On this page are listed cycad-related articles and publications not originally printed in the Cycad Newsletter. Click a link below to go directly to the subject area of your choice.

[ Biology/Anatomy/General Info ]  [ Taxonomy/Phylogenetics ]  [ Ethnobotany ]
[ Toxins ]  [ Pests ]  [ Endangered Species Listings/Management Plans ]  [ Books for Sale ]

Cycad Specialist Group Publications

The most important publication of the IUCN/SSC Cycad Specialist Group (CSG) released to date is the Cycad Action Plan. Published in 2003, this 86-page document brings together a collection of data and opinions on one of the world's most threatened plant groups and presents a series of action plans to promote their conservation. The Cycad Action Plan is available in PDF format from the IUCN/SSC website. The full citation for this document is as follows:

Donaldson, J.S. (ed.). 2003. Cycads: Status Survey and Conservation Action Plan. IUCN/SSC Cycad Specialist Group. IUCN, Gland, Switzerland, and Cambridge, UK. ix + 86 pp.

Click the links below to download the entire document or parts thereof.

In 2003, CSG Chair, Dr. John Donaldson, produced the following article on cycad reserves for the 5^th World Parks Congress in Durban, South Africa:

• CSG Article on Cycad Reserves

Also in 2003, the CSG provided most of the data and analysis for the CITES Significant Trade Review of Cycads, compiled by TRAFFIC East/Southern Africa. The report is as follows:

• CSG 2003 Cycad Trade Review

The following news release on cycads was issued by the IUCN in 2003:

• IUCN 2003 News Release: Modern lifestyle threatens oldest seed plants on Earth

The following is a report submitted by the CSG Chair, Dr. John Donaldson, to the IUCN in 2004:

• CSG 2004 Report to IUCN

The "World List of Cycads" is maintained by the CSG and is published every three years. Click the link below to go to the most recent version.

• Hill, K. D., D. W. Stevenson & R. Osborne. 2004. The world list of cycads. Pp. 195-212, In: J.A. Lindström (ed.), Proceedings of the Sixth International Conference on Cycad Biology (CYCAD 2002). Nong Nooch Tropical Garden, Pattaya, Thailand.

In 2005, on the 20th anniversary of the "World List of Cycads," Jody Haynes conducted a historical review of all ten world lists and compiled a comprehensive comparison table listing every taxon ever included on a world list and indicating when each name was added and, in some cases, later removed. This document has never been published, but the most recent update (May 2006) is available for download using the link provided below.

• Haynes, J. L. 2006. Historical Review of the World List of Cycads. Unpublished. 37 pp. (1.1 Mb)

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Non-CSG Publications

Cycad Biology, Anatomy & General Information

Below are some articles that provide either general information on cycads or more specific information on their biology or anatomy.

• Caldwell, O. W., and C. F. Baker. 1907. The identity of Microcycas calocoma. Botanical Gazette 43:330-335.

• Chamberlain, C. J. 1907. Preliminary note on Ceratozamia. Botanical Gazette 43:137.

• Chamberlain, C.J. 1919. The Living Cycads. University of Chicago Press, Chicago, IL. (5.8 Mb)

• Fisher, J. B., and T. E. Marler. 2006. Eccentric growth but no compression wood in a horizontal stem of Cycas micronesica (Cycadales). IAWA Journal 27(4):377-382. (450 Kb)

• Niklas, K. J., E. D. Cobb, and T. Marler. 2006. A comparison between the record height-to-stem diameter allometries of pachycaulis and leptocaulis species. Annals of Botany 97:79-83.

• Siegel, L. 2007. Living fossils have hot sex. University of Utah news release referring to Dr. Irene Terry's article in Science (below).

• Smith, F. G. 1907. Morphology of the trunk and development of the microsporanium of cycads: Contributions from the Hull Botanical Laboratory XCI. Botanical Gazette 43:187-204. (1.2 Mb)

• Terry, I., G. H. Walter, C. Moore, R. Roemer & C. Hull. 2007. Odor-mediated push-pull pollination in cycads. Science 318:70.

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Cycad Taxonomy & Phylogenetics

In 2002, a Cycad Classification Concepts workshop was held at Montgomery Botanical Center in Miami, FL, USA. The primary goal of the workshop was to ensure that cycad taxonomy and classification efforts are scientifically robust and standardized as much as possible. A 300-page volume entitled Cycad Classification: Concepts and Recommendations was published in 2004 from papers presented at this workshop. The full citation for this volume is as follows:

Below are links to an unpublished manuscript entitled A Taxonomic Monograph of the Cycads, written by Charles J. Chamberlain just prior to his death in 1943. This is the first time this manuscript has ever been made available to the general public. It is obviously a draft and contains hand-written editorial comments by Arthrur W. Haupt of UCLA.

• Chamberlain, C. J. n.d. A taxonomic monograph of the cycads. Unpublished. (entire manuscript: 225 pages, 10 Mb)

  • Title Page / Preface / Contents (6 pages, 200 Kb)

  • Introduction (11 pages, 500 Kb)

  • Relationships and Distribution (3 pages, 160 Kb)

  • Morphology and Anatomy (22 pages, 1 Mb)

  • Introduction to Cycadaceae / Key to the Genera (3 pages, 120 Kb)

  • Cycas (30 pages, 1.3 Mb)

  • Stangeria (4 pages, 200 Kb)

  • Bowenia (6 pages, 280 Kb)

  • Dioon (12 pages, 520 Kb)

  • Macrozamia (26 pages, 1.1 Mb)

  • Encephalartos (35 pages, 1.4 Mb)

  • Microcycas (3 pages, 140 Kb)

  • Ceratozamia (10 pages, 420 Kb)

  • Zamia (43 pages, 1.7 Mb)

  • List of Figures (12 pages, 470 Kb)

The following document was associated with Chamberlain's manuscript. It was obviously written after his death, but the author is unknown.

• Synopsis of Living and Fossil Cycadaceae (5 pages, 250 Kb)

The following was one of the earliest attempts at addressing the phylogenetic relationships of the modern cycads.

• Chamberlain, C. J. 1915. A phylogenetic study of cycads. Proceedings of the National Academy of Sciences U.S.A. 1(2):86–90.

The link below is one of the most controversial works on cycads ever published. According to Dr. Paolo DeLuca, Schuster's lack of familiarity of both the living cycads and modern nomenclatural concepts caused “great confusion and instability in the nomenclature of cycads” (DeLuca, 1990). Written entirely in German, it is presented here solely as a historical piece. Be forewarned that the entire manuscript file is quite large and may not open properly; but it can be downloaded in its entirety. For reading online, it is recommended to use the individual page files.

• Schuster, J. 1932. Cycadaceae. Pp. 1-168, in A. Engler (ed.), Das Pflanzenreich 99(IV. 1). (entire manuscript: 174 pages, 27 Mb)

  • Title page - p. 24 (4 Mb)

  • Pp. 25-49 (4 Mb)

  • Pp. 50-74 (4 Mb)

  • Pp. 75-98 (4 Mb)

  • Pp. 99-121 (3.8 Mb)

  • Pp. 122-144 (3.6 Mb)

  • Pp. 145-168 (3.6 Mb)
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Below is an emended description of Dioon mejiae presented at the CYCAD 2005 conference in Xalapa, Veracruz, Mexico.

• Haynes, J. L. & M. A. Bonta. 2007. An emended description of Dioon mejiae Standl. & L.O. Williams (Zamiaceae). Proceedings of the 7th International Conference on Cycad Biology, Xalapa, Mexico, January 2005. Memoirs of the New York Botanical Garden 97: 418-443. (2.1 Mb)

In 2005, Tim Gregory, Jody Haynes, and Jeff Chemnick created a wonderfully illustrated poster and identification key that summarizes the vegetative traits most useful in identifying the known species of Dioon (right). Originally presented at the 7th International Conference on Cycad Biology (CYCAD 2005) in Xalapa, Veracruz, Mexico, this one-of-a-kind poster (which measures 24" x 36") has been printed on heavy glossy paper and made available for sale by TCS. Proceeds from the sale of the posters will go toward cycad research, conservation, and/or education. Click the link below to order a copy today! Key to the Species of Dioon

In 2008, Jeff Chemnick, Tim Gregory, Jody Haynes, Miguel Angel Perez-Farrera, and Loran Whitelock created another wonderfully illustrated poster and identification key that summarizes the vegetative traits most useful in identifying the known species of Ceratozamia (see below). Originally presented at the 8th International Conference on Cycad Biology (CYCAD 2008) in Panama City, Panama, this one-of-a-kind poster (which measures 24" x 36") has been printed on heavy glossy paper and made available for sale by TCS. Proceeds from the sale of the posters will go toward cycad research, conservation, and/or education. Click the link below to order a copy today! Key to the Species of Ceratozamia

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Cycad Ethnobotany

The following is a preliminary assessment of the impacts of harvest on the Indian endemic, Cycas circinalis, in the Nilgiri Biosphere Reserve in Western Ghats, South India. It was prepared in a collaborative effort by representatives of People and Plants International and Keystone Foundation:

• Harvest, Trade, and Conservation of the Endemic Multi-use Cycad, Cycas circinalis L., in the Nilgiri Biosphere Reserve, South India. (600 Kb)

And below is a Cycad Newsletter article written by the same authors and describing their continuing work on Cycas circinalis in India:

• Saneesh, C.S. & A. Varghese. 2007. Mutualistic relationships involving the endemic Cycas circinalis L.: Field notes from the Appankappu Forests, Nilambur, Kerala, India. Cycad Newsletter 30(4):28-29.

Below is a recent paper on the ethnobotany and conservation status of Dioon mejiae—known locally as ‘tiusinte’—in Honduras:

• Bonta, M. A., O. Flores Pinot, D. Graham, J. Haynes & G. Sandoval. 2006. Ethnobotany and conservation of tiusinte (Dioon mejiae Standl. & L.O. Williams, Zamiaceae) in Northeastern Honduras. Journal of Ethnobiology 26(2):228-257. (2.6 Mb)

The following is a recent article on the ethnobotany of Dioon merolae in Mexico:

• Pérez-Farrera, M.A. & A.P. Vovides. 2006. The ceremonial use of the threatened “espadaña” cycad (Dioon merolae, Zamiaceae) by a community of the central depression of Chiapas, Mexico. Bol. Soc. Bot. Méx. 78:107-113. (1.6 Mb)

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Cycad Toxins

• Brenner, E.D., D.W. Stevenson & R. W. Twigg. 2003. Cycads: Evolutionary innovations and the role of plant-derived neurotoxins. Trends in Plant Science 8:446-452.

• Marler, T. E., V. Lee, and C. A. Shaw. 2005. Cycad toxins and neurological diseases in Guam: Defining theoretical and experimental standards for correlating human disease with environmental toxins. HortScience 40(6):1598-1606. (1 Mb)

• Marler, T. E., V. Lee, and C. A. Shaw. 2005. Spatial variation of steryl glucosides in Cycas micronesica plants: Within- and among-plant sampling procedures. HortScience 40(6):1607-1611. (1 Mb)

• Marler, T. E., V. Lee, J. Chung, and C. A. Shaw. 2006. Steryl glucoside concentration declines with Cycas micronesica seed age. Functional Plant Biology 33:857-862.

• Marler, T. E., V. Lee, and C. A. Shaw. 2007. Storage does not influence steryl glucoside concentration in Cycas micronesica seeds. HortScience 42(3):626-628.

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Cycad Pests

• Marler, T. E., and R. Muniappan. 2006. Pests of Cycas micronesica leaf, stem, and male reproductive tissues with notes on current threat status. Micronesica 39(1):1–9.

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Endangered Species Listings & Management Plans

Below is an "endangered species" listing for a cycad species in Australia.

• Macrozamia humilis (a perennial cycad) - endangered species listing. New South Wales Scientific Committee, Australia, 2002.

Below is the 2004 cycad management plan for the province of KwaZulu-Natal, South Africa. Specific locality information has been removed. Thanks go to Ms. Brigitte Church of KZN Wildlife for submitting the plan for inclusion on this page.

• KwaZulu-Natal Cycad Management Plan 2004

Below is the 2005 cycad ordinance for the Transvaal region of South Africa.

• Transvaal Cycad Ordinance 2005

Below is a draft recovery plan for Cycas megacarpa, C. ophiolitica, Macrozamia cranei, M. lomandroides, M. pauli-guilielmi, and M. platyrhachis in Queensland, Australia.

• Draft Recovery Plan for Cycas megacarpa, C. ophiolitica, Macrozamia cranei, M. lomandroides, M. pauli-guilielmi, and M. platyrhachis: 2006 - 2011. Queensland Herbarium, Environmental Protection Agency, Queensland, Australia.

On 17-21 July 2006, a population and habitat viability assessment (PHVA) workshop was held for Encephalartos latifrons in Bathurst, Eastern Cape, South Africa. The workshop was organized in association with the IUCN/SSC Conservation Breeding Specialist Group and the Endangered Wildlife Trust. Below is the report produced following this workshop:

• Albany Cycad PHVA Workshop 2006 Report (1.6 Mb)

(Note: The previous file may not open properly in older versions of Acrobat or Acrobat Reader. If this happens, please click the "Get Acrobat Reader" button below to download the most recent version of the free Adobe Acrobat Reader.)

In June 2007, the government of Northern Territory, Australia, released a draft management plan document for public comment. It can be accessed using the link below.

• Draft Management Program for Cycads in the Northern Territory, Australia

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Books for Sale

Various books on cycads are available.

Note: To read or print PDF documents, you will need Adobe Acrobat or the free Adobe Acrobat Reader software installed on your computer. If you do not have Acrobat Reader, you may download a free copy from the Adobe website by clicking on the following button: 
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