|Flower of Lacandonia schismatica|
E.Martínez & Ramos
E.Martínez & Ramos
Lacandonia schismatica is the sole species of the genus Lacandonia and is endemic to the Lacandon Jungle in the Mexican state of Chiapas. It is known from very few populations and is considered endangered by the researchers who investigate this species. L. schismatica is a mycoheterotrophic species that contains no chlorophyll and has the unique characteristic of inverted positions of the male (androecium) and female (gynoecium) floral parts, something that had not been seen in any other plants with the occasional exception of some individuals of the related Triuris brevistylis.
L. schismatica is a small saprophytic plant that lacks chlorophyll and has a rhizomatous, mycotrophic habit. This species exhibits racemous inflorescences and bract-like leaves. The flowers are actinomorphic and are considered "inverted" from the typical flower arrangement–usually 3 (but sometimes two to four) stamens are in the center of the flower surrounded by 60 to 80 pistils. This characteristic where the position of the androecium and the gynoecium are inverted is unique in the known and described taxa of flowering plants.
Flowers of L. schismatica are bisexual and self-pollinate and fertilize before the flower opens (preanthesis cleistogamy). They are true flowers as opposed to pseudanthia as had been suggested earlier in the literature. The three-celled pollen grains germinate within the anthers and the pollen tube grows through the receptacle to reach the ovaries. L. schismatica can be found flowering year-round when its environment is moist enough, with a particularly active flowering period in November and December. Owing to the prenathesis cleistogamy, a form of autogamy (self-pollination), the known population of L. schismatica lacks genetic variability and has a high instance of homozygosity. The haploid chromosome number of this species is n = 9.
Distribution and habitat
L. schismatica is known from several small populations at altitudes around 200 m (656 ft) in the Lacandon Jungle, a rainforest in southeastern Mexico. It grows in shady sites within this rainforest. Gerrit Davidse and Esteban Martínez noted in 1990 how the plants are "extremely localized and highly endangered" due to encroaching habitat conversion to cattle pasture. They also explain that the species is difficult to cultivate and thus encourage other scientists to study this unique organism's biology before it can no longer be found in the wild.
Taxonomy and botanical history
L. schismatica was first described by Martínez and Clara Hilda Ramos in 1989, who placed the species in its own family, Lacandoniaceae, which itself was placed in the Triuridales. In 1991, Traudel Rübsamen-Weustenfeld suggested that L. schismatica be included in the family Triuridaceae within the genus Sciaphila, Peltophyllum, or its own genus. Another study in 1998 presented data that supports the separation of L. schismatica into its own, monotypic family. The APG II system transferred the genus to the Triuridaceae and placed that family in the Pandanales.
The difficulty expressed in correctly placing the species in the proper family is due to the unique floral morphology. How this inverted position of the androecium and gynoecium evolved is unknown, but some studies have posed hypotheses. Davidse and Martínez suggested that L. schismatica could be one of Richard Goldschmidt's "hopeful monsters", meaning that the inverted floral morphology could have arisen from a macromutation in the genes that control floral development. It is also possible that chromosomal repatterning, also known as chromosomal rearrangement, was the origin of this species.
Since the original description and early work on this species in the 1990s, other field work has revealed some instances of L. schismatica flowers that were unisexual. The closely related species Triuris brevistylis was discovered to be mostly dioecious but a few individuals were located that had bisexual flowers with the flower arrangement inverted, just like that of the normal L. schismatica flowers. This discovery led the authors of the study to conclude that the inverted floral morphology evolved before L. schismatica and T. brevistylis diverged. Isolated populations during the Quaternary Period (around five million years ago) when temperatures in the Lacandon lowland rainforest were six to eight °C (10.8 to 14.4 °F) cooler than today. This hypothesis is supported by the geographic distribution where L. schismatica is restricted to the warmer lowlands and T. brevistylis has a distribution in the cooler highlands.
- Vázquez-Santana, S., Engleman, E. M., Martínez-Mena, A., and Márquez-Guzmán, J. (1998). Ovule and seed development of Lacandonia schismatica (Lacandoniaceae). American Journal of Botany, 85(3): 299-304.
- Márquez-Guzmán, J., Vázquez-Santana, S., Engleman, E. M., Martínez-Mena, A., and Martínez, E. (1993). Pollen development and fertilization in Lacandonia schismatica (Lacandoniaceae). Annals of the Missouri Botanical Garden, 80(4): 891-897.
- Davidse, G. and Martínez, E. (1990). The chromosome number of Lacandonia schismatica (Lacandoniaceae). Systematic Botany, 15(4): 635-637.
- Barbara A. Ambrose, Silvia Espinosa-Matías, Sonia Vázquez-Santana, Francisco Vergara-Silva, Esteban Martínez, Judith Márquez-Guzmán and Elena R. Alvarez-Buylla. (2006). Comparative developmental series of the Mexican triurids support a euanthial interpretation for the unusual reproductive axes of Lacandonia schismatica (Triuridaceae). American Journal of Botany, 93(1): 15-35.
- Coello, G., Escalante, A., and Soberon, J. (1993). Lack of genetic variation in Lacandonia schismatica in its only known locality. Annals of the Missouri Botanical Garden, 80(4): 898-901.
- Francisco Vergara-Silva, Silvia Espinosa-Matías, Barbara A. Ambrose, Sonia Vázquez-Santana, Alejandro Martínez-Mena, Judith Márquez-Guzmán, Esteban Martínez, Elliot M. Meyerowitz, and Elena R. Alvarez-Buylla. (2003). Inside-out flowers characteristic of Lacandonia schismatica evolved at least before its divergence from a closely related taxon, Triuris brevistylis. International Journal of Plant Sciences, 164(3): 345-357.
- Martínez, E. and Ramos, C. H. (1989). Lacandoniaceae (Triuridales): Una neuva familia de Mexico. Annals of the Missouri Botanical Garden, 76(1): 128-135.
- The Angiosperm Phylogeny Group. (2003). An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II. Botanical Journal of the Linnean Society, 141(4): 399-436.