Biology:Juglandaceae

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The Juglandaceae are an angiosperm family known as the walnuts. They are trees, or sometimes shrubs, in the order Fagales. Members of this family are native to the Americas, Eurasia, and Southeast Asia.

The nine or ten genera in the family have a total of around 63 species,[1] and include the commercially important nut-producing trees walnut (Juglans), pecan (Carya illinoinensis), and hickory (Carya). The Persian walnut, Juglans regia, is one of the major nut crops of the world. Walnut, hickory, and gaulin are also valuable timber trees while pecan wood is also valued as cooking fuel.

The genetic relationships between different members of the family have recently been studied in more detail.[1][2][3][4]

Description

Members of the walnut family are trees (rarely large shrubs)[5] with aromatic leaves that are usually alternate, but opposite in Alfaroa and Oreomunnea. The leaves are pinnately compound or ternate, and usually 20–100 cm long. The trees are wind-pollinated, and both the male and female the flowers are usually arranged in catkins. [6][5]Members of this family are often monoecious, but can be dioecious. [5][7]

The fruits of the Juglandaceae are often confused with drupes but are accessory fruit because the outer covering of the fruit is technically an involucre and thus not morphologically part of the carpel; this means it cannot be a drupe but is instead a drupe-like nut.[6]

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Catkin of Juglans cinera (White walnut or Butternut)

Ecology

Climate change

Members of Juglandaceae have been projected to experience a range shift as the climate changes.[8] Species distribution models show that within Europe Juglans regia will experience a northward shift to stay within a climatically suitable habitat. Different climate scenarios were modeled, revealing that the severity of range shift will increase with greater temperature changes. Expansion could extend beyond the native range, making this species invasive.[8]

Some members of Juglandaceae have been documented to have formed associations with both ectomycorrhizal fungi and arbuscular mycorrhizal fungi. [2] Dual mycorrhizal associations and their functional significance are still being studied, but they may confer ecological advantages.[9] Within Juglandaceae, these relationships remain an ongoing area of research.[2]

Biogeography

Juglandaceae distribution ranges from the temperate regions of North American, Europe, and Asia into the tropical regions of Asia and parts of Central and South America. [10]The subfamily Juglandoideae is thought to have originated and rapidly diversified in the early Cenozoic in North America. The history of the distribution of this family was determined through phylogenetic studies and the fossil record.[10][11] It then spread to Asia and Europe across the Bering and North Atlantic land bridges.[1]

Seed dispersal

Seeds are dispersed by rodents (mostly squirrels) and members of the crow family. Squirrels will bury nuts for winter and forget where they put them, aiding in seed dispersal. Humans have also dispersed seeds for farming purposes.[12]

Toxicity

Members of this family, particularly the Black walnut (Juglans nigara) produce juglone (5 hydroxy-1, 4-napthoquinone), a toxic chemical compound responsible for hindering the growth of surrounding plants. This is present in most parts of the plants that produce it. The kernel of the nut is edible.[13][14]

Taxonomy

The known living genera are grouped into subfamilies, tribes, and subtribes as follows:[3]

Systematics

Modern molecular phylogenetics suggest the following relationships:[11][17]

 
 

Myricaceae (outgroup)

 
Juglandaceae
Rhoipteleoideae

Rhoiptelea

 
 
Engelhardioideae
 

Engelhardia

 
 
 

Alfaroa

 
 

Oreomunnea

 
 
 
Juglandoideae
 
 

Carya

 
 

Annamocarya

 
 
 
 

Platycarya

 
 
 

Cyclocarya

 
 
 

Pterocarya

 
 

Juglans

 
 
 
 
 
 
 
 

Recent research

Recent molecular studies, by analyzing chloroplast and nuclear DNA sequences, have clarified the relationships in Juglandaceae.[17] There is evidence that whole genome duplication happened in an ancestral species.[18] Studies have shown there to be two new species within Engelhardia. These two species are Engelhardia anminiana H.H Meng 2018 and Engelhardia borneensis H.H Meng 2018.[4]

Human uses

Food

Members of this family, such as walnuts (Juglans) and pecans (Carya illinoinensis) are often farmed for their fruits which are eaten across the world. The United States of America produces the most pecans, followed by Mexico.[19] China produces the majority of the worlds walnuts, followed by the United States of America. [1] Sap from the hickory trees (Carya) was collected by Native-Americans to be used as a sweetener. [20]

Timber

Walnut wood is generally known for its rich dark brown color, which makes it popular amongst woodworkers. It is often used for smaller wooden objects, such as cabinets and furniture. The wood of butternut (Juglans cinerea) is usually a lighter tan color.[21]

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This is a jewelry box made out of black walnut wood

Hickory (Carya) wood is also popular. It is known for being flexible, making it often used to make various sporting equipment. Bows, ax handles, skis, and lacrosse sticks are commonly made out of hickory wood. Its shock resistant qualities made it popular amongst the American pioneers for building their cabins.[20]

References

  1. ↑ 1.0 1.1 1.2 Mu, Xian-Yun; Tong, Ling; Sun, Miao; Zhu, Yi-Xuan; Wen, Jun; Lin, Qin-Wen; Liu, Bing (2020-06-01). "Phylogeny and divergence time estimation of the walnut family (Juglandaceae) based on nuclear RAD-Seq and chloroplast genome data". Molecular Phylogenetics and Evolution 147. doi:10.1016/j.ympev.2020.106802. ISSN 1055-7903. https://www.sciencedirect.com/science/article/pii/S1055790320300749. 
  2. ↑ 2.0 2.1 2.2 Wilgan, Robin (January 2025). "Phylogenetic Determinants Behind the Ecological Traits of Relic Tree Family Juglandaceae, Their Root-Associated Symbionts, and Response to Climate Change" (in en). International Journal of Molecular Sciences 26 (14). doi:10.3390/ijm. ISSN 1422-0067. https://www.mdpi.com/1422-0067/26/14/6866. 
  3. ↑ 3.0 3.1 Manos, P. S.; D. E. Stone (2001). "Evolution, phylogeny and systematics of the Juglandaceae". Annals of the Missouri Botanical Garden 88 (2): 231–269. doi:10.2307/2666226. Bibcode: 2001AnMBG..88..231M. http://www.biodiversitylibrary.org/pdf3/008608200086343.pdf. 
  4. ↑ 4.0 4.1 Meng, Hong-Hu; Zhang, Can-Yu; Low, Shook Ling; Li, Lang; Shen, Jian-Yong; Nurainas; Zhang, Yu; Huang, Pei-Han et al. (2022-11-01). "Two new species from Sulawesi and Borneo facilitate phylogeny and taxonomic revision of Engelhardia (Juglandaceae)". Plant Diversity 44 (6): 552–564. doi:10.1016/j.pld.2022.08.003. ISSN 2468-2659. https://www.sciencedirect.com/science/article/pii/S2468265922000786. 
  5. ↑ 5.0 5.1 5.2 Manning, Wayne E. (1978). "The Classification Within the Juglandaceae". Annals of the Missouri Botanical Garden 65 (4): 1058–1087. doi:10.2307/2398782. ISSN 0026-6493. https://www.jstor.org/stable/2398782. 
  6. ↑ 6.0 6.1 John Derek Bewley, Michael Black, Peter Halmer (2006) The Encyclopedia of Seeds: Science, Technology And Uses
  7. ↑ Manning, Wayne E. (1940). "THE MORPHOLOGY OF THE FLOWERS OF THE JUGLANDACEAE. II. THE PISTILLATE FLOWERS AND FRUIT" (in en). American Journal of Botany 27 (10): 839–852. doi:10.1002/j.1537-2197.1940.tb13945.x. ISSN 0002-9122. https://bsapubs.onlinelibrary.wiley.com/doi/10.1002/j.1537-2197.1940.tb13945.x. 
  8. ↑ 8.0 8.1 Paź-Dyderska, Sonia; Jagodziński, Andrzej M.; Dyderski, Marcin K. (2021-02-01). "Possible changes in spatial distribution of walnut (Juglans regia L.) in Europe under warming climate" (in en). Regional Environmental Change 21 (1): 18. doi:10.1007/s10113-020-01745-z. ISSN 1436-378X. https://doi.org/10.1007/s10113-020-01745-z. 
  9. ↑ Teste, François P.; Jones, Melanie D.; Dickie, Ian A. (March 2020). "Dual‐mycorrhizal plants: their ecology and relevance" (in en). New Phytologist 225 (5): 1835–1851. doi:10.1111/nph.16190. ISSN 0028-646X. https://nph.onlinelibrary.wiley.com/doi/10.1111/nph.16190. 
  10. ↑ 10.0 10.1 10.2 Zhou, Huijuan; Hu, Yiheng; Ebrahimi, Aziz; Liu, Peiliang; Woeste, Keith; Zhao, Peng; Zhang, Shuoxin (2021-10-21). "Whole genome based insights into the phylogeny and evolution of the Juglandaceae" (in en). BMC Ecology and Evolution 21 (1): 191. doi:10.1186/s12862-021-01917-3. ISSN 2730-7182. PMID 34674641. PMC 8529855. https://doi.org/10.1186/s12862-021-01917-3. 
  11. ↑ 11.0 11.1 "Large-scale phylogenetic analyses reveal fagalean diversification promoted by the interplay of diaspores and environments in the Paleogene". Perspectives in Plant Ecology, Evolution and Systematics 16 (3): 101–110. 2014. doi:10.1016/j.ppees.2014.03.001. Bibcode: 2014PPEES..16..101X. 
  12. ↑ Frei, Jonas (2021). "A Brief History of Juglandaceae". Arnoldia 78 (3): 10–17. ISSN 0004-2633. https://www.jstor.org/stable/27085389. 
  13. ↑ Sharma, Munish; Sharma, Munit; Sharma, Munish (December 2022). "A comprehensive review on ethnobotanical, medicinal and nutritional potential of walnut (Juglans regia L.)" (in en). Proceedings of the Indian National Science Academy 88 (4): 601–616. doi:10.1007/s43538-022-00119-9. ISSN 0370-0046. PMC 9510174. https://link.springer.com/10.1007/s43538-022-00119-9. 
  14. ↑ "Black Walnut Toxicity". http://www.wvu.edu/~agexten/hortcult/fruits/blkwalnt.htm. 
  15. ↑ "GRIN Genera of Juglandaceae". Germplasm Resources Information Network. United States Department of Agriculture. http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?595. 
  16. ↑ Luo, Jie; Chen, Junhao; Guo, Wenlei; Yang, Zhengfu; Lim, Kean-Jin; Wang, Zhengjia (January 2022). "Reassessment of Annamocarya sinesis (Carya sinensis) Taxonomy through Concatenation and Coalescence Phylogenetic Analysis" (in en). Plants 11 (1). doi:10.3390/plant. ISSN 2223-7747. https://www.mdpi.com/2223-7747/11/1/52. 
  17. ↑ 17.0 17.1 Mu, Xian-Yun; Tong, Ling; Sun, Miao; Zhu, Yi-Xuan; Wen, Jun; Lin, Qin-Wen; Liu, Bing (2020-06-01). "Phylogeny and divergence time estimation of the walnut family (Juglandaceae) based on nuclear RAD-Seq and chloroplast genome data". Molecular Phylogenetics and Evolution 147. doi:10.1016/j.ympev.2020.106802. ISSN 1055-7903. https://www.sciencedirect.com/science/article/pii/S1055790320300749. 
  18. ↑ Trouern-Trend, Alexander J.; Falk, Taylor; Zaman, Sumaira; Caballero, Madison; Neale, David B.; Langley, Charles H.; Dandekar, Abhaya M.; Stevens, Kristian A. et al. (April 2020). "Comparative genomics of six Juglans species reveals disease-associated gene family contractions". The Plant Journal: For Cell and Molecular Biology 102 (2): 410–423. doi:10.1111/tpj.14630. ISSN 1365-313X. PMID 31823432. https://pubmed.ncbi.nlm.nih.gov/31823432. 
  19. ↑ Grauke, L.J; Wood, Bruce; Harris, Marvin. "Crop Vulnerability: Carya". https://journals.ashs.org/view/journals/hortsci/51/6/article-p653.xml. 
  20. ↑ 20.0 20.1 "Kentucky Energy and Environment Cabinet". https://eec.ky.gov/Pages/index.aspx. 
  21. ↑ "Butternut | The Wood Database (Hardwood)" (in en-US). https://www.wood-database.com/butternut/. 

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