Phoradendron

Phoradendron is a genus of mistletoe, native to warm temperate and tropical regions of the Americas. The center of diversity is the Amazon rainforest.[3] Phoradendron is the largest genus of mistletoe in the Americas, and possibly the largest genus of mistletoes in the world.[4] Traditionally, the genus has been placed in the family Viscaceae, but recent genetic research acknowledged by the Angiosperm Phylogeny Group shows this family to be correctly placed within a larger circumscription of the sandalwood family, Santalaceae.

Phoradendron
Phoradendron californicum fruit
Scientific classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Order: Santalales
Family: Santalaceae
Genus: Phoradendron
Nutt.[1][2]
Species and Distribution

about 235-240 species; see text

Phoradendron distribution by state or province.
Synonyms
  • Allobium Miers
  • Spiciviscum Engelm.[1]
Phoradendron strongyloclados showing leathery leaves and an inflorescence resulting from one apical meristem.
Phoradendron macrophylla in Arizona sycamore
Multiple Phoradendron individuals parasitizing an oak tree. While lower branches show new leaf growth, the mistletoe appears to be negatively affecting the tree's budding in the upper branches.
Phoradendron californicum (Desert Mistletoe), Granite Mountains, Mojave Desert, California

They are woody hemi-parasitic shrubs with branches 10–80 cm (3.9–31.5 in) long, which grow on other trees. The foliage is dichotomously branching, with opposite pairs of leaves; these are fairly large, 2–5 cm (0.79–1.97 in) long, green and photosynthetic in some species (e.g. P. leucarpum), but minimal in some others (e.g. P. californicum). Although they are able to photosynthesize the plant relies on its host for some nutrients. The plant draws its mineral and water needs, and some of its energy needs, from the host tree using a haustorium which grows into the stems of the host.[5]

The flowers are inconspicuous and incomplete, no petals and 3-4 greenish-yellow sepals,[6] 1–3 mm (0.039–0.118 in) diameter. The fruit is a berry, white, yellow, orange, or red when mature, containing one to several seeds embedded in very sticky juice, called viscin.[4] The flowers are unisexual, and depending on the species, the plant will be monoecious or dioecious (both male and female flowers on a single plant or male and female plants with only one sex of flowers).[4] The seeds are dispersed when birds eat the fruit and remove the sticky seeds from their bills by wiping them on tree branches where they can germinate.

The foliage and berries of some species are toxic. Leafy mistletoes seldom kill but they cause stress reducing crop productions in fruits and nut trees.[7]

Phoradendron plants can be distinguished from mistletoes in other genera in Viscaceae by their inflorescences, which lack leaves and come from a single branching point or apical meristem.[4] However, it can be difficult to identify species within Phoradendron, because leaf shape and color can vary greatly even within species.[6]

Phoradendron sp. on Spanish oak

Host and symptoms

Phoradendron species can infest many taxa of plants including hackberry (Celtis spp.) mesquite (Prosopsis spp.), cedar, elm (Ulmus spp.), and Osage-orange.[8] Certain species of Phoradendron are host-specific; for example, in Arizona, Phoradendron tomentosum infests cottonwood (Populus fremontii), sycamore (Platanus wrightii), ash (Fraxinus spp.), walnut (Juglans spp.) and willow (Salix spp.).[9] P. californicum infests shrubs and trees such as acacia (Acacia spp.) and blue palo verde (Parkinsonia florida). Some species infest junipers (Juniperus spp.) and oaks (Quercus spp.).[10] Branches often become swollen and distorted, forming burls and making the tree more susceptible to insect attack.[8] Phoradendron presents serious problems along rivers, streams, parks, and golf courses with large cottonwood trees. Deciduous trees can be mistaken for evergreens during the winter if infection is heavy.[10] Other common symptoms include swelling formations of witch's broom, dieback, and weakened branches.[11]


Life cycle

Phoradendron species are hemiparasites which produce their own chlorophyll but rely on the host plant to provide water, minerals, and other nutrients.[12] Birds are the primary means of dispersal of the parasite.[13] Birds consume the drupes of the mistletoe and excrete or regurgitate the seeds onto the branches of the host plant. The seeds do not need to be ingested to germinate. Germinating seeds produce a radicle, a holdfast, and eventually the germinated seeds produce haustoria. The haustorium is a root-like structure that penetrates the host plant's bark and cambium, reaching the xylem and phloem where it extracts water and minerals, primarily carbon and nitrogen compounds. A study on the nutrient ratio between Phoradendron and their hosts found that the parasite have higher concentrations of nitrogen and minerals, especially in leguminous hosts. This suggests that the parasite actively draws nutrients from the host plant through both the xylem and phloem, challenging the alternative theory of the passive uptake of nutrients by the parasite from the host xylem only.[14] The most important birds for effective dispersal include the cedar waxwing, euphonias, silky-flycatchers, bluebirds, and thrushes.[9]

Phoradendron caripense

Management

Leafy mistletoe can adversely affect trees growing in urban environments and in forests.[12] It is considered a nuisance in urban environments because of its appearance on deciduous trees during winter. Severe colonization of mistletoe can affect the health of an individual tree, and a tree already stressed by other factors can be killed. Forest fragmentation can increase Phoradendron infection rates in some oak trees, as trees in lower density forests and those closer to the forests' edges are more likely to be colonized by the mistletoe.[15] Control and management regimes include watering the host plant to improve its vigor, and removing infested vegetation. Pruning infested branches is not generally effective because the haustoria can infiltrate deeply. Plucking the mistletoe herbage is a temporary treatment because it easily resprouts, but keeping its herbage sparse can help to reduce its seed production.[9]

Wildlife & human uses

Phorodendron species are larval host plants for a number of Lepidoptera. The buckthorn duskywing (Erynnis pacuvius) feeds on Phoradendron californicum.[16]

Phoradendron are the preferred food of the Phainopepla, a silky-flycatcher. The male defends territories where fruiting mistletoe is abundant.[17]

There may actually be some mutualistic interactions between the parasite and the host in some Phoradendron species. The presence of Phoradendron juniperinum on host Juniperus monosperma, for example, has been suggested to increase dispersal of the host's seeds by birds. The mistletoe berries may attract frugivorous birds (e.g. thrushes) to eat the host juniper's seeds and disperse them. The net benefit on the hosts is difficult to quantify, however. The birds dispersing the host's seeds and benefiting the host, while at the same time dispersing the mistletoe's seeds and spreading the Phoradendron infection.[18]

Phoradendron californicum was used by Native American peoples as a food and medicinal remedy.[19]

Some Phoradendron species, such as Phoradendron serotinum and Phoradendron flavescens, are used in North America as Christmas decoration, substituting for the European mistletoe Viscum album.[20] They are harvested commercially for that purpose.

Etymology

Nuttall named the genus Phoradendron from the Greek, phor (a thief) and dendron (tree), alluding to the genus' parasitic habit.[2]

Taxonomy

There are roughly 235[3] to 240[21] species in the genus.

Species include:[22][23][24][25]

References
  1. "Genus: Phoradendron Nutt". Germplasm Resources Information Network. United States Department of Agriculture. 2010-08-25. Archived from the original on 2012-10-10. Retrieved 2011-03-08.
  2. Nuttall, T. 1848. Journal of the Academy of Natural Sciences of Philadelphia ser. 2, 1 185.
  3. Coder, K. D. American mistletoe (Phoradendron serotinum var. serotinum) infection in trees. WSFNR08-25. Tree Health Series. University of Georgia. 2008.
  4. Kujit, Job (2003). "Monograph of Phoradendron (Viscaceae)". Systematic Botany Monographs. 66: 1–643. doi:10.2307/25011253. JSTOR 25011253.
  5. Hiller, J. B. (2010). Marvelling at mistletoe. Highlights for Children. 65(12) 40-41.
  6. Kellogg, Elizabeth A. (1991). "Why Study Mistletoes?". Arnoldia. 51 (3): 11–17. JSTOR 42954407.
  7. Krautwurst, T. (2003) The truth about mistletoe. Mother Earth News 195 26-29.
  8. Turner, M. W. Remarkable Plants of Texas: Uncommon Accounts of our Common Natives. University of Texas Press. 2009. pg. 261.
  9. Geils, B. W., et al. Mistletoes of North American conifers. Gen. Tech. Rep. RMRS-GTR-98. Ogden, Utah: USDA, Forest Service, Rocky Mountain Research Station. 2002.
  10. Hine, R. Diseases of Urban Plants. Adapted by M. Olsen. Plant Disease Publications. University of Arizona Cooperative Extension. Revised 1999.
  11. Call, R. E. Mistletoe. Archived 2010-07-26 at the Wayback Machine Home Horticulture. University of Arizona Cooperative Extension, Cochise County.
  12. Watson, W. T. and T. Martinez-Trinidad. (2006). Strategies and treatments for leafy mistletoe (Phoradendron tomentosum (DC.) Engelm ex. Gray) suppression on cedar elm (Ulmus crassifolia Nutt.). Arboriculture and Urban Forestry 32(6) 265-70.
  13. Aukema, J. E. (2004). Distribution and dispersal of desert mistletoe is scale-dependent, hierarchically nested. Ecography 27 137-44.
  14. Panvini, A. Darlene; Eickmeier, William G. (1993). "Nutrient and Water Relations of the Mistletoe Phoradendron leucarpum (Viscaceae): How Tightly are they Integrated?". American Journal of Botany. 80 (8): 872–878. doi:10.1002/j.1537-2197.1993.tb15307.x. JSTOR 2445506.
  15. Barbosa, Jomar M.; Sebastián-González, Esther (2016). "Hemiparasite--host plant interactions in a fragmented landscape assessed via imaging spectroscopy and LiDAR". Ecological Applications. 26 (1): 55–66. doi:10.1890/14-2429. JSTOR 24701220. PMID 27039509.
  16. Soule, J. A. Butterfly Gardening in Southern Arizona. Tierra del Sol Institute. Tucson, Arizona. 2012. ISBN 0-9758554-3-3
  17. Soule, J. A. Father Kino's Herbs: Growing Them & Using Them Today. Tierra del Sol Institute. Tucson, Arizona. 2010. ISBN 0-9758554-2-5
  18. van Ommeren, Ron J. (January 1, 2002). "Changes in interactions between juniper and mistletoe mediated by shared avian frugivores: parasitism to potential mutualism". Oecologia. 130 (2): 281–288. Bibcode:2002Oecol.130..281V. doi:10.1007/s004420100792. PMID 28547152.
  19. Phoradendron californicum. Native American Ethnobotany. University of Michigan, Dearborn.
  20. Solomon, J. D., et al. Ash pests: A guide to major insects, diseases, air pollution injury and chemical injury. Gen. Tech. Rep. SO-96. New Orleans, Louisiana. USDA Forest Service, Southern Forest Experiment Station. 1993.
  21. Phoradendron. The Jepson Manual eFlora 2012.
  22. "GRIN Species Records of Phoradendron". Germplasm Resources Information Network. United States Department of Agriculture. Retrieved 2011-03-08.
  23. "Phoradendron". Integrated Taxonomic Information System. Retrieved 2011-03-08.
  24. Phoradendron. Bolivia Checklist. eFloras.org
  25. "Phoradendron Nutt. | Plants of the World Online | Kew Science". Plants of the World Online. Retrieved 2019-12-03.
  26. Wiens, D. and C. L. Calvin. (2011). Two epiparasitic species of Phoradendron (Viscaceae) from Honduras: one new and for the other a range extension and host determination. Aliso 29(2) 119-23.
  27. Kuijt, J. (1997). Phoradendron olae Kuijt, a new species from Mexico pivotal in the taxonomy of the genus, with comments on P. californicum Nutt. Brittonia 49(2), 181-88.
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