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Ceratonia siliqua Phylogenetic Tree

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Leguminosae Node Chloe Fuson
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1. Ceratonia siliqua Phylogenetic Tree
Caesalpinioideae Subfamily Divergence Chloe Fuson
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1. Ceratonia siliqua Phylogenetic Tree
Umtiza Clade Chloe Fuson
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Missing Sister Species (C. oreothauma) Chloe Fuson
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Statistical Predictions Chloe Fuson
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Carob: From Pod to Pantry Chloe Fuson Ceratonia siliqua, commonly known as carob, St. John’s Bread, and locust bean, is a flowering deciduous tree in the legume family that produces large seed pods with a wide range of commercial uses. Carob has existed long before the start of agriculture, with origins across the Mediterranean basin dating back to the Paleolithic age. Recent phylogenetic data suggests domestication of carob occurred through localized selection of wild genotypes, with some long-distance westward dispersal events by the Romans or Arabs. Carob trees are valued ecologically for their low maintenance, high adaptability to poor soil conditions, and drought resistance, with some even living for 100 years or more. They feature thick, dark green and broad foliage, thus ornamental trees can be found throughout California, Australia, and other parts of the world with Mediterranean climate. However the main attraction of Ceratonia siliqua is its brown, leathery pod. The carob pod can be split into two main components: the pulp and the seeds. The naturally sweet pulp has a sugar composition of up to 48-56%, though it also contains cyclitols, fiber, and polyphenols such as tannins. Carob pulp is a nutritionally significant source of amino acids, minerals, and vitamins. The seeds contain a galactomannan gum also known as carob bean gum (CBG) or locust bean gum (LBG) used as a thickening agent in many food products. Carob pods are associated with the prevention and treatment of a wide variety of diseases, including diabetes, hyperlipidemia, irritable bowel syndrome, and colon cancer. In the past few decades carob has been marketed simply as a healthier chocolate alternative, but its rich history, health benefits, and potential applications in areas affected by climate change should allow it to stand on its own as a staple in the pantry and beyond. By: Chloe Fuson and Skylar Yee #Carob
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Carob: From Pod to Pantry Ceratonia siliqua, commonly known as carob, St. John’s Bread, and locust bean, is a flowering deciduous tree in the legume family that produces large seed pods with a wide range of commercial uses. Carob has existed long before the start of agriculture, with origins across the Mediterranean basin dating back to the Paleolithic age. Recent phylogenetic data suggests domestication of carob occurred through localized selection of wild genotypes, with some long-distance westward dispersal events by the Romans or Arabs. Carob trees are valued ecologically for their low maintenance, high adaptability to poor soil conditions, and drought resistance, with some even living for 100 years or more. They feature thick, dark green and broad foliage, thus ornamental trees can be found throughout California, Australia, and other parts of the world with Mediterranean climate. However the main attraction of Ceratonia siliqua is its brown, leathery pod. The carob pod can be split into two main components: the pulp and the seeds. The naturally sweet pulp has a sugar composition of up to 48-56%, though it also contains cyclitols, fiber, and polyphenols such as tannins. Carob pulp is a nutritionally significant source of amino acids, minerals, and vitamins. The seeds contain a galactomannan gum also known as carob bean gum (CBG) or locust bean gum (LBG) used as a thickening agent in many food products. Carob pods are associated with the prevention and treatment of a wide variety of diseases, including diabetes, hyperlipidemia, irritable bowel syndrome, and colon cancer. In the past few decades carob has been marketed simply as a healthier chocolate alternative, but its rich history, health benefits, and potential applications in areas affected by climate change should allow it to stand on its own as a staple in the pantry and beyond. By: Chloe Fuson and Skylar Yee #Carob
Nomenclatural History
The scientific name Ceratonia siliqua L. is derived from the Greek work keras, meaning “horn”, and the Latin word siliqua, which refers to the shape and fitness of the pod. And, of course, the L. is in reference to Carl Linnaeus. The binomial name was first recorded by Linnaeus in Species Plantarum 2 in May 1753, with a locality of Apulia and Sicily (Italy), Crete (Greece), Cyprus, Syria, and Palestine. The designated lectotype is located at the Linnean Society of London Herbarium (Missouri Botanical Garden, n.d.). The common name for this plant is carob, and has its roots in ancient Mediterranean/Middle Eastern cultures. Carob originates from the Hebrew word kharuv, which means “locust”. This relates to two other common colloquial names for carob: St. John’s bread and locust bean. These two names stem from the biblical references to St. John the Baptist’s consumption of locusts during his travels. The “locusts” he was consuming were likely actually the pods from carob trees, hence the adoption of common names that have to do with the word locust. As the carob tree spread throughout the Mediterranean and other parts of the globe, other names were adopted in the new regions, such as the Arabic kharrūb, Spanish algarrobo or garrofero, Italian carrubo, and many more (Batlle & Tous, 1997, p. 9).
Evolution & Domestication
Evolutionary History
Ceratonia siliqua is a member of the Leguminosae family, the third largest angiosperm family and possibly originating in the late Cretaceous period (110.5-66 mya). (Wojciechowski et al, 2004 ). Within this classification, carob belongs to the subfamily Caesalpinioideae which, along with the other two Leguminosae clades (Mimosoideae and Papilionoideae), is evidenced to have been present since the Eocene (55.8-33.9 mya) (Wojciechowski et al, 2004). C. siliqua is then a member of the Caesalpinioideae subfamily that diverged even further into a smaller group known as the Umtiza Clade, to which C. siliqua belongs. The use of fossil records has allowed researchers to estimate that the Umtiza Clade was likely present as far back as the Late Paleocene, dating it to about 56.5 mya, making it one of the oldest Caesalpinioid clades. (Bruneau et al, 2008).
It was long thought that C. siliqua was the only member of the genus Ceratonia, but in the mid-1900s another species of the Ceratonia genus was discovered: C. oreothauma, which is native to Arabia and Somalia. C. oreothauma has been proposed as the potential wild ancestor of the cultivated C. siliqua due to its smaller, less evolved pollen form. It is distinct from the cultivated species since its flowers are strictly unisexual. (Batlle & Tous, 1997, p. 9, 14).
Domestication History
The origin and domestication history of carob remains unclear. Paleobotanical and archaeological evidence of pollen grains, seeds, and wood charcoal dating back to the Paleolithic (as far as 40,000 BCE) and Neolithic ages led researchers to suspect that carob originated in the East Mediterranean basin long before the start of agriculture. Notably, little to no prehistoric evidence has been found in other parts of the Mediterranean basin. Linguistic evidence and historical literature led to the proposition that the domestication and spread of carob originated in the East Mediterranean and occurred with the introduction of scion grafting during Roman times (Zohary, 2002).
However, comprehensive phylogeographic research based on genetic data done over the past several years has deduced that the carob likely originated in a biogeographic region in southwest Morocco. Overall, it was found that carob domestication has been propelled mostly by the local selection and dissemination of wild genotypes, with some long-distance westward dispersal events of domesticated varieties by humans (likely Romans or Arabs), in contrast to what was previously proposed. Plastid data revealed a low human influence on the main current patterns of genetic diversity and structure of the carob tree across the Mediterranean. Genome-wide diversity and phylogenetic reconstructions have revealed that the carob tree followed two main routes of migration from its origin in southern Morocco. One migration went northward, reaching western north Africa and south Spain. The other migration path went east and gave rise to central-eastern carob evolutionary units, or geographically homogenous genetic groups of carob populations (Baumel et al., 2021).
Though researchers are still uncovering the origins and history of carob domestication, its spread in recent times is known. In the early to mid-1800s, it was introduced to Mediterranean-like regions such as California, Arizona, Mexico, Chile, and Argentina by Spanish missionaries, to Australia by Mediterranean emigrants, and to South Africa and India by the English. Cultivars of carob were brought on a larger scale to the US and California in particular in 1854 by the US Patent Patent Office and again in the 1950s to San Diego County (Batlle & Tous, 1997, p. 7, 20, 21).
Ecology & Life History
Carobs are well suited to warm temperate or subtropical Mediterranean areas with temperatures ranging from 30-40°C, and can tolerate more hot and humid coastal climates. Consequently they are sensitive to the cold. Their main limitation is their low frost-tolerance, as young trees can be killed by frosts reaching temperatures of at least -4°C or 24.8°F. They can adapt to a wide range of soils including poor, sandy soils, rocky, calcareous hillsides, or deep soils. They are highly drought resistant, though like all crops require irrigation if grown commercially (Batlle & Tous, 1997, p. 48, 49, 63). In the wild, carob shelter, foliage, and pods attract browsing animals, which in turn spread seeds to other areas (Batlle & Tous, 1997, p. 26). In California, the carob tree thrives in areas such as the Imperial and Coachella Valleys that experience extreme heat and mild winters. In southern Europe it has been reported that carob thrives when in vicinity of the sea, but this does not hold in California, where it has been observed that trees closer to the coast do not produce as much fruit (Condit et al., 1919).
It generally takes about 6-7 years before a carob tree produces its first seed crop, so they do not necessarily begin making consumable products right away. However, they have great longevity, living and producing for 100 years or more (LEAF Network, 2016). Carobs are evergreen trees, so they do not have a season when they lose their leaves. Interestingly, carob is the only Mediterranean tree with a fall flowering season. The main flowering season of the carob is in autumn, usually from about September to November, sometimes longer. Further, this time and length can differ depending on local climate conditions These flowers are then shed mainly from October to December, slowing during January to February, and then almost never occurring from June to August (Batlle & Tous, 1997, p. 16, 18).
Pods normally ripen from October to December, remaining on the tree for weeks if they are not gathered (Condit et al., 1919). If one is interested in harvesting the pods to be used for their moist pulp, they should be picked in the spring while they are still green and not yet fully ripe because at this stage the pulp will still be very sweet. On the other hand, if the desire is for the pods to be milled and made into a powder, then one should observe the tree in late summer and harvest pods that easily snap, indicating that they are dry enough to be milled (“Carob- …”, 2022).
Morphology
Vegetative Morphology
Carob is a sclerophyllous evergreen shrub or tree with a thick trunk with rough bark, and sturdy branches that may grow up to approximately 10 m high, though wild trees may grow a few meters taller (Batlle & Tous, 1997, p. 10). Some features of sclerophyllous trees include hard, leathery leaves and short internodal distances, which allow them to adapt to long periods of drought and heat (Alm, 2016). The leaves of the carob tree are 10-20 cm long, alternate, pinnately compound with 4-10 oppositely arranged pairs of leaflets, with or without a terminal leaflet. Each leaflet is 3-7 cm long, ovate or elliptic, dark green on the top with slightly undulate margins. The sclerophyllous leaves have a thick epidermis layer to protect from water loss (Batlle & Tous, 1997, p. 10). Carob trees also have a broad, semi-spherical crown that requires low maintenance and are valued as an ornamental shade tree throughout California, Australia, and elsewhere. However, their strong, extensive root system that allows them to adapt to dry conditions in the wild are considered invasive in landscaping (Batlle & Tous, 1997, p. 28).

Reproductive Morphology
Most carob trees are dioecious with female and male flowers on separate trees, though a small number are hermaphroditic. The prevalence of flower types varies by region. For example, hermaphrodite trees are found more frequently in the Mediterranean Spanish coast than in Southern Spain, though they are still rare with respect to male and female trees. Generally, the flowers are 6-12 mm long, arranged along an inflorescence axis in racemes, with coloration that ranges from green to red. They display radial symmetry with no petals and a disc-shaped calyx consisting of 5 hairy sepals. Flowers typically begin bisexual but one sex is repressed during later development, hence the presence of rudimentary stamens or pistils on females or males, respectively. All three types have a nectarial disc, though females produce the highest volume of nectar (Batlle & Tous, 1997, p. 12).
Female flowers feature a fully developed pistil that is 6-8.5 cm long with 2 stigmatic lobes, 2 carpals, and 5 rudimentary stamens. Male flowers have 5 stamens and a central rudimentary pistil. The filaments may either be of either short or long type, while the anthers may range from yellow to red. Hermaphrodite flowers are a combination of both male and female types with a fully developed pistil and 5 fully developed stamens (Batlle & Tous, 1997, p. 12, 14).
The carob fruit is an indehiscent pod or legume 10-30 cm long, 1.5-3.5 cm wide, and about 1 cm thick; they are compressed, with a straight or curved shape depending on the cultivar. Pods are soft and green but become hard, brown, and leathery upon ripening. Carob pods have two main components: the pulp and the seeds. The pulp consists of an outer, dark and leathery pericarp and an inner, soft mesocarp that separates the seeds. The seeds range from 8-10 mm long, 7-8 mm wide, and 3-5 mm thick. They are composed of a hard external coat, an embryo/germ, and the endosperm (Batlle & Tous, 1997, p. 12, 24).
Cultivation Practices
Cultivars
There are less than 50 cultivars described in the world, many of which are locally distributed. Over 18 carob cultivars are grown throughout Spain, while others are located in Italy, Portugal, Morocco, Greece, Cyprus, Turkey, Israel, USA, and Australia. Regional cultivars show a high degree of genetic variation for traits such as pod size, shape, and quality, seed yield, pest and disease resistance, productivity, sugar content, or gum content. However, most commercially grown cultivars have traditionally been selected for a large pod size as well as a high pulp and sugar content, which distinguishes them from their wild counterparts. Pulp and seed content are negatively correlated so growers have selected against seed yield. The most common cultivars in orchards are female, as hermaphrodites are never the main producing trees. Thus, male and hermaphrodite trees are typically planted in orchards as sources of pollen (Batlle & Tous, 1997, p. 22, 30-32).
Cultivation Methods
Going hand in hand with the carob’s success in semi-arid climates, carob trees prefer well-drained soil. The maximum precipitation requirement for carob trees is 20-23 inches, but they can thrive and produce high yields when receiving as low as 12-14 inches of rain (Hills, 1980). With that said, even though the tree does not need much water, best results are achieved when some additional irrigation water is provided.
Like many other cultivated fruit trees, domesticated carob trees are typically propagated asexually as opposed to the sexual cross-pollination that occurs in wild counterparts (Batlle & Tous, 1997, p. 22). Carob may be propagated by seed, budding, veneer grafting, cutting, or air layering. However carob seeds have a very hard coat which must be removed in order for germination to occur successfully. Seedlings are used as rootstocks for grafting, which is currently the most common method of propagation. The seedling rootstocks should be budded 1 year after germination in a nursery or 2 years after being planted in an orchard. Vegetative propagation with cuttings is not commercially available, and they have been described as extremely difficult to root (Gubbuk et al., 2011).
Harvesting Methods
Carob pods may be harvested at different times of the year depending on the cultivar and their intended post-harvest use. Most are harvested at the end of summer or beginning of autumn (Batlle & Tous, 1997, p. 60). However, the methods for gathering the pods are pretty much the same and are fairly simplistic though they require much labor. In order to gather the pods from the trees, they should be prodded off the tree with a pole and are usually collected in nets or sheeting placed under the tree to catch them as they fall on the ground. If the harvested pods are dried properly, they can be stored for up to a year. One simply must lay them out in the sun and some sort of sheet for about two days to lower the moisture content. Then it is simply a matter of storing the pods in dry conditions while they await milling or any other intended use (“Carob”, 2022).
Traditionally, carob orchards were hardly fertilized as carob is extremely well-adapted to poor soils. However in more recent times, mineral or organic fertilizers have been applied (Batlle & Tous, 1997, p. 57). Intercropping with other early-bearing fruit and nut trees such as peach or almond are sometimes implemented in orchard design. Unlike many other fruit and nut trees, carob trees require minimal pruning after the basic framework of the tree is established. However, carob has a slow growth rate with a long vegetative phase (Batlle & Tous, 1997, p. 54, 55, 57). In fact, carob trees do not reach their full bearing production until they are around 20-25 years old, after which pod yield stabilizes. Additionally carob shows biennial or alternate bearing, meaning each year it alternates between a high yield or a low yield (Batlle & Tous, 1997, p. 59).
Consumption & Nutritional Value
Carob is commonly known and utilized as a cacao substitute to its high sugar content (48-56%) and lack of theobromine and caffeine. Currently, the main use is for galactomannan gum extraction. However, carob pods are an excellent source of bioactive compounds and contain low levels of fat (0.2-0.6%). To get a better idea of the nutritional value carob offers and its wide variety of uses in food, it’s convenient to split it into its two components. As mentioned previously, the carob fruit can be split upon crushing the pod into the pulp and the seeds, which approximately make up 90% and 10% of the pod’s weight, respectively (Batlle & Tous, 1997, p. 23).
Pulp
The pulp can be ground into large pieces called kibbles, or ground into powder. Kibbles are typically used for animal feed, carob syrup or liqueur, while the powder is used as a cacao substitute or flour for baked goods or sweets. The pulp can also be roasted whole and eaten as a snack (Batlle & Tous, 1997, p. 26-27). Carob pulp is high in sugar content (48-56%), but also contains cyclitols, fiber, polyphenols such as tannins, amino acids, minerals, and vitamins. The main cyclitol in the pulp is D-pinitol, which has anti-diabetic effects due to the compound's ability to increase insulin sensitivity and regulate blood sugar levels. The fiber and polyphenols in carob are also known to enhance lipid metabolism, lower total and LDL cholesterol to counter hyperlipidemia, and produce anticancer effects that involve anti-tumor, anti-proliferative, and proapoptotic activity. The main polyphenols are tannins, which have anti-diarrheal effects. Carob pulp is also considered a good source of amino acids by the World Health Organization as it contains all 7 essential amino acids at levels that meet their standards. Carob is also a strong source of calcium, and potassium, though it also contains other nutritionally important minerals such as iron, copper, zinc, or manganese (Goulas et al., 2016). Some prominent vitamins in carob pulp include vitamin E, D, C, niacin (B3), B6, and folic acid (B9), with lower levels of A, B2, and B12 (Papaefstathiou et al., 2018).
Seeds
Upon de-hulling the seeds, they are composed of the embryo, which is used in germ meal, and the endosperm. The endosperm is ground into carob bean gum (CBG) also known as locust bean gum (LBG), which is the most common use of commercially grown carob. LBG is used as a stabilizer, thickener, or a binder and gelling agent in a wide variety of food products such as ice creams, soups, sauces, cheese, pies, canned meats, or pet food (Batlle & Tous, 1997, p. 26-28). Germ meal is high in protein content (50%), while LBG is also used for pharmaceutical purposes due to its inhibition of gastrointestinal diseases like gastroesophageal reflux (Batlle & Tous, 1997, p. 24; Goulas et al., 2016).

Ultimately, carob pods are associated with the prevention and treatment of a wide variety of diseases, including diabetes, hyperlipidemia, irritable bowel syndrome, and colon cancer. More clinical research on its mechanisms and benefits in humans is needed, though the results of previous and ongoing studies are encouraging (Goulas et al., 2016).
Human Experience
Historical Applications
As mentioned earlier, carob is a very ancient species, with not only genetic and fossil evidence of its existence, but also cultural evidence as well. For example, carob was present in Mesopotamian culture and was even referenced in “The Epic of Gilgamesh”, the oldest existing work of literature known to exist. Part of the Epic tells of Gilgamesh’s quest to find Utnapishtim, a once-mortal man who was made immortal by the gods, and learn the secret to eternal life. Part of his journey involves venturing through a tunnel that runs under Mount Mashu and emerging into the “Garden of the Gods”. This garden is full of “jewel trees” that bear precious stones instead of fruit, and in certain versions of the story, one of the types of trees he encountered in this garden was a carob with “abashmu-stone, agate and haematite” (Daniel, 2012).
This mention of the carob tree in the “Epic of Gilgamesh” is only one example of carob’s rich history. Carob also has strong historical roots in ancient Egyptian culture. Carob pods and seeds have been found in Egyptian tombs, while the gummy pectin from carob seeds was used in the process of mummification (Cretacarob, 2020). Additionally, the Egyptian hieroglyph meaning “sweet” is a depiction of a carob pod (Vakulenko, 2019). Additionally, most everyone has heard of the term “carat” that is used as a measurement for precious stones, but what you may not know is that this term actually traces back to the Middle Eastern practice of weighing precious gemstones against carob seeds as a measure of purity (Vakulenko, 2019). Because carob seeds are generally very uniform in size and weight (about 0.20 g), they served as the perfect reliable object to which items of unknown mass could be compared. Thus, the term “carat” was created as a unit of measurement and has endured to this day.
Future Applications
The carob tree has great potential for expanded utilization in the future. As aforementioned, carob trees are suited to dry, arid regions and require very little water to survive and produce fruit. This makes them the ideal candidate for application as an ecosystem-restoring plant and combatant to the effects of climate change. Researchers have already begun to see this potential and have started looking into the prospects of using carob trees to restore degraded and marginalized lands in the Mediterranean. According to one study, an estimated 1.32 million km² of land in the Mediterranean region is undergoing desertification and groundwater reserves are significantly diminishing (Winer, 1980). This is where carob would come into play. Carob trees require light to no irrigation, increase soil moisture levels, reduce surface water runoff and soil erosion, and, as a perk, offer economic incentives as they produce fruit, timber, and livestock fodder (Winer, 1980). As the effects of global warming and climate change continue to grow and affect more and more regions, carob trees offer a very feasible option for habitat restoration in regions with low water supply.
In addition to needing little water, carob trees are also well-suited for dry regions because they act as a very effective fire barrier. Because the carob tree is an evergreen species, their xylems contain large quantities of water in order to keep the plant alive during dry seasons. This, along with its thick, leathery sclerophyllous leaves, makes it more difficult to catch fire and burn, and thus a great tree for areas prone to wildfires (Srečec et al, 2017). It is a very well known fact that the frequency and severity of wildfires across the globe -including here in California- are increasing significantly. Thus, the idea of planting less flammable species in fire-prone regions is something that should be taken into great consideration and is another example of a future use for carob trees.
Further, while carob trees are currently being taken advantage of for a wide variety of commercial uses, there is still room for an expanded market. Over the past fifty years, carob production worldwide has decreased quite significantly, due partly to low pod prices making the cultivation less economically worthwhile (“Carob …”, 2022). World production of carob is around 310,000 tons and is mainly isolated to countries in the Mediterranean region (Spain, Italy, Portugal, Morocco, Greece, etc.), with a few other scattered producing regions in locations such as Australia, California, and South Africa (“Carob …”, 2022). However, popularity of carob products is beginning to rise as more manufacturers and consumers are becoming aware of its health benefits and plethora of commercial applications, so this leaves the door open to market expansion and may hopefully lead to an increase in carob tree cultivation in the coming years.
References
1. Alm, M. (2016, September 2). Sclerophyllous plants. What are sclerophyllous plants. Retrieved May 26, 2022, from https://www.jardineriaon.com/en/sclerophyll-plants.html
2. Batlle, I., & Tous, J. (1997). Carob tree: Ceratonia Siliqua L. International Plant Genetic Resources Institute.
3. Baumel, A., Feliner, G. N., Medail, F., Malfa, S. L., Diguardo, M., Boudagher-Kharrat, M., Mirleau, F., Frelon, V., Ouahmane, L., Diadema, K., Sanguin, H., & Viruel, J. (2021). Genome-wide footprints in the carob tree (Ceratonia Siliqua) unveil a new domestication pattern of fruit trees in the Mediterranean. https://doi.org/10.22541/au.163344873.38613513/v1
4. Bruneau, A., Mercure, M., Lewis, G. P., & Herendeen, P. S. (2008). Phylogenetic patterns and diversification in the caesalpinioid legumes. Botany, 86(7), 697-718. https://doi.org/10.1139/B08-058
5. Carob- An Ancient Food Tree. (2022). Gardening in South Africa. Retrieved from https://www.gardeninginsouthafrica.co.za/carob-an-ancient-food-tree?highlight=WyJjYXJvYiIsInRyZWUiLCJ0cmVlJyIsInRyZWUnLCIsInRyZWUncyIsIid0cmVlIiwiY2Fyb2IgdHJlZSJd
6. Condit, I. J., Jaffa, M. E., Albro, F. W. . (1919). The carob in California. Berkeley, Calif.: University of California Press.
7. Cretacarob (2020). Carob: The Black Gold of History. Retrieved from https://cretacarob.com/en/blog/news/to-charoypi-o-mayros-chrysos-tis-istorias/
8. Daniel, Mark (2012). Inheritance: Covenants, Kingdoms, Bodies and Nations. (n.d.). (n.p.): Lulu.com, 45
9. Goulas, V., Stylos, E., Chatziathanasiadou, M. V., Mavromoustakos, T., & Tzakos, A. G. (2016). Functional Components of Carob Fruit: Linking the Chemical and Biological Space. International journal of molecular sciences, 17(11), 1875. https://doi.org/10.3390/ijms17111875
10. Gubbuk, H., Gunes, E., Ayala-Silva, T., & Ercisli, S. (2011). Rapid vegetative propagation method for carob. Notulae Botanicae Horti Agrobotanici Cluj-Napoca, 39(1), 251–254. https://doi.org/10.15835/nbha3916074
11. Hills, L. D. (1980) The Cultivation Of The Carob Tree (Ceratonia Siliqua), International Tree Crops Journal, 1:1, 27-36, https://doi.org/10.1080/01435698.1980.9752712
12. LEAF Network. (November 2016) Carob: Ceratonia siliqua L. https://leafnetworkaz.org/resources/PLANT%20PROFILES/Carob_profile.pdf
13. Missouri Botanical Garden. (n.d.). Tropicos. Retrieved May 26, 2022, from https://tropicos.org/name/13028551
14. Papaefstathiou, E., Agapiou, A., Giannopoulos, S., & Kokkinofta, R. (2018). Nutritional characterization of carobs and traditional carob products. Food Science & Nutrition, 6(8), 2151–2161. https://doi.org/10.1002/fsn3.776
15. Srečec, S., Kremer, D., Karlović, K., Purgar, D. D., & Erhatić, R. (2017). Possible Role of Carob Tree (ceratonia siliqua L.) in Fire Protection of Agro-Forest Systems of Croatian South Adriatic Islands Regarding the Similarities With Other Mediterranean Countries. Sofia: Surveying Geology & Mining Ecology Management (SGEM). https://doi.org/10.5593/sgem2017H/33
16. Vakulenko, Galyna (2019). Tree Spotlight: Carob tree. Canopy Blog. Retrieved from https://canopy.org/blog/tree-spotlight-carob-tree/
17. Winer, N. (1980) The Potential of the Carob (Ceratonia siliqua), International Tree Crops Journal, 1:1, 15-26, https://doi.org/10.1080/01435698.1980.9752711
18. Wojciechowski, M.F., Lavin, M. and Sanderson, M.J. (2004), A phylogeny of legumes (Leguminosae) based on analysis of the plastid matK gene resolves many well-supported subclades within the family. Am. J. Bot., 91: 1846-1862. https://doi.org/10.3732/ajb.91.11.1846
19. Zohary, D. (2002). Domestication of the carob (Ceratonia siliqua L.). Israel Journal of Plant Sciences, 50(1). https://doi.org/10.1560/bw6b-4m9p-u2ua-c6nn

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Nomenclatural History

Evolution & Domestication

Ecology & Life History

Morphology

Cultivation Practices

Consumption & Nutritional Value

Human Experience

References