Annotated List of Boletes and Amanita in the Hawaiian Islands

North American Fungi

Volume 3, Number 7, Pages 167-176 Published August 29, 2008 Formerly Pacific Northwest Fungi

Don E. Hemmes1 and Dennis E. Desjardin2

1Biology Department, University of Hawai’i at Hilo, 200 W. Kawili St., Hilo, HI 96720 and 2Department of Biology, San Francisco State University, 1600 Holloway Avenue, San Francisco, CA 94132

Hemmes, D. E., and D. E. Desjardin. 2008. Annotated List of Boletes and Amanita in the Hawaiian Islands. North American Fungi 3(7): 167-176. doi: 10.2509/naf2008.003.00710

Abstract: Five species of boletes, Chalciporus piperatus, Suillus brevipes, S. granulatus, S. salmonicolor, and Pulveroboletus xylophilus and three species of Amanita, A. marmorata subsp. myrtacearum, A. muscaria var. formosa, and A. manicata have been collected in the Hawaiian Islands. The boletes are associated with introduced pines and the Amanita are associated with introduced pines, Myrtaceae and Casuarina. Pulveroboletus xylophilus and Amanita manicata appear to be saprotrophic instead of mycorrhizal.

Key Words: boletes, Amanita, Hawai’i 168 Hemmes and Desjardin. Boletes and Amanita species in Hawaiian Islands. North American Fungi 3(7): 165-174

Introduction: Since there are no native spp.) and Kukuiolono Park (Eucalyptus spp.); ectomycorrhizal trees in Hawai’i (Koske, Gemma, O’ahu, Aiea Loop Trail (Eucalyptus spp., and Flynn, 1992), mushrooms found in the native Melaleuca, Casuarina) ; Moloka’i, Pala’au State forests are largely small, saprotrophic species. Park (Eucalyptus spp., Melaleuca, Casuarina) Large ectomycorrhizal mushrooms obviously and Kamakou Forest Preserve (Eucalyptus spp., started appearing with the introduction of Pinus Pinus); Maui, Polipoli Springs State Recreation species in the late 1880’s. The introduction of Area (Pinus spp., Sequoia, Cupressus), Waihou conifers, including more Pinus species, Cupressus Springs State Recreation Area (Pinus spp. species, Sequoia sempervirens, and Cryptomeria Melaleuca spp., Cupressus), Waikamoi, (Pinus japonica, continued from 1910 through 1960 with spp.), Maluhia Boy Scout Camp (Eucalyptus spp.) the large scale plantings of millions of seedlings and Kaumahina Wayside Park (Eucalyptus spp, at different elevations on all the major islands by Melaleuca); Lana’i, the Munro Trail (Eucalyptus the Division of Forestry (Little and Skolmen, spp., Pinus spp.); and Hawai’i, MacKenzie Park 1989). Along with conifers, extensive plantations (Casuarina), Tree Planting Road (Pinus spp.), of alien Myrtaceae including Eucalyptus (more Mana Road (Cryptomeria), Honokaia Boy Scout than 90 species) and Melaleuca were established Camp (Eucalyptus spp.). Fruiting bodies were during the same period. Over a million trees of photographed in the field, important characters Melaleuca quinquenervia have been planted were recorded, and representative specimens are since their introduction to the islands in 1917. stored at the University of Hawai’i at Hilo (UH- Finally, Casuarina equisetifolia was introduced Hilo). to the islands in the late 1800’s and now dominates the windward coastlines in many areas of the islands. These trees are associated with Results: Annotated List of Boletaceae some ectomycorrhizal fungi and the duff under these “ironwoods” or “Australian pines” supports Chalciporus piperatus (Bull.:Fr.) Bataille a wide variety of saprotrophic fungi. In this paper Specimens examined: Kaua’i, Makaha Ridge we document the species of Boletaceae and Road, scattered under pines, 5 January 1995, Amanitaceae found under introduced conifers, DEH-690 (Fig. 1). Myrtaceae, and Casuarina in Hawai’i and include Pulveroboletus xylophilus and Amanita Suillus brevipes (Peck) Kuntze manicata that appear to be saprotrophic rather Specimens examined: Hawai’i Island, Saddle than mycorrhizal species. Most of these species Road, scattered in duff under Monterey Pine, 9 were pictured previously in Mushrooms of November 1993, DEH-322; same location, 15 Hawai’i (Hemmes and Desjardin, 2002). March, 1997, DEH-1473. Maui, Waihou Springs State Park, scattered under Loblolly pines, 22 Materials and Methods: The species of November 1996, DEH-1340 (Fig. 2). boletes and Amanita listed in this paper have been recorded over the past fifteen years in our Suillus granulatus (L.:Fr.) Kuntze surveys and inventories project documenting the Specimens examined: Hawai’i Island, Saddle Agaricales and other fleshy fungi of the Hawaiian Road, scattered in duff under Monterey Pines, 29 Islands. Trips to the various islands were August 1995, DEH-879; Waimea, County Public scheduled from September through January Works Headquarters, in grass under slash pine, when the rains were most frequent. The 13 November 1999, DEH-1866; same location, in following conifer and Eucalyptus plantations grass under slash pine, 10 August 2001, DEH- were visited: Kaua’i, Koke’e State Park/Makaha 2115. Maui, Waihou Springs, Olinda, scattered in Ridge Road (Pinus spp., Sequoia, Eucalyptus duff under pines, 15 January, 1995, DEH-734; Hemmes and Desjardin. Boletes and Amanita species in Hawaiian Islands. North American Fungi 3(7): 165-174 169

Polipoli Springs, scattered under pines, 13 campus, scattered in grass under Melaleuca, 17 January 2000, DEH-1921 (Fig. 3). April 2003, DEH-2390; same location, 26 April 2003, DEH-2409; same location, scattered Suillus salmonicolor (Frost) Halling under Melaleuca, 22 October 2004, DEH-2519; Specimens examined: Hawai’i Island, Volcano same location, scattered under Melaleuca, 15 Golf Course, under pines, 4 January 1997, DEH- September 2000, DEH-1953. Lana’i, Munro 1392; Kaua’i, Makaha Ridge Road, in duff under Trail, scattered under Eucalyptus robusta, 25 pines, 3 October 1994, DEH-604; Lana’i, Munro January, 1997, DEH-1450; same location, Trail, scattered under slash pine, 5 January scattered under Eucalyptus robusta, 5 January 2002, DEH-2231 (Fig. 4). 2002, DEH-2232. Maui, Makawao, scattered under Eucalyptus robusta, 22 November 1996, Pulveroboletus xylophilus (Petch) Singer DEH-1330. Moloka’i, Kamakou Forest Specimens examined: Hawai’i Island, Preserve, scattered under Eucalyptus robusta, 9 MacKenzie Park, scattered in black sand at berm January 2000, DEH-1906; Pala’au State Park, of beach under coconut, 28 July 1993, DEH-272; scattered under Casuarina/Eucalyptus, 7 same location, 17 July 2000, DEH-1964; January 2000, DEH-1898. Oahu: University St. Hawaiian Beaches subdivision, scattered in lawn Louis Heights, scattered under Norfolk Island grass, 6 September 2001, DEH-2130; Ookala, Pines, Araucaria columnaris 14 September, scattered in Guinee Grass, Panicum maximum, 1996, DEH-1236 (Fig. 6). 5 December 2004, DEH-2543 (Fig. 5). Amanita muscaria var. formosa (Pers.:Fr.) Bertillon Annotated List of Amanitaceae Specimens examined: Kaua’i, Makaha Ridge Road, scattered under loblolly pine and slash Amanita marmorata subsp. myrtacearum pine, 18 February 1995, DEH 741; same location O. K. Miller, Hemmes & Wong 7 January 1996, DEH-950; same location, 10 Specimens examined: Hawai’i Island, January 1996, DEH-973; same location 6 MacKenzie Park, scattered in duff under January 1997, DEH-1411; same location, 4 Casuarina, 20 September 1994, DEH 559; January 2000, DEH-1902. Moloka’i, Kamakou, University of Hawaii at Hilo campus, scattered under pines, 3 January 2007, DEH-2667 (Fig. under Melaleuca, 12 May 1995, DEH-782; 7). Kapoho, scattered under Casuarina, 1 September 1995, DEH-884; UH-Hilo campus, Amanita manicata (Berk. & Broome) Pegler scattered under Melaleuca, 27 June, 1996, DEH- Specimens examined: Hawai’i Island, Waipio 1127; Hawai’i Community College campus, Valley, solitary specimen in black sand at berm scattered in grass under Melaleuca, 19 August of beach, 1 October 1996, DEH-1390; Church of 1999, DEH-1831; Kalopa St. Park, scattered Jesus Christ of Latter-Day Saints, Kilauea under Eucalyptus robusta, 19 November 1996, Avenue, Hilo, scattered in grass, 29 April, 2003, DEH-1321; Mackenzie Park, scattered under DEH-2416; Bay Front, Hilo, Hawai’i Island, Casuarina, 17 September 2000, DEH-1962. scattered in composted woodchips, 12 Kaua’i, Koke’e, in dirt under Eucalyptus, 4 September 2005, DEH-2571; same location, 22 October 1994, DEH-616; MacKenzie Park, September 2005, DEH-2586; same location, 29 Hawai’i Island, scattered in duff under September 2006, DEH-2627 (Fig. 8). Casuarina, 9 May 2002, DEH-2278; Keaukaha, Hilo, Hawai’i Island, scattered in duff under Discussion:Three species of Suillus are Casuarina, 18 April 2003, DEH-2389; UH-Hilo commonly found under pines in Hawai’i. S. 170 Hemmes and Desjardin. Boletes and Amanita species in Hawaiian Islands. North American Fungi 3(10): 165-174 brevipes is typically found under Monterey monitored for years, A. muscaria var. formosa pines, P. radiata, whereas S. granulatus and S. has never been seen fruiting under these trees as salmonicolor are typically found under slash in New Zealand (Ridley and Dick, 2001). pines, P. elliottii and loblolly pines, P. taeda. The preliminary identification of Amanita Chalciporus piperata has been found only under manicata was made by Ross Tulloss (personal P. elliottii at Koke’e Kaua’i in the same area as A. communication). Solitary specimens of this muscaria. Pulveroboletus xylophilus is widely species have been collected on lawns at the spread over Hawai’i Island. The first specimens National Tropical Botanical Garden at Lawai, were collected from black sand under coconut Kauai, at Hoomaluhia Botanical Gardens on trees at the berm of the beach at MacKenzie park Oahu, at the Keauhou Beach Hotel on the Kona in the Puna District of Hawai’i Island. Since Coast of Hawai’i Island, and in black sand at the then additional collections have come from berm of the beach at Waipio Valley on Hawai’i coffee farms on the Kona Coast, from cattle Island. In 2005 a large fruiting of A. manicata pastures along the Hamakua Coast, and from appeared on a lawn at the Church of Jesus Christ grassy vacant lots in the Puna District. This of Latter-Day Saints on Kilauea Avenue in Hilo obviously saprotropic species stains deep blue and then in 2006 in composted wood chip piles when bruised or cut in half and usually grows on along Bay Front in downtown Hilo. The large the ground but has been seen fruiting on fruiting bodies have no volva, have uplifted coconut tree stumps. scales on the stipe, and have a distinctly Amanita marmorata var. myrtacearum is by far unpleasant odor. The closest associated trees in the most common Amanita in the Hawaiian most of these fruitings have been Monkey Pod islands and is found in Eucalyptus plantations, trees, Pithecellobium saman, but this Amanita and especially around Melaleuca, “bottle brush” is presumed to be saprotropic. or “paper bark” trees, used in landscaping in urban areas. The association of A. marmorata Acknowlegements: The authors dedicate this var. myrtacearum with introduced Myrtaceae article to our friend, Jack Rogers, in recognition was obvious when the grass surrounding a large of his outstanding contributions to mycology planting of various species of Australian over his entire career, and in fond memory of Myrtaceae at the National Tropical Garden at wonderful collecting trips together in the Lawai on Kaua’i was white with hundreds of Hawaiian Islands. This research was funded in fruiting bodies. A. marmorata is also commonly part by NSF Biotic Surveys and Inventories found under coastal Casuarina and is Grant #DEB-9300874. presumably mycorrhizal with these trees too. Amanita muscaria var.formosa is much more Literature Cited limited in its distribution within the islands. Hemmes, D. E., and D. E. Desjardin. 2002. Fruiting bodies of this species are common Mushrooms of Hawai’i. Ten Speed Press. under slash pine pines and loblolly pines along Berkeley, CA, pp. 212. the Makaha Ridge Road at Koke’e, Kaua’i, during December and January, and recently Koske, R. E., J. N. Gemma, and T. Flynn. 1992. several fruiting bodies have been collected at Mycotrophy in Hawaiian angiosperms: A survey Kamakou on Moloka’i under the same pines. with implications for the origin of the native Amanita muscaria var. formosa has not been flora. Am. J. Bot. 79: 853-862. seen under the same pines on Maui or Hawai’i http://dx.doi:2307/2444994 Island. Even though groves of Monterey pines, Pinus radiata, were planted extensively, Little Jr., E. L. and R. G. Skolmen. 1989. especially on Hawai’i Island and have been Common Forest Trees of Hawaii (Native and Hemmes and Desjardin. Boletes and Amanita species in Hawaiian Islands. North American Fungi 3(7): 165-174 171

Introduced). United States Department of Agriculture, Forest Service, Agriculture Handbook No. 679, Washington DC, pp. 321.

Ridley, G. S., and M. A. Dick. 2001. An Introduction to the diseases of forest and amenity trees in New Zealand. Forest Research Bulletin 220: 110 p. Rotorua: Forest Research. 172 Hemmes and Desjardin. Boletes and Amanita species in Hawaiian Islands. North American Fungi 3(10): 165-174

Fig. 1. Chalciporus piperatus under slash pine along Makaha Ridge Road at Koke’e, Kaua’i.

Fig. 2. Suillus brevipes is commonly found under pines on all the Hawaiian Islands. Hemmes and Desjardin. Boletes and Amanita species in Hawaiian Islands. North American Fungi 3(7): 165-174 173

Fig. 3. Suillus granulatus under slash pine. 174 Hemmes and Desjardin. Boletes and Amanita species in Hawaiian Islands. North American Fungi 3(10): 165-174

Fig. 4. Suillus salmonicolor under pines along Makaha Ridge Road, Koke’e, Kaua’i.

Fig. 5. Pulveroboletus xylophilous from the berm of a black sand beach on Hawai’i Island. Hemmes and Desjardin. Boletes and Amanita species in Hawaiian Islands. North American Fungi 3(7): 165-174 175

Fig. 6. Amanita marmorata subsp. myrtacearum from under Melaleuca on the UH-Hilo campus.

Fig. 7. Amanita muscaria var. formosa at Makaha Ridge Road, Koke’e, Kaua’i. 176 Hemmes and Desjardin. Boletes and Amanita species in Hawaiian Islands. North American Fungi 3(10): 165-174

Fig. 8. Amanita manicata from composted wood chip piles at the bay front of Hilo, Hawai’i.