Ctenophorus caudicinctus

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Ctenophorus caudicinctus
Ring-tailed Dragon (Ctenophorus caudicinctus) (8851937562).jpg
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Order: Squamata
Suborder: Iguania
Family: Agamidae
Genus: Ctenophorus
Species:
C. caudicinctus
Binomial name
Ctenophorus caudicinctus
Synonyms [1] [2] [3]
  • Amphibolurus caudicinctus
    (Günther, 1875)
  • Grammatophora caudicincta
    (Günther, 1875)
  • Tachyon caudicinctus
    (Günther, 1875)
  • Amphibolurus imbricatus
    (Peters, 1876)

Ctenophorus caudicinctus, commonly known as the ring-tailed dragon or ring-tailed bicycle-dragon is a native species of agamid lizard occurring in rocky ranges and outcrops of Australia. [4] [5] Ctenophorus caudicinctus is most commonly found in the Pilbara region and offshore islands of Western Australia. [6] The ctenophorus has 28 known species in the northern, southern, and western parts of Australia. It is recognized to be the most speciose group of Australian agamids. [7]

Contents

Description

Ctenophorus caudicinctus are recognized for their bright colours, their survival tactics, and sexual dimorphism. Males display brighter colours than females to make themselves more attractive to a mate. The colour of the lizard is also impacted by their age, season, and temperature of their body. [8] The ring-tailed dragon is about 25–35 cm long and is differentiated from other lizards by the line of spines that curves beneath the eyes. The pattern of the ctenophorus caudicinctus can range from a pale beige to dark orange. They have distinctive banding around the tail and a white to yellow underbelly. They have a short crest of nuchal spines that varies in colour. [9] [10]

Etymology

Ctenephorus is the largest group of lizards in Australia. Ctenephorus means comb-bearing dragons and caudicinctus means ring-tailed. Many of these species have been grouped by similar morphology and have been given informal names derived from mythological creatures. [10]

Range of where ctenophorus caudicinctus can be found in Western Australia Australian map of Ctneophorus caudicinctus location.png
Range of where ctenophorus caudicinctus can be found in Western Australia

Anatomy

Excretion

The environment in Western Australia where Ctenophorus Caudicinctus is commonly found is arid and dry. [11] Ctneophorus caudicinctus and many other lizards in dry tropics, do not produce hyperosmotic urine. The wall of the colon absorbs water and salts so that the body can retain water to prevent dehydration. [12]

Sensory characteristics

Ctenophorus caudicinctus has vestibular hearing. [13] The hair cells located in the neuroepithelial structures in the inner ear acts as receptors. They convert mechanical energy by the displacement of their surroundings into electrochemical energy. [14] The displacement of their surroundings are caused by sound waves and they use the balance of their head to perceive their surroundings. [15]

Maintenance of water

Ctenophorus caudicinctus has scale hinges that overlap each other and it creates an enclosed spaced between the scales. The scale hinges are flexible and able to stretch due to its thin layer of β-keratin in the hinge joint. These scale hinges create a smooth surface texture. [16] The water flows into the mouth and over the scales not between the scales. The purpose of this is to promote cutaneous water transport to the body. It allows water to be absorbed into the deeper layers of the body through capillary action and hinge joint channels. [17]

Osmoregulation

In spring where there is no breeding, ctenophorus caudicinctus has more sodium and potassium in their plasma and lower water influx and efflux. However, in autumn there is an increase in water metabolism. There are higher water influx and efflux but less sodium and potassium in their body. Their total body water is higher in autumn than in spring. This is due to the mating season and heavy rainfall in autumn. [18] Sodium and potassium content is important because it shows the rehydration or dehydration of any lizard. The nasal salt gland contributes to the potassium excretion for rehydration. Therefore, in autumn ctenophorus caudicinctus is more hydrated than in spring. Osmoregulation is important in lizards because of hormonal control, maintaining bodily functions and survival. [19]

Autotomy

Most lizards can regrow their tail and they can lose it voluntarily. However, most agamid lizards, including the ctenophorus caudicinctus do not lose their tail. Their tail is very thick compared to other lizards that can regrow their tail. [20] Therefore, it is hard to regenerate if they lose their tail. In addition, ctenophorus caudicinctus uses their tail for balance when they bipedal. [21]

Evolution

Ctenophorus caudicinctus is part of a large genus that shares similar phenotypes. But are still distinct from each other. Ctenophorus is a type of convergent evolution that both the ecological and biological factors that cause the evolution of a species to repeat. [22]

The ctenophorus genera have an evolution rate of approximately 1.3% divergence between lineages per million years of evolutionary separation. This rate of evolution is common in vertebrates because of their rapid reproduction, survival tactics, and natural selection. [23] The ctenophorus has two lineages that arose during the late Miocene and Pliocene. Which caused a separation of species to be separately distributed to Northern and southern parts of Australia. The ancestorial ecology of ctenophorus has features similar to the current day ctenophorus species of rock sheltering, shrub sheltering, and digging. These features are derived from a burrowing ancestor. [24] Through the geographic separation of the original dragons and different climates, the ctenophorus caudicinctus developed their own characteristics for survival. [25]

Plasticity also plays a role in the evolution of the ctenophorus caudicinctus. The ctenophorus can develop phenotypes depending on the environment that they face and have similar morphologies. [25] [26] Ctenophorus caudicinctus has a larger head than other species. They have favourable limb and snout length for hunting resulting in bigger and flatter heads. Along with pressure from predators, they gained relatively long limbs to move more quickly. To manage the heat, the ctenophorus caudicinctus is lighter in colour to absorb less heat. [27]

Adaptation

The morphology of ctenophorus caudicinctus is highly impacted by their habitat. It lives in rocky ranges, whilst other species live in burrows, with no burrows, and vegetation. Therefore, the bone structure of this species adapts to its surroundings and defers from other species of the ctenophorus. Ctenophorus caudicinctus has a longer and narrower spout to catch its prey and a more flattened body to fit through horizontal spaces. [28]

Behaviour

Thermoregulation

Ctenophorus caudicinctus with pale beige body and spotting. Ctenophorus Caudicinctus.jpg
Ctenophorus caudicinctus with pale beige body and spotting.

Thermoregulation behaviour is important for ctenophorus caudicinctus to carry out daily activities. The availability of proper thermal radiation is limited to species like ctenophorus caudicinctus because of extreme seasons. [29] Ctenophorus caudicinctus body temperature in shade is 34.2 7 °C. They start their day on the ground and move higher throughout the day. Then return to the ground at the end of the day. Their body temperatures are mostly kept cool because of higher atmospheres and less absorbance of the sun. If the weather isn't appropriate, ctenophorus caudicinctus can use psychological changes to be active during the day. It can increase their absorbance or loss of heat up to 77–87.7% through changing the colour of their skin. This would help ctenophorus caudicinctus during lower climates like spring and winter to increase their body temperature. Vice versa, they can lower their body temperatures in hotter seasons. [24]

Retreat

Ctenophorus caudicinctus has highspeed locomotion and retreats to rocks. They have bipedalism and quadrupedalism types of retreat. Ctenophorus caudicinctus is able to be bipedal due to their morphology. Ctneophorus caudicinctus has a narrow body, long hindlimbs, short forelimbs and a long tail. These are adaptations to bipedalism. [30]

Communication

Ctenophorus caudicinctus communicates with animals and things around them through motion-based signalling. Their motion-based signalling is heavily impacted by their surroundings. [31] They can identify things through the sound, seismic, and colours. They choose to not move to blend into their surroundings to hide from predators. Therefore, their running speed is not as fast as other species in their genera. One of the ways ctenophores caudicinctus communicates with other species is being territorial and use of motor patterns to recognize other species. There are many other lizard species in their habitat that are in their genera. Therefore, the ctenophores caudicinctus' unique behaviour allows them to be able to differentiate their kind from others. [32]

Ecology

Habitat

Ctenophorus caudicinctus is found in places with low vegetation, rocky and arid. Ctenophorus caudicinctus is highly territorial. [32] They are mostly found in Hameresly ranges in Western Australia, the Pilbara, MacDonnell, James and Musgrave Ranges. [33]

Diet

Owing to the ctenophorus caudicinctus being a diurnal and saxicolous lizard species living in rocky areas, the species is insectivorous. They consume mostly arthropods, termites and ants. Occasionally they will eat some vegetative material. Termites can be found in large numbers in one spot. Ctenophorus caudicinctus needs to be wary about ant eating lizards. They eat the skin of the ants. [34] Ctenophorus caudicinctus primarily waits for their prey in sheltered rocks and burrows. [25] Ctenophorus caudicinctus are ambush predators. They catch their predators with strategy, rather than attacking. [35]

Effects of extreme weathers

Ctenophorus caudicinctus does not dig burrows and lives in an arid area. Therefore, it is more difficult to find shelter. Extreme weather conditions such as lightning and heavy storms that can cause fires, often kill them. Periods of limited rainfall can also be the cause of death for lizards. Too much heat causes them to be dehydrated or burns them. The lack of shade and shelter exposes ctneophorus caudicinctus to these harsh environments. [36]

Predators

Ctenophorus caudicinctus is high in abundance and has very few predators. Therefore, they are not an endangered species. Their main predators are snakes, dugites and pythons. [37]

Parasites

Ctenophorus caudicinctus has a symbiotic relationship with skrjabinoptera phrynosoma. [38] It is stomach parasite and their hosts are commonly lizards. [39]

Reproduction

Red-brown coloured ctenophorus caudicinctus Ring-tailed Dragon (Ctenophorus caudicinctus) (8851342177).jpg
Red-brown coloured ctenophorus caudicinctus

The ring tailed dragon's reproductive cycle favours conditions that are best for their survival and growth. Thus, reproductive cycles vary over the years due to rainfall and the availability of food. [8] Most commonly, ctenophorus caudicinctus breeds during spring, September to October. Then in the following months of winter, there is winter rainfall. This rainfall is important as it stimulates insect abundance to provide abundant food for ctenophorus caudicinctus. Adequate food helps with fertilization, spermatogenesis, and healthy reproduction. [40] For males, the main season for spermiogenesis is from February to August as this when they generate the maximum amount of sperm and conditions are optimal for breeding. However, some males do not mate until they are matured. They would hold it in until the next mating season. [41] Ctenophorus caudicinctus is oviparous. [42] The number of offspring is lesser than the number of eggs produced because of a lack of maternal protection. [43]

Physical attributes in mating

Physical attributes also impact how males and females mate. Males have bigger heads to assert dominance and win intrasexual competitions. In addition, colour plays an important role. [8] During mating season, males change colour adopting yellow markings on the flanks, and a dark chest patch. The yellow colour emphasizes the banding on their tails. However, they lose their original camouflage and this makes them easier to be caught by predators. [44] Bright colours and the pattern of their body makes the males more attractive. Most females have a single coloured pattern over their body, whereas males have more complex colour patterns. [45]

Brain structure and sexual selection

Ctenophorus caudicinctus has two regions in the brain that regulate sexual behaviour. These regions are the medial preoptic nucleus (MPON), which controls male reproductive behaviour, and the ventromedial hypothalamic nucleus (VMN), which controls the female reproductive behaviour and is involved in male aggression. Males have larger MPON and smaller VMN. In females they have similar MPON and VMN. The volumes of both these vertebrae are correlated to their brain volume and differ from one lizard to another. [46]

Conservation status

It is highly abundant and has no conservation significance. Ctenophorus caudicinctus are assessed as being of the least concern in the Nature Conservation Act 1992 (NCA). [47]

Subspecies

Ctenophorus caudicinctus has two subspecies: [48] [49]

The subspecies of Ctenophorus caudicinctus all have the same band pattern and vary in the number of spotting and colour. Some are brighter more vibrant colours like green and red. However, Ctenophorus caudicinctus is composed of brown and dark red colours to blend into the rocky habitat. [50]

Related Research Articles

<span class="mw-page-title-main">Lizard</span> Informal group of reptiles

Lizard is the common name used for all squamate reptiles other than snakes, encompassing over 7,000 species, ranging across all continents except Antarctica, as well as most oceanic island chains. The grouping is paraphyletic as some lizards are more closely related to snakes than they are to other lizards. Lizards range in size from chameleons and geckos a few centimeters long to the 3-meter-long Komodo dragon.

<span class="mw-page-title-main">Agamidae</span> Family of lizards

Agamidae is a family of over 550 species of iguanian lizards indigenous to Africa, Asia, Australia, and a few in Southern Europe. Many species are commonly called dragons or dragon lizards.

<i>Ctenophorus</i> Genus of lizards

Ctenophorus is a genus of lizards, commonly known as comb-bearing dragons, found in Australia. They are in the dragon lizard family, known as Agamidae.

<span class="mw-page-title-main">Frilled lizard</span> Species of reptile

The frilled lizard, also known as the frillneck lizard, frill-necked lizard or frilled dragon, is a species of lizard in the family Agamidae. It is native to northern Australia and southern New Guinea. This species is the only member of the genus Chlamydosaurus. Its common names come from the large frill around its neck, which usually stays folded against the lizard's body. It reaches 90 cm (35 in) from head to tail and can weigh 600 g (1.3 lb). Males are larger and more robust than females. The lizard's body is generally grey, brown, orangish-brown, or black in colour. The frills have red, orange, yellow, or white colours.

<span class="mw-page-title-main">Jacky dragon</span> Species of lizard

The jacky dragon is a type of lizard native to south-eastern Australia. Other common names include blood-sucker, stonewalker, and tree dragon. It was one of the first Australian reptiles to be named by Europeans, originally described by English zoologist George Shaw in Surgeon-General John White's Journal of a Voyage to New South Wales, published in London in 1790. The lizard is well-known for its bright yellow mouth and well-developed vertebral crest, as well as the temperature-dependent sex determination of its offspring.

<i>Ctenophorus maculosus</i> Species of lizard

Ctenophorus maculosus, commonly known as the Lake Eyre dragon or salt-lake ground-dragon, is a species of agamid lizard endemic to South Australia. C. maculosus mainly inhabits the edges of salt lakes in South Australia, from which its common names are derived. It survives in this harsh habitat through adaptive mechanisms to tolerate high temperatures and lack of free water. The female C. maculosus possesses unique male rejection techniques which are currently the focus of further research.

<span class="mw-page-title-main">Central netted dragon</span> Species of lizard

The central netted dragon or central netted ground dragon is a species of agamid lizard occurring in a wide range of arid to semiarid regions of Australia. It is widespread across the continent, commonly found in open, sandy, desert habitats. It is a popular pet and can often be found in zoos.

<i>Ctenophorus pictus</i> Species of lizard

Ctenophorus pictus, commonly known as the painted ground-dragon or painted dragon, is a species of lizard from the family Agamidae. It is endemic to the drier areas of southern and central Australia.

<i>Ctenophorus mirrityana</i> Species of lizard

The Barrier Range dragon is an agamid lizard which has been newly described (2013) as a separate species from the tawny dragon. This species is endemic to Australia, restricted to just three sites in western New South Wales (NSW).

<i>Ctenophorus fionni</i> Species of lizard

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<i>Ctenophorus decresii</i> Species of lizard

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<i>Tympanocryptis tetraporophora</i> Species of lizard

Tympanocryptis tetraporophora, also known as Eyrean earless dragon or long-tailed earless dragon, is one of a documented species of a relatively small dragon belonging to the genus Tympanocryptis. Tympanocryptis is differentiated from other genera within the family Agamidae by a tympanum covered with scales and a missing phalange in the fifth toe of the rear foot. T. tetraporophora is a ground dwelling dragon inhabiting semi arid regions of central New South Wales, arid regions of South Australia, Northern Territory, Queensland and tropical grasslands of Northern Queensland.

The gibber earless dragon also known as the smooth-snouted earless dragon, is a species of agamid lizard endemic to Australia. It is one of a documented species of the genus Tympanocryptis, a group of small terrestrial lizards that feed off invertebrates and are characterised by the absence of an external ear structure.

<span class="mw-page-title-main">Mallee military dragon</span> Species of lizard

The Mallee dragon, also commonly known as the Mallee dragon and the Mallee sand-dragon, is a species of lizard in the family Agamidae. The species is native to the arid parts of southern Australia.

<i>Ctenophorus gibba</i> Species of lizard

Ctenophorus gibba, commonly known as the bulldust ground-dragon or gibber dragon, is a species of agamid lizard occurring in the sparsely vegetated gibber plains of northern South Australia.

Ctenophorus mckenziei, more commonly known as the dwarf-bicycle dragon, is a species of endemic Australian lizard within the family Agamidae and genus Ctenophorus. Originally identified as the agamid Amphibolurus mckenziei, the lizard had been identified within the regions of Western Australia and South Australia in which it occupied the shrubbery and woodland areas as its habitat. It was subsequently transferred to the genus Ctenophorus along with other Agamid species in which it shared similar morphology and characteristics. The name mckenziei is in reference to Norman Leslie Mckenzie, who was a zoologist and discovered the existence of the lizard. Listed on the IUCN red list page, threats to its population numbers are evaluated as least concern; however, their numbers are threatened by habitat loss, climate change, and feral predators.

<i>Ctenophorus parviceps</i> Species of lizard

Ctenophorus parviceps, commonly known as the Gnaraloo heath dragon or northwestern heath dragon is a species of agamid lizard occurring in pale coastal sands and shell grit with open heaths and beach spinifex, between the North West Cape and Carnarvon, Western Australia and on Bernier Island. The Gnaraloo heath dragon is a lizard that can be found along the coast of Western Australia between Exmouth Gulf and Shark Bay, and is also known as the northwestern heath dragon. It is native to Australia and usually inhabits sandy coastal dunes. The species' longevity is 3–50 years and its population density is extremely low. The Gnaraloo heath dragon is a member of the Agamidae family, which contains 15 genera. The lizard is under the Ctenophorus genus which has up to 33 species. This genus shows the most morphological and ecological diversity out of the three large agamid genera. 83% of the lizards in this genus lack a crest, while 17% possess crests. They are smaller than most agamids but do have relatively large heads. The Gnaraloo heath dragon can be differentiated from related species by a series of spines on the tail's base, a pale-grey brown broad vertebral band along its back, and hour-glass bars extending upwards to meet the pale vertebral band. It is usually 45mm in terms of length, measuring from snout to vent.

<i>Ctenophorus tjantjalka</i> Species of lizard

Ctenophorus tjantjalka, also known as the ochre dragon, is a species of agamid lizard occurring on low, weathered, crumbling outcrops and stony hills in arid South Australia, from the Painted Hills north-west to the base of the Everard Ranges.

Ctenophorus infans, the Laverton ring-tailed dragon, is a species of agamid lizard occurring around Laverton and the Mount Margaret Goldfield of Western Australia.

<i>Ctenophorus spinodomus</i> Species of lizard

Ctenophorus spinodomus commonly known as Eastern Mallee Dragon, is a species of agamid lizard endemic to the arid and semi-arid regions of southeastern Australia. Belonging to the genus Ctenophorus, which comprises a diverse group of 35 species this small lizard is characterised by striking colours and unique behaviours. Commonly known as dragon lizards, spinodomus is derived from the Greek words "spinos" (thorn) and "domos" (house), referring to its preference for burrowing habitats among spiny vegetation. They are curious creatures and can be found darting between the safety of Triodia grass clumps or basking on the red sand.

References

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  28. ↑ Thompson, Graham; Withers, Philip (2005). "The relationship between size-free body shape and choice of retreat for Western Australian Ctenophorus (Agamidae) dragon lizards". Amphibia-Reptilia. 26 (1): 65–72. doi: 10.1163/1568538053693323 . ISSN   0173-5373.
  29. ↑ Smith, Kathleen R.; Cadena, Viviana; Endler, John A.; Porter, Warren P.; Kearney, Michael R.; Stuart-Fox, Devi (15 June 2016). "Colour change on different body regions provides thermal and signalling advantages in bearded dragon lizards". Proceedings of the Royal Society B: Biological Sciences. 283 (1832): 20160626. doi:10.1098/rspb.2016.0626. ISSN   0962-8452. PMC   4920320 .
  30. ↑ Clemente, Christofer J.; Withers, Philip C.; Thompson, Graham; Lloyd, David (1 July 2008). "Why go bipedal? Locomotion and morphology in Australian agamid lizards". Journal of Experimental Biology. 211 (13): 2058–2065. doi: 10.1242/jeb.018044 . ISSN   1477-9145. PMID   18552294.
  31. ↑ Witter, Geoffrey. "FAUNA of Australia" (PDF). 26: 26–30.{{cite journal}}: Cite journal requires |journal= (help)
  32. 1 2 Ramos, Jose A.; Peters, Richard A. (August 2017). "Motion-based signaling in sympatric species of Australian agamid lizards". Journal of Comparative Physiology A. 203 (8): 661–671. doi:10.1007/s00359-017-1185-5 (inactive 3 May 2024). ISSN   0340-7594. PMID   28573349. S2CID   43838080.{{cite journal}}: CS1 maint: DOI inactive as of May 2024 (link)
  33. ↑ Wilson, Steve (2012). Australian lizards : a natural history. Collingwood, Vic.: CSIRO Pub. ISBN   978-0-643-10641-3. OCLC   794058548.
  34. ↑ Pianka, Eric R.; Pianka, Helen D. (2 March 1970). "The Ecology of Moloch horridus (Lacertilia: Agamidae) in Western Australia". Copeia. 1970 (1): 90. doi:10.2307/1441978. JSTOR   1441978.
  35. ↑ "Encyclopedia of Life". eol.org. Retrieved 14 May 2021.
  36. ↑ Light, Paul; Dawson, William R.; Shoemaker, Vaughan H.; Main, A. R. (22 March 1966). "Observations on the Thermal Relations of Western Australian Lizards". Copeia. 1966 (1): 97. doi:10.2307/1440766. JSTOR   1440766.
  37. ↑ "Herpetological Review". Society for the Study of Amphibians and Reptiles. 8 November 2016. Retrieved 14 May 2021.
  38. ↑ "Taxonomy browser (Ctenophorus caudicinctus)". ncbi.nlm.nih.gov. Retrieved 14 May 2021.
  39. ↑ Zoological Society of London. (1833). Proceedings of the Zoological Society of London. Vol. 1922:pt.3-4 [pp.483-1276]. London: Academic Press, [etc.]
  40. ↑ On the ecology of Australia's arid zone. H. Lambers. Cham, Switzerland. 2018. ISBN   978-3-319-93943-8. OCLC   1049568222.{{cite book}}: CS1 maint: location missing publisher (link) CS1 maint: others (link)
  41. ↑ Goldberg, Stephen (1 January 2009). "Male cycle of the military sand-dragon Ctenophorus isolepis, from Western Australia". Herpetofauna. 39: 43–45 – via ResearchGate.
  42. ↑ "Ctenophorus caudicinctus". The Reptile Database. Retrieved 14 May 2021.
  43. ↑ "Oviparity | biology". Encyclopædia Britannica. Retrieved 14 May 2021.
  44. ↑ "Spinelesswonders". spinelesswonders.smugmug.com. Retrieved 14 May 2021.
  45. ↑ Chen, I-Ping; Stuart-Fox, Devi; Hugall, Andrew F.; Symonds, Matthew R. E. (November 2012). "Sexual Selection and the Evolution of Complex Color Patterns in Dragon Lizards". Evolution. 66 (11): 3605–3614. doi:10.1111/j.1558-5646.2012.01698.x. PMID   23106722. S2CID   14242593.
  46. ↑ Hoops, D.; Ullmann, J. F. P.; Janke, A. L.; Vidal-Garcia, M.; Stait-Gardner, T.; Dwihapsari, Y.; Merkling, T.; Price, W. S.; Endler, J. A.; Whiting, M. J.; Keogh, J. S. (February 2017). "Sexual selection predicts brain structure in dragon lizards". Journal of Evolutionary Biology. 30 (2): 244–256. doi: 10.1111/jeb.12984 . hdl: 10536/DRO/DU:30088478 . ISSN   1420-9101. PMID   27696584.
  47. ↑ Department of Environment and Science (20 October 2014). "Species profile | Environment, land and water". apps.des.qld.gov.au. The State of Queensland. Retrieved 14 May 2021.
  48. ↑ Cogger, H. (2000) Reptiles and Amphibians of Australia, Reed New Holland, Sydney, New South Wales, ISBN   1876334339
  49. ↑ Wilson, S., Swan, G. (2013) A Complete Guide to Reptiles of Australia, New Holland Publishers, Sydney, New South Wales, ISBN   9781921517280
  50. ↑ Melville, Jane; Haines, Margaret L.; Hale, Joshua; Chapple, Stephanie; Ritchie, Euan G. (2016). "Concordance in phylogeography and ecological niche modelling identify dispersal corridors for reptiles in arid Australia". Journal of Biogeography. 43 (9): 1844–1855. doi: 10.1111/jbi.12739 . ISSN   1365-2699.
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