Taxonomic notes on the Geastraceae , Tulostomataceae , Nidulari-aceae and Sphaerobolaceae ( Gasteromycetes ) sensu Bottomley , in southern Africa

Bottomley’s (1948) Gasteromycetes o f South Africa is still widely used for identification purposes. However, as a result of developments since 1948, the work has become outdated in many respects. Entries in the Geastreae (Lycoperdaceae), Tulostomataceae, Nidulariaceae and Sphaerobolaceae sensu Bottomley (1948) that require updating are listed and briefly


INTRODUCTION
Although Bottomley's (1948) Gasteromycetes o f South Africa was, in her own words, not in any sense a criti cal revision...' but '... merely an assembling of the known records of these fungi in Southern Africa', it still remains the standard source of reference with regard to the identi fication and classification of the Gasteromycetes of the re gion.However, errors in the original work, changes to the International Code of Botanical Nomenclature (ICBN) and particularly, considerable changes in Gasteromycete systematics since 1948, contributed to an unfortunate situ ation in which current users of Bottomley (1948) are at considerable risk of ending up with incorrect identifica tions or outdated names.In the families considered in this paper more than 70% of the entries in Bottomley (1948) are affected to a greater or lesser degree.
Since a com prehensive revision o f the southern African Gasteromycetes is still some years in the offing, there is a need for an interim guide listing those entries in Bottomley (1948) which may lead to the inaccurate iden tification and classification of specimens.In this first in stalment, the Geastreae (Lycoperdaceae), Tulostoma taceae, N idulariaceae and Sphaerobolaceae sensu Bottomley (1948) are considered.It is emphasized that this paper is primarily a reflection of views and ideas ex pressed in the literature since 1948, and that it does not claim to be a critical re-appraisal of any of the taxa con cerned.The principle objective is to provide an interim aid towards the more effective use o f Bottomley (1948).
The order of arrangement of the taxa listed below fol lows Bottomley (1948), and the taxon name and author citation heading each entry have been taken, unchanged, from that publication.The number in brackets following each heading refers to the relevant page number in Bottomley (1948).Entries in Bottomley (1948) which, to our present knowledge, do not require comment, are not included in the list.This also applies to names in which ANNOTATED LIST OF TAXA 1. Geastreae (586) The tribe Geastreae sensu Bottomley (1948) has sub sequently been treated at the family level (G eastraceae Corda\ order Lycoperdales) by most em inent gas teromycete taxonomists (Zeller 1949;Eckblad 1955;Kreisel 1962;Demoulin 1968;Ponce de Leon 1968;Dring 1973;Calonge & Demoulin 1975;Demoulin & Dring 1975;Demoulin & Marriott 1981;Sunhede 1989;Mornand 1993).Exceptions include Dorfelt and co workers who place these organism s in the order Geastrales (Dorfelt & M uller-Uri 1984;Dorfelt & Bumzaa 1986;Dorfelt & Heklau 1987).

Geastrum Persoon (586)
In accordance with the changes to the ICBN enacted in 1981 (Korf 1983), the appropriate author citation, in dicating the sanctioned status of this name, is G eastrum Per s.: Pers.
De Villiers (1994) has recently completed a revision of the genus Geastrum in South Africa, providing an up dated key to their identification.Persoon (588) According to Korf (1983) the author citation as used in Bottomley (1948) is acceptable in non-taxonomic works only.In taxonomic treatments authors are strong ly advised to use the more informative G eastru m pecti natum Pers.: Pers.
1.1.4Geastrum ambiguum Montague (591) Bottomley (1948) and Ponce de Leon (1968) regard Geastrum drum m ondii Berk, and Geastrum ambiguum Mont.as the same species.Dring (1964) and Sunhede (1989), however, both expressed the view that more m aterial should be exam ined before such a conclusion is drawn.A ccording to Sunhede (1989) both of these species are very sim ilar to G eastrum cam pestre Morgan.Dem oulin & Dring (1975), on the other hand, state that G. am biguum in Bottomley (1948) is the same as the G. drum m ondii of Dring (1964) and Dring & Rayner (1967), and that it differs from the type speci mens of G. am biguum and G. drummondii.They adopt the name Geastrum schweinfurthii Henn.for this fun gus.Bottom ley (1948) and Ponce de Leon (1968) re garded G. schw einfurthii as conspecific with G. am biguum.De V illiers (1994) does not follow Demoulin & Dring (1975) in recognizing G. schweinfurthii as a sep arate species, stating that '...there is not (sic) doubt that the southern A frican collections of G. ambiguum have been correctly identified by Bottomley The last word on the identity o f this fungus has probably not been spoken.1.1.5Geastrum quadrifidum Persoon (591) According to Korf (1983) the author citation as used in Bottomley (1948) is acceptable in non-taxonomic works only.In taxonomic treatments authors are strong ly advised to use the more informative Geastrum quadrifidum Pers.: Pers.
According to Sunhede (1989), Geastrum limbatum sensu Coker & Couch is the same fungus as the one de scribed by Lloyd as Geaster limbatus, cited by Bottomley (1948) as a good description of G. limbatum Fr.G. lim batum sensu Coker & Couch, however, is generally ac cepted to be a synonym of Geastrum smardae V.J.Stanek (Ponce de Leon 1968;Sunhede 1989).
A comparison of the descriptions in Bottomley (1948) and Sunhede (1989) indicates that Bottom ley's fungus might be G. coronatum rather than G. smardae.This view has recently been confirmed by De Villiers (1994).

Geastrum velutinum Morgan (599)
Ponce de Leon (1968) regarded Geastrum velutinum Morgan as synonym of the earlier described Geaster javanicus Lev., and created the new combination Geastrum javanicum (Lev.)P.Ponce de Leon.As explained in Demoulin (1984), however, Geaster is merely an ortho graphic variant of Geastrum, which makes Ponce de Leon's combination superfluous.According to Dring & Rayner (1967), as well as Sunhede (1989), a thorough re vision of G. velutinum and related taxa is desirable.De Villiers (1994) accepts G. velutinum as the correct name, but if this fungus is indeed conspecific with G. javanicum, the correct spelling and citation will be Geastrum javanicum Lev.This fungus must have confused Bottomley (1948) as well, since, while she regards it as a good species on p. 599, she also lists the name Geastrum velutinum as a synomym under G. saccatum on p. 597.

Geasteropsis Conrathi Hollos (606)
According to Sunhede (1989) and article 60.11 of the ICBN, the correct orthography should be Geasteropsis conrathii Hollos.Long (1945) placed this fungus in the genus Trichaster Czer.as Trichaster conrathii (Hollos) Long, while Ponce de Leon (1968) created the new combina tion Geastrum conrathii (Hollos) PPonce de Leon.In his authoritative revision of the G eastraceae, however, Sunhede (1989) is absolutely convinced that this fungus has no place in any of the above two genera and that it should be retained in the genus Geasteropsis Hollos.Sunhede (1989) warns, however, that his inclusion of Geasteropsis in the Geastraceae, as accepted also by De Villiers (1994), is tentative, pending further studies.
In her key to the genera of the Tulostomataceae, Bottomley (1948) included the genus Schizostoma Ehrenb.ex Lev.emend.Lloyd, although it had not yet been recorded from southern Africa at the time.Schizostoma laceratum (Fr.)Lev.has, however, been recorded since then (Talbot 1958) and appears to be fairly common.

Tulostoma Persoon (608)
The appropriate author citation, indicating the sanc tioned status of this name, is Tulostoma Pers.: Pers.
From Wright (1987) it is evident that serious short comings exist in Bottomley's descriptions of the southern African Tulostoma species and that numerous specimens cited in Bottomley (1948) have been incorrectly identi fied.A taxonomic reassessment of the Tulostoma speci mens at PREM should therefore be a worthwhile exer cise.
2.1.1Tulostoma album Massee (610) Wright (1987) regards this as a doubtful species, but accepts Tulostom a m acalpinianum Lloyd, which Bottomley (1948) cites as a synonym of Tulostoma album Massee, as a validly published species.As far as could be ascertained, T. macalpinianum has not been recorded in southern Africa yet.The specimen listed by Bottomley (1948) as T. album (PREM 28528) is, howev er, Tulostoma lesliei Van der By I (Wright 1987).

Tulostom a albicans White (611)
This name is to be cited as Tulostoma albicans V.S. White. W right (1987), however, regards T. albicans as '...an ill-defined species, easy to mistake for oth ers...', and excludes Africa from its distributional range.PREM 8764, listed in Bottomley (1948) as T. albicans, has been described as a new species under the name Tulostoma exasperatosporum J.E.W right (Wright 1983), while PREM 28638 is Tulostoma involucratum Long (W right 1987).
However, with two exceptions, Wright (1987) referred all o f the specim ens cited under T. bonianum in Bottomley (1948) (PREM 1344;1969;20378 & 30617), to Tulostoma verrucosum Morgan, which, in turn, is a synonym of Tulostoma squamosum (J.F.Gmel.: Pers.)Pers.(M oreno et al. 1992a).Wright (1987) regards T. pusillum as a fungus of tropical rain forests and does not include southern Africa in its distributional range.In the light of this, the status of T. pusillum in southern Africa requires verification.

Tulostom a brumale Persoon (613)
In accordance with the changes to the ICBN enacted in 1981 (K orf 1983), the appropriate author citation, in dicating the sanctioned status of this name, should be Tulostoma brum ale Pers.: Pers.Tulostoma nanum (Pat.)J.E.Wright respectively.In the light of this the identity of the other specimens cited as T. brumale in Bottomley (1948) is suspect and requires ver ification.
According to K orf's (1983) interpretation of the changes to the ICBN that were enacted in 1981, the au thorship of this name would be more appropriately cited as Tulostoma squamosum (J.F.Gmel.: Pers.)Pers.Lloyd (615) According to Wright (1987) the specimens cited by Bottomley as Lloyd Myc. Coll. 28934, 28958 are in fact Tulostoma purpusii Henn.The identity of the rest of the material cited in Bottomley (1948) requires verifica tion.
Although the closely related Tulostoma volvulatum var.elatum Har.& Pat. has been reported from Namibia, Africa is not included in the distributional range of T. volvulatum var.obesum (W right 1987).Bottom ley (1948) mentions a single collection of this fungus from southern Africa, but hastens to add that no material had been available for examination.Unless B ottom ley's (1948) record can be verified, the status of this fungus in southern Africa should be regarded as doubtful.
2.1.9Tulostoma MacOwani Bresadola (617) Orthographic error.According to ICBN article 60.11 and recommendation 60C .l.(b), the spelling of the epi thet should be corrected.The correct spelling and author citation for this name, first published in Petri (1904), therefore is Tulostoma macowanii Bres.ex Petri.

Batarrea Persoon (619)
As discussed in Coetzee & Eicker (1992) this name has been spelled in various ways.Bottomley (1948) used Persoon's original spelling which lends itself to correc tion as provided for by article 60 of the ICBN.In a num ber of recent works, Battarraea has been the preferred form of spelling (Rauschert 1986;Wright 1987;Mornand 1993 and several other authors cited in Martin & Llimona 1994).Another school of thought, however, strongly opposes this (Martin & Llimona 1994), recog nizing Battarrea as the correct spelling.Until this matter is resolved, we prefer to use the original spelling of this name, to be cited as Batarrea Pers.: Pers.Persoon (619) According to K orf's (1983) interpretation of the changes to the ICBN enacted in 1981, the authorship of this name would be more appropriately cited as Batarrea phalloides (Dicks.: Pers.)Pers.
2.4 Dictyocephalos Underwood (626) White (1901) attributed the name Dictyocephalos to Underwood.It is, however, not easy to determine from the original publication whether this should be treated as an "in' or 'ex' case as discussed in ICBN article 46 (Greuter et al. 1994).We therefore accept the citation suggested in Greuter et al. (1993), namely Dictyo cephalos Underwood ex V.S. White.
One of the specimens listed in Bottomley (1948)

Nidulariaceae Fries (631)
According to David (1993) this family name is not at tributable to Fries.It should be cited as Nidulariaceae Dumort.

Cyathus Haller ex
3.2.1 Cyathus dasypus Nees (634) Brodie (1975) regards this as a synonym of Cyathus olla (Batsch: Pers.)Pers., stating that 'Cyathus dasypus from South Africa is surely a form of C. olla with extra large irregular peridioles'.The peridiole measurements given in Verwoerd (1928) and Bottomley (1948) are, however, considerably smaller than the dimensions given for C. olla in Bottomley (1948), Eckblad (1955), Brodie (1975) and others.In the light of this, Brodie's statement does not make sense and might be worth investigating.Lloyd emend. Verwoerd (634) In his authoritative monograph of the Nidulariaceae, Brodie (1975), apparently unaware of the amplified de scription o f this fungus by Verwoerd (1928), lists it as a doubtful species which, according to him, cannot legally be recognized as a valid species, but if found again it should be easily recognized by collectors of African material because of the minute spores'.
According to W right (1987), T. brumale is a typical European species not occurring in southern Africa.He diagnosed PREM 20946 & 31371, cited as T. brumale in Bottom ley (1948), as Tulostoma rufum Lloyd and as P. pistillaris (PREM 27280), has been described as a new species by De Villiers et al. (1989), namely Podaxis rugospora De Villiers et al.