Search

Intraspecific host discrimination is widespread in solitary parasitoids whose adult females forage for and evaluate host suitability, whereas interspecific discrimination is less common. In some parasitoid species, mostly Diptera and Coleoptera, the larva performs the last step of host searching. It has been suggested that host discrimination will rarely occur in such host-seeking larvae because their low mobility results in a low host encounter rate. We determined the extent to which the larvae of Aleochara bilineata Gyllenhal (Coleoptera: Staphylinidae), a solitary parasitoid of aggregated Diptera pupae: (1) discriminated between unparasitized hosts and hosts parasitized by conspecifics; (2) used semiochemical cues to discriminate; (3) were influenced by life expectancy, presence of conspecifics and host availability in their host acceptance decision; and the extent to which (4) A. bilineata and A. bipustulata L., a species exploiting the same hosts and occurring sympatrically, showed interspecific host discrimination. A. bilineata larvae were able to discriminate between unparasitized hosts and hosts parasitized by conspecifics in a choice experiment. Such behavior has never previously been described for a coleopteran parasitoid or for a parasitoid species whose larvae perform host searching. Host discrimination in this species was not based on the presence of visual or tactile cues (e.g., entrance holes) but rather on chemical cues. The life expectancy of A. bilineata larvae was significantly shorter in the presence than in absence of hosts, and older larvae had lower parasitism success than young larvae in a 24-h experiment. However, the host acceptance decision of A. bilineata larvae was not influenced by larval age or the presence of conspecifics when the ratio of hosts per larva was greater than or equal to 1. When hosts were scarce, the degree of superparasitism increased significantly with the number of foraging conspecifics and the age of the larvae. Both species of Aleochara showed intra- and interspecific host discrimination in a choice experiment. In contrast to A. bipustulata, A. bilineata larvae more frequently parasitized hosts parasitized by A. bipustulata than those parasitized by conspecifics. We suggest that host discrimination will be frequent in solitary parasitoids with host-seeking larvae when hosts are aggregated.

Fimicolous Staphylinidae prey on rearing dipterous in cattle dung, acting as their natural controllers, including pests such as horn fly. To survey the abundance and succession of these coleopterans in cattle dung deposited in pasture, six experiments were conducted from March to October 1995 in Uberlândia, State of Minas Gerais, Brazil. Cattle dung pats were exposed at a pasture for 1 hr, 8 hr, 24 hr, 2, 3, 7 and 10 days and were than taken to laboratory separate from each other, for Staphylinidae extraction. A total of 156 dung pats were exposed at pasture, from which 6225 Staphylinidae were recovered. Representing at least 30 species, Staphylinidae sp.1 (29.6%), Philonthus flavolimbatus (22.2%), Heterothops sp.1 (16.6%), Oxytelus sp.2 (7.6%), Aleochara sp.2 (7.6%) and Criptobium sp.1 (4.4%) were the most abundant, representing 87.8% from the total. The increased frequency of the majority of these species along the dung exposition time at pasture, indicated that they would be preying on at all the immature stages of the dipterous, or eggs and first instar larvae of species that lay eggs on the dung after its second exposition day at the pasture.

A taxonomic revision of the subgenus Coprochara Mulsant & Rey, 1874 of the genus Aleochara Gravenhorst, 1802 in Japan is presented. The following three species are recognized: Aleochara (C.) verna Say, 1833, A. (C.) binotata Kraatz, 1856 and A. (C.) squalithorax Sharp, 1888, of which A. binotata is new to Japan. All previous records of “A. (C.) bipustulata (Linnaeus, 1760)” should be regarded as misidentifications of either A. verna or A. binotata. New records are added for, A. squalithorax, a littoral species. All species are redescribed, figured, keyed and mapped.

The Japanese species of the seashore-dwelling subgenera Emplenota Casey and Triochara Bernhauer of the genus Aleochara Gravenhorst are revised. Five species are recognised in Emplenota, of which three are described as new species: Aleochara (Emplenota) segregata n. sp., A. (E.) hayamai n. sp. and A. (E.) yamato n. sp. The remaining known species A. (E.) fucicola Sharp and A. (E.) puetzi (Assing) are redescribed. Three species recognised in Triochara, Aleochara (Triochara) trisulcata Weise, A. (T.) zerchei (Assing) and A. (T.) nubis (Assing) are redescribed. All species are keyed. For some species ecological data are reported. The phylogenetic relationships of the Japanese species are discussed, and the distributions of all species are mapped.

Study of 41 species provided information as follows: (a) a setigerous lobe is located at the costal margin in every species in the subf. Staphylininae; (b) a setal comb occupies the same margin in Xantholinini and O maliinae; (c) one or more spinulae do line the anal fi eld of all O maliinae and most O xytelinae, Tachyporinae and A leocharinae; (d) number of these spinulae in A leocharinae ranges from 1 up to about 100 and is null in two species. Hypothetically, a functional importance may be attributed to both the setigerous lobe, which suggests a mechanical receptor for wing folding control, and to the f labellum-like anal fi eld of the Aleochara, which looks as a device affecting the flying trim.

A succession of insect species associate with decaying bodies, and because of the relatively predictable arrival and departure times of many species, this process is routinely used to estimate minimum post-mortem interval. Corpse fauna are compared with baseline data on succession rates, which are usually taken from decomposing animal carcasses. Baseline data are traditionally collected over a single year only; however, annual differences in weather patterns and insect population parameters probably alter succession rates. Surprisingly, this potential source of error has not previously been examined. Therefore, a two-year succession experiment was performed to investigate annual variation in succession rates. Sets of five still-born piglet carcasses were placed in a forest each season of both years. Six insects that occur year-round were selected: adults of Aleochara brachialis Jekel, Anotylus spp., Creophilus erythrocephalus Fabricius (Staphylinidae: Coleoptera), Ptomaphila lacrymosa Schreibers (Silphidae: Coleoptera), Pseudonemadus spp. (Leiodidae: Coleoptera), and larvae of Calliphora spp. (Calliphoridae: Diptera). Arrival and departure weeks to and from carcasses were measured in each season of both years. Two-way ANOVAs revealed significant season and year interactions on the arrival weeks of Anotylus spp., C. erythrocephalus, and Pseudonemadus spp., and significant differences in seasonal and yearly arrival times for A. brachialis, and P. lacrymosa. For larvae of Calliphora spp., there was a significant seasonal effect on arrival times. A significant interaction between season and year affected the departure weeks of all taxa. These results show that error in the estimates of minimum post-mortem interval can result if baseline succession data are collected only over one year.

Two new species, Aleochara (Aleochara) pseudolustrica Klimaszewski, and Aleochara (Calochara) beckeri Klimaszewski, from Florida, are described on the basis of structures of adults. A key is given to adults of the eleven species of Aleochara now known to occur in Florida. Additional distributional data are given for Aleochara (Coprochara) notula Erichson and A. (A.) lustrica Say.

A revision of the genusAleocharaof America north of Mexico is presented. Seven subgenera are redefined with 47 currently recognized species, of which nine are described as new (A.arizonica,A.cavernicola,A.curtidens,A.lobata,A.ocularis,A.nidicola,A.rufobmnnea,A.rufonigra,A.unicolor). The species are arranged according to their relationships into species groups within each subgenus. Sixty-one synonymies are newly established (the first specific name is valid): Subg. COPROCHARA,A.bimaculataGravenhorst =B.deserticolaCasey =B.innocuaCasey =B. obsolescens Casey =B. recta Casey;A.notulaErichson =A.duplicataErichson =B.nanellaCasey =B.nitidicollisCasey;A.suffusa(Casey) =B.acuminataCasey =A.verna brundiniBernhauer;A.vernaSay =A.alticolaSharp. Subg.XENOCHARA,A.lacertinaSharp =B.salicolaCasey =B.imbricataCasey =B.idoneaCasey =A.densiventrisBernhauer;A.sallaeiSharp =A.torquataSharp;A.puberulaKlug =B.bipartitaCasey;A.fumataGravenhorst =A.languidaSachse =B.defectaCasey =B.affluensCasey;A.castaneipennisMannerheim =A.glenoranaCasey =B.rotundicollisCasey =B.mannerheimiCasey =B.acomanaCasey =B.insulanaCasey =B.concurrensCasey =B.eludensCasey =A.oregonaHatch;A.quadrataSharp =B.uvidulaCasey =A.sparsicollisBernhauer. Subg.CALOCHARA,A.rubripennisCasey =A.rubripesBlatchley =A.wallawallaeHatch. Subg.ALEOCHARA,A.centralisSharp =A.stygialisSharp;A.gracilicornisBernhauer =A.kansanaCasey =A.pleuralisCasey =A.americanaCasey =A.ellipsicollisCasey;A.lustricaSay =A.pauperSharp =A.serrataSharp =A.texanaCasey =A.fusicornisCasey =A.sternalisCasey =A.algonquinaCasey =A.medialisCasey;A.tahoensisCasey =A.postpictaCasey =A.montanicaCasey;A.thoracicaCasey =A.collusorCasey. Subg.ECHOCHARA,A.lucifuga(Casey) =A.cavicola(Garman). Subg.EMPLENOTA,A.littoralis(Mäklin) =A.maritima(Casey) =P.arenariaCasey =E.quadriferCasey =E.trilimbataCasey =E.longicepsCasey. Subg.MASEOCHARA,A.validaLeConte =M.ponderosaCasey =M.robustaSharp =M.ruficaudaCasey;A.opacella(Sharp) =M.hogeiSharp =M.decipiensCasey;A.depressa(Sharp) =M.puberulaCasey =M.basalisCasey =M.mustaCasey.Ninety-three lectotypes and three neotypes are designated (neotypes are designated for:C.caviolaGarman,A.lustricaSay, andA.vernaSay).Keys to subgenera and species are provided. All species are described, each illustrated with line drawings and scanning electron microscope photomicrographs, and all available bionomic and distributional data are given. Emphasis was put on host records of thoseAleocharaspecies which may be used in biological control of some pest species of Diptera. The systematics and phylogeny of the subgenera and zoogeography of species are discussed.A checklist of NearcticAleocharaand a table listing the known host records for some species is added.

Aleochara sp. from southern Africa is a potential candidate for introduction to Australia for control of the Australian buffalo fly,Haematobia irritans exigua De Meijere. Aspects of its host searching and acceptance behaviour were studied in South Africa usingHaematobia thirouxi potans (Bezzi), to assess, in part, its potential value as a biological control agent and to provide a basis for the development of a mass rearing technique. First instars ofAleochara sp. were found to parasitize most effectively those hosts that had buried themselves in the substrate, probably by orienting to a chemical cue and using the tunnels made by the post-feeding larvae. Lower rates of parasitization occurred in pupae placed on the surface of the substrate or buried by the experimenters. They would accept, parasitize and develop on all ages ofH. thirouxi potans pupae, but showed reduced survival in the very oldest ones. Although there was no difference in relative acceptability among hosts of different ages, a higher proportion of younger hosts were parasitized when these had buried themselves, probably due to a tendency of the parasitoids to follow fresher tunnels into the substrate. Survival was high (>80%) at relative humidities from 10–91% but was less than 20% at an R.H. of 100%. In rearing trials, the most economical ratio of parasitoid eggs: hosts was 1∶1.5, with 70% of the parasitoids developing successfully to adults.

A simple ovipositon device, containing inert, expanded, clay granules, has been developed for both the house fly, Musca domestica L., and the staphylinid beetle, Aleochara bilineata Gyllenhal. For the house fly and the Aleochara beetles, the optimum particle size of the granules is 2.0 to 2.8 mm and 1.0 to 2.0 mm, respectively. A quick method for extracting the eggs of both species from the oviposition medium, by washing, is also described.

A review of David Sharp's types of Aleochara from Mexico, and Central and South America is presented. Nineteen species are treated here, one of which is described as new (A. felixiana), seven are redescribed [A. angusticeps Sharp, A. funestior Sharp, A. mexicana Sharp, A. oxypodia Sharp, A. simulatrix Sharp, A. gracilis (Sharp), and A. duplicata (Sharp)] and one [A. miradoris Sharp] is put in synonymy under A. mexicana Sharp. The remaining ten species were treated in my 1984-revision of North American Aleochara but are here included in a checklist with references to description, redescription, and illustrations. Some additional information is provided for previously synonymized A. torquata Sharp (= A. sallaei Sharp), and for A. quadrata Sharp. Illustrations of genitalic structures of both sexes are provided for the first time for the seven redescribed species. Lectotypes are designated for A. angusticeps, A. duplicata, A. funestior, A. mexicana, and A. oxypodia.

The native parasitoid complexes associated with four key muscids, Musca domestica L., Stomoxys calcitrans (L.), Fannia canicularis (L.), and F. femoralis Stein—and two predatory Diptera, Muscina stabulans (Fallen) and Ophyra leucostoma (Wiedemann)—are described and shown graphically for the major climatic zones of the southwestern United States. Three geographical groups of the parasitoid species identified were (1) those whose populations extended into most climatic zones below 2,000 meters (Muscidifurax species and Spalangia nigroaenea Curtis); (2) those whose populations were restricted to the warmer climatic areas (S. endius Walker and S. cameroni Perkins); and (3) those that were largely restricted to higher latitudes and colder winter climates (S. nigra Latreille and Aleochara spp.). Considerable yearly, seasonal, and climatic variations were shown in the abundance of both host and parasitoid demes. These factors, combined with others, probably account for the high degree of instability among demes. This instability may be important in determining the panmictic nature of most populations. Some parasitoids such as the Diapriidae and Staphylinidae (Aleochara spp.), which exist in small demes, appeared to be local species that may have adapted to invading Diptera. The Stilpnus species were specific to Fannia. Parasitoid species appear to be irreplaceable mortality factors in the natural control of populations of their dipterous hosts.

Six species of Staphylinidae under the genera Philonthus Curtis, Coenonica Kraatz, Aleochara Grouvelle and Oligota Mannerheim are reported from 18 different commodities in Kolkata area. All these species are new to the stored product habitat from India. Their diagnostic features with illustrations, distribution and habitat records are dealt with and an identification key to these species is given. Key words: Coleoptera, Staphylinidae, Systematics, Stored products, Kolkata J. bio-sci. 14: 49-55, 2006

Aleochara taeniata Erichson, a staphylinid parasite of the house fly, Musca domestica L., was cultured in California after its introduction from Jamaica. Adults of the species are voracious predators of house fly eggs and young larvae. Eggs hatch in about 4 days in the laboratory, and larvae search actively through the substrate for fly pupae, the parasite-susceptible host stage. Pupae are entered through a hole gnawed in the puparium wall by the young larvae; the hole is closed with what appears to be fecal material. The 3 larval instars are ectoparasitic on the fly pupae within the puparium. Mature larvae emerge into the substrate where pupation occurs. Larval development requires 6-7 days, and pupal development another 14-16 days at 75°F. The total life cycle at this temperature approximates 25 days from egg to adult.

Field parasitism of the face fly, Musca autumnalis De Geer, by insect parasites and the nematode, Heterotylenchus autumnalis Nickle, was investigated in south central Nebraska. Four species of insect parasites were collected from face fly puparia: Aphaereta pallipes Say, Muscidifurax sp., Eucoila sp., and Aleochara sp. A. pallipes was found only during the first 4 wk of the study (6–24 June), whereas the other three parasites were intermittently found, at low levels, for the remainder of the study (24 June–26 September). Total seasonal insect parasitism averaged 4.3%. Seasonal parasitism by the nematode H. autumnalis averaged 5.8% and peaked at 18% on 9 September.

This work was carried out in order to determinate the local fimiculous Staphylinidae species due to these predators and or parasites importance on the natural control of the dung associated bovine parasites. For that, fecal masses 1, 2 and 3 days old were weekly collected on a pasture of Brachiaria decumbens Stapf, from May 1990 through April 1992. The feces were conditioned in 15-liter opaque plastic buckets, containing lateral and top openings, where flasks were fastened for capturing Staphylinidae beetles present in these masses, during a 40-day period. After this period the soil and fecal masses from the buckets were examined and the remaining insects collected. A total of 13215 specimens were collected, belonging to 34 species. The following twelve Genera were observed: Oxytelus (3 species; 70.1% of the specimens collected); Falagria (1 sp.; 7.9); Aleochara (4 sp.; 5.8); Philonthus (3 sp.; 5.1); Atheta (2 sp.; 4.0); Cilea (2 sp.; 1.2); Neohypnus (1 sp.; 0.7); Lithocharis (1 sp.; 0.7); Heterothops (2 sp.; 0.6); Somoleptus (1 sp.; 0.08); Dibelonetes (1 sp.; 0.06) and, Dysanellus (1 sp.; 0.04). The seven most abundant, constant, and or frequent species, followed respectively by the percentage of collected specimens, were: Oxytelus sp.1 (38.8%); Oxytelus sp.2 (28.1); Falagria sp. (7.9); Philonthus sp.1 (4.1); Atheta sp.1 (3.7); Aleochara sp.1 (3.6), and Oxytelus sp.3 (3.2). Twelve species (3.6% of the collected specimens) were determined as far as family only. Este trabalho foi executado com o objetivo de determinar as espécies locais de estafilinídeos fimícolas, devido à importância destes predadores e ou parasitóides no controle natural de parasitos de bovinos associadas às fezes. Para tanto, massas fecais com 1, 2 e 3 dias de idade foram coletadas semanalmente em uma pastagem de Brachiaria decumbens Stapf, no período de maio de 1990 a abril de 1992. As fezes foram acondicionadas em baldes plásticos, opacos, com capacidade para 15 litros, contendo aberturas lateral e no topo, onde foram fixados frascos para a captura, por um período de 40 dias, dos besouros estafilinídeos presentes nas massas fecais. Após este período a massa fecal e o solo existente nos baldes eram examinados e os insetos remanescentes recolhidos. Foi coletado um total de 13.215 exemplares, pertencendo a 34 espécies e/ou morfo espécies. Foram observados os seguintes doze gêneros: Oxytelus (3 espécies; 70,1%); Falagria (1 sp.; 7,9); Aleochara (4 sp.; 5,8); Philonthus (3 sp.; 5,1); Atheta (2 sp.; 4,0); Cilea (2 sp.; 1,2); Neohypnus (1 sp.; 0,7); Lithocharis (1 sp.; 0,7); Heterothops (2 sp.; 0,6); Somoleptus (1 sp.; 0,08); Dibelonetes (1 sp.; 0,06) e, Dysanellus (1 sp.; 0,04). As sete espécies mais abundantes, constantes e ou freqüentes, seguidas, respectivamente, pelo percentual de indivíduos coletado, foram: Oxytelus sp.1 (38,8%); Oxytelus sp.2 (28,1); Falagria sp. (7,9); Philonthus sp.1 (4,1); Atheta sp.1 (3,7); Aleochara sp.1 (3,6) e Oxytelus sp.3 (3,2). Doze presumíveis espécies (3,6% do total coletado) foram identificadas apenas até família.

The species composition and abundance of coprophilous predatory Staphylinidae in two sites of temperate and subtropical Argentina are presented and discussed in terms of their hypothetical effect on the dung breeding fly community. Specimens of Staphylinidae were collected from cow manure by flotation. Dung breeding fly puparia were also collected to obtain parasitoidal Staphylinidae. Roughly the same number of species, and the same species of common staphylinids were found in both sites. Also, the dominant species were almost the same: Philonthus pauxillus Solsky, P. flavolimbatus Erichson, and Aleochara verberans Erichson. However, Philonthus sp. 1, and Aleochara notula Erichson were very abundant in Buenos Aires, but rare in Tucuman, and Philonthus nv. sp., was common in Tucuman but not found in Buenos Aires. Of the parasitoid species in the genus Aleochara Gravenhorst, A. notula was recovered from Oxysarcodexia varia (Walker), Sarcophagula occidua (F.), and Palaeosepsis discolor (Bigot) (Seps...

The larval stages of all species of Aleochara in which life histories have been investigated are ectoparasites on the pupae of Diptera within the puparium. The adults are active predators of dipteran eggs and larvae, often destroying enormous numbers. This fact has been demonstrated conclusively with Aleochara bilineata , a parasite of the cabbage maggot, Hylemya brassicae (Wadsaworth 1915, Colhoun 1953). Other species of Aleochara have not been so extensively investigated, but some host records are known. A table of these is given in which new host records are reported.

The coastal staphylinids are introduced for the first time from Dokdo Island. Three coastal species, Aleochara (Emplenota) fucicola Sharp, Atheta (Badura) tokiokai (Sawada) and Cafius histrio (Sharp), were identified and are reported newly from Dokdo island. Their habitus and information about species distribution are provided.

SUMMARY (1) Host acceptance and suitability of a range of dipterous hosts of the predator/ parasitoid Aleochara sp. (Coleoptera: Staphylinidae, Aleocharinae) were studied in South Africa to assess the likelihood of the parasitoid attacking the novel target pest Haematobia irritans exigua De Meijere, in Australia. (2) In laboratory experiments, first instar parasitoids were presented with puparia of Haematobia thirouxi potans (Bezzi) and other candidate host species in choice and nochoice tests. Parasitized puparia were reared to determine survival and parasitoid size. (3) Some species were not acceptable as hosts. The parasitoid larvae were apparently attracted to, but could not penetrate the puparium of Musca xanthomelas Wiedemann. They penetrated, but did not feed on, Musca sorbens Weidemann and died in the first instar. (4) The remaining hosts tested, Borborillus marginatus (Adams), Sepsis thoracica Robineau-Desvoidy, H. thirouxi potans, Musca domestic L. and Orthellia peronii (Robineau-Desvoidy), were penetrated and consumed, with resulting development to a range of parasitoid sizes and with variable survival rates. The relationship between a preference index and the loglo of host mass was quadratic, with preference relative to H. thirouxipotans increasing with increasing host mass up to 8 mg and decreasing thereafter. Parasitoid mass increased but survival decreased with increasing host mass. (5) From the size of field-collected Aleochara sp., the relationships determined in the laboratory, and the ecology and mass of H. irritans exigua in Australia, it is considered likely that Aleochara sp. would accept and develop successfully on the target species in Australia.

Review of Erichson's types from Mexico, West Indies and South America is presented. Nine species are treated herein, of which three are redescribed (A. breviuscula Er., A. chrysorrhoa Er. and A. lateralis Er.), one has been previously synonymized by Klimaszewski (1984), with first name being valid (A. notula Er. = A. duplicata Er.), two are newly transferred to different genera (A. peltata Er. to Hoplandria Dr., and A. restricta Er. to Tricolpochila Bernh.), and three species were previously revised and redescribed by Klimaszewski (1984) and Klimaszewski and Genier (1986), A. notula Er., A. taeniata Er. and A. verberans Er.). Six lectotypes are designated here. A checklist of species is presented with reference to the original description, original material and the main source of reference. A discussion, illustrations and/or reference to the most reliable redescription are provided for each species.

Recent observations (Wright 1956, 1957, 1958) have shown that certain chemical control measures may increase the infestation of the roots of brassica crops by the larvae of Erioischia brassicae (the cabbage root fly). This work led to the conclusion that the insecticides were interfering with a system of natural control of the larval population. To aid the interpretation of these results detailed investigations of the population dynamics of E. brassicae were begun in 1955, together with studies of the biology and habits of associated parasites and possible predators. There are many references in the literature to organisms responsible for mortality at different stages in the development of E. brassicae. Predation of the egg stage by trombidiid mites (Schoene 1916, Gibson & Treherne 1916, and Lowry 1915), ants (Schoene 1916) and by carabid and staphylinid beetles (Wishart, Colhoun & Monteith 1957) has been recorded. The last team of workers stained eggs with a solution of Bengal rose and placed groups of marked eggs around plants in the field. Collections made after periods of 24, 48 and 72 hours showed large daily losses of marked eggs. Mortality in the larval stage has been caused by predatory ants (Schoene 1916), beetles (Gibson & Treherne 1916, de Wilde 1947) and other anthomyiid larvae (Smith 1927). Many hymenopterous parasites attack the larval stages but the mortality occurs after the puparium is formed. Five species of the Braconidae, three of the Cynipidae, and four of the Ichneumonidae, have been reared from E. brassicae puparia (Wadsworth 1915, James 1928, Paillot 1914, Schoene 1916, Vodinskaya 1928, Colhoun 1953, de Wilde 1947, Nixon 1939, and Wishart, Colhoun & Monteith 1957). Of these parasites only the cynipid Idiomoryha rapae (Westwood) is of major importance. Parasitism of the pupal stage by species of Aleochara (Coleoptera, Staphylinidae) has been recorded by most of the authors cited above. Predation of the adult flies by anthomyiids (Schoene 1916) and by cordylurids (Hobby 1931, Reid 1958) has been recorded. The adult stage is also attacked by the entomogenous fungus, Empusa musca (Schoene 1916). Various techniques have been developed to estimate the populations of the immature stages of Erioischia brassicae, and while the present paper deals only with the population dynamics of the first generation, the same techniques have also been used in investigations of the later generations of the life-cycle. The