Plant hybrid zones as centers of biodiversity: the herbivore community of two endemic Tasmanian eucalypts

We found the hybrid zone between Eucalyptus amygdalina and Eucalyptus risdonii to be a center of insect and fungal species richness and abundance. Of 40 taxa examined, 73% were significantly more abundant in the hybrid zone than in pure zones, 25% showed on significant differences, and 2% were most abundant on a pure host species. The average hybrid tree supported 53% more insect and fungal species, and relative abundances were, on average, 4 times greater on hybrids than on either eucalypt species growing in pure stands. Hybrids may act as refugia for rare species: 5 of 40 species were largely restricted to the hybrid zone. Also, 50% of the species coexisted only in the hybrid zone, making for mique species assemblages. Although hybrids support more species and greater abundances, all hybrids are not equal: 68% of the 40 taxa examined were significantly more abundant on one hybrid phenotype than another. While herbivore concentrations on F1 type intermediates were rare, concentrations were common on phenotypes resembling backcrosses either to E. amygdalina or E. risdonii. For specialist herbivores, the hybrid phenotype most heavily utilized appears to be determined by its phenotypic affinity to its host species. Generalists exhibit an overall greater abundance on hybrids, but are less likely to utilize one hybrid phenotype over another. Mechanistic explanations for these distributions are numerous and probably species specific, but are likely to include: increased genetic susceptibility of hybrids due to hybrid breakdown; increased stress in the hybrid zone resulting in greater plant susceptibility; and a greater diversity of resources in the hybrid zone which could support more species. Seed capsule production by hybrids and their parental species is negatively correlated with herbivory. However, it is difficult to determine whether herbivores cause this pattern as hybrids may have inherently lower sexual reproduction. Laws enacted to protect rare and endangered species do not include hybrids. We argue that a re-examination of our current hybrid policy is warranted. Plant hybrid zones are centers of plant evolution and speciation, sources of economically important plants and potential biocontrol agents, and, as our study suggests, also provide essential habitats for phytophagous communities.     

Enemy-free space? Host preference and larval performance of a willow leaf beetle

To examine whether enemy-free space is an important factor determining the host utilization pattern of a leaf beetle Plagiodera versicolora, we investigated the relationship between adult preference and offspring performance on three co-occurring willow species, Salix sachalinensis, S. miyabeana and S. integra. Salix sachalinensis was by far the most preferred host plant of feeding adults, while both S. miyabeana and S. integra were rarely fed upon. The fact that most oviposition was observed on S. sachalinensis also suggested that P. versicolora preferred S. sachalinensis to other willows for oviposition. This adult preference did not correspond well to patterns of larval performance on the three willow species in the absence of enemies. Higher survivorship, shorter developmental time and larger adult size were achieved on S. sachalinensis and S. miyabeana than on S. integra. Performance as indicated by female adult size and development time on S. miyabeana were higher than on S. sachalinensis. In the presence of enemies, however, the survivorship of first-instar larvae on S. miyabeana was much lower than on other willows. Adults of P. versicolora apparently avoided S. miyabeana as an oviposition and feeding host and preferred S. sachalinensis as an enemy-free space. This was not because larvae had poorer performance on S. miyabeana, but because predation pressure on eggs and early instar larvae was more severe on S. miyabeana.     

Interspecific and temporal variation in herbivore responses to hybrid willows

We studied herbivory of two species of willows (Salix sericea and S. eriocephala) and their interspecific hybrids to test alternative hypotheses concerning the effects of hybridization on plant resistance. Individually marked plants were identified using morphological traits in the field and random amplified polymorphic DNA (RAPD) band analysis was used to verify the genetic status of many parental and hybrid plants. The desities of 12 herbivore species on plants in the field were compared between two parents and their F₂-type hybrids. We found about equal support for the additive, dominance, and hybrid susceptibility hypotheses over 4 years. In one year, one species supported the hybrid resistance hypothesis. Guild membership was not a good predictor of similar responses of species to hybrid versus parental plants. There were marked differences in support for particular hypotheses among years for four herbivore species. This study demonstrates the diversity of responses of phytophages in response to interspecific hybridization, and indicates that year-to-year variation in relative resistance of hybrid plants can be important.     

Plant genetics affects arthropod community richness and composition: evidence from a synthetic eucalypt hybrid population

To examine how genetic variation in a plant population affects arthropod community richness and composition, we quantified the arthropod communities on a synthetic population of Eucalyptus amygdalina, E. risdonii, and their F1 and advanced-generation hybrids. Five major patterns emerged. First, the pure species and hybrid populations supported significantly different communities. Second, species richness was significantly greatest on hybrids (F1 > F2 > E. amygdalina > E. risdonii). These results are similar to those from a wild population of the same species and represent the first case in which both synthetic and wild population studies confirm a genetic component to community structure. Hybrids also acted as centers of biodiversity by accumulating both the common and specialist taxa of both parental species (100% in the wild and 80% in the synthetic population). Third, species richness was significantly greater on F1s than the single F2 family, suggesting that the increased insect abundance on hybrids may not be caused by the breakup of coadapted gene complexes. Fourth, specialist arthropod taxa were most likely to show a dominance response to F1 hybrids, whereas generalist taxa exhibited a susceptible response. Fifth, in an analysis of 31 leaf terpenoids that are thought to play a role in plant defense, hybrids were generally intermediate to the parental chemotypes. Within the single F2 family, we found significant associations between the communities of individual trees and five individual oil components, including oil yield, demonstrating that there is a genetic effect on plant defensive chemistry that, in turn, may affect community structure. These studies argue that hybridization has important community-level consequences and that the genetic variation present in hybrid zones can be used to explore the genetic-based mechanisms that structure communities.     

Plant genetic determinants of arthropod community structure and diversity

To test the hypothesis that genes have extended phenotypes on the community, we quantified how genetic differences among cottonwoods affect the diversity, abundance, and composition of the dependent arthropod community. Over two years, five major patterns were observed in both field and common-garden studies that focused on two species of cottonwoods and their naturally occurring F1 and backcross hybrids (collectively referred to as four different cross types). We did not find overall significant differences in arthropod species richness or abundance among cottonwood cross types. We found significant differences in arthropod community composition among all cross types except backcross and narrowleaf cottonwoods. Thus, even though we found similar richness among cross types, the species that composed the community were significantly different. Using vector analysis, we found that the shift in arthropod community composition was correlated with percent Fremont alleles in the host plant, which suggests that the arthropod community responds to the underlying genetic differences among trees. We found 13 arthropod species representing different trophic levels that were significant indicators of the four different cross types. Even though arthropod communities changed in species composition from one year to the next, the overall patterns of community differences remained remarkably stable, suggesting that the genetic differences among cross types exert a strong organizing influence on the arthropod community. Together, these results support the extended phenotype concept. Few studies have observationally and experimentally shown that entire arthropod communities can be structured by genetic differences in their host plants. These findings contribute to the developing field of community genetics and suggest a strategy for conserving arthropod diversity by promoting genetic diversity in their host plants.     

Community structure of insect herbivores in a hybrid system: examining the effects of browsing damage and plant genetic variation

Abstract.  1. Using three genetic classes of willows, Salix eriocephala , Salix sericea , and their interspecific F1 hybrid, the influence of browsing damage and the importance of genetic class on insect community structure were evaluated.
2. Three-year-old plants grown from seeds generated from controlled crosses were placed in a common garden after a damage treatment was imposed on them (plants were either left undamaged during the previous winter or they had 50% of the previous year's growth removed). Clipping damage caused large increases in mean shoot length for plants.
3. The abundance of eight species of insect herbivores was determined for every plant to evaluate community structure for three genetic classes across the two damage levels. Based on manova , damage treatment had a modest effect on the relative abundance of herbivores (i.e. their proportional representation). In contrast, dramatic differences were detected among genetic classes for relative abundance; in cases where damage treatment influenced relative abundance of herbivores, the importance of genetic class was at least 20-fold greater than that of damage treatment. No interaction between genetic class and browsing treatment was detected for community structure.
4. The weak response of the herbivore community to clipping damage, contrasted to the large response to genetic class, was very surprising because mean shoot length was greatly altered by damage treatment. These findings, coupled together with previous research, suggest that plant genetic differences can act as the primary basis for herbivore community structure, while the effects of browsing may not be as common.     

Induced resistance in mountain birch: defence against leaf-chewing insect guild and herbivore competition

Summary The effect of leaf damage simulating the feeding of early season insect herbivore species, e.g. Epirrita autumnata, to mountain birch, Betula pubescens ssp. tortuosa, on the performance of insect larvae was studied with eleven leaf-chewing sawfly species. I found variation in the results that was due to short- and long-term inducible responses and to the phenology of herbivore species. In general, early and mid-season species were more strongly affected by induced reactions than late-season species. This finding is in accordance with earlier results but I could show that the persistance of induced reactions rather than the influence of timing of damage is responsible for the result. The growth of the larvae of mid-season sawfly species was affected by both short- and long-term induced reactions. This result shows that early season species may escape short-term induced reactions of mountain birch in current year but may not avoid long-term effects. It is supposed that seasonal deterioration of leaf quality either masks the effects of induced defences or late-season species are better adapted to low-quality leaves. Some species show variation in their response to induced defence in different years. This may be due to yearly differences in induced reactions as well as to species-specific responses. Induced defence reactions may play a role in competitive interactions between herbivore species in leaf-chewing guild of mountain birch.     

Genetic and soil-nutrient effects on the abundance of herbivores on willow

The effects of soil-nutrient environment, plant genotype, and the interaction between the two on the resistance of the willow, Salix sericea, to insect species in a diverse herbivore community was measured. We found that soil-nutrient environment influenced plant growth and the abundance of most herbivores of S. sericea. However, environmental effects on herbivore abundance were often modified by plant genetics; the abundance of four of seven herbivores exhibited significant genotypeby-environment interaction effects. Pure genotype effects were mostly small and non-significant. The effects of fertilization differed among herbivores. Several herbivores were more abundant on fertilized plants, one was less abundant, and the abundance of others did not change. We found that feeding guild was a poor predictor of herbivore response. Finally we found significant phenotypic and genetic correlations among growth rate, internode length, and the abundances of several herbivores.     

Tree growth indicates resource quality for foliage-feeding insects: Pattern and structure of herbivore diversity in response to productivity

In forests, local site conditions can affect both trees and herbivores and hence site-related factors act indirectly on herbivores mediated by tree growth rates. Here, tree foliage represents a fundamental prerequisite for insect herbivore development providing energy in the form of plant tissue quality. Resource-based theories, on the other hand, assume that the synthesis of defensive compounds is a trade-off with growth and peaks at low resource availability. However, the extent to which plant tissue quality in response to site productivity is relevant in the species-energy relationship is unknown. Therefore, we aimed at a better understanding of the form and structure of the species-energy relationship in forest insects. We used census data of foliage-feeding insects along a productivity gradient of Scots pine forests defined by relative growth rates of trees (RGR). As a result, diversity monotonically increases with decreasing RGR (as a proxy for energy) during almost two decades of sampling. Herbivore assemblages become more similar with available energy as species turnover linearly decreases and proportions of sites occupied by individual species rise. The results suggest that tree growth rate influences herbivore dynamics in this system by altering the chemical composition of needles, without necessarily affecting the form in the relationship. The site-specific resource availability requires trees to adjust their allocation to synthesis of carbon-based secondary metabolites or growth, which then results in fundamental differences in herbivore dynamics at low vs. lowest RGR (regular cycles (dominance) vs. dampened cycles (evenness)). However, these differences inevitably demonstrate that species richness is not necessarily a result of more individuals and implicate that different mechanisms are involved (facilitation vs. competition/temporal heterogeneity). The resulting pattern and structure of foliage-feeding insects advance our understanding of herbivore dynamics in response to site quality and tree growth, which may ultimately improve our knowledge of plant-insect interactions in the face of environmental change.     

Fertility and sexual structure in a polygamous willow population

This note reports on an extraordinary polygamous population of Salix acmophylla from Nahal Dishon, Israel. Remarkably, all individuals in this population are bisexuals, that is, they all contain typically female catkins (with some or without any male florets), typically male catkins (with some or without any female florets) and mixed catkins. The proportions of these three catkin types in populations are 36.3%, 46.8% and 16.9% respectively. The fertility of these willows was found to be poor: no mature seeds were found at all and mean pollen stainability was low as 15.6%. The possible reasons for this semi-sterility are discussed.     

Molecular evidence indicates that subarctic willow communities in Scotland support a diversity of host-associated Melampsora rust taxa

Rare and threatened subarctic willow scrub communities in the UK are the subject of ongoing conservation programmes, yet little is known about the diversity of fungal taxa that they support. Isolates of the rust genus Melampsora were sampled from 112 leaves of eight subarctic willow (Salix) taxa and their hybrids from twelve sites in the UK. In order to determine the number of Melampsora taxa present in the samples, isolates were sequenced for the Internal Transcribed Spacer (ITS) region of rDNA and data were subject to phylogenetic analysis. Maximum likelihood and Bayesian analysis indicated that the isolates fell into three strongly supported host-associated clades. Clade I contained only isolates from Salix herbacea and was distinguished morphologically by dense urediniospore echinulation and thin cell walls. Clade II contained isolates from Salix arbuscula and Salix reticulata only. These could not be distinguished morphologically from isolates in Clade III which were found on Salix lapponum, Salix myrsinites, Salix myrsinifolia, Salix aurita, Salix lanata, and their hybrids. Clade II was most distinct in ITS sequence, differing by 50 bases from Clades I and III, while the latter clades differed in sequence by only 24 bases on average. Clades I and III are likely to represent the previously recognised taxa Melampsora alpina Juel 1894 and Melampsora epitea Thüm. 1879 respectively, but Clade II has not apparently been described before. Significant differences in the intensity of infection by isolates of Clade III were found among different Salix species at a single site, suggesting either differences in resistance among Salix taxa, or the presence of further cryptic taxa within Clade III. The study illustrates the power of molecular phylogenetic analysis to reveal cryptic biodiversity within Melampsora, and suggests that conserving Salix host diversity within subarctic willow communities will ensure that a diversity of associated Melampsora taxa is maintained.     

Plant regrowth response to a stem-boring insect: a swift moth-willow system

To examine plastic willow regrowth response to herbivory, we studied the effect of a boring insect, the swift moth Endoclita excrescens (Hepialidae: Lepidoptera), which does not remove apical meristems, on shoot growth in three willow species—Salix gilgiana, S. eriocarpa, and S. serissaefolia−by direct observations and experiments in the field. We hypothesized that the stem-boring could initiate new lateral bud activation, and result in secondary shoot regrowth without the removal of the primary apical meristems. There were significantly more lateral shoots on naturally attacked than unattacked stems, and a significant positive correlation between lateral shoot density and the number of swift moth tunnels per tree was observed for all three willow species. Artificial boring, and larval infestation, resulted in an increase in the number of lateral shoots, but did not affect growth of current-year shoots. The length of lateral shoots differed between species, being significantly longer in S. gilgiana than S. eriocarpa and S. serissaefolia. The results of this study show that compensatory regrowth can result even if herbivory does not remove the apical meristem. We argue that this type of plant compensatory response is probably widespread, given that the stem-boring is a common feeding type of insect herbivores.     

Host plants affect predator fitness via the nutritional value of herbivore prey: Investigation of a plant-aphid-ladybeetle system

The interactions among host plants(Medicago sativa L., cv. `OKO8' and Vicia faba L., cv. `Windsor'), aphid prey(Acyrthosiphon pisum Harris, Homoptera:Aphididae), and Coccinella septempunctata L. (Coleoptera: Coccinellidae) preimaginal biology were evaluated. Interactions were measured over a range of limiting daily prey levels (1.2 mg–16.4 mg)from each host plant colony. Compared withA. pisum reared on V. faba, A. pisum reared on M. sativa storedsignificantly more fatty acids which resultedin a 1.17-fold increase in available caloriesfor developing C. septempunctata. Theincreased survival, decreased developmentaltimes, and larger size of C.septempunctata supplied with A. pisumreared on M. sativa clearly demonstratehost plant effects at the third trophic level. At low very limiting daily prey levels, A. pisum reared on M. sativa were moresuitable prey for C. septempunctatasurvival, development, and adult size thanA. pisum reared on V. faba. Coccinella septempunctata survival ratios(larval), developmental times, and adult sizeconverged (were not statistically different)between host plants at higher daily A.pisum levels. These convergence's supportthe hypothesis that there were quantitativedifferences in the nutritional value ofaphids, as influenced by differences in fattyacids and subsequent nutritional levels(calories), between aphids reared on separateplant hosts. The observed tritrophicinteractions appear to be modulated by thebiochemical response of A. pisum to hostplants.     

Fungal colonization of shrub willow roots at the forefront of a receding glacier

Shrub willows (Salix spp.) form associations with arbuscular mycorrhizal (AM), ectomycorrhizal (EM) and dark septate endophytic (DSE) fungi. Willow root colonization by these three types of fungi was studied on a deglaciated forefront of Lyman Glacier, Washington, USA. Root colonization was low; less than 1% of the root length was colonized by AM and 25.6% by DSE. EM colonized 25% of the root tips and 19.4% of the root length. AM and DSE colonization were not related to distance from the present glacier terminus or to canopy cover. EM colonization increased with distance from the glacier terminus based on gridline intercept data but not on root tip frequency data. Availability of propagules in the substrate was low, but numbers of propagules increased with distance from the glacier terminus. The EM communities were dominated by three ascomycetes showing affinity to Sordariaceae in BLAST analyses. Other frequent taxa on the glacier forefront included species of Cortinariaceae, Pezizaceae, Russulaceae, Thelephoraceae and Tricholomataceae. When occurrence of individual taxa was used as a response variable to canopy cover, distance from the glacier terminus, and their interaction, four different fungal guilds were identified: 1) fungi that did not respond to these environmental variables; 2) fungi that occurred mainly in intercanopy areas and decreased with distance from the glacier terminus; 3) fungi that were insensitive to canopy cover but increased with distance from the glacier terminus; 4) fungi that occurred mainly under willow canopies and increased with distance from the glacier terminus. We suggest that fungal colonization is mainly limited by fungal propagule availability. Environmental conditions may also limit successful establishment of plant-fungus associations. We propose that the four EM guilds partly explain successional dynamics. The initial EM community comprises fungi that tolerate low organic matter and nitrogen environment (first and second guilds above). During later community development, these fungi are replaced by those that benefit from an increased organic matter and nitrogen environment (third and fourth guilds above).     

Host plant preference based on salicylate chemistry in a willow leaf beetle (Chrysomela aeneicollis)

Summary Chrysomela aeneicollis (Coleoptera: Chrysomelidae) uses salicin from its host plant (Salix spp.) to produce a defensive secretion, salicylaldehyde. Because it requires salicin for this secretion, I predicted that C. aeneicollis should be attracted to willows which possess salicin and other salicylates. To test this prediction, I determined the host-plant preferences of C. aeneicollis among four potential hosts which occur in the Sierra Nevada range of eastern California. These species have very different salicylate chemistries but do not differ in nutritional quality for C. aeneicollis. In oviposition-preference tests, gravid females showed no preference between a salicylate-poor species, S. lutea, and a salicylate-rich species, S. orestera. However in feeding-choice tests, both larvae and adults preferred S. orestera over S. lutea. This preference was not affected by the species on which the larvae were reared. In other feeding tests, adults preferred S. orestera over two medium-salicylate species, S. boothi and S. geyeriana, regardless of which host species they had been feeding on in nature. In a final feeding test, adults were stimulated to feed by salicin itself. In nature, the relative abundances of C. aeneicollis adults and egg clutches among these species correspond to the adult feeding preference in the laboratory. Additionally, multiple regression analyses showed that adult abundance was not related to among-clone differences in leaf toughness or nutritional quality, but rather to salicin content and plant size. Thus for C. aeneicollis, both laboratory and field results demonstrate a preference for salicylate-rich willows which is partly responsible for the increased level of attack on them.     

Estimation of the effect of photoinhibition on the carbon gain in leaves of a willow canopy

The occurrence of photoinhibition of photosynthesis in leaves of a willow canopy was examined by measuring the chlorophyll-a fluorescence ratio of F _V/F _M (F_M is the maximum fluorescence level of the induction curve, and F_V is the variable fluorescence, F _V=F _M–F ₀, where F₀ is the minimal fluorescence). The majority of the leaves situated on the upper parts of peripheral shoots showed an afternoon inhibition of this ratio on clear days. This was the consequence of both a decrease in F _M and a rise in F _O. In the same leaves the diurnal variation in intercepted photosynthetic photon flux density (PPFD) was monitored using leaf-mounted sensors. Using the multivariate method, partial least squares in latent variables, it is shown that the dose of PPFD, integrated and linearly weighted over the last 6-h period, best predicts photoinhibition. Photoinhibition occurred even among leaves that did not intercept PPFDs above 1000 mol·m^–2·s^–1. Exposure of leaves to a standard photoinhibitory treatment demonstrated that the depression in the F _V/F _M ratio was paralleled by an equal depression in the maximal quantum yield of CO₂ uptake and a nearly equal depression in the rate of bending (convexity) of the light-response curve of CO₂ uptake. As a result, the rate of net photosynthesis is depressed over the whole natural range of PPFD. By simulating the daily course in the rate of net photosynthesis, it is estimated that in the order of one-tenth of the potential carbon gain of peripheral willow shoots is lost on clear days as a result of photoinhibition. This applies to conditions of optimal temperatures. Photoinhibition is even more pronounced at air temperatures below 23° C, as judged from measurements of the F_V/F_M ratio on clear days: the afternoon inhibition of this ratio increased in a curvilinear manner from 15% to 25% with a temperature decrease from 23° to 14° C.Abbreviations and Symbols F_O minimum fluorescence - F_V variable fluorescence - F_M maximum fluorescence - PLS partial least squares in latent variables - PPFD photosynthetic photon flux density - VPD water vapour-pressure deficit This study was supported by the Swedish Natural Science Research Council. We are indebted to Dr. Jerry Leverenz (Department of Plant Physiology, University of Umeå, Sweden) for guidance with the modelling of the photosynthesis data.     

Community heritability measures the evolutionary consequences of indirect genetic effects on community structure

The evolutionary analysis of community organization is considered a major frontier in biology. Nevertheless, current explanations for community structure exclude the effects of genes and selection at levels above the individual. Here, we demonstrate a genetic basis for community structure, arising from the fitness consequences of genetic interactions among species (i.e., interspecific indirect genetic effects or IIGEs). Using simulated and natural communities of arthropods inhabiting North American cottonwoods (Populus), we show that when species comprising ecological communities are summarized using a multivariate statistical method, nonmetric multidimensional scaling (NMDS), the resulting univariate scores can be analyzed using standard techniques for estimating the heritability of quantitative traits. Our estimates of the broad-sense heritability of arthropod communities on known genotypes of cottonwood trees in common gardens explained 56-63% of the total variation in community phenotype. To justify and help interpret our empirical approach, we modeled synthetic communities in which the number, intensity, and fitness consequences of the genetic interactions among species comprising the community were explicitly known. Results from the model suggest that our empirical estimates of broad-sense community heritability arise from heritable variation in a host tree trait and the fitness consequences of IGEs that extend from tree trait to arthropods. When arthropod traits are heritable, interspecific IGEs cause species interactions to change, and community evolution occurs. Our results have implications for establishing the genetic foundations of communities and ecosystems.     

Secretory avocado idioblast oil cells: evidence of their defensive role against a non-adapted insect herbivore

We tested the hypothesis that avocado idioblast oil cells play a defensive role against herbivorous insects. Toxicities of the intact avocado idioblast oil cells and the extracted idioblast oil were compared for three insect herbivores. Spodoptera exigua (Hübner) larvae are generalists that do not feed on avocados. By contrast, Sabulodes aegrotata (Guenée) and Pseudoplusia includens (Walker) larvae are generalist herbivores that readily feed on avocados. All bioassays were performed at a naturally occurring concentration of idioblast oil cells (2% w/w). Choice experiments showed that S. exigua larvae avoided diet treated with avocado idioblast oil cells and consume more control than treated diet. In contrast, idioblast oil cells had no significant antifeedant effects on the adapted S. aegrotata and P. includens larvae. Subsequent experiments designed to assess resistance mechanisms separated pre-ingestive (behavioral) and post-ingestive (physiological) effects of the avocado idioblast oil cells, and the extracted idioblast oil, on the two adapted herbivores. Post-ingestive adaptation was the mechanism that allows feeding. Because the impact of the avocado idioblast oil cells was greatest on the performance of non-adapted S. exigua, additional experiments determined that larvae fed diet containing the oil cells had higher mortality and reduced larval growth compared to controls. Developmental times were significantly prolonged for the survivors. Thus, increased mortality, reduced developmental rates, and antifeedant activity in the non-adapted insect indicate that defense against non-adapted herbivores may be an important function of idioblast cells in avocados.     

Associational plant refuges: convergent patterns in marine and terrestrial communities result from differing mechanisms

Summary An associational plant refuge occurs when a plant that is susceptible to herbivory gains protection from herbivory when it is associated with another plant. In coastal North Carolina, the abundance of the palatable red alga Gracilaria tikvahiae is positively correlated with the abundance of the unpalatable brown alga Sargassum filipendula during times of increased herbivore activity. To see if grazing by the sea urchin Arbacia punctulata could generate this pattern, controlled experiments were conducted in out-door microcosms and in the laboratory. Gracilaria beneath a canopy of Sargassum was eaten significantly less than Gracilaria alone. When Arbacia were excluded, Gracilaria alone grew significantly more than Gracilaria beneath Sargassum, demonstrating that Sargassum is a competitor of Gracilaria. Experiments investigating Sargassum's deterrent role indicated that Sargassum decreased the foraging range of Arbacia and the rate at which it fed on Gracilaria. Additional experiments with plastic Sargassum mimics indicated that the decreased grazing on Gracilaria was not a result of Sargassum morphology, but was probably attributable to some chemical characteristic of Sargassum. The pattern of increased grazing in monocultures (only Gracilaria present) versus polycultures (both Gracilaria and Sargassum present) demonstrated in this study also has been demonstrated for plant-insect interactions in terrestrial communities. In these communities, insect density is higher in monocultures than in polycultures because insects find and immigrate to monocultures more rapidly, and once in a monoculture, they emigrate from them less often than from polycultures. In this study, urchins did not find and immigrate to monocultures more rapidly, nor did they tend to stay in them once they were found; in fact, they emigrated from monocultures of Gracilaria more rapidly than from Gracilaria and Sargassum polycultures. Increased grazing in Gracilaria monocultures resulted from increased rates of movement and feeding of individual herbivores, not from increased herbivore density as has been reported for terrestrial systems.     

Population structure and genetic diversity in two species of Hawaiian picture-winged Drosophila

Over the last several decades many picture-winged Drosophila have become less common in both geographical distribution and local population size (pers. obs., Foote pers. comm., Montgomerey pers. comm.). Here we report on a study of two Hawaiian Drosophila species, D. engyochracea, and D. hawaiiensis, to determine the impact that changes in population sizes over the past thirty years have had on the genetic diversity of these species. D. engyochracea is known from only two locations on the Island of Hawai'i (Kipuka Ki and Kipuka Pua'ulu), while D. hawaiiensis is currently more wide spread across Hawai'i Island. We collected 65 D. hawaiiensis and 66 D. engyochracea from two forest patches (kipuka) isolated by a 400 year old volcanic ash deposit. DNA sequence data for 515 bases of the mitochondrial gene COII was analyzed for both species to estimate relative total genetic diversity as well as inter-kipuka gene flow. The more wide spread species, D. hawaiiensis, has more genetic diversity (23 vs. 11 unique haplotypes) than the rarer species, D. engyochracea. The distribution of haplotypes in the kipuka is consistent with more gene flow in D. engyochracea than in D. hawaiiensis. Phylogenetic analysis indicates a small number of individuals morphologically identified as one species but have DNA sequence diagnostic for the other species. These results are consistent with these individuals being descendant from hybrids between species.