Tegeticula antithetica Yucca Moth Tegeticula antithetica belongs to the order Lepidoptera and suborder Glossata.

. It is a member of the Prodoxidae family (superfamily: Incurvarioidea), which is commonly called Yucca Moths because of their close association with Yucca plants. The Tegeticula and Parategeticula genera are the only two genera of the family that are obligate pollinators as well as herbivores of Yucca plants. Yucca moths are famous for being one of the best-studied examples of coevolution between a plant and insect, as the plants rely on adult moths for their pollination and the moth larvae depend on developing seeds to complete their development [3].

Figure 1: A Yucca moth on a Yucca flower

Geographic Range Yucca moths can be found wherever their host plants are located. In the case of T. antithetica, its host plant is the Joshua tree (Yucca brevifolia), located in the Mojave Desert of the North American southwest, specifically southern California, southern Nevada, southwestern Utah, and western Arizona (Figure 2)[1]. Other related yucca plants and moths (Tegeticula and Parategeticula) are generally located in the American west and northern Mexico.

Figure 2: Geographic distribution of Y. brevifolia and its two Tegeticula pollinators. Map includes southern California, southern Nevada, southwestern Utah, and western Arizona. Y. brevifolia distribution is outlined in white line, with the dashed line indicating the split in the two Yucca varieties, brevifolia and vespertina. Black circles represent T. synthetica and black squares represent T. antithetica

Morphology T. antithetica is most closely related to T. synthetica because the two species are sister taxa and diverged from a single pollinator species of Yucca brevifolia [1]. The two moths may have allopatrically speciated in response to the Bouse embayment, an estuarine extension that inundated low-lying areas of the Mojave Desert region ~6.5 million years ago [2]. This may have caused the considerable differences in stature and branching architecture between the eastern and western versions of the Joshua tree (Figure 3) and its pollinators T. antithetica and T. synthetica, respectively. T. antithetica has a wingspan of 13.5-15 mm in males and 13-16 mm in females. T. antithetica can be distinguished by arrow-shaped mark in the discal cell of its forewing (Figure 4). T. antithetica is significantly smaller than its close relative T. synthetica (Figure 5). The integuments are dark brown and the ventral side of the female head has 20-25 sensory setae (the male having none) and the antennae are about a third the length of the forewing. The genitalia of the males also allow for distinguishing between the two closely related species. The valvae of T. antithetica have much smaller ventral protrusions and a different number and distribution of spines on those protrusions (Figure 6).

Figure 3: (A) Yucca brevifolia with high branching, asymmetric crown, and long narrow pistil (inset) typical of Yuccas pollinated by T. synthetica. (B) Y. brevifolia with low branching, symmetric crown, and short thick pistil (inset) typical of Yuccas pollinated by T. antithetica. Photos not to scale

Figure 4: (A) Male T. antithetica and (B) Male T. synthetica. (C, inset) shows an enlarged image of the arrow-shaped mark in the discal (central) cell of the T. antithetica forewing. The arrows are pointing to the notable differences in the moths valvae (claspers). The scale bars in both A and B are 2.5 mm. The hind wing in A is actually darker, but appears brighter because of a reflecting glare in the photo.

Figure 5: Half-images of female T. synthetica (left) and T. antithetica (right). Note the arrow-shaped make on the forewing of T. antithetica.

Figure 6: Male genitalia of (A) T. antithetica and (B) T. synthetica. The right valvae are shown for both. Arrows indicate the differences in ventral protrusions of the valvae and differences of spine number and distribution. Scale bars represent 0.5 mm.

Life Cycle, Ecology & Behavior The female Tegeticula moth oviposits Joshua tree flowers by cutting through the ovary wall and extending her ovipositor down the stylar canal to lay eggs atop the ovules. Then, the female pollinates the flower to ensure the production/ availability of seeds for her larvae to feed on (Figure 7) [2]. The female does this by using unique tentacular mouthparts to collect pollen by dragging them across the anthers. The moth compacts the pollen using the tentacular mouthparts and stores it underneath her head. Recurring pollen collection through the moths lifetime results in multiple pollen genotypes. After oviposition of the eggs, the female walks up to the stigma, scrapes some pollen off of her batch using her tentacles, and places the pollen on the style (see Video Clip) [4]. The eggs of Tegeticula hatch within a few days and then the larvae begin immediately feeding on the developing yucca seeds, eating only a small proportion [5]. After feeding, the larva creates an exit path so it can escape the flower to find a location to burrow itself. The larva waits inside the fruit for optimal weather conditionsuntil rainy and usually at nightthen burrows into the ground where it create a silk-lined cocoon covered with soil or sand. Inside the cocoon, the larva enters diapause and will pupate a few weeks before emergence. Interestingly, the larvas diapause can range from one to four years to help its emergence synchronize with host flowering. Based on Pellmyrs unpublished data, very high fruit set during mass flowering episodes in yucca populations that then effectively cease flowering almost completely for several years suggests that the moth larvae are capable of diapausing for several years as well, and that there are unidentified cues that trigger completion of the moths development and adult emergence [4]. Most Tegeticula species, including antithetica, are monophagous, and the adult moths only live for a few days, so they must access the plant during the short flowering period. This indicates that moth populations would have to be locally adapted for the flowering periods their specific hosts [4].

Figure 7: Female Tegeticula oviposition into a Joshua tree flower and a cross section of the floral pistil showing the path of the moths ovipositor. The female ovipositor first cuts through the stylar wall then pushed down the stylar canal to the ovules. (Video)

http://www.denimandtweed.com/2009/10/video-of-yucca-pollination.html
Video: A yucca moth laying eggs, then actively pollinating a yucca flower.

Interesting Biology As mentioned above, Yucca moths have played a significant role in scientific research because their unique relationship with yucca plants provides a rare opportunity to study coevolution. While there are many examples in nature of mutualistic relationships between plants and insects, coevolution is particularly rare to find (and prove) because it requires reciprocal selective forces between two species. If the specific, beneficial traits are to signify coevolution, they have to have developed as a result of the two species interactions and not any extrinsic forces. Tegeticula and Yucca brevifolia are believed to have coevolved because the yucca plants rely exclusively on the moths for pollinationwhich actively pollinate the yucca flowers they will ovipositand the larvae feed exclusively on the developing seeds. The moths and plants physiological traits are intertwined because the moth has an exclusively sized ovipositor and only an active pollinator like Tegeticula can pollinate the plant. The females ovipositor must be long enough to reach the ovules but not so long as to injure them, as seeds would not develop to feed the larvae [5]. Coevolution acting on the partners should favor matching between the length of the ovipositor and the flowers stylar canal [2]. This is because the female moth only pollinates the flower after successfully ovipositing her eggs. The two species coevolve because any morphological change in the reproductive structures of one species forces a reciprocal selective force on the other. For example, if genetic drift permits the ovipositor length in a population of moths to shorten, then the only yucca plants that will get pollinated and reproduce are the ones with shorter stylar canals. Here, the change in one species population forces a change in the other species population, and vice versa. Overtime, enough changes can occur where two species become entirely dependent on each other for

survival and if the two species dont evolve with each other, than the populations will go extinct. Godsoe, et al. (2008) presented convincing evidence for this coevolution by showing the exclusive relationship between Tegeticula antithetica (and synthetica) and their respective populations of Yucca brevifolia. Perhaps most importantly, Godsoe et al. showed that only the reproductive features of the moths and plants have been evolvingovipositor length and floral characters, respectivelyand not body size or vegetative features, respectively. This indicates that only reciprocal sexual selection, and not extrinsic forces (such as climate, etc.), has been acting on the evolution of the two species. Like other organisms with specialized hosts, the ability to specialize can reduce competition. Tegeticula moths dont have to compete with numerous other populations of varying species for locations to lay their eggs, and Yucca brevifolia plants dont have the stress of too many insect populations exploiting its seeds.

References [1] Pellmyr, O. and Segraves, K. A. 2003. Pollinator Divergence within an Obligate Mutualism: Two Yucca Moth Species (Lepidoptera; Prodoxidae: Tegeticula) on the Joshua Tree (Yucca brevifolia; Agavaceae). Ann. Entomol. Soc. Am. 96(6): 716-722 [2] Godsoe, W., Yoder, J. B., Smith, C. I., and Pellmyr, O. 2008. Coevolution and Divergence in the Joshua Tree/ Yucca Moth Mutualism. The American Naturalist. 171(6): 816-823. [3] Pellmyr, O., Thompson, J. N., Brown, J. M., Harrison, R. G. 1996. Evolution of Pollination and Mutualism in the Yucca Moth Lineage. The American Naturalist. 148(5): 827-847. [4] Pellmyr, O. 2003. Yuccas, Yucca Moths, and Coevolution: A Review. Annals of the Missouri Botanical Garden. 90(1): 35-55

[5] Pellmyr, O., and C. J. Huth. 1994. Evolutionary stability of mutualism between yuccas and yucca moths. Nature. 372: 257-260. Figure 1: The yucca moth by: Grand Canyon Trust Volunteers (Flickr) http://www.flickr.com/photos/grand_canyon_trust/4627481885/ Figure 2: Pellmyr, O. and Segraves, K. A. 2003. Figure 3: Godsoe, et al. 2008. Figure 4: Pellmyr, O. and Segraves, K. A. 2003. Figure 5: Godsoe, et al. 2008. Figure 6: Pellmyr, O. and Segraves, K. A. 2003. Figure 7: Godsoe, et al. 2008.