Laboratory and field evaluation of predator odors as repellents for kiore (Rattus exulans) and ship rats (R. rattus)

Predator odors may serve to stop rats from entering conservation areas or to decrease predation, food consumption, and other damage by rats in areas tainted with predator odor. We compared the efficacy of real predator odors and synthetic odors (derived from the urine and feces of carnivores) as rat repellents with real herbivore odors as controls in a Y maze. We tested six predator odors: cat (Felis catus) urine and feces, mongoose (Herpestes auropunctatus) feces, n-propylthietane, S-methyl, methyl butanol, and isopentyl-methyl sulphide. The herbivore odors we used were: red deer (Cervus elaphus) urine, guinea pig (Cavia porcellus) feces, and white rabbit (Oryctolagus cuniculus) urine. Ship rats (Rattus rattus) and kiore or Polynesian rats (R. exulans) showed no aversion to any of the six predator odors when compared with herbivore odors. Ship rats, however, may have avoided synthesized odors more than real ones. We applied two odors (S-methyl, methyl butanol and n-propylthietane) to purpose-built feeders in native forest but recorded no change in either visitation rate or duration of visits for rodents [rats and mice (Mus musculus)] or possums (Trichosurus vulpecula). The consumption of maize at feeders was correlated with the number and duration of possum visits, but only weakly correlated with the number of visits by rodents. Consumption of maize was unaffected by the odor associated with the feeder. It is unlikely that the odors we tested will be useful in deterring rodents or possums from areas where they have been removed for economic, public health or conservation reasons.     

Responses of Wild Norway Rats (Rattus norvegicus) to Predator Odors

If rats could be shown to avoid the odors of predators, then conservation managers could manipulate this behavior to exclude rats from important conservation sites. We evaluated the ability of six predator odors to elicit avoidance responses by wild-caught Norway rats (Rattus norvegicus) from two New Zealand populations (Kapiti Island and North Island). Kapiti Island is free of mammalian predators, while the North Island has established populations of felids, canids, and mustelids. Three of the predator odors were real and three were synthesized volatile ingredients of real animal feces or urine. We compared the rats' responses to predator odors with their responses to three natural herbivore odors. We used a Y maze, and rats were offered the choice of an odor in one arm of the maze and distilled water in the other arm. Each choice arm was ducted at the entrance to remove air and the odor. We recorded: (1) the time until the rat left the first arm of the maze, (2) the time until the rat visited each arm, (3) the number of visits to each arm, (4) the amount of time spent in each arm, and (5) a total activity score for each rat. Kapiti rats showed an aversion to five of the six predator odors, despite never having encountered them before. Kapiti rats visited herbivore odors more often than carnivore odors and were less active in the presence of carnivore odors than they were when tested with herbivore odors. In addition, Kapiti rats approached some herbivore odors more quickly than they approached carnivore odors. North Island rats appeared to avoid cat urine, but despite being predator experienced, did not show a consistent aversion to the carnivore odors we tested. Both samples of rats, but particularly the North Island group, showed high individual variation. We conclude that predator odors are unlikely to be an effective management tool for all populations of this species of rat because of this high individual variability and the likelihood that each island population will differ in its responses to a given odor.     

Olfactory recognition of individuals by male cavies (Cavia aperea)

In one study, using a habituation procedure, male South American cavies,Cavia aperea, distinguished individual differences in odors collected from three sources: perineal gland secretions, urine, and supracaudal gland secretions. In a second study, male cavies spent more noninvestigatory time and rested more on the side of a cage containing the odor of a familiar subordinate male as compared to the cage side containing the odor of a familiar dominant male. Since the odor source was a glass plate which had been left in the home cage of the donor males for three days, the actual odorous cue to which the test males responded is not known. These studies demonstrate that male cavies distinguish odors of individuals, that individual differences in odors are found in at least three different sources, and that cavies use chemical cues to distinguish between known individuals.     

Odor Preferences of Weanling and Mature Male and Female Pine Voles

Olfactory cues from conspecifics have strong influences on behavior. Both female and male juveniles are attracted to familiar odors while adults show preferences for odors of opposite-sex conspecifics. Females show an increased preference for male odors as they mature, but no study has examined whether change with age is seen in males. In this experiment, responses of weanling and mature male and female pine voles (Microtus pinetorum) to familiar odor cues from their family versus those of an unfamiliar, opposite-sex conspecific were examined. Voles were tested in a three-chambered cage with end chambers containing the stimuli. There was a significant difference in odor preference between weanling and mature females. Weanling females preferred odor cues from their families, and mature females preferred the odors of unfamiliar mature males. The pattern was similar in males but the effect was not as strong. The results support the hypothesis that responses to odor cues change with age in pine voles. At least for females, the change in response to odor cues may be involved in the timing of reproduction and thus dispersal.     

Use of odor baits in traps to test reactions to intra- and interspecific chemical cues in house mice living in outdoor enclosures

House mice (Mus musculus) living in outdoor enclosures were tested for urinary chemical cue preferences using odor-baited traps. In the first experiment, with only volatile cues available, odors from conspecific males and females of various age classes and reproductive conditions were tested; no preferences were exhibited. In the second experiment mice had both nonvolatile and volatile cues available from the same sources as in experiment I. All age and sex class and female reproductive condition groups exhibited odor cue preferences except juvenile females. There were no specific odor cue preferences exhibited by any of the responder types with regard to odors from juvenile females. In the third and fifth experiments, mice were presented with nonvolatile plus volatile or only volatile urine odor cues, respectively, from four genera,Mus, Peromyscus, Microtus, andHomo. Mice of all age classes and both sexes preferredMus musculus odor, were neutral towardMicrotus ochrogaster odor, and avoided odors fromPeromyscus leucopus andHomo sapiens; these patterns were the same regardless of whether only volatile or both volatile and nonvolatile cues were presented. The fourth and sixth experiments involved testing volatile cues only and volatile cues plus nonvolatile cues from human sweat or feces from dogs, cats, or shrews. Mice avoided the human sweat and feces from cats and shrews, but were neutral toward the odor of dog feces. There were effects on whether mice were trapped in the interior of the enclosure or on the perimeter for some odors tested in these six experiments. The findings provide insights regarding possible functions of odor cues in the behavioral ecology of house mice. Odor-baiting traps can be an effective tool with respect to testing some, but not all questions pertaining to olfactory cues and house mouse social biology.     

Phytopathogen Lures Its Insect Vector by Altering Host Plant Odor

Many phytopathogens that cause worldwide losses of agricultural yield are vectored by herbivorous insects. Limited information is available about the interactions among phytopathogens, host plants, and insect vectors. In this paper, we report that the cell wall-lacking bacterium Candidatus Phytoplasma mali can alter both the odor of its host plant (apple) and behavior of its vector, the univoltine psyllid Cacopsylla picta. Apple trees infected by this phytoplasma emitted higher amounts of beta-caryophyllene when compared to uninfected ones. Psyllids that had no previous contact with Ca. P. mali, as well as infected pyllids, are more attracted by volatiles emitted from phytoplasma-infected apple plants than from uninfected ones. Psyllids that had developed on infected plants without getting infected showed the opposite behavior. These results suggest that the pathogen modifies host plant odor that lures its vector to infected plants. This may result in higher numbers of transmitting vector insects within the population.     

How contact foraging experiences affect preferences for host-related odors in the larval parasitoidCotesia marginiventris (Cresson) (Hymenoptera: Braconidae)

Responses of individual females of the parasitoidCotesia marginiventris to the odors of four different complexes of host larvae feeding on leaves were observed in a four-arm olfactometer. The plant-host complexes were composed of fall armyworm (FAW) larvae or cabbage looper (CL) larvae feeding on either corn or cotton seedlings. Prior to testing, each female was given a brief foraging experience on a plant-host complex and was then exposed to the odors of the same complex in the olfactometer. The experienced females responded to familiar odors in a dose-related manner, and these responses were virtually identical to all four complexes. Preferences for the odors of one of two plant-host complexes were tested in dual choice situations. Generally, FAW odors were preferred over CL odors and corn odors over cotton odors. A short foraging experience significantly affected the females' odor preferences in favor of the odors released by the experienced complex. Additional experiments revealed that neither longer bouts of experience nor bouts that included ovipositions resulted in a stronger change in preference. Experience affected preference in combinations where only the host species was varied as well as in combinations where only the plant species was varied. The results, therefore, strongly indicate that both the plants and the hosts somehow are involved in the production and/or release of the semiochemicals that attractC. marginiventris.Mention of a proprietary product does not constitute an endorsement or the recommendation for its use by the USDA.     

Unraveling the Nature of Individual Recognition by Odor in Hermit Crabs

Individual recognition is a key element in the social life of many invertebrates. However, most studies conducted so far document that several species are capable of a “binary” discrimination among conspecifics, but not of a “true individual recognition.” Our objective was to learn more about the mechanisms that underlie individual recognition by odor in hermit crabs by individuating some of its properties. Using Pagurus longicarpus Say 1817 as a model species, we conducted four series of experiments in which the response of every test crab (the “receiver”) to the different odor treatments (emitted by a “sender”) was evaluated from its investigative behavior toward an empty, high-quality shell. After having excluded the possibility that crabs chemically recognize familiar/unfamiliar shells and/or shells of high/low quality, we explored whether the receivers discriminate odors from two familiar senders and whether this discrimination also occurs with unfamiliar crabs. We also asked whether crabs form an association between the odor of a familiar sender and some of its relevant attributes, i.e., rank, size, and shell quality. Finally, the shells inhabited by familiar individuals were manipulated to modify the association between odor and shell quality. Results showed that: (1) there is no odor specific of a rank; (2) individual crabs discriminate their own odor from the odor of other individuals; (3) they can chemically discriminate between larger crabs inhabiting higher-quality shells and smaller crabs inhabiting lower-quality shells, provided that these crabs are familiar to them; (4) they associate the odor of an individual crab with the quality of the shell it inhabits; and (5) this association quickly changes when social partners switch to shells of different quality. These results indicate that the nature of chemical recognition in P. longicarpus is more refined than a simple binary system. The receiver appears able to associate a type of information from the sender with memories of past experiences, therefore suggesting the hermit crab's potential for relatively high-order knowledge about conspecifics.     

An evaluation of repellents to suppress browsing by possums

The efficacy in feeding suppression trials with possums (Trichosorus vulpecula Kerr) of six synthetic odorous chemicals uniquely associated with various of the Mustelidae species and of the red fox (Vulpes vulpes) has been evaluated with respect to several proprietary repellents. The predator odors, two proprietary repellents, and two other repellent formulations (olive oil in petroleum jelly and a synthetic fermented egg odor cocktail) were applied as 5% w/v solutions in paraffin to the foliage ofPinus radiata seedlings and the extent of browse compared with that on seedlings treated with Treepel, an effective egg-based repellent already available commercially. All four of the mustelid predator odors [()-2-n-propylthietane, ()-3-n-propyl-1,2-dithiolane, 2,2-dimethylthietane, and 3,3-dimethyl-1,2-dithiolane] and the volatile compound from fox urine (³-isopentenyl methyl sulfide) were shown to be more or at least equally effective compared to Treepel. This was true also of one commercial repellent Gori, which contained bitrex and the synthetic fermented egg formulation. The most effective compounds will be evaluated when administered with adhesives and surfactants in the further development of possum repellents with longer field lives.     

Scent lures from anal sac secretions of the ferretMustela furo L.

A bioassay of eight synthetic anal sac compounds showed that ferrets were most attracted to a mixture of 2-propylthietane andtrans- andcis-2,3-dimethylthietane. This mixture was used as the basis of an artificial scent lure. Trapping experiments used two traps at each site. In comparing trap success, ferrets chose artificial lure in preference to no odor, and chose both food baits and natural-product anal sac odor over artificial lure. However, the effectiveness of artificial lure was demonstrated by comparison between trap sites with bait-lure pairs and those with lure only. The comparison indicated that scent lures should be as effective as bait in attracting ferrets, and this was confirmed in a field program. We conclude that scent lures are valuable additions to the current techniques used in mustelid control operations, but more work is needed to increase the inherent species specificity of scent lures for mustelids and to develop a simple means of making a lure that will release odor for a long time.     

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.     

Trirhabda canadensis (Coleoptera: Chrysomelidae) responses to plant odors

The responses of the goldenrod leaf beetleTrirhabda canadensis to host and nonhost volatile odors were tested in a Y-tube olfactometer in the laboratory. Beetles preferred host to nonhost odors and were sensitive to concentrations of host odor. Beetles distinguished between host and nonhost volatiles of only one of the two nonhostSolidago species; host volatiles were preferred to all nonhost volatiles at the family and order levels. In other words, all nonhosts above the genus level had similar effects on beetle responses. Although the odors of most nonhosts were neutral (i.e., neither attractive nor repellent) to the beetles as tested against air, this neutrality disappeared if the odors of two or more nonhosts were added to the host odor and beetles were given a choice between this mixture and pure host odor. Given this choice, they strongly preferred pure host odor, which suggests that diversity of odors per se is unattractive to the beetles. Beetles walked rather than flew to locate their hosts in the field, and their movements suggest that they used olfactory cues to locate hosts.     

Odors in traps: Does most recent occupant influence capture rates for house mice?

Responses of house mice (Mus domesticus) to odors in live traps were studied in a series of eight 0.1-ha outdoor field enclosures. It was assumed that the most recent mouse capture would provide the predominant odor in a trap for at least one week. Three different populations were tested, one in 1989 and two in 1992, involving over 800 different mice. Similar response patterns were recorded from all three groups. Two types of questions were tested: (1) Were there any biases contingent upon what had been previously caught? (2) Were there consistent responses of mice of particular age, sex, or reproductive classes to trap odors? Traps soiled by juvenile females caught adult females significantly less often than expected, but there were no consistent relationships in terms of the effects of specific residual odors on the subsequent capture at a particular trap. For various age, sex, and reproductive classes, (1) adult males preferred odors from juvenile and estrous females and avoided odors of other males significantly more than expected, (2) juvenile females selected traps with odors of other juvenile females and avoided all other types of female odors significantly more than expected, (3) nonestrous females exhibited a significant preference for adult male odor, and (4) estrous females selected traps containing odors from adult males but avoided those that had previously contained either nonestrous or pregnant/lactating females significantly more than expected. These findings have potential implications with regard to both the methods used for trapping small rodents and the social biology of house mice.     

The Role of Fresh <Emphasis Type="Italic">versus</Emphasis> Old Leaf Damage in the Attraction of Parasitic Wasps to Herbivore-Induced Maize Volatiles

The odor produced by a plant under herbivore attack is often used by parasitic wasps to locate hosts. Any type of surface damage commonly causes plant leaves to release so-called green leaf volatiles, whereas blends of inducible compounds are more specific for herbivore attack and can vary considerably among plant genotypes. We compared the responses of naïve and experienced parasitoids of the species Cotesia marginiventris and Microplitis rufiventris to volatiles from maize leaves with fresh damage (mainly green leaf volatiles) vs. old damage (mainly terpenoids) in a six-arm olfactometer. These braconid wasps are both solitary endoparasitoids of lepidopteran larvae, but differ in geographical origin and host range. In choice experiments with odor blends from maize plants with fresh damage vs. blends from plants with old damage, inexperienced C. marginiventris showed a preference for the volatiles from freshly damaged leaves. No such preference was observed for inexperienced M. rufiventris. After an oviposition experience in hosts feeding on maize plants, C. marginiventris females were more attracted by a mixture of volatiles from fresh and old damage. Apparently, C. marginiventris has an innate preference for the odor of freshly damaged leaves, and this preference shifts in favor of a blend containing a mixture of green leaf volatiles plus terpenoids, after experiencing the latter blend in association with hosts. M. rufiventris responded poorly after experience and preferred fresh damage odors. Possibly, after associative learning, this species uses cues that are more directly related with the host presence, such as volatiles from host feces, which were not present in the odor sources offered in the olfactometer. The results demonstrate the complexity of the use of plant volatiles by parasitoids and show that different parasitoid species have evolved different strategies to exploit these signals.     

The effect of short-range host odor stimuli on host fruit finding and feeding behavior of plum curculio adults (Coleoptera: Curculionidae)

In laboratory assays, we investigated responses of female plum curculios (PCs),Conotrachelus nenuphar (Herbst), to host and nonhost fruit or leaf odor when PCs were crawling on experimental tree branchlets or twigs. In choice tests where test specimens were hung from the ends of a wooden crosspiece, PCs made significantly more visits to host plum fruit than to plum leaves, nonhost tomato fruit, wax models of plum fruit, or blanks (wire). In similar tests, PCs made significantly more visits to plum leaves compared to nonhost maple leaves or to blanks. PCs in test chambers that contained host or nonhost odor were significantly more prone to feed on wax plum models in the presence of odor from host fruit or host leaves compared to odor from nonhost fruit or leaves or a water blank. In choice tests offering alternating cluster types on an apple branchlet, PCs visited leaf clusters bearing a host apple fruit more than leaf clusters without a fruit. In tests to assay the distance at which PCs can detect an individual host fruit, PCs crawled from the central stem of an apple branchlet onto a side stem significantly more often when an apple fruit on a side stem was hung 2 cm from the central stem compared to 4 or 8 cm away. Our combined results suggest that PCs use host fruit odor to locate host fruit at close range.     

Ability of honeybee,Apis mellifera,to detect and discriminate odors of varieties of canola (Brassica rapa and Brassica napus) and snapdragon flowers (Antirrhinum majus)

Honeybees (Apis mellifera) use odors to identify and discriminate among flowers during foraging. This series of experiments examined the ability of bees to detect and discriminate among the floral odors of different varieties of two species of canola (Brassica rapa and Brassica napus) and also among three varieties of snapdragons (Antirhinnum majus). Individual worker honeybees were trained using a proboscis extension assay. The ability of bees to distinguish a floral odor from an air stimulus during training increased as the number of flowers used during training increased. Bees conditioned to the odor of one variety of flower were asked to discriminate it from the odors of other flowers in two different training assays. Bees were unable to discriminate among flowers at the level of variety in a randomized presentation of a reinforced floral odor and an unreinforced floral odor. In the second type of assay, bees were trained with one floral variety for 40 trials without reinforcement and then tested with the same variety or with other varieties and species. If a bee had been trained with a variety of canola, it was unable to differentiate the odor of one canola flower from the odor of other canola flowers, but it could differentiate canola from the odor of a snapdragon flower. Bees trained with the odor of snapdragon flowers readily differentiated the odor of one variety of a snapdragon from the odor of other varieties of snapdragons and also canola flowers. Our study suggests that both intensity and odor quality affect the ability of honeybees to differentiate among floral perfumes.     

Plant Volatiles Mediate Host-Finding Behavior of the Apple Leafcurling Midge

The behavioral responses of apple leafcurling midge,Dasineura mali Kieffer, mated females to foliage from host and nonhost trees were investigated in a wind tunnel. When released downwind of apple or pear foliage, females exposed to apple were more likely to exhibit upwind flight and to approach and land on foliage. On apple foliage, landings were concentrated on buds and immature leaves. Probability of taking flight and latency to flight did not differ for females exposed to apple and pear. When foliage was placed behind screens to obscure plant visual stimuli, females again distinguished between apple and pear, with more of the females exposed to apple odors flying upwind and landing. Females exposed to pear odors were more likely to fly upwind, approach, and land than females exposed to clean air. Odors from immature apple foliage triggered orientation responses in a larger percentage of females than odors from mature apple foliage. A dichloromethane extract of immature apple foliage also triggered orientation responses.     

Associative Learning of Food Odor by Social Wasps in a Natural Ecosystem

The ability of insects to associate olfactory cues with food from their environment has been well documented with various insect orders. However, these studies were based on prior training of insects to associate odors with food sources in the laboratory or in the field with almost no evidence for the development of this phenomenon in natural ecosystems. In New Zealand’s ancient Fuscospora spp. or beech forests, invasive Vespula social wasps were attracted to odor from honeydew (benzaldehyde and n-octanol) but did not respond to a known wasp attractant (isobutanol and acetic acid). On the other hand, wasps in a rural/suburban area in New Zealand did not respond to honeydew odor but responded instead to the known wasp attractant. Similarly, social wasps in Hungary did not respond to honeydew odor, but responded to the known wasp attractant. DNA sequences of Vespula vulgaris from the two locations in New Zealand were 100% identical. Similarly, DNA sequences of V. germanica from the two locations in New Zealand were 100% identical, indicating little or no intra-specific variation. On the other hand, DNA sequences of V. vulgaris and V. germanica from New Zealand were 99.56 and 99.78% matches with V. vulgaris and V. germanica samples from Hungary, respectively. Electroantennogram (EAG) response profiles of wasps from the three locations to benzaldehyde, isobutanol, acetic acid, n-octanol and heptyl butyrate were similar. The high similarity in DNA sequences and EAG profiles, with only a behavioral difference in the response to odor sources, suggests that social wasps in New Zealand’s beech forest have naturally developed associative learning for the odor of the carbohydrate-rich honeydew. This is the first study to provide conclusive evidence of the occurrence and the development of associative learning of food odor by social insects in a natural ecosystem free of human interference. The ability of social wasps to naturally develop associative learning of food odor might in part explain why they are extremely successful invaders of new ecosystems, especially those with rich honeydew resources. The finding of our study will have significant implications for the use of attractants in wasp monitoring and control in different habitats.     

Alert odor from skin gland in deer

Black-tailed deer (Odocoileus hemionus columbianus) discharge an odor from the metatarsal (MT) gland, located on the hindleg, when disturbed or alarmed. Freely moving, captive deer were exposed to the MT odor by means of a remote-release apparatus. Responses by males and females to MT odor from both sexes were recorded with a coding system and a video camera. When the odor was present, females became more alert and left the site more often than in the presence of control odors, odorless air, or deer urine. It is concluded that the MT secretion provides an alert signal, placing the odor into the class of alarm pheromones. There is no evidence that the closely related white-tailed deer,O. virginianus, possesses this alert (or alarm) odor system to the same degree.     

Pulsed Odors from Maize or Spinach Elicit Orientation in European Corn Borer Neonate Larvae

Lepidoptera larvae are capable of orienting towards or away from plants by using odors as cues but whether this attraction is innate or secondarily acquired remains unknown. We tested the hypothesis that European corn borer (ECB) neonate larvae express an innate attraction towards odors released from maize, and avoidance towards odors from spinach. Neonate larvae were placed on a locomotion compensator within a constant stream of humidified air that was loaded intermittently with airborne odors drawn from potted plants. The odor stream was delivered continuously or pulsed (1 to 10 sec pulses) at 40 ml/min. ECB larvae oriented toward maize odors pulsed at 2 to 6 sec but walked away from maize odors delivered at lower frequencies (9 and 10 sec pulses or to continuous ones). They consistently walked away from spinach odors, irrespective of the pulsing regime except at 1 sec pulses that did not elicit orientation. We further explored odor intensity on orientation towards maize odors by adjusting the odor stream intensity. At higher intensity (60 ml/min), the direction reversal started at the 6 sec half period, while at lower intensity (20 ml/min), it showed up only for the continuous stimulus. ECB larvae exhibit a striking ability to lock on to a direction, which they maintained despite gaps of up to 10 sec in the odor stream. Our results demonstrate that ECB neonate larvae express innate orientation preferences towards natural odors from plants. These reactions correlate well with the biological value of these plants for ECB: maize generally is accepted by ECB larvae and adults, while spinach represents a poor host because it produces (non-volatile) phytoecdysteroids that are toxic and deterrent.