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Bethylidae = Link 1
Biology and Behavior
Bethylidae. -- Bethylids are small to medium-sized, dark-colored wasps; the females of many species are apterous and antlike in appearance. In some there are both winged and wingless forms. They are parasitoids of the larvae of Lepidoptera and Coleoptera and several species attack moths or beetles that infest grain or flour. A few species will sting humans. Goniozus legneri Gordh has been deployed successfully in the biological control of navel orangeworm, Amyelois transitella (Walker).
. Bethylidae are of special interest because of their mating and food habits, the maternal care of the brood and a tendency in some species toward community life. Extended summaries of the habits of the family were presented by Bridwell (1919, 1920) and Wheeler (1928), but the most thorough study of a species was that by van Emden (1931) on Cephalonamia gallicola Ashm., a gregarious external parasitoid of the larvae and pupae of the anobiid beetle, Stegobium paniceum L. in Europe. Kearns (1934a) studied the same species as a parasitoid of the cigarette beetle, Lasioderma serricorne F. in North America.
Regarding adult habits in the gregarious species of the family, the males of the brood usually emerge a little in advance of females. They have the habit of then tearing open the female cocoons and, often, of entering them in order to mate with the newly transformed females. This habit of males assisting in the emergence of females results in very extensive inbreeding, which is normal in many species. However, its effects are offset by the repeated mating that is necessary during the life of individual females, and the matings following the first are in most cases with males of unrelated broods. The closest inbreeding occurs in species that are able to produce several successive broods on a single host individual. Under such conditions, a female may mate with males of several generations of her own descendants (Clausen 1940/1962).
Behavior of the adult female in her attack on prey, the method of feeding and the tendency toward maternal care of the progeny are distinctive. In the initial attempt to overcome the host larvae, the female parasitoid usually crawls over the dorsum and curves her abdomen beneath the thorax, inserting the sting first in the throat or in one of the thoracic segments. This obviously is for the purpose of paralyzing the mandibles and legs. In Sclerodermus chilonellae Brid., the initial stinging is by a backward thrust toward the mouth. Repeated stinging takes place, the later insertions being in the abdominal region. In Laelius anthrenivorus Trani, the female then chews the throat, apparently with the object of injuring the cervical ganglion. The complete process of subduing the host may take several hours (Clausen 1940/1962).
. Some species attack their hosts in the open. After stinging, they transport the host to a suitable crevice or cavity in the manner of fossorial wasps. Bethylus fulvicornis Curt. stores its caterpillar hosts in hollow stalks. The host is usually much larger then the female bethylid and thus is dragged along the surface, although Epyris extraneus Brid and Cephalonomia mycetophila Kieff. carry it across the dorsum (Clausen 1940/1962). Host paralysis is usually complete and permanent, though in the above species and in several species of Goniozus it is only temporary (1/2 hr duration). This is not true of G. legneri and G. emigratus.
. Adult females feed on the body fluids of the host which is apparently essential for egg development as witnessed by the frequent lapse of several days between such feeding and the deposition of first eggs. In many species females spend their entire lives in the burrow, cell or cocoon of the host where the body fluids are the only food available. Females of the genus Laelius have not been observed to host feed. Bridwell believed that all members of the Goniozus group require sugary foods, Sclerodermus fed only on host fluids, and the Epyris group fed on both. A preoviposition period has been found to be a minimum of 5 days. The number of hosts, which are attacked and paralyzed by a female, is often considerably in excess of that which receive eggs. Ramachandra Rao & Cherian (1928) stated that females of Perisierola nephantidis Mues. were observed to mob full-grown caterpillars and kill them. In Prorops nasuta, the female frequently feeds on young coffee borer larvae, though oviposition is entirely on mature larvae. It seems that the total host mortality effected by bethylids may be considerably in excess of that resulting from successful parasitization. The females of the genus Laelius, attacking larvae of Dermestidae, bite away the covering of long hairs on the venter of the abdomen before depositing their eggs (Howard 1901).
. Complete and permanent paralysis of the host is the usual case in Bethylidae. In one exception, Goniozus sp. (Taylor), parasitizing Nacoleia octosema Meyr in Java, the caterpillar recovers from the sting and continues normal feeding until parasitoid eggs hatch, when it spins a scanty cocoon (Clausen 1940/1962). The caterpillar host of G. claripennis Foerst (Voukassovitch 1925b) also recovers within two hours after stinging and may on occasion molt. Sometimes a light cocoon may be spun before death.
The position of the egg on the host varies, though there is some uniformity within genera. Eggs of solitary species lay longitudinally, with the anterior end directed caudad. The eggs of Prorops nasuta are ventral on thorax; that of Parascleroderma berlandi Man. (Maneval 1930) is attached at the side; in E. extraneus, the egg is on the venter of the first abdominal segment; and in Holepyris hawaiiensis Ashm. it is on the dorsum of the posterior portion of the abdomen. Among the species of gregarious habits, the eggs are usually placed transversely, exceptions to this being Perisierola gallicola Kieff, where eggs lie longitudinally on one side of the dorsum, and Laelius spp., which lays them in two rows on the venter of the abdomen. Cephalonomia gallicola Ashm. and Bethylus cephalotes Foerst. place them in two rows on the venter of the abdomen of host larvae, but in the case of Cephalonomia the position is changed to the intersegmental grooves of the dorsum when the pupal stage of the host is attacked. C. mycetophila Kieff. differs in habit from other species of the genus by placing its eggs on the dorsum of the thorax of host larvae (Clausen 1940/1962). A variation found in C. tarsalis Ashm., parasitic on Oryzaephilus larvae, is that only two eggs are placed on each host; the first one deposited, which is destined to develop into a female, is placed on the side of the prothorax, and the second, which is always male, is deposited on the mesothorax (Powell 1938). Some Goniozus deposit their eggs transversely in the intersegmental grooves of the mid-dorsal region.
. Females accomplish oviposition in various ways. In species placing eggs longitudinally with respect to the host body, the female takes the venter-to-venter position during egg laying, whereas those which place the egg transversely in an intersegmental groove encircle the body of the host in a manner identical with that of the Tiphiidae (Clausen 1940/1962). Evidence suggests that females of the different gregarious species regulate to some extent in relation to its size, the number of eggs deposited on a single host. Perisierola emigrata Roh. deposits only two eggs on small Ereunetis larvae and an average of ca. eight on larger larvae of Cryptophlebia.
. The total number of eggs deposited by a single female ranges to a max. of 150, although Willard (1927) recorded 236 from one female of P. emigrata in 44 days. A female of Cephalonomia gallicola laid 158 on 76 hosts during 36 days; and 42 Lasioderma larvae were killed but not parasitized (Kearns 1934a,b). Mated females of C. tarsalis deposit ca. 85 eggs, whereas virgin females produce only 50. Solitary species probably produce smaller numbers of eggs. In the gregarious species an interval of several days must elapse between the deposition of successive batches of eggs.
. Regarding larval development, the newly hatched larvae are for the most part unable to move and therefore make their feeding punctures and fix themselves before they are completely freed from the eggshell. A single feeding puncture is used throughout their life. In the late first stage, the larva of Goniozus claripennis is U-shaped, with a fold of host skin firmly held between the two ends of the body, which anchors it firmly in position. In the intermediate and late larval stages of Laelius anthrenivorus, the head and thoracic segments are embedded in the wound, and often more than half the body of the larva is within the host. During the first portion of the feeding period the larva lies horizontally on the body of the host, but as development proceeds the parasitoid body becomes more spindle-shaped, and in several species and genera, it assumes a vertical position (Clausen 1940/1962). This usually does not happen until the last stage. The larvae which lie in the horizontal position retain the exuviae as a pad beneath the body, whereas whose that stand vertically carry them at the posterior end of the body. Hyslop (1916) mentioned an "irregular shield of rugose lemon-yellow skin bearing two hairs," in Pristocera armifera Say, which Clausen (1940) believed to be a cast skin.
. Maternal care of developing offspring is exhibited in a number of species, ranging from attention for only a few days after oviposition in C. gallicola up to the completion of larval development in S. immigrans and S. macrogaster (Clausen 1940/1962). Wheeler mentioned that the females of the latter often stand over the brood and lick their bodies, at which time they are held between the forefeet. They will at times eat their own eggs, but not their larvae, and this habit is thought to be associated with a tendency toward regulation of the size of the brood rather than being the result of a food shortage (Clausen 1940/1962). Bridwell mentioned that sometimes several females of Sclerodermus were found living together and rearing their progeny on a single host larva, and without any indications of interference or cannibalism, even though the females were sometimes of different species.
. The larval feeding period is very short, being complete in 2-3 days in several species of Goniozus and Cephalonomia, while the majority most likely require ca.a 5 days. The maximum duration of this period was 10 days by B. cephalotes.
. Larvae which have completed development are quite active and able in some species to move several centimeters away for pupation. Most species spin cocoons, which in the gregarious species are usually matted together. Parascleroderma berlandi spins a transverse operculum across the burrow of the host before forming the cocoon. An unusual pupation habit was recorded for Parasierola sp., parasitic on the larva of the strawberry leaf roller, Ancylis comptana Froel. in North America (Fink 1932). No cocoon is formed, but the mature larva fastens the tip of its abdomen to the leaf surface and then transforms to a pupa within the last larval skin, which becomes jet black in color (Clausen 1940/1962).
. There is a lot of variation in size among the individuals that develop on a single host, due largely to crowding. Where the number of eggs deposited is above that which can be brought to maturity, the surplus larvae succumb, presumably by being forced away from the available food supply. However, superparasitism has not been officially reported in the Bethylidae (Legner & Warkentin 1988).
. Reported life cycles of various Bethylidae range from 9-12 days minimum to a more frequent 30-35 days. In C. gallicola 34 days at 22.6°C and 64 days at 18.4°C are required. However, S. domesticus Kieff. seems to require at least two months for development. The egg stage covers 1-4 days with an extreme of 7 days in the case of Parascleroderma berlandi, though that of C. gallicola extends to 13 days at the minimum temperature shown above. The larval feeding period is usually followed by a prolonged resting period in the cocoon prior to pupation. The time spent in the cocoon may range from 10-20 days, though Perisierola nephantidis on Nephantis serinopa Meyr. in India is said to require only four days (Ramachandra Rao & Cherian 1928). In C. gallicola Kearns found the females, winged males and apterous males required 11-18, 9-15 and 7-12 days, respectively in the cocoon. Adult life of females ranges from one to three months under summer conditions with hosts available for feeding and reproduction. The life of adult males is much shorter, being only 6 days in C. tarsalis, and no feeding takes place during this period. The winter may be passed either in the adult stage in sheltered places or in the mature larval or pupal stage in cocoons. In some species two generations are produced during the time required for one of the host.
Regarding sex ratio, females predominate numerically in all species studied. This ranges from 2 to 1 in S. domesticus to 5 to 1 in S. immigrans. Rearings of S. immigrans by Keeler (1929a,b) showed that only female progeny resulted from unmated wingless females. This study revealed two biological races, and Clausen (1940) thought it surprising that thelytoky should have developed in a race of one species and not elsewhere in the family. Taylor found that the brood of 3-5 Goniozus sp., which develops on the larva of the banana scab moth, usually includes a single male. Wheeler (1928) stated that oviposition by virgin females of S. macrogaster is conspicuously delayed, and Soika (1932) asserted that parthenogenetic reproduction cannot occur in S. domesticus.
. A study of Goniozus legneri Gordh by Legner & Warkentin (1988) showed that oviposition behavior was significantly influenced by variations in host/parasitoid densities. Oviposition rates tended to increase with increases in host density. Sex ratios, although strongly female biased (>82%), were significantly influenced by host/parasitoid density within a range of ca. 10%.
Clausen (1940) discussed in some detail the morphological characteristics of immature stages of Bethylidae.
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