Characterizing the interaction between the bogus yucca moth and yuccas: do bogus yucca moths impact yucca reproductive success?

Yucca moths are most well known for their obligate pollination mutualism with yuccas, where pollinator moths provide yuccas with pollen and, in exchange, the moth larvae feed on a subset of the developing yucca seeds. The pollinators, however, comprise only two of the three genera of yucca moths. Members of the third genus, Prodoxus, are the bogus yucca moths and are sister to the pollinator moths. Adult Prodoxus lack the specialized mouthparts used for pollination and the larvae feed on plant tissues other than seeds. Prodoxus larvae feed within the same plants as pollinator larvae and have the potential to influence yucca reproductive success directly by drawing resources away from flowers and fruit, or indirectly by modifying the costs of the mutualism with pollinators. We examined the interaction between the scape-feeding bogus yucca moth, Prodoxus decipiens, and one of its yucca hosts, Yucca filamentosa, by comparing female reproductive success of plants with and without moth larvae. We determined reproductive success by measuring a set of common reproductive traits such as flowering characteristics, seed set, and seed germination. In addition, we also quantified the percent total nitrogen in the seeds to determine whether the presence of larvae could potentially reduce seed quality. Flowering characteristics, seed set, and seed germination were not significantly different between plants with and without bogus yucca moth larvae. In contrast, the percent total nitrogen content of seeds was significantly lower in plants with P. decipiens larvae, and nitrogen content was negatively correlated with the number of larvae feeding within the inflorescence scape. Surveys of percent total nitrogen at three time periods during the flowering and fruiting of Y. filamentosa also showed that larval feeding decreased the amount of nitrogen in fruit tissue. Taken together, the results suggest that although P. decipiens influences nitrogen distribution in Y. filamentosa, this physiological effect does not appear to impact the female components of reproductive success.     

Cheating in mutualism: defection of Yucca baccata against its yucca moths

Yucca baccata cheats in its obligate pollination/seed predation mutualism with yucca moths. Although all individuals use the pollination services of yucca moths, many individuals do not reciprocate in sustaining yucca moth larvae. Cheating is associated with the morphology of Y. baccata pistils. In Y. baccata , the apex of the ovary contains only inviable ovules, and there are two distinct flower types, one of which has twice as many potentially viable ovules as the other. Because yucca moths oviposit at the apex of Y. baccata ovaries, larvae in flowers with few viable ovules fail to encounter viable ovules and therefore perish. Inflorescences generally have just one flower type, implying that some individuals cheat whereas others maintain the yucca moth population. Our most surprising observation, however, is that although the proportion of cheaters should be low, over 70% of Y. baccata individuals cheat. We hypothesize that both density- and frequency-dependent processes maintain a balance of cheaters and noncheaters in this system.     

Chemistry and geographic variation of floral scent in Yucca filamentosa (Agavaceae)

We identified volatiles from the floral headspace of Yucca filamentosa using gas chromatography and mass spectrometry and analyzed floral scent composition and variation among populations pollinated by different yucca moth species. Twenty-one scent compounds were repeatedly identified and most could be categorized into two major classes: (1) homoterpenes derived from the sesquiterpene alcohol nerolidol and (2) long chain aliphatic hydrocarbons. Two biosynthetic pathways are thus responsible for the majority of floral volatiles in Y. filamentosa. The homoterpene E-4,8-dimethylnona-1,3,7-triene, which is released systemically by higher plants upon herbivory, was the most abundant compound. Two di-oxygenated compounds not previously reported as floral compounds also were detected. No differentiation in floral scent was observed between populations pollinated by different yucca moths, nor was there any correlation between chemical distance and geographic distance among populations. The total release rate of volatiles differed significantly among populations, but not between populations with different pollinators. The combination of unique compounds and low variation in the fragrance blend may reflect highly selective attraction of obligate pollinators to flowers. The observed lack of differentiation in floral scent can putatively explain high moth-mediated gene flow among sites, but it does not explain conservation of odor composition across populations with different pollinators.     

Oviposition and pollination behavior of the yucca moth,Tegeticula maculata (Lepidoptera: Prodoxidae), and its relation to the reproductive biology of Yucca whipplei (Agavaceae)

Summary The adult behavior of the yucca moth, Tegeticula maculata Riley, is finely tuned to the reproductive biology of its specific host plant, Yucca whipplei Torr. The female moths oviposit in the ovaries of the yucca flowers and actively pollinate the same flowers with pollen which they have collected previously. The selective pressures imposed on the moths by 1) the plant's need for pollen transfer via an insect pollinating agent, 2) its partial self-incompatibility, and 3) its ability to regulate seed set by aborting excess fruits, have molded the pollinator's behavior in such a way that its offspring have the greatest possible chance of surviving through the early larval stages. The evolutionary responses of the pollinator include the following: 1) the female moths consistently fly to a different plant after collecting pollen, thus insuring cross-fertilization of the flowers, 2) they always pollinate after depositing the first egg in a flower, but not necessarily after subsequent ovipositions, and 3) females emerging near the end of the flowering season frequently oviposit in developing seed pods, as opposed to open flowers which are more likely to be aborted by the plants.     

Pollination ecology of Yucca elata

The pollination biology of a population of 250 Yucca elata (Liliaceae) plants was studied in southern New Mexico. Yucca elata and the prodoxid yucca moth Tegeticula yuccasella have a mutualistic association that is essential for the successful sexual reproduction of both species. However, a wide range of other invertebrate species visit flowers during the day and at night. Our aim was to quantify the role of yucca moths and other invertebrate visitors in pollination and fruit set, using manipulative field experiments. Inflorescences were bagged during the day or night (N=12 inflorescences) to restrict flower visitors to either nocturnal or diurnal groups. Yucca moths were active exclusively nocturnally during the flowering period and thus did not visit inflorescences that were unbagged during the day. None of the 4022 flowers exposed only to diurnal visitors set fruit, whereas 4.6% of the 4974 flowers exposed only to nocturnal visitors (including yucca moths) produced mature fruit. The proportion of flowers producing fruit in the latter treatment was not significantly different from unbagged control inflorescences. In a series of experimental manipulations we also determined that: (1) flowers opened at dusk and were open for two days on average, but were only receptive to pollen on the first night of opening; (2) pollen must be pushed down the stigmatic tube to affect pollination; and (3) most plants require out-cross pollination to produce fruit. The combination of these results strongly suggests that yucca moths are the only species affecting pollination in Y. elata, and that if another species was to affect pollination, it would be a rare event.     

Effect of pollinator-inflicted ovule damage on floral abscission in the yucca-yucca moth mutualism: the role of mechanical and chemical factors

The long-term persistence of obligate mutualisms (over 40 Mya in both fig/fig wasps and yucca/yucca moths) raises the question of how one species limits exploitation by the other species, even though there is selection pressure on individuals to maximize fitness. In the case of yuccas, moths serve as the plant's only pollinator, but eggs laid by the moths before pollination hatch into larvae that consume seeds. Previous studies have shown that flowers with high egg loads are more likely to abscise. This suggests that yucca flowers can select against moths that lay many eggs per flower through selective abscission of flowers; however, it is not known how yucca moths trigger floral abscission. We tested how the moth Tegeticula yuccasella triggers floral abscission during oviposition in Yucca filamentosa by examining the effects of ovipositor insertion and egg laying on ovule viability and floral abscission. Eggs are not laid at the site of ovipositor insertion: we used this separation to test whether wounded ovules were more closely associated with the ovipositor site or an egg's location. Using a tetrazolium stain to detect injured ovules, we determined whether the number of ovipositions affected the number of wounded ovules in naturally pollinated flowers. Two wounding experiments were used to test the effect of mechanical damage on the probability of floral abscission. The types of wounds in these experiments mimicked two types of oviposition-superficial oviposition in the ovary wall and oviposition into the locular cavity-that have been observed in species of Tegeticula. The effect of moth eggs on ovule viability was experimentally tested by culturing ovules in vitro, placing moth eggs on the ovules, and measuring changes in ovule viability with a tetrazolium stain. We found that ovules were physically wounded during natural oviposition. Ovules showed a visible wounding response in moth-pollinated flowers collected 7-12 h after oviposition. Exact location of wounded ovules relative to eggs and oviposition scars, as well as results from the artificial wounding experiments, showed that the moth ovipositor inflicts mechanical damage on the ovules. Significantly higher abscission rates were observed in artificially wounded flowers in which only 4-8% of the ovules were injured. Eggs did not affect ovule viability as measured by the tetrazolium stain. These results suggest that physical damage to ovules caused by ovipositing is sufficient to explain selective fruit abscission. Whether injury as a mechanism of selective abscission in yuccas is novel or a preadaptation will require further study.     

Facultative non-mutualistic behaviour by an “Obligate” mutualist: “Cheating” by Yucca moths

The interaction between yucca moths (Tegeticula spp., Incurvariidae) and yuccas (Yucca spp., Agavaceae) is an obligate pollination/seed predation mutualism in which adult female yucca moths pollinate yuccas, and yucca moth larvae feed on yucca seeds. In this paper we document that individual yucca moths, which are capable of acting as mutualists, facultatively cheat by ovipositing in yucca pistils without attempting to transfer pollen. Additionally, a high proportion of flowers are unlikely to receive pollen even when pollination is attempted, because many yucca moths carry little or no pollen. The probability of occurrence of non-mutualistic behaviour is not affected by the amount of pollen a moth carries: moths with full pollen loads are just as likely to act non-mutualistically as moths carrying little or no pollen. We propose four hypotheses that could explain facultative non-mutualistic behaviour in yucca moths.Present address: Department of Biology, Simon Fraser University, Burnaby, British Columbia V5A 1S6, Canada     

Patterns of genetic structure among populations of an oligophagous pollinating yucca moth (Tegeticula yuccasella)

Plant-insect associations have served as models for investigations of coevolution and the influence of biotic interactions on diversification. The pollination association between yuccas and yucca moths is a classic example of an obligate mutualism often suggested to have been affected by coevolution. Recent work has shown high host specificity in pollinating yucca moths, and here we use Tegeticula yuccasella, the species with the widest diet breadth, to ask how host specificity and isolation by distance contribute to specialization. Isolation by distance at a regional scale was observed in nucleotide variation within the mitochondrial gene cytochrome oxidase I (COI) (r =.294; P =.003). Host-related genetic structure (F(ct) = 0.08) was found to be slightly lower than the level of structure observed between eastern and western moth populations (F(ct) = 0.096). However, 56% of the COI haplotypes sampled from moths on Yucca filamentosa mapped to a host-specific clade in the haplotype network. Taken together, these results suggest that differentiation among T. yuccasella populations on alternative hosts is slight, but gene flow is influenced by both host association and geographic distance.     

Genetic consequences of specialization: yucca moth behavior and self-pollination in yuccas

Reciprocal specialization in interspecific interactions, such as plant-pollinator mutualisms, increases the probability that either party can have detrimental effects on the other without the interaction being dissolved. This should be particularly apparent in obligate mutualisms, such as those that exist between yucca and yucca moths. Female moths collect pollen from yucca flowers, oviposit into floral ovaries, and then pollinate those flowers. Yucca moths, which are the sole pollinators of yuccas, impose a cost in the form of seed consumption by the moth larvae. Here we ask whether there also is a genetic cost through selfish moth behavior that may lead to high levels of self fertilization in the yuccas. Historically, it has been assumed that females leave a plant immediately after collecting pollen, but few data are available. Observations of a member of the Tegeticula yuccasella complex on Yucca filamentosa revealed that females remained on the plant and oviposited in 66% of all instances after observed pollen collections, and 51% of all moths were observed to pollinate the same plant as well. Manual cross and self pollinations showed equal development and retention of fruits. Subsequent trials to assess inbreeding depression by measuring seed weight, germination date, growth rate, and plant mass at 5 months revealed significant negative effects on seed weight and germination frequency in selfed progeny arrays. Cumulative inbreeding depression was 0.475, i.e., fitness of selfed seeds was expected to be less than half that of outcrossed seeds. Single and multilocus estimates of outcrossing rates based on allozyme analyses of open-pollinated progeny arrays did not differ from 1.0. The discrepancy between high levels of behavioral self-pollination by the moths and nearly complete outcrossing in mature seeds can be explained through selective foreign pollen use by the females, or, more likely, pollen competition or selective abortion of self-pollinated flowers during early stages of fruit development. Thus, whenever the proportion of pollinated flowers exceeds the proportion that can be matured to ripe fruit based on resource availability, the potential detrimental genetic effects imposed through geitonogamous pollinations can be avoided in the plants. Because self-pollinated flowers have a lower probability of retention, selection should act on female moths to move among plants whenever moth density is high enough to trigger abortion. Received: 18 March 1996 \Accepted: 30 July 1996     

Genetic diversity and population structure of Yucca filamentosa (Agavaceae)

Using 19 allozyme loci we studied genetic diversity in 18 populations of Yucca filamentosa (Agavaceae) from the southeastern United States. Of the 19 loci surveyed, 17 (89.5%) were polymorphic in at least one of the populations sampled. There was considerable variation among populations in the percentage of polymorphic loci (range = 31.6-84.2%, mean = 67.6%). Similar heterogeneity among populations was observed for mean number of alleles per polymorphic locus (range = 2.0-3.0; mean = 2.48) and mean expected heterozygosity (range = 0.113-0.288; mean = 0.213). On average, 83% of the total genetic diversity was found within populations. Duplications of three allozyme loci were detected in several populations. The life-history characteristics of Y. filamentosa (a long-lived, semiwoody, predominantly outcrossing monocot with a large geographical range) may contribute to the maintenance of such high levels of genetic diversity. These results contradict expectations of the genetic structure of Y. filamentosa based on observations of the dispersal and pollination behavior of its sole pollinator, Tegeticula yuccasella, the yucca moth.     

Inbreeding and outcrossing in Yucca whipplei: consequences for the reproductive success of plant and pollinator

Unlike most pollinators, yucca moths are active pollinators of their host plants. Females lay their eggs in the flowers they pollinate, and their larvae feed solely on the resulting seeds. Previous evidence suggests that the yucca moth Tegeticula maculata avoids self-pollinating their host Yucca whipplei . Other yucca moths may self-pollinate more frequently. When pollinating, yucca moths are also reported to fly large distances between plants, bypassing neighbouring plants in the process. We experimentally verify the suggestion of Pellmyr et al . that yucca is more likely to retain fruits from self-pollination if overall fruit set is low. Thus, selection on moths to avoid self-pollinating should be density dependent. We found no evidence that mating with close neighbours resulted in inbreeding depression, thus the moth's long-distance flights between plants are yet to be explained.     

Variation in the costs and benefits of mutualism: the interaction between yuccas and yucca moths

Summary Yucca moths are both obligate pollinators and obligate seed predators of yuccas. I measured the costs and net benefits per fruit arising for eight species of yuccas from their interaction with the yucca moth Tegeticula yuccasella. Yucca moths decrease the production of viable seeds as a result of oviposition by adults and feeding by larvae. Oviposition through the ovary wall caused 2.3–28.6% of ovules per locule to fail to develop, leaving fruit with constrictions, and overall, 0.6–6.6% of ovules per fruit were lost to oviposition by yucca moths. Individual yucca moth larvae ate 18.0–43.6% of the ovules in a locule. However, because of the number of larvae per fruit and the proportion of viable seeds, yucca moth larvae consumed only 0.0–13.6% of potentially viable ovules per fruit. Given both oviposition and feeding effects, yucca moths decreased viable seed production by 0.6–19.5%. The ratio of costs to (gross) benefits varied from 0% to 30%, indicating that up to 30% of the benefits available to yuccas are subsequently lost to yucca moths. The costs are both lower and more variable than in a similar pollinator-seed predator mutualism involving figs and fig wasps.There were differences between species of yuccas in the costs of associating with yucca moths. Yuccas with baccate fruit experienced lower costs than species with capsular fruit. There were also differences in costs between populations within species and high variation in costs between fruit within populations. High variability was the result of no yucca moth larvae being present in over 50% of the fruit in some populations, while other fruit produced up to 24 larvae. I present hypotheses explaining both the absence and high numbers of larvae per fruit.     

RAD‐seq phylogenomics recovers a well‐resolved phylogeny of a rapid radiation of mutualistic and antagonistic yucca moths

The interaction between yucca plants and yucca moths has been one of the focal model systems investigated in the study of pollination mutualism and coevolution, especially in terms of understanding the prevention of overexploitation by mutualist partners. Yuccas have the ability to assess the number of eggs placed by pollinators into their ovaries, and can preferentially abort those flowers that would have many moth larvae consuming yucca seeds. Previous phylogenetic research identified a rapid radiation of moth species that corresponded with shifts in the interaction with their host plants. These shifts led to the evolution of moth species that circumvent the egg detection method used by yuccas to limit seed damage. In particular, some pollinator species deposit their eggs so that they are undetectable by the plants, whereas other species are ‘cheaters’ that have lost the ability to pollinate, yet deposit eggs into developing fruit rather than flowers. The evolution of these new species happened so quickly that the phylogeny of the moths has remained unresolved despite repeated attempts with standard Sanger sequencing of mtDNA loci and AFLP marker generation. Here, we use extensive analyses of RAD‐seq data to determine the phylogenetic relationships among yucca moth species. The results provide a robust phylogenetic framework that identifies the evolutionary relationships among shifts in egg‐laying strategies, as well as determining the closest pollinating relatives to the cheater species. Based on the obtained phylogeny, a shift in egg‐laying strategy that avoided the overexploitation regulatory mechanism used by yucca plants was a precursor for the evolution of two species with cheating behaviour.     

The evolution of obligate pollination mutualisms: senita cactus and senita moth

We report a new obligate pollination mutualism involving the senita cactus, Lophocereus schottii (Cactaceae, Pachyceereae), and the senita moth, Upiga virescens (Pyralidae, Glaphyriinae) in the Sonoran Desert and discuss the evolution of specialized pollination mutualisms. L. schottii is a night-blooming, self-incompatible columnar cactus. Beginning at sunset, its flowers are visited by U. virescens females, which collect pollen on specialized abdominal scales, actively deposit pollen on flower stigmas, and oviposit a single egg on a flower petal. Larvae spend 6 days eating ovules before exiting the fruit and pupating in a cactus branch. Hand-pollination and pollinator exclusion experiments at our study site near Bahia Kino, Sonora, Mexico, revealed that fruit set in L. schottii is likely to be resource limited. About 50% of hand-outcrossed and open-pollinated senita flowers abort by day 6 after flower opening. Results of exclusion experiments indicated that senita moths accounted for 75% of open-pollinated fruit set in 1995 with two species of halictid bees accounting for the remaining fruit set. In 1996, flowers usually closed before sunrise, and senita moths accounted for at least 90% of open-pollinated fruit set. The net outcome of the senita/senita moth interaction is mutualistic, with senita larvae destroying about 30% of the seeds resulting from pollination by senita moths. Comparison of the senita system with the yucca/yucca moth mutualism reveals many similarities, including reduced nectar production, active pollination, and limited seed destruction. The independent evolution of many of the same features in the two systems suggests that a common pathway exists for the evolution of these highly specialized pollination mutualisms. Nocturnal flower opening, self-incompatible breeding systems, and resource-limited fruit production appear to be important during this evolution. Received: 19 August 1997 / Accepted: 24 November 1997     

Florivore impacts on plant reproductive success and pollinator mortality in an obligate pollination mutualism

Florivores are present in many pollination systems and can have direct and indirect effects on both plants and pollinators. Although the impact of florivores are commonly examined in facultative pollination mutualisms, their effects on obligate mutualism remain relatively unstudied. Here, we used experimental manipulations and surveys of naturally occurring plants to assess the effect of florivory on the obligate pollination mutualism between yuccas and yucca moths. Yucca filamentosa (Agavaceae) is pollinated by the moth Tegeticula cassandra (Lepidoptera: Prodoxidae), and the mutualism also attracts two florivores: a generalist, the leaf-footed bug Leptoglossus phyllopus (Hemiptera: Coreidae), and a specialist, the beetle Hymenorus densus (Coleoptera: Tenebrionidae). Experimental manipulations of leaf-footed bug densities on side branches of Y. filamentosa inflorescences demonstrated that feeding causes floral abscission but does not reduce pollen or seed production in the remaining flowers. Similar to the leaf-footed bugs, experimental manipulations of beetle densities within individual flowers demonstrated that beetle feeding also causes floral abscission, but, in addition, the beetles also cause a significant reduction in pollen availability. Path analyses of phenotypic selection based on surveys of naturally occurring plants revealed temporal variation in the plant traits important to plant fitness and the effects of the florivores on fitness. Leaf-footed bugs negatively impacted fitness when fewer plants were flowering and leaf-footed bug density was high, whereas beetles had a positive effect on fitness when there were many plants flowering and their densities were low. This positive effect was likely due to adult beetles consuming yucca moth eggs while having a negligible effect on floral abscission. Together, the actions of both florivores either augmented the relationship of plant traits and fitness or slightly weakened the relationship. Overall, the results suggest that, although florivores are always present during flowering, the impact of florivores on phenotypic selection in yuccas is strongly mitigated by changes in their densities on plants from year to year. In contrast, both florivores consistently influenced pollinator larval mortality through floral abscission, and H. densus beetles additionally via the consumption of pollinator eggs.     

Pollinator attraction to volatiles from virgin and pollinated host flowers in a yucca/moth obligate mutualism

The classic obligate pollination–seed consumption mutualism between yuccas and yucca moths has been thought to be mediated by chemical cues, but empirical data on pollinator attraction to host floral volatiles in this association have been lacking. Here we show that the scent from virgin flowers of the host Yucca glauca is sufficient to attract its obligate pollinator Tegeticula yuccasella in Y‐tube olfactometer tests. Interestingly, both sexes of moths were attracted to the scent stimulus. Because yucca moths mate inside host flowers, the attraction of both females and males to host floral volatiles is likely to increase encounter rates. In a second test, female moths did not discriminate between virgin and hand‐pollinated flowers, indicating no post‐pollination change in scent production by the host that would lead to a reduction in pollinator attraction and thereby limit exploitation of the available seeds in host flowers. However, other mechanisms that could stabilise the mutualism between T. yuccasella and its yucca hosts have already been documented, i.e. selective abortion of heavily infested flowers, and a female‐derived host‐marking pheromone. Headspace collection and GC–MS were used to identify the blend of floral volatiles emitted by Y. glauca, which was found to be very similar to those of two other allopatric capsular‐fruited species, Y. elata and Y. filamentosa, revealing strong conservation of this trait within Yucca section Chaenocarpa.     

The role of abiotic and biotic factors in determining coexistence of multiple pollinators in the yucca–yucca moth mutualism

The determinants of a species' geographic distribution are a combination of both abiotic and biotic factors. Environmental niche modeling of climatic factors has been instrumental in documenting the role of abiotic factors in a species' niche. Integrating this approach with data from species interactions provides a means to assess the relative roles of abiotic and biotic components. Here, we examine whether the high host specificity typically exhibited in the active pollination mutualism between yuccas and yucca moths is the result of differences in climatic niche requirements that limit yucca moth distributions or the result of competition among mutualistic moths that would co‐occur on the same yucca species. We compared the species distribution models of two Tegeticula pollinator moths that use the geographically widespread plant Yucca filamentosa. Tegeticula yuccasella occurs throughout eastern North America whereas T. cassandra is restricted to the southeastern portion of the range, primarily occurring in Florida. Species distribution models demonstrate that T. cassandra is restricted climatically to the southeastern United States and T. yuccasella is predicted to be able to live across all of eastern North America. Data on moth abundances in Florida demonstrate that both moth species are present on Y. filamentosa; however, T. cassandra is numerically dominant. Taken together, the results suggest that moth geographic distributions are heavily influenced by climate, but competition among pollinating congeners will act to restrict populations of moth species that co‐occur.     

Geographic isolation trumps coevolution as a driver of yucca and yucca moth diversification

Coevolution is thought to be especially important in diversification of obligate mutualistic interactions such as the one between yuccas and pollinating yucca moths. We took a three-step approach to examine if plant and pollinator speciation events were likely driven by coevolution. First, we tested whether there has been co-speciation between yuccas and pollinator yucca moths in the genus Tegeticula (Prodoxidae). Second, we tested whether co-speciation also occurred between yuccas and commensalistic yucca moths in the genus Prodoxus (Prodoxidae) in which reciprocal evolutionary change is unlikely. Finally, we examined the current range distributions of yuccas in relationship to pollinator speciation events to determine if plant and moth speciation events likely occurred in sympatry or allopatry. Co-speciation analyses of yuccas with their coexisting Tegeticula pollinator and commensalistic Prodoxus lineages demonstrated phylogenetic congruence between both groups of moths and yuccas, even though moth lineages differ in the type of interaction with yuccas. Furthermore, Yucca species within a lineage occur primarily in allopatry rather than sympatry. We conclude that biogeographic factors are the overriding force in plant and pollinator moth speciation and significant phylogenetic congruence between the moth and plant lineages is likely due to shared biogeography rather than coevolution.     

Reproductive consequences of interactions between Yucca glauca (Agavaceae) and Tegeticula yuccasella (Lepidoptera) in Colorado

For over 100 years the association between Yucca (Agavaceae) and Tegeticula (Lepidoptera: Incurvariidae) has been accepted as a quintessential example of an obligate mutualism. The yucca moth is purported to be the sole pollinator of Yucca, while Yucca flowers provide courtship and mating arenas, and Yucca seeds provide food for developing Tegeticula larvae. We studied Yucca glauca in northern Colorado, comparing the reproductive ecology and breeding systems of Yucca in plains populations, the “preferred” habitat of Yucca, with “marginal” sites at the edge of the local elevational distribution. Tegeticula are abundant at plains sites, and fruit set is significantly higher than in the foothills, where fruit set is limited by the paucity of moths. The low frequency of moths at high elevation, coupled with behaviors of adult female Tegeticula which lead to self-pollination, failure to pollinate, and periodic overloading of fruits with eggs, may help explain why Yucca glauca appears to maintain alternative pollinators. Some fruits lack evidence of Tegeticula infestation, suggesting that larvae die before completing development, or that fertilization of Yucca sometimes occurs without the intervention of Tegeticula. Biochemical analyses of nectar and observations of floral visitors revealed that it is highly likely that the fly Pseudocalliope sp. nov. (Lauxaniidae), which congregates and mates on Yucca glauca blossoms, acts as a secondary pollinator. Autogamy appears to occur infrequently in natural populations. We therefore propose that the yucca-yucca moth symbiosis be viewed as a facultative mutualism.     

Defection by plants in the yucca–yucca moth association: a test of the cheater plant hypothesis for Yucca treculeana

In mutualisms, an underlying conflict of interests may select for defection from providing benefits. In the obligate mutualism between yuccas and yucca moths, where pollination service and seeds for pollinator larvae are traded, it has been suggested that some individuals in a population of Y. baccata may defect by preventing pollinator egg or larvae from development. We tested this hypothesis in Y. treculeana , another species suggested to contain cheater plants. Five specific predictions were tested during two years of study. A prediction that a surplus of plants without pollinator larvae should be present was met. Predicted existence of two distinct fruit morphs was rejected, and none of several highly variable morphological traits were linked to presence/absence of larvae. Predicted excess of intact seeds in the fruits of plants without larvae was not found; in fact, such plants produced fewer seeds, contrary to the hypothesis. A suggestion that inverse frequency-dependent fitness could explain the pattern was rejected. Contrary to prediction, distribution of larvae of a closely related cheater yucca moth was positively associated with pollinator larvae, even though it would not be affected by the proposed killing mechanism. The results together provide strong support against the existence of cheater plants in Y. treculeana .