BIOLOGY semipunctata (Coleoptera: Cerambycidae), a Serious Pest of in California: Biology and Laboratory-Rearing Procedures

LAWRENCE M. HANKS, J. STEVEN McELFRESH, JOCELYN G. MILLAR, AND TIMOTHY D. PAINE Department of Entomology, University of California, Riverside, CA 92521

Ann. Entomol. Soc. Am. 86(1): 96-102 (1993) ABSTRACT Procedures are described for establishing a laboratory colony of the euca- lyptus longhorned borer, F., and rearing the adult on a continual basis. Adult beetles reared from naturally infested Eucalyptus logs were caged and provided with oviposition substrates (folded sheets of plastic). Techniques for han- dling and caring for eggs and neonate larvae are discussed. Larvae were individually transferred into shallow incisions in the bark of fresh logs. Total survivorship from neonate larvae to adult was =35%. Our procedures yielded an average of 63 adult progeny for every adult female, with a generation time of =2 mo during the summer. colonies were protected from pyemotid mites by dusting rearing logs with sulfur and from ants with granular diazinon. An experiment on adult beetle diet showed that the provision of sucrose water greatly increased longevity and fecundity over a distilled water control, but increas- ing the concentration of sucrose >5% did not significantly improve beetle performance.

KEY WORDS rearing, Eucalyptus, Phoracantha semipunctata

DIFFICULTY IN REARING cerambycid beetles has len of Eucalyptus flowers (Chararas 1969a, long hindered research on these (e.g., Scriven et al. 1986, Hanks et al. 1990). However, Linsley 1959, Payne et al. 1975). Maintaining adults of both sexes are strongly attracted to cerambycids on artificial diet has sometimes stressed Eucalyptus trees and logs of any size as proven effective for taxonomic or cytogenetic oviposition sites (Chararas 1969b, Drinkwater studies for which relatively small numbers of 1975, Ivory 1977, Gonzalez-Tirado 1986, Scriven individuals were needed (e.g., Gardiner 1970, et al. 1986, Mendel 1987, personal observation). De Viedma et al. 1985). However, rearing ceram- Females oviposit under loose bark and in bark bycids on artificial diet for behavioral or ecolog- fissures in batches of up to 40 eggs. Neonate ical studies may have three drawbacks: (1) it may larvae then penetrate the bark and feed along the be difficult to develop a diet that beetles will eat cambium. and that provides all the necessary nutrients; (2) because cerambycids typically have pro- Heavy infestations of P. semipunctata larvae longed larval stages (Linsley 1962), rearing on an result in destruction of virtually the entire cam- artificial diet may be very labor intensive be- bium layer and the rapid death of the tree cause of the need to transfer larvae to fresh me- (Chararas 1969b, Drinkwater 1975, Scriven et al. dia periodically (Linit 1985); (3) artificial diet 1986, personal observation). Fully grown larvae may influence adult physiology and behavior, burrow into the sapwood where they construct a resulting in differences between field and labo- pupal chamber, plugging the passage to the log ratory insects (e.g., Stanic" et al. 1989). Rearing surface with wood chips. During the warm sea- cerambycids in their natural hosts eliminates son, the larvae rapidly pupate but will become these dietary artifacts and can be simpler and quiescent over the winter. Overwintering larvae more productive than using an artificial diet. pupate in spring and then emerge as adults, The eucalyptus longhorned borer, Phoracan- chewing their way out through the pupal plug in tha semipunctata (F.), is a severe pest of euca- late April to mid-May in southern California lyptus in many countries (Chararas 1969a, Ivory (personal observation). Emergence is not tightly 1977, Gonzalez-Tirado 1986, Mendel 1987) and synchronized, and adult beetles are present con- appeared in southern California in 1984 (Scriven tinually from spring until late fall. Flight activity et al. 1986). Adults are nocturnal, resting under of the adult beetles is restricted to warm nights bark during the day, and do not damage host when the temperature at dusk is above =15^ trees themselves but feed on the nectar and pol- (Chararas 1969b, Loyttyniemi 1983). In southern

0013-8746/93/0096-O102$02.00/0 © 1993 Entomological Society of America January 1993 HANKS ET AL.: REARING EUCALYPTUS LONGHORNED BORER 97

California, P. semipunctata completes two over- The counter top on which cages were kept was lapping generations per year (Hanks et al. 1990). lightly dusted with talc (Humco Laboratory, Tex- Artificial diets have been developed for P. arkana, TX) to inhibit the movement of pyemotid semipunctata (Chararas 1969a, De Viedma et al. mites (Hanks et al. 1992). As many as 20 beetles 1985), but for the reasons described above were were kept in each aquarium. Each cage was pro- unsuitable for the large-scale, continuous pro- vided with 30% sucrose solution in 8-ml vials duction of adults that we required. We present with a 4-cm-long number 2 cotton roll (Patterson here information on techniques for rearing P. Dental, South Edina, MN) inserted halfway into semipunctata on its natural host which are effec- the vial. We provided one of these feeders for tive in producing large numbers of beetles, and every three beetles. A small amount of honeybee we provide new information on the biology of pollen (0.5 g per cage) was also provided. this important pest. Oviposition substrates consisted of sheets of 6-mil polyethylene sheeting cut into strips (10 cm long, 6 cm wide) that were stacked five Materials and Methods strips thick, folded once lengthwise, and stapled. Rearing and Handling Adult Beetles. We ob- These folded sheets were laid on the top screen tained logs from several species of Eucalyptus of the cage with their loose-leaf sides down, pro- that were naturally infested by P. semipunctata viding an abundance of folds into which a fe- from campuses of the University of California at male could insert her ovipositor up through the Riverside and San Diego. These logs were <50 screen. These oviposition substrates worked best cm in circumference and were cut into lengths of when weighted down slightly to keep them in =0.5 m for ease of handling. On 19 March 1990, place. We replaced the oviposition substrates we placed these logs upright in screened cages daily so that all eggs in each substrate were of the (1.3 m high, 0.65 m deep, 0.55 m wide). Because same age, thereby synchronizing hatching. adult P. semipunctata are negatively geotactic, Handling Eggs and Rearing Larvae. To de- we could collect emerging adult beetles by cap- termine survivorship of eggs to hatching, 200 ping the cage with a pyramidal screen top termi- freshly laid eggs were monitored until they nating in an inverted plastic funnel over which hatched under ambient laboratory conditions was positioned a 2-liter cardboard container. (=20°C and =40% RH) in March 1992. Adult beetles were collected from cardboard Neonate larvae were manually transferred to containers daily to reduce injury from fighting. logs to control larval density and to provide co- Cages for rearing adults were kept in a con- horts of the same age. Logs for rearing P. semi- trolled environment chamber (=30 ± 5°C under punctata larvae were cut into lengths of 0.5 m natural daylight; humidity was not controlled from freshly cut and was near ambient with a mean of =40% RH). Dehnhardt trees having an average trunk basal Sulfur dust (F.M.C., Philadelphia, PA) was lib- circumference of 40 cm. P. semipunctata larvae erally applied to the logs (=20 ml per cage) to require logs that are fresh to complete develop- inhibit attack by a pyemotid mite that rapidly ment (Chararas 1969b, Mendel 1985). To pre- paralyzed emerging beetles (Hanks et al. 1992). serve the moisture content of logs, we waxed the Cages were examined frequently for moribund ends of the logs by dipping them in paraffin wax or dead beetles that might be paralyzed by mites (Bromar, Newport Beach, CA) heated until just and later become a source of mite infestations. smoking. Allowing the logs to dry out for 24 h To examine the temporal pattern of emergence before waxing reduces bark turgidity which can of adult beetles from logs, we recorded the num- inhibit the movement of neonate larvae in the bers of beetles emerging in five cages on a bark and prevent their establishment (Hanks et weekly basis until logs were no longer produc- al. 1991a). Waxed Eucalyptus logs stored at 5°C tive. Each cage contained logs originating from remained viable hosts for the larvae for as long as the same location (either University of California 2 mo. Riverside or San Diego) on the same date (during Larvae were introduced into logs by cutting a spring 1990). slit in the bark a few millimeters wide and deep Adult beetles captured in cage traps were han- enough to reach the cambium, then transferring dled by their heavily chitinized antennae with the larvae into this slit with a damp fine-tipped plastic-tipped forceps. This handling technique paintbrush (size 00000). Once in place, the lar- prevented the beetles from holding onto the for- vae were protected by a cover of paper (such as ceps and did not appear to damage the antennae. paper toweling) taped over the slit, and the logs Adult beetles were maintained under laboratory were left in place for 24 h to prevent the larvae conditions (18-24°C, =40% RH, natural day- from being dislodged before they bored into the light). cambium. Because survivorship of larvae de- Egg Production. To obtain P. semipunctata creases with their density in the log (Hanks et al. eggs in quantity, adult beetles were housed in 1991b), the number of larvae introduced into the glass aquaria (50 cm long, 26 cm wide, 30 cm logs should be such that the resulting density is deep) covered with aluminum window screen. low (=50 mature larvae per square meter of bark 98 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 86, no. 1 represents an optimal density for E. camaldulen- tom. A 12.5-cm-diameter circle of no. 1 filter sis [L.M.H., personal observation]). Using an ini- paper (Whatman International, Maidstone, En- tial density only slightly in excess of this density gland) was placed in the bottom of the cage. ensures that the maximum density will be Distilled water and sucrose or honey solutions reached but without significant mortality from were presented in glass vials (as described competition. We infested nine logs of E. camald- above). Two vials were put into each cage, laid ulensis (surface area 0.24 ± 0.02 m2 with 20 lar- on their sides in a smaller petri dish (8.5 cm vae (average initial density of 83 larvae per diameter) to keep them from rolling onto beetles. square meter of bark) on 16 August 1990 and Both water solutions and pollen were changed wrapped each log in organdy to capture emerg- every other day. ing adult beetles. Logs were maintained in a Phoracantha semipunctata mated readily and temperature-controlled rearing facility (=30 ± repeatedly during the active period (at night), 5°C, =40% RH, natural daylight) until emer- and all eggs produced were fertile. In the screen gence ceased. It was crucial that ants be ex- cages, females oviposited readily between the cluded from rearing facilities because the bark of filter paper and the smaller petri dish, making it heavily infested logs often cracks, exposing the easy to determine fecundity. At least five pairs of defenseless larvae or pupae to predation. We beetles were used in each treatment, and longev- sprinkled granular diazinon (Ortho Consumer ity and fecundity were recorded. Analysis of vari- Products, San Ramon, CA) lightly around the ance was used to examine the effects of diet on legs of the log shelves to discourage ant attack. longevity and fecundity (SAS Institute 1988), After adult emergence ceased, we removed the and treatment means were compared using mean bark of each study log to measure the following separation tests (Waller-Duncan t tests) (SAS In- three variables: (1) the number of first instars stitute 1988). No voucher specimens from this reaching the cambium, determined by counting study were retained. larval galleries; (2) the number of larvae reaching the prepupal stage, determined by counting Results and Discussion holes where mature larvae had entered the sap- wood to pupate; and (3) the number of pupae Rearing and Handling Adult Beetles. The fun- reaching adulthood, determined by counting nel trap at the top of the cages was effective in opened emergence holes. These data allowed us capturing adult beetles that emerged from logs. to calculate survivorship of larvae during coloni- Adult P. semipunctata began emerging from the zation, larvae to pupal survivorship, pupal survi- naturally infested logs beginning on 30 March vorship, and total survivorship from first instar to 1990, 5 wk after caging. In total, 975 adults adult. emerged over a 16-wk period (Fig. 1). Although Adult Feeding Experiment. Adult P. semi- there were clear peaks in emergence between punctata have been maintained in the laboratory weeks 11-15, emergence was not tightly syn- with varying results using a variety of diets in- chronized and beetles emerged continually over cluding honey (Duffy 1963), simple sugars or a 16-wk period. These findings support the ob- simple sugars plus leaf extracts (Chararas et al. servation that adults in the field emerge from 1971), and malt extract (Ivory 1977). We have logs for extended periods with no distinct gener- observed the adult beetles feeding on Eucalyp- ations (Chararas 1969a; Loyttyniemi 1983; Men- tus flowers in the field and attempted to develop del 1985). Emergence of adult P. semipunctata a diet for adults that would provide similar nu- from logs containing prepupae can be more trients. The experiment was conducted begin- closely synchronized if the logs are cold-treated ning 18 June 1991 to determine the concentra- (—5 to + 10°C for 30 d) before they are transferred tions of sucrose with or without pollen that to warm (25°C) rearing conditions (Hanks et al. would maximize the reproductive rate. Individ- 1991b). ual mated pairs of newly emerged adult beetles Adult P. semipunctata in the laboratory cages were presented with eight diets: 5, 10, or 30% were quiescent during the day but walked rap- sucrose in distilled water, honeybee pollen idly around the cage from shortly after dusk until (0.1 g; primarily pollen from asteraceous flowers) dawn. Because P. semipunctata males will fight (Clark's Nutrition, Riverside, CA) and distilled among themselves for possession of females, the water, pollen (0.1 g) and 10% sucrose water (dis- sex ratio in the laboratory cages was adjusted to tilled), pollen (0.1 g) and 30% sucrose water, strongly favor females. A ratio of one male to 10% honey water (distilled), and distilled water three females was effective in reducing conflicts alone. The amount of pollen used in the pollen and injuries while maintaining high fecundity. treatments (0.1 g) exceeded the amount that bee- Egg Production. Female P. semipunctata laid tles would consume before replacement. Beetle eggs in the oviposition substrates in batches of pairs (male and female) were caged in cylinders up to 30 eggs. Eggs adhered to the plastic sheets made from aluminum window screen that were and so could be easily moved to glass jars. Eggs 12 cm high and 12 cm in diameter with 14-cm- within a batch hatched synchronously within diameter plastic petri dishes on the top and bot- one day. P. semipunctata eggs could be stored January 1993 HANKS ET AL.: REARING EUCALYPTUS LONGHORNED BORER 99

90i 80" 70- 60-

Beetl e 50- t 40-

< 30- 20-

No . 10" 0 5 6 7 8 9 10 1112 13 14 15 16 17 18 19 20 21 Week Fig. 1. Number of adult P. semipunctata emerging per week from Eucalyptus logs in five cages. for as long as 7 d at 16°C in covered petri dishes The number of pupating beetles emerging as containing a moist Kimwipe (Kimberly-Clark, adults was 7.0 ±2.1 adults per log, or 55.5% Roswell, GA) to maintain high humidity. survivorship. Nonemerging beetles died in pre- Handling Eggs and Rearing Larvae. From a pupal, pupal, and teneral adult stages for un- batch of 200 P. semipunctata eggs, 196 (98%) known reasons. This level of mortality in the successfully hatched. Larvae could be easily col- pupal chambers is similar to that observed in the lected by keeping oviposition substrates contain- field (L.M.H., personal observation). On average, ing eggs in a 1-liter glass jar. Larvae hatched the survivorship of the 20 larvae per log from first from eggs and fell to the bottom of the jars in instar to adulthood was 35%, even in the absence =5 d at 20°C. In the absence of food and under of predators and parasites. laboratory conditions, first instars remained via- With an average fecundity of =180 eggs per ble for only =3 d. However, the longevity of female (see below), we were able to infest nine neonates may be prolonged to 7 d when stored at logs with 20 larvae each using progeny of a sin- 16°C. gle adult female. Given a survivorship of first There is a strong host plant species effect on instars to adulthood of 35%, our culturing tech- the performance of P. semipunctata larvae (Pow- niques allowed us to produce an average of 63 ell 1978, Loyttyniemi 1983). Therefore, it is crit- adult P. semipunctata for every female from the ical that the host log species be considered in previous generation. rearing this cerambycid. We have found E. cam- Adult Feeding Experiment. Longevity among aldulensis to be among the highest quality hosts. beetles provided with only distilled water was Infesting logs with P. semipunctata larvae by shorter than for beetles provided with 10% hand makes it possible to control larval density honey water (F = 3.26; df = 7, 78; P = 0.0043) in logs. By using lower densities, we could max- (Fig. 2A). A diet of only sucrose solutions (5, 10, imize productivity by limiting competition 30%) resulted in survival times that were com- among larvae, thus decreasing development time parable with those of the honey water diet. Bee- and increasing both body size and fecundity. tles provided with only pollen had a mean sur- Larvae required —40 d to complete development vival time not significantly different from those to the prepupal stage under laboratory condi- fed only distilled water, suggesting that they tions. When 20 larvae were introduced by hand were obtaining insufficient nutrition from pollen. into each of nine E. camaldulensis logs, an aver- A combination of pollen and sucrose solution age of 17.1 ± 3.7 (mean ± SEM) succeeded in resulted in longevity similar to that of beetles fed colonizing the logs, for a survivorship of 85.5%. on honey or sucrose solutions. The average number of these larvae reaching These results suggest that the addition of su- maturity (i.e., constructing a pupal cell) was crose greatly improves longevity, but there is no 12.6 ± 4.0, or 73.4% survivorship. advantage in increasing the concentration of su- 100 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 86, no. 1

i i

Distilled Sucrose Sucrose Honey 100^ Water • Pollen <0 1 80: 70

50 40 c 30 o 20 10 0 10 30 10 30 10

250i

200- a a

. 150- I. O ab _k

50 O IL 10 30 10 30 10 % Solution Fig. 2. Longevity (A) and fecundity (B) of P. semipunctata on eight diet treatments. Means ± SEM marked with different letters are significantly different (P < 0.05; Waller-Duncan * tests). January 1993 HANKS ET AL.: REARING EUCALYPTUS LONGHORNED BORER 101 crose above 5%; that the addition of pollen to a California, San Diego); and Ralph Eberhard and the sucrose diet did not improve longevity over a tree crew (University of California, Riverside) for the sucrose solution alone. Microscopic examination provision of Eucalyptus logs. We also thank the Ran- of frass produced by study beetles showed par- cho Santa Fe Association, the Elvenia Slosson Fund for Ornamental Horticulture, the California Department of tially digested pollen, confirming that beetles Forestry and Fire Protection, and the California De- did eat the honeybee pollen. We have also found partment of Food and Agriculture for their financial partially digested Eucalyptus pollen in the frass support. of freshly caught feral adult P. semipunctata, in- dicating that pollen is a natural food source. The results described above indicate that honeybee References Cited pollen is not of significant nutritional value to the Candy, D. J. 1989. Utilization of fuels by the flight beetles in the laboratory, because pollen-fed muscles, pp. 305-319. In G. J. Goldsworthy & C. H. beetles had longevities equal to those main- Wheeler [eds.], flight. CRC, Boca Raton, FL. tained on distilled water alone. However, pollen Chararas, C. 1969a. Biologie et ecologie de Phora- may provide amino acids that are used in flight cantha semipunctata F. (Coleoptere Cerambycidae (Candy 1989). xylophage) ravageur des Eucalyptus en Tunisie, et The 30% sucrose, 10 and 30% sucrose + pol- methodes de protection des peuplements. Ann. len, and 10% honey water treatments all resulted Inst. Nat. Rech. For. Tunie 2: 1-37. in significantly higher fecundity than did the dis- 1969b. Etude biologique de Phoracantha semi- tilled water control (F = 2.55; df = 7, 37; P = punctata F. (Coleoptere Cerambycidae xylophage) specifique des Eucalyptus en Tunisie et recherches 0.03) (Fig. 2B). Although fecundity in the 5% sur la vitalite et l'adaptation de ces essences. C. R. sucrose treatment (116 ± 45.5 eggs per female Seances Acad. Agric. Fr. 55: 47-57. [mean ± SEM]) was >16 times that of the dis- Chararas, C, J. E. Courtois, A. Lefay & A. Thuillier. tilled water control (7.7 ± 7.5 eggs per female) 1971. Biologie, evolution et nutrition de Phora- treatment, this difference was not statistically cantha semipunctata, coleoptere cerambycidae, significant because of the great amount of varia- specifique des Eucalyptus. C. R. Seances Soc. Biol. tion in fecundity within treatments. However, Fil. 165: 1565-1568. when the sucrose data (5, 10, and 30% sucrose) De Viedma, M. G., A. Notario & J. R. Barngano. 1985. were combined into one "sucrose" treatment, Laboratory rearing of Iignicolous Coleoptera (Ce- and the pollen plus sucrose data (10 and 30% rambycidae). J. Econ. Entomol. 78: 1149-1150. Drinkwater, T. W. 1975. The present pest status of sucrose plus pollen) were combined into a single eucalyptus borers Phoracantha spp. in South Africa, "sucrose plus pollen" treatment, there were no pp. 119-129. In H.J.R. Durr, J. H. Giliomee & significant differences between the honey water, S. Neser [eds.], Proceedings, I Congress of the sucrose, or sucrose plus pollen treatments (re- Entomological Society of Southern Africa, 1974. spective means ± SEM, 188.7 ± 46, 123 ± 22, Entomological Society of Southern Africa, Pretoria. and 186 ±31 eggs per female), but all three were Duffy, E.AJ. 1963. A monograph of the immature significantly higher than the distilled water treat- stages of Australasian timber beetles (Ceramby- ment (7.7 ± 7.5 eggs per female; F = 4.66; df = cidae). British Museum (Natural History), London. 4, 40; P = 0.0035). These data strongly indicate Gardiner, L. M. 1970. Rearing wood-boring beetles (Cerambycidae) on artificial diet. Can. Entomol. that fecundity can be improved nearly 20-fold 102: 113-117. over a distilled water diet when sucrose or honey Gonzalez Tirado, L. 1986. Phoracantha semipunc- are provided. As with longevity, fecundity was tata F.: damages caused in the province of Huelva not improved by increasing the concentration of during 1983 and 1984. Bol. San. Veg. Plagas 12: sucrose above 5%. We have found honey water to 147-162. be less practical than sucrose for rearing beetles Hanks, L. M., J. G. Millar & T. D. Paine. 1990. Bi- because honey water ferments after 24 h whereas ology and ecology of the eucalyptus longhorned a sucrose solution remains potable to the beetles borer (Phoracantha semipunctata F.) in southern for as long as 3 d. California, pp. 12-16. In D. Adams & J. Rios [eds.], Proceedings, 39th Meeting of the California Forest Using these methods, we have maintained a Pest Council, 14-15 November 1990. California large colony of P. semipunctata in the laboratory Department of Forestry and Fire Protection, Sacra- for >3 yr. Although the production of adult bee- mento, CA. tles can be somewhat cyclical because of the Hanks, L. M., T. D. Paine & J. G. Millar. 1991a. long emergence period of the larvae, our colony Mechanisms of resistance in Eucalyptus against has yielded as many as 25-50 adult beetles per larvae of the eucalyptus longhorned borer day for a total of > 10,000 beetles in a single year (Coleoptera: Cerambycidae). Environ. Entomol. 20: with a sex ratio of 57% female. 1583-1588. Hanks, L. M., J. G. Millar & T. D. Paine. 1991b. An Acknowledgments evaluation of cold temperatures and density as mor- tality factors of the eucalyptus longhorned borer We thank students and part-time assistants Jun Isoe, (Coleoptera: Cerambycidae) in California. Environ. Kristin Kund, Ta Promlee, and Mi Suk Yu (University Entomol. 20: 1653-1658. of California at Riverside) for their technical assis- Hanks, L. M., J. S. McElfresh, J. G. Millar & T. D. tance; Phil Peters and the tree crew (University of Paine. 1992. Control of the straw itch mite 102 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 86, no. 1

(Acari: Pyemotidae) in an insect rearing facility, gin, biology, damage and control. Phytoparasitica j. Econ. Entomol. 85: 683-686. 15: 131-137. Ivory, M. H. 1977. Preliminary investigations of the Payne, J. A., H. Lowman & R. R. Pate. 1975. Artifi- pests of exotic forest trees in Zambia. Commonw. cial diet for rearing the tilehorned Prionus. Ann. For. Rev. 56: 47-56. Entomol. Soc. Am. 68: 680-682. Linit, M. J. 1985. Continuous laboratory culture of Powell, W. 1978. Colonization of twelve species of Monochamus carolinensis (Coleoptera: Ceramby- Eucalyptus by Phoracantha semipunctata (F.) (Co- cidae) with notes on larval development. Ann. En- leoptera: Cerambycidae) in Malawi. Bull. Entomol. tomol. Soc. Am. 78: 212-213. Res. 68: 621-626. Linsley, E. G. 1959. Ecology of Cerambycidae. SAS Institute. 1988. SAS/STAT user's guide, re- Annu. lease 6.03 ed. SAS Institute, Cary, NC. Rev. Entomol. 4: 99-138. Scriven, G. T., E. L. Reeves & R. F. Luck. 1986. 1962. Cerambycidae of North America. Part I. Intro- Beetle from Australia threatens eucalyptus. Calif. duction. Univ. Calif. Publ. Entomol. 19: 1-75. Agric. 40: 4-6. Loyttyniemi, K. 1983. Flight pattern and voltinism Stanil, V., M. Jankovid-Hladni, J. Ivanovid & V. Nena- of Phoracantha beetles (Coleoptera, Cerambycidae) dovic\ 1989. Joint effects of temperature, food in a semihumid tropical climate in Zambia. Ann. quality, and season on the development of the ce- Entomol. Fenn. 49: 49-53. rambycid Morimus funereus under laboratory con- Mendel, Z. 1985. Seasonal development of the eu- ditions. Entomol. Exp. Appl. 51: 261-267. calypt borer, Phoracantha semipunctata, in Israel. Phytoparasitica 13: 85-93. Received for publication 28 April 1992; accepted 26 1987. Major pests of man-made forests in Israel: ori- August 1992.