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DISTRIBUTION

H. detritum is an Asiatic tick that ranges from Manchuria through China and India, much of southern Russia, southeastern Europe, and the Middle East; into Asia Minor, the Near East, the Mediterranean littoral of Africa westwards to Algeria; and into northcentral Sudan, where it occurs only in small, localized populations. It also occurs in Spain, probably having been introduced from northwestern Africa.

In Egypt and apparently in Libya, H. detritum is considerably less common than it is to the west in Algeria and in Near Eastern countries. This distributional pattern is common for animal groups that have invaded North Africa from the east.

NOTE: Specimens referred to this species by Rageau (1951) from the Cameroons were later determined by him (1953) as H. truncatum (= H. transiens).

NORTH AFRICA: *ALGERIA (All as H. mauritanicum or as H. mauritanicum annulatum: Senevet 1922B,C,192ZA,I;25, 1928A, 1929B, I937. Senevet and Rossi 1924. Sergent, Donatien, Parrot, Lestoquard, and Plantureux 1926, 1927A,B,C,D,E. Sergent, Donatien, Parrot, and Lestoquard 1928A,B,C,1931A,B,C,D,E,F,1932A, B, 1933A,B, 1935A,B,1936A,B,C, 1945. A. Sergent 1930. Sergent and Poncet 1937, 1940, 194l. Sergent, Donatien, and Parrot 1945. E. Sergent 1948. Blanc and Brunneau 1949. d'Arces, 1952. As H. detritum mauritanicum: Schulze 1930. Kratz 1940).

MOROCCO (As H. mauritanicum: Desportes 1938). TUNISIA (As H. detritum: "Senevet 1937. Colas-Belcour and Rageau 1951). LIBYA (As H. mauritanicum: Franchini 1927, 1929A, E. Hoogstraal, ms.). EGYPT (Present but rare: Hoogstraal, m.)

East AFRICA: SUDAN (As H. detritum: Hoogstraal 1954B).

Note: H. mauritanicum has been reported from Somalia, without precise Locality data, by Niro (1935), but this record has not been subsequently repeated by Italian workers.

*Algerian specimens of H. mauritanicum, kindly presented by Dr. E. Sergent and Dr. Senevet, conform to H. detritum.

NEAR EAST: PALESTINE (All as H. detritum: Adler and Feldman Muhsam T946,1948. Feldman Muhsam 1948. Adler 1952). SYRIA (As H. detritum damascenium: Schulze and Schlottke 1930. Schulze 1930. Kratz ISO. As H. *mauritanicum: Pigoury 1937). TURKEY (As H. steineri steineri: Schulze I:36D. Kratz 1940. As H. detritum: Mimi oglu Iš52. Kurtpinar 1954. Hoogstraal, ms." As H. mauritanicum: Yasarol 1954). IRAN (As H. detritum: Delpy

EUROPE: *BULGARIA (As H. scupense and as H. detritum dardanicum: Pavlov 1947). #, either H. Scupense or H. savignyi by different local workers according to # 1937. As H. scupense and H. detritum dardanicum: Oswald 1937,1938A, £ As H. Gürttum: "Angelovsky 1954).

GREECE (As H. scupense: Knuth, Behn, and Schulze 1918. Schulze 1919, 1930. # and Schlottke 1930. Kratz 1940. Pandazis 1947. As H. detritum scupense: Delpy l946. As H. detritum dardanicum: Schulze and #t'. 1930. Schulze T93UTKratzTSZUT Pandazis 1947).

SPAIN (As H. steineri codinai: Schulze 1936D. Kratz 1949. As H. mauritanicum. GIT CoITado Tø48A. Miranda-Entrenas 1954).

RUSSIA: As H. detritum: Olenev l929B, 1931A. Pavlovsky 1940. Kurchatov I94 I. Pavlovsky, Galuzo, and Lototsky 1941. Galuzo 1941,1943,1944. Lototsky and Pokrovsky 1946. Tselishcheva 1953. Viazkova and Bernadskaia l954. Petrisheva 1955. Zhmaeva, Pchelkina, Mishchenko, and Karulin 1955.

As H. detritum detritum: Olenev 1929A, 1931C. Schulze and Schlottke 193OTSchulze TS30.

As H. detritum rubrum: Schulze 1930. Olenev 1931A,C,1934. Pomerantzev 19%. Galuzo 1935. Galuzo and Bespalov 1935. Kratz 1940.

*The hosts of immature stages listed by authors in these countries indicate that they are quite possibly dealing with a different species of tick.

As H. detritum Pavlovskyi. Olenev l929A.

As H. transcaucasicum: Olenev 1934 (the synonymy of this name appears to have been overlooked by subsequent workers).

As both H. detritum and H. scupense: Pomerantzev 1937. Pomerantzev, Matikashvily, and Lototsky 1940. Markov, Gildenblat, Kurchatov, and Petunin 1948. Note: Pomerantzev 1950, in his work on Soviet ticks, considers these two as distinct species. See BIOLOGY and IDENTIFICATION below.

As H. scupense: Olenev 1934. Nikolsky 1948. Petunin 1948.

Pomerantzev . Alfeev l95l. Shatas l952. Melnikova 1953. Rementsova 1953. Shatas and Bustrova 1954. Note: Pomerantzev 1950 considers H. volgense and H. uralense to be synonyms of H. scupense, while Delpy places them under either H. detritum or " H. excavatum (see two paragraphs below). Note the references To wr: scupense" in the section on European distribution above. See also paragraph above and below.

H. verae Olenev, 1931B, is also placed in synonymy under H. # by Pomerantzev 1950; Delpy did not consider H. verae in s list of synonyms.

As H. volgense and/or H. uralense: Schulze and Schlottke 1930. s# . Olenev 1929A, 1931A,C,1934. Zasukhin 1932, l935. Borzenkov and Donskov 1934. Zolotarev 1934. Galuzo l935. Kochetkov 1935. Artjukh 1936. Kurchatov 1940B. Markov, Abramov, and Dzasokhov 1940. Enigk 1947. See paragraph on H. scupense above and paragraph below.

Delpy (1949B) was not certain whether H. uralense Schulze and Schlottke, 1930, and H. volgense Schulze and Schlottke, 1930, are synonyms of H. detritum or o ... excavatum, and stated that Russian workers may have included both species under these two names.

In Schulze's collection, now in Rocky Mountain Laboratory, there are 236 and 292 from Ukrainia, identified by Schulze as H. volgense. These are typical H. detritum. The same institution possesses 336 and 39Q from Crimea, determined by Schulze first as H. 7marginatum, and later crossed out and identified by him as H.

uralense. The males are, all but one, H. detritum”; the exception appears to be H. marginatum; the females are in poor condition. It is reasonable to assume that what is now considered to be H. detritum (or, in part, "H. scupense") was treated by Schulze in part as H. uralense and in # as H. volgense, though, he overlooked other species in the same collection and referred to them by the Same name •

As H. tunesiacum #: Schulze and Schlottke l930. Described and iTrustrated as H. detritum Pavlovskyi by Kratz l940.

According to Galuzo 1935, the H. asiaticum of Olenev is H. detritum (= H. detritum rubrum). H. asiaticum is usually con

SIGGFed as a synonym of H. Römedarii.

MIDDLE EAST: INDIA (As H. aegyptium ferozedini and as H. a. isaaci: arif 1928. As H. Sharifi : " Schulze and Schlottke 1930. Schulze and Gossel 1936. Kratz 1940. As H. isaaci: Kratz 1940). PORTUGESE INDIA (As H. detritum: Santos Dias 1954J).

FAR EAST: CHINA including MONGOLIA (As H. detritum albiictum: "Schulze 1919, 1930. Schulze and Schlottke T930. "YamaShita 1939. Kratz l940. As H. detritum perstrigatum: Schulze 1930. Schulze and Schlottke 1930. Hoep###and Feng 1933. Olenev 1934. Kratz 1940).

HOSTS

Domestic cattle and horses are the most common hosts of H.

detritum, all stages of which feed on the same kind of animal. Sheep and goats are sometimes attacked. For the Soviet Union, Pomerantzev (1950) lists cattle, horses, donkeys, pigs, camels, sheep, and hares; and, for nymphs, especially cattle and horses. Man is apparently commonly attacked under local conditions.

Oswald's (1939) and Pavlov's (1947) remarks for parasitism by immature stages in Yugoslavia and Bulgaria of various birds and

*At the time of checking this material, I did not realize the significance of "H. scupense". It cannot, therefore, be said that these specimens # not resemble the latter form.

lizards, and of hares and dogs are either incorrect or refer to
a different species of tick, most probably H. marginatum.7 A.
Sergent (1930) noted a nymph of H. detritum parasitizing another
nymph of the same species. Hosts of specimens in British Museum
(Natural History) are domestic cattle and camels (Palestine),
domestic buffalo and pony (India), hare (India), and deer
(Romania).

"H. scupense" attacks the same hosts as H. detritum. It has also been found on the Persian or goitred gazelle, Gazella subgutterosa, and on the red deer, Cervus elaphus bactrianus (Pomerantzev 1950). In the Crimean National # (Melnikova 1953), "H. scupense" is common on red deer and occurs in smaller numbers on roe deer. It is present but not common on hares but absent on squirrels and jays. Wild foxes may also be attacked. Domestic cattle are heavily infested, collections from single animals in various localities in and near the forest averaging from 78 to 756 ticks, with individual maxima ranging from 350 to 5000 ticks per animal. Domestic pigs in the same forest averaged 21 ticks per host. Zolotarev (1934) listed this tick (as H. volgense) from camels.

BIOLOGY
Introduction

Because of the interesting biological and taxonomic principles involved, separate reviews of life cycle and ecology are devoted to H. detritum and "H. scupense". H. detritum is a twohost species whose adults feed in the summer and whose nymphs undergo an extensive winter diapause; this feature is common throughout the range of H. detritum though the overwintering habits in Algeria and (usually) in Russia differ markedly; it should be determined what factors account for this variation in habits. H. scupense is confined to parts of the Soviet Union and possibly to Greece and Yugoslavia; it is said to be distinguishable from H. detritum by slight morphological differences through parts of its geographic range (see IDENTIFICATION below), and is a one-host

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