Despite long-standing interest in latitudinal variation in ecological patterns and processes, there is to date weak and conflicting evidence that herbivore pressure varies with latitude. We used three approaches to examine latitudinal variation in herbivore pressure in Atlantic Coast salt marshes, focusing on five abundant plant taxa: the grass Spartina alterniflora, the congeneric rushes Juncus gerardii and J. roemerianus, the forb Solidago sempervirens, and the shrubs Iva frutescens and Baccharis halimifolia. Herbivore counts indicated that chewing and gall-making herbivores were typically > or = 10 times more abundant at low-latitude sites than at high-latitude sites, but sucking herbivores did not show a clear pattern. For two herbivore taxa (snails and tettigoniid grasshoppers), correctly interpreting latitudinal patterns required an understanding of the feeding ecology of the species, because the species common at high latitudes did not feed heavily on plant leaves whereas the related species common at low latitudes did. Damage to plants from chewing herbivores was 2-10 times greater at low-latitude sites than at high-latitude sites. Damage to transplanted "phytometer" plants was 100 times greater for plants transplanted to low- than to high-latitude sites, and two to three times greater for plants originating from high- vs. low-latitude sites. Taken together, these results provide compelling evidence that pressure from chewing and gall-making herbivores is greater at low vs. high latitudes in Atlantic Coast salt marshes. Sucking herbivores do not show this pattern and deserve greater study. Selective pressure due to greater herbivore damage at low latitudes is likely to partially explain documented patterns of low plant palatability to chewing herbivores and greater plant defenses at low latitudes, but other factors may also play a role in mediating these geographic patterns.
Volatile organic chemical (VOC) emission by plants may serve as an adaptive plant defense by attracting the natural enemies of herbivores. For plant VOC emission to evolve as an adaptive defense, plants must show genetic variability for the trait. To date, such variability has been investigated primarily in agricultural systems, yet relatively little is known about genetic variation in VOCs emitted by natural populations of native plants. Here, we investigate intraspecific variation in constitutive and herbivore-induced plant VOC emission using the native common milkweed plant (Asclepias syriaca) and its monarch caterpillar herbivore (Danaus plexippus) in complementary field and common garden greenhouse experiments. In addition, we used a common garden field experiment to gauge natural enemy attraction to milkweed VOCs induced by monarch damage. We found evidence of genetic variation in the total constitutive and induced concentrations of VOCs and the composition of VOC blends emitted by milkweed plants. However, all milkweed genotypes responded similarly to induction by monarchs in terms of their relative change in VOC concentration and blend. Natural enemies attacked decoy caterpillars more frequently on damaged than on undamaged milkweed, and natural enemy visitation was associated with higher total VOC concentrations and with VOC blend. Thus, we present evidence that induced VOCs emitted by milkweed may function as a defense against herbivores. However, plant genotypes were equally attractive to natural enemies. Although milkweed genotypes diverge phenotypically in their VOC concentrations and blends, they converge into similar phenotypes with regard to magnitude of induction and enemy attraction.
Although grasshoppers are common salt marsh herbivores, we know little about geographic variation in their species composition. We documented latitudinal variation in species composition of the tettigoniid grasshopper fauna of Atlantic Coast salt marshes. Tettigoniids (N=740 adults) were collected from the Spartina alterniflora zone of 31 salt marsh sites across a latitudinal range of 13.19°(Florida to Maine), with an additional 52 individuals collected from the Juncus roemerianus zone of low-latitude marshes for comparative purposes. Eight species were collected, but some were common only at a few sites or rare throughout the entire collection range. The tettigoniid community was dominated by Orchelimum fidicinium at low latitudes and Conocephalus spartinae at high latitudes. Several factors might explain this shift, including changes in climate, plant phenology, and plant zonation patterns. O. fidicinium and C. spartinae increased in body size toward low latitudes. In laboratory feeding assays, O. fidicinium readily ate S. alterniflora and J. roemerianus leaves, Orchelimum concinnum, which is largely restricted to the J. roemerianus zone, ate only J. roemerianus leaves, and Conocephalus spp. ate neither, consistent with literature suggestions that they mainly consume seeds and flowers.Geographic variation in species composition and body size of grasshoppers may help explain documented patterns of geographic variation in plant palatability and plant-herbivore interactions in Atlantic Coast salt marshes. Because it can be difficult to identify tettigoniids to species, we present a guide to aid future workers in identifying the tettigoniid species common in these marshes.
The existence of predictable latitudinal variation in plant defense against herbivores remains controversial. A prevailing view holds that higher levels of plant defense evolve at low latitudes compared to high latitudes as an adaptive plant response to higher herbivore pressure on low-latitude plants. To date, this prediction has not been examined with respect to volatile organic compounds (VOCs) that many plants emit, often thus attracting the natural enemies of herbivores. Here, we compared genetically-based constitutive and herbivore-induced aboveground vegetative VOC emissions from plants originating across a gradient of more than 10° of latitude (>1,500 km). We collected headspace VOCs from Asclepias syriaca (common milkweed) originating from 20 populations across its natural range and grown in a common garden near the range center. Feeding by specialist Danaus plexippus (monarch) larvae induced VOCs, and field environmental conditions (temperature, light, and humidity) also influenced emissions. Monarch damage increased plant VOC concentrations and altered VOC blends. We found that genetically-based induced VOC emissions varied with the latitude of plant population origin, although the pattern followed the reverse of that predicted-induced VOC concentration increased with increasing latitude. This pattern appeared to be driven by a greater induction of sesquiterpenoids at higher latitudes. In contrast, constitutive VOC emission did not vary systematically with latitude, and the induction of green leafy volatiles declined with latitude. Our results do not support the prevailing view that plant defense is greater at lower than at higher latitudes. That the pattern holds only for herbivore-induced VOC emission, and not constitutive emission, suggests that latitudinal variation in VOCs is not a simple adaptive response to climatic factors.
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