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WEST AFRICA: LIBERIA (Bequaert 1930A). SIERRA LEONE (Simpson 1913. Robinson 1926). IVORY COAST (Rousselot 1951,1953B. Villiers 1955).

CENTRAL AFRICA: CAMEROONS (Neumann 1901,1911. Ziemann 1905, 1912A. Rageau 1951,1953A,B). RIO MUNI (Robinson 1926). FRENCH EQUATORIAL AFRICA (Neumann 1899. Tonelli_Rondelli 1930A. Fiasson 1943B. Rousselot 1951,1953B. Rageau 1953B).

BELGIAN CONGO (Neumann 1899,1911. Nuttall and Warburton 1916. Roubaud and Van Saceghem 1916. Robinson 1926. Schwetz 1927A,B,C, 1932. Schouteden 1929. Tonelli Rondelli 1930A. Bequaert 1930A,B, 1931. Rodhain 1936. Fain 1949. Theiler and Robinson 1954. Van Vaerenbergh 1954).

NOTE: According to Theiler (correspondence), the record for Ruanda Urundi by Santos Dias (1954D) is in error.

EAST AFRICA: SUDAN (King 1911,1926. Robinson 1926. Hoogstraal 1954B).

KENYA (Neumann 1922. Lewis 19310,1932. Mulligan 1938). UGANDA (Neave 1912. Robinson 1926. Robinson 1926. Tonelli Rondelli 1930A. Mettam 1932. Wilson 1950. See HOSTS below). TANGANYIKA (Neumann 1899,1907, 1910B,1911. Morstatt 1913. Robinson 1926. Hoogstraal 1954C. J. B. Walker, unpublished; see HOSTS below).

SOUTHERN AFRICA: ANGOLA (Gamble 1914. Robinson 1926. Santos Dias 1950). MOZAMBIQUE (Santos Dias 1947A,B,1948,1949‚D,1950B, 1953B,1955A. Bacelar 1950. Tendeiro 1952). Bacelar 1950. Tendeiro 1952F). NYASALAND (Neumann 1899*. Neave 1912. Robinson 1926. Wilson 1950B).

A. tholloni has not yet been recorded from the UNION OF SOUTH AFRICA, but Theiler (correspondence) believes that this is probably because it has not been looked for and that it possibly occurs on Kruger Park elephants and on remote herds in Southwest Africa. possibility that this tick is incapable of following its normal

The

*The reference to "region du Nyassa" by Neumann (1899), for specimens collected by Ed. Foa, may refer to Niassa Province, Mozambique.

host into the southern periphery of the host's range should be considered in view of its apparent total absence on elephants in Bahr El Ghazal Province of the Sudan.7

HOSTS

All authors list the African elephant, Loxodonta africana subspp., as chief host. Uncommon hosts that have been reported for a few specimens are: gazelle (Neumann 1901,1907,1910B), antelope and domestic horse (Neumann 1911), rhinoceros (Neumann 1922), leopard (Robinson 1926), bushpig and large lizard (Schwetz 1927B), domestic dog (Santos Dias 1953B) and buffalo (Hoogstraal 1954C). Dr. Theiler has larvae, nymphs, and a male specimen from a bird, Pitta reichenowi, another indication of ground birds as hosts of immature Amblyomma species. In our collection are several nymphs and a male with massive legs taken from a hippopo tamus in Kazinga Channel of Lake Edward by Lt. Colonel Don Davis, U.S. Army. Miss J. B. Walker has a collection consisting of seven males and four females from a Tanganyika black rhinoceros; also others from an elephant there.

The small nymph collected at Lokila, Equatoria, from a chameleon by E. T. M. Reid is an unusual record. Santos Dias (1948) states that larvae and nymphs are rarely found on elephants. The only other records for nymphal hosts in nature are those of the bird and hippopotamus listed above, and one nymph in the Onderstepoort collection (Theiler, unpublished) with adults, from an elephant at Toro, Uganda.

BIOLOGY

Adult specimens of the elephant amblyomma may be found on any part of the host's body, Immature stages previously have been reported only by Santos Dias (1948), who states that larvae and nymphs are rarely taken on elephants. Santos Dias experimentally reared larvae and nymphs on guinea pigs and reported that six months were required to complete the three host life cycle. He observed a chalcid wasp parasite, Hunterellus hookeri, infesting nymphs. Fiasson (1943B) reported 3000 eggs from an engorged female.

Mr. J. Owen, who furnished the 197 specimens from a single elephant in a plains herd passing through Lotti Forest, reported that his "boys" could have collected at least twice as many from this elephant if they had had more containers for them. No specimens other than the few listed could be found on the three other Equatoria Province elephants noted above. Numerous other newly killed elephants in this Province have been examined with out finding ticks of any sort.

DISEASE RELATIONS

A. tholloni is possibly a vector of Nuttallia loxodontis of elephants.

REMARKS

The stage to stage growth of A. tholloni has been charted by Campana Rouget (1954). Misshapen specimens have been described and illustrated by Santos Dias (19478,1949,1955A).

Larval and nymphal stages of A. tholloni were described and illustrated by Santos Dias (1949D).

Variations in male scutal patterns from Mozambique were il lustrated by Santos Dias (1947A). Within the geographical range of the elephant amblyomma there are two scutal color patterns, one drab and lightly pigmented with small areas of color, the other brightly marked, usually with more extensive pigmented areas. The bright form is particularly common among numerous specimens seen from West Africa and rare among those from East Africa; the drab form is common in East African specimen and rare in West African material. After having examined all of the numerous A. tholloni specimens in British Museum (Natural History) collec tions, which represent almost all areas of the geographic range of this species, one may only conclude that these two color patterns do not appear to be genetic variants and are probably not associated with temperature or rainfall factors or with methods of preservation. They may possibly derive from nutritional factors. A biological study of living specimens is the

only means of determining the reason for these two color patterns. Since the above was written, Rageau (1953B) has reported that Cameroons specimens and others that he has seen from French Equatorial Africa all show the reduction in ornamentation that has appeared to me to be more common in East than in West African specimens.7

Robinson (1926) stated that a large pale spot in each lateral field of the scutum and a stout spur on coxa IV are female diagnostic characters. I have examined Nuttall's lot 3381 in British Museum (Natural History), on which Robinson's species definition and illustrations were based, and find it to be the most heavily and liberally ornamented material, along with a few others from Sierra Leone, of any representatives of this species in the collection. Actually, lateral field pale spots are absent in most of these specimens. The stoutness of the spur on coxa IV is also a variable character and the specimen selected by Robinson is an extreme example. In most specimens, this spur is merely a small pointed projection from or near the posterior coxal margin.

All Sudan specimens at hand are drably colored and their pigmented areas are no more extensive than those illustrated (Figure 90). On some, ornamentation is almost obsolete.

IDENTIFICATION

Males. No other African amblyomma can be confused with this species because of the smallness of its pigmented areas, flat eyes, and absence of both lateral grooves and of large scutal punctations. Males measure about 5.0 mm. long and 4.0 mm. wide.

Females are equally easily distinguished from all others in Africa by the absence of large scutal punctations, triangular scutal shape with narrow posterior margin, and distribution of color pattern that is usually only an irregular spot in the pos terior point but sometimes also has small lateral spots. The cervical grooves are short and eyes are large, flat, and pale. Females, unengorged, are usually about 6.0 mm. long and 4.7 mm. wide.

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Figures 92 and 93, o, dorsal and ventral views Figures 94 and 95, o, dorsal and ventral views

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