Chemically Mediated Species Recognition in Closely Related Podarcis Wall Lizards

In many animals, chemical signals play an important role in species recognition and may contribute to reproductive isolation and speciation. The Iberian lizards of the genus Podarcis, with up to nine currently recognized lineages that are often sympatric, are highly chemosensory and provide an excellent model for the study of chemically mediated species recognition in closely related taxa. In this study, we tested the ability of male and female lizards of two sister species with widely overlapping distribution ranges (Podarcis bocagei and P. hispanica type 1) to discriminate between conspecific and heterospecific mates by using only substrate-borne chemical cues. We scored the number of tongue flicks directed at the paper substrate by each individual in a terrarium previously occupied by a conspecific or a heterospecific lizard of the opposite sex. Results show that males of P. bocagei and P. hispanica type 1 are capable of discriminating chemically between conspecifics and heterospecifics of the opposite sex, but females are not. These results suggest that differences in female, but not male, chemical cues may underlie species recognition and contribute to reproductive isolation in these species. The apparent inability of females to discriminate conspecific from heterospecific males, which is not because of reduced baseline exploration rates, is discussed in the context of sexual selection theory and species discrimination.     

Interspecific pairing between two siblingIps species (Coleoptera: Scolytidae)

Host and conspecific discrimination were tested as reproductive isolating mechanisms betweenIps paraconfusus Lanier infestingPinus coulteri (Torrey) andI. confusus (Le Conte) infestingP. monophylla (Fremont). In two areas (one area largely Coulter pine and the other largely pinyon pine) where these bark beetles and hosts cooccur in southern California, we induced pheromone production in host and nonhost logs using males of each species.Ips paraconfusus females joined both heterospecific and conspecific males tunneling in both tree species in each area.Ips confusus females failed to joinI. paraconfusus males in Coulter pine, but joinedI. paraconfusus in pinyon pine. Sympatry was demonstrated when females of both sibling species joined conspecific males in their respective hosts. Males attacked all four beetlehost treatment combinations in both areas. Laboratory tests confirmed these results. Males did not displace heterospecific males from nuptial chambers in hosts, and they did not occupy a gallery in which heterospecific males produced frass. Females that left conspecific males in a host were readily accepted by heterospecific males and oviposited in a nonhost.     

Allelopathic properties of α-terthienyl and phenylheptatriyne,naturally occurring compounds from species of asteraceae

Alpha-terthienyl (-T), a naturally occurring polyacetylene derivative from roots ofTagetes erecta L., and phenylheptatriyne (PHT), from leaves ofBidens pilosa L., were tested as possible allelopathic agents against four seedling species (Asclepias syriaca L.,Chenopodium album L.,Phleum pratense L.,Trifolium pratense L.).Asclepias was the most sensitive of the species. Allelopathic activity was enhanced in the presence of sunlight or sources of near-UV, with LC_50s forA. syriaca of 0.15 ppm and 0.66 ppm with -T and PHT, respectively; 0.27 and 0.85 forC. album; 0.79 and 1.43 forP. pratense, and 1.93 and 1.82 forT. pratense. Near-UV exposure was saturating but never more than found in summer sunlight at Ottawa, Canada. Growth inhibition was observed with seedlings treated with -T and PHT but without near-UV irradiation. Germination of seedlings was also sensitive to -T and PHT with or without near-UV treatment. -T was extracted from soil surrounding the roots ofTagetes. Concentrations calculated for the soil (0.4 ppm) indicate that seedling growth could be significantly hindered. The activity and specificity of -T was sufficiently high to warrant future field trials to assess its potential as a natural weed-control agent.     

Interspecific Pheromone Plume Interference Among Sympatric Heliothine Moths: A Wind Tunnel Test Using Live, Calling Females

Three species of North American heliothine moths were used to determine the level at which interspecific female interference of male attraction to conspecific females occurs. We used live calling females of Heliothis virescens, H. subflexa, and Helicoverpa zea, as lures for conspecific males in a wind tunnel, and then placed heterospecific females on either side of the original species such that the plumes of the three females overlapped downwind. In nearly all combinations, in the presence of heterospecific females, fewer males flew upwind and contacted or courted the source than when only conspecific females were used in the same spatial arrangement. Males did not initiate upwind flight to solely heterospecific female arrangements. Our results show that the naturally emitted pheromone plumes from heterospecific females of these three species can interfere with the ability of females to attract conspecific males when multiple females are in close proximity. However, the fact that some males still located their calling, conspecific females attests to the ability of these male moths to discriminate point source odors by processing the conflicting information from interleaved strands of attractive and antagonistic odor filaments on a split-second basis.     

Role of allelopathy as a possible factor associated with the rising dominance ofBunias orientalis L. (Brassicaceae) in some native plant assemblages

Leaf extracts ofBunias orientalis were shown to inhibit seed germination of a variety of cultivar plant species and of species cooccurring withB. orientalis in the field. Root exudate solutions and leaf litter leachates ofB. orientalis were tested for their allelopathic activity using seedling growth assays. Additionally, in comparative seedling growth assays soil cores removed from denseB. orientalis stands were tested bimonthly for elevated allelopathic effects. The impact of root exudates on seedling growth was generally weak and varied between species. Similar results were obtained for the effect ofB. orientalis leaf litter leachates on seedlings grown in sand culture relative to the effect of leaf litter leachates of a plant species mixture. When soil as a growth substrate was used, no consistent differences in seedling growth were obtained between the two litter leachate treatments. In the soil core experiment seedlings grown in soil cores collected from a denseB. orientalis stand unexpectedly showed better performance than seedlings grown in soil cores collected from a nearby mixed plant stand withoutB. orientalis, at least in early spring and late autumn. Predominating nutrient effects are, therefore, assumed to conceal a potentially increased allelopathic effect of soil beneath denseB. orientalis stands. It is concluded that other factors than allelopathy must be investigated to explain the rapid establishment of dense stands of this alien plant species.     

Perspectives on allelopathy in mexican traditional agroecosystems: A case study in tlaxcala

Agroecosystems in Tlaxcala, Mexico, are surrounded by trees and water channels and have a great variety of cultivated and noncultivated plants. The main results of a study carried out on a traditional agroecosystem in Santa Inés, Tlaxcala are presented. Some ecological aspects of polycultures, plant covers (dry leaves ofAlnus firmifolia, Berula erecta, andJuncus sp.), and the allelopathic potential of crops and noncultivated plants (fresh and dry material) were analyzed. The main plants (trees, shrubs, and herbs) present in the agroecosystem were identified. The total number of weeds in plots where plant covers were added was reduced. The number of nodules ofRhizobium phaseoli and the production of bean and squash increased with plant covers. Corn, beans, and squash showed a clear allelopathic effect, as well asChenopodium murale, Tradescantia crassifolia, Melilotus indicus, andAmaranthus hybridus, among other weeds. The contribution of allelopathy in studies of traditional agroecosystems is of great importance for the management of species in space and time. Allelopathy can be the basis of biological control of pests and weeds and of the discovery of new useful substances.     

Asymmetric small-scale distribution and allelopathy: Interaction betweenRumex obtusifolius L. and meadow species

Analysis of small-scale distribution may allow allelopathic interaction among plant species to be distinguished from competitive interaction. If all environmental factors except slope are uniform, allelopathy produces spatial asymmetry in the association with a hypothetical receptor species and the corresponding hypothetical emitter species in the neighborhood of the latter. The interaction betweenRumex obtusifolius and the meadow speciesLolium perenne, Poa pratensis, Dactylis glomerata, andTrifolium repens is analyzed by way of example.     

In search of allelopathy in the Florida scrub: The role of terpenoids

The hypothesis was tested that allelopathic agents released from fire-sensitive plants of the Florida scrub community deter the invasion of fireprone sandhill grasses. The structures of the constituents of four endemic scrub species,Conradina canescens, Calamintha ashei, Chrysoma pauciflosculosa, andCeratiola ericiodes, were established and their phytotoxic activity against two grasses of the sandhill was examined. Effects of the secondary metabolites from the above scrub species and their degradation products upon the germination and radicle growth of little bluestem (Schizachyrium scoparium) and green sprangletop (Leptochloa dubia), two native grasses of the Florida sandhill community, were determined. The studies included determination of the water solubility and release mechanism of terpenes and other allelopathic agents from the source plants and their aqueous transport to the target species. Some of the natural products were nontoxic until activated by light and/or oxidation after release from the source plant into the environment.Dedicated to one of the pioneers in allelopathy, Cornelius H. Muller.     

Effects of pine-produced chemicals on selected understory species in aPinus ponderosa community

Pinus ponderosa accounted for more than 98% of all tree and shrub stratum stems in a climax community with low herb coverage and aboveground biomass, 35% and 60 g/m², respectively. Because of our previous report that nitrification and nitrifying bacteria in the same community were allelopathically inhibited, we speculated that the pine-produced allelochemics might also directly influence the development and growth of the herb stratum. In most cases decaying needles, needle leachate, and field soils significantly reduced germination and radicle growth ofAndropogon gerardii andA. scoparius, pine-associated herbaceous species. Additionally, growth ofAndropogon scoparius seedling radicles was reduced 28–56% by pine needle extracts, 33% by pine bark extracts, and 67% by soil hydrolysate extracts.Andropogon seed germination was reduced 20–25% by pine needles and soil. Phytotoxins identified in various plant parts and associated soils were caffeic acid, chlorogenic acid, quercetin, and condensed tannins. Pine needle water and soil hydrolysate extracts were most inhibitory to the radicle growth of the test species. Thus it appears that the limited growth of the herbaceous stratum in the pine community may be accounted for, in part, by allelopathy. Such allelopathic interactions may have an adaptive ecological significance in various forest and other plant communities.     

Allelopathic Potential of Aquatic Plants Associated with Wild Rice (Zizania palustris): I. Bioassay with Plant and Lake Sediment Samples

The allelopathic potential of eight aquatic plants associated with wild rice was investigated using lettuce and wild rice seedling bioassays. Rhizome aqueous extracts of Scirpus acutus, Potamogeton natans, Nymphaea odorata, Nuphar variegatum; shoot extract of Eleocharis smallii; whole plant extract of Myriophyllum verticillatum; and leaf extract of P. natans significantly reduced the root length of lettuce and wild rice seedlings. The lettuce seedling bioassay was more sensitive than the wild rice bioassay. Shoot growth was less affected than the root growth. Water extract of sediments associated with the aquatic plants had little growth inhibitory effect on wild rice. Our study did not yield any conclusive evidence that the wild rice-associated aquatic plants have allelopathic effects on wild rice. We emphasize the use of target species as a bioassay material in allelopathic studies. Further investigation on allelopathic effects of lake sediments associated with the neighboring plants of wild rice is necessary to evaluate their ecological significance.     

Role of sex pheromone components in behavioral reproductive isolation betweenAutographa gamma (L.) and eitherTrichoplusia ni (Hübner) ORChrysodeixis chalcites (Esp.) (Lepidoptera: Noctuidae: Plusiinae)

The composition of theAutographa gamma sex pheromone was reexamined and only (Z)-7-dodecenyl acetate and (Z)-7-dodecenol were identified by capillary GC, GC-MS, and dimethyl disulfide derivatization and subsequent GC-MS analysis. The fatty acid content of the pheromone glands was also studied, and a series of saturated and unsaturated acids was identified. However, most of the related pheromonal compounds were not detected. The male response to the pheromone components was studied in a flight tunnel and compared with the response to calling females. The best synthetic baits evoked a response similar to that observed to the virgin females, but males spent significantly more time at calling females than at the synthetic baits. The preferred synthetic baits consisted of (Z)-7-dodecenyl acetate alone or of a blend with 5% (Z)-7-dodecenol. Increasing the relative amount of the alcohol caused a gradual reduction in male response, particularly in the last steps of the courtship sequence. The addition of the minor sex pheromone components of the sympatric Plusiinae species,Trichoplusia ni andChrysodeixis chalcites, to theA. gamma pheromone was also investigated in the flight tunnel. Some of these components exhibited a significantly antagonistic effect on theA. gamma male courtship behavior. The most potent antagonists were (Z)-5-dodecenyl acetate and (Z)-9-tetradecenyl acetate. The response ofA. gamma andT. ni males to conspecific and heterospecific females was also compared in the flight tunnel. WhereasA. gamma males were attracted only to their conspecific females, a small percentage ofT. ni males were also attracted toA. gamma females and 11% performed the whole courtship sequence.Contribution from the Agricultural Research Organization (ARO), No. 3459-E, 1992 series.     

Use of a comparative approach to identify allelopathic potential and relationship between allelopathy bioassays and “competition” experiments for ten grassland and plant species

Various allelopathy bioassays were used to evaluate the allelopathic potential of 10 grassland forage species against a common test (phytometer) species,Carduus nutans L. Aqueous extracts did not influenceC. nutans germination, although radicle elongation was often severely inhibited.C. nutans was strongly affected by shoot, but not root, leachates. Decomposing ground tissue had mixed effects, and often stimulated shoot production ofC. nutans. Calculation ofR ² (coefficient of determination) values between these results, and the results of previous experiments investigating the effects of the same 10 species onC. nutans emergence and development in field plots and glasshouse competition experiments frequently revealed strong, statistically significant relationships. Our results therefore provide correlative evidence for the importance of allelopathy in field conditions.     

Comparative study of allelopathy as exhibited byProsopis Juliflora swartz andProsopis cineraria (L) druce

The allelopathic effects ofProsopis juliflora were studied both in the laboratory and in nature and compared with that ofProsopis cineraria to understand the chemical nature of allelochemics. Both species occupy the same habitats butP. cineraria does not appear to have any toxic effect on other plants under its canopy.P. juliflora is highly allelopathic and does not allow the growth of any other species. Leaf extracts and leaf leachates ofP. juliflora were inhibitory. Decaying leaves were also inhibitory at early stages of decomposition. Live roots were not found to be inhibitory in cogermination and interplanting of seeds. Chemical investigation of the extracts showed the allelopathic compounds to be phenolic in nature in both the species. Slow decomposition and heavy accumulation of leaf litter belowP. juliflora may possibly result in accumulation of toxic substances in soil layers, inhibiting growth of other species.     

Allelopathic interactions among algae

Five species of green algae isolated from the Cleveland County, Oklahoma, area and a commercially obtained yellow-green alga were tested for allelopathic interactions. Axenic cultures ofPandorina morum Bory were stimulated in three sterile filtrates from nonaxenic cultures ofScenedesmus incrassatulus var.mononae G.M. Smith, with the oldest filtrate showing the least stimulation. By measuring the growth of axenic cultures inoculated into sterile filtrates of old cultures, instances of inhibition and stimulation were recorded in three screening experiments.Cosmarium vexatum West filtrate was inhibitory to the five other species, a characteristic probably important in its role of producing waterblooms in ponds and swamps. The allelopathic effects of filtrates of the five other species onBolrydium becherianum Vischer could be a key to its restriction to terrestrial sites. Instances of heat lability of the active substances were noted. These interactions, both allelopathic and stimulatory, probably play a significant role in the succession of algal blooms in nature.     

Shifts in Life-History Traits as a Response to Cannibalism in Larval Long-Toed Salamanders (Ambystoma macrodactylum)

We examined the potential influence of cannibalism on life-history characteristics of larval long-toed salamanders (Ambystoma macrodactylum). Using a 2 × 2 factorial design, crossing morphology with diet, we exposed typical morph larvae to one of four types of stimulus animals: cannibal morphs fed a conspecific diet, cannibal morphs fed a heterospecific diet (i.e., Tubifex), typical morphs fed a conspecific diet, and typical morphs fed a heterospecific diet. Test larvae exposed to stimulus animals fed a conspecific diet exhibited a slower growth rate and an increase in the time taken to reach metamorphosis. These changes in life history likely represent a cost of antipredator behavior.     

Responses of Cambarid Crayfish to Predator Odor

The responses of individuals of four sympatric species of cambarid crayfish to the introduction of the odor of a common predator, the snapping turtle Chelydra serpentina, were recorded in the laboratory. Adult Orconectes virilis spent significantly more time in a lowered posture and reduced the frequency of nonlocomotory movements following introduction of snapping turtle odor but showed no change in behavior upon introduction of the odor of painted turtle (Chrysemys picta). Recently released young O. virilis did not respond to snapping turtle odor initially but did so after turtle odor and conspecific alarm odor had been paired. Individuals of O. propinquus did not respond to snapping turtle odor. Initial tests with O. rusticus did not yield any response to snapping turtle odor but after experience with paired turtle and alarm odor, individuals showed a decrease in nonlocomotory movements when just snapping turtle odor was introduced. Individuals of Cambarus robustus spent less time in the lowered posture, less time in their burrow, and more frequently executed nonlocomotory movements, in response to snapping turtle odor. The differences in responses to the odor of a common predator are correlated with ecological differences among the crayfish species.     

Attraction and inhibition in moth species responding to sex-attractant lures containing Z-11-hexadecen-1-yl acetate

Sex attractants for several sympatric noctuid moths required Z-11-hexadecen-1-yl acetate and additional olefinic compounds (coattractants) for effective, species-specific operation. In nearly all cases at least one of the coattractant compounds for each species functioned as a strong inhibitor of one or more of the other species in the group. It was concluded that species specificity in sex attractants can be achieved through conspecific coattractants which are at the same time transspecific inhibitors.Issued as NRCC No. 16150.     

Chemically mediated behavior in Acari: Adapations for finding hosts and mates

Ticks and mites respond to a limited spectrum of stimuli in their search for hosts and mates. Airborne chemical signals include carbon dioxide, ammonia, organic acids, terpenoids, 2,6-dichlorophenol, and other phenolic compounds. These are detected primarily by sensilla in and adjacent to Haller's organ. Most ixodid species examined have one or more multiporose sensilla that detect such volatiles. These olfactoreceptors enable the ticks to respond to remote volatile chemicals from hosts and from the other ticks, e.g., sex pheromones. Other sensilla, probably mechanogustatory in function, also occur on the tarsi. Gustatory sensilla on the palps detect assembly pheromones that enable ticks and mites to respond to conspecific or heterospecific chemical stimuli in their environment. Responses to those stimuli in ticks result in clustering, i.e., arrestant behavior. Arrestant behavior also occurs in certain mites. Finally, cheliceral chemosensilla enable ticks to recognize specific phagostimulants in host blood, e.g., ATP and glutathione, that stimulate feeding. InDennacentor variabilis andD. andersoni, these same cheliceral chemosensilla recognize species-specific genital sex pheromones in the vulvae of conspecific mates, without which they do not copulate.     

Rye residues contribute weed suppression in no-tillage cropping systems

The use of allelopathic cover crops in reduced tillage cropping systems may provide an ecologically sound and environmentally safe management strategy for weed control. Growers often plant winter rye (Secale cereale L.) for increased soil organic matter and soil protection. Spring-planted living rye reduced weed biomass by 93% over plots without rye. Residues of fall-planted/spring-killed rye reduced total weed biomass over bare-ground controls. Rye residues also reduced total weed biomass by 63% when poplar excelsior was used as a control for the mulch effect, suggesting that allelopathy, in addition to the physical effects of the mulch, did contribute to weed control in these systems. In greenhouse studies, rye root leachates reduced tomato dry weight by 25–30%, which is additional evidence that rye is allelopathic to other plant species.Journal Article No. 10712 of the Michigan Agricultural Experiment Station. Supported by United States Department of Agriculture Competition Grant No. 78-59-2261-0-1-060-1.     

Alarm response by a plethodontid salamander (Desmognathus ochrophaeus): Conspecific and heterospecific “Schreckstoff”

The detection of chemical alarm cues plays an important role for predator avoidance in many taxonomic groups, but little is known about the presence of such chemical cues in adult or caudate amphibians. We investigated the response (i.e., aversion or nonaversion) to chemical cues from damaged salamander skin and mealworms (Tenebrio molitor) in the plethodontid salamander,Desmognathus ochrophaeus. Avoidance responses were demonstrated to skin extracts of both conspecific and heterospecific salamanders. However, salamanders (D. ochrophaeus) did not avoid heated conspecific skin, fresh conspecific viscera, fresh mealworm, or freshPlethodon richmondi skin extracts. These results indicate that chemical alarm cues are: (1) present in the skin ofDesmognathus salamanders, (2) not present in mealworm or the viscera ofDesmognathus salamanders, and (3) denatured or deactivated by heating. These results also suggest that an avoidance response to chemical cues from damaged conspecifics has adaptive value in predator avoidance in terrestrial as well as aquatic vertebrates.