The honey locust (''Gleditsia triacanthos''), also known as the thorny locust or thorny honeylocust, is a

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 ...

tree In botany, a tree is a perennial plant with an elongated stem, or trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, including only woody plants with secondary growth, plants that are ...

in the family Fabaceae, native to central

North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...

where it is mostly found in the moist soil of river valleys. Honey locust is highly adaptable to different environments, has been introduced worldwide, and can be an aggressive,

invasive species An invasive species otherwise known as an alien is an introduced organism that becomes overpopulated and harms its new environment. Although most introduced species are neutral or beneficial with respect to other species, invasive species ad ...

outside of its native range.

Description

The honey locust, ''Gleditsia triacanthos'', can reach a height of . They exhibit fast growth, but live a medium life span of about 120 years. The leaves are

pinnate Pinnation (also called pennation) is the arrangement of feather-like or multi-divided features arising from both sides of a common axis. Pinnation occurs in biological morphology, in crystals, such as some forms of ice or metal crystals, and in ...

compound Compound may refer to: Architecture and built environments * Compound (enclosure), a cluster of buildings having a shared purpose, usually inside a fence or wall ** Compound (fortification), a version of the above fortified with defensive struct ...

on older trees but

bipinnate The following is a list of terms which are used to describe leaf morphology in the description and taxonomy of plants. Leaves may be simple (a single leaf blade or lamina) or compound (with several leaflets). The edge of the leaf may be regular o ...

ly compound on vigorous young trees. The leaflets are (smaller on bipinnate leaves) and bright green. They turn yellow in the autumn. Honey locusts leaf out relatively late in spring, but generally slightly earlier than the

black locust ''Robinia pseudoacacia'', commonly known in its native territory as black locust, is a medium-sized hardwood deciduous tree, belonging to the tribe Robinieae of the legume family Fabaceae. It is endemic to a few small areas of the United States ...

(''

Robinia pseudoacacia ''Robinia pseudoacacia'', commonly known in its native territory as black locust, is a medium-sized hardwood deciduous tree, belonging to the tribe Robinieae of the legume family Fabaceae. It is endemic to a few small areas of the United States, ...

''). The strongly scented, cream-colored flowers appear in late spring, in clusters emerging from the base of the

leaf axil A leaf ( : leaves) is any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, ste ...

s. The trees are

polygamodioecious Plant reproductive morphology is the study of the physical form and structure (the morphology) of those parts of plants directly or indirectly concerned with sexual reproduction. Among all living organisms, flowers, which are the reproductive st ...

: most are strictly

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 ...

with male and female flowers on separate trees though some have bisexual flowers with a few male or female flowers on the same tree. The fruit of the honey locust is a flat

legume A legume () is a plant in the family Fabaceae (or Leguminosae), or the fruit or seed of such a plant. When used as a dry grain, the seed is also called a pulse. Legumes are grown agriculturally, primarily for human consumption, for livestock f ...

(pod) that matures in early autumn. The pods are generally between . The seeds are dispersed by grazing

herbivore A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...

s such as cattle and horses, which eat the pod pulp and excrete the seeds in droppings; the animal's digestive system assists in breaking down the hard seed coat, making germination easier. In addition, the seeds are released in the host's manure, providing fertilizer for them. Honey locust seed pods ripen in late spring and germinate rapidly when temperatures are warm enough. Honey locusts commonly have

thorns Thorn(s) or The Thorn(s) may refer to: Botany * Thorns, spines, and prickles, sharp structures on plants * ''Crataegus monogyna'', or common hawthorn, a plant species Comics and literature * Rose and Thorn, the two personalities of two DC Com ...

long growing out of the branches, some reaching lengths over ; these may be single, or branched into several points, and commonly form dense clusters. The thorns are fairly soft and green when young, harden and turn red as they age, then fade to ash grey and turn brittle when mature. These thorns are thought to have evolved to protect the trees from browsing

Pleistocene megafauna Pleistocene megafauna is the set of large animals that lived on Earth during the Pleistocene epoch. Pleistocene megafauna became extinct during the Quaternary extinction event resulting in substantial changes to ecosystems globally. The role of hu ...

, which may also have been involved in seed dispersal, but the size and spacing of them is less useful in defending against smaller extant herbivores such as deer. Thornless forms (''Gleditsia triacanthos'' var. ''inermis'') are occasionally found growing wild and are available as nursery plants. Hybridization of honey locust with

water locust ''Gleditsia aquatica'', commonly called water locust or swamp locust after its habitat of river swamps and slough margins, is a tree native to the Southeastern United States and adjacent regions. Description ''Gleditsia aquatica'' often grows 50 ...

(''G. aquatica'') has been reported. File:HoneyLocustThorn.JPG, Detail of thorns File:Honey Locust Tree Thorns in Kansas.jpg, Honey locust tree thorns in Kansas File:Gleditsia triacanthos Flower HW1.jpg, Detail of flowers File:2014-10-11 12 49 16 Honey Locust foliage during autumn in Elko, Nevada.JPG, Autumn leaf color File:Honeylocust fruits.jpg, Mature honeylocust fruits File:Old growth honeylocust tree.jpg, Old growth honeylocust tree in Tennessee, US, 1941

Cultivation and history

Its

cultivar A cultivar is a type of cultivated plant that people have selected for desired traits and when propagated retain those traits. Methods used to propagate cultivars include: division, root and stem cuttings, offsets, grafting, tissue culture, ...

s are popular

ornamental plant Ornamental plants or garden plants are plants that are primarily grown for their beauty but also for qualities such as scent or how they shape physical space. Many flowering plants and garden varieties tend to be specially bred cultivars that i ...

s, especially in the northern plains of North America where few other trees can survive and prosper. It tolerates urban conditions, compacted soil, road salt, alkaline soil, heat, and drought. The popularity is in part due to the fact that it transplants so easily. The fast growth rate and tolerance of poor site conditions make it valued in areas where shade is wanted quickly, such as new parks or housing developments, and in disturbed and reclaimed environments, such as mine tailings. Cultivars with narrow upright growth habit, such as ''Northern Sentinel,'' are especially prized as street trees. It is resistant to

spongy moth ''Lymantria dispar'', also known as the gypsy moth or the spongy moth, is a species of moth in the family Erebidae. ''Lymantria dispar'' is subdivided into several subspecies, with subspecies such as ''L. d. dispar'' and ''L. d. japonica'' bei ...

s but is defoliated by another pest, the

mimosa webworm ''Homadaula anisocentra'', also known as the mimosa webworm, is a species of moth in the family Galacticidae. It is considered a pest of ornamental plants. They attack the leaves of mimosa ('' Albizia julibrissin'') and honeylocust (''Gleditsia t ...

. Spider mites,

canker A plant canker is a small area of dead tissue, which grows slowly, often over years. Some cankers are of only minor consequence, but others are ultimately lethal and therefore can have major economic implications for agriculture and horticultur ...

s, and

gall Galls (from the Latin , 'oak-apple') or ''cecidia'' (from the Greek , anything gushing out) are a kind of swelling growth on the external tissues of plants, fungi, or animals. Plant galls are abnormal outgrowths of plant tissues, similar to be ...

s are a problem with some trees. Many cultivated varieties do not have thorns.

Agriculture

The species is a major invasive environmental and economic weed in agricultural regions of Australia. The plant forms thickets and destroys the pasture required for livestock to survive. The thickets choke waterways and prevent both domestic and native animals from drinking and also harbour

vermin Vermin (colloquially varmint(s) or varmit(s)) are pests or nuisance animals that spread diseases or destroy crops or livestock. Since the term is defined in relation to human activities, which species are included vary by region and enterpr ...

. The spines cause damage to both people and domestic and native wildlife and puncture vehicle tires. In much of the Midwest of the United States the honey locust is also considered a weed tree and a pest that establishes itself in farm fields. In other regions of the world, ranchers and farmers who employ

monocropping In agriculture, monocropping is the practice of growing a single crop year after year on the same land. Maize, soybeans, and wheat are three common crops often monocropped. Monocropping is also referred to as continuous cropping, as in "continuous ...

deem honey locust a nuisance weed; its fast growth allows it to out-compete grasses and other crops.

Uses

Food

The pulp on the inside of the pods is edible (unlike the

, which is toxic) and consumed by wildlife and livestock. Despite its name, the honey locust is not a significant

honey Honey is a sweet and viscous substance made by several bees, the best-known of which are honey bees. Honey is made and stored to nourish bee colonies. Bees produce honey by gathering and then refining the sugary secretions of plants (primar ...

plant. The name derives from the sweet taste of the

pulp, which was used for food and

traditional medicine Traditional medicine (also known as indigenous medicine or folk medicine) comprises medical aspects of traditional knowledge that developed over generations within the folk beliefs of various societies, including indigenous peoples, before the ...

Native American people Native Americans, also known as American Indians, First Americans, Indigenous Americans, and other terms, are the Indigenous peoples of the mainland United States (Indigenous peoples of Hawaii, Alaska and territories of the United States are ...

, and can also be used to make

tea Tea is an aromatic beverage prepared by pouring hot or boiling water over cured or fresh leaves of ''Camellia sinensis'', an evergreen shrub native to East Asia which probably originated in the borderlands of southwestern China and north ...

. The long pods, which eventually dry and ripen to brown or maroon, are surrounded in a tough, leathery skin that adheres strongly to the pulp within. The pulp—bright green in unripe pods—is strongly sweet, crisp and succulent in ripe pods. Dark brown tannin-rich beans are found in slots within the pulp. Likewise, its edible seed has nutritional potential, and the

flour Flour is a powder made by grinding raw grains, roots, beans, nuts, or seeds. Flours are used to make many different foods. Cereal flour, particularly wheat flour, is the main ingredient of bread, which is a staple food for many culture ...

made from its

cotyledon A cotyledon (; ; ; , gen. (), ) is a significant part of the embryo within the seed of a plant, and is defined as "the embryonic leaf in seed-bearing plants, one or more of which are the first to appear from a germinating seed." The numb ...

s constitutes a food source with various potential uses for pastry and bakery, among other gastronomic uses.

Timber

Honey locusts produce a high quality, durable wood that polishes well, but the tree does not grow in sufficient numbers to support a bulk industry. However, a niche market exists for honey locust furniture. It is also used for posts and rails because of the dense, rot-resistant nature of the wood. In the past, the hard thorns of the younger trees were used as nails and the wood itself was used to fashion

treenail A treenail, also trenail, trennel, or trunnel, is a wooden peg, pin, or dowel used to fasten pieces of wood together, especially in timber frames, covered bridges, wooden shipbuilding and boat building. It is driven into a hole bored through two ...

s for shipbuilding.

Nitrogen fixation

The ability of ''Gleditsia'' to fix nitrogen is disputed. Many scientific sources state that ''Gleditsia'' does not fix nitrogen. Some support this statement with the fact that ''Gleditsia'' does not form

root nodule Root nodules are found on the roots of plants, primarily legumes, that form a symbiosis with nitrogen-fixing bacteria. Under nitrogen-limiting conditions, capable plants form a symbiotic relationship with a host-specific strain of bacteria known a ...

s with

symbiotic bacteria Symbiotic bacteria are bacteria living in symbiosis with another organism or each other. For example, rhizobia living in root nodules of legumes provide nitrogen fixing activity for these plants. Symbiosis was first defined by Marko de Bary in 186 ...

, the assumption being that without nodulation, no nitrogen fixation can occur. In contrast, many popular sources,

permaculture Permaculture is an approach to land management and settlement design that adopts arrangements observed in flourishing natural ecosystems. It includes a set of design principles derived using whole-systems thinking. It applies these principle ...

publications in particular, claim that ''Gleditsia'' does fix nitrogen but by some other mechanism. There are anatomical, ecological, and taxonomic indications of nitrogen fixation in non-nodulating legumes. Both nodulating and non-nodulating species have been observed to grow well in nitrogen-poor soil with non-nodulating legumes even dominating some sites. The litter and seeds of non-nodulating species contain levels of nitrogen higher than non-legumes and sometimes even higher than nodulating legumes growing on the same site. How this happens is not yet well understood but there have been some observations of

nitrogenase Nitrogenases are enzymes () that are produced by certain bacteria, such as cyanobacteria (blue-green bacteria) and rhizobacteria. These enzymes are responsible for the Organic redox reaction, reduction of nitrogen (N2) to ammonia (NH3). Nitrog ...

activity in non-nodulating leguminous plants, including honey locust. Electron microscopy indicates the presence of clusters around the inner cortex of roots, just outside the xylem, that resemble colonies of rhizobial bacterioids. These may well constitute the evolutionary precursors in legumes for nitrogen fixation through nodulation. It is not known whether the non-nodulating nitrogen fixation, if it exists, benefits neighboring plants as is said to be the case with nodulating legumes.

Research

In research using databases, more than 60

phytochemical Phytochemicals are chemical compounds produced by plants, generally to help them resist fungi, bacteria and plant virus infections, and also consumption by insects and other animals. The name comes . Some phytochemicals have been used as poisons ...

s were identified from honey locust, including

polyphenol Polyphenols () are a large family of naturally occurring organic compounds characterized by multiples of phenol units. They are abundant in plants and structurally diverse. Polyphenols include flavonoids, tannic acid, and ellagitannin, some of ...

triterpene Triterpenes are a class of chemical compounds composed of three terpene units with the molecular formula C30H48; they may also be thought of as consisting of six isoprene units. Animals, plants and fungi all produce triterpenes, including squale ...

sterol Sterol is an organic compound with formula , whose molecule is derived from that of gonane by replacement of a hydrogen atom in position 3 by a hydroxyl group. It is therefore an alcohol of gonane. More generally, any compounds that contain the go ...

s, and

saponin Saponins (Latin "sapon", soap + "-in", one of), also selectively referred to as triterpene glycosides, are bitter-tasting usually toxic plant-derived organic chemicals that have a foamy quality when agitated in water. They are widely distributed ...

s, with

in vitro ''In vitro'' (meaning in glass, or ''in the glass'') studies are performed with microorganisms, cells, or biological molecules outside their normal biological context. Colloquially called "test-tube experiments", these studies in biology an ...

studies assessing for possible biological activity.

References

External links

*
''Gleditsia triacanthos'' images
at bioimages.vanderbilt.edu
''Gleditsia triacanthos'' images
at Forestry Images
''Gleditsia triacanthos''
at the USDA Plants Database {{Taxonbar, from=Q157650 Edible legumes Garden plants of North America Gleditsia Nearctic realm flora Ornamental trees Plants described in 1753 Plants used in Native American cuisine Plants used in traditional Native American medicine Trees of humid continental climate Trees of Ontario Trees of the Eastern United States Trees of the North-Central United States Trees of the Northeastern United States Trees of the Plains-Midwest (United States) Trees of the Southeastern United States Trees of the United States