shiny black spots in the lateral groove, one pair above coxa I and one pair between coxae III and IV, there should be no real difficulty in distinguishing the species. Tarsal differences mentioned in Nuttall et al (1908) are too variable for applica tion. Adults average about 8.0 mm. long by 6.0 mm. or 7.0 mm. wide, although freshly engorged females may reach 11.0 m. in length. 0. savignyi is usually somewhat larger, averaging about 11.0 mm. Tong. The mammillated integument, conspicuous tarsal dorsal protuberances, and absence of cheeks easily distinguish both 0. moubata and O. savignyi from all other African species. Males are often a little smaller than females and their genital aperture is a short, rounded opening as compared with the broad slit of the female. Nymphs have no genital aperture, but in larger instars a small round depression is present in its place.

In a study of the sexual differences in this species, Nuttall (in Cunliffe 1921) has shown that the average male is smaller than the average female, though extremes overlap, and that the same is true for the size of the genital aperture of the two sexes.

An estimate of the stage of development of immature stages can be made on the basis of differences in mouthparts, legs, and spiracles (Cunliffe 1921).

ORNITHODOROS (ORNITHODOROS) SAVIGNYI (Audouin, 1827).

(Figures 8 to 12)

THE EYED TAMPAN*

DISTRIBUTION IN THE SUDAN

Central and northern Provinces (King 1908,1911,1926).

Localities reported in literature and from which I have seen specimens are:

Darfur: 15 miles northwest of Fasher (SGC). Nyala (horse; SVS). No locality (BMNH).

Kordofan: El Obeid (camel yard; SGC, SVS). "Several local

ities (King 1911, Balfour 1911E); SGC, SVS).

Blue Nile: Wad Medani (camel yard; SGC, HH). Wad Raiya and Kosti (SGC).

Kassala: Karora and Goz Rageb (on human; SGC). Khor Mashi hills south of Tokar, Bir Qui Tiri (under tamarisk tree near well; SGC).

Khartoum: Khartoum (cattle quarantine station; HH). Khartoum, Shambat (H. W. Bedford 1939). "Oasis near Khartoum (King 1911 Balfour 1911E).

Northern: Dongola (Dönitz 1906). Wadi Halfa and Abu Hamed (camel yards; HH).

DISTRIBUTION

0. savignyi is distributed locally through arid parts of North, East, and South Africa, the Near East, India, and Ceylon. The Near

*In South Africa, called "The Sand Tampan"" (Theiler 1952A,B).

or Middle East probably was its original home. Transportation by caravans, lack of field search and of literature reports, and con fused identification in Africa have combined to provide a still uncertain picture of this tampan's actual distribution within the noted range.

During Neumann's time 0. savignyi was frequently confused with 0. moubata and acceptance of many early records and of some even more recent reports is questionable. These two species never occur in the same ecological niches. They are close neighbors in some areas, as Somaliland, where 0. moubata inhabits huts next to trees under which O. savignyi hides.

Brumpt (1936) summarized the known geographical distribution and medical relations of O. savignyi. He noted especially that even though this tampan is frequently found along remote camel trails, it is not known from Morocco in spite of considerable search for it there.

Brumpt (loc. cit.) also considered it surprising that 0. savignyi has not been carried to Madagascar but that 0. moubata is common in some areas of that island. Once the very different biology of these two species is understood, a reasonable explana tion for this distribution pattern may be offered.

0. moubata is a highly domesticated parasite that inhabits man's dwellings and frequently hides among his personal effects. It was probably transported from Africa to nearby Madagascar among gear in seagoing vessels. This tick has not been able to survive elsewhere outside of tropical and southern Africa, where it is endemic.

0. savignyi, on the other hand, appears to have erratically invaded Africa from the East. It prefers more arid outdoor conditions than are found in most parts of Madagascar, and part of its spread probably has been by camels, which are not used in Madagascar. In southern Africa, the range of 0. savignyi appears to be related to environment and wild animals rather than to the comparative recent introduction of camels, movements of domestic stock, or treks of hunters (Theiler, unpublished). In African regions where 0. savignyi does occur, populations are often spotty,

even in areas that appear favorable. This tampan's predilection for resting outdoors in the soil probably more closely confines its apread to overland routes than does 0. moubata's propensity for hiding in goods or personal effects.

Koch

NORTH AFRICA: EGYPT (Savigny 1826. Audouin 1827. 1875. Brumpt 1908A. Neumann 1911. Yakoub 1945. Halawani 1946. Davis 1947. Taylor and Hurlbut 1953. Hoogstraal 1954A. Davis and Hoogstraal 1954. Theiler and Hoogstraal 1955. Taylor et al 1955. Hurlbut 1956. Taylor, Work, Hurlbut, and Rizk 1956). LIBYA (Franchini 1927,1929A,E,1932B,1933A,B,C,D,1934B,1935A,B,C, 1937,1938. Zavattari 1930,1932,1933,1934.

Tonelli-Rondelli

Theiler

1930B,1932A,B,D,1935. Gaspare 1933,1934. Garibaldi 1935. and Hoogstraal 1955). TUNISIA, including Ile de Djerba (Weiss 1911B,1912. Galli-Valerio 1911A. Nicolle, Blaizot, and Conseil 1912,1913A,B. Absence in oases: Langeron 1921. Nicolle and Anderson 1927. Colas Belcour 1928,1929A,C,1930,1931). ALGERIA (Chalon 1923. Catanei 1929). [Apparently absent in Morocco (Brumpt 1936).7

WEST AFRICA: NIGERIA (Northern Province only: Simpson 1912A. Alcock 1915. Leeson 1953). Leeson 1953). FRENCH WEST AFRICA (Brumpt 1936. Kone 1949. Rousselot 1951,1953B). [Absent in Gold Coast: Selwyn Clark, Le Fanu, and Ingram (1923) and Corson and Ingram (1923). Absent in Liberia: Bequaert (1930A).7

CENTRAL AFRICA: No substantiated records. Not known from Belgian Congo, but should be searched for there (Bequaert 1930A, 1931). 0. savignyi caecus listed by Schwetz (1927) is a synonym of 0. moubata. Leeson (1953) lists 0. savignyi, probably after Nuttall et al (1908) or Howard (1908) but these records are most likely repetitions of early misidentifications.]

EAST AFRICA: SUDAN (Not Koch 1875 or Neumann 1896, "Upper Nubia, cf. Eritrea. Dönitz 1906. Brumpt 1908A. King 1908,1911, 1926. Neumann 1911. Balfour 1911E,1912. Brumpt 1936. Kirk 1938B,1939. H. W. Bedford 1939. Hoogstraal 1954B. Theiler and Hoogstraal 1955).

ETHIOPIA (Nuttall et al 1908. Neumann 1911,1922. Wenyon 1926. Bruns 1937 statements questionable). ERITREA (Koch 1875

and Neumann 1896 from River Anseba, "Upper Nubia". Franchini 1929D, E. Tonelli-Rondelli 1930A. Niro 1935. Stella 1938A,1939A,1940). FRENCH SOMALILAND (Neumann 1922. Brumpt 1936. Stella 1938A,1939A, 1940). BRITISH SOMALILAND (Pocock 1900. Drake-Brockman 1913A,B, 1914,1915A,B,1920, biology and medical implications in part mixed with 0. moubata. Neumann 1922. Stella 1938A,1939A,1940. Anderson 1947. Heisch 1950A. Falcone 1952, erroneous disease rela tions). ITALIAN SOMALILAND (Brumpt 1908B. Lees 1914. Paoli 1916. Franchini 1925,1927,19290,E,1934. Tonelli Rondelli 1930A, 1931. Mattei 1933. Niro 1935. Massa 1936A,B, medical implica tions questionable. Moise 1938,1950. Stella 1938A,1939A,1940. Lipparoni 1951,1954. Giordano 1953).

KENYA (The O. savignyi of Karsch 1878 refers to 0. moubata. Dönitz 1906. Neave 1912. Neumann 1912. Anderson 1924A,B. Mackie 1927. Brassey-Edwards 1932. Lewis 1931A,C,1939A. Heisch 1937,1951A. Walton 1951). UGANDA (Bruce et al 1911. Neave 1912. Mettam 1932).

[?TANGANYIKA:

?As 0. morbillosus: Gerstacker (1873). Neumann (1901,1907,1910B,1911). Howard (1908) ?quoting Neumann. Absent in Bukoba: Horstatt (1914). The presence of this tick in Tanganyika remains questionable._7

SOUTHERN AFRICA: ANGOLA: Neumann (1896) listed specimens from Landana that Bequaert (1930A) quite logically believes were 0. moubata. Subsequent statements (Santos Dias 1950C) are merely a repetition of Neumann and disease relations are incorrect. "RHODESIA": Report of Lounsbury (1900), obviously referring to 0. moubata, quoted by other authors. No subsequent reports of 0. savignyi. ?MOZAMBIQUE: Neumann (1896) ?should be 0. moubata. Lounsbury (1900) should be 0. moubata. Theiler (1943B) states 0. savignyi doubtfully recorded but possibly present. Santos Dias (1953B) evidence not presented. Mozambique records are considered incorrect by Theiler and Robinson (1954).7

NYASALAND (Wilson 1950B). BECHUANALAND (Bedford 1926,1927, 1932B,1934. Theiler and Robinson 1954). SOUTHWEST AFRICA (Louns bury 1900C, possibly referring to 0. moubata. As O. pavimentosus: Neumann 1901,1911. Dönitz 19070,1910B. Howard 1908. Trommsdorff 1914. Bedford 1926,1927, 1932B,1934. As 0. moubata: Mönnig 1949.