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Introductory

SPECIES: Mahonia trifoliolata | Agarito
ABBREVIATION : MAHTRI SYNONYMS : Berberis trifoliata Moric. SCS PLANT CODE : MATR3 COMMON NAMES : agarito algerita TAXONOMY : The currently accepted scientific name of this taxon is Mahonia trifoliolata (Moric.) Fedde (Berberidae) [1,24]. LIFE FORM : Shrub FEDERAL LEGAL STATUS : No special status OTHER STATUS : NO-ENTRY COMPILED BY AND DATE : D. Tirmenstein November, 1990 LAST REVISED BY AND DATE : NO-ENTRY AUTHORSHIP AND CITATION : Tirmenstein, D. A. 1990. Mahonia trifoliolata. In: Remainder of Citation

DISTRIBUTION AND OCCURRENCE

SPECIES: Mahonia trifoliolata | Agarito
GENERAL DISTRIBUTION : Agarito grows throughout most of Texas except for the eastern and southeastern portions of the state. It extends westward into New Mexico and Arizona and southward into Mexico [52]. The variety trifoliolata grows only in Texas, south of Austin to Corpus Christi. The variety glauca occurs from west Texas through New Mexico into Arizona and Mexico [1]. ECOSYSTEMS : FRES15 Oak - hickory FRES30 Desert shrub FRES31 Shinnery FRES32 Texas savanna FRES33 Southwestern shrubsteppe FRES34 Chaparral - mountain shrub FRES35 Pinyon - juniper FRES38 Plains grasslands FRES39 Prairie FRES40 Desert grasslands STATES : AZ NM TX MEXICO ADMINISTRATIVE UNITS : AMIS BIBE CACA GUMO BLM PHYSIOGRAPHIC REGIONS : 7 Lower Basin and Range 13 Rocky Mountain Piedmont 14 Great Plains KUCHLER PLANT ASSOCIATIONS : K031 Oak - juniper woodlands K041 Creosotebush K044 Creosotebush - tarbush K045 Ceniza shrub K054 Grama - tobosa prairie K059 Trans-Pecos shrub savanna K060 Mesquite savanna K071 Shinnery K084 Cross Timbers K085 Mesquite - buffalograss K086 Juniper - oak savanna K087 Mesquite - oak savanna SAF COVER TYPES : 66 Ashe juniper - redberry (Pinchot) juniper 67 Mohrs ("shin") oak 68 Mesquite 239 Pinyon - juniper 242 Mesquite SRM (RANGELAND) COVER TYPES : NO-ENTRY HABITAT TYPES AND PLANT COMMUNITIES : Agarito grows as a dominant or codominant in a number of desert shrub and grassland communities. It is on lands that have changed from mixed prairie to open scrub oak-juniper (Quercus-Juniperus spp.) woodlands [42] and where honey mesquite (Prosopis glandulosa)) is expanding its range in Texas savannas [2]. It is also a component of south Texas and Arizona chaparral, semiarid Texas grasslands, and pinyon (Pinus spp.)-juniper woodlands [9,29,39]. Common codominants include creosotebush (Larrea tridentata), tarbush (Flourensia cernua), lecheguilla (Agave lechuguilla), juniper, sotol (Dasylirion spp.), yucca (Yucca spp.), and black grama (Bouteloua eriopoda). Agarito has been listed as a dominant in the following community type classification: Natural terrestrial communites of Brewster County, Texas [21] Plant associates: Common associates of agarito in bottomland communities include little walnut (Juglans microcarpa), netleaf hackberry (Celtis reticulata), oneseed juniper (J. monosperma), littleleaf sumac (Rhus microphylla), and lotebush condalia (Condalia obovata) [20]. Pungent oak (Quercus pungens), live oak (Q. virginiana), mesquite, sugarberry (Celtis laevigata), Texas persimmon (Diospyros texana), spiny hackberry (Celtis pallida), curly mesquite (Hilaria belangeri), Texas needlegrass (Stipa leucotricha), grama (Bouteloua spp.), and lotebush (Ziziphus obtusifolia) occur with agarito in scrub oak-juniper woodlands and Texas savannas [2,6,42]. Ashe juniper (J. ashei) and redberry juniper are common overstory dominants [39,42]. In chaparral communities, honey mesquite, curly mesquite, acacia (Acacia spp.), threeawn (Aristida spp.) and buffalograss (Buchloe dactyloides) are common associates [8].

VALUE AND USE

SPECIES: Mahonia trifoliolata | Agarito
WOOD PRODUCTS VALUE : NO-ENTRY IMPORTANCE TO LIVESTOCK AND WILDLIFE : Browse: Agarito reportedly has no forage value for cattle, horses, or domestic sheep, and has little value for domestic goats [29]. Less than 1 percent of the tender new sprouts were browsed by Spanish goats following fire on the Edwards Plateau of Texas [49]. Agarito is browsed by deer in some areas [35] but is generally considered an "undesirable deer food" [5]. In some locations, twigs, bark, and leaves of grape-hollies (Mahonia spp.) are eaten by the ringtail, and various species of hares and rabbits [37]. Fruit: Agarito berries are readily eaten by many species of small birds [52] and mammals. PALATABILITY : Leaves of agarito are spiny and unpalatable [29] and toughen with age [53]. Mature leaves may remain palatable to some insects, such as foraging ants, but often become too tough for the ants to cut [53]. Increasing toughness may deter large herbivores as well. Berries are highly palatable to many species of small birds and mammals. NUTRITIONAL VALUE : Nutrient value of agarito varies by plant part [30] and phenological development. The chemical composition of mature foliage has been documented as follows [26]: percent protein ether crude N-free H20 ash potash lime magnesia phosphoric extract fiber extract acid 10.32 2.38 30.59 46.61 6.13 2.97 0.91 0.88 0.24 0.34 Nutrient content by plant part and date is listed below [30]: plant part date water (%) ash (%) P (%) protein (%) shoots 3/27 69 4 0.27 16 shoots 4/13 48 3 0.25 13 shoots 10/25 71 4 0.27 16 leaves+stems 3/28 71 4 0.29 15 COVER VALUE : Agarito provides valuable cover for many wildlife species [43]. Clumps or thickets serve as important hiding cover for white-tailed deer on the rolling Texas plains [20]. Agarito forms good hiding, nesting, and resting cover for a variety of small birds and mammals [35]. In parts of Texas, wild turkeys occasionally nest beneath agarito [17]. VALUE FOR REHABILITATION OF DISTURBED SITES : Rehabilitation value of agarito is unknown. However, plants can be easily propagated from seed [24]. Seed can be planted in the fall, or stratified and planted in the spring [43]. Agarito can also be propagated by suckers, cuttings, and layering. Properly treated cuttings taken in early summer will usually root by fall [24]. OTHER USES AND VALUES : Roots of agarito contain berberine and other alkaloids and were traditionally used to make preparations for treating toothaches and stomach ailments [40,52]. However, these alkaloids are poisonous in high concentrations [40]. Yellow dyes were made from the roots and wood [52]. Fruit of agarito is edible [51] and can be used to make wine and jelly [52]. However, Durand [24] cautions that when collecting the fruit of Texas mahonias, "it is always a good precaution to poke around the bush with a stick before threshing to make sure there are no rattlesnakes." Seeds of agarito can be roasted and used as a coffee substitute [52]. Flowers provide nectar for honeybees, and consequently agarito is considered to be a good honey plant [51,52]. The attractive leaves and flowers make agarito well-suited for use as an ornamental [45]. It can be planted as a hedge or used singly [24,45]. When mixed with Amur privit, it forms an excellent and attractive combination hedge [24]. The variety glauca has been cultivated in England [1]. Agarito is relatively intolerant of cold temperatures and in North America is not hardy north of zone 6 [46]. MANAGEMENT CONSIDERATIONS : Brushfield expansion: During the past century, shrubs such as mesquite (Prosopis spp.) and agarito have been increasing on Texas rangelands [2,3,10,39]. An estimated 736,744 acres (1.82 million ha) of Texas rangeland is now "infested" with agarito. The situation is particularly pronounced on the Edwards Plateau where agarito density can reach up to 121 plants per acre (300/ha) [19]. Possible causes of this shrub expansion include fire suppression, overgrazing, and/or climatic change [2,10]. Grazing: In semiarid grasslands of Texas, heavy grazing apparently favors the spread of redberry juniper (Juniperus pinchotii) which in turn facilitates the establishment of agarito. Frequency of agarito is typically highest on grazed sites occupied by redberry juniper, as illustrated below [39]: high plains rolling plains grazed sites ungrazed sites redberry redberry present absent present absent j. present j. absent (percent frequency) 29 0 63 20 25 0 Mechanical removal: Ranchers commonly view agarito as a nuisance [24], and numerous studies have focused on various means of mechanical removal [19,22,23]. Soil penetration to a depth of 4 to 6 inches (10-15 cm) is generally necessary to sever the taproot from the crown and to uproot all lateral roots capable of resprouting. Resprouting often occurs where blade penetration is insufficient. Small plants with shallow roots are most readily killed by mechanical means [19]. On the Edwards Plateau of Texas, various types of mechanical grubbing have produced erratic mortality [19]. Grubbing, using a modified blade with fins on top, killed up to 93 percent of the plants where pretreatment densities had ranged from 17 to 79 plants per acre (42-195/ha). Consequently, this method of control was judged both effective and economical [19]. On rocky outcrops with shallow soil, the blade was prevented from eliminating all of the lateral roots. An estimated 88 percent of agarito was killed on these sites. Root plowing is also effective, although often prohibitively expensive. Posttreatment burning, when used in combination with mechanical removal, can also aid in reducing agarito density [see Fire Management Considerations]. Chemical control: Agarito is resistant to many herbicides including 2,4,5-T and picloram [14,19,22]. Although canopies were reduced by 24 to 30 percent 13 months after large amounts of picloram were applied to the soil, plants recovered within 24 months [33]. No plants were actually killed by the herbicide. Details on response to various rates of herbicide application are available [14,29,22]. Damage/disease: Agarito is susceptible to black stem rust [52]. However, roots contain large amounts of the alkaloid berberine, a substance known to inhibit some root fungi. Roots may, therefore, be relatively resistant to a number of pathogens [24]. Biomass: Models have been developed for predicting biomass estimates of agarito. Current growth is primarily leaves and includes little twig elongation. Studies indicate that new growth on young plants is greater per unit volume than is growth on older plants [16].

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Mahonia trifoliolata | Agarito
GENERAL BOTANICAL CHARACTERISTICS : Agarito is a dense, thicket-forming evergreen shrub which grows 3 to 10 feet (1-3 m) in height [1,46,51,52]. Twigs are smooth and reddish-green when young but turn gray to reddish-brown with age [52]. Bark is gray to reddish-brown and often exfoliating [28,52]. The alternate, trifoliolate leaves are stiff, spiny, and hollylike [1,35,52]. Leaflets are thick and coriaceous, lanceolate-oblong to elliptic, and have coarsely serrate or spinose margins [1,28,52]. Leaflets are pale green to glaucous [28]. Yellowish, perfect flowers are borne in few-flowered racemes at the upper axils or terminally on short shoots [28,52]. Fruit is a subglobose to globose berry, 0.3 to 0.5 inch (8-12 mm) in diameter [1,28]. Berries are lustrous, and red or black to pruinose blue [1,28,35]. The pulpy fruit is acidic and aromatic [52]. Fruit is borne on short pedicels which are tightly appressed to the stem axis [24]. Each fruit contains one to several seeds [52]. RAUNKIAER LIFE FORM : Phanerophyte REGENERATION PROCESSES : Agarito reproduces through seed and sprouts vigorously following most types of disturbance. Seed: Agarito produces an abundance of seed nearly every year [43]. In related species of mahonia, annual fruit production is sometimes reduced by poor pollination and adverse weather conditions. The specific genetic composition of individual plants can also influence fruit production [24]. Seed of agarito is dispersed during the summer by a variety of birds and mammals. Under natural conditions, seed germinates the following spring [43]. Vegetative regeneration: Agarito typically sprouts vigorously from the roots or root crown after aboveground vegetation is removed or damaged [11,19]. Box and others [11] reported that agarito sprouts from or near ground level after fire. After mechanical removal, Cross and Wiedemann [19] observed sprouting from lateral roots and from crown tissue attached to the taproot. The majority of regrowth (56 percent) was attributed to lateral root sprouting, whereas 13 percent was derived from crown tissue. No sprouts were observed on the taproot itself. SITE CHARACTERISTICS : Agarito grows in a variety of habitats including flat pastureland, lower alluvial flats, in ephemeral drainage channels, on mesa sides, and on dry, stony hills [3,18,35,40,52]. Agarito grows well on sunny sites [51]. It is often well represented in riparian areas and in bottomland communities of the rolling Texas plains [18,20]. Soils: Agarito grows on a variety of soil textures including loam, clay, shallow clay-loam, and gravelly soil [19,22,39,51]. Soils are commonly dry and well drained [51]. Agarito often occurs on soils derived from limestone parent material [15,50]. Climate: Agarito grows in semiarid climates with average annual precipitation estimated at 22 to 30 inches (55-76 cm) [19,22]. Winters are typically short and mild, with as many as 283 frost-free days per year [9]. Elevation: Agarito grows at approximately 3,000 feet (914 m) in Arizona [32]. SUCCESSIONAL STATUS : Grasslands: Agarito is a common woody invader in semiarid grasslands of Texas [19,22]. On the Edwards Plateau, mixed prairie has given rise to a scrub oak-juniper disclimax in which agarito occurs as a prominent woody species [42]. In many semiarid grasslands of Texas, redberry juniper is a common invader on overgrazed sites. The presence of large junipers consequently facilitates the establishment and spread of agarito [39]. Texas savannas: In south Texas savannas which have been gradually invaded by honey mesquite, species such as Texas pricklypear (Opuntia lindheimeri) and prickly ash (Zanthoxylum fagara) assume dominance during years 12 to 26. Sugarberry, Texas persimmon, and lotebush condalia commonly dominate 29- to 39-year-old stands, while agarito, desert yaupon (Schaefferia cuneifolia), and lotebush dominate 36- to 45-year-old stands [2]. Agarito may eventually be lost from climax stands [3]. Desert shrub communities: Agarito invades disturbed desert shrub communities including those in the Guadalupe Mountains of New Mexico. It commonly appears in the "last stages of community degradation" [54]. Once vegetation has reached this level of degradation, recovery may be unlikely. SEASONAL DEVELOPMENT : Fruit ripens from April through July [51,52], but reportedly reaches peak ripeness on approximately May 1 [24]. Generalized flowering and fruiting dates by geographic location are as follows: Location Flowering Fruiting Authority Southwest ---- July Vines 1960 Great Plains March-April June Great Plains Flora Association 1986

FIRE ECOLOGY

SPECIES: Mahonia trifoliolata | Agarito
FIRE ECOLOGY OR ADAPTATIONS : Relatively frequent fires in western Texas rangeland tend to favor climax grass species [10]. Fire suppression and vegetative changes brought about by overgrazing have contributed to the invasion of woody species such as agarito [12]. Fine fuels have been reduced by both grazing and fire suppression and competition from grasses has decreased. Agarito generally sprouts from the roots or root crown after a single fire [11,19]. Growth in large mottes affords some protection from fire. Although the outer portion is commonly destroyed, centers of large mottes are often undamaged. Seed from adjacent unburned areas may be dispersed onto burned sites by birds and mammals. Some postfire seedling establishment is possible. POSTFIRE REGENERATION STRATEGY : Tall shrub, adventitious-bud root crown Small shrub, adventitious-bud root crown Geophyte, growing points deep in soil

FIRE EFFECTS

SPECIES: Mahonia trifoliolata | Agarito
IMMEDIATE FIRE EFFECT ON PLANT : Agarito is described as "very resistant to fire" [49]. It is readily top-killed by fire [5,49], but underground regenerative structures such as roots often survive [49]. In south Texas chaparral, mortality is typically greatest on the windward side of large mottes, and least on the leeward side. Centers of large mottes are often undamaged [11]. Few plants were root-killed by a prescribed burn conducted in Tom Green County, Texas [49]. However, 33 percent of individual agarito plants were killed after a fall burn in south Texas chaparral [11]. Burned plants generally exhibit damage such as split stems and shredding bark [11]. DISCUSSION AND QUALIFICATION OF FIRE EFFECT : NO-ENTRY PLANT RESPONSE TO FIRE : Agarito commonly sprouts from the roots or root crown after aboveground vegetation is consumed by fire [5,11,19]. Lateral sprouting is most common where the motte growth pattern existed prior to fire. Centers of large mottes often survive and subsequently increase in size in the postfire community [11]. Individual plants and small mottes are generally less likely to sprout. Fire can cause reductions in canopy cover and relative abundance of agarito [10,12]. A single burn reduced cover by as much as 58 percent in south Texas chaparral [10]. Following fire, many of the sprouts were of poor vigor and some plants produced only a single sprout. Approximately 40 percent of the plants that eventually died produced new sprouts during the first spring [11]. However, many sprouts died during the summer. Some postfire establishment from seed may occur as birds and mammals disperse seed from adjacent unburned sites. DISCUSSION AND QUALIFICATION OF PLANT RESPONSE : NO-ENTRY FIRE MANAGEMENT CONSIDERATIONS : Prescribed fire: On west Texas rangeland, fires tend to favor climax grass species and, when used in combination with other methods, can help to reduce the cover of woody colonizers such as mesquite and agarito [10]. Fire is most effective in reducing agarito when large mottes are first mechanically treated [11]. In southern Texas grasslands, density of agarito was reduced by 28 percent on plots which had been roller chopped, shredded and burned, or roller-chopped, treated with herbicides, shredded, and burned. Density was unchanged on plots which had been roller-chopped and shredded but not burned [22,23]. Without pretreatment, burns are often patchy and uneven and leave many large mottes intact [12]. Best results can often be obtained by waiting up to 3 years after the brush is chopped or shredded and allowing grass and forbs to grow vigorously among the drying woody fuels. The herbaceous growth and chopped portions of old brush tops can provide fuel for a relatively hot fire [11,12], which is presumably more effective in reducing agarito. The following canopy cover of agarito was reported 1 year after a fall burn in south Texas chaparral [11]: unburned burned control 5.6 1.1 shredded 1.2 .9 chopped .7 .7 scalped .3 .4 Evidence suggests that both fall and winter burns can reduce agarito. However, winter burns tend to favor forbs, whereas fall burns often decrease forb production and increase grass production[12]. In experiments in south Texas chaparral, a fall fire with a winter reburn was more effective in reducing agarito than either a single fall or winter fire [12]. Response by season of burn was as follows on the Welder Wildlife Refuge of Texas [12]: control fall winter fall and winter (percent composition) 4.1 2.8 1.6 2.2 Wildlife: Agarito is often replaced by more desirable browse species such as honeysuckle (Lonicera spp.) and hackberry (Celtis spp.) after fire in Texas shinoak rangeland [44].

REFERENCES

SPECIES: Mahonia trifoliolata | Agarito
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Prescribed range burning in the Edwards Plateau of Texas: Proceedings of a symposium; 1980 October 23; Junction, TX. College Station, TX: Texas Agricultural Extension Service, The Texas A&M University System: 22-26. [11430] 6. Bernard, Stephen R.; Brown, Kenneth F. 1977. Distribution of mammals, reptiles, and amphibians by BLM physiographic regions and A.W. Kuchler's associations for the eleven western states. Tech. Note 301. Denver, CO: U.S. Department of the Interior, Bureau of Land Management. 169 p. [434] 7. Best, Troy L.; Skupski, Marian P.; Smartt, Richard A. 1993. Food habits of sympatric rodents in the shinnery oak-mesquite grasslands of southeastern New Mexico. Southwestern Naturalist. 38(3): 224-235. [22136] 8. Box, Thadis W. 1960. Herbage production in four range plant communities in south Texas. Journal of Range Management. 13: 72-76. [3939] 9. Box, Thadis W. 1961. Relationships between plants and soils of four range plant communities in south Texas. Ecology. 42: 794-810. 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[5746] 27. Garrison, George A.; Bjugstad, Ardell J.; Duncan, Don A.; [and others]. 1977. Vegetation and environmental features of forest and range ecosystems. Agric. Handb. 475. Washington, DC: U.S. Department of Agriculture, Forest Service. 68 p. [998] 28. Great Plains Flora Association. 1986. Flora of the Great Plains. Lawrence, KS: University Press of Kansas. 1392 p. [1603] 29. Humphrey, Robert R. 1953. Forage production on Arizona ranges. III. Mohave County: A study in range condition. Bulletin 244. Tucson, AZ: University of Arizona, Agricultural Experiment Station. 79 p. [4440] 30. Huston, J. E.; Rector, B. S.; Merrill, L. B.; Engdahl, B. S. 1981. Nutritional value of range plants in the Edwards Plateau region of Texas. Report B-1375. College Station, TX: Texas A&M University System, Texas Agricultural Experiment Station. 16 p. [4565] 31. Kartesz, John T.; Kartesz, Rosemarie. 1980. A synonymized checklist of the vascular flora of the United States, Canada, and Greenland. Volume II: The biota of North America. Chapel Hill, NC: The University of North Carolina Press; in confederation with Anne H. Lindsey and C. Richie Bell, North Carolina Botanical Garden. 500 p. [6954] 32. Kearney, Thomas H.; Peebles, Robert H.; Howell, John Thomas; McClintock, Elizabeth. 1960. Arizona flora. 2d ed. Berkeley, CA: University of California Press. 1085 p. [6563] 33. Kitchen, Lynn M.; Scifres, C. J.; Mutz, J. L. 1980. Susceptibility of selected woody plants to pelleted picloram. Journal of Range Management. 33(5): 349-353. [10287] 34. Kuchler, A. W. 1964. Manual to accompany the map of potential vegetation of the conterminous United States. Special Publication No. 36. New York: American Geographical Society. 77 p. [1384] 35. Lamb, S. H. 1971. Woody plants of New Mexico and their value to wildlife. Bull. 14. Albuquerque, NM: New Mexico Department of Game and Fish. 80 p. [9818] 36. Lyon, L. Jack; Stickney, Peter F. 1976. Early vegetal succession following large northern Rocky Mountain wildfires. In: Proceedings, Tall Timbers fire ecology conference and Intermountain Fire Research Council fire and land management symposium; 1974 October 8-10; Missoula, MT. No. 14. Tallahassee, FL: Tall Timbers Research Station: 355-373. [1496] 37. Martin, Alexander C.; Zim, Herbert S.; Nelson, Arnold L. 1951. American wildlife and plants. New York: McGraw-Hill Book Company, Inc. 500 p. [4021] 38. McCain, John W., Hennen, Joe F. 1982. Is the taxonomy of Berberis and Mahonia (Berberidaceae) supported by their rust pathogens Cumminsiella santa sp. nov. and other Cumminsiella. Systematic Botany. 7(1): 48-59. [10286] 39. McPherson, Guy R.; Wright, Henry A.; Wester, David B. 1988. Patterns of shrub invasion in semiarid Texas grasslands. American Midland Naturalist. 120(2): 391-397. [7197] 40. Powell, A. Michael. 1988. Trees & shrubs of Trans-Pecos Texas including Big Bend and Guadalupe Mountains National Parks. 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Index

Related categories for Species: Mahonia trifoliolata | Agarito

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