Armenian Cochineal (Porphyrophora hamelii)
If I am asked about the most wonderful animal of Armenia, I would answer without delay - the Armenian cochineal. Indeed, one of the most interesting insects of Armenia is Porphyrophora hamelii, commonly known as Armenian cochineal, or Ararat cochineal (Արարատյան որդան կարմիր). The insect has original looking, remarkable life cycle, and profound impact on our culture.
History of description.
The Armenian cochineal was known from the Ancient times. We can find the very first mentions about it in the works of Ancient Armenian historians. Movses Khorenatsi, the founder of the Armenian History, wrote in V century AD, "Ararat mountain with its valley has abundandce of everything, as well as Gaylot lake and the worm of the roots of reeds for garnishing to red color." Another prominent historian, Ghazar Parpetsi mentioned the cochineal in his History of Armenia as well. The author writes, "Also not without good the welcomed valley of Ararat produced roots of reeds on which a worm is grown used for adornment in red color and for profit and luxury for people." There are many other mentions about the cochineal from the works of Medieval historians, particularly Arab, but it does not seem expedient to quote them all in this posting.
In the contemporary scientific literature, the cochineal appears in 1833 when it was described as a new species by Johann Friedrich von Brandt. Brandt names the species after Joseph Christian Hamel, and that is why the scientific name of the insect is Porphyrophora hamelii (Brandt, 1833). Here is the first description by Brandt. Interesting off-top, when in 1853 Hamel was commanded to the USA, he was forced to give a wriiten promise "not to eat human meat in America" :) Why he was commanded such a stupid order remains unclear. Okay, let's continue to our topic. Hamel receives the insects from Armenia, and makes a lot of historical, linguistic, and biology researh, and in 1833 prints his fundamental work about the Armenian cochineal. The article is in German, and in 1835 it was translated into Russian. Both versions can be found in the end of this posting in the works cited part.
The picture below is taken from the Hamel's article, it is the first known scientific depiction of the cochineal.
Once again let me be distracted. The picture below is taken from Ghevond Alishan's book Ayrarat written in 1890. In the book, we can also find the picture of the insect signed as Red worm, aka Araratian cochineal. As a matter of fact, there is depicted another insect, most probably Mexican cochineal. It seems that Alishan didn't have the actual picture of Armenian cochineal, and used another that was available to him in the remote island of San Lazzaro :)
Taxonomy.
Armenian cochineal is included in the family of Margarodidae, the superfamily of Coccoidea, the suborder of Sternorrhyncha, and the order of the Homoptera. The family of Margarodidae consists, according to the various authors, of about 10 to 70 genera. In Armenia, there are three species of the genus Porphyrophora: Porphyrophora hamelii (Brandt), Porphyrophora tritici (Bodenheimer), and Porphyrophora monticola (Borchsenius). It is worth to notice that the genus and the family in general has not been well researched in Armenia.
Distribution.
Armenian cochineal distributed only on the Armenian Highland, in the territory of Armenia and adjacent areas of Turkey and Iran. In Armenia, its distibution is extremely fragmented. It can be found only on a few patches of the salt marshes on Ararat valley, at altitudes of 800-900 meters above sea level.
The typical landscape where the Armenian cochineal lives.
The Ararat cochineal is a sucking parasite on roots of two plants from the grass family (Poaceae) - Aeluropus littoralis and Phragmites australis.
The close view of the main host plant, Aeluropus littoralis.
Aeluropus littoralis
All parts of the cochineal contain red color pigment - Carminic acid (formula C22H20O13), which the pigment Carmine is produced from. It is suggested that Carminic acid produced by the insects as a mean of defence.
You can appreciate the majectic color of the insects on the picture below.
The insects received their name because of their red color. Porphyrophora translated from Latin means giving porphyry. The term porphyry is from Ancient Greek (πορφύρα porphúra) and means "purple". (Wikipedia). As I noticed in the begining, the Armenian cochineal was used for producing Carmine, the expencive red dye. Carmine has been used by Armenians for centuries to dye textile, carpets, paints. Now, It is used as color additive for food. There are some rumors that one of the secret ingredients of Coca Cola is Carmine. In addition, there are some mentions about the medicinal using of Carmine in medieval Armenia. Surprisingly, in the nature, there can be found (although very rare - frequency 1 out of 100,000) absolutely white individuals, albinos. (Mkrtchyan L. P., Sarkisov R. N., 1985).
Porphyrophora hamelii (female)
Morphology and life cycle.
Araratian cochineal is characterized by the sexual dimorphism. That means that males and females have totally different morphology. Another interesting fact is that insects go through some stages before the maturation, and in the early stages, males and females are very similar. There is suggestion that females are kind of stay "frozen" in the larval stage during the evolution.
Cochineals spend almost whole life underground. Only in the fall, during a couple of weeks, the adults crawl up to the surface in the morning hours for just an hour ot two to mate. That time is the only time that we can observe them on the ground and appreciate their uniqueness. Males die on the same day. Fertilized females dig into the soil, lay eggs, and die soon thereafter. Armenian cochineal is charachterized by both bisexual and parthenogenetic development. Diploid set of females' chromosome is 14, while males is 13. The sexual differentiation system is XX-X0.
The ova of the cochineal are elongated, dark-red in color, 0.4-0.7 mm in length and 0.2-0.3 mm in width. They are laid inside the ovum sac. In one sac, there are about 500 ova.
Porphyrophora hamelii (оvorum saccum)
The first-instar nymphs, or crawlers hatch in the spring, in the April-May. They crawl (hence their name) until find the host plant and attach to it. The body of the crawlers is elongated, dark-red in color, 0.2-0.8 mm long and 0.2 mm wide. The body is more or less segmented. Antennae are 6-segmented. The last segment of the body has long hair-like structures called caudal setae. The first-instar nymphs feed on the roots of the host plant, change the form of the body from elongated to pear-shape and become the second-instar nymphs.
The second-instar nymphs differentiate from the first-instars by the body shape and that their body is not segmented. They are also red colored, 0.7 mm long and 0.3-0.8 mm wide. Antennae are 6-segmented. Unfortunately, I dont have the actual pictures of the first and second-instar nymphs. In the end of May, the second-instar nymphs transform into the cysts
The third-instar nymphs, or cysts have apodous (without legs and antennae) wide-oval body, with hard skin, 1-3 mm long and 1-2 mm wide. Below is the picture of a cyst attached to the root of the host plant.
Porphyrophora hamelii (larval cysta)
Some third-instar nymphs increase in size and become the female four-instar nymphs.
The female four-instar nymph is also a cyst similar to the previous stage cyst but with a larger body. The length of the body is 3-7 mm, and the widht is 3-5 mm.
Porphyrophora hamelii (larval cysta)
In the end of August-beginning of September from the other cysts come out to the surface the male four-instar nymphs.
The male four-instar nymphs are mobile, 2-5 mm long and 1-3 mm wide. The body is segmented, skin is soft, antennae are 11-12 segmented. As you will see later, the male four-instar nymphs are very similar to the adult females! Those nymphs can be distinguish from the adult females by the mean of size and body shape. The nymphs are a little bit smaller and have a slightly flatten body. Otherwise, they cannot be separated from the adult females! After a few days, the nymphs dig in the soil and transform into the adults. Again sadly, I don't have the actual pictures of the male four-instar nymphs.
While male nymphs go through some transfpormations, their female counterparts continue feeding on the host plants' roots and accumulate energy for the reproduction. In September with the first rays of the sun, the matured insects come out to the surface.
The adult females are oval, wingless, lethargic dark-red colored insects. The body is segmented, with the soft skin. Mouthparts absent, that is to say they don't feed. The eyes are simple, antennae are 11-12-segmented. The length of the body is 2-12 mm, with the width of 1-6 mm. The average weight of the insect is 27 mg.
Porphyrophora hamelii (female), ventral and dorsal view
Prothoracic legs are fossorial, highly developed with strengthened claws. They are required for burrowing down into the soil.
Porphyrophora hamelii (female), detail of the leg.
The adult males, in contrast, are winged, very active insects 2-4 mm long and 0.5-1 mm wide. The average weight is only 2 mg. Forewings are transparent with the purple rim on the anterior side, legs are well developed. Eyes are compound. Antennae are 13-segmented. From the end of the body, there are long hair-like waxy filaments.
Porphyrophora hamelii (male)
As it was noticed earlier, the insects come out to the surface in the morning to mate. Below are the intriguing pictures of the copulation.
Porphyrophora hamelii (in copula)
Porphyrophora hamelii (in copula)
No less fascinating video of mating.
After mating, the male goes to another female, while the fertilized female burrows down into the soil. If a female is not fertilized, it digs into the soil as well, but on the next day appears to the surface again for the another try.
The mated female produces white silky ovisac. Here it is.
Porphyrophora hamelii (оvorum saccum)
In this way, the development of the cochineal is complited and repeated every year.
A couple of words about cochineal's enemies. In the nature there are many enemies of the cochineal, such as ticks (Tyrophagus putrescentiae), beetles (Nephus bipunctatus, Lithophilus bipustulatus, Sepedophilus bipustulatus), ants (Tapinoma erraticum, Messor sp., Pheidole pallidula), and some birds (common starling, crested lark). On the picture below is shown the regular foe of the cochineal - an ant.
In the last edition of the Red Book of Armenia, cochineal is registered under the Critically Endangered status, hence it highly needs to be protected.
One more video about our Porphyry adornment :)
I used mostly the two books: Mkrtchyan L. P., Sarkisov R. N. (1985) The biology and reproduction of the Ararat cochineal Porphyrophora hamelii Brandt (Homoptera, Coccinea, Margarodidae), Yerevan, USSR, 157 pp., (in Russian) and Ter-Grigoryan M. A. (1976) The morphology of Ararat cochineal Porphyrophora hamelii. In: Biological Journal of Armenia, v. XXIX, 3, (in Russian).
Sources:
History of description.
The Armenian cochineal was known from the Ancient times. We can find the very first mentions about it in the works of Ancient Armenian historians. Movses Khorenatsi, the founder of the Armenian History, wrote in V century AD, "Ararat mountain with its valley has abundandce of everything, as well as Gaylot lake and the worm of the roots of reeds for garnishing to red color." Another prominent historian, Ghazar Parpetsi mentioned the cochineal in his History of Armenia as well. The author writes, "Also not without good the welcomed valley of Ararat produced roots of reeds on which a worm is grown used for adornment in red color and for profit and luxury for people." There are many other mentions about the cochineal from the works of Medieval historians, particularly Arab, but it does not seem expedient to quote them all in this posting.
In the contemporary scientific literature, the cochineal appears in 1833 when it was described as a new species by Johann Friedrich von Brandt. Brandt names the species after Joseph Christian Hamel, and that is why the scientific name of the insect is Porphyrophora hamelii (Brandt, 1833). Here is the first description by Brandt. Interesting off-top, when in 1853 Hamel was commanded to the USA, he was forced to give a wriiten promise "not to eat human meat in America" :) Why he was commanded such a stupid order remains unclear. Okay, let's continue to our topic. Hamel receives the insects from Armenia, and makes a lot of historical, linguistic, and biology researh, and in 1833 prints his fundamental work about the Armenian cochineal. The article is in German, and in 1835 it was translated into Russian. Both versions can be found in the end of this posting in the works cited part.
The picture below is taken from the Hamel's article, it is the first known scientific depiction of the cochineal.
Once again let me be distracted. The picture below is taken from Ghevond Alishan's book Ayrarat written in 1890. In the book, we can also find the picture of the insect signed as Red worm, aka Araratian cochineal. As a matter of fact, there is depicted another insect, most probably Mexican cochineal. It seems that Alishan didn't have the actual picture of Armenian cochineal, and used another that was available to him in the remote island of San Lazzaro :)
Taxonomy.
Armenian cochineal is included in the family of Margarodidae, the superfamily of Coccoidea, the suborder of Sternorrhyncha, and the order of the Homoptera. The family of Margarodidae consists, according to the various authors, of about 10 to 70 genera. In Armenia, there are three species of the genus Porphyrophora: Porphyrophora hamelii (Brandt), Porphyrophora tritici (Bodenheimer), and Porphyrophora monticola (Borchsenius). It is worth to notice that the genus and the family in general has not been well researched in Armenia.
Distribution.
Armenian cochineal distributed only on the Armenian Highland, in the territory of Armenia and adjacent areas of Turkey and Iran. In Armenia, its distibution is extremely fragmented. It can be found only on a few patches of the salt marshes on Ararat valley, at altitudes of 800-900 meters above sea level.
The typical landscape where the Armenian cochineal lives.
The Ararat cochineal is a sucking parasite on roots of two plants from the grass family (Poaceae) - Aeluropus littoralis and Phragmites australis.
The close view of the main host plant, Aeluropus littoralis.
Aeluropus littoralis
All parts of the cochineal contain red color pigment - Carminic acid (formula C22H20O13), which the pigment Carmine is produced from. It is suggested that Carminic acid produced by the insects as a mean of defence.
You can appreciate the majectic color of the insects on the picture below.
The insects received their name because of their red color. Porphyrophora translated from Latin means giving porphyry. The term porphyry is from Ancient Greek (πορφύρα porphúra) and means "purple". (Wikipedia). As I noticed in the begining, the Armenian cochineal was used for producing Carmine, the expencive red dye. Carmine has been used by Armenians for centuries to dye textile, carpets, paints. Now, It is used as color additive for food. There are some rumors that one of the secret ingredients of Coca Cola is Carmine. In addition, there are some mentions about the medicinal using of Carmine in medieval Armenia. Surprisingly, in the nature, there can be found (although very rare - frequency 1 out of 100,000) absolutely white individuals, albinos. (Mkrtchyan L. P., Sarkisov R. N., 1985).
Porphyrophora hamelii (female)
Morphology and life cycle.
Araratian cochineal is characterized by the sexual dimorphism. That means that males and females have totally different morphology. Another interesting fact is that insects go through some stages before the maturation, and in the early stages, males and females are very similar. There is suggestion that females are kind of stay "frozen" in the larval stage during the evolution.
Cochineals spend almost whole life underground. Only in the fall, during a couple of weeks, the adults crawl up to the surface in the morning hours for just an hour ot two to mate. That time is the only time that we can observe them on the ground and appreciate their uniqueness. Males die on the same day. Fertilized females dig into the soil, lay eggs, and die soon thereafter. Armenian cochineal is charachterized by both bisexual and parthenogenetic development. Diploid set of females' chromosome is 14, while males is 13. The sexual differentiation system is XX-X0.
The ova of the cochineal are elongated, dark-red in color, 0.4-0.7 mm in length and 0.2-0.3 mm in width. They are laid inside the ovum sac. In one sac, there are about 500 ova.
Porphyrophora hamelii (оvorum saccum)
The first-instar nymphs, or crawlers hatch in the spring, in the April-May. They crawl (hence their name) until find the host plant and attach to it. The body of the crawlers is elongated, dark-red in color, 0.2-0.8 mm long and 0.2 mm wide. The body is more or less segmented. Antennae are 6-segmented. The last segment of the body has long hair-like structures called caudal setae. The first-instar nymphs feed on the roots of the host plant, change the form of the body from elongated to pear-shape and become the second-instar nymphs.
The second-instar nymphs differentiate from the first-instars by the body shape and that their body is not segmented. They are also red colored, 0.7 mm long and 0.3-0.8 mm wide. Antennae are 6-segmented. Unfortunately, I dont have the actual pictures of the first and second-instar nymphs. In the end of May, the second-instar nymphs transform into the cysts
The third-instar nymphs, or cysts have apodous (without legs and antennae) wide-oval body, with hard skin, 1-3 mm long and 1-2 mm wide. Below is the picture of a cyst attached to the root of the host plant.
Porphyrophora hamelii (larval cysta)
Some third-instar nymphs increase in size and become the female four-instar nymphs.
The female four-instar nymph is also a cyst similar to the previous stage cyst but with a larger body. The length of the body is 3-7 mm, and the widht is 3-5 mm.
Porphyrophora hamelii (larval cysta)
In the end of August-beginning of September from the other cysts come out to the surface the male four-instar nymphs.
The male four-instar nymphs are mobile, 2-5 mm long and 1-3 mm wide. The body is segmented, skin is soft, antennae are 11-12 segmented. As you will see later, the male four-instar nymphs are very similar to the adult females! Those nymphs can be distinguish from the adult females by the mean of size and body shape. The nymphs are a little bit smaller and have a slightly flatten body. Otherwise, they cannot be separated from the adult females! After a few days, the nymphs dig in the soil and transform into the adults. Again sadly, I don't have the actual pictures of the male four-instar nymphs.
While male nymphs go through some transfpormations, their female counterparts continue feeding on the host plants' roots and accumulate energy for the reproduction. In September with the first rays of the sun, the matured insects come out to the surface.
The adult females are oval, wingless, lethargic dark-red colored insects. The body is segmented, with the soft skin. Mouthparts absent, that is to say they don't feed. The eyes are simple, antennae are 11-12-segmented. The length of the body is 2-12 mm, with the width of 1-6 mm. The average weight of the insect is 27 mg.
Porphyrophora hamelii (female), ventral and dorsal view
Prothoracic legs are fossorial, highly developed with strengthened claws. They are required for burrowing down into the soil.
Porphyrophora hamelii (female), detail of the leg.
The adult males, in contrast, are winged, very active insects 2-4 mm long and 0.5-1 mm wide. The average weight is only 2 mg. Forewings are transparent with the purple rim on the anterior side, legs are well developed. Eyes are compound. Antennae are 13-segmented. From the end of the body, there are long hair-like waxy filaments.
Porphyrophora hamelii (male)
As it was noticed earlier, the insects come out to the surface in the morning to mate. Below are the intriguing pictures of the copulation.
Porphyrophora hamelii (in copula)
Porphyrophora hamelii (in copula)
No less fascinating video of mating.
Click to viewAfter mating, the male goes to another female, while the fertilized female burrows down into the soil. If a female is not fertilized, it digs into the soil as well, but on the next day appears to the surface again for the another try.
The mated female produces white silky ovisac. Here it is.
Porphyrophora hamelii (оvorum saccum)
In this way, the development of the cochineal is complited and repeated every year.
A couple of words about cochineal's enemies. In the nature there are many enemies of the cochineal, such as ticks (Tyrophagus putrescentiae), beetles (Nephus bipunctatus, Lithophilus bipustulatus, Sepedophilus bipustulatus), ants (Tapinoma erraticum, Messor sp., Pheidole pallidula), and some birds (common starling, crested lark). On the picture below is shown the regular foe of the cochineal - an ant.
In the last edition of the Red Book of Armenia, cochineal is registered under the Critically Endangered status, hence it highly needs to be protected.
One more video about our Porphyry adornment :)
I used mostly the two books: Mkrtchyan L. P., Sarkisov R. N. (1985) The biology and reproduction of the Ararat cochineal Porphyrophora hamelii Brandt (Homoptera, Coccinea, Margarodidae), Yerevan, USSR, 157 pp., (in Russian) and Ter-Grigoryan M. A. (1976) The morphology of Ararat cochineal Porphyrophora hamelii. In: Biological Journal of Armenia, v. XXIX, 3, (in Russian).
Sources:
- Brandt J. F. (1833) Zur Naturgeschichte der Cochenille. Ueber die Cochenille Armeniens. In Brandt J. F., Ratzeburg J. T. C. (1833) Medizinische Zoologie, pp. 355-356.
- Nerses Archbishop (1833) About Ararat cochineal. In: Collection of reviews related to Armenian history, v. 2, 1838. (in Russian)
- Hamel J. (1835) Über Cochenille am Ararat und über Wurzelcochenille im Allgemeinen. In Mémoires de l'Académie impériale des sciences de St.-Pétersbourg. sér. 6, pp. 9-64.
- Hamel J. (1835) About Ararat cochineal; An abstract from works of Hamel, printed in the Memoirs of the Imperial Academy of Sciences (in Russian).
- Chopin I. (1852) Cochineal. In: Historical state and condition of Armenian province (in Russian).
- Alishan G. (1890) Cochineal from Masis. In: Ayrarat (in Armenian).
- Porphyrophora hameli. In: Blanchard R. (1883) Les Coccides Utiles. In Bulletin de la Societe Zoologique de France.
- Bekhovsepyan L. (1932) Ararat Cochineal and the Perspectives of Its Industrial Using. Socialist Agronomy, vol. 1 (in Armenian).
- Borchsenius N. S. (1950) Genus Porphyrophora. In: Coccoidea of USSR (in Russian).
- Ter-Grigoryan M. A. (1976) The morphology of Ararat cochineal Porphyrophora hamelii. In: Biological Journal of Armenia, v. XXIX, 3, (in Russian).
- Mkrtchyan L. P., Sarkisov R. N. (1985) The biology and reproduction of the Ararat cochineal Porphyrophora hamelii Brandt (Homoptera, Coccinea, Margarodidae), Yerevan, USSR, 157 pp. (in Russian).
- Sarkisov R. N. et all (1991) Ararat scale and its reproduction ex situ. Yerevan (in Armenian).
- Ben-Dov Y (1993) A Systematic Catalogue of the Soft Scale Insects of the World (Homoptera: Coccoidea: Coccidae). Gainesville, FL: Sandhill Crane Press, 536 pp.
- Vahedi H. A. (2002) A Revision of the Genus Porphyrophora (Hemiptera: Coccoidea: Margarodidae) with Particular Reference to the Middle East and with a Discussion on the Relationships of the Hypogaeic Margarodids. Ph.D. thesis. Department of Agricultural Sciences, Imperial College at Wye, Wye, Ashford, Kent, UK, 358 pp.
- Foldi I. (2005) A generic revision of Margarodidae. In: Annales de la Société Entomologique de France (Nouvelle série).
- Porphyrophora hamelii In: García Morales M., Denno B. D., Miller D. R., Miller G. L., Ben-Dov Y., Hardy N.B. (2016) ScaleNet: A literature-based model of scale insect biology and systematics. Database. doi: 10.1093/database/bav118. http://scalenet.info.