i7961 ~ar vio~aoao ‘va~triiv ioo’IoIa ~o Vc!~ ~tVITII~ MOflt~W ~IVJs~OO ~31~E~IO~ ~O ~J~V1AI dTO ~O~K~t ~HJ, ~!O~ ~ ~ ~o j~N~rniflflA ‘wIJ~vc! MI ISH~KAIMf1 VJ~t~tWI1V ~O Nh1flDY~ ~H~Ii OJ~ iwan~ ~I~H~L V IOMEM ~nO~oV~IHawIo ~IO V~T~N~fJ !‘O s~aictn~ ~ tt 017 ‘. ~I~LIO aUfl1V~EJ~I’I ...•...•...••• .c.~IVWJT~flS A ii: ••••••••~•‘••••‘‘ MOIS~flO~I~ ~INY sMoI~vA~asaO A1 9 ~ ~OH ~t1~W VI~~1Th 111 . ‘ . ~ ~o ~tIA~U • II t ••••••••••••• ..•.•s•e•e•••q••••• NoI~OfltO~~LNI i At •••••••••••••••••••••••••~••••••••••• ~Unott~ ~ao ~~i’i ttt ...........................~!aV1 ~O J~SI’I gJ~N~J~NOO ~O ~‘I~VJi ttt 91 ‘‘~~‘ ~ ~flOQO t~.8tO .XU03 JO ~tU~OJ Ot~o!ot~&OW ~ue~t~ jo ~o~-~X~dWOO peq.~~uc~~1 j 9 tq~ a ri~i~ ~o ~r~r’r LIST OF FIGURES Figure Page 1. Photograph of sporophore of C1ath~ fisoberi.... 12 2. Photograph of sporophore of Colus hirudinOsUs ... 12 3. Photograph of sporophore of Colonnarià o olumnata. • • • • • • • • • • . • , • • . • . 20 4. “Latern&t glebal position of Colozinarla ......... 23 5. Photograph of sporophore of Pseudooo~ ~~y~nicuS ~ 29 6. Photograph of transect ions of It~gg~tt of pseudooo1~ javanious showing three arms ........ 34 7. PhotographS of transactions of Itegg&t of Pseudocolus javanicUs showing four arms ......... 34 8. Basidia and basidiospOres of Pseudoco].uS j aVafliCUs . 35 iv CHAPTER I INTRODU~flON Several collections of a elath~aceous fungus were made during the summer of 1963 in a wooded area off Boulder Park Drive just outside the city limits of Atlanta, Georgia. Attempts to identify the fungus revealed the existence of confusion regarding the taxo— nomic position of certain clathraceous genera. After a more critical survey of the literature, it was evident that diverse opinions prevail relative to the nomenclatorial status of certain generic names and taxonomic delimitations of these genera. The olathraoeous fungi belong to the class Basidiomycetes, the subclass Homobasidiomycetes, the order Phallales, and the family Clathraoeae. Opinions vary concerning the recognition of orders in the Homobasidiomycetes and this affects the number of families recognized. The order Phallales, however, and the fami lies Phallaoeae and Clathraoeae are recognized by most mycologists. Fungi belonging to Phallales are commonly loiown as phalloids, so called beóause of the excessively fetid odor of the exposed gleba present on a horn—like receptaöJ.e. The term “pha].loid”, however, has a more valid reference to the family Phallaoeae, to which Phallus belongs and upon which the family was established. The family Olathraceae, on the other hand, is based on the genus Clathrus and those related stinkhorns that have a receptacle composed of obliquely anastomosing branches or a columnar recep— taclo. The lattice—work pattern of branching is more character istic of the group. This pattern may be confined to an apical 2 position or it may be completely lacking, with the receptacle consisting of simple vertical branches. ~Unoertainty regarding generic limits and status prevail espeoiàlly among those clathraceous members with the receptacle consisting of columnar branches. The problem still appears to be unresolved although several studies of fresh and preserved collec tions of clathraceous fungi assigned to different genera are reported. in the literature. When an attempt was made to identify the locally collected specimens, five generic names were encoun tered. These included Colonnaria, Colus, Clathrus, Latornea aM Pseudocolus. Three of these genera, Laternea, Colonnaria and. Pseudocolus, were clearly based on the presence of columnar receptacles, and two, Colus and Clathrus, were based on a partial or full lattice—work receptacle pattern. Since it was found that the status of Colus and Clathrus required no further clarification, it became the objective of these studies to attempt to clarify the status and relationship of the remaining three genera, namely, Colonnaria, Laternea and Pseudooolus. The principal delimiting features of the genera were found to involve variations in global position, the morphology of the receptacle, and the presence or absence of a stipe. The approach used in an attempt to resolve this problem was to criti cally study and evaluate the morphologic characteristics possessed by specimens assigned to these genera, in order to ascertain the limits of generic boundaries and the extent of species differences within these taxa. CHAPTER II REVIEW OF LITERATURE Traditionally the clathraoeous fungi have been pl~cod in the Gasteromycetes, a subdivision of the subclass Homobasidiomycetes. Today, the term Gasterornyoetes is not generally recognized as a taxonomic category. Martin (1961) has included Gasteromycetes as one of the two earlier series of the Homobasidiomycetes. At one time, all fungi in this series were grouped into the single order, Lycoperdales. Fischer (1890) recognized the order Lycoperdales and included as one of the families was Phallaoeae, the taxon to which the genera Colus and Clathrus were assigned. Excluding Colus, he placed all columnar olathraoeous fungi in the genus Clathrus. C]~ents and Shear (1931) followed Fischer’s treatment of the clathraceous fungi in their volume on known genera of fungi. In the earlier work of Fries (1823), the Gasteromycetes were recognized as a valid class. He included in this class, the order Angiogastres and the suborder Phalloideae, to which four genera were assigned. Clathrus was among the included genera, Laternea. being listed under it as a tribe. Saccardo (1888) elevated Phalloideae to family status and designated Clathreae and Phalleas as subfamilies. The exact status of Laternea and Pseudocolus was not indicated. In Coker and Couches (1928) study of the Gasteromycetes, the family Phallaceae, with subfamilies Clathreae and Phalleae, was recognized. Genera included in Clathreae were Clathrus and Colus; 3 4 Pseudocolus was only mentioned as a generic name and no further comment was made. Fischer (1933) established Phallineae as a suborder and Clathraoeae, including Clathrus and related forms, and Phallaoeae, including Phallus and related forms, as subfamilies. Among the genera of Clathraceae recognized were Colonnaria, Laternea and Psoudocolus. •Zeller (1949) published keys to the Gasteromycetes based on Fischer’s treatment. Differences extend beyond these categories .to the genera. In Glathraceae, the morphology of the receptacle, variation in the gleba in relation to the receptacle, and the presence or absence of a. stipe represent the features providing the taxonomic bases for generic delimitations. The genus Clathrus established by Micheli (1729) was described as having a receptaole without a single distinct branch, forming an inflated hollow network. Colus, erected by Cavalier and Sechier (1835), was indicated as having 6 to 9 vertical branches of equal length, attached to an apical dome—shape lattice—work, to which the gleba adhered. Clathraceous fungi with simple columns joined at the apex said free at the base have been placed in various taxa. Colonnaria, a genus erected by Rafinesque (i808), was described as having a receptacle divided “into four pillars united at the top which bear the seeds in the margin”, Turpin (1822), apparently unaware of Rafinesque’s Colonnaria, described the genus Laternea. The genus Laternea was described as having a receptacle consisting of slender, 5 smooth columns united at the apex with the gleba restricted to this apical region. Linder (1928), in his treatment of clathraceous genera, recognized the validity of Laternea. This genus was not recognized by Fischer (1890) in an earlier treatment, for ho incor porated all columnar clathraceous fungi, except Colus, in the genus Clathrus. It was further suggested by Linder (1928), that Colonnaria, on the basis of priority, be recognized and include those clathraoeous fungi with simple columnar receptacles having no hhlaternett gleba. Linder suggested that Clathrus be reserved for those forms having aaaastomosed receptacles. Cunningham (1931) separated the columnar species, previously included in Clathrus by Fischer (1890), and established for them the genus Linderia, differing from Laternea in having its globa on the inner surface of the arms rather than in the etlaternett position. Linderia was later changed by Cunningham (1942) to Linderiella, since the name Linderia was similar in appearance to Lindera Thunberg, a member of Lauraceae. Cunningham indicated that investigators recognizing Colonnaria were operating on mere guess work in assuming that Rafinesque was dealing with a. fungus. Lloyd (1907) described Rafinescjue’s report of a new fungus as being “Rafinesque raving”. The opinions of both Lloyd and Cunningham were shared by Long and Stouffer (1948), who rejeoted Fischer’s later treatment (1933) of Colonnaria and Laternea. According to Santesson (1943), the rejection of Rafinesque’s description of Colonnaria cannot be accepted in light of the present rules of nomenclature. Santesson expressed the opinion 6 that because of the wording of the description, the only fungus to which such a description could apply would be to a olathraceoUs fuiagus. Santesson therefore made Linderia a synonym of 001 onnaria. In 1933 Fischer recognized the genera Laternea and Coloimaria, and indicated the thain difference delimiting these genera as being ?tfistutous columns” and “massive columns” respectively. Santessofl (1943) reported that the receptacle of Colorinaria was not “massive” but was “lacunose”; the arms of La,ternea were not “fistutoUs” but were ~~lacunosett. Th~.s differing with the work of Fischer, he further indicated only a difference in the size of the cavities in the lacunose tissue, a difference deemed insufficient to warrant separation of the genera. Santesson considered Laternea to be a synonym of Colonnaria, recognizing the latter as the valid genus. Linder (1928) indicated that the “laterne” gleba]. position was common to Laternea. For his studies, he only had comparative material preserved in alcohol of Clathrus columna.tus, and therefore the gleba was misinterpreted, according to Santesson (1943), who did a detailed study of Clathrus co1ur~.atu~ and compared this to Linder’s photographs of Laternea triscapa.