Effects of density on the reproductive success of pine engravers: is aggregation in dead trees beneficial?

1. The pine engraver bark beetle Ips pini (Say) (Coleoptera: Scolytidae), aggregates primarily on dead or dying pine trees. In this study pine engravers were laboratory-reared on logs at a range of low densities to determine whether there was a fecundity advantage of breeding aggregations.
2. Mean reproductive success for both males and females declined exponentially with increasing density.
3. Female pine engravers had shorter egg galleries at higher densities, suggesting that they left high-density breeding sites earlier. This would reduce the number of eggs that failed to survive due to larval competition.
4. Some pine engravers colonized the logs voluntarily during the experiment. These volunteers settled independently of the original density.
5. The fungus Ophiostoma sp. was present on the logs and may be competing with the pine engravers for limited bark area.
6. Aggregation resulted in a considerable cost to pine engraver reproductive success even at low densities. Thus, it remains perplexing why pine engravers aggregate actively in nature.     

Death of vigorous trees benefits bark beetles

Bark beetles (Coleoptera: Scolytidae) are commonly associated with live host trees that are stressed, a relationship that has been attributed to lower host defenses or greater nutritional quality of these trees. However, most bark beetle species commonly inhabit freshly dead trees where induced host defenses are absent. In this study, we investigate the role of tree vigor at the time of death for pine engraver bark beetles, Ips pini (Say), breeding in freshly dead jack pine, Pinus banksiana Lamb. As indices of tree vigor, we considered tree size, phloem thickness, and several measures of recent growth rate (last year's growth increment, mean annual increment and basal area increment in the past 5 and 10 years, and periodic growth ratio). We examined the relationship between these indices in three stands, aged 60, 77, and 126 years, and found that phloem thickness, previously shown to have a strong positive effect on bark beetle reproduction, was only weakly associated with tree growth rate and inconsistently related to tree size among the three stands. To examine the effects of tree vigor on pine engraver reproduction, we felled 20 trees of various sizes from the 77-year-old stand, and experimentally established breeding males and females in 25-cm-long sections. Offspring were collected and characteristics of breeding galleries were measured. Using stepwise regression, we consistently found that indices associated with tree growth rate best explained beetle reproductive performance, as they were positively related to parental male and female establishment on logs, female reproductive success, length of egg galleries, proportion of eggs resulting in emerged offspring, and negatively related to the length of the post-egg gallery. Surprisingly, phloem thickness had no unique effect on pine engraver reproduction, except for a weak negative effect on the establishment success of parental females. The strong effect of tree vigor observed in this study suggests that substantial mortality of vigorous trees, such as caused by windthrow, can contribute to significant increases in bark beetle populations that could trigger outbreaks in living trees. Received: 3 February 1999 / Accepted: 27 April 1999     

The evolution of mating systems in bark and ambrosia beetles (Coleoptera: Scolytidae and Platypodidae)

The extraordinary array of mating systems in the Scolytidae and Platypodidae has been largely overlooked by researchers interested in the evolution of sexual behaviour. This paper provides the first overview of reproductive behaviour in this important and widespread group, known to most biologists only by the reputations of tree-killing taxa. Referred to generally as ‘bark beetles’, these insects chew egg tunnels inside a variety of (usually dead) plant tissues, though most species are either phloeophagous (breeding in the inner bark of woody plants) or xylomycetophagous (all stages feeding on mutualistic fungi growing on sapwood or heartwood). In most species, permanent records of many aspects of reproductive behaviour are etched in the host; in many, engravings reveal female fecundity, eggs sired per male, hatching success, and offspring survivorship. Each gallery arm represents a good portion of a given female's lifetime reproduction, but in many species females commonly re-emerge to reproduce in one or two additional sites. In most species of bark beetles, each female initiates her own gallery, to be joined later by a male. These monogynous gallery systems are associated with mating systems defined by how long males stay with females: in a few species, males seldom if ever join females under the bark; in the vast majority of species, males stay for part or all of the oviposition period then leave to seek other mates; and a few groups exhibit permanent monogamy, in that both sexes die in their only gallery system. While these patterns emerge from an overview of the world scolytid fauna, the length of male residency has seldom been quantified, and the costs and benefits associated with male mating strategies have not been measured for any bark beetle. Male-initiated monogyny is uncommon in Scolytidae, though the rule in Platypodidae; all instances of which I am aware are summarized from a phylogenetic perspective. Inbreeding polygyny with highly biased sex ratios has arisen at least seven times in Scolytidae. These taxa are usually characterized by males being dwarfed, flightless, and uncommon. Sex determination is known for only a few examples, but both haplodiploidy and diplodiploidy have been reported. Multiple origins of harem polygyny (otherwise rare in invertebrates) add an exciting dimension to the comparative and experimental study of scolytid mating systems. In harem polygynous taxa, males initiate gallery construction. I summarize what little can be learned from the literature about the fine structure of harem polygynous mating systems in bark beetles, and the problem of measuring reproductive success. Data on the nature of harem polygyny in Pityophthorus lautus are presented, illustrating (a) the fluidity of harems; (b) that average eggs laid per gallery arm is relatively unaffected by harem size, but strongly influenced by resource quality; (c) that male egg-gain is strongly correlated with territory quality (a consequence of (b) above); and (d) the temporal patterning of immigration and emigration and its effects on gallery system sex ratios. The second half of this paper is a discussion of the evolution of bark and ambrosia beetle mating Mating systems, emphasizing sexual selection and the role of resources. Male, residence is interpreted as postcopulalory guarding—preventing sexual liaisons with wandering males. Operational sex ratio, encounter rate, synchrony of breeding, ejaculate competition, and spatiotemporal distribution of resources are discussed as evolutionary forces moulding scolytid and platypodid male postmating behaviour. The nature of male male competition is reviewed. The paucity of information on male behaviour in gallery systems is mentioned; whether or not males significantly aid females is not known. Three hypotheses are presented for why females re-emerge, a feature which strongly affects operational sex ratios. Finally, I summarize features of bark beetle existence predisposing them to the evolution of post-inseminative guarding. Male-initiated monogyny presents a puzzle. I propose that most uncontested examples can be explained by monogyny re-evolving from (male-initiated) harem polygyny, and I present an argument for the evolution of harem polygyny leading to the development of male gallery initiation. The evolution of harem polygyny in birds and mammals has attracted considerable attention. The Verner Willson Orians polygyny threshold model is discussed with respect to bark beetles in general and P. lautus in particular. Resource quality is a major factor in P. lautus harem dynamics: the cost to females of joining harems is apparently slight compared to benefits accrued from moving into sites with higher quality inner bark. Female-biased adult sex ratios have been suggested to lead to harem polygyny, and literature and original data pertinent to this hypothesis are examined. The geometric constraints model, based on the polygyny threshold concept but tailored to bark beetles, is proposed to account for the failure of most species to evolve harem polygyny, and testable predictions are derived that interrelate breeding systems, habitat quality, and progeny size. The evolution of Inbreeding is briefly covered, and two routes to inbreeding polygyny are suggested.     

Density-mediated responses of bark beetles to host allelochemicals: a link between individual behaviour and population dynamics

Abstract  1. Bark beetles (Coleoptera: Scolytidae) accept or reject host conifers based partly on concentrations of phloem monoterpenes. They colonise trees in aggregations, in response to pheromones that attract flying beetles to trees undergoing colonisation. A series of entry and gallery construction assays was conducted to determine whether responses by individual beetles to monoterpenes are altered by pheromones and/or the presence of other beetles.
2. Entry into the amended media by Ips pini and the length of time until entry were not influenced by the presence of aggregation pheromones.
3. Entry into amended media was influenced by the presence of other beetles on the surface of, or constructing galleries in, the substrate. The effects of alpha-pinene and limonene on host entry behaviour were mediated by the density of beetles on the surface of the assay arena, and by the density of beetles constructing galleries within the medium.
4. The percentage of beetles entering medium amended with higher concentrations of monoterpenes increased with increased density of beetles on the surface of the assay arena, until a threshold density of three or four beetles per assay arena, after which entrance rate declined.
5. The presence of other beetles constructing galleries elicited more rapid entry by the test beetles.
6. Gallery lengths were generally higher in the presence of aggregation pheromones.
7. Gallery lengths increased with increased density of beetles within the assay arena.
8. These results suggest a link between the density of bark beetles and responses of individuals. This linkage may partially explain behavioural changes observed during population eruptions.     

Phylogeny of ips DeGeer species (Coleoptera: scolytidae) inferred from mitochondrial cytochrome oxidase I DNA sequence

We used 766 bp of DNA sequence data from the mitochondrial cytochrome oxidase I gene to reconstruct a phylogeny for 39 of 43 Ips species, many of which are economically important bark beetles. The phylogeny was reconstructed using equally weighted and weighted parsimony. In both analyses, peripheral clades were well supported while internal clades were poorly supported. Phylogenetic analysis of translated amino acids produced a poorly resolved tree that was discordant with trees reconstructed with nucleotide sequence data. Two main conclusions are drawn about the monophyly of Ips and traditional systematic groups within Ips. First, Ips is monophyletic only when I. mannsfeldi, I. nobilis, and the concinnus and latidens species groups are excluded. The latidens group, I. mannsfeldi, and I. nobilis form a monophyletic group with 3 Orthotomicus species, while the concinnus group has a more basal position. Second, the majority of the species groups in the current classification for Ips are not monophyletic. European Ips species do not form a monophyletic group, contrary to common usage, and are dispersed on the phylogenetic tree among North American species. These results indicate that a formal systematic revision of Ips is needed.     

Movement patterns and gallery use by the sexually dimorphic Wellington tree weta

The Wellington tree weta, Hemideina crassidens, is a harem-polygynous nocturnal insect whose males defend and mate groups of females residing in cavities in trees. In this study I examined sexual differences in gallery use (number of galleries occupied per unit time), distance travelled per night and activity patterns after sunset. In addition, I investigated how gallery size affected each of these variables. On average, males and females did not differ in the number of galleries visited per night, or in the distance they travelled per night; however, adult males were more likely to be seen earlier in the night than later. Although males and females did not differ in their gallery use or distance travelled, adult males occupied a significantly greater number of galleries and travelled greater distances per night at sites with small galleries than did males at sites with large galleries. These results suggest that habitat structure of a forest patch influences intrasexual tree weta behaviour.     

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.     

Evaluation of funnel traps for characterizing the bark beetle (Coleoptera: Scolytidae) communities in ponderosa pine forests of north-central Arizona

Lindgren funnel traps baited with aggregation pheromones are widely used to monitor and manage populations of economically important bark beetles (Coleoptera: Scolytidae). This study was designed to advance our understanding of how funnel trap catches assess bark beetle communities and relative abundance of individual species. In the second year (2005) of a 3-yr study of the bark beetle community structure in north-central Arizona pine (Pinus spp.) forests, we collected data on stand structure, site conditions, and local bark beetle-induced tree mortality at each trap site. We also collected samples of bark from infested (brood) trees near trap sites to identify and determine the population density of bark beetles that were attacking ponderosa pine, Pinus ponderosa Douglas ex Lawson, in the area surrounding the traps. Multiple regression models indicated that the number of Dendroctonus and Ips beetles captured in 2005 was inversely related to elevation of the trap site, and positively associated with the amount of ponderosa pine in the stand surrounding the site. Traps located closer to brood trees also captured more beetles. The relationship between trap catches and host tree mortality was weak and inconsistent in forest stands surrounding the funnel traps, suggesting that trap catches do not provide a good estimate of local beetle-induced tree mortality. However, pheromone-baited funnel trap data and data from gallery identification in bark samples produced statistically similar relative abundance profiles for the five species of bark beetles that we examined, indicating that funnel trap data provided a good assessment of species presence and relative abundance.     

Mutual mate choice by mountain pine beetles: size‐dependence but not size‐assortative mating

1. Mutual mate choice may be rare, occurring when both sexes invest heavily in reproduction, mating opportunities are abundant, and individuals differ in quality. 2. Mountain pine beetles, Dendroctonus ponderosae (Curculionidae: Scolytinae) appear to meet the conditions for mutual mate choice. We introduced males to females in breeding sites and observed the occurrence and speed of a male entering a female's gallery. We tested for consequences of mutual mate choice, namely condition‐dependent choosiness and assortative mating. 3. Males were more likely to enter a female's gallery when the gallery was in a smaller tree with less resin production and when the gallery was larger. Female body size and condition did not influence the probability of entry. Larger males were less likely to enter a gallery than were smaller males, probably because of size‐dependent choosiness rather than physical limitations. 4. Small males took longer to enter galleries of large females than of small females, whereas large males entered as quickly into galleries of large females as small females. This suggests size‐dependent choosiness by females. 5. No assortative mating by body size was detected, probably because males appeared to choose on the basis of female‐associated resources rather than on female traits.     

Observations on sex ratio and behavior of males in Xyleborinus saxesenii Ratzeburg (Scolytinae, Coleoptera)

Strongly female-biased sex ratios are typical for the fungalfeeding haplodiploid Xyleborini (Scolytinae, Coleoptera), and are a result of inbreeding and local mate competition (LMC). These ambrosia beetles are hardly ever found outside of trees, and thus male frequency and behavior have not been addressed in any empirical studies to date. In fact, for most species the males remain undescribed. Data on sex ratios and male behavior could, however, provide important insights into the Xyleborini's mating system and the evolution of inbreeding and LMC in general.In this study, I used in vitro rearing methods to obtain the first observational data on sex ratio, male production, male and female dispersal, and mating behavior in a xyleborine ambrosia beetle. Females of Xyleborinus saxesenii Ratzeburg produced between 0 and 3 sons per brood, and the absence of males was relatively independent of the number of daughters to be fertilized and the maternal brood sex ratio. Both conformed to a strict LMC strategy with a relatively precise and constant number of males. If males were present they eclosed just before the first females dispersed, and stayed in the gallery until all female offspring had matured. They constantly wandered through the gallery system, presumably in search of unfertilized females, and attempted to mate with larvae, other males, and females of all ages. Copulations, however, only occurred with immature females. From galleries with males, nearly all females dispersed fertilized. Only a few left the natal gallery without being fertilized, and subsequently went on to produce large and solely male broods. If broods were male-less, dispersing females always failed to found new galleries.     

Impacts of silvicultural thinning treatments on beetle trap captures and tree attacks during low bark beetle populations in ponderosa pine forests of northern Arizona

Our research used a combination of passive traps, funnel traps with lures, baited trees, and surveys of long-term thinning plots to assess the impacts of different levels of stand basal area (BA) on bark beetle tree attack and on trap captures of Ips spp., Dendroctonus spp., and their predators. The study occurred at two sites in ponderosa pine, Pinus ponderosa Dougl. ex Laws., forests, from 2004 to 2007 during low bark beetle populations. Residual stand BA ranged from 9.0 to 37.0 m2/ha. More predators and bark beetles were collected in passive traps in stands of lower BA than in stands of higher BA; however, significance varied by species and site, and total number of beetles collected was low. Height of the clear panel passive traps affected trap catches for some species at some sites and years. When pheromone lures were used with funnel traps [Ips pini (Say) lure: lanierone, +03/-97 ipsdienol], we found no significant difference in trap catches among basal area treatments for bark beetles and their predators. Similarly, when trees were baited (Dendroctonus brevicomis LeConte lure: myrcene, exo-brevicomin and frontalin), we found no significant difference for days to first bark beetle attack. Surveys of long-term thinning treatments found evidence of bark beetle attacks only in unthinned plots (approximately 37 m2/ha basal area). We discuss our results in terms of management implications for bark beetle trapping and control.     

Behavior of male and female spruce bark beetles,Ips typographus,on the bark of host trees during mass attack

The behavior of 118 spruce bark beetles, Ips typographus,was observed on trees under colonization. Most individuals were followed from when they landed until they entered or left the tree. Both males and females spent most time inspecting crevices and searching for a place to start boring or for a hole to enter. These behaviors accounted for 87 and 70% of all behavioral acts recorded for males and females, respectively. Females entered galleries with males only after a period of pushing at the gallery entrance. Males spent on average 3 min and females 4 min on the bark before entering or leaving the tree. Thirty-three percent of the beetles eventually entered the tree, 31% flew away, 35% dropped from the host, and one beetle was eaten by a predator. The results are discussed in relation to the question of mate choice in bark beetles and to studies on attack dynamics of spruce bark beetle populations.     

The Relationship Between Resource Control, Association with Females and Male Weapon Size in a Male Dominance Insect

In species with a resource‐defence (male dominance) mating system, males are expected to maximize fitness by controlling resources deemed more valuable by sexually receptive females because these sites attract more mates. Furthermore, males, which control more valuable resources should themselves be of high quality. I experimentally tested these predictions in the laboratory using the sexually dimorphic Wellington tree weta, Hemideina crassidens (Blanchard) (Orthoptera: Tettigonioidea: Anostostomatidae). Male H. crassidens use their mandibular weaponry to fight for control of harems (groups of adult females) that seek shelter in trees cavities (galleries). As predicted, larger galleries housed significantly larger groups of females and males with larger weaponry controlled large galleries significantly more often. Therefore, galleries with a larger volume are likely considered more valuable by males because they house larger harems. However, contrary to prediction, males with larger weaponry did not reside with significantly more females overall because females did not always form the largest possible groups in galleries and males with smaller weaponry were able to reside with single females in small galleries. The latter observation suggests a possible alternative mating strategy by disadvantaged males.     

RESPONSE OF IPS GRANDICOLLIS (EICHHOFF) ADULTS (COLEOPTERA;SCOLYTIDAE) TO SITES IN BARK INITIATED BY OTHERS OF THE SAME SPECIES

b
Dispersing males and females of Ips grandicollis , whether mated or virgin, initiated galleries on experimental logs held in a naturally infested P. radiata plantation. A few females initiated galleries despite the presence of adjacent male-initiated galleries. Males responded to logs bearing females by entering 41% of the female-initiated galleries. The pattern of entry of these males differed depending on whether the entrance to the female-initiated gallery was blocked by frass or not. The findings of this study has led to interesting speculations about the behaviour and biology of this species.     

Inbreeding depression and its effect on intrinsic population dynamics in engraver beetles

Abstract.  1. Phloem-feeding bark beetles (Coleoptera: Curculionidae, Scolytinae) generally disperse before mating, leading to expectations of outbreeding. New York and British Columbia populations of engraver beetles ( Ips pini ) were tested for inbreeding depression using different methods. Among several traits measured, only the number of offspring surviving to adulthood was strongly reduced by inbreeding.
2. There was no evidence of avoidance of inbreeding depression in two possible mechanisms considered: differential male and female emergence times within full sib broods, and early termination of brood construction in forced sib mating.
3. Sib-mated females lay more eggs and have longer galleries than those in outbred crosses, despite a low rate of survival to adulthood for such eggs. This difference may be due to the ability of engraver beetles to assess crowding in broods as larvae begin to feed, and allows partial compensation for the effects of inbreed depression.
4. Population models assuming density-dependent generational effects were modified to account for inbreeding depression. Inbreeding depression makes populations less prone to cyclical behaviour, particularly at lower carrying capacities.
5. Inbreeding depression has not been previously measured in scolytids, nor has inbreeding-related behaviour been explicitly considered outside of exclusively inbreeding tribes.     

Reproductive Consequences of Male Arrival Order in the Bark Beetle, Ips grandicollis

For group-living animals the choice of whether to join aggregations or initiate their own is influenced by potential benefits such as group protection and reduced energetic expenditure, as well as costs such as competition for food and mates. The bark beetle Ips grandicollis is an invasive pest species that colonises recently felled timber in Australian pine (Pinus spp.) plantations. Male beetles initiate colonies by burrowing under the bark of trees and emitting an aggregation pheromone which attracts conspecifics, including a harem of females with whom they mate. We predicted that males that initiated colonies, or who arrived early, would have larger harems than later arrivals (due to decreased competition for females). However, we found the opposite effect with early-arriving males actually associated with fewer females than later arriving males, although this may have resulted from some females leaving harems as they get older. We conclude that pioneering does not improve male likelihood of attracting females in Ips grandicollis, at least initially, but it may provide advantages for offspring when competing for food during development.     

Number of gallery arms of the tree-killing bark beetle Polygraphus proximus (Coleoptera: Curculionidae: Scolytinae)

Polygraphus proximus Blandford (Coleoptera: Curculionidae: Scolytinae) is a non-aggressive monogynous bark beetle that has caused mass mortality in fir (Abies spp.) forests in the last decade. Although gallery structures made by polygynous beetles may influence their reproductive success, the gallery structure and the number of eggs laid by the monogynous tree-killing bark beetle P. proximus has not been investigated in detail in the natural setting of their native range. We, therefore, investigated the length of mother galleries of P. proximus and the number of eggs oviposited by the beetles in relation to gallery systems with different numbers of arms. The number of eggs oviposited in two-armed galleries was significantly greater than that observed in one- and three-armed galleries. Additionally, the length of one- or two-armed galleries with no oviposition was significantly shorter than that in galleries consisting of more than two arms. Our data may suggest that the optimal number of arms in a gallery system (in terms of the number of eggs oviposited) was two.     

Notes on the mating system of Hylesinus varius (F.)(Col., Scolytidae), a putatively bigynous bark beetle

Abstract: Previous research has left confusion about central features of the mating system of the common ash bark beetle Hylesinus varius; in particular, reports conflict as to whether this species is monogynous or bigynous. It was found that females initiated egg tunnel construction; copulation takes place either on the bark in the initial phase of egg tunnel construction, or inside the tunnel system. Observations of one male copulating with several females and of solitary females in galleries with eggs suggested that the males sometimes leave after copulation. In contrast to findings for populations in Slovakia, Norwegian male H. varius were most commonly paired with only one female. Bigynous galleries contained significantly more eggs than monogynous galleries, but bigynous males did not double their reproductive success as found in some other bigynous bark beetles. This suggests that bigynous galleries at first are the work of one female but some males are later joined by a second female.     

Effects of chronic gamma radiation on populations of Ips (Coleoptera: Scolytidae) in trap logs

Abstract. 1. Population dynamics of Ips calligraphus and I. grandicollis (Coleoptera: Scolytidae) were studied in a forest exposed to high doses of gamma radiation.
2. Populations were replicated at eleven dose rates by using 'trap logs'.
3. Adult beetles did not avoid high radiation intensities and attacked all sites equally.
4. Utilization of the cambium-inner bark area varied from c. 2% at a dose rate of 2520 R/20 h day to 90% at dose rates of 63 and 105 R/20 h day.
5. Developmental success of the new brood was directly dependent on the dose rate, and developing progeny were often killed by accumulated exposure.
6. Larger, slower developing species of subcortical beetles were eliminated at dose levels that allowed successful development of the two species of Ips bark beetles.     

Within-harem competition among Ips females, an overlooked component of density-dependent larval mortality

Intraspecific larval competition is commonly believed to be the major component of egg-to-adult mortality in bark beetles. Larval mortality is usually shown to be density dependent. In this study I looked specifically at competition among females in the same gallery system, in a sparse but highly female-biased population of Ips acuminatus breeding in cut pine branches near Kongsberg, southeastern Norway. Supplementary data from a slightly female-biased population in western Norway and a population with intermediate female bias in W. Germany are also provided. Survivorship from egg to adult or pupa was measured in laboratory experiments, for the area in which larvae from different arms potentially competed relative to survivorship in areas away from competition. Survivorship was significantly correlated with both average interarm distance (r²= 0.76) and food per larva (cm² per larva: r²= 0.73). There was no evidence that females could detect the presence of other, nearby egg arms: neither egg spacing (on the arm sides nearest each other) nor interarm distance increased when two arms were near each other. Nor did females distribute themselves optimally within harems, though the distribution of egg arms within harems was significantly different from that expected from random settling. analysis of interarm distances and harem sizes in 1984 branches revealed that a minimum of 34% of arm sides could be expected to suffer considerably reduced larval survivorship due to within-harem competition. Harem size in Ips typographus is reviewed, and data on the distribution of females within harems is presented for Norwegian populations of I. Typographus, Orthotomicus erosus and O. laricis. Evidence from the literature is presented to show that uneven distribution of females between harem sides is not uncommon. I suggest that the findings for Ips acuminatus are applicable to a wide range of harem polygynous scolytids, including I. typographus: the factors influencing density cannot be fully understood without taking into account the distribution of females among males and the distribution of females within harems.