Importance of log size on host selection and reproductive success of Ips pini (Coleoptera: Scolytidae) in ponderosa pine slash of northern Arizona and western Montana

Pine engraver, Ips pini (Say), often use thinning slash, and their populations are known to be influenced by the condition of this material. In our study, we evaluated the importance of three log diameters (5, 10, and 20 cm) and three lengths (60,120, and 240 cm) on various parameters of bark beetle host attack, development, and emergence. Evaluation of slash colonization in northern Arizona (near Flagstaff) and western Montana (near Missoula) over several years, during both spring and summer reproductive periods, indicated that the size of material selected by pine engraver differed by state, year, and season. However, within individual trials and over all trials, a trend in preference for larger diameter logs was apparent, whereas log length was of little effect. When evaluating only attacked logs, results indicated that the apparent preference for larger logs was not followed by significantly greater reproductive performance in the larger log sizes. Log origin was tested as a potential factor for the difference in attack densities between Arizona and Montana trials. Although male beetles in Montana and female beetles in Arizona seemed to distinguish between local-source and foreign-source logs during the attack phase, the gallery development and reproductive success of egg, larval, and new adult stages were not significantly different. Ultimately, differences among the trials both in host size selection and in reproductive success within attacked logs were closely tied to the overall population density of adult beetles. The implications of these findings for resource managers are discussed.     

Seasonal pheromone response by Ips pini in northern Arizona and western Montana, U.S.A.

1 Populations of Ips pini (Say) in northern Arizona and western Montana, U.S.A., were studied to determine regional pheromone response and to evaluate seasonal shifts in that response. A range of enantiomeric blends of the attractant ipsdienol, alone and in the presence of the synergist lanierone, were tested during spring and summer seasons over several years.
2 Both populations were most attracted to high levels of ( R )-(–)-ipsdienol, and lanierone was highly synergistic.
3 A significant seasonal shift in pheromone response between spring and summer seasons was found in both regions in both years. Shifts resulted in a more specific preference for the pheromone treatment of 97% ( R )-(–)-ipsdienol with lanierone.
4 Several coleopteran insect associates of I. pini also displayed responses to the ipsdienol and lanierone treatments. Temnochila chlorodia (Mannerheim) (Trogositidae), Enoclerus sphegeus (F.) (Cleridae) and, to a limited extent, Lasconotus laqueatus (LeConte) (Colydiidae) were attracted to higher proportions of ( R )-(–)-ipsdienol with no apparent reaction to the presence of lanierone. Orthotomicus latidens (LeConte) (Curculionidae: Scolytinae) was strongly attracted to ( S )-(+)-ipsdienol with Enoclerus lecontei (Wolcott) (Cleridae), Pityogenes carinulatus (LeConte) (Curculionidae: Scolytinae) and Hylurgops porosus (LeConte) (Curculionidae: Scolytinae) demonstrating some preferences for the ( S )-(+)-enantiomer. However, lanierone was synergistic for E. lecontei and P. carinulatus , inhibitory for O. latidens , and produced no significant reaction for H. porosus . Elacatis sp. (Salpingidae, previously Othniidae) was attracted to the presence of ipsdienol but displayed no preference to the enantiomeric ratios of ipsdienol or the presence of lanierone.     

The enemy of my enemy is still my enemy: competitors add to predator load of a tree‐killing bark beetle

1 The mountain pine beetle Dendroctonus ponderosae is a major tree‐killing bark beetle in North America. We evaluated how the subsequent arrival of a competing bark beetle Ips pini influences the arrival of predators and their impact on both species. 2 The predators Temnochila chlorodia and Enoclerus sphegeus were strongly attracted to pheromones of D. ponderosae. By contrast, Enoclerus lecontei was mostly attracted to I. pini pheromones. The host compound myrcene synergized attraction of both D. ponderosae and E. sphegeus to the pheromone of D. ponderosae. However, it inhibited attraction of both I. pini and E. lecontei to I. pini’s pheromone. 3 Dendroctonus ponderosae were more attracted to trees than logs treated with its pheromones, whereas I. pini were more attracted to logs than trees treated with its pheromones. Some 78% of T. chlorodia were captured at hosts baited with D. ponderosae pheromones, whereas 83% of E. lecontei were captured at hosts baited with I. pini pheromones. We characterized the sequence of arrival to live trees baited with pheromones of D. ponderosae as: D. ponderosae, T. chlorodia, E. sphegeus, I. pini, E. lecontei. 4 Various combinations of I. pini and predators were added to logs colonized by D. ponderosae in the above sequence of arrival observed in live trees baited with D. ponderosae aggregation pheromones. Ips pini reduced D. ponderosae adult brood production. However, the combination of I. pini and E. lecontei did not raise D. ponderosae brood production above that observed with only I. pini present. Similarly, the combination of I. pini and T. chlorodia did not reduce D. ponderosae brood production below that observed with I. pini alone. By contrast, the combination of I. pini, T. chlorodia and E. lecontei caused more brood loss to D. ponderosae than I. pini alone. 5 Enoclerus lecontei did not reduce brood production by T. chlorodia, whereas T. chlorodia substantially reduced brood production by E. lecontei. 6 Secondary bark beetles that exploit the resource created by primary tree‐killing species exert negative effects through both competition and increased predator load. Implications to the population dynamics, ecology and evolution of tree‐killing bark beetles are discussed.     

Effect of host tree seasonal phenology on substrate suitability for the pine engraver (Coleoptera: Scolytidae): implications for population dynamics and enemy free space

This study evaluated the effects of seasonal phenology on the substrate quality of susceptible hosts to the pine engraver, Ips pini (Say). We also determined the effects of the duration and method of storage on host quality for purposes of laboratory rearing. Live red pine trees were felled at various times during the season, and I. pini adults from a laboratory colony were established on the logs. Subsamples of logs were stored for various intervals, and then provided to beetles. Subsamples of stored logs were waxed at both ends to prevent water loss before being submitted to the same assays. Suitability of red pine phloem tissue in susceptible hosts declined for I. pini throughout the growing season. As the season progressed, the number of beetle progeny that emerged from colonized hosts dropped substantially. This decline was associated with simultaneous reductions in phloem moisture content. Reduction in host suitability may partially offset any advantage I. pini may gain from colonizing trees after the major predators have become less abundant. Bark beetle brood production decreased significantly with length of storage, regardless of the month of tree felling or the method of storing. Implications for bark beetle population dynamics and laboratory rearing systems are discussed.     

Influence of elevation on bark beetle (Coleoptera: Curculionidae, Scolytinae) community structure and flight periodicity in ponderosa pine forests of Arizona

We examined abundance and flight periodicity of five Ips and six Dendroctonus species (Coleoptera: Curculionidae, Scolytinae) among three different elevation bands in ponderosa pine (Pinus ponderosa Douglas ex. Lawson) forests of northcentral Arizona. Bark beetle populations were monitored at 10 sites in each of three elevation bands (low: 1,600-1,736 m; middle: 2,058-2,230 m; high: 2,505-2,651 m) for 3 yr (2004-2006) using pheromone-baited Lindgren funnel traps. Trap contents were collected weekly from March to December. We also studied temperature differences among the elevation bands and what role this may play in beetle flight behavior. Bark beetles, regardless of species, showed no consistent elevational trend in abundance among the three bands. The higher abundances of Ips lecontei Swaine, I. calligraphus ponderosae Swaine, Dendroctonus frontalis Zimmerman, and D. brevicomis LeConte at low and middle elevations offset the greater abundance of I. knausi Swaine, D. adjunctus Blandford, D. approximatus Dietz, and D. valens LeConte at high elevations. I. pini (Say) and I. latidens LeConte were found in similar numbers across the three bands. Flight periodicity of several species varied among elevation bands. In general, the flight period shortened as elevation increased; flight initiated later and terminated earlier in the year. The timing, number, and magnitude of peaks in flight activity also varied among the elevation bands. These results suggest that abundance and flight seasonality of several bark beetles are related to elevation and the associated temperature differences. The implications of these results are discussed in relation to bark beetle management and population dynamics.     

Attraction of southern pine engravers and associated bark beetles (Coleoptera: Scolytidae) to ipsenol, ipsdienol, and lanierone in southeastern United States

We determined the response of the small southern pine engraver, Ips avulsus (Eichhoff); eastern fivespined ips, Ips grandicollis (Eichhoff); sixspined ips, Ips calligraphus (Germar); and pine engraver, Ips pini (Say) to the pheromones (+/-)-ipsenol, (+/-)-ipsdienol, and lanierone in the southeastern United States. Catches of I. avulsus and I. grandicollis to baited multiple-funnel traps were increased by (+/-)-ipsenol and (+/-)-ipsdienol in Florida, Georgia, Louisiana, and North Carolina. In all four localities, the highest numbers of I. avulsus were caught in traps baited with the combination of (+/-)-ipsenol, (+/-)-ipsdienol, and lanierone. In Florida, the highest numbers of I. grandicollis were captured in traps baited with the combination of (+/-)-ipsenol and (+/-)-ipsdienol (with or without lanierone). In the remaining three localities, the largest catches of I. grandicollis occurred in traps baited with (+/-)-ipsenol alone or the combination of (+/-)-ipsenol and (+/-)-ipsdienol (with or without lanierone). (+/-)-Ipsdienol was the only consistent attractant for I. calligraphus and I. pini. Attraction of I. pini in North Carolina to (+/-)-ipsdienol-baited traps was synergized by lanierone but interrupted with (+/-)-ipsenol. The interruptive effect of (+/-)-ipsenol on attraction of I. pini to (+/-)-ipsdienol was negated by lanierone. (+/-)-Ipsdienol was attractive to black turpentine beetle, Dendroctonus terebrans (Olivier), in Florida but not North Carolina, whereas (+/-)-ipsdienol was attractive to I. calligraphus in Louisiana, Georgia, and Florida. Both (+/-)-ipsenol and (+/-)-ipsdienol affected catches of Gnathotrichus materiarus (Fitch) in North Carolina. Trap catches of Hylurgops rugipennis pinifex (Fitch), Hylastes salebrosus Eichhoff, and Hylastes tenuis Eichhoff were unaffected by the pheromone treatments. The combination of (+/-)-ipsenol, (+/-)-ipsdienol, and lanierone may be a cost-effective general lure for I. avulsus, I. grandicollis, and I. pini.     

Effects of sunlight exposure and log size on pine engraver (Coleoptera: Curculionidae) reproduction in ponderosa pine slash in Northern Arizona, U.S.A.

1 Abiotic conditions within logs can affect pine engraver Ips pini (Say) reproductive success, and slash management techniques have been developed that exploit these relationships to reduce brood production. In the present study, we investigated the affect of sunlight exposure on phloem temperature and moisture in logs of two diameters and the subsequent effects on pine engraver reproduction.
2 Logs, 30 cm in length, with diameters of 10 and 15 cm, were cut, left in the field for natural colonization by I pini , and then placed in an open meadow and under shade cloth, providing 27% and 66% shade, until offspring beetles had left the logs. Phloem temperature and moisture were recorded over the duration of the experiment and, at the end of the field experiment, logs were dissected and galleries were measured to gauge beetle reproductive success.
3 As sunlight exposure increased, phloem temperatures increased and potentially lethal temperatures were often reached in the high-sunlight exposure but seldom in the low-sunlight. Smaller diameter logs had drier phloem than larger diameter logs. All logs dried with time but sunlight level did not affect desiccation rates. Ips pini preferred attacking larger logs and the bottom side of logs. Sunlight exposure had a significant effect on net reproductive success in smaller diameter logs, with very little net reproductive success in high-sunlight exposed logs, and the highest reproductive success was found in small diameter logs in the low-sunlight treatments.
4 Management implications of these results are discussed.     

Season-specific directional movement in migratory Australian butterflies

Large numbers of adults of certain species of butterfly flying in an apparently 'purposeful' manner are often noted by entomologists and the general public. Occasionally, these are recorded in the literature. Using these records we summarise information regarding the direction of movement in Australian butterflies and test whether there are consistent patterns that could account for known seasonal shifts in geographical range. The data were analysed using contingency tables and directionality statistics. Vanessa itea, Vanessa kershawi, Danaus plexippus , Danaus chrysippus and Badamia exclamationis flew predominately south in the spring–summer and north in the autumn–winter. Tirumala hamata has a strong southern component to its flight in spring but, as in Euploea core, appears non-directional in the autumn. For many supposedly known migratory species, the number of literature records are few, particularly in one season (mainly autumn). Thus, for Appias paulina , four of seven records were south in the spring–summer, as were six of nine records for Catopsilia pomona, and three of five for Zizina labradus. For Belenois java , flight records were only available for the spring and these showed geographical differences; predominantly north-west in northern Australia (Queensland) and south-west in southern Australia (Victoria, New South Wales). There were too few records for Papilio demoleus in the literature (four only) to draw any conclusions. Major exceptions to the seasonal trend of south in the spring and north in the autumn were Junonia villida , which showed a predominant north-westward direction in both seasons, and Eurema smilax, with a predominant southern or western flight in both seasons. We discuss these species specific trends in migration direction in relation to seasonal shifts in suitable habitat conditions, possible cues used in orientation and in timing changes in direction.     

Interactions among intraspecific competition, emergence patterns, and host selection behaviour in Ips pini (Coleoptera: Scolytinae)

Abstract.  1. Variations in developmental conditions of bark beetles, particularly intraspecific competition, can induce morphological and physiological modifications in the offspring. It is hypothesised that intraspecific competition could also affect host selection behaviour. Such behavioural changes might be manifested in response to host (alpha-pinene) or beetle (verbenone) compounds.
2.  Ips pini were bred at different densities. The offspring were measured for size, weight, and lipid concentration, and then subjected to tunnelling bioassays in agar media amended with varying amounts of alpha-pinene or verbenone.
3. High parental colonisation densities reduced emergence time of parents and offspring. Increasing colonisation density and emergence time had a negative influence on offspring vigour, resulting in a reduction of the distance tunnelled by the beetles.
4. Both alpha-pinene and verbenone were repellent. Surprisingly, verbenone was also toxic at high concentrations, with its effect being greater on beetles arising from high densities. The repellent effect of these compounds did not vary according to colonisation density, but for both compounds, at a 1.5 mg g^–1 concentration, it increased with emergence time.
5. The consequences of varying vigour and behaviour in relation to developmental conditions and emergence patterns on population dynamics of I. pini are discussed.     

Functional expression of a bark beetle cytochrome P450 that hydroxylates myrcene to ipsdienol

The final steps in the pheromone-biosynthetic pathway of the pine engraver beetle, Ips pini (Say) (Coleoptera: Scolytidae) are unknown, but likely involve myrcene (7-methyl-3-methylene-1,6-octadiene) hydroxylation to produce the aggregation pheromone component, ipsdienol (2-methyl-6-methylene-2,7-octadien-4-ol). We have isolated a full-length I. pini cDNA encoding a cytochrome P450, CYP9T2. The recovered cDNA is 1.83kb and the open reading frame encodes a 532 amino acid protein. CYP9T2 is regulated by the same physiological factors that induce pheromone production. Quantitative real-time PCR experiments showed that feeding on host phloem induced CYP9T2 expression in males, but not females, and that basal expression levels are highest in male midguts, similar to other I. pini pheromone-biosynthetic genes. Microsomes prepared from Sf9 cells co-expressing baculoviral-mediated recombinant CYP9T2 and housefly (Musca domestica) NADPH-cytochrome P450 reductase converted myrcene to ipsdienol. The product identified by coupled GC-MS was mostly (4R)-(-)-ipsdienol, an important aggregation pheromone component for western North American I. pini. These results are consistent with CYP9T2 encoding a myrcene hydroxylase that functions near the end of the pheromone-biosynthetic pathway.     

进口美国松木中截获齿小蠹的鉴定(鞘翅目:小蠹科)

本文对2010—2015年全国口岸从美国进境松木中截获的齿小蠹进行了分析,分别对美云杉齿小蠹Ips borealis、美雕齿小蠹I.calligraphus、混点齿小蠹I.confusus、重齿小蠹I.duplicatus、南部松齿小蠹I.grandicollis、蒙大拿齿小蠹I.montanus、似混齿小蠹I.paraconfusus、白云杉齿小蠹I.perturbatus、美松齿小蠹I.pini、平额重齿小蠹I.plastographus、十二齿小蠹I.sexdentatus、云杉八齿小蠹I.typographus 12种昆虫的进境寄主进行统计,对截获频次进行分析对其成虫的主要形态特征进行了描述,编制了分种检索表,对检疫鉴定工作具有指导意义。     

The role of grasslands in the yearly life-cycle of Rhopalosiphum padi (Homoptera: Aphididae) in Sweden.

The population density of the bird cherry-oat aphid (Rhopalosiphum padi) was monitored in spring sown barley and in grasslands (leys and pastures) and a suction trap was used to monitor the flight periodicity of the aphids. Emigrants from the primary host (bird cherry) colonised both grass and cereals in spring and a migration from cereals to grasses took place in mid-summer. There was a negative correlation between the sizes of summer and autumn migration. There was a positive relationship between late summer growth in leys and the size of autumn migrations. It is concluded that the size of the autumn migration is mainly dependent on aphid population growth in grasslands during late summer and autumn.     

Seasonality of recruitment and reproduction of Acanthocephalus tumescens (Acanthocephala) in fishes from Lake Moreno (Patagonia, Argentina)

The seasonality of recruitment and reproduction of Acanthocephalus tumescens at the component population level was investigated in Lake Moreno (Patagonia, Argentina). Seasonal samples of the principal fish host species were taken between spring 1999 and spring 2000. Prevalence, mean intensity, coefficient of dispersion, sex ratio, proportion of maturity stages of females, and percentages of the 2 sources of infection in the stomach of fishes were calculated. Overall prevalence, mean intensity, and coefficient of dispersion showed the same pattern of seasonal changes. The seasonal feeding patterns of fishes affect the occurrence of A. tumescens, producing 1 peak in spring and the other peak in autumn. The lower temperature in winter delay reproductive processes after the autumn period of recruitment.     

Test of nonhost angiosperm volatiles and verbenone to protect trap trees for Sirex noctilio (Hymenoptera: Siricidae) from attacks by bark beetles (Coleoptera: Scolytidae) in the northeastern United States

Sirex noctilio F. (Hymenoptera: Siricidae) is an invasive woodwasp, currently established in northeastern North America. In other regions of the world, stressed trap trees are used to monitor populations of S. noctilio and to provide inoculation points for the biological control nematode Deladenus siricidicola Bedding. However, the operational use of trap trees for S. noctilio in North America may be compromised by the large community of native organisms that inhabit stressed and dying pine trees. Common bark beetles such as Ips pini (Say) and Ips grandicollis (Eichhoff) (Coleoptera: Scolytidae) could potentially compete with S. noctilio and associates for resources on trap trees, possibly reducing the efficacy of trap trees as habitats for the woodwasp. In an attempt to develop a technology to mitigate this potential issue, three common semiochemical interruptants--conophthorin, green leaf volatile mix, and verbenone--were tested for effectiveness in reducing arrivals of I. pini and I. grandicollis on trap trees treated with herbicides in northeastern United States. In addition, the effects of these compounds were determined independently with pheromone-baited multiple-funnel traps. None of the interruptants reduced numbers of I. pini or I. grandicollis either arriving on trap trees or caught in pheromone-baited traps. However, verbenone increased catches of I. grandicollis in traps baited with its pheromone, ipsenol. The mix of green leaf volatiles reduced catches of a native ambrosia beetle, Gnathotrichus materiarius (Fitch), whereas verbenone reduced trap catches of an exotic ambrosia beetle, Xylosandrus germanus (Blandford). Catches of X. germanus in traps adjacent to trap trees were enhanced with conophthorin.     

Seasonal changes in the flight apparatus of winged females and sexual males of the aphid <Emphasis Type="Italic">Tuberculatus quercicola</Emphasis> (Hemiptera: Aphididae)

Tuberculatus quercicola (Matsumura) feeds on Quercus dentata Thunberg, and has mutualistic interactions with ants. Tuberculatus quercicola has two winged morphs in its life cycle, winged females appear in summer and sexual males appear in autumn. Previous studies have shown that wing loading (ratio of body volume to wing area) is higher for the winged females, because of ant attendance, resulting in extremely low dispersal. It is known that the nutritional quality of host plants is high in spring and autumn, when leaves are growing or senescent, and low in summer when leaves are mature. This study examined the effects of seasonal plant deterioration on the development of flight apparatus (wing size and flight muscle) of winged females and males. Moreover, field intercept traps were used to examine the extent of dispersal of males. The results showed that seasonal plant deterioration affected development of the flight apparatus of winged females, particularly flight muscle. Flight muscle development was significantly higher in winged males in autumn than in winged females. However, winged males were not caught in any of the traps. The different resource allocation to the flight apparatus of winged females and males is discussed.     

Parasitoids and dipteran predators exploit volatiles from microbial symbionts to locate bark beetles

Host location by parasitoids and dipteran predators of bark beetles is poorly understood. Unlike coleopteran predators that locate prey by orienting to prey pheromones, wasps and flies often attack life stages not present until after pheromone production ceases. Bark beetles have important microbial symbionts, which could provide sources of cues. We tested host trees, trees colonized by beetles and symbionts, and trees colonized by symbionts alone for attractiveness to hymenopteran parasitoids and dipteran predators. Field studies were conducted with Ips pini in Montana. Three pteromalid wasps were predominant. All were associated with the second and third instars of I. pini. Heydenia unica was more attracted to logs colonized by either I. pini or the fungus Ophiostoma ips than logs alone or blank controls (screen with no log). Rhopalicus pulchripennis was more attracted to logs colonized by I. pini than logs alone or blank controls. Dibrachys cavus was attracted to logs but did not distinguish whether or not they were colonized. Two dolichopodid predators were predominant. A Medetera species was more attracted to colonized than uncolonized logs and more attracted to logs than blank controls. It was also more attracted to logs colonized with the yeast Pichia scolyti than uncolonized logs, but attraction was less consistent. An unidentified dolichopodid was more attracted to logs colonized with I. pini, O. ips, and the bacteria Burkholderia sp., than to uncolonized logs. It was also attracted to uncolonized logs. Its responses were less consistent and pronounced than H. unica. These results suggest some parasitoids and dipteran predators exploit microbial symbionts of bark beetles to locate hosts. Overall, specialists showed strong attraction to fungal cues, whereas generalists were more attracted by plant volatiles. These results also show how microbial symbionts can have conflicting effects on host fitness.     

Flight speeds and climb rates of Brent Geese: mass-dependent differences between spring and autumn migration

Aerodynamic theories of bird flight predict that horizontal flight speed will increase with increasing load whereas vertical flight speed will decrease. Horizontal flight speed for birds minimizing overall time on migration is predicted to be higher than flight speed for birds minimizing energy expenditure. In this study we compare flight speeds of Brent Geese Branta b. bernicla recorded by tracking radar and optical range finder during spring and autumn migration in southernmost Sweden, testing the above-mentioned predictions. Geese passing Sweden in spring are substantially heavier than in autumn and there might also be a stronger element of time-selection in spring than in autumn. Recorded airspeeds were significantly higher in spring (mean 19.0 m s⁻¹) than in autumn (mean 17.3 m s⁻¹), the average difference being slightly larger than predicted due to the mass difference alone. The effects on airspeed of wind, vertical speed, flock size and altitude were also analysed, but none of these factors could explain the seasonal difference in airspeed. Hence, the results support the hypothesis of mass-dependent flight speed adjustment. The difference between the two seasons was not large enough to corroborate the hypothesis of a stronger element of time-selection in spring, but this hypothesis cannot be rejected. Vertical flight speeds were lower in spring than in autumn, supporting a negative effect of load on birds' flight power margin.     

RAPD‐PCR DETECTION OF DNA POLYMORPHISM OF COTTON APHID*

Abstract The RAPDPCR technique was used to determine the DNA polymorphism of the aphid Aphis gossypii(Glover) collected from different host plants and in different seasons. Three primers were selected from 20 primers and used for cluster analysis based on the data of Nei's genetic distance (D). The results showed that the aphids on Chinese prickly ash differentiated obviously from the aphids on the other four host plants at DNA level. The seasonal population of cotton aphid might be clustered into three groups, i.e. the spring and autumn yellow colored aphids, the spring and autumn green colored aphids and the yellow dwarf form aphid in summer. However, the genetic relationship of dwarf form was more closely to the spring and autumn green colored aphid.     

Specialization of midgut cells for synthesis of male isoprenoid pheromone components in two scolytid beetles,Dendroctonus jeffreyi and Ips pini

Endodermal or midgut cells have only recently been recognized as the site of pheromone synthesis in bark beetles. Midgut cells are not only specialized for digestion, but they have also been recruited to form isoprenoid compounds that function as pheromone components in Ips pini and Dendroctonus jeffreyi. Male bark beetle midgut cells are competent to produce isoprenoid pheromones after feeding or stimulation by juvenile hormone (JH) III. Competent midgut cells share many ultrastructural features with cells that do not secrete isoprenoid pheromone, but they are distinguished from these by abundant and highly ordered arrays of smooth endoplasmic reticula. During secretion, both midgut cells that produce pheromone and cells that do not are characterized by the presence of apical extrusions (apocrine secretion) rather than the presence of vesicles that fuse with the apical membrane and undergo exocytosis (eccrine secretion). Pheromone-producing cells of the midgut do not represent a population of cells that are distinct from cells involved in digestion. All, or most, midgut cells of male I. pini and D. jeffreyi can secrete pheromones as well as digestive enzymes.     

Lack of genetic differentiation in aggressive and secondary bark beetles (Coleoptera: Curculionidae, Scolytinae) from Arizona

Bark beetles (Coleoptera: Curculionidae, Scolytinae) play an important role as disturbance agents in ponderosa pine (Pinus ponderosa Douglas ex Lawson) forests of Arizona. However, from 2001 to 2003, elevated bark beetle activity caused unprecedented levels of ponderosa pine mortality. A better understanding of the population structure of these species will facilitate analysis of their dispersal patterns and improve management strategies. Here, we use fluorescently labeled amplified fragment length polymorphism (fAFLP) analysis to resolve genetic variation among and within sampling locations in northcentral Arizona of Ips pini (Say), Dendroctonus brevicomis LeConte, and D. frontalis Zimmermann. We generated genetic fingerprints for >500 beetle specimens and analyzed genetic diversity. For all species, gene flow estimates among sampling locations were high, and significant population subdivision was not discernible across a large portion of ponderosa pine forests in Arizona. However, a weak relationship was detected with I. pini population structure and elevation. Because of the lack of genetic differentiation detected throughout the large study area, our findings suggest these insects are capable of long distance dispersal and exhibit a high degree of gene flow across a broad region. We conclude that our results are consistent with strong dispersal patterns and large population sizes of all three species.