Hostname: page-component-cd9895bd7-mkpzs Total loading time: 0 Render date: 2024-12-26T16:56:52.608Z Has data issue: false hasContentIssue false
  • English
  • Français

The Anopheles hyrcanus Group in South-East Asia (Diptera: Culicidae)

Published online by Cambridge University Press:  10 July 2009

J. A. Reid
J. A. Reid
Affiliation:
Entomologist, Institute for Medical Research, Kuala Lumpur, Malaya.
Get access Rights & Permissions [Opens in a new window]

Extract

This paper is a study of the southeast Asian forms of the Anopheles hyrcanus complex with special reference to those of the Malay Peninsula, and it is to the latter forms that the detailed descriptions apply. The group is distinguished from all but a few other species by the presence in the adult female of a tuft of scales on the clypeus on each side, together with pale bands on the palps and a ventral tuft of scales on the seventh abdominal segment. For a complete diagnosis see page 6.

The usual practice has been to recognise only two oriental forms of hyrcanus: varieties sinensis and nigerrimus. It is shown that these varieties as recognised up till now are heterogeneous, and that at least in southeast Asia, ‘hyrcanus’, is a group of sibling species of which there are no less than seven or eight in the Malay Peninsula. For convenience, and where the exact species is not known, the name ‘hyrcanus’ in inverted commas is used, meaning hyrcanus sibling species group.

There is no doubt that these sympatric forms are distinct species for although individual characters, such as the width of the pale bands on the hind tarsi, are variable and the range of variation between two or more species may overlap, by using a combination of characters there is no difficulty in identifying individual adult specimens of either sex. Furthermore, offspring reared from eggs are always of the same form as the female parent. Larvae, pupae, and eggs can usually be identified, and keys are given for the identification of these stages as well as the adults.

Seven named species are described, but only one of these, A. crawfordi, is regarded as new, the others have all been described at various times, and the names later sunk as synonyms of sinensis and nigerrimus.

All seven species have been recognised also outside the Malay Peninsula, and some have a wide range from western India to the Philippines or Moluccas. Except for sinensis, the species belong to southeast Asia, where Assam, Burma, Siam, the Malay Peninsula and Sumatra support the largest number. A. sinensis also occurs in these countries, but the centre of its range is more to the northeast towards China and it does not appear to have penetrated further into the archipelago than the Malay Peninsula and Sumatra. Towards the limits of the Oriental region in northwest India and central and northern China, the southeast Asian species of hyrcanus are few or absent, and are replaced by other forms of hyrcanus of palaearctic type. In China, the name sinensis probably covers two overlapping forms, a southerly oriental one presumed to be Wiedemann's type form which lays a wide decked egg, and a northerly palaearctic form laying a narrow decked egg.

Species of which sufficient numbers have been seen from different countries show geographical variation. The effects of this variation are most marked in the relatively isolated Philippine Islands, where out of three forms occurring there, one, pseudosinensis, is regarded as a species peculiar to the Philippines, though clearly derived from the same stock as the widespread nigerrimus which does not occur there. A second, lesteri, is probably a subspecies of the form of lesteri found in Borneo and the Malay Peninsula. The group thus seems to conform to the classical pattern of speciation by geographical isolation. Much of the evolution and distribution of the species probably took place in pleistocene times.

Broadly speaking A. ‘hyrcanus’ can be described as a swamp breeder, with a preference for animal blood. But it is evident that the individual species show important differences in their biology. Some breed in sunlit waters and their larvae are often green; others seem to prefer shaded places, though not in jungle, and their larvae are usually dark coloured. A. sinensis and lesteri at least, can breed in moderately saline waters, sometimes in company with A. sundaicus and baezai.

It appears that most of the species will bite man freely out of doors shortly after dark, but are reluctant to enter rooms in search of human blood. However, the degree of reluctance to bite man probably varies to an important extent between different species. All species can usually be captured at night at cattle sheds. The adults do not often rest indoors by day, and the outdoor resting places of indiensis and lesteri are described.

In southeast Asia, though ‘ hyrcanus ’ is not one of the principal mosquito vectors of malaria or filariasis, it has been found infected on a number of occasions. In most cases the exact species cannot be determined now, but where this is possible, it has been A. nigerrimus Giles in every instance. Venhuis (1939) showed that his A. hyrcanus var. X (=nigerrimus) was a vector of malaria in Celebes and Java, and Kariadi (1941) implicated it in the transmission of malayan filariasis in southeast Borneo. In Malaya, examination of preserved specimens showed that nigerrimus was the principal or only vector during an outbreak of malaria at Kuala Lumpur which Hodgkin (1933) had shown to be due to hyrcanus, and recently nigerrimus has been found to play a part in the transmission of malayan filariasis in Kedah and Province Wellesley. Experimental infections with Wuchereria malayi, the worm which causes malayan filariasis, showed that nigerrimus was less readily infected than the principal vectors (Mansonia spp.), and some other members of the hyrcanus group, but that it bit the human carrier more readily than the latter. A. sinensis, though probably harmless in Malaya, is a vector of malaria and bancroftian filariasis in parts of China, but the exact identification of the vector is now in doubt, since there appear to be at least two forms in China included under the name sinensis.

Type
Original Articles
Information
Bulletin of Entomological Research , Volume 44 , Issue 1 , May 1953 , pp. 5 - 76
Copyright
Copyright © Cambridge University Press 1953

Access options

Get access to the full version of this content by using one of the access options below. (Log in options will check for institutional or personal access. Content may require purchase if you do not have access.)

References

D'Abrera, V. St. E. (1944). The eggs of the Ceylon Anopheline mosquitoes.—J. Malar. Inst. India, 5, pp. 337–359.Google Scholar
Baisas, F. E. (1931). The barbirostris-hyrcanus group of the Philippine Anopheles.—Philipp. J. Sci., 44, pp. 425–448.Google Scholar
Baisas, F. E. & Hu, S. M. K. (1936). Anopheles hyrcanus var. sinensis of the Philippines and certain parts of China, with some comments on Anopheles hyrcanus var. nigerrimus of the Philippines.—Mon. Bull. Bur. Hlth Philipp., 16, pp. 205–242.Google Scholar
Barraud, P. J. & Christophers, S. R. (1931). On a collection of Anopheline and Culicine mosquitoes from Siam.—Rec. Malar. Surv. India, 2, pp. 269–285.Google Scholar
Bates, M., Beklemishev, W. N. & La Face, L. (1949). Anophelines of the Palearctic region.— In Boyd, M. F.Ed. Malariology, 1, pp. 419–442. Philadelphia, Pa., Saunders.Google Scholar
Bentley, A. (1902). Anopheles mosquitoes in Tezpur, Assam.—Indian med. Gaz., 37, pp. 15–16.Google ScholarPubMed
Bonne-Wepster, J. (1951). Anopheles venhuisi n. sp.—Docum. neerl. indones. Morb. trop., 3, p. 284.Google Scholar
Brug, S. L. & Bonne-Wepster, J. (1947). The geographical distribution of the mosquitoes of the Malay Archipelago.—Chron. nat., 103, 19 pp.Google Scholar
Chow, C. Y. (1950). Mosquito studies in China, past and present.—Mosq. News, 10, pp. 134–137.Google Scholar
Christophers, S. R. (1924). Provisional list and reference catalogue of the Anophelini.—Indian med. Res. Mem., 3, 105 pp.Google Scholar
Christophers, S. R. (1933). The fauna of British India. Diptera, Vol. IV. Family Culicidae. Tribe Anophelini.—371 pp. London.Google Scholar
Christophers, S. R. & Barraud, P. J. (1931). The eggs of Indian Anopheles, with descriptions of the hitherto undescribed eggs of a number of species.—Rec. Malar. Surv. India, 2, pp. 61–192.Google Scholar
Christophers, S. R. & Chand, K. (1915). Notes on some Anophelines from Arabia and Mesopotamia.—Indian J. med. Res., 3, pp. 180–200.Google Scholar
Colless, D. H. (1948). The Anopheline mosquitoes of north-west Borneo.—Proc. Linn. Soc. N.S.W., 73, pp. 71–119.Google Scholar
Covell, G. (1944). Notes on the distribution, breeding places, adult habits, and relation to malaria of the Anopheline mosquitoes of India and the Far East.—J. Malar. Inst. India, 5, pp. 399–434.Google Scholar
Crawford, R. (1938). Some Anopheline pupae of Malaya with a note on pupal structure.—110 pp. Singapore, Govt S.S. & Malar. Adv. Bd F.M.S.Google Scholar
Dobby, E. G. H. (1950). Southeast Asia. London.Google Scholar
Edwards, F. W. (1921). A revision of the mosquitos of the Palaearctic region.—Bull. ent. Res., 18, pp. 263–351.CrossRefGoogle Scholar
Edwards, F. W. (1929). Mosquito notes. VIII.—Bull. ent. Res., 20, pp. 321–343.CrossRefGoogle Scholar
Edwards, F. W. (1932). Diptera. Fam. Culicidae.—Genera Insect., 184, 245 pp. Brussels.Google Scholar
Evans, A. M. (1938). Mosquitoes of the Ethiopian region. II. Anophelini.—404 pp. London, Brit. Mus. (Nat. Hist.).Google Scholar
Feng, L. C. (1938). A critical review of the literature regarding the records of mosquitoes in China.—Peking nat. Hist. Bull., 12, pp. 169–181, 285–318.Google Scholar
Fowler, H. W. (1949). A dictionary of modern English usage.—London.Google Scholar
Gater, B. A. R. (1933). Notes on Malayan mosquitoes, I. The genus Anopheles.—Malay. med. J., 8, pp. 39–42.Google Scholar
Gater, B. A. R. (1935). Aids to the identification of Anopheline imagines in Malaya.—242 pp. Singapore, Govt. S.S. & Malar. Adv. Bd F.M.S.Google Scholar
Giles, G. M. (1900). A handbook of the gnats or mosquitoes.—374 pp. London.Google ScholarPubMed
Giles, G. M. (1902). A handbook of the gnats or mosquitoes. 2nd edn., 530 pp.Google Scholar
Giles, G. M. (1904). A revision of the Anophelinae.—48 pp. London.Google Scholar
van Hell, J. C. (1950). De betekenis van A. (A.) hyrcanus var. X als malaria-overbrenger op Zuid-Celebes.—Med. Maandbl., 3, pp. 557–567. In English in Docum. neerl. indones. Morb. trop., 3, pp. 373–380.Google Scholar
Ho, Ch'i. (1938). On a collection of Anopheline mosquitoes from the island of Hainan.—Ann. trop. Med. Parasit., 32, pp. 387–411.CrossRefGoogle Scholar
Hodgkin, E. P. (1933). Anopheles hyrcanus Pallas as a malaria carrier in Malaya.—Bull. Inst. med. Res. F.M.S. no. 1 of 1932, pp. 1–6.Google Scholar
Hodgkin, E. P. (1937). Division of Entomology.—Rep. Inst. med. Res. F.M.S., 1936, pp. 79–92.Google Scholar
Hodgkin, E. P. (1939). Division of Entomology.—Rep. Inst. med. Res. F.M.S., 1938, pp. 64–79.Google Scholar
Hodgkin, E. P. (1940). The breeding of certain species of Anopheles in saline waters.—Trans Far-East Ass. trop. Med., 10th Congr., 2, pp. 839–871.Google Scholar
Hodgkin, E. P. & Rajamoney, P. D. (1933). A descriptive and biological note on the Malayan varieties of Anopheles hyrcanus Pallas.—Bull. Inst. med. Res. F.M.S., no. 1 of 1933, pp. 7–18.Google Scholar
Institute for Medical Research, Malaya. (1950). Division of Entomology.—Rep. Inst. med. Res. Malaya, 1949, pp. 25–32.Google Scholar
Institute for Medical Research, Malaya. (1951). Division of Entomology.—Rep. Inst. med. Res. Malaya, 1950, pp. 14–20.Google Scholar
James, S. P. & Liston, W. G. (1904). A monograph of the Anopheline mosquitoes of India. Calcutta.CrossRefGoogle Scholar
James, S. P. & Liston, W. G. (1911). A monograph of the Anopheline mosquitoes of India., 2nd edn., 128 pp.CrossRefGoogle Scholar
James, S. P. & Stanton, A. T. (1912). Revision of the names of Malayan Anophelines.—Paludism, 5, pp. 59–63.Google Scholar
Kariadi, S. P. (1941). A. hyrcanus X en filariasis malayi te Martapoera.—Geneesk. Tijdschr. Ned.-Ind., 81, pp. 107–118.Google Scholar
Knight, K. L. & Chamberlain, R. W. (1948). A new nomenclature for the chaetotaxy of the mosquito pupa, based on a comparative study of the genera (Diptera: Culicidae).—Proc. helminth. Soc. Wash., 15, pp. 1–10.Google Scholar
Lamborn, W. A. (1921). Federated Malay States Malaria Bureau Report, 1920.—Suppl. F.M.S. Govt. Gaz., 4th 11 1921, pp. 8–13.Google Scholar
Lamborn, W. A. (1922 a). The mosquitos of some ports of China and Japan.—Bull. ent. Res., 12, pp. 401–409.CrossRefGoogle Scholar
Lamborn, W. A. (1922 b). The bionomics of some Malayan Anophelines.—Bull. ent. Res., 13, pp. 129–149.CrossRefGoogle Scholar
Laveran, A. (1902). Sur des culicides du Cambodge.—C.R. Soc. Biol., Paris, 54, pp. 906–908.Google Scholar
Leicester, G. F. (1908). The Culicidae of Malaya.—Stud. Inst. med. Res. F.M.S., 3, no. 3, pp. 18–261.Google Scholar
McArthur, J. (1950). Malaria and its vectors in Borneo.—Indian J. Malariol., 4, pp. 1–90.Google ScholarPubMed
Mayr, E. (1944). Wallace's line in the light of recent zoogeographic studies.—Quart. Rev. Biol., 19, pp. 1–14; also in Science and Scientists in the Netherlands Indies. New York, 1945.CrossRefGoogle Scholar
Mayr, E. (1947). Systematics and the origin of species.—3rd edn.New York.Google Scholar
Otsuru, M. & Miyagawa, M. (1950). The eggs of Japanese Anopheles.—Jap. J. sanit. Zool., 1, pp. 3–4 (Abstract only seen).CrossRefGoogle Scholar
Pittendrigh, C. S. (1950). The ecotype specialization of Anopheles homunculus, and its relation to competition with A. bellator.—Evolution, 4, pp. 64–78.CrossRefGoogle Scholar
Poynton, J. O. & Hodgkin, E. P. (1938). Endemic filariasis in the Federated Malaya States.—Bull. Inst. med. Res. F.M.S., no. 1 of 1938, 67 pp.Google Scholar
Reid, J. A. (1947). Type specimens of Culicidae described by Laveran. (Diptera: Culicidae).—Proc. R. ent. Soc. Lond., (B) 16, pp. 86–91.Google Scholar
Reid, J. A. (1949). A preliminary account of the forms of Anopheles leucosphyrus Dönitz (Diptera: Culicidae).—Proc. R. ent. Soc. Lond., (B) 18, pp. 42–53.Google Scholar
Reid, J. A. (1950). Some new records of Anopheline mosquitoes from the Malay Peninsula with remarks on geographical distribution.—Bull. Raffles Mus., 21, pp. 48–58.Google Scholar
Reid, J. A. & Hodgkin, E. P. (1950). The Anopheles umbrosus group (Diptera: Culicidae).—Trans. R. ent. Soc. Lond., 101, pp. 281–334.CrossRefGoogle Scholar
Russell, P. F. & Baisas, F. E. (1934). A practical illustrated key to the larvae of Philippine Anopheles.—Philipp. J. Sci., 55, pp. 307–336.Google Scholar
Russell, P. F. & Baisas, F. E. (1936). A practical illustrated key to adults of Philippine Anopheles.—Philipp. J. Sci., 58, pp. 15–64.Google Scholar
Scrivenor, J. B. & others. (1943). A discussion on the biogeographic division of the Indo-Australian archipelago, with criticism of the Wallace and Weber lines…—Proc. Linn. Soc. Lond., 154, pp. 120–165.CrossRefGoogle Scholar
van Someren, E. C. C. (1947). The description of a new Anopheles of the Myzorhynchus series from Madagascar, with notes on its systematic position in relation to the Ethiopian species of this group.—E. Afr. med. J., 24 pp. 42–46.Google Scholar
Stanton, A. T. (1915). Notes on Sumatran Culicidae.—Indian J. med. Res., 3, pp. 251–258.Google Scholar
Stanton, A. T. & Hacker, H. P. (1917). The Anopheles of Malaya. III. A new variety of A. albotaeniatus, Theo..—Bull. ent. Res., 7, pp. 273–275.CrossRefGoogle Scholar
Stoker, W. J. (1931). Anopheles montanus (Anopheles albotaeniatus var. montanus Stanton and Hacker, 1917).—Meded. Dienst Volksgezondh. Ned.-Ind., 20, pp. 129–132.Google Scholar
Sweet, W. C., Feng, L. C., Chow, C. Y. & Hsu, S. C. (1942). Anophelines of southwestern Yunnan and their relation to malaria.—J. nat. Malar. Soc., 1, pp. 25–32.Google Scholar
Swellengrebel, N. H. (1914). Een nieuwe anopheline voor Deli, Myzorhynchus argyropus n. sp.—Geneesk. Tijdschr. Ned.-Ind., 54, p. 334.Google Scholar
Swellengrebel, N. H. (1921). De Anophelinen van Nederlandsch Oost-Indië. 2nd edn.—Meded. kolon. Inst. Amst., (Trop. Hyg.) no. 10, 155 pp.Google Scholar
Swellengrebel, N. H. & Rodenwaldt, E. (1932). Die Anophelen von Niederländisch-Ostindien.—242 pp. Jena, Fischer.Google Scholar
Theobald, F. V. (1901). A monograph of the Culicidae, 1. London, Brit. Mus. (Nat. Hist.).Google Scholar
Theobald, F. V. (1903). A monograph of the Culicidae, 3.Google Scholar
Theobald, F. V. (1907). A monograph of the Culicidae, 4.Google Scholar
Thomson, R. C. M. (1940). Studies on the behaviour of Anopheles minimus. Part III. The influence of water temperature on the choice and suitability of the breeding place.—J. Malar. Inst. India, 3, pp. 323–348.Google Scholar
Thomson, R. C. M. (1951). Studies on salt-water and fresh-water Anopheles gambiae on the East African coast.—Bull. ent. Res., 41, pp. 487–502.CrossRefGoogle Scholar
Venhuis, W. G. (1939). The hyrcanus problem in the Netherlands East Indies, with description of a widespread malaria carrying variety: An. hyrcanus X. (First communication).—Meded. Dienst Volksgezondh. Ned.-Ind., 28, pp. 376–389.Google Scholar
Walch, E. W. (1930). The larva of Anopheles peditaeniatus (Leicester).—Meded. Dienst Volksgezondh. Ned.-Ind., 19, pp. 44–45.Google Scholar
Walch, E. W. & Walch-Sorgdrager, G. B. (1935). The eggs of some Netherlands-Indian Anophelines.—Trans. Far East Ass. trop. Med., 9th Cong., pp. 65–81. In Dutch in Geneesk. Tijdschr. Ned.-Ind., 75, pp. 1700–1730, with plates.Google Scholar
Wharton, R. H. (1950). Daytime resting places of Anopheles maculatus and other Anophelines in Malaya, with results of precipitin tests.—Med. J. Malaya, 4, pp. 260–271.Google Scholar
Wiedemann, C. R. W. (1828). Aussereuropäische zweiflügelige Insekten, 1, p. 547. Hamm.Google Scholar
Wigglesworth, V. B. (1942). The principles of insect physiology. 2nd edn.London, Methuen.Google Scholar
Wilson, T. & Reid, J. A. (1951). Filariasis.—Stud. Inst. med. Res. Malaya, 25, pp. 209–227.Google Scholar
Yamada, M. (1927). A new species of Anopheles in Chosen (Korea).—Keijo J. Med., 8, pp. 237–255.Google Scholar
Yamada, S. (1924). A revision of the adult Anopheline mosquitoes of Japan. Part 1.—Sci. Rep. Inst. infect. Dis. Tokyo Univ., 1924, 3, pp. 215–241.Google Scholar
Yao, Y. T. & Wu, C. C. (1935). One year's observation of Anopheles hyrcanus var. sinensis in Nanking, 1933.—Trans. Far-East Ass. trop. Med., 9th Congr., 2, pp. 3–26.Google Scholar
Yao, Y. T. & Wu, C. C. (1936). Some abnormalities of the morphology of the male hypopygia of Anopheles hyrcanus var. sinensis Wied., in Nanking.—Peking nat. Hist. Bull., 11, pp. 27–34.Google Scholar
Zeuner, F. E. (1941). Geology, climate and faunal distribution in the Malaya archipelago.—Proc. R. ent. Soc. Lond., (A) 16, pp. 117–123.Google Scholar