Studies in Schizaea Fistulosa Labill
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STUDIES IN SOHIZAF~ FISTUTJOSALABILL. A Thesis presented for the Degree of Mas of Science and Honours in Botany In University of Oanterbury, Ohristchurch, New Zealand. Ian ~ash December 1966. CH.. l\P'llER L Introduction 1 ClIAPl'EH I L External Morpho 11 CH.AP'I'1!~R I I I • Gametophytes. 40 Chromosome Number 48 S'oore M:orpholo 52 VI. Di tion and Ecological tors. CfIAPTEH VI I. Conclusions. 59 ReGord imens. BIBLIOGR.4.PIIY. a: Ins ----- accounts ~or about 30 (Sell 1944) 9 r-:'he Genus THE U[JRARY Ut-.liVERSITY OF CANTERBURY CHRJSTCHURCH, 1'II.z. CHAP'rER I. INTRODUCTION. The Schizaeaceae is one o~ the most primitive ~amilies in the }'ilicales and one o~ the oldest surviving ~ern ~amilies (Arnold, 19LJ-7). It is divided into ~our genera: Schizaea, Lygodium, Anemia and Ivlohria. These ~our genera include elL approximately 175 species, o~ which Schizaea"was ~irst described by Smith in 1793. Since then there have been attempts to divide it into more genera, but it is now divided intc sections and sub-sections which as listed below are ~rom Selling (19LJL~). Section I. Euschizaea, Hook. Sub. section 1. Pectinatae, Prantl. Sub. section 2. Bi~idae, Prantl. Section II. Actinostachys, Wall. (Synonym. Section I." Digitatae, Prantl). Section III. Lophidium, Rich, Sub. section 1. Dichotomae, Prantl. Sub. section 2. Elegantes, Prantl. All the species, except one, Which are mentioned in this thesis are in Section I, Sub. section 1, the exception being Schizaea. bi~ida Sw. which is in the 2. The systematic classification of Schizaea fistulosa sensa lat. has been far from clear since \ , the original d.escription by de Labillardiere in 1806. The object of this thesis is to endeavour to clarify this situation and to either confirm or disprove the valldity of the grouping into S. fistulosa Labill. (Plate No.1) and S. fistulosa vare australis Hook. f. (Plate No.2). To carry out the study on S. fistulosa sense lat. four lines of observation have been followed:- 1. General external IT;orphology. 2. Spore morphology. 3. Development of gametophytes. 4. Chromosome nurrillers. The first two of these have been carried out on all available material from the total geographic range of the species~ while for the latter two live material collected in New Zealand has been used. To all material, live and herbarium, I have given a number of my own for convenience when quoting in the text. A list with corresponding Herbaria numbers appears in the appendix at the end. CLASSIFICATION. To understand fully the problem and reasons for the investigations a summary of the systematics as they Plate No. L A specimen typical of the species. Plate No.2. A specimen typical of the variety. (No. 66). ~ , ", ~ , ~ . \ . I , ) ! J ,'. , ~: .. ' I \ \ 1. I . ~ ~.' ~ , r o , . r ' ... " J . ,. ~ ., .... - --:.-~--::--~ .- ..- .-: -- ............. -- ~ .... ... - 3. appear at present, which includes all synonyms and classifications that seem to have occurred, is gi ven below. Allan (19 ) es only S, fistulosa Labill. wi th two synonyms So propingua A. Cunn. and (Lab ill.) Reed. In his discussion he quotes Hooker (1867) in connection with var australis ...;;....;...;;;.;;;.::=~;;;.;; Hook. f. To this he (Hooker) gives two synonyms s. australis Gaud. and .=.;;..--r;;=...::::..;;;..:=::c:...::..:;;,; Hombr. and JacQ.. Hooker ( ), in fact, originally classified them as var australis. These he later changed to and S. fistulosa and pointed out that the conf'usion had come from his reliance on Cunnlnghams herbarium mater which both S. bif'ida and s. f'istulosa were labelled ~~~~~~= Hooker and (1874) reverted to the idea of two species and S. australis, the f'ormer with and S. propingua in part as synony~s, and the latter with S. palmata as a synonym. The rest of he put into the ..;;;.,;;;-,.j;::..=....:.....:;,.;=~= unbranched section It will be noted here that there been a change from S. pectinata to as synonyms: this point will now be discussed. 4. There appears to be confusion over the use of the names S. pectinata and S. palmata. From what can be found out one is correct and the other is used out of context. S. palmata is the only Schizaea described by Hombr. and Jacq. in 1872 and is equated with S. fistulosa var australis by Cheeseman (1925) and Christensen (1906), and is equated with S. australis by Thompson (1882). It appears that its association with this section is the correct usage. I have not been able to find.. where 'rhompson (1882) and Hooker and Baker (1874) got the notion that the description or s. pectinata came from the same source~ i.e. Hombr. and Jacq. Christensen (1906) uses the name but for an entirely unrelated species from South Africa~ as does Selling (1944). Richter (1916) refers to S. pectinata J.E.Sm. This is the same authority as that for the genus. Simm (1915) and Christensen (1906) both give the authority as (Linn.) Sw. This apparently just being a genus change from Acrostichum to Schizaea and it may well be that the Sw. should be Sm. Thompson (1882), gives two species in New Zealand, S. fistulosa and S. australis. To S. fistulosa he gives the synonyms S. valdiviana, 5. s. birida in part and S. propinqua in part. This is probably the results from a combination of the racts from Hooker (1867) and Hooker and Baker (1874). To S. australis he gives three synonyms, S. palmata, S. propingua and S. fistulosa var australis. Cheeseman (1925) reverts to the classification of Hooker (1867) with the exception that in the synonyms he substitutes S. palmata Hombr. and Jacq. for S. pectinata. He holds that the variety is just a depauperated form connected to the species by transitional stages. The next classification of note is that of Allan (1961) which is quoted at the beginning of this section. This then gives the classification within New Zealand and shows a differing or views of the systematics, i.e. one species with a variety, Hooker and Cheeseman: two species, Thompson and Hooker and Baker: one species only, Allan. Moving to a wider field we find that all of the type specimens are non New Zealand. The species type is from Tasmania and S. australis, described by Gaudichaud in 1825, is from the Falkland Islands. S. valdiviana type is from Corral and S. chilensis from the Archipielago de Chones, both in Southern Chile. 6. Philippi first described S. valdiviana and S. chilensis as different species but on receiving §,nother specimen from between the two type localities he put them all into the one species S. chilensis (Looser 1937). In the other areas of the species range there was also confusion over the groupings. Prantl (1881) included S. malaccana Bak. (from New Guinea, Malaya and Borneo), and S. robusta Bak. (from the Hawaiian Islands) as varieties of S. fistulosa. In 1934 Christensen and Holttum, in their paper on the ferns of Mt. Kinabalu, give S. malaccana as a synonym of S. fistulosa. In the same paper they describe S. malaccana var robustior which they hold is superficially similar to S. fistulosa and S. robusta. Since then Selling (1944), Holttum (1959) and other workers have been convinced that these are all separate species. This fact, along with the grouping of other synonyms, is discussed below. The inclusion of S. robusta within S. australis drew support from Braclcenridge, Mann, Derby and others (Selling 1944) who regarded it as a tropical form of s. australis, and in 1875 Luerssen gave it varietal status, S. australis var robusta. Richter (Selling 1944) considered S. australis should be a separate species with as a synonym, and within included S. chilensis and as synonyms. This would seem out of keeping with the descriptions of these Chilean c , there appearing to be one from of the two groups. Rosenstock (Selling 1944) appears to have been the first to have suggested that the Chilean species be included within S. fistulosa when he identif one of de Labellardi~res specimens from as S. ristulosa var chilensis. Further cor~usion is 0 d by the suggestion from Fernald (S ing 1944) that S. australis and Pursh. (North American) could well be one species. s is not the case as is shown at the end of this er. Sell (1944) is convinced that by spore morphology there is only need for the spec with no variety., At one stage it was thought that one group of specimens from Madagascar were S. fistulosa. This is but one of the stions on the classification of this group. was not happy with the classification in 1944 and after a thorough survey of the group described these ecimens as a separate species, S. confusa, which he published in 1947. This, he pointed out, is only 8. remotely and indirectly related to S.fistulosa through several other species. The result of all this, I feel, is that the synonynti ty should be:- S. fistulosa Labill. sensa Lat. a Microschizaea fistulosa (Labill.) Reed. 1947. ~ Yare Malaccana Prantl. 1881. a S. valdiviana Phil. 1868. a var. australis Hook.f. 1867. a S. chilensis Phil. 1850-60. ex S. palmata Hombr. and ,JacQ. 1852. ex S. propinqua ~t:.~. Cunn. 1852. a S. australis Gaud. 1825. ~ s. bifida Willd. (in Hooker 1867) • ~ yare robusta Prantl. 1881. f3 S. pectinata (Linn) J.E. 8m or Sw. (in Hooker and Baker 1874). f3 S. australis var. robusta Luerssen 1875. a completely included within S. fistulosa sensa lat. now. ~ Once included within S. fistulosa sense lat. by oversight or genuine description but are now recognized as separate species. S. fistulosa was once used for the Madagascar species which is now S. con:fusa Selling (1947). 9. STI~ONYMS WITH SPECIFIC RANK.