NEAR EAST: PALESTINE (All as H. detritum: Adler and Feldman Muhsam 1946,1948. Feldman Muhsam 1948. Adler 1952). SYRIA (As H. detritum damascenium: Schulze and Schlottke 1930. Schulze 1930. Kratz 1940. As H. ?mauritanicum: Pigoury 1937). (As H. steineri steineri: detritum: Mimioglu 1954.

Schulze 1936D.

Schulze 1936D. Kratz 1940.

TURKEY

As H.

Kurtpinar 1954. Hoogstraal, ms. As
IRAN (As H. detritum: Delpy

H. mauritanicum: Yasarol 1954). 19368,1949).

EUROPE: *BULGARIA (As H. scupense and as H. detritum dar danicum: Pavlov 1947). *YUGOSLAVIA (As either H. scupense or H. savignyi by different local workers according to Oswald 1937. As H. scupense and H. detritum dardanicum: Oswald 1937,1938A, B,C,1939A,B,1940. As H. detritum: Angelovsky 1954).

GREECE (As H. scupense: Knuth, Behn, and Schulze 1918. Schulze 1919,1930. Schulze and Schlottke 1930. Kratz 1940. Pandazis 1947. As H. detritum scupense: Delpy 1946. As H. detritum dardanicum: Schulze and Schlottke 1930. Schulze 1930. Kratz 1940. Pandazis 1947).

SPAIN (As H. steineri codinai: Schulze 1936D. Kratz 1940. As H. mauritanicum: Gil Collado 1948A. Miranda Entrenas 1954).

RUSSIA: As H. detritum: Olenev 19298,1931A. Pavlovsky 1940. Kurchatov 1941. Pavlovsky, Galuzo, and Lototsky 1941. Galuzo 1941,1943,1944. Lototsky and Pokrovsky 1946. Tselish cheva 1953. Viazkova and Bernadskaia 1954. Petrisheva 1955. Zhmaeva, Pchelkina, Mishchenko, and Karulin 1955.

As H. detritum detritum: Olenev 1929A,1931C. Schulze and Schlottke 1930. Schulze 1930.

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

1940.

Kratz

*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 1929A.

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, Gilden blat, 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 1950. Alfeev 1951. Shatas 1952. Melnikova 1953. Rement sova 1953. Shatas and Bustrova 1954. Note: Pomerant zev 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 H. 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. scupense by Pomerantzev 1950; Delpy did not consider H. verae In his list of synonyms.

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

Delpy (19498) was not certain whether H. uralense Schulze and Schlottke, 1930, and H. volgense Schulze and Schlottke, 1930, are synonyms of H. detritum or of H. 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 20 and 200 from Ukrainia, identified by Schulze as H. volgense. These are typical H. detritum. The same institution possesses 8 and 300 from Crimea, determined by Schulze first as H. ?marginatum, 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. detri tum (or, in part, "H. scupense") was treated by Schulze in part as H. uralense and in part as H. volgense, though, he overlooked other species in the same collection and referred to them by the

same name.

As H. tunesiacum pavlovskyi: Schulze and Schlottke 1930. Described and illustrated as H. detritum pavlovskyi by Kratz 1940.

According to Galuzo 1935, the H. asiaticum of Olenev is H. detritum (H. detritum rubrum). H. asiaticum is usually con sidered as a synonym of H. dromedarii.

MIDDLE EAST: INDIA (As H. aegyptium ferozedini and as H.
sharifi: Schulze and Schlottke
Kratz 1940.
Kratz 1940. As H. isaaci:

a. isaaci: Sharif 1928. As H.
1930. Schulze and Gossel 1936.

Kratz 1940). PORTUGESE INDIA (As H. detritum: Santos Dias 1954J).

FAR EAST: CHINA including MONGOLIA (As H. detritum albipictum: Schulze 1919,1930. Schulze and Schlottke 1930. Yama shita 1939. Kratz 1940. As H. detritum perstrigatum: Schulze 1930. Schulze and Schlottke 1930. Hoeppli and Feng 1933. Olenev 1934. Kratz 1940).

HOST S

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 did 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 Forest (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 prin ciples 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

tick whose adults feed in the winter and early spring. Pomerant zev (1950) considers "H. scupense" to be a "biological race" of H. detritum.

Life Cycle of H. detritum

The life cycle of H. detritum (= H. mauritanicum) in Algeria has been studied by Sergent, Donatien, Parrot, and Lestoquard (1931B and subsequent works). Larvae hatch in the late autumn and feed and molt to nymphs on the same host, remaining attached for approximately sixteen days. Nymphs hibernate for approximate ly eight months either in groups in cracks and crevices of farm yard walls about six feet above the ground, or under boulders. They are never found in fields without trees and boulders. Hibernation sites are generally those with a warm, sunny exposure, and those nymphs in the warmest places molt earliest in the year (June). Some days after molting, young adults start out in search of a host; they leave their hibernation place at night and travel towards stables, sometimes covering as much as thirty yards a week to reach cattle or horses.

Adults commence attaching to the host in mid June. They mate and feed there, females dropping off after ten to twelve days. Feeding females are most common in July and August; afterwards fewer are found on animals. Those that feed later do not oviposit the same season. Eggs are laid on the ground near animals and hatch in about six weeks. Females die after laying eggs. There appears to be quite a little variation from this typical life cycle.

Males remain on the host for a much longer time than fe males and may move from one host to the other. Nymphs some. times move from one host to another.

Note that according to the Algerian reports mentioned above, nymphs hibernate in cracks or crevices of farmyard walls but that almost all Soviet workers, mentioned below, find nymphs alone or with larvae on cattle during the winter. This raises the question whether the Russian and Algerian ticks are actually the same species, and, if so, whether climatic or other factors modify their choice of niches for hibernating in these far-flung areas.