Biology:Armenian cochineal

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Short description: Species of true bug

Armenian cochineal
Porphyrophora hamelii, female.jpg
Porphyrophora hamelii (female)
Porphyrophora hamelii, male.jpg
Porphyrophora hamelii (male)
Scientific classification edit
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hemiptera
Suborder: Sternorrhyncha
Family: Margarodidae
Genus: Porphyrophora
Species:
P. hamelii
Binomial name
Porphyrophora hamelii
Brandt, 1833
Synonyms [1]
  • Porphyrophora hameli Brandt, 1834 (revived by Jakubski, 1965)
  • Porphyrophora armeniaca Burmeister, 1835
  • Coccus radicis Meyer, 1856
  • Margarodes hameli Cockerell, 1902 and Fernald, 1903
  • Coccionella armeniaca Lindinger, 1954
  • Coccionella hamelii Lindinger, 1954
  • Coccionella hameli Lindinger, 1954

The Armenian cochineal (Porphyrophora hamelii), also known as the Ararat cochineal or Ararat scale, is a scale insect indigenous to the Ararat plain and Aras (Araks) River valley in the Armenian Highlands and in Turkey. It was formerly used to produce an eponymous crimson carmine dyestuff known in Armenia as vordan karmir (Armenian: որդան կարմիր, literally "worm's red") and historically in Persia as kirmiz.[1][2][3][4][5][6] The species is critically endangered within Armenia.[7]

The Armenian cochineal scale insect, Porphyrophora hamelii, is in a different taxonomic family from the cochineal found in the Americas. Both insects produce red dyestuffs that are also commonly called cochineal.[8]

History and art

Porphyrophora hamelii is one of the ancient natural sources of red dye in the Middle East and Europe, along with the insect dyes kermes (from Kermes vermilio and related species), lac (from Kerria lacca and related species), and carmine from other Porphyrophora species such as the Polish cochineal (Porphyrophora polonica), and the plant dye madder (from Rubia tinctorum and related species).[3][4][6][9][10][11][12][13] It is possible that Armenian cochineal dye was in use as early as 714 BC, when the Neo-Assyrian king Sargon II was recorded as seizing red textiles as spoils of war from the kingdoms of Urartu (the geographic predecessor of Armenia) and Kilhu.[3][4][14][15][16] The Roman-era physician and pharmacologist Dioscorides, writing in the 1st century AD, noted that the best kokkos baphike, the kermes shrub and its "grain" (kermes insect) that some ancient writers likely confused with Porphyrophora hamelii, came from Galatia and Armenia.[3][4][17] In the Early Middle Ages the Armenian historians Ghazar Parpetsi and Movses Khorenatsi wrote specifically of a worm-produced dyestuff from the Ararat region.[4][16]

During the Middle Ages the Armenian cochineal dyestuff vordan karmir, also known in Persia as kirmiz, was widely celebrated in the Near East.[4][5][6][16] Kirmiz is not to be confused with dyer's kermes, which was derived from another insect.[6] The Armenian cities Artashat and Dvin were early centers of the production of kirmiz: during the 8th through 10th centuries Arab and Persian historians even referred to Artashat as "the town of kirmiz".[4][16] The Arabs and Persians regarded kirmiz as one of the most valuable commodities exported from Armenia.[6] The Armenians themselves used vordan karmir to produce dyes for textiles (including oriental rugs) and pigments for illuminated manuscripts and church frescos.[4][16][18][19] Chemical analyses have identified the dye of Porphyrophora hamelii in Coptic textiles of the 3rd through 10th centuries, a cashmere cloth used in a kaftan from Sassanid Persia in the 6th or 7th century, silk liturgical gloves from 15th-century France , Ottoman fabrics such as velvets and lampas of the 15th through 17th centuries, and a 16th-century velvet cap of maintenance that belonged to Henry VIII of England.[6][20][21]

At the time of the Renaissance in Europe, Porphyrophora insects were so valuable that in Constantinople during the 1430s, one kilogram (2.2 pounds) of Porphyrophora hamelii insects was worth more than 5 grams (0.18 ounces) of gold.[6][22][23] The crimson Porphyrophora-based dyes were especially prized in Europe for dyeing silk, as the scarlet dye kermes was more plentiful, cheaper, and more effective for dyeing woolen textiles, which are heavier than silk and require more dye.[6] It has been estimated that on the order of a half million dried Porphyrophora hamelii insects were required to dye one kilogram (2.2 lb) of silk crimson during this period.[6][24] On the comparison between Armenian and Polish cochineal, the author of a 15th-century treatise on silks in Florence wrote that "two pounds of the large [Armenian cochineal insects] will dye as much [silk] as one pound of small [Polish cochineal insects]; it is true that it gives a more noble and brighter colour than the small, but it gives less dye."[6][25]

Around the end of the 16th century the Old World Porphyrophora dyes were supplanted by dyes of the Dactylopius coccus cochineal species from the Americas, which could be harvested several times per year and yielded a much more concentrated dye.[15]

The carmine dyestuff of Porphyrophora hamelii owes its red color almost entirely to carminic acid, making it difficult to distinguish chemically from the dyestuff of cochineal from the Americas.[6][9][10][15][26] The dyestuff of Porphyrophora polonica can be distinguished by its small admixture of kermesic acid, which is the major constituent of kermes from Kermes vermilio.[citation needed]

In 1833 the German naturalist Johann Friedrich von Brandt suggested the scientific name Porphyrophora hamelii after the Russian physician, traveler, and historian of German descent Iosif Khristianovich Gamel (Josef Hamel) (ru), who visited the Ararat plain in the early 1830s and wrote a report about the "cochineal" insects living there.[27][28]

  • Vordan karmir ceiling art at Noravank Monastery

  • Khachkars (cross-stones) at Geghard Monastery painted with vordan karmir[29]

  • Silk liturgical gloves (left) of Cardinal Pierre de Foix (15th century), dyed crimson with P. hamelii carmine[6]

  • A 2006 Armenian postage stamp depicting P. hamelii

Biology

Porphyrophora hamelii cysts around the root of Aeluropus littoralis

Porphyrophora hamelii is a sexually dimorphic species.[30][31][32] The adult female, from which carmine is extracted, is oval-shaped, soft-bodied, crimson in color, and has large forelegs for digging. The females can be quite large for a Porphyrophora species: up to 10–12 mm (0.39–0.47 in) long and 7 mm (0.28 in) wide.[30][32] It has been noted that one troy pound (360 grams) of cochineal insects requires 18,000–23,000 specimens of Porphyrophora hamelii, but 100,000–130,000 specimens of the sister species Porphyrophora polonica (or 20,000–25,000 specimens of Dactylopius coccus).[33][34] The adult male Porphyrophora hamelii is a winged insect.

The life cycle of Porphyrophora hamelii is mostly subterranean.[7][30][31] Newly hatched nymphs emerge from the ground in the springtime and crawl until they find the roots of certain grassy plants that grow in saline soil, such as Aeluropus littoralis (Armenian: որդանխոտ (genus Aeluropus),[18][35] literally "worm's grass") and the common reed Phragmites australis.[7][30] The nymphs continue to feed on these roots throughout the spring and summer, forming protective pearl-like cysts in the process. From mid-September to mid-October adults emerge from the ground between 5 a.m. and 10 a.m. to mate.[6][7][35][36] The adult insects, lacking mouthparts, do not feed.[31] Adult males live for only a few days, but adult females can live longer, burrowing into the ground to lay their eggs.[31]

Habitat and conservation

The historic habitat of Porphyrophora hamelii. Vordan Karmir State Reservation is in red and the historic dye-producing cities Artashat and Dvin are in purple.

The red dye-producing insects of the Ararat plain were once plentiful: a 19th-century French traveler wrote that shepherds' flocks, when led to drink from the Araxes (Araks) River, would appear bloody from the insects.[4] In the mid-20th century the extent of occurrence in Armenia was 100 km2 (39 sq mi) with a recorded distribution that included the Ararat and Armavir provinces in Armenia as well as the Turkish, Iranian, and Russian Caucasus, but by the 1990s the extent of occurrence in Armenia had shrunk to about 20 km2 (7.7 sq mi), mostly in Armavir Province.[7][30][37] During the Soviet period, desalination of the Armenian salt marshes to create "economic and agricultural regions", and the creation of lakes for fisheries, "severely restricted the [habitable] area for the insects and [endangered their] existence."[35]

Vordan Karmir State Reservation, Armenia

The population in Armenia resides almost entirely in the Vordan Karmir State Reservation, a salt meadow habitat of 198.33 ha (490.1 acres) northwest of Arazap village and 21.52 ha (53.2 acres) in the north of Jrarat village established in 1987 near the Araks River border with Turkey, plus a site southeast of Ararat village and a few patches of several hectares elsewhere.[7][38][39] There have been no recent scientific reports on populations of Porphyrophora hamelii outside the surroundings of Mount Ararat.[6]

Porphyrophora hamelii is considered critically endangered within Armenia by meeting the following conditions: an area of occupancy of less than 10 km2 (3.9 sq mi), plus severely fragmented occupancy or known to exist at only a single location, plus continued decline (observed, inferred, or projected) in the area of occurrence, area of occupancy, and area, extent, and/or quality of habitat; and an extent of occurrence of less than 100 km2 (39 sq mi) with the aforementioned conditions of continued decline.[7][40]

Threats to the Porphyrophora hamelii population in Armenia include the development of saline lands, agricultural improvements, uncontrolled livestock grazing, and possibly climate change.[7] Natural foes of the species include mold mites, lady beetles, harvester ants, and erratic ants.[1][2]

See also

References

  1. ↑ 1.0 1.1 1.2 Ben-Dov, Y.; Miller, D.R.; Gibson, G.A.P. (9 October 2014). "ScaleNet, Porphyrophora hamelii". http://scalenet.info/validname/Porphyrophora/hamelii/. 
  2. ↑ 2.0 2.1 Ben-Dov, Yair (2005). A Systematic Catalogue of the Scale Insect Family Magarodidae (Hemiptera: Coccoidea) of the World. United Kingdom: Intercept (Lavoisier). ISBN 978-1-84585-000-5. 
  3. ↑ 3.0 3.1 3.2 3.3 Forbes, R.J. (1964). Studies in Ancient Technology. IV (2nd ed.). Leiden, Netherlands: E.J. Brill. pp. 102–103. ISBN 90-04-08307-3. https://books.google.com/books?id=ua83AAAAIAAJ&pg=PA102. 
  4. ↑ 4.0 4.1 4.2 4.3 4.4 4.5 4.6 4.7 4.8 Donkin, R.A. (1977). "The Insect Dyes of Western and West-Central Asia". Anthropos (Anthropos Institute) 72 (5/6): 847–880. 
  5. ↑ 5.0 5.1 Vedeler, Marianne (2014). Silk for the Vikings. Oxford, United Kingdom: OXBOW BOOKS. p. 52. ISBN 978-1-78297-215-0.  Vedeler, citing Cardon (2007), notes that "the Persian name Kirmiz originally referred to the Armenian carmine, a parasitic insect living on Gramineae grass, but the same name was also used by Arab geographers for insects living on oak trees in Maghreb and Al-Andalus, probably referring to Kermes vermilio", although "[i]t is ... not clear whether the 'Kirmiz' dyestuff mentioned in early Arab texts always refers to the use of the insect Kermes Vermilio."
  6. ↑ 6.00 6.01 6.02 6.03 6.04 6.05 6.06 6.07 6.08 6.09 6.10 6.11 6.12 6.13 Cardon, Dominique (2007). Natural Dyes: Sources, Tradition, Technology and Science. London, United Kingdom: Archetype Books. ISBN 978-1-904982-00-5.  English translation by Caroline Higgitt of Cardon's French-language book Le monde des teintures naturelles (Éditions Belin, Paris, 2003).
  7. ↑ 7.0 7.1 7.2 7.3 7.4 7.5 7.6 7.7 Khachatryan, H.. "Porphyrophora ham melii Brandt, 1833". Republic of Armenia Ministry of Environment. http://www.mnp.am/red_book_fauna/eng/a65.html. 
  8. ↑ Eastaugh, Nicholas; Walsh, Valentine; Chaplin, Tracey; Siddall, Ruth (2004). Pigment Compendium: A Dictionary of Historical Pigments. Oxford, UK and Burlington, MA: Elsevier Butterwoth-Heinemann. pp. 118–119. ISBN 0-7506-5749-9. https://books.google.com/books?id=2OH28vpzzMsC&pg=PA118. Retrieved 4 December 2015. 
  9. ↑ 9.0 9.1 Robinson, Stuart (1969). A History of Dyed Textiles. London, United Kingdom: Studio Vista. ISBN 978-0-289-79644-3. 
  10. ↑ 10.0 10.1 Böhmer, Harald; Thompson, Jon (1991). "The Pazyryk Carpet: A Technical Discussion". Source: Notes in the History of Art (Ars Brevis Foundation) 10 (4): 30–36. doi:10.1086/sou.10.4.23203293. 
  11. ↑ Koren, Zvi C. (2005), "Chromatographic analyses of selected historic dyeings from ancient Israel", in Janaway, R.; Wyeth, P., Scientific Analysis of Ancient and Historic Textiles: Informing, Preservation, Display and Interpretation, London, United Kingdom: Archetype Publications, pp. 194–201, http://www.tekhelet.com/pdf/KOREN_2005_194-201.pdf, retrieved 8 October 2014 
  12. ↑ Cardon, Dominique (2010), "Natural Dyes, Our Global Heritage of Colours", Proceedings of the 12th Biennial Symposium of the Textile Society of America ("Textiles and Settlement: From Plains Space to Cyber Space") (October 6–9, 2010), Lincoln, Nebraska, USA: Textile Society of America, http://digitalcommons.unl.edu/tsaconf/12/ 
  13. ↑ Kirby, Jo (2011), "Dyes, Dyeing and Lake Pigments – Historical Background", Back to the Roots – Workshop on the Preparation of Historical Lake Pigments (March 23–25, 2011), Munich, Germany: Doerner Institut, http://www.doernerinstitut.de/downloads/Back_to_the_Roots/Back_to_the_roots_Kirby_I.pdf, retrieved 9 October 2014 
  14. ↑ Thureau-Dangin, François (1912) (in fr). Une Relation de la Hutième Campagne de Sargon (714 av. J.-C.) texte Assyrien inédit, publié et traduit. Paris, France: Librairie Paul Geuthner. p. 57. https://archive.org/details/unerelationdelah00thuruoft. Retrieved 8 October 2014. 
  15. ↑ 15.0 15.1 15.2 Phipps, Elena (2010). Cochineal Red: The Art History of a Color [adapted from The Metropolitan Museum of Art Bulletin, v. 67, no. 3 (Winter 2010)]. New York City and New Haven, USA and London, UK: The Metropolitan Museum of Art and Yale University Press. ISBN 978-0-300-19560-6. http://www.metmuseum.org/research/metpublications/Cochineal_Red_the_art_history_of_a_color#. Retrieved 8 October 2014.  (Also at archive.org.)
  16. ↑ 16.0 16.1 16.2 16.3 16.4 Kurdian, H. (1941). "Kirmiz". Journal of the American Oriental Society (American Oriental Society) 61 (2): 105–107. doi:10.2307/594255. 
  17. ↑ Obaldeston, T.A. (2000). Dioscorides, De Materia Medica. Five books in one volume: new modern English translation. 4. Johannesburg, South Africa: IBIDIS Press. pp. 588–591. ISBN 978-0-620-23435-1. https://archive.org/details/de-materia-medica. Retrieved 4 January 2015.  English translation by T.A. Obaldeston with introductory notes by R.P. Wood.
  18. ↑ 18.0 18.1 Babenko, Vitali (1988). "Vordan Karmir or Armenian Cochineal". Oriental Rug Review (Oriental Rug Auction Review) VIII (5): 40–41. http://www.rugreview.com/orr/8-5-40.htm. Retrieved 9 October 2014. 
  19. ↑ Mushak, Paul (1988). "The use of insect dyes in Oriental rugs and textiles: Some unresolved issues". Oriental Rug Review (Oriental Rug Auction Review) VIII (5): 33–39. http://www.rugreview.com/orr/8-5-33.htm. Retrieved 9 October 2014. 
  20. ↑ Petroviciu, Irina; Crețu, Ileana; Vanden Berghe, Ina; Wouters, Jan; Medvedovici, Andrei; Albu, Florin; Creanga, Doina (2012). "A discussion on the red anthraquinone dyes detected in historic textiles from Romanian collections". E-Preservation Science (Morana RTD) 9: 90–96. http://www.morana-rtd.com/e-preservationscience/2012/Petroviciu-29-02-2012.pdf. Retrieved 4 January 2015. 
  21. ↑ "Henry VIII's Cap of Maintenance". Treasures of Medieval Waterford, Ireland: Medieval Museum. http://www.waterfordtreasures.com/medieval-museum/whats-inside/king-henry-viiis-cap-of-maintenance. 
  22. ↑ Cardon, Dominique (2000), "Du verme cremexe au veluto chremesino: une filierè vénitienne du cramoisi au XVe siécle", in Molà, L.; Mueller, R.C.; Zanier, C. (in fr), La Seta in Italia dal Medioevo al Seicento, Venice, Italy: Fondazione Giorgio Cini, pp. 63–73 
  23. ↑ Some articles improperly cite Cardon to suggest, incorrectly, that Armenian cochineal insects were more valuable, by weight, than gold (i.e., one gram of insects was worth several grams of gold) during this era. Cardon (2007) does, however, note that according to the records of a Venetian merchant trading in Constantinople during the 1430s, even the cheapest Armenian cochineal insects were still worth more, pound-for-pound, than some live slaves (Circassian women and adolescents) that he had bought.
  24. ↑ 400,000 to 560,000 dried P. hamelii insects were required to dye 1 kg of silk according to the figures of Cardon (2007): 1,000 g to 1,400 g of dried insects per 100 g of silk, with 40 adult females per gram of dried insects. Note that Virey (1840) reports 18,000–23,000 insects per 360-gram troy pound (50–64 insects per gram; not stated whether they were dried).
  25. ↑ Gargiolli, Girolamo, ed (1868) (in it). L'arte della seta in Firenze. Trattato del Secolo XV, pubblicato per la prima volta, e dialoghi raccolti da Girolamo Gargiolli.. Florence, Italy: G. Barbèra. p. 32. https://archive.org/details/lartedellasetai00garggoog. Retrieved 16 October 2014. 
  26. ↑ Wouters, Jan; Verhecken, André (1989). "The Coccid Insect Dyes: HPLC and Computerized Diode-Array Analysis of Dyed Yarns". Studies in Conservation (Maney Publishing) 34 (4): 189–200. doi:10.1179/sic.1989.34.4.189. 
  27. ↑ Brandt, Johann Friedrich; Ratzeburg, Julius Theodor Christian (1833) (in de). Medizinische Zoologie oder getreue Darstellung und Beschreibung der Thiere, in der Arzneimitellehre in Betracht kommen, in systematischer Folge herausgegeben.. 2. Berlin, Germany: Trowitzsch and Sohn. p. 356. https://books.google.com/books?id=ZQPnKys-3mkC&pg=PA356. Retrieved 11 October 2014. 
  28. ↑ Hamel, J. (1833), "Über Cochenille am Ararat und über Wurzelcochenille im Allgemeinen" (in de), Mémoires de l'Académie Impériale des Sciences de St. Pétersbourg / Sciences mathématiques, physiques et naturelles, Tome III (1835) [publishing the second part of Tome I (1833)], Frankfurt, Germany: Universitätsbibliothek Frankfurt am Main, pp. 9–64, http://publikationen.ub.uni-frankfurt.de/files/21947/E000014948.pdf, retrieved 11 October 2014 . Publication of Hamel's 4 May 1833 report on the Ararat cochineal. Hamel's report mentions Brandt.
  29. ↑ "Geghard 7: Monk cells, Khachkar Wall. Geghard's Khachkars.". Armenian Monuments Awareness Project. http://www.armenianheritage.org/en/monument/Geghard/78. "The red color found on some of the cross stones is a result of their being painted with Vortan Karmir, a red dye made from beetles native to Armenia. The red dye was among the more famous exports of the kingdom, and was valued more than gold in Europe and the Near East. Its resilience has long since proved itself; the color you see now is more than 800 years old." 
  30. ↑ 30.0 30.1 30.2 30.3 30.4 Vahedi, Hassan-Ali; Hodgson, C.J. (2007). "Some species of the hypogeal scale insect Porphyrophora Brandt (Hemiptera: Sternorrhyncha: Coccoidea: Margarodidae) from Europe, the Middle East and Africa". Systematics and Biodiversity (Taylor & Francis) 5 (1): 23–122. doi:10.1017/s1477200006002039. 
  31. ↑ 31.0 31.1 31.2 31.3 Foldi, Imre (2005). "Ground pearls: a generic revision of the Margarodidae sensu stricto (Hemiptera: Sternorrhyncha: Coccoidea)". Annales de la Société Entomologique de France. New Series (Taylor & Francis) 41 (1): 81–125. doi:10.1080/00379271.2005.10697442. http://zoologie.umh.ac.be/asef/pdf/2005_41_01/compact/Foldi_ASEF_2005_41_1_81_125_compact.pdf. Retrieved 9 October 2014. 
  32. ↑ 32.0 32.1 Jakubski, Antoni Władysław (1965). A Critical Revision of the Families Margarodidae and Termitococcidae (Hemiptera, Coccoidea). London, United Kingdom: Trustees of the British Museum (Natural History). 
  33. ↑ Virey, M.J.J. (1840), "Remarks on the former uses of purple or scarlet colouring insects. New cochineal of Persia and Armenia.", in Watt, Charles; Watt, John Jr., The Chemist: Or, Reporter of Chemical Discoveries and Improvements, and Protector of the Rights of the Chemist and Chemical Manufacturer, I, London, United Kingdom: R. Hastings, pp. 209–210, https://books.google.com/books?id=LKQwAAAAYAAJ&pg=PA209 
  34. ↑ Fennel, James H. (1842), "On useful insects and their products", in Watt, Charles; Watt, John Jr., The Mirror of Literature, Amusement, and Instruction, I, London, United Kingdom: J. Limbird, pp. 295–296, https://books.google.com/books?id=wTo5AQAAMAAJ&pg=PA295 
  35. ↑ 35.0 35.1 35.2 "Vordan Karmir: The Red Worm (From the Soviet Armenia Encyclopedia, Vol. 8, 1982, pp. 642–43)". Oriental Rug Review (Oriental Rug Auction Review) VIII (5): 42–43. 1988. http://www.rugreview.com/orr/8-5-42.htm. Retrieved 9 October 2014. 
  36. ↑ Cardon (2007) cites her own field mission in Armenia in 1989 as well as the papers by Jakubski (1965) and Mktrtchian and Sarkisov (1985) for her description of Porphyrophora hamelii biology, which states that the mating time is from 5 a.m. to 10 a.m. The Soviet Armenian Encyclopedia (1982) and the online Red Book of Armenia (which cites Mktchyan and Sarkisov (1985) and others) state that the mating time is from 6 a.m. to 11 a.m. Note that in 1982 and 1985 the emergence of the insects in early September would have been in Armenian Summer Time (UTC+5), whereas the Republic of Armenia has been on UTC+4 time year-round since 2012.
  37. ↑ "Political Administrative Region (marz): Armavir". http://www.rec-caucasus.org/Flash/PDF/AR/AR7.pdf. 
  38. ↑ "Error: no |title= specified when using {{Cite web}}" (in hy). Government of the Republic of Armenia. July 12, 2003. http://www.arlis.am/DocumentView.aspx?DocID=48076. "1. «Որդան կարմիր» պետական արգելավայրը (այսուհետ` արգելավայր) ստեղծվել է Հայկական Սովետական Սոցիալիստական Հանրապետության Մինիստրների խորհրդի 1987 թվականի փետրվարի 2-ի N 61 որոշմամբ` Հայաստանի Հանրապետության Արմավիրի մարզի աղուտ հողերի վրա: Արգելավայրն զբաղեցնում է 219.85 հեկտար տարածք, բաղկացած է երկու առանձին տեղամասերից` Արազափի գյուղական համայնքի հյուսիսարևմտյան մասում (198.33 հեկտար) և Ջրառատի գյուղական համայնքի հյուսիսային մասում (21.52 հեկտար)` Արարատյան հարթավայրում, ծովի մակերևույթից 835–850 մետր բարձրության վրա:" 
  39. ↑ "Էնդեմիկ Տեսակներ: Կենդանիներ [Endemic Species: Animals"] (in hy), I, Yerevan, Armenia: "Geodeziayi ev Kʻartezagrutʻyan Kentron" POAK, 2007, pp. 83, ISBN 978-99941-0-176-4, http://www.cadastre.am/resources/atlasA/images/83.jpg, retrieved 3 January 2015 
  40. ↑ IUCN Red List Categories and Criteria, Version 3.1, Second Edition (Report). International Union for Conservation of Nature (IUCN) Species Survival Commission. 9 February 2000. pp. 16–17. http://jr.iucnredlist.org/documents/redlist_cats_crit_en.pdf. Retrieved 10 October 2014. 

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