DISTRIBUTION IN THE SUDAN

Khartoum: Omdurman (King 1926). Khartoum (Nuttall 1914A, Robinson 1926). Specimens from Omdurman and Khartoum in Sudan Government collections are from captive tortoises from unstated localities and probably represent the records or parts of the collections on which King's, Nuttall's, and Robinson's statements were made. No evidence is at hand to show that this tick occurs in nature in Khartoum Province.7

DISTRIBUTION

A. marmoreum ranges throughout the Ethiopian Faunal Region, except in the Arabian extension of this area. It appears to be more common in eastern and in southern Africa than it is in western and central Africa.

NORTH AFRICA: ALGERIA (As A. sparsum: Neumann 1899. This specimen was actually collected in the Paris Zoological Garden* (Bequaert, correspondence). The occurrence of A. marmoreum in Algeria and North Africa is questionable. See footnote, page 228).7

WEST AFRICA: FRENCH WEST AFRICA (Neumann 1911). SIERRA LEONE (Entomological Report 1916. Hoogstraal 1954C ).

CENTRAL AFRICA: BELGIAN CONGO (Neumann 1911. Nuttall and Warburton 1916. Bequaert 1930A,1931).

Note: According to Theiler (correspondence), the record

for Ruanda Urundi in Santos Dias (1954D) is incorrect.

EAST AFRICA: SUDAN (King 1911,1926. Nuttall 1914A. Robin son 1926. Hoogstraal 1954B).

*Hesse (1920) reported a female from the Leipzig Zoological Gardens and Hoogstraal (19540) another from the London Zoological Gardens.

ETHIOPIA (Neumann 1922. Stella 1938A,1939A,1940. Charters 1946). ERITREA (Tonelli Rondelli 1930A. Stella 1940). "SOMALI LAND, Gueldessa" (Robinson 1926). BRITISH SOMALILAND (Neumann 1922. Stella 1938A,1939A). ITALIAN SOMALILAND (Paoli 1916. Tonelli Rondelli 1932E, see also Tonelli-Rondelli 1935 as A. falsomarmoreum sp. nov. Niro 1935. Stella 1940).

KENYA (Neumann 1899,1901,1911,1912. Neave 1912. Anderson 1924A,B. Robinson 1926. Bequaert 1930A. Lewis 1931A,C,1932A, 1934,1939A. Loveridge 1936B). UGANDA (Hirst 1909. Robinson 1926. Mettam 1932. Wilson 1950. Binns 1952). TANGANYIKA (Howard 1908. Neumann 1907,1910B,1911. Morstatt 1913. Love ridge 1923C, as A. marmoreum is actually A. nuttalli according to Bequaert 1930A. Robinson 1926. Loveridge 1928. J. B. Walker, unpublished; see HOSTS below).

SOUTHERN AFRICA: ANGOLA (Specimens in HH collection). MOZAMBIQUE (Howard 1908. Neumann 1911. Robinson 1926. Santos Dias 1947A,1951A,1952D,1953B. Hoogstraal 1954C).

NORTHERN RHODESIA (Neave 1912. Robinson 1926). SOUTHERN RHODESIA (Jack 1921,1928,1937,1942). NYASALAND (Neave 1912. Robinson 1926. Wilson 1950B).

BECHUANALAND (Robinson 1926). SOUTHWEST AFRICA (Tromsdorff 1914). UNION OF SOUTH AFRICA (Koch 1844. Neumann 1899,1901, 1911. Lounsbury 1905*. Howard 1908. Dönitz 1910B. Bedford 1920,1926,1927,1932B. Robinson 1926. Curson 1928. Alexander 1931. Bedford and Graf 1934,1939. Neitz 1948).

Moore 1912.

OUTLYING ISLANDS: ZANZIBAR (Neumann 1899,1901,1911).

HOSTS

The chief hosts of the components of the A. marmoreum group await to be determined. Tortoises and the rhinoceros were men tioned by all early workers, mostly without further data (see REMARKS).

*See Robinson (1926).

Contemporary reports of the rhinoceros, Rhinoceros bicornis subspp. as a host are chiefly those of Robinson (1926) who re corded half a dozen collections from Kenya, Nyasaland, and Rhode sia. My collection and that of the Museum of Comparative Zoology contain specimens from rhinoceros in Kenya. Miss Walker's Tanga nyika collections (correspondence) contain 47 males and seven females from four rhinoceros hosts. The white, or square-lipped, rhinoceros, Ceratotherium s. simum, is a host in Zululand (Curson 1928).

Recent records from tortoises, Testudo spp. or Kinixys belliana, are those of Neumann (1922), Robinson (1926) with numerous collections from throughout the tick's range, and Hoogstraal (1954C) from Sierra Leone. Bedford's (1932B) statements and Theiler's unpublished records from South Africa indicate that tortoises are commonly infested in Transvaal and are the tick's chief host there. Other scattered records for tortoises are those of the Sudan specimens above, Mettam (1932) for Uganda, Wilson (1950B) for Nyasaland, and Santos Dias (1953B) summary of Mozambique ticks in which no other hosts are listed for A. marmoreum. A single collection from Tanganyika consists of six males and nine females (J. B. Walker, unpublished).

The warrener or leguan lizard, Varanus spp., is sometimes attacked. More recent reports are the single collection of Robinson (1926) and that from the Sudan listed above, Mettam's (1932) Uganda note, Loveridge's (1936A) Kenya record, a few lots in the Onderstepoort collection (Theiler, unpublished), and a few lots in the HH collection including one from Angola.

Some snakes are hosts, apparently only exceptionally*. Neumann's (1911) A. sparsum (said to be a synonym of A. marmoreum *The hosts of A. s. sparsum Neumann, 1899 (according to Robinson 1926 a synonym of A. marmoreum) were reported by Neumann to be Spilotes variabilis and Testudo mauritanica from Algeria and East Africa. S. variabilis is a synonym of S. p. pullatus, a large South American tree snake (Loveridge, correspondence). Therefore, since A. marmoreum (= A. sparsum) does not occur in South America, either Neumann's locality record or host record is incorrect. The fact that the "Algerian" material of A. sparsum was collected in a European zoological garden (cf. page 226) would suggest that both data are difficult to assess. It further suggests that the validity of the synonymy of A. sparsum should be reinvestigated.

came from Spilotes p. pullatus (= variabilis). Puff adders, Bitis spp., have been reported by Hirst (1909), Robinson (1926), Love_ ridge (1928), Bequaert (1930A), and Hoogstraal (1954C); Miss Walker's Tanganyika collection (correspondence) contains four males and a female from one puff adder. Python sp. has been listed by Neumann (1911) and Lewis (1934).

Mammals, other than the rhinoceros, are also occasional hosts. Theiler's unpublished records include some adults from domestic cattle and sheep. Lewis (1939A) found specimens rarely on buffalo and on domestic cattle. Two buffaloes were listed by Bequaert (1930A,1931) and one by Robinson (1926); my collection contains 37 specimens from a single buffalo in Kenya. Neumann (1911) recorded a genet, Genetta pardina, Robinson (1926) an eland, and Tonelli Rondelli (1930A) a bushpig. In my collection are numerous adults from giraffes in Kenya. Alexander (1931) was unable to induce adults to feed on domestic animals.

Among birds, the guineafowl has been recorded as a Uganda host by Mettam (1932).

Man was reported as an actual host by Charters (1946) in Ethiopia.

Nymphs are sometimes taken with adults from tortoises. Dr. Theiler (unpublished records) has numerous nymphs from fowls. Seven nymphs were removed from the African hoopoe, Upupa africana, in Mozambique (Hoogstraal 1954C). Lounsbury (1905*) and Howard (1908) reported that larvae and nymphs feed readily on lizards, cattle, goats, tortoises, and birds.

In the Onderstepoort laboratory, larvae and nymphs feed readily on guineapigs but adults do not (Theiler, correspondence). Kenya larvae and nymphs from females from tortoises feed well on the ears of rabbits, and resulting adults on the scrotum of a ram. This is a hardy species, and sixteen month old nymphs feed quite well while adults remain alive for 23 months without feeding (J. B. Walker, correspondence). See REMARKS below.

*See Robinson (1926).

The sorting out of these host records awaits a thorough bio logical and taxonomic study on this interesting group of ticks.

BIOLOGY

(See also REMARKS below)

Life Cycle

Lounsbury (1905*) stated that adults will not feed on goats or on oxen if they have not first fed from a tortoise in an earlier stage. Adults attached to the host before seeking the other sex, unlike many other species of Amblyomma. Lounsbury also provided observations on time required for each stage and for feeding. These data are not abstracted here for in the light of confused nomenclature it is questionable whether the species with which Lounsbury worked is the same as that in the Sudan. Note that Alexander (1931) was unable to induce South African adults of A. marmoreum to feed on cattle.

Ecology

In the Ethiopian Faunal Region, this group of ticks occurs in a variety of faunal areas. Where tortoises are common these ticks are often abundant, but this incidence is by no means universal in Africa.

On tortoises, these ticks are usually deep in the host's axillae and it is necessary to kill or anesthetize the animal to see or secure all of the specimens.

DISEASE RELATIONS

Textbook statements that A. marmoreum actually has been in criminated as a vector of boutonneuse fever of man refer merely to a remark that this was one of several tick species found on patients.

*See Robinson (1926).