Revised May 1, 2002; August 29, 2007; January 2, 2011; May 2, 2013; and April 23, 2017.
Copyright 1999-2017 by John W. Allen.



Lady Shroomer on Kualoa Ranch, Kahuku, O'ahu. Photo: J. W. Allen.


2. Methodology  
Fungi specimens referred to in this paper were collected from several locations in the Hawaiian Islands. These specimens were later deposited in three institutions listed below. The fungi species referred to in Tables 1-5 were identified by Ewald Gerhardt in Berlin, Germany, and Gastón Guzmán in Xalapa, Mexico. Some of these same species were later analyzed chemically for suspected psychoactive compounds using TLC and HPTLC techniques by T. Stijve in Vevey, Switzerland (Table 1).

Non-Psychoactive Species Collected from O'ahu

3. RESULTS 3.1. Specimens Collected for Herbarium Deposit  
Several specimens of Hawaiian fungi (primarily various collections of Panaeolus and Copelandia spp.) were collected from various locations on O'ahu island; including two collections from Maui. The specimens (Isotypus in BISH, M. Burton, Sn #626837, Sn #626836; J. W. Allen, Sn #616451, Sn #626827, Sn #626828, Sn #626829, Sn #626831, Sn #616832, Sn #626833, Sn #626834, Sn #626835, Sn #626839) were sun-dried and deposited at the Herbarium Pacificum at the Bernice P. Bishop Museum, Honolulu, Hawai'i. Hawaiian specimens of Panaeolus spp. and Copelandia spp. deposited in the Bernice P. Bishop Museum previous to this study included collections by M. C. Neal (Sn #498521, Sn #498522, Sn #615638) and A. Puharich (Sn #146101).

Additional specimens collected in this study were deposited in the Instituto de Ecologia, Xalapa, Veracruz, Mexico (XAL) and the Institut für Biotechnologie, Leipzig, Germany.

Collected specimens of A. muscaria have been deposited at the National Tropical Botanical Gardens at Lawai, Kaua'i and at the Herbarium Pacificum at the Bernice P. Bishop Museum in Honolulu, Hawai'i (Isotypus in BISH, Montgomery Sn #594747, Sn #594748, Sn #594749).

Specimens Collected on O'ahu

3.2. Species Identification  
On November 14, 1989, Guzmán identified the following species based on specimens from Hawai'i: C. cyanescens, C. bispora, P. subbalteatus, P. campanulatus, and Psilocybe coprophila. The specimens were deposited at the Institut de Ecologia, Xalapa, Veracruz, Mexico ( see Table 3).

The specimens of C. cyanescens identified by Guzmán were collected on October 25, 1987 from cattle pastures in manure at the U of Hawaii's livestock and Experimental Farm on the North Shore of O'ahu.

Copelandia cyanescens is known from the tropics of both hemispheres. It was originally reported and described from Sri Lanka by Berkeley and Broome in 1871 as Agaricus cyanescens.

A few years later it was identified from the Philippines as Copelandia papilionacea by Bresadola (1881-92).

Copelandia cyanescens, one of eight binomials published in the genus Copelandia, has also been classified as Panaeolus (subgenus Copelandia) cyanescens (see Allen et al., 1992 and Weeks et al. 1979). According to Guzmán, (pers. comm., 1988), P. cyanescens and C. cyanescens are the same species following two different taxonomic positions. Weeks et al. (1979) pointed out that "Copelandia has been reduced to an infragentic taxon by some authors who had preferred to include this Copelandia in Panaeolus (Fr.) Quel. (Ola'h, 1968, 1969, 1970)." They argued that "this treatment seemed to have some merit as long as only one species was believed to belong in Copelandia and a broader genus concept was thought to be desirable." Weeks et al. asserted that this concept is not valid since each Copelandia specimen they studied were "specifically different from one another and all showed the characteristic colored metalloid cystidia and bluing context, without any forms known intermediate between these and Panaeolus." Therefore, in accordance with Singer (1975), Horak (1968), and Malençon et Bertault (1970), Weeks et al. maintained "that a narrower genetic concept in the Panaeoloideae is warranted (see Stijve, 1992)."

Existing mycological literature describes various species of Copelandia from India, Indonesia, Thailand, Cambodia, Philippines, Australia, New Zealand, Samoa, Hawai'i, various areas of North America (including British Columbia, Washington, Oregon, California, and Florida), Jamaica, Mexico, various areas of Central America and South America (including Colombia, Bolivia, Brazil, and Peru), Madagascar, France, and Italy (Allen and Merlin 1992, Allen et al., 1991; Cox 1981; Schultes and Hofmann 1980; Weeks et al., 1979; Guzmán 1978; Lincoff and Mitchell 1977; Pollock, 1974, 1976, 1977-1978; Ola'h, 1968, 1969, 1970).

Copelandia species are common in numerous cattle pastures on O'ahu. For example, species described in this paper were observed in or near various grassland areas of O'ahu including Mokule'ia, Haleiwa, Kahuku, La'ie, Kualoa, Kahalu'u and Kailua.

On Maui, C. cyanescens can be observed in many grassland locations throughout the island, e. g., near Pa'ia, Hana, Makawao, Kula, 'Ulupalakua Ranch Along Kula Highway on Maui on Haleakala Volcano, Ho'okipa and northwest of Wailuku. Copelandia fungi have also been observed fruiting from manure in pastures as high as 2,000 m on the slopes of Haleakala volcano, East Maui. Another species, Panaeolus subbalteatus, is also common in several pastures and wooded areas on the slopes of Haleakala volcano at approximately 3,000 foot elevation.

On Hawai'i island, in November of 1988, one of the authors (JWA), along with Don Hemmis, a professor of biology at the University of Hawai'i at Hilo, observed C. cyanescens, P. subbalteatus, P. campanulatus, A. semiovata, and Psilocybe coprophila at approximately 2,000 m on the windward slopes of Mauna Kea volcano. Copelandia spp. and P. subbalteatus are also known to occur in dung found in the cattle pastures located in the northern region of Hawai'i island, as well as in the grasslands of cattle ranches in the southern and western areas of that island. Recently JWA observed Copelandia fungi in the Paradise Park residential sub-division of Kea'ua on Hawai'i island. These fungi were growing at 10 to 15 m above sea level in areas grazed by horses and cattle, and did not appear to be adversely affected by the salt spray from the ocean nearby.

Pollock (1974, 1976), reported Copelandia species from the island of Kaua'i. These fungi are also undoubtedly found in pastures on the islands of Moloka'i, and possibly in grazed grasslands on Lana'i and the privately-owned island of Ni'ihau. Copelandia tropicalis was originally reported as occurring on the dung of cattle and other wild or domesticated ruminants in Cambodia, the Republic of Central Africa, and Mexico (Pollock, 1976). Ola'h (1969), Pollock (1976) and Rhoades (1975) all reported this species from Hawai'i.

Specimens of C. bispora (Malençon et Bertault) Singer and Weeks were collected on October 25, 1987 from pastures of a livestock and experimental farm on the North Shore of O'ahu. They were identified in 1989 by Guzmán (pers. comm.). This species was previously recognized only from North Africa (Malençon et Bertault 1970), and is now reported for the first time from the Hawaiian Islands. Copelandia anomala was identified from Hawai'i in 1975 by F. Rhoades (Pollock, 1976).

Panaeolus subbalteatus (Berk. et Br.) Sacc., collected on October 5, 1988 (Isotypus in XAL, Mexico) and November 4, 1992 Isotypus in BISH, Allen Sn #626950, SN #627400), were also collected in pastures of the same livestock and experimental farm on the North Shore, O'ahu refered to above. Specimens of this species were also identified by Guzmán. One collection from Maui is on deposit at the Herbarium Pacificum in Honolulu (Isotypus at BISH, Burton and Pence SN #627399.

Panaeolus subbalteatus, infrequently collected with Copelandia specimens, is sometimes used as a recreational drug in Hawai'i. Although this fungi can be observed in large numbers in some areas on Maui, it is not as common as the Copelandia spp. found in many pastures of the Hawaiian Islands.

The geographical distribution of P. subbalteatus is cosmopolitan. It has been reported from Australia, Asia, India, North America, Central America, South America, Western Europe, Scandinavia, and Africa (Stijve and Kuyper 1985; Miller et al., 1982; Weil, 1980, 1977; Pollock, 1976; Shepherd and Hall 1973; Anonymous, 1972; Enos, 1970)and in the State of Alaska. Panaeolus subbalteatus is not only coprophilous, but is also known to occur in lawns, grassy areas, compost heaps, and in rotted hay which has been left out to decay. It can also be found at horseback riding stables.

Psilocybe coprophila Guzman was collected from horse manure on 2 September 1987 in cattle pastures near Kahalu'u, Oahu. This species was identified by Guzman. It has also been observed on both Hawaii's and Maui Islands. In February 1991, Gerhardt identified Panaeolus goosensiae Beeli from specimens which were collected in 1989 from a pasture also near Kahalu'u, O'ahu (Isotypus in BISH, Allen, 626827).

All of the dung-inhabiting species referred to in this paper were probably introduced into the Hawaiian Islands by cattle (Bos Taurus) and/or horses (Equus caballus) or perhaps other alien ruminant herbivores. Cattle were first introduced to Hawaii by the British captain, George Vancouver in 1793 and 1794; hor4ses were first introduced to Hawai'i in 1803 aboard a merchant ship under the command of Captain George Vancouver (Tomich, 1986).

Results of Chemical Analysis of Fungi Collected on O'ahu Island July 17-24, 1990.
Analysis by T. Stijve, Nestec Ltd. (Plate 1-2. Fig. 13-14)

3.3. Specimens Collected for Chemical Analyses  
Fungi specimens gathered in the field for chemical analyses included nine collections of psychoactive mushrooms identified as either C. cyanescens or C. cambodginiensis. Three other non-psychoactive species belonging to the genus Panaeolus were also gathered for chemical analyses.

The Copelandia collections were labeled A-I, while the Panaeolus collections were identified separately. All of these collections were forwarded to T. Stijve of Nestec Ltd., Vevey, Switzerland for chemical analyses (see Table 4 for results of chemical analyses of A-I; see Table 5 for results of chemical analyses of Panaeolus).

Species identification of these specimens was made by Ewald Gerhardt of Berlin. Duplicate collections of Hawaiian Copelandia spp. were deposited at the Institut fur Biotechnologie, Leipzig, Germany. The collections analyzed chemically are described below. A. Two specimens collected on July 17, 1990 near Kahalu'u, O'ahu were identified by Gerhardt in Berlin as C. cambodginiensis. B. Numerous primordial size specimens collected on July 22, 1990 near Kualoa, O'ahu were identified in 1991 by Gerhardt in Berlin as C. cambodginiensis; this species has been reported previously from Cambodia, France, and Africa (Ola'h, 1968, 1969), Colombia (Pollock, 1975, 1976; Ott and Guzmán 1976), and recently from Mexico and Peru by Guzmán (pers. comm., 1990); this is the first report of C. cambodginiensis from Hawai'i.

Collections A and B differ from C. cyanescens (collections C-I) by having smaller spores and thin-walled metuloids; and in collection A the cystidia had a somewhat greenish color, which has an uncertain taxonomical value.

The other collections analyzed chemically, and labeled C-I, are described below. Specimens in these collections were all identified by Gerhardt as C. cyanescens. C. Fifty specimens were collected on July 23, 1990 in a pasture from a livestock and Experimental Farm on the North Shore of O'ahu (Isotypus in BISH, Allen Sn #626839); D. Seven specimens were collected on July 23, 1990 in a pasture near Kahalu'u, O'ahu (Isotypus in BISH, Allen Sn #626829); E. Four specimens with blue-green stained stems were collected on July 23, 1990 in a pasture near Kahalu'u, O'ahu (Isotypus in BISH, Allen Sn #626831);

F. One very large specimen and seven other larger than average specimens were collected on July 23, 1990 near Kahalu'u, O'ahu; G. Five specimens were collected on July 24, 1990 near Kahalu'u, O'ahu; H. Eight specimens were collected on July 24, 1990 near the Kahalu'u, O'ahu; I. Thirteen specimens were collected on July 24,1990 near the Coral Kingdom on Kahalu'u, O'ahu. Several additional collections of Copelandia spp. and other dung-inhabiting fungi have been deposited in the Herbarium Pacificum at the Bernice P. Bishop Museum, Honolulu, Hawai'i.

Analytical Results of Non-Psychoactive Dung Fungi From O'ahu, Hawaii  

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