The importance of Olneya tesota as a nurse plant in the Sonoran Desert

Abstract. The function of Olneya tesota (ironwood) as a nurse plant and habitat modifier species in the Sonoran Desert was evaluated at five study sites (using 75 250-m² sample plots) from Bahia Kino, Sonora to Organ Pipe Cactus National Monument, Arizona. Beneath the canopy of O. tesota trees 75 perennial plant species were found. A principal component ordination of the plots created three groups: southern, perturbed and protected sites. A strong triple association of columnar cacti, big shrubs with berry type fruits and O. tesota was detected. The relationships of Lophocereus schottii and Peniocereus striatus with O. tesota were studied in more detail. Significant differences in soil surface temperatures and stem temperatures of P. striatus were detected beneath the shade of O. tesota trees compared with soils and plants in open spaces. Olneya tesota must be considered as a habitat modifier species with ecological and conservational importance to the plant communities in the Sonoran Desert.     

GENETIC DIVERSITY AND CLONAL STRUCTURE IN A COLUMNAR CACTUS,LOPHOCEREUS SCHOTTII

In southern Arizona the columnar cactus, Lophocereus schottii, inhabits desert riparian environments. Reproduction in this part of its range is predominantly asexual and occurs by either the dispersal of stems in the immediate vicinity of parents or the long-distance transport of detached stem pieces downstream by floodwaters. Genetic diversity and clonal structure of eight populations of L. schottii in Organ Pipe Cactus National Monument, Arizona, were examined. In all populations ramets were mapped and stem tissue from each ramet was examined electrophoretically. At the species level, 44.4% of the loci were polymorphic and the genetic diversity was 0.145. Within populations, the mean proportion of polymorphic loci and genetic diversity were 34.4% and 0.126%, respectively. Although most of the allozyme variation was within populations, appreciable heterogeneity was found among populations (G_ST = 0.130). Genetic and genotypic diversity was greatest in three of the four populations located in the principal area of L. schottii occurrence in the monument. Genotypic diversity was lowest in the smallest population and in the most isolated population. Ramets in all populations were spatially aggregated. Plant pairs with identical multilocus genotypes were usually ≤ 10 m apart, but some widely separated individuals had identical genotypes. Occasional long-distance dispersal of stems and the periodic recruitment of seedlings have caused genets to intermingle, promoting outcrossing and maintaining genetic diversity.     

BRANCHING PATTERNS OF COLUMNAR CACTI: INFLUENCES ON PAR INTERCEPTION AND CO2 UPTAKE

Field measurements and a computer model were used to determine how stem shape and arrangement of stems in space affect interception of photosynthetically active radiation (PAR) and CO₂ uptake under otherwise optimal conditions for four species of columnar cacti (Carnegiea gigantea, Lophocereus schottii, Pachycereus pringlei, and Stenocereus thurberi). In simulations where the number of widely spaced stems was increased from 1 to 19 but plant volume remained constant, surface area and PAR interception increased, leading to 3-fold increases in whole-plant CO₂ uptake. Increasing the distance between stems from 0 cm to infinity decreased self-shading and increased predicted CO₂ uptake 4-fold. Stem length, diam, ribbing characteristics, and spine coverage also influenced PAR interception. The model indicated that the observed higher frequency of branches on the south side of the trunk of C. gigantea had only a slight, though positive, effect on CO₂ uptake for single-branched plants. Because of its greater surface area (A), a five-stemmed plant of C. gigantea typical for a field site near Tucson, Arizona was predicted to have 52% more CO₂ uptake than a single-stemmed plant of the same volume (V). Although large A/V decreases water storage per unit transpiring area, whole-plant CO₂ uptake can be increased when A/V is increased by branching for these constant-volume plants. However, the stems must be arranged to avoid excessive self-shading and thus keep the area below PAR compensation small.     

Analysis of the community structure of yeasts associated with the decaying stems of cactus. III.Stenocereus thurberi

Yeast communities in necroses of organpipe cactus (Stenocereus thurberi) were surveyed at 3 localities in Arizona. Quantitative analysis of random samples allows comparisons of the types and numbers of yeasts at 3 levels: within plants, between plants within a locality, and between localities. The analysis shows that the major source of variability is between plants. This pattern is identical with the pattern shown by agria cactus (Stenocereus gummosus) and is thought to be due to sampling different successional stages. No significant differences in estimates of the effective number of yeast species (ENS) in agria and organpipe samples were found. Comparisons of agria, organpipe, and prickly pear (Opuntia) cacti support the hypothesis that cactus chemistry is an important determinant of the yeast community structure which, in turn, is an important determinant of the diversity ofDrosophila species which utilize necrotic cacti as feeding and breeding substrates.     

Yeast communities from host plants and associated Drosophila in southern arizona: new isolations and analysis of the relative importance of hosts and vectors on comunity composition

Summary The yeast communities from slime fluxes of three deciduous trees (Prosopis juliflora, Populus fremontii and Quercus emoryi) and the necroses of two cacti (Opuntia phaeacantha and Carnegiea gigantea) were surveyed in the region of Tucson, Arizona. In addition, the yeasts carried by dipterans associated with the fluxes or necroses (Drosophila carbonaria, D. brooksae, D. nigrospiracula, D. mettleri, and Aulacigaster leucopeza) were sampled. The results indicate that each host sampled had a distinct community of yeasts associated with it. The dipterans, which can act as vectors of the yeasts, deposited yeasts from other sources in addition to those found on their associated hosts. It is argued that host plant physiology is relatively more important than the activity of the vector in determining yeast community composition. Furthermore, the average number of yeast species per flux or necrosis is not different from the average number of yeast species per fly. It is hypothesized that the vector may affect the number of species per individual flux or not, and that the number is lower than the rot or necrosis could potentially support.     

Acetylene reduction and indoleacetic acid production by Azospirillum isolates from Cactaceous plants

Azospirillum isolates were obtained from rhizosphere soil and roots of three cactaceae species growing under arid conditions. All Azospirillum isolates from rhizosphere and roots ofStenocereus pruinosus andStenocereus stellatus were identified asA. brasilense; isolates of surface-sterilized roots fromOpuntia ficus-indica were bothA. brasilense andA. lipoferum. Azospirilla per g of fresh root in the three species ranged from 70×10³ to 11×10³. The most active strains in terms of C₂H₂ reduction (25–49.6 nmol/h·ml) and indoleacetic acid (IAA) production (36.5–77 μg/ml) were those identified asA. brasilense and isolated from Stenocereus roots.A. lipoferum isolated from Opuntia roots produced low amounts of IAA (6.5–17.5 μg/ml) and low C₂H₂-reduction activity (17.8–21.2 nmol/h·ml).     

Morphological consequences of the trade-off between growth and reproduction in a columnar cactus (Lophocereus schottii)

Since cacti have very small surface/volume ratios, the amount of light they intercept is limited, and so is photosynthesis. However, many cacti shade themselves by means of dense hairy mats, called cephalia, which protect the floral buds. There is a trade-off between allocating a surface to photosynthesis or to the protection of reproductive structures. We found that proportion of the stem covered by cephalium affected fitness in the columnar cactus Lophocereus schottii. Too little cephalium holds few floral buds while too much reduces fecundity – probably through diminished photosynthesis – with an intermediate amount maximising reproduction. The optimal proportion of cephalium becomes smaller as the plant grows, probably because self-shading by new branches causes a further reduction in light interception. Smaller plants had significantly less cephalium than the optimal amount to maximise current reproduction. Since the largest individuals produced more reproductive structures, small cacti may find it more profitable to grow fast to achieve a large fecundity, than to assign resources to immediate reproduction via cephalium development.     

Alterations in growth and crassulacean acid metabolism (CAM) activity of in vitro cultured cactus

Unlike C-3 plants, cacti possess a crassulacean acid metabolism (CAM) physiology that can alter the pattern of carbon uptake and affect plant growth under artificial environmental conditions, especially in tissue culture. In vitro-derived plantlets of Coryphantha minima grew 7-fold larger than plants cultured under similar ex vitro conditions. Growth regulators incorporated into the culture media during shoot proliferation stage of micropropagation had a strong influence on this increased growth. Other important factors that contributed to increased growth under in vitro conditions were high relative humidity and sugar in the culture medium. An analysis of gas exchange and daily fluctuations of malic acid levels revealed an increase in net photosynthetic rate, in terms of carbon assimilation, by in vitro plants compared with that of ex vitro plants. This stimulated photosynthesis in the presence of an external carbon source was unexpected but apparently true for cacti exhibiting CAM physiology. Unlike CAM plants grown in ex vitro conditions, net CO₂ uptake by in vitro-cultured cacti occurred continuously in the light as well as the dark. Once regenerated, cacti were transferred to ex vitro conditions where the normal CAM pathway resumed with a concomitant reduction in growth and CO₂ uptake. These results showed that growth of cacti can be considerably accelerated by in vitro culture. This revised version was published online in June 2006 with corrections to the Cover Date.     

Developmental Time and Thorax Length Differences Between the Cactophilic Species Drosophila Buzzatii and D. Koepferae Reared in Different Natural Hosts

Drosophila buzzatii and Drosophila koepferae are two cactophilic sibling species whose ranges partially overlap in Northwestern and Western Argentina. Both species can utilize the decaying tissues of both Opuntia and columnar cacti as breeding sites. Though D. buzzatii and D. koepferae are not differentially attracted to Opuntia and columnar hosts, the composition of the communities of flies emerging from natural substrates of both cacti differed significantly in a natural population. The objective of this paper is to analyze whether intra and/or interspecific competition affects development time and thorax length in D. buzzatii and D. koepferae when both species are reared in single and mixed species culture and fed with semi-natural media prepared with fermenting materials of Opuntia sulphurea(tuna) and Trichocereus terschekii(cardón). Our results showed that both traits differ significantly between flies raised in different hosts and that differences between D. koepferae and D. buzzatii species for both thorax length and development time depend on the type of culture (mixed vs. single species). In addition, the host by type of culture interaction was significant. We also observed thorax length differences between Drosophila species and type of culture. Our present data suggest that the effect of intra and interspecific competition varied between the two traits investigated and between species. However, competition alone cannot explain the differential pattern of resource utilization shown by D. buzzatii and D. koepferae in the natural population studied.     

Soil O2 and CO2 effects on root respiration of cacti

Through use of a recently developed technique that can measure CO₂ exchange by individual attached roots, the influences of soil O₂ and CO₂ concentrations on root respiration were determined for two species of shallow-rooted cacti that typically occur in porous, well-drained soils. Although soil O₂ concentrations in the rooting zone in the field were indistinguishable from that in the ambient air (21% by volume), the CO₂ concentrations 10 cm below the soil surface averaged 540 μLL⁻¹ for the barrel cactusFerocactus acanthodes under dry conditions and 2400 μLL⁻¹ under wet conditions in a loamy sand. For the widely cultivated platyopuntiaOpuntia ficus-indica in a sandy clay loam, the CO₂ concentration at 10 cm averaged 1080 μLL⁻¹ under dry conditions and 4170 μLL⁻¹ under wet conditions. For both species, the respiration rate in the laboratory was zero at 0% O₂ and increased to its maximum value at 5% O₂ for rain roots (roots induced by watering) and 16% O₂ for established roots. Established roots ofO. ficus-indica were slightly more tolerant of elevated CO₂ than were those ofF. acanthodes, 5000 μLL⁻¹ inhibiting respiration by 35% and 46%, respectively. For both species, root respiration was reduced to zero at 20,000 μLL⁻¹ (2%) CO₂. In contrast to the reversible effects of 0% O₂, inhibition by 2% CO₂ was irreversible and led to the death of cortical cells in established roots in 6 h. Although the restriction of various cacti and other CAM plants to porous soils has generally been attributed to their requirement for high O₂ concentrations, the present results indicate that susceptibility of root respiration to elevated soil CO₂ concentrations may be more important.     

Molecular evolution and population genetics of two Drosophila mettleri cytochrome P450 genes involved in host plant utilization

Understanding the genetic basis of adaptation is one of the primary goals of evolutionary biology. The evolution of xenobiotic resistance in insects has proven to be an especially suitable arena for studying the genetics of adaptation, and resistant phenotypes are known to result from both coding and regulatory changes. In this study, we examine the evolutionary history and population genetics of two Drosophila mettleri cytochrome P450 genes that are putatively involved in the detoxification of alkaloids present in two of its cactus hosts: saguaro (Carnegiea gigantea) and senita (Lophocereus schottii). Previous studies demonstrated that Cyp28A1 was highly up-regulated following exposure to rotting senita tissue while Cyp4D10 was highly up-regulated following exposure to rotting saguaro tissue. Here, we show that a subset of sites in Cyp28A1 experienced adaptive evolution specifically in the D. mettleri lineage. Moreover, neutrality tests in several populations were also consistent with a history of selection on Cyp28A1. In contrast, we did not find evidence for positive selection on Cyp4D10, although this certainly does not preclude its involvement in host plant use. A surprising result that emerged from our population genetic analyses was the presence of significant genetic differentiation between flies collected from different host plant species (saguaro and senita) at Organ Pipe National Monument, Arizona, USA. This preliminary evidence suggests that D. mettleri may have evolved into distinctive host races that specialize on different hosts, a possibility that warrants further investigation.     

GENETICS OF HOST-CACTUS RESPONSE AND LIFE-HISTORY EVOLUTION AMONG ANCESTRAL AND DERIVED POPULATIONS OF CACTOPHILIC DROSOPHILA MOJAVENSIS

The extent of host-specific genetic variation for two life-history traits, egg to adult developmental time and viability, and one morphological trait closely tied to fitness, adult thorax size, was exposed by employing a nested half-sib/full-sib breeding design with Baja and mainland populations of Drosophila mojavensis recently extracted from nature. This study was motivated by the presence of substantial variation in life histories among populations of D. mojavensis that use the fermenting tissues of particular species of columnar cacti for feeding and breeding in the Sonoran Desert. Full-sib progeny from all sire-dam crosses were split into cultures of agria cactus, Stenocereus gummosus, and organ pipe cactus, S. thurberi, to examine patterns of genotype-by-environment interaction for these fitness components. Baja flies expressed shorter egg-to-adult developmental times, higher viabilities, and smaller body sizes than mainland flies consistent with previous studies. Significant sire and dam components of variance were exposed for developmental time and thorax size. Genotype-by-environment interactions were significant at the level of dams for developmental time and nearly significant for viability (P = 0.09). Narrow- and broad-sense heritabilities were influenced by host cactus, sex, and population. No strong pattern of genetic correlation emerged among fitness components suggesting that host-range expansion has not been accompanied by formation of coadapted life histories, yet the ability to estimate genetic correlations and their standard errors was compromised by the unbalanced nature of the data set. Genetic correlations in performance across cacti were slightly positive, evidence for ecological generalism among populations explaining the observed pattern of multiple host cactus use within the species range of D. mojavensis.     

Prey refugia and the distributions of two Sonoran Desert cacti

Summary Two small Sonoran Desert cacti, Mammillaria microcarpa and Echinocereus englemannii, are commonly found beneath canopies of the larger, tree-like cactus Opuntia fulgida. The mechanism leading to this distribution pattern is incidental to the mode of reproduction in O. fulgida. Opuntia fulgida propagates by means of easily-detached, spine-covered stem joints that accumulate beneath the parent plant. These accumulations of spines apparently deter mammalian herbivores that otherwise consume succulent tissues of the smaller cacti. Such incidental effects are little studied, but they may contribute substantially to structure within plant communities.     

Identification of self-incompatibility groups in<Emphasis Type="Italic"> Hatiora</Emphasis> and<Emphasis Type="Italic"> Schlumbergera</Emphasis> (Cactaceae)

The Cactaceae, a family of about 1,800 species of succulent perennials, contains numerous species that exhibit self-incompatibility (SI). The objective of the current study was to determine the number of incompatibility groups present among diploid (2n=2x=22) cultivars of the genera Schlumbergera Lem. (Christmas cacti) and Hatiora Britton & Rose (Easter cacti). Two partial diallel crosses were performed, one with 19 cultivars of Christmas cacti [= S. truncata (Haworth) Moran and S. × buckleyi (Buckley) Tjaden] and the other with 10 cultivars of Easter cacti [= H. gaertneri (Regel) Barthlott, H. rosea (Lagerheim) Barthlott, and H. × graeseri Barthlott ex D. Hunt]. The compatibility/incompatibility status of crosses was determined by percent fruit set and presence of seed in mature fruit. None of the cultivars set fruit when selfed or crossed with a cultivar in the same incompatibility group, but fruit set ranged from 35% to 100% following compatible crosses. For the Christmas cacti, eight intra-incompatible but reciprocally compatible groups were identified, with 13 of the 19 cultivars assigned to three incompatibility groups (68%). The ten cultivars of Easter cacti yielded nine intra-incompatible but reciprocally compatible groups, with two cultivars in one incompatibility group and the other eight cultivars each assigned to a unique group. One cultivar of Christmas cactus ('Abendroth 6') was incompatible when crossed as a male with cultivars in incompatibility group 2 but was compatible in reciprocal crosses, results that suggest that this cultivar is an S-allele homozygote. The crossing relationships are consistent with a one-locus, gametophytic SI system with multiple alleles. Allozyme locus Lap-1, shown previously to be linked with the S locus (recombination frequency 7%) in Schlumbergera, exhibited insufficient allelic diversity for determining the S genotypes of the 19 cultivars of Christmas cacti. Based on the number of incompatibility groups in each diallel, at least five S-alleles occur in the 19 Christmas cacti and the 10 Easter cacti.Publication 3337 of the Massachusetts Agricultural Experiment Station. This material is based on work supported in part by the Cooperative State Research, Extension, Education Service, United States Department of Agriculture, Massachusetts Agricultural Experiment Station, under Project No. 746     

Variation in age and height of onset of reproduction in the saguaro cactus (<Emphasis Type="Italic">Carnegiea gigantea</Emphasis>) in the Sonoran Desert

The objective of this study is to determine the height and age at which reproduction begins (i.e., production of flowers and fruits; the transition to adulthood) in the giant saguaro cactus (Carnegiea gigantea) in four geographically and environmentally distinct populations, and to relate observed variability to environmental differences. The onset of reproduction has been estimated at a height of 2.2 m in near optimal conditions. This value has been widely accepted and applied to populations in less optimal conditions, although variations under less ideal conditions have not been investigated. In addition, previous research has demonstrated that Carnegiea growth rates are highly variable over their range. Thus, even if 2.2 m is a consistent transition height to adulthood over their range, the age of individuals in different populations would be different. I investigate the age and height at which this transition occurs. The author sampled the heights of the shortest reproductive individuals and the tallest non-reproductive individuals to estimate the mean height of the onset of flowering in each of four locales in the northern Sonoran Desert. Using a previously published age-height-growth model, the mean age of the start of reproduction was also computed for the four sites. ANOVA and t-tests were used to compare the average transition to adulthood across sites by both age and height. Statistical results are robust and significant variations in the onset of reproduction are observed by both age and height across the four sites. Saguaro National Park and Organ Pipe Cactus National Monument individuals are transitioning to adulthood, on average, at younger ages and shorter heights than the other two locales. At the arid and marginal Kofa site, individuals that established during the regeneration peak of the late 1800s-early 1900s are only now becoming reproductive (individuals that established around 1899), while at Saguaro National Park, on average, individuals that established in the 1950s are already transitioning to adulthood. These results have implications for regeneration, particularly in marginal locales where regeneration is already limited.     

NUTRIENT LEVELS IN CACTI—RELATION TO NOCTURNAL ACID ACCUMULATION AND GROWTH

The levels of ten essential nutrients and Na in the chlorenchyma and subjacent parenchyma of ten species of cacti were measured along with the maximal rates of nocturnal acid accumulation. Nutrient levels varied considerably among species; also, soil differences between sites affected levels within Opuntia ficus-indica and O. chlorotica. Compared to most agronomic plants, chlorenchyma levels of Ca, Mg, and Mn in cacti tended to be higher and Na lower. Moreover, Ca tended to accumulate in the chlorenchyma with age. The strongest correlation between nutrient level and a metabolic process for the 11 elements tested was with N, where nocturnal acid accumulation tended to be greater when the N level in the chlorenchyma was higher (r² = 0.39). Hydroponically grown seedlings of Carnegiea gigantea, Ferocactus acanthodes, and Trichocereus chilensis responded to N fertilization, reaching about 90% of their maximal growth rates when provided with N at 0.25 x that in Hoagland's solution (namely, 4 mm nitrate). Nocturnal acid accumulation was negatively correlated with the chlorenchyma Na (r² = 0.32), which averaged only 28 ppm for O. ficus-indica (the root contained considerably more) and 234 ppm for the other species. Growth of seedlings was 50% reduced at about 100 mm NaCl for F. acanthodes, T. chilensis, and C. gigantea, while variations in P had a relatively small effect.     

Plant temperatures and heat flux in a Sonoran Desert ecosystem

Summary In the extreme desert environment the potential energy load is high, consequently high temperatures might be a limiting factor for plant survival. Field measurements of plant temperatures in a Sonoran Desert ecosystem were made using fine thermocouples. Temperatures of six desert species were measured: Opuntia engelmannii, Opuntia bigelovii, Opuntia acanthocarpa, Echinocereus engelmannii, Larrea tridentata and Franseria deltoidea. Daily temperature profiles were used to compare the different responses of cacti and shrubs to the desert heat load and also to compare spring and summer responses. Leaf temperature of shrubs was at or near air temperature during both the mild, spring season and the hotter dry season. The cacti, on the other hand, absorbed and stored heat, thus temperatures were often above air temperature. The energy absorbed is determined largely by plant orientation and surface area exposed to the sun. Actual energy absorbed by the plants was estimated from energy diagrams.The flat stem pads of Opuntia engelmannii plants are oriented to receive maximum sunlight without long periods of continuous heating. Opuntia bigelovii spines reflect and absorb much of the environmental energy load, thereby protecting the thick, succulent stems from overheating. The smaller stems of Opuntia acanthocarpa dissipate heat more effectively by their large surface area exposed to convective air currents. Leaves on desert shrubs remain nearer to air temperature than do succulent stems of cacti, because their very large surface to volume ratio allows them to dissipate much heat by convection.     

Pollination ofOpuntia lindheimeri and related species

The large, yellow, bowl-shaped flowers ofOpuntia lindheimeri, O. discata, O. phaeacantha major, andO. compressa in Texas are visited by various species of beetles and bees. The beetles and small bees (Perdita, Dialictus) are pollen thieves. The pollinators are the medium-sized and larger bees (Melissodes, Diadasia, Lithurge, Megachile, Agapostemon, etc.). Different species of theOpuntia lindheimeri group have the same pollination system and there is no evidence of any floral isolation between them. The pollination system of these species ofOpuntia in Texas is essentially the same as that ofEchinocereus fasciculatus andFerocactus wislizeni in Arizona.Pollination of North American Cacti, II.—SeeGrant & Grant (1979).     

Endangered Cactus Restoration: Mitigating the Non-Target Effects of a Biological Control Agent (Cactoblastis cactorum) in Florida

The moth Cactoblastis cactorum (Berg), the poster child of weed biological control in Australia, has recently invaded the United States and threatens native cacti. Concern is greatest for the endangered semaphore cactus, Opuntia corallicola, of which only two known populations exist in the wild. We made three separate outplantings of O. corallicola, designed to bolster the number of extant cacti and to test the effectiveness of three different treatments to protect the cacti from Cactoblastis. In one outplanting, we tested the associational susceptibility hypothesis and found that cacti planted more than 20 m away from the common prickly pear cactus, Opuntia stricta, which act as a reservoir of Cactoblastis, were just as frequently attacked and killed by Cactoblastis as cacti planted within 5 m. In addition, Cactoblastis attack was greater in the shade than in the sun. In the second outplanting, we minimized the attack from Cactoblastis by using protective cages planted at least 500 m from O. stricta in areas not inhabited by cacti. Cages attracted the attention of local animals, which destroyed the cages and trampled the cacti inside to death. Crown rot caused high mortality in this outplanting. In the third outplanting, again conducted at least 500 m away from O. stricta, fertilization did not reduce crown rot mortality. We suggest that increasing populations of O. corallicola in Florida, by means of outplantings, will remain a challenge because of death from Cactoblastis when planted in areas where cacti normally grow and because of death from crown rot in areas where they do not. Because Cactoblastis is moving rapidly northward and westward and has already reached Charleston, South Carolina, rare cacti in the rest of the U.S. Southeast may be in danger. Eventually, many cactus species in the U.S. South, Southwest, and Mexico will likely be threatened by this moth.     

Constant association of ophiostomatoid fungi with the bark beetle <Emphasis Type="Italic">Ips subelongatus</Emphasis> invading Japanese larch logs

Ceratocystiopsis, Ceratocystis, Grosmannia, and Ophiostoma species were isolated from Ips subelongatus and beetle-infested Japanese larch logs collected at several areas in central and northern Honshu Island, Japan, to determine constant associates of I. subelongatus. Ceratocystiopsis minuta, two species of Ceratocystis, three species of Grosmannia, and four species of Ophiostoma were isolated. Of the fungi isolated in the present study, G. laricis, O. brunneociliatum, and O. piceae were constant associates of the beetles. Ceratocystis fujiensis, Ceratocystiopsis minuta, and Ophiostoma sp. F were occasionally isolated with high frequencies of occurrence but were not consistent associates. Ceratocystis fujiensis was most often isolated as the leading fungal invasion in the sapwood of Japanese larch logs invaded by I. subelongatus, confirming that the fungus acts as a primary invader of sapwood in beetle-attacked logs. Contribution No. 221, Laboratory of Plant Parasitic Mycology, Graduate School of Life and Environmental Sciences, University of Tsukuba