doi:10.5281/zenodo.26629.
  3. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 Taylor, R.W. (1977). "Nothomyrmecia macrops: a living-fossil ant rediscovered". Science 201 (4360): 979–985. Bibcode:1978Sci...201..979T. doi:10.1126/science.201.4360.979. JSTOR 1746819. PMID 17743619.
  4. ↑ McArthur, A. (2012). "Report on a search for the dinosaur ant in Western Australia". South Australian Naturalist 86 (2): 89–92. ISSN 0038-2965.
  5. ↑ "Biodiversity". State of the Environment, South Australia. Environment Protection Authority. Retrieved 23 November 2015.
  6. ↑ "CSIRO team finds world's most primitive ant.". The Canberra Times (Canberra, ACT: National Library of Australia). 18 November 1977. p. 3. Retrieved 22 November 2015.
  7. ↑ "'Dinosaur' that still lives.". The Canberra Times (Canberra, ACT: National Library of Australia). 30 June 1982. p. 18. Retrieved 22 November 2015.
  8. ↑ Donisthorpe, H. (1944). "Notes arising out of Mr. W. D. Hincks's review of Mr. Donisthorpe's 'A list of the type-species of the genera and subgenera of Formicidae'". Entomologists Monthly Magazine 80: 59. doi:10.5281/zenodo.26492.
  9. ↑ Clark, J.S. (1951). The Formicidae of Australia (Volume 1). Subfamily Myrmeciinae. (PDF). Melbourne: Commonwealth Scientific and Industrial Research Organisation, Australia. p. 16.
  10. ↑ Wheeler, G.C.; Wheeler, J. (1985). "A simplified conspectus of the Formicidae" (PDF). Transactions of the American Entomological Society 111: 255–264. doi:10.5281/zenodo.25105.
  11. ↑ Dlussky, G.M.; Fedoseeva, E.B. (1988). The origin and early stages of evolution of ants (Hymenoptera: Formicidae in The Creaceous biocenotic crisis and the evolution of insects. Moscow: Nauka. pp. 70–144.
  12. 1 2 3 Jaisson, P.; Fresneau, D.; Taylor, R. W.; Lenoir, A. (1992). "Social organization in some primitive Australian ants. I. Nothomyrmecia macrops Clark" (PDF). Insectes Sociaux 39 (4): 425–438. doi:10.1007/BF01240625.
  13. ↑ Bolton, B. (1990). "Army ants reassessed: the phylogeny and classification of the doryline section (Hymenoptera, Formicidae)". Journal of Natural History 24 (6): 1339–1364. doi:10.1080/00222939000770811.
  14. ↑ Baroni Urbani, C. (2000). "Rediscovery of the Baltic amber ant genus Prionomyrmex (Hymenoptera, Formicidae) and its taxonomic consequences". Eclogae geologicae Helveticae 93 (3): 471–480.
  15. 1 2 Dlussky, G.M.; Perfilieva, K.S. (2003). "Paleogene ants of the genus Archimyrmex Cockerell, 1923 (Hymenoptera, Formicidae, Myrmeciinae)" (PDF). Paleontological Journal 37 (1): 39–47.
  16. 1 2 3 Ward, P.S.; Brady, S.G. (2003). "Phylogeny and biogeography of the ant subfamily Myrmeciinae (Hymenoptera : Formicidae)" (PDF). Invertebrate Systematics 17 (3): 361–386. doi:10.1071/IS02046.
  17. 1 2 Dlussky, G. M. (2012). "New fossil ants of the subfamily Myrmeciinae (Hymenoptera, Formicidae) from Germany". Paleontological Journal 46: 288–292. doi:10.1134/s0031030111050054.
  18. ↑ Baroni Urbani, C. (2005). "Phylogeny and biogeography of the ant subfamily Prionomyrmecinae (Hymenoptera: Formicidae)" (PDF). Annali del Museo Civico di Storia Naturale di Genova 96: 581–595.
  19. ↑ Baroni Urbani, C. (2008). "Orthotaxonomy and parataxonomy of true and presumed bulldog ants (Hymenoptera, Formicidae)" (PDF). Doriana (suppl. to Annali del Museo civico di storia naturale Giacomo Doria) 8 (358): 1–10. ISSN 0417-9927.
  20. 1 2 Wilson, E. O.; Hölldobler, B. (2005). "The rise of the ants: A phylogenetic and ecological explanation". Proceedings of the National Academy of Sciences 102 (21): 7411–7414. Bibcode:2005PNAS..102.7411W. doi:10.1073/pnas.0502264102. PMC 1140440. PMID 15899976.
  21. ↑ Ward, P.S. (2007). "Phylogeny, classification, and species-level taxonomy of ants (Hymenoptera: Formicidae)". Zootaxa 1668: 549–563.
  22. ↑ Moreau, C.S.; Bell, C.D.; Vila, R.; Archibald, S.B.; Pierce, N.E. (2006). "Phylogeny of the ants: diversification in the age of angiosperms". Science 312 (5770): 101–104. Bibcode:2006Sci...312..101M. doi:10.1126/science.1124891. PMID 16601190.
  23. 1 2 3 Taylor, R.W. (9 January 2014). "Australian endangered species: Dinosaur Ant". The Conversation. Retrieved 3 August 2015.
  24. 1 2 3 4 5 6 Clarke, G.M.; Spier-Ashcroft, F. "A Review of the Conservation Status of Selected: Australian Non-Marine Invertebrates" (PDF). Natural Heritage Trust. Government of Australia. pp. 81–83. Retrieved 6 December 2015.
  25. ↑ Harper, Douglas. "Macro-". Online Etymology Dictionary.
  26. ↑ Senning, A. (2006). Elsevier's dictionary of chemoetymology the whies and whences of chemical nomenclature and terminology (1st ed.). Amsterdam: Elsevier. p. 283. ISBN 978-0-08-048881-3.
  27. ↑ Imai, H.T.; Taylor, R.W.; Kubota, M.; Ogata, K.; Wada, M.Y. (1990). "Notes on the Remarkable Karyology of the Primitive Ant Nothomyrmecia macrops, and of the Related Genus Myrmecia (Hymenoptera: Formicidae)" (PDF). Psyche: A Journal of Entomology 97 (3–4): 133–140. doi:10.1155/1990/91237.
  28. ↑ Veeresh, G.K.; Mallik, B.; Viraktamath, C.A. (1990). Social insects and the environment. Leiden: E.J. Brill. p. 315. ISBN 978-90-04-09316-4.
  29. ↑ Hölldobler & Wilson 1990, p. 27.
  30. ↑ Archibald, S.B.; Cover, S. P.; Moreau, C. S. (2006). "Bulldog Ants of the Eocene Okanagan Highlands and History of the Subfamily (Hymenoptera: Formicidae: Myrmeciinae)" (PDF). Annals of the Entomological Society of America 99 (3): 487–523. doi:10.1603/0013-8746(2006)99[487:BAOTEO]2.0.CO;2.
  31. 1 2 3 4 5 Taylor, R.W. (2014). "Evidence for the Absence of Worker Behavioral Subcastes in the Sociobiologically Primitive Australian Ant Nothomyrmecia macrops Clark (Hymenoptera: Formicidae: Myrmeciinae)". Psyche: A Journal of Entomology 2014 (4707): 1–7. doi:10.1155/2014/232057.
  32. ↑ Brown, W.B.; Wilson, E.O. (1959). "The search for Nothomyrmecia" (PDF). Western Australian Naturalist 7 (2): 25–30.
  33. ↑ Billen, J. (1990). "The sting bulb gland in Myrmecia and Nothomyrmecia (Hymenoptera : Formicidae): new exocrine gland in ants" (PDF). International Journal of Insect Morphology and Embryology 19 (2): 133–139. doi:10.1016/0020-7322(90)90023-I.
  34. ↑ Kugler, C. (1980). "The sting apparatus in the primitive ants Nothomyrmecia and Myrmecia". Australian Journal of Entomology 19 (4): 263–267. doi:10.1111/j.1440-6055.1980.tb00983.x.
  35. ↑ Hölldobler & Wilson 1990, p. 302.
  36. ↑ Wheeler, G.C.; Wheeler, J.; Taylor, R.W. (1980). "The larval and egg stages of the primitive ant Nothomyrmecia macrops Clark (Hymenoptera: Formicidae)". Australian Journal of Entomology 19 (2): 131–137. doi:10.1111/j.1440-6055.1980.tb00974.x.
  37. ↑ Brown, W.V.; Jaisson, P.; Taylor, R.W.; Lacey, M.J. (1990). "Novel internally branched, internal alkenes as major components of the cuticular hydrocarbons of the primitive australian ant Nothomyrmecia macrops Clark (Hymenoptera: Formicidae)". Journal of Chemical Ecology 16 (9): 2623–2635. doi:10.1007/BF00988074.
  38. 1 2 3 4 5 Threatened Species Scientific Committee. "Dinosaur Ant, Fossil Ant (Nothomyrmecia macrops)". Department of Sustainability, Environment, Water, Population and Communities. Government of Australia. Retrieved 15 August 2010.
  39. 1 2 "Species: Nothomyrmecia macrops". AntWeb. The California Academy of Sciences. Retrieved 3 April 2015.
  40. 1 2 Foster, R. (1998). "Notes on the distribution of the dinosaur ant Nothomyrmecia macrops Clark (Hymenoptera: Formicidae) in South Australia". Australian Entomologist 25 (1): 29–31. ISSN 1320-6133.
  41. ↑ Taylor, R. W.; Brown, D.R.; Cardale, J.C. (1985). Hymenoptera, Formicoidea, Vespoidea and Sphecoidea. Zoological catalogue of Australia 2. Canberra: Australian Government Publishing Service. p. 6. ISBN 978-0-644-03922-2.
  42. 1 2 Choe, Jae C.; Crespi, Bernard J. (1997). The evolution of social behavior in insects and arachnids (1st ed.). Cambridge: Cambridge University Press. pp. 377–378. ISBN 978-0-521-58977-2.
  43. 1 2 3 4 5 6 7 8 9 10 Hölldobler, B.; Taylor, R. W. (1983). "A behavioral study of the primitive ant Nothomyrmecia macrops Clark". Insectes Sociaux 30 (4): 384–401. doi:10.1007/BF02223970.
  44. ↑ Shattuck, S.O.; Barnett, N. (June 2010). "Nothomyrmecia Clark, 1934". Ants Down Under. CSIRO Entomology. Archived from the original on 21 November 2015. Retrieved 21 November 2015.
  45. ↑ Ward, P.S.; Taylor, R.W. (1981). "Allozyme variation, colony structure and genetic relatedness in the primitive ant Nothomyrmecia macrops Clark (Hymenoptera: Formicidae)". Australian Journal of Entomology 20 (3): 177–183. doi:10.1111/j.1440-6055.1981.tb01026.x.
  46. ↑ Hölldobler & Wilson 1990, pp. 145–146.
  47. ↑ Sanetra, M.; Crozier, R. H. (2001). "Polyandry and colony genetic structure in the primitive ant Nothomyrmecia macrops". Journal of Evolutionary Biology 14 (3): 368–378. doi:10.1046/j.1420-9101.2001.00294.x.
  48. ↑ Hölldobler & Wilson 1990, p. 187.
  49. ↑ Hölldobler & Wilson 1990, p. 217.
  50. ↑ Trager, J.C. (1988). Advances in myrmecology (1st ed.). Leiden, Netherlands: E.J. Brill. p. 164. ISBN 978-0-916846-38-1.
  51. ↑ Hölldobler & Wilson 1990, p. 161.
  52. ↑ Sanetra, M.; Crozier, R.H. (2002). "Daughters inherit colonies from mothers in the 'living-fossil' ant Nothomyrmecia macrops". Die Naturwissenschaften 89 (2): 71–74. doi:10.1007/s00114-001-0288-5. PMID 12046624.
  53. ↑ Dietemann, Vincent; Peeters, Christian; Hölldobler, Bert (2004). "Gamergates in the Australian ant subfamily Myrmeciinae". Naturwissenschaften 91 (9): 432–435. Bibcode:2004NW.....91..432D. doi:10.1007/s00114-004-0549-1. PMID 15278223.
  54. ↑ Hoyt, E. (1997). The earth dwellers: adventures in the land of ants (1st ed.). New York: Touchstone. p. 108. ISBN 978-0-684-83045-2.
  55. ↑ Sanetra, M.; Crozier, R. H. (2003). "Patterns of population subdivision and gene flow in the ant Nothomyrmecia macrops reflected in microsatellite and mitochondrial DNA markers". Molecular Ecology 12 (9): 2281–2295. doi:10.1046/j.1365-294X.2003.01900.x. PMID 12919468.
  56. 1 2 3 Michael Pyle, R.; Mark Collins, N. (1983). The IUCN invertebrate red data book. Michigan: IUCN. p. 508.

Cited literature

External links

External identifiers for Nothomyrmecia
Encyclopedia of Life 620300
ITIS 582824
NCBI 147889
This article is issued from Wikipedia - version of the Saturday, January 30, 2016. The text is available under the Creative Commons Attribution/Share Alike but additional terms may apply for the media files.