The order Phallales, as outlined by Cunningham
(1942) and Zeller (1949), contains three
families and approximately 20 genera of fleshy
basidiomycetes, which, because of their usually
stalked fruiting bodies and foetid odor, are
commonly called "the stinkhorns." The group
includes some of the most bizarre and unusual
of the fungi

Goos, R.D.

Pacific Science

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Phalloid Fungi in Hawaii!R. D. GOOS2THE ORDER Phallales, as outlined by Cunning-ham (1942) and Zeller (1949), contains threefamilies and approximately 20 genera of fleshybasidiomycetes, which, because of their usuallystalked fruiting bodies and foetid odor, arecommonly called "the stinkhorns." The groupincludes some of the most bizarre and unusualof the fungi .The Phallales appear to be poorly representedin Hawaii, at least insofar as present records in-dicate. Five species, representing five genera intwo families, have been collected in the Islands .It is the purpose of this report to summarizecurrent knowledge of the group in Hawaii, inthe hope that this will stimulate further col-lecting and reporting of these interesting fungi .We are indebted to N . A. Cobb, formerlychief plant pathologist with the Hawaiian SugarPlanters' Association Experiment Station, formuch of our knowledge of the phalloids inHawaii. Cobb's observations on fruit body de-velopment and spore dispersal in these fungiare among the most detailed to be found inmycological literature (Cobb, 1906, 1909). Un-fortunately, they appear to have been overlookedby many mycologists.Because he associated these fungi with rootdisease in sugar cane, Cobb devoted much at-tention to them, and was particularly concernedwith factors affecting their growth, dispersal,and survival in sugar cane fields. He very care-fully studied the feeding habits of the sarcoph-agous flies which feed on the spores of thesefungi, and which are the agents of spore dis-persal, and demonstrated that the fungus sporescan pass unharmed through the gut of the in-sect. In light of present knowedge of the be-havior of many coprophilous fungi, this factdoes not seem surprising, but it appears to be! This investigation was supported by PHS ResearchGrant no. 15198 from the N ational Institutes ofHealth . Manuscript received June 13, 1969.2 Department of Botany, University of Hawaii,Honolulu, Hawaii 96822.one of the few documented observations of thisphenomenon in the phalloids. Cobb (1906)showed that an adult fly could consume up to80 percent of its body weight in fungal sporesin a single day, and that a single "speck" of flyexcrement could contain as many as 22,400,000fungus spores.Statements in current mycological literature(Ingold, 1965) indicate that the spores of thePhallales have never been observed to germi-nate; however, Cobb and his associate, 1. Lew-ton-Brain, reported and illustrated germinatingspores of Phallus rttbictmdttJ (Cobb, 1906) .The fruiting body of the phalloids is initiallyenclosed within a whitish, egg-shaped structure.Development of the sporophore takes placecompletely within the "egg," and when mature,the pressure caused by the enlargement of theinternal structures causes the outer wall, orperidium, to break. The spongy, spore-bearingstructure then emerges. Emergence of thesporophore usually occurs quite rapidly. Alexo-poulos (1962) states that in Dictyophora dttp-licata, the stalk is fully expanded 5 hours afterthe peridium splits . Cobb (1906) illustratedthe rapid development of P. rubicundltS in aseries of photographs, portrayed here as Fig-ure 1.The fruiting body is extremely ephemeraland wilts soon after its emergence from the"egg." Cobb (1906) stated that fructificationsof P. mbictmdttJ that were fully expanded by6 :30 A.M. had begun to wilt by 10:00 A.M.A checklist of Hawaiian fungi published in1940 (Parris, 1940) listed three species of phal -loid fungi. Two additional species have beenfound in the Islands since the publication ofthat report. Phalloids now known to occur inHawaii are discussed in the subsequent para-graphs.Antburus [auanicns (Penzig) G. H. Cunning-ham. Proceedings of the Linnean Society ofNew South Wales, vol. 56, p. 186, 1931.282Phall oid Fungi in Hawaii-Goos~* ... . ' ~. . . . .'. ... .•'.•283A.M.FIG. 1. Phallus rubicundus . D evelopment from the "egg stage." (A fter Cobb, 1906) .Cunningham, G, H. , Th e Gasteromycetesof Australia and New Zealand , p. 103,1942,The fungus upon which this record is basedwas collected near Punaluu, Oahu, on Augu st10, 1969, by D. Herbst, und er a stand ofEucalyptus. The collection consisted of one ex-pand ed fru ctification and two '"eggs,'" both ofwhich subsequently developed to maturity in amoist chamber. Mature fructifi cations consistedof a shor t stalk subtended by a volva and bear-ing above two to four simple reddish arms,united at the apex (Fig. 2) . Overall height ofthe fruiting bodies ranged from 3,0 to 4.0 cm;width at the base was 7 to 8 mm, Spore masseswere produced in a greenish mucoid mass on theinner surface of the receptacle arms. The sporeswere elliptical, and measured 4-5 X 1.5-2f1.The fungus agrees fairl y well with the de-scription and illustration of A. [auanicus , asgiven by Cunningham ( 1942), alth ough ther eis a slight difference in spore size. Spores of theHawaiian fungus measure 4- 5f1 in length ,whereas Cunningham gives spore length as 5- 6f1.There is great confusion surro unding thename Anthnrns, making placement of a fungusin this genus less than satisfa ctory. Zeller(1948) treats A utbnrns as a synonym of Ly-FIG. 2. An /hums [aranicus, Two fully deve lopedsporocarps and an "egg,284Slims, and this treatment is accepted by Ains-worth (1961). By implication, Rea andHeidenhain (1955) also suggest that the twogenera are synonymous, but an adequate tax-onomic treatment is lacking in their paper.Exact placement of the Hawaiian fungus isimpossible without the examination of appro-priate herbarium material and possibly a reviewof the several genera that may be involved,which is beyond the scope of the present in-vestigation. Inasmuch as the fungus shows closeagreement with Antburus javanims, it is pro-visionally assigned to that species.Aseroe rubra La Billardiere ex Fries. SystemaMycologicum, vol. 2, p. 285, 1822. Cun-ningham, G. H ., The Gasteromycetes ofAustralia and New Zealand., p . 107, 1942.Aseroe rubra is one of the showiest of thephalloid fungi . The mature fruiting body con-sists of a basal volva (the ruptured peridium)subtending a spongy stalk, which bears at itsapex an expanded, brightly colored receptaclefrom whose margin arise a number (5 to 9)of bifurcated arms (Fig. 3). The gloeoid,olivaceous spore mass is borne on the uppersurface of the expanded receptacle. The stalkand volva are usually pale pinkish in color,. while the receptacle may vary from pink to red.A definite foetid odor is usually detectable, andin fresh specimens may be pronounced.According to Cunningham, A. rubra is com-mon in Australia and New Zealand, and it hasbeen reported from Tasmania, New Caledonia,the Malay Archipelago and Ceylon. There is asingle report of it from England (Transactionsof the British Mycological Society, vol. 1, p.FIG. 3. Aseroe rubra. (After Ingold, 1965) .PACIFIC SCIENCE, Vol. 24, April 197025, 1896--1901) , where it was found in KewGardens, associated with plants introduced fromAustralia. It was concluded that the fungusalso was "surely introduced. "The following records of A . rubra in Hawaiihave been obtained: (1) Oahu, 1965; asingle specimen found by D . E. Hardy; no otherdata available. (2) Oahu, 12/31 /68; two speci-mens collected under ElIcalyptllS at an elevationof 1,800 feet, on the trail to Puu Hapapa,Waianae, Mts., by J. Obata. ( 3) Oahu,1/16/69 ; remnants of two decaying sporophoresfound on decaying wood under Eucalyptus,Helemano Forest Reserve, by R. Goos. (4)Oahu, 4/12/69; single sporophore collectedunder Eucalyptus , Aiea Loop Trail, by JaneMihata. (5) Maui, 12/18/67; specimen col-lected at an elevation of approximately 2,500feet, under Cytisus palmensis, 2.3 miles east ofthe junction of highways 377 and 37, by D.Herbst. (6) Maui , 4/ 9/ 67; specimen collectedat an elevation of 700 feet under bamboo, nearthe point where Oopu ola stream intersects HanaRoad, by J. Obata . (7) Molokai , 2/7/69; twosporophores collected in Palaau State Park, onein an open grassy area used for campsites, theother on a foresters' road leading from thecamp area, by B. K. H. Lee and R. Goos.A seroe rubra appears to be well establishedin Hawaii, and its occurrence on three of theprincipal islands suggests that it may occurthroughout the Archipelago. Its distribution inHawaii does not appear to be greatly affected byaltitude, since it has been found at elevationsranging from 700 to 2,500 feet. Whether thefungus is indigenous to Hawaii or has been in-troduced with forest trees brought into theIslands from Australia or other South Pacificlocations, is a matter of speculation . That suchintroduction could occur is evidenced by thesingle report of this fungus from England.Dictyoph ora indusiata (Ventenat ex Persoon)Desvaux. Journal de Botanique, vol. 2,1809. Cunningham, G. H., The Gastero-mycetes of Australia and New Zealand, p.96, 1942. Cobb, N. A., HSPA ExperimentStation, Division of Pathology and Phy-siology, Bulletin 6, pp. 8-14, 1909.The genus Dictyophora is, perhaps, one ofthe most attractive of all the fungi. The fruitingPhalloid Fungi In Hawaii-Goos 285Linderiella columnat« (Bosc) G. H. Cunning-ham. New Zealand Journal of Science andbody consists of a pileus borne at the apex of ahollow cylindrical stem, which emerges, as inAseroe, from a ruptured peridium that remainsas a volva. The spore mass is olivaceous, muc-ilaginous, and foetid, and it covers the exteriorof the pileus. The distinctive characteristic ofthe genus is the production of an elegant netlikeskirt, the indusium, that subtends the pileus(Fig. 4) .Dictyophora indusiai« was reported fromHawaii by Cobb, based on a collection madenear Pepeekeo on the island of Hawaii in 1907by J. W . Webster. The fungus was found onsugar cane trash. The description given by Cobbfor these specimens agrees well with that givenby Cunningham, and his illustrations (Fig. 4)leave no doubt regarding the ident ity of thefungus. In the words of Webster , as quoted byCobb, "In this pile of trash we have the finestincubator of these 'potatoes' and fungus . 'Pota-toes' 1Yz inches diameter. Toadstools 8 to 10inches in height with a large yellowish 'mos-quito net' hung like an umbrella from under thehead."This fungus does not appear to have beenrecorded in Hawaii since Cobb's report. Parris(1940) cites only the single record, and noothers have come to my attention.(Af ter Cobb,Technology, vol. 23, p. 171, 1942.= Clathms trilobatus Cobb, HSPA Experi-ment Station, Division of Pathology andPhysiology, Bulletin 5, p. 209, 1906.The receptacle of Linderiella consists ofsimple columns, united apically, but free anddivided basally (Fig. 5). The spore mass isolivaceous and gloeoid as in the two pr ecedingspecies, and is borne on the inner surface of thereceptacle. There is no stalk in this genus, butthe receptacle arms loop upward to form anupright fructification. Cunningham differen-tiates this genus from Clatbrus on the basisof the basally free arms of the receptacle; inClatbrus they are attached. Zeller (1948)did not include Linderiella in his key to thePhallales, indicating that he probably did notconsider the genus valid .FIG. 5. Linde riella columnate.1909) .•• · 10(After Cobb,. .. .FIG. 4. Dictyopbora indusiata.1909 ) .286Linderiella columnata was reported from Ha-waii by Cobb (1909), who described it as a newspecies in the genus Clatbru s, The fungus wasfound in sugar cane fields on Kauai , and wasassumed by Cobb to be parasitic on the roots ofthe cane plants. According to Cunningham, thefungus is widely distributed, occurring in Northand South America, the West Indies and NewZealand, as well as in Hawaii .There are no additional records of this fun-gus in Hawaii since Cobb's report.Phallus rubicandus (Bose) Fries. Systema My-cologicum, vol. 2, p. 284, 1822. Cunning-ham, G. H., Gasteromycetes of Australiaand New Zealand, pp. 94-95, 1942.= Ithyphallus coralloides Cobb. HSPA Ex-periment Station, Division of Pathologyand Physiology, Bulletin 5, p. 208, 1906.The genus Phallus morphologically resemblesDictyoph ora, but lacks the conspicuous indus-ium. The sporophore consists of a hollow fusoidstem, bearing apically a pileus on which themucilaginous, olivaceous spore mass is produced(Fig. 1).Cobb (1906) reported this fungus as oc-curring on all of the principal islands of Hawaii,associated with root disease in sugar cane. Thereare no more recent records of it in Hawaii,but its apparently common and widespread oc-currence in cane fields in 1906 suggests that thefungus must surely still be extant in the islands.Cunningham reports it as widely distributed intropical and subtropical areas.Phallus rubicundns was the fungus that stimu-lated Cobb's intense interest in the phalloidfungi associated with sugar cane in Hawaii. Hewas convinced that P. rabicundns and Lin-deriella colnmnata were parasitic on the rootsof cane and were causing severe damage to thecrop. This hypothesis did not prove valid how-ever, and recent treatises on sugar cane dis-eases (Martin, 1938; Martin et al., 1961, 1964)make no mention of either fungus. Dlctvopborais ' mentioned as the cause of root disease incane in Australia by Martin et al., (1961),but the slight treatment accorded the fungusindicates that it is not considered a serious prob-lem. While Cobb's opinion of the economic im-portance of these fungi may have been invalid,PACIFIC SCIENCE, Vol. 24, April 1970it did provide the stimulus for some outstandingresearch, and we are indebted to him for somesignificant basic information about them.It is of interest to mycologists to question whythese rather conspicuous and unusual fungi, atone time of such concern to plant pathologists,have been recorded so infrequently in Hawaii .A part of the answer unquestionably lies in theephemeral nature of their fruiting bodies. If notobserved soon after they elongate, they will beoverlooked. A further explanation may lie inthe fact that there are simply not enough collec-tors in Hawaii, and also that those phalloidsthat are seen go unreported.I would like to thank Dr. Gladys Baker forpermission to include her distribution records ofAseroe rubra, for helpful suggestions, and forreviewing the manuscript.LITERATURE CITEDAINSWORTH, G. C. 1961. Ainsworth and Bisby'sdictionary of the fungi . 5th ed. Common-wealth Mycological Institute, Kew, Surrey.ALEXOPOULOS, C. J. 1962. Introductory my-cology. 2nd ed. John Wiley & Sons, NewYork.COBB, N . A. 1906. Fungus maladies of thesugar cane. Hawaiian Sugar Planters' Associa-tion Experiment Station, Division of Pathol-ogy and Physiology, Bulletin 5, pp. 1-254.- - - 1909. Fungus maladies of the sugarcane. Hawaiian Sugar Planters' AssociationExperiment Station, Division of Pathologyand Physiology, Bulletin 6, pp . 5-30.CUNNINGHAM, G. H. 1942. The Gasteromycetesof Australia and New Zealand. Printed pri-vately by John McIndoe, 76 Vogel Street,Dunedin, New Zealand.INGOLD, C. T. 1965. Spore liberation. Claren-don Press, Oxford.MARTIN, J. P. 1938. Sugar cane diseases inHawaii. Advertiser Publishing Co., Honolulu.MARTIN, J. P., E. V. ABBOT, and G. C.HUGHES, eds. 1961-64. Sugar cane diseasesof the world. Vols. I and II. Elsevier Pub-lishing Co., Amsterdam and New York.PARRIS, G. K. 1940. A check list of fungi ,bacteria, nematodes, and viruses occurring inPhalloid Fungi In Hawaii- GoosHawaii, and their hosts. Plant Disease Re-porter, Supplement 121.REA, P. M., and BERTA H EIDENHAIN. 1955.The genus Lyss rus. Papers of the MichiganAcademy of Science, Arts, and Letters, vol.60, pp. 49-66.287ZELLER, S. M. 1948. N otes on certain Gastero-mycetes, including two new orders. Mycolo-gia, vol. 40, pp . 639-668.- - - 1949. Keys to the orders, families, andgener a of the Gasteromycetes. Mycologia, vol.41, pp. 36-58.

Phalloid Fungi in Hawaii

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Phalloid Fungi in Hawaii

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