Lindera Melissaefolia
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''Lindera melissifolia'', common name pondberry or southern spicebush, is a
stolon In biology, stolons (from Latin '' stolō'', genitive ''stolōnis'' – "branch"), also known as runners, are horizontal connections between organisms. They may be part of the organism, or of its skeleton; typically, animal stolons are external s ...
iferous,
deciduous In the fields of horticulture and Botany, the term ''deciduous'' () means "falling off at maturity" and "tending to fall off", in reference to trees and shrubs that seasonally shed leaves, usually in the autumn; to the shedding of petals, aft ...
, aromatic shrub in the
laurel family Lauraceae, or the laurels, is a plant family that includes the true laurel and its closest relatives. This family comprises about 2850 known species in about 45 genera worldwide (Christenhusz & Byng 2016 ). They are dicotyledons, and occur m ...
. This endangered species is native to the southeastern United States, and its demise is associated with habitat loss from extensive drainage of
wetland A wetland is a distinct ecosystem that is flooded or saturated by water, either permanently (for years or decades) or seasonally (for weeks or months). Flooding results in oxygen-free (anoxic) processes prevailing, especially in the soils. The ...
s for agriculture and forestry.US Fish and Wildlife Service: Species Recovery Plan: ''Lindera melissifolia''.
/ref>
/ref> Restoration efforts are currently being conducted.


Description

Form: Pondberry occurs in dense thickets with erect or ascending shoots up to tall and few branches; stems are connected underground by stolons. Thickets of female plants tend to be smaller than those of males and are sometimes absent from populations.
/ref> Die-back of stems is a fairly common occurrence. Foliage: The drooping, alternate leaves are oblong-elliptic to narrowly ovate, long, wide, and tend to be strongly tapered to a point at the tip.Patrick, T.S.; Allison, J.R.; Krakow, G.A..1995. Protected Plants of Georgia, An Information Manual on Plants Designated by the State of Georgia as Endangered, Threatened, Rare or Unusual. Georgia Department of Natural Resources.
/ref> Undersides are strongly net-veined and covered with short, soft hairs. When crushed, the leaves strongly resemble sassafras (''
Sassafras albidum ''Sassafras albidum'' (sassafras, white sassafras, red sassafras, or silky sassafras) is a species of ''Sassafras'' native to eastern North America, from southern Maine and southern Ontario west to Iowa, and south to central Florida and eastern T ...
'') in fragrance. Foliage is deciduous. Flowers: Stems flower after two to four years of growth.Devall, Margaret; Schiff, Nathan; Boyette, Douglas. 2001. Ecology and reproductive biology of the endangered pondberry, ''Lindera melissifolia'' (Walt) Blume. Natural Areas Journal. 21:250-258.
/ref> Male and female flowers, each across, are produced on different plants (
dioecious Dioecy (; ; adj. dioecious , ) is a characteristic of a species, meaning that it has distinct individual organisms (unisexual) that produce male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproductio ...
). Flowers appear before the leaves (February to mid-March) in tight, stalkless clusters. The petal-like
tepals A tepal is one of the outer parts of a flower (collectively the perianth). The term is used when these parts cannot easily be classified as either sepals or petals. This may be because the parts of the perianth are undifferentiated (i.e. of very ...
are pale to bright yellow, oblong, and long. Male flowers occur in dense clusters, with 9-12 stamens surrounded by two whorls of tepals. Female flowers are less conspicuous, with fewer flowers per cluster and a single pistil surrounded by two whorls of tepals; the outer whorl is petal-like and the inner whorl is reduced to nectar-producing scales. Flowers remain open for about 1 week making thickets conspicuous. Flowers are thought to be insect pollinated. Late season frosts occasionally damage flowers, resulting in reduced fruit set. Fruit: A bright red, single-seeded drupe, ellipsoid, long matures in late summer or fall (August to early October). Individual fruit stalks are long, thick, and appear swollen at the apex. Stalks persist beyond fruit fall; their presence indicates the plant's sex and past level of fruit production. Fruit production is highly variable from year-to-year, ranging from 0 to 150 fruit per stem.


Habitat and range

Pondberry occurs in shallow depression ponds in wetland habitats with
hydric soil Hydric soil is soil which is permanently or seasonally saturated by water, resulting in anaerobic conditions, as found in wetlands. Overview Most soils are aerobic. This is important because plant roots respire (that is, they consume oxygen and ...
s, along margins of cypress ponds, and in seasonally wet, low areas among bottomland hardwoods. At present there are some 36 populations in Alabama, Arkansas, Georgia, Mississippi, Missouri, North Carolina, and South Carolina.Hoyle, Zoe. Pondberry: Modest But Mysterious. Southern Research Station: Compass, Issue 6.
/ref> It has apparently been extirpated from Louisiana and possibly Florida. Most of these populations are located in Lower
Mississippi Alluvial Plain The Mississippi River Alluvial Plain is an alluvial plain created by the Mississippi River on which lie parts of seven U.S. states, from southern Louisiana to southern Illinois (Illinois, Missouri, Kentucky, Tennessee, Arkansas, Mississippi, Lou ...
, with the largest population being in the Delta National Forest in western Mississippi.


Ecology

Pondberry has probably always been a rare species, and knowledge of its ecology is limited. In Mississippi, pondberry occurs in bottomland hardwood forests. In northeastern Arkansas and southeastern Missouri pondberry is found on the bottoms and edges of shallow seasonal ponds in old dune fields, but in southeastern Arkansas it occurs in low habitat along a river. In South Carolina the species occurs in areas with
karst Karst is a topography formed from the dissolution of soluble rocks such as limestone, dolomite, and gypsum. It is characterized by underground drainage systems with sinkholes and caves. It has also been documented for more weathering-resistant ro ...
topography, around the edges of sinkholes, and in Georgia it occurs along the borders of sphagnum bogs. Sunlight at the different sites ranges from deep shade to almost full sun. Most pondberry colonies occur in light shade beneath a forest canopy, but a few grow in almost full sunlight. Pondberry appears to be able to occupy widely different habitats as long as its requirements for water are met. In open conditions, competition from other plant species may be a problem. Many of the remaining populations consist only of male plants and are apparently the sprouts of a single individual. Habitat fragmentation severely affects dioecious species like pondberry because populations with plants of a single sex can only vegetatively reproduce. With significant habitat loss, plants become ever more isolated, lessening the likelihood that pollinators will travel from male to female plants.


Propagation

Clones expand vegetatively through stolons, and this mechanism of
vegetative reproduction Vegetative reproduction (also known as vegetative propagation, vegetative multiplication or cloning) is any form of asexual reproduction occurring in plants in which a new plant grows from a fragment or cutting of the parent plant or spec ...
is the principal way that colonies develop. Stems usually live 6 or 7 years, and when a stem dies it is usually replaced by a new stem that grows from the base of the plant. Thus, mature colonies often include some dead stems intermingled with numerous live stems. Despite the regular production of mature fruit, virtually no
seedling A seedling is a young sporophyte developing out of a plant embryo from a seed. Seedling development starts with germination of the seed. A typical young seedling consists of three main parts: the radicle (embryonic root), the hypocotyl (embry ...
s of pondberry have been observed at any of the known sites. The cause of this apparent lack of natural reproduction is not currently known, but the consequences are clear—it severely reduces the species' chance for long-term survival. Sexual reproduction of pondberry is critical for long-range dispersal and genetic diversity. Before modern flood control was imposed along the Mississippi River and its tributaries, historic floods may have been an important mechanism in fruit and/or
seed dispersal In Spermatophyte plants, seed dispersal is the movement, spread or transport of seeds away from the parent plant. Plants have limited mobility and rely upon a variety of dispersal vectors to transport their seeds, including both abiotic vectors, ...
over long distances or for creating suitable conditions for seedling establishment. Although the fruit of pondberry sinks in water after a short time, the seed with the pulp removed will float for a day or sometimes longer. Characteristics of pondberry's fruit—the showy color, fleshy pulp, and its persistent on stems—suggest that animals, particularly birds, may be important dispersal agents. Of 82 bird species observed in the vicinity of pondberry thickets in fruit, only two species were observed to eat the fruit—hermit thrushes ('' Catharus guttatus'') and northern cardinals (''
Cardinalis cardinalis The northern cardinal (''Cardinalis cardinalis'') is a bird in the genus ''Cardinalis''; it is also known colloquially as the redbird, common cardinal, red cardinal, or just cardinal (which was its name prior to 1985). It can be found in southea ...
''). Of these, the cardinals destroyed the seed by crushing and is thus considered a seed predator. However, the hermit thrush swallows the whole fruit and later regurgitates the seed, indicating that it is an important
seed disperser In Spermatophyte plants, seed dispersal is the movement, spread or transport of seeds away from the parent plant. Plants have limited mobility and rely upon a variety of dispersal vectors to transport their seeds, including both abiotic vectors, ...
. The foraging habits of the thrush suggests that most of the seeds would be dispersed with 100 m of existing female colonies. Mammals may also be potential dispersers of pondberry seeds, including the raccoon (''
Procyon lotor The raccoon ( or , ''Procyon lotor''), sometimes called the common raccoon to distinguish it from other species, is a mammal native to North America. It is the largest of the procyonid family, having a body length of , and a body weight of ...
'') and opossum (''
Didelphis virginiana The Virginia opossum (''Didelphis virginiana''), also known as the North American opossum, is the only opossum living north of Mexico, its range extending south into Central America. It is the northernmost marsupial in the world. In the United ...
''). Historically, the black bear (''
Ursus americanus Ursus is Latin for bear. It may also refer to: Animals * ''Ursus'' (mammal), a genus of bears People * Ursus of Aosta, 6th-century evangelist * Ursus of Auxerre, 6th-century bishop * Ursus of Solothurn, 3rd-century martyr * Ursus (''praefectus ...
'') may have been important to seed dispersal. Seed germination does not appear to be a deterrent to seedling establishment. Fairly high rates of germination have been reported under both controlled and field conditions.Connor, Kristina; and others. 2006. A study of the early fruit characteristics of pondberry. Gen. Tech. Rep. SRS-92. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southern Research Station: 564-568.
/ref>Aleric, Katherine M; Kirkman, L. Katherine. 2005. Seed germination observations of the federally endangered ''Lindera melissifolia''. Castanea 70: 157-160. Removal of the fruit's pulp and sowing seeds into the soil favor germination. Under field conditions, germination of sown seeds has been observed to occur over a number of years suggesting some form of
dormancy Dormancy is a period in an organism's life cycle when growth, development, and (in animals) physical activity are temporarily stopped. This minimizes metabolic activity and therefore helps an organism to conserve energy. Dormancy tends to be clo ...
.Smith, Tim E. 2003. Observation on the experimental planting of ''Lindera melissifolia'' (Walter) Blume in southeastern Missouri after 10 years. Castanea 68: 75-80. The artificial establishment of pondberry to new areas may be needed if the species is to recover. Plants have been successfully transplanted from existing colonies to suitable locations. The transplants seem to do well in some locations although survivorship and growth has been low in others.Devall, Margaret S.; Schiff, Nathan M.; Skojac, Stephanie A. 2004 Outplanting of the Endangered Pondberry Gen. Tech. Rep. SRS–71. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southern Research Station: 574-577.
/ref> Both transplants and seedlings do well under cultivation in a nursery setting, which has been used to provide planting stock for creating new colonies in field locations.


Uses

Pondberry fruit are eaten by hermit thrushes, northern cardinals, and perhaps other birds. Animals observed to consume pondberry seeds located on a cleared soil surface in a hardwood forest included: northern cardinal, brown thrasher (''
Toxostoma rufum The brown thrasher (''Toxostoma rufum''), sometimes erroneously called the brown thrush or fox-coloured thrush, is a bird in the family Mimidae, which also includes the New World catbirds and mockingbirds. The brown thrasher is abundant through ...
''), swamp rabbit (''
Sylvilagus aquaticus The swamp rabbit (''Sylvilagus aquaticus''), also called the cane-cutter, is a large cottontail rabbit found in the swamps and wetlands of the southern United States. The species has a strong preference for wet areas, and it will take to the wate ...
''), armadillo (''
Dasypus novemcinctus The nine-banded armadillo (''Dasypus novemcinctus''), also known as the nine-banded long-nosed armadillo or common long-nosed armadillo, is a mammal found in North, Central, and South America, making it the most widespread of the armadillos. It ...
''), and gray squirrel (''
Sciurus carolinensis The eastern gray squirrel (''Sciurus carolinensis''), also known, particularly outside of North America, as simply the grey squirrel, is a tree squirrel in the genus ''Sciurus''. It is native to eastern North America, where it is the most prodi ...
'').Abilio, Fernanda Maria; and others. 2008. Pondberry (''Lindera melissifolia'') seed predators. IV Congreso Forestal Latinoamericano.
/ref> Swamp rabbits have been observed browsing on stems. The spicebush swallowtail (''
Papilio troilus ''Papilio troilus'', the spicebush swallowtail or green-clouded butterfly, is a common black swallowtail butterfly found in North America."Simply Butterflies," Accessed March 17, 2011, http://www.simplybutterflies.com/Backyard_Butterflies.html I ...
'') larvae feed on the leaves and roll themselves in a leaf making a tent. Leaf cutter bees (''
Megachilidae Megachilidae is a cosmopolitan family of mostly solitary bees. Both that their pollen-carrying structure (called a ''scopa'') is restricted to the ventral surface of the abdomen (rather than mostly or exclusively on the hind legs as in other b ...
'' spp.) cut circular sections from the leaf margins, sometimes removing most of the leaf. A vintage use of pondberry fruit in the rural South was as projectiles in toy pop guns constructed by children from hollowed-out elderberry (''
Sambucus canadensis ''Sambucus canadensis'', the American black elderberry, Canada elderberry, or common elderberry, is a species of elderberry native to a large area of North America east of the Rocky Mountains, south to Bolivia. It grows in a variety of conditions ...
'') stems.Devall, Margaret S.;Schiff, Nathan M. U.S. Forest Service. ''Lindera melissifolia'', Pondberry.
/ref>


Threats

A large part of pondberry habitat disappeared when forests were cut for timber or for conversion to agricultural fields, and as wetlands were drained. In some cases, wetlands were permanently flooded to construct lakes. Many of the existing colonies of pondberry are small, and occupy only a portion of the apparently suitable habitat. There are indications that pondberry, which is in the Laurel family, is susceptible to laurel wilt (''Raffaelea lauricola'')."Laurel Wilt", This fungal disease is introduced into host plants by a non-native insect, the redbay ambrosia beetle (''Xyleborus glabratus'') and kills the plant by plugging the water-conducting cells of the afflicted individual, causing it to wilt and eventually die. There is no known cure for this disease, which has quickly spread through other members of the laurel family (especially redbay, ''Persia borbonia'') from the coast of South Carolina inland towards the native habitat of pondberry.


Restoration

Pondberry was listed as endangered in 1986. The
U.S. Fish and Wildlife Service The United States Fish and Wildlife Service (USFWS or FWS) is an agency within the United States Department of the Interior dedicated to the management of fish, wildlife, and natural habitats. The mission of the agency is "working with othe ...
's restoration plan states: existing pondberry populations should be protected from forestry and agricultural management actions and protected from grazing and browsing animals; searches for new populations should be continued; areas where pondberry has not been located but provide suitable habitat should be protected; and, new populations should be established or reestablish extirpated populations at suitable sites. A critical part of the restoration effort is to increase the knowledge about pondberry's ecology and reproduction. A team of researchers from the
U.S. Forest Service The United States Forest Service (USFS) is an agency of the U.S. Department of Agriculture that administers the nation's 154 national forests and 20 national grasslands. The Forest Service manages of land. Major divisions of the agency in ...
's Southern Research Station are working with the
U.S. Army Corps of Engineers , colors = , anniversaries = 16 June (Organization Day) , battles = , battles_label = Wars , website = , commander1 = ...
and the U.S. Fish and Wildlife Service to learn as much as they can about the biology and ecology of pondberry. Scientists are investigating the role of flooding and light availability on pondberry at a large-scale impoundment facility, and they have set up controlled
experiment An experiment is a procedure carried out to support or refute a hypothesis, or determine the efficacy or likelihood of something previously untried. Experiments provide insight into Causality, cause-and-effect by demonstrating what outcome oc ...
s to study competition, seed germination, seed storage, and seed persistence in the
soil seed bank The soil seed bank is the natural storage of seeds, often dormant, within the soil of most ecosystems. The study of soil seed banks started in 1859 when Charles Darwin observed the emergence of seedlings using soil samples from the bottom of a lak ...
. An integrated approach is being used to learn more about pondberry's ecology, insect predators, fungal pathogens, physiological responses to light availability and flooding, population genetics, seed physiology, and seed dispersal.


References


External links


Lower Mississippi River Conservation Committee: Pondberry.


* ttps://web.archive.org/web/20090321002712/http://www.ncnhp.org/Images/97.pdf North Carolina Natural Heritage Program: Pondberry.
USGS: Southern Wetland Flora: Field Office Guide to Plant Species: Southern Spicebush.

United States Department of Agriculture Natural Resources Conservation Service: Plant Profile. Southern Spicebush.



Glitzenstein, Jeff. Native Plant Society and Forest Service re-invigorate endangered pondberry at Honey Hill. Journal of the South Carolina Native Plant Society. Fall 2007.
{{Taxonbar, from=Q6552425 melissifolia Endangered plants Flora of the Southeastern United States Plants described in 1788