• GENERAL BOTANICAL CHARACTERISTICS
• RAUNKIAER LIFE FORM
• REGENERATION PROCESSES
• SITE CHARACTERISTICS
• SUCCESSIONAL STATUS
• SEASONAL DEVELOPMENT

GENERAL BOTANICAL CHARACTERISTICS:

Inkberry is a stoloniferous or rhizomatous shrub forming extensive, dense colonies [45,48,63,69,110,116,153]. Inkberry may comprise 67 to 75% of shrub cover on coastal plain sites [116]. Several aerial stems arise singly or in groups from a rhizome that ranges in diameter from 0.2 to 0.5 inch (0.5-1.3 cm). A few small fibrous or woody roots occur along the entire rhizome, and longer roots (up to 48 inches or 120 cm) occur at infrequent intervals.  These longer roots may extend to the water table if it is near the surface; the rhizome itself grows in the upper 2 inches (5 cm) of soil [114]. In coastal plain loblolly forests, inkberry was estimated to have 660 to 1,459 rootstocks per acre (1,650-3,648 rootstocks/ha) [132].

Male plants have flowers occurring in clumps of 3 on long peduncles, while female plants have flowers occurring singly [45,69] or in clumps of up to 3 flowers [45]. Inkberry fruits grow individually or in small clusters, with 3 to 9 seeds [69].

RAUNKIAER [154] LIFE FORM:

REGENERATION PROCESSES:

Breeding system: Inkberry is dioecious [45,62,63,69,153].

Pollination: Inkberry is pollinated by insects [123].

Seed production: In general, inkberry produces good seed crops annually, though individual plants or clumps may bear heavily one year and be barren the next [69]. Immediately following fire, inkberry fruit production drops off, but within 4 years high fruit yields occur [187]. The pruning of older inkberry growth may cause inkberry to flower with renewed vigor, though 2 seasons are required [116]. Inkberry does not flower or produce fruit until the 2nd year after injury, at which point it often bears profusely [69,80,116]. In Georgia slash pine plantations, inkberry fruited heavily in all successional stages except recently burned stands, and produced maximum yields in 4-year-old plantations. Fruit yields were generally higher in young stands than in older stands [97]. After the 3rd or 4th season, the yield of inkberry fruits gradually declines, though fruit production occurs for many years [116].

Seed dispersal: Inkberry seeds are likely dispersed by birds [19].

Seed banking: No information

Germination: In general, members of the genus Ilex have immature embryos at time of ripeness, necessitating a period of after-ripening before germination will occur [45].

Seedling establishment/growth: Natural inkberry seedlings are rare; virtually all new growth originates from sprouting [69]. A study of a slash pine savannah in Mississippi found a greater frequency of established inkberry adults and a greater density of inkberry seedlings near the base of trees (<6.5 ft or 2 m). This trend is attributed to a higher seed rain underneath trees, presumably due to the dispersion of seeds by birds [19].

Asexual regeneration: Inkberry regenerates from readily sprouting rhizomes [62,63,116,119]. Mechanical damage to inkberry plants or simply exposure of a rhizome by removing soil may stimulate a new shoot to grow [116].

SITE CHARACTERISTICS:

Elevation and topography: Inkberry commonly occurs on the lower elevations of the coastal plain [203]. It occurs at elevations from 14 to 249 feet (4.3-76 m) [8,112,121,131,152]. Inkberry grows on sites with flat to slightly undulating topography [1,48,65,97,118,131,144,194,202].

Site types: Inkberry is found in wetlands [100,103,157,174], bogs [33,45,62,63,103,157,208], seeps [45,62,63], pocosins [7,153,156,157,176,206], swamps [72,134,135,148,170,171,174], lake and pond borders [4,72,164], and creek bottoms [45,55,62,63]. It also occurs in upland areas [55,103] including pine flatwoods [33,45,48,62,63,72,153,207,208], pine barrens [153], prairies [62,63,196], terraces [184], and savannahs [38,45,48,62,63,72,153]. Inkberry is also found on coastal sand dunes [38,196] and swales [33,208].

In Nova Scotia, inkberry is commonly found on rocky barrens and dry hillsides, in addition to swampy or mesic areas [165,185].

Soils: Soils supporting inkberry include coarse till [185,186], fine sand [4,21,38,54,65,68,69,70,72,77,78,121,125,131,150,196], loam [11,68,70,131], clay [54,70,125], marl [38,54,196], and peat [41,69,72,126,148,176]. These soils may have an underlying, impervious clay hardpan [4,114]. Soils often have low organic matter [4,54,72] and nutrient content [4,38,121,126,131,184,185], and are often acidic [1,4,45,62,63,69,121,126,131,134,152,176,184,186]. Soils may be calcareous [43,196], though Halls [69] reports that inkberry grows poorly on soils with high lime content.

Though it occurs on soils with low levels of extractable phosphorus (<2.0 ppm) [70], inkberry growth may be enhanced by pine needle fall due to increased phosphorus concentrations [21].

Inkberry is common on both well-drained [1,38,185,188] and poorly drained sites [1,20,38,56,57,65,68,72,77,97,176] with moist or wet soils [4,11,17,32,56,77,102,133,144,151] and a shallow or fluctuating water table [17,57,72,148,176,206]. The soils are often seasonally wet [4,72,152] or periodically flooded [1,134,184,196], saturated in winter and droughty during the growing season [54,72,148]. In New Jersey, inkberry may be restricted to areas where the roots can extend to the water table, and inkberry seldom occurs in abundance on upland sites [114].

SUCCESSIONAL STATUS:

SEASONAL DEVELOPMENT:

Related categories for SPECIES: Ilex glabra | Inkberry