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.
In view of the mention of S. bifida, S. malaccana,
s. robusta and S. pus ilIa being associated with
S. fistulosa a brief examination of the external
morphology of them was undertaken in order to ensure
that they are distinct.
S. bifida (Plate No.3) Specimen No. 100.
There is no doubt that this species is distinct.
The rhizome scales are very irregular in shape and
less dense on the rhizome than in S. fistulosa and the
colour of the scales is a very light brovm. Division
of the frond, the presence of many fine projections
from the sides of the sorophores and the nature of the
spores are also distinct.
S. malaccana (Plate No.4) Specimen No. 48.
Stomatal arrangement in grooves on the frond,
scale colour and the shape of the frond in cross
section are all distinctive enough to make it a separate
species without the aid of any other characters.
All these differences are clearly described by
Holttwn (1959). Plate No.3. s. bifida Willd. (No. 100)
Plate No.4. s. malaccana Bak. (No. 48) 1 . ' .
'"" , "
.. 'LOI .. ,.. ~ .....tfil.W JlALAHO "",1- _ .------',~ ...... L· "'J'\ ' 10.
S. pusilla (Plate No.5) Specimen Nos, 94, 95,
96,97,98,99.
The most obvious distinguishing feature here is the presence of many short sterile fronds. These are flat in contrast to the round nature of the fertile fronds. Added to this is the presence of fine projections from the sides of the sorophores and an almost complete absence of scales on the rhizome~
s. robusta (Plate No.6) Specimen Nos. 91, 92, 93.
Frond shape, stomatal position on the frond, and the presence of many sterile fronds are definite distinguishing features of this species.
These four specimens along with S. pectinata anci S. confusa, which were not seen but are well described b;y Simm (1915) and Selling (1947) can all be seen to be distinct from one another and from
S. fistulosa. These differences are further added to by Selling (1944) in his work on the spore morphology of the genus Schizaea. PI e No.5. Pursh. (No. 94)
te No. Bak. (No. 93) ,.
\ \ ( I ! ) .1 "'~ \~ _ j, f , "f
\
11111 . " I'. " ...... - \ .... . 11.
CHAP'rER II.
EX'l'E:Rl'~P.L MORPdOLOGY
MORPHOLOGI CAL CHAllAC'l'EF S OBSERVED.
In all the specimens from New Zealand and its associated Islands, as well as in the non New Zealand specimens, fourteen characters have been selected for study and comparison. These will be dealt with in order generally and then divided into Geographical groups, the variations discussed and finally all the groups compared.
1. Frond length.
This is tho ehtire length of the frond from the rhizome to the top of the fertile zone.
2. Fertile zone length.
This is the length of frond, at the top, that is occupied by the sorophores (as used by Holttum
(1959) ); these lengths va'ried greatly as is shoV'm in
Plate No.7.
3. Fertile zone width.
This is the width of the fertile zone taken at the widest part. It is in effect the length of the longest sorophore.
4. Sorophore width.
The width of the sorophore in all specimens was almost constant varying little either side of 1 mm. Plate No.7. Variation in fertile zone 1 (Nos. 23, 4 and 37).
Plate No. 8~ tile zone division. (No. 20)
12. throughout. This feature is the same all along the sorophore.
5. Sorophore nrunber.
This is the number of sorophore pairs per frond.
6. Sporangia number.
This is the number of sporangia on each sorophore (an average for each frond).
7. Fronds per rhizome.
This is a count of the number of fronds on a given length of rhizome, 1 cm., as near the apex as possible. In many this was hard to determine as specimens had Eot been cleaned before mounting.
86 Stomata nwnber.
For this areas on the frond were selected at random and over a distance of 3 mm. the number of stomata were counted.
96 Sorophore division (Fig. No.1)
In many it was noted that the sorophores di vid.ed, especially in the lower half' of the fertile zone. In some this occurred very clearly while others showed ju£t a slight fusion near the base of the sorophores. In one specimen, No. 87, there was a tripinnate diviSion but this was the only occurrence of this feature noted.
Other specimens observed had a complete division of the Fi g. N~ 1.
Point of Sporangial ~..a.----- attachment.
pical ra ore Division from Specimen N?1. The shape of ch individual is simil to aU Sorophores. Size: 1mm. x 3mm.
Fine projections/hair VQry prominent in S. biUda. fertile zone ( ate No.8).
10. Stomata arrangement (Fig. No.2).
The stomata extent in all is from the base to just below the ferti zone and in most specimens they occur in two single rows. In some cases over a short distance the stomata were arranged more than one wide. This was either in two rows side by side or just an indiscriminate grouping of stomata three or four wide.
11. Sporangia arrangement.
This is in two rows with the sporangia roughly alternate ther side of the sorophore mid rib.
In some the is much more regular than in others.
12. Frond diameter.
This is a variable character and one inf'luenced by drying of' herbarium eCimens, but does give some indication of extent of variation that there is within specimens assuming they all contracted by a proportionate amount.
13. Frond scales (Fig. No.3).
These are situated the full length of the frond but are ]~os t with so that there are generally only a few left at the base by the time the frond has reached maturity. They are brown with the apex pointing up the frond and they are borne on special support cells that mayor may not be obvious. In many cases these support Plate No.9. Scale density and orientation o~ the apical end of a rhizome.
Plate No. 10. Dichotomous division of the rhizome. (No. 20).
Fig. N~ 2. Stomata from N? 27.
Normal Stomatal arrangement with a detailed diagram of a stomata. Density: 14pcr 3mm.
Base and. 14.
cells become more obvious after the scales have fallen off~
14. Rhizome scales. (Fig. No.4).
The rhizome is clothed, especially that of the
last two seasons growth, with long scales. These are
densely packed and all directed upwards (Plate No.9).
They have features, colour, length and shape, which
vary to a degree that is felt to be of significance for
the seperation into groups, if this is possible.
In the text the scales length is referred to as units,
each unit being equal to 0.15 mm.
One feature that was very common in growing
specimens and in some herbarium sheets was the
dichotomous division of the rhizome as 1s shown in
Plate No. 10.
An abnormality was observed in Spa No. 20 where the frond was divided (Plate No. 11).
PART I.
NON-NEW ZEALAND SPOROPHYTES.
To discuss these I will divide them into geographical groups 1. e. V'Ies tAus tralia, Tasmania, New
Guinea, New Caledonia, Borneo and Chile. Two specimens from West Australia were examined, five from Tasmania, nine from New Guinea, five from New Caledonia, one from
Borneo and five from Chile. Plate No. 11. Abnormal division the frond. (No. 20).
Fig. N~ 3. Frond Scales,
c ~ "> ,/ / / / / / / / / // / / / / / / /-
Side v'lew of a scale and support cell trom N? 21.
support cell base end scale, •
- - -c: - --
Scale a nd support co II sh ow n in relati on to the C?pidermal cell s. Note the diffC2r<2nCa in call size. (N?27) 15.
These have all been observed for the fourteen characters listed at the beginning of this chapter.
The variations, with the possible exception of two, I feel can be related to ecological conditions or are not consistent in their differences.
The two main exceptions are the nature of the scales on the fronds and their support cells, and the rhizome scales. These characters vary to a degree that seems to be of importance; the others vary so widely that they are of no significance as characters to divide the species into varieties.
LOCALI'nES OF 'rUE NON NF.:w Z-P~A[,pj\m SPECIES
To help relate characters to their habitats there is now included a list of the specimens with, where possible, a note on the area where collected.
I have had to rely on the herbarium labels and these are not always very informative.
WEST AUSTRllliIA
No. 17, among sedges in surface of deep peat.
No. 37, no information.
TASMANIA
l'Io. 38, on button grass plain. No. 39, no information. No. 40, no information. Fig. N'.' 4. Rhizome Las.
a
Nt? -Typi cal of tho main NawCalodonia group:!- Langth: ',Smm.
b
, .. N?l1. Normal scale typo. Typical of all specimens but the New Caledonia group. Length: 3 mm. 16.
No. 54, no information.
No. 55, no information.
1'1 EW GUINE..l\
No. 18, alpine grassland, 3590 m.
No. 19, alpine shrubbery - on peat soil, it
would seem sheltered, 3270 m.
No. 41, open rocky slope, 3750 m.
No. 42, alpine grassland in boggy ground, 10,300 ft. No. 43, bog grassland, 8,800 ft. No. 44, open rocky slope, 3,750 m. No. 45, alpine grassland, 10,000 ft. No. 47, open places sub alpine scrub vegetation, 3380 m.
No. 49, bog grassland, 8,800 ft.
NEW CALEDONIA AND BORNEO
No. 46, stunted gnarled mossy forest, ultra
basic sub soil, 2500-2900 m.
No. 50, in bog beside stream, 160-170 m. ,
appear slightly sheltered.
No. 51, forest floor in mist zone, just inside forest, 3,700 ft.
No. 52, open ridge in scrub t 2,500 ft.
No. 53, among scru-o on steep hill, 2,500 ft, -. , No. 90, :8'oret a Nothofagus. 17.
CHILE
No. 59, bogs at sea leveL
No. 60, 2,000 :ft.
No. bl t amongst -tussoc:kS.
No. 62, swamp at sea leveL
No. 63, in bog.
WESrE AUSTRALIA
Both specimens observed from here, Nos. 17 and 37,
(Plate J.\o. 12) were limi ted in the material available although general observations were possible. The most conspicuous feature was the long (28 em. and 48 em.) fronds and the long fertile zones on them (35 mm. and
Although the fertile zones are long they do not have a correspondingly great width,- being on an average
3 mm. wide and never more than 4mm. The sorophores are all close together giving a very regular appearance to the fertile zone. The sorophores, about 25 pairs per frond, all support 14~15 sporangia in two rows.
In 1'1"0. 17 some of the sorophores showed signs of a bipinnate condition. This was not confined to the few basal sorophores but occurred all the way up the frond.
Sorophore width and frond diameter were constant in both specimens i.e. 1 mm. and 0.75 mrn. respectively.
Stomatal density was not great, both having an average of 10/3 mm. In No. 17 it was noted that in Plate No. 12. A specimen from West Australia, (No. 37).
Plate .No. 13. A specimen from Tasmania. (No. 38). z ~ Z 11"-:1- '"~'1 !' l> I o.n<>.~ ;:.19 -" 0., c ~ ,:....~ -I ~IiHuuulJ L..I 0 117 .. ,I'tCM$ if ....." ~ ~ z . scale 't ~ r l> , 1 :- r- ~ \ ::z:: '" :;'. ~ '"0 m ti" I' r- 1f -< ::= rl r-> Q' ~ l> ~
Cotf. e. "",nt-•• I If•. JlsJ. Notes J .. - f,..A"t: .. t,.~ ~./I.. ..,m ; ".",1<1 •• -__ ~vl ..... ,., ...... - ' 180
sections the stomata were in a double row. The frond
scales are obvious in both, being larger in No. 37 than
in No. 17, although both have visible support cells
that are also apparent after the scales have fallen offo
The rhizome scales were only available from No. 17.
These were brown and an average of 21 lUlits in length,
which is slightly longer than in other specimens observed.
They are of the normal form as shown in Fig. No. ~.
and consist mostly of two cells with an occasional one
of three cells.
These plants are different from the Ne.w Zealand
specimens in their great frond length and slightly
darker colour of the rhizome scales.
TASIVLA}jIA
There are five specimens collected from this area
Nos. 38, 39, 40, 54 and 55 (Plate No. 13). They are, with the exception of No. 40, all of
similar height averaging around 16 cm. in length.
No. 40 has plants which are much longer (30 cm.). The fertile zones are of a consistent length (13 mm)., bearing 10-14 pairs of sorophores, that are on an average 3 mm. long and 1 mm. wide, close together to give a compact appearance. The sorophores bear an average of 13 sporangia each, arranged roughly alternately in two rows. In Nos. 38, 39 and 54 there are definite signs of sorophore division. 19.
Stomata are consistent in all at 16-17/3 mm. with NOSe 39, 40 and 55 all having double rows in places.
In all but No. 40 the frond scales are obvious with support cells that can be easily seen after the scales have fallen off. These support cells project above the level of the rrond surface. In No. 40 the scales are present and of a similar size to the rest but the supports are not quite so obvious.
Frond diameter was similar to other groups being between 0.5 and 0.75 mm.
Only two records of frond density on the rhizome were possible and these were both 14-15 fronds per 1 cm.
Rhizome scales were only available from Nos. 54 and 55. There is a variation in colour, with No. 54 being brown and No. 55 dark brown. There is also a variation in their length, No. 54 being 17 units and
No. 55 about 25 units. However there is no variation from the normal shape, No. 54 generally having two cells and No. 55 mostly three cells but an occasional one with rour or two.
These do not vary greatly, except in Size, rrom the specimens of West Australia. 20.
NEW GUINEA
There are nine specimens from this area, four
from Australian New Guinea, four from Papua and one
from the Indonesian sector. The specimens are in
order Nos. 18, 19, 41, 44, 42, 43, 45, 49 and 47 (Plate No. 14). Two pairs of these specimens are
duplicates from the same collection but different herbaria, Nos. h3 and 49 and Nos. 41 and Lt4.
COLsistent characters evident in this group of
specimens are the arrangement of sporangia in two
rows and stomata in a single row in all observed
specimens. In all but No. 42 the density of
stomata is from ll-l~ per 3 mm. while in No. 42 it
is 22 per 3 mm. It is possible that this is not a
true indication as the material was limited. As is the
case with all other specimens the width of the
sorophore is just over 1 mme
Four specimens showed no division of the
sorophore but there is no significant combination of specimens to be of importance~ It is of interest here that 1-;os. 41 and 44 (duplicates) differ on this point.
Frond diameter in all is between 0.5 mm. and
0.75 mm. The scales borne on the frond vary in number, a factor that I feel is influenced by the way and time they have been pressed and age of the collection, from many to very few. They are all of Plate No. A specimen from New Guinea. (No. 42).
Plate No. 1.5. A specimen similar to S. bifida fro~ New Caledonia. (No. 50). }'bllh of l\w 5mlt.1\\<;,.h~v!l PadJi.~ ";1;""- C"'lU.UV!l>IA 21.
a normal size except in Nos. 18 and 19 v{here they are just slightly longer. The support cells in all specimens are of medium size and in young and old material all are obvious.
:Prond length varied from just lli~der 6 cm. in length to nearly 20 cm. This factor appears more and more to be ini'luenced by ecological condi tions especially altitude and exposure, (See :Pig, No, 5).
In most cases there is some correlation between the length of the fertile zone and the frond length.
The lengths of the fertile zones being from 5.6-15.6 mm. on an average. These are, as would be expected, related to the number of' sorophore pairs, 7-12, and nUlnber of sporangia per sorophore, 8-16~ There is no suc~ similar relationship between these features and the width of the fertile zone, which ranges from
2-3.1 mm. except in No. 49 which averages 4.1 mm ..
However they all give a similar visual appearance of being short and broad with the sorophores close together.
The density of fronds varies from 9-20 per
1 cm. of rhizome.
fu'lizome scales vary in colour and length, this latter being from 12-15 units on an average with the extremes at 10 and 20 units. With the exception Fig N°S Frond lQngth v Altitude for New Guinea Mat~riaL
20 ><49 19 18 17 III 1.6 E ><43 (.) 15 .-c .- ..c. 14 0 13 ><45 c (!,I 12 E 11 C!J 0. (J) 10 x42 9 ><19 8 x18 x44
7 )( 41 6 )(47 5
4 3 2 1
8 9 10 11 12 13 ALtitude in 1000's of feet. 22. of Nos. 43~ 45 and 46 they are all brown in colour,
the former two being slightly lighter and the latter
is darker than the rest. They are all of the normal shape and consist of two cells with a few of either one, three or four cells. No. 47 shows a slight difference in that most of the scales are of three cells.
The results once again show similarity to the
Australian material, with no significant features to distinguish varieties from the species.
NEW' CALEDONIA J\ED BORNEO
From this area six specimens, No. 46 (Plate
No. 17), 50 (Plate No. 15), 51, 52 (Plate No. 16),
53 and 90, were examined. All these specimens differed, in varying degrees, from the Australian and New Guinea material as well as from one another.
They can be grouped into three sections, a. No. 50 b. No. 90 and c. Nos. 46, 51, 52 and 53. There is no doubt they are all S. fistulosa as keyed from Holttum (1959), but they exhibit small differences that could cause confusion.
No. 50, at first glance, gives the impression of being an unbranched specimen of S. bifida, the deception being brought about by the nature of the fertile zone which is short, 8-12 mm., but comparatively P1a te No, 16. A specimen from the main New Caledonia group. (No. 51).
A suecimen from Borneo. (No: 46). FLORA OF 'v
., (; ..,;l ...· ·. '11
#1J7 ~ Of :iC ~ f 0 "'''' --- ,~o: I. ""- n. __ ~.I,. "t. iIf. . ,:~O,"":, 1._,
·.:rr.- ':: ~;-.... t •. " ::&I$~ ~iec
,.~U\l't~ . "l!~. ~ 1)' - .)t.r"'b. 1" .....11
L:.:. . LJ~_.L~_~ L 1 ~ ~ J .. " /~ - ' • ...... f , +. , ;",-- C, ...... I, 23. wide, 4-5 mm. sorophores are slightly less than 1 mm. wide and are ely packed to give the appearance (See c.r. in Plate No.3).
Rhi~me scales are of the shape, size and colour seen in Australian and other mate The rrond scales are ver-;l fine and rew in number. Their t cells are indistinguishable rrom epidermal c t not b sed or of a dirferent shape as was no d previously.
s reature is common to all six specimens in this group.
Spo a size nQmber, stomatal density and sorophore rs vari from Australian and other material but not
No. 90 is separated out because of its greater size ~ fronds 38 cm. f'ert zone 1 , 28 mm., and width, 6-7 mm. making it much 1 r, but correspondingly so, than the next four. The stomata are only exception being 12 per 3 mm. which seems normal r a 1 frond, while in the others it is up to 24 per 3mrn. This specimen lacked rhizome sc a factor that would have been of major assistance here.
Nos. 46, 51, 52 have black to near black rhizome scales. latter three these are not of the normal shape but more bead like (Fig. No.4).
'fhey are 10-11 units long and consist 5 cells c. f.
No. 46 where they are 28 units, 3 c s the normal shape. Frond scales are as described for No. 50.
The fertile zones are 12~14 mm. long and 4 mm. wide
on fronds from 7-33 cm. long. The sorophore width is
approximately 1 mm. and they are more widely spaced
giving an open appearance as against the neat compact
form in No. 50 or No. 17.
Sporangia are small and nwnerous neatly arranged
on the sorophores. The number of sorophores per frond
is 11 pairs in all four with signs of sorophore
division in all but No. 46.
One final factor that was noted in all six
specimens was the slightly flattened nature of the frond.
This may have been caused by pressing but has not been
seen in any other material.
'l'hese six leave the impression that 11vhile they
are defini tely s. fistlJ.los~ they are slightly distinct
from the Australian and other material observed and
may form a distinct geographical race.
CHILE
The material observed from this region was all
collected by Dr. Godley during the 1958/59 Royal
Society Expedition to the area, and consists of five 8uecimens Nos. 59, 60, 61, 62 and 63
(Plate No. 18). Plate No. 180 A specimen from Chile. (No. 60). _._ ...J _ _-W
~ .: o ' t.:. .• ,
; ;I .... ~ .....,;..t·r? 1) "'1": ,: :- "" 10 ,.: "I .. ' >"~ ~,, ;'.. ~ _ (.~ ...
1- :~":" ...... " _ • .:;' r':,:" .. . -~. -"7 ...... 25.
These show a remarkable uniformity with the exception of No. 62. This is interesting as it once again raises the Question of the varieties being
jus t tne result of ecological influences. 'fne other four specimens are all on an average 8 cm. high with the fertile zone 7.3 mm. in length but No. 62 is
13 cm. high and the fertile zone is 10 mm. long.
The first four also have denser fronds on the rhizome which average 30 per 1 cm., and denser stomata,
20 per 3 mm. of frond. In No. 62 these figures are
16 fronds per 1 cm. of rhizome and 13 stomata per
3 mm. of frond. No. 59 was the only specimen with a double row of stomata.
The rertile zone width, 2 mm., and frond diameter, 0.5 mm., are corrtant in all five specimens.
Sorophores per frond and sporangia per sorophore are also constant in all, being 8-9 sorophore pairs per frond and 8-9 sporangia per sorophore. All specimens showed a tendency to division in the sorophores.
Frond scales are all of the normal size borne on slightly smaller, but no less obvious, support cells than in the Australian and other material.
Rhizome scales were observed on all but No. 63.
These were all brown varying slightly in length and number of cells .. Nos. 59 and 60 were an average of 26.
11 units long with Nos. 61 and 62, 16 units long. Cell number in No. 59 was 1-2, in Nos. 60 and 61 all were either 2 or 3 and in No. 62 most were 3 with some
2 or 4 celled.
Once again the conclusion is that there is no single character or group of characters that can be used to distinguish varieties within this geographical region of the species.
From the study of all the material from outside
New Zealand there appears to be a noticeable uniformity of features with the possible exception of the New
Caledonian material. The major variation is the frond height and with it, as would be expected, the fertile zone length. (See Fig. No. 6 for averages of all characters). This I feel is purely the result of envirollQental influences of shelter, altitude, latitude and to some extent soil nature, i.e. peat or well drained, but at all times, poor soils, Fig. No. 7 shows several characters, for all the non New Zealand material, plotted with their altitude shown. 1. Frond length.
2 .. Fert zone 1 ength .. 3. Fert e zone width. 5. Sorophore number. 6. Sporangia per s reo
7. Fronds per 1 em. of zome. 8. Stoma per 3 mm. of
9. Sorophol"'c 3ion~ sion..
n "'" no di ion.
10. Stomata arrangement.
si = s e rO'NS, d = double rows. Fig. No.6. TilliLE OJ? AVERA.GE SIZj,;S _!'L'm lWMBFJtS ------POrt N(}N FE~W ZEAI,~lI2\:D hl/,-'rERIAJ--,"
---- Species Character number number. --"."~. 1 2 3 5 6 7 8 9 10
------~--. WEST AUSTRAllIA 17 40 20.3 3 20 14.5 10.1 9 s d 37 21 2Ll- 2.5 29 15.4 15 n si
'r AS M..--'l.N I A 38 16. 7 13 2.8 10.2 17.3 15 s si 39 15. 7 12·7 3.2 12.8 13 27 s d )-\.0 30 20 4 15 15 21 n d 54 13.4 11. 2 3.3 8.9 13.9 15 16.4 s si 55 17.6 15 3.3 1=, 14.8 U~. 16.3 n d t,~F~:!v (j·UIN:H:A 18 8.2 8 2.3 8 12.2 10 13 s si 19 9.1 7 ') 7.7 8.3 12 12 n si 41 7 5.6 2 7. u 8 1~ 12 n si ~L1 8.3 10.1 3.1 9.8 8.5 12 11 s si ,.-, 1.1·7 5.8 6.9 ~ 6 6. LL 9.5 20 U! n si 42 9.6 8 2.4 8.9 11. 7 17 22 n si 43 1.5.9 7.2 2.8 e 10 8 13 s 8t 45 12.7 11 3 9.3 10 13 13 8 si 49 19.5 15.5 4.1 11. 6 16 9 11 s si NEVI C)'I..LEDONI A j\j:;JD BORNEO 46 27.8 13 w- 10 13 10 13 n si 50 21 9.6 4.4 7.8 15.2 9 19 s si 51 31 20.5 L-+ 14.8 26.4 15 14 s si 52 10.8 9.4 2.6 10·5 14 16 18 s si "2: 53 14 14 -' 12 18.7 15 19.5 s si 90 38 28 6.5 15 35 12 s si CHIL:E 59 8 7.5 2.4 8.3 9.3 25 19 s d 60 8.6 7. 6 2.6 9 7.6 30 19.6 s si 61 7.8 7.1 2 9.1 9.5 hO 21.2 s si 62 12.6 10 2 8 9.3 ].6 13.6 s si
~ 63 7.9 7.2 2 8 8.5 35 23.2 0 SJ 40 Fig. N'? 7. Non NGlw Zealand material. 38 --Frond length in ems. 36 -·--Fertile 'lOM lG' ngth in mm. 34 ---Stomata PQf 3 mm. 32 30 28 26 .A. A 1\ i \ I \ 24 I \ \ J \ ~ 22 i I \ 1\ ;' " J \ 1\ I , 20 I 'L 1\ :. \ I \ I \ '7) I \ \ / I A I I HI \ I \ II I \ / \ ,: :/\ \ I \ /1 I \ I' \ \ \ / \ 16 \ I 1/ \ I I : \ i ;':\ \ r··!: ...." I \ / I, \ I \ 14 / \f/ X I '. ... I \ ...... ,.1 \ / \ ) \ ,/ ,/ lI- : I '\. '''I .' ~ I '" \/ 12 /" \/\/..! V' 10 , V \ / a " \I 6
4
2
O~~17 ______90 51 40 46 ------~ 37 50 4S 55 38 43 39 53 54 45 62 52 42 19 60 44 18 59 63 61 41 47
SpocimGn NumbsF 27.
PART
NEVI Z:F~ALP.Jm SPOIWPHYTES
li'or convenience the Hew Zealand specimens have
been divided into eight geographic groups. They
will be described within these groups then the whole
range summarised. The groups are:-
AUCKLAND AN]) CAMPBELL I SL.AJ'IDS
Nos. 64, 65, 66, 67.
S'rEWAR'l' ISLi\.ND
Nos. 13, 15, 16, 34, 35, 81, 85, 86, 87, 88, 89.
SOUrNI-SOU'fH ISLAND - ]'IORDLAHD 80U'l'H
Nos. 14, 75, 76, 78, 79, 82, 83.
SOTITil CEJliTRAL-SOU'l'lj ISLJli'm - FICRDLAi.m '1'0 CHARLESTON
Nos. 2, 3, 27, 28, 29, 30, 56, 57.
Nos. 1, 20, 21, 22, 23, 24, 25, 26, 84.
FORTH-sourrE ISLAI~D - KP..RAMKII.. NORTH
Nos. 4,5,6,7,8,9,10,11,31,32,33,58,77,80.
SOUTI.! - NORTH ISLAND
Nos. 12, 36, 68, 69, 70, 74.
NOE'fH - NOwrT-I ISL.AND
Nos. 71, 72, 73.
AUCK..J~AND P.ND CAMPBl!iLL I SLMTDS
The four specimens from these two islands, Nos.
6L!., 65, 66 and 67, were all very similar. Constant 28.
features in them were the frond diameter of 5 mm~, sorophore width of 1 mm. the arrangement
sporangia in two rows. Almost const is the arrangement
of stomata in a s e row except for • 66, and general lack of sorophore division whi was only evident in • 65.
The fronds were all short wi th N.o. 66 having an average 1ength of 3.2 cm. and the rest between 5.4 em. and
6 cm. length. • 66 also had a shorter fertile zone
5.8 mm. with the rest from 6.2 to 7.1 mm. length. width of the fert zones averaged 2.5 mm. and ea.ch consis of between 5 and 8 rs of sorophores v.:hich bore, on an average, 8 sporangia per sorophore.
The density of fronds on rhizome was constant averaging 18 r cm.
Stomata arrangement has been mentioned but not density. This varied with Nos .. 64 and being on an average per 3 mm. frond Nos. 66 and 67 averaging
per 3 mm.
Scales on fronds of all imens were normal in size with obvious support cells, although No. 66 had slightly smaller but no less obvious support cells ..
Rhizome scales were all slightly lighter in colour than those of non New Z materi There was a variation in cell nun:ber in the scales but not in shape which was normal. The cell number was predominantly
3 wi th .No. 65 and 67 having a noticeable number of
2 celled scales.
STEVV.AI{T ISLAl~D
There were eleven specimens~ Nos. 13, 15, 16, 34,
35, 81, 85, 86, 87, 88 and 89 from this area three of which
Nos. 16, 34 and 35, were kept growing at the University.
Constant features were the frond diameter of 0.75 mm., sorophore width of 1 mm. and s:porangial arrangement in two rows.
The frond length varied from an average of 6 to
17 em. with the extremes of 5 and 20 em. A similar but not so marked variation was observed in the length of the fertile zone which averaged from 6.8 to 11. 8 mm. wi th extremes of 5 and 17 mm. The width of the fertile zone was more stable but comparatively large being from 3 to 3.9 mm. except for Nos. 34, 35 and 86 where it was
2 mm.
Five specimens, Nos. 13, 15, 16, S7 and 89 had divisions in the sorophores with, in No. 87, a second division instead of the normal single one. The number of sorophores varied from 6 to 10 pairs per frond with extremes of 5 and 12 pairs. The sorophores bore on an 30. average 9 to 12 sporangia.
Stomata were in single rows excepting in Nos. 15, 87 and 89 where there were small sections which were double.
The concentration of stomata varied from 12 to 20 per 3 mm. of frond with the extremes 11 and 22 per
3 mm.
Frond density ranged from 10 to 35 per 1 cm. of rhizome.
Frond scales in all but No. 87 were normal in size with obvious sllpport cells. No. 87 had scales slightly shorter than the rest and along with No. 89 had slightly shorter support cells.
The rhizome scales were of the normal shape and were similar in colour to the scales of the specimens from the more Southern Islands. Most of the scales vrere three celled wi th the occasional two and four celled ones.
SOU'l'H-SOU'J.'H ISLAND
This region takes in all specimens from the area south of ,Tacl{sons Bay. Only one of the seven specimens No. 14, was kept growing at the University.
The other specimens were Nos. 75, 76, 78, 79, 82 and 83.
'l'here was a vdde range in height which averaged from 5 to 17.7 em. with extremes of 4.5 and 22 em. 31.
The width of the fertile zone varied only slightly between 2 and 3 mm. but the same cannot be said of the length which averaged between 7.6 and 1292 mm. with extremes of 5 and 17 mm.
The number of sorophores was more constant with
6 to 9 pairs per frond each one of which bears between
7 and 10 sporangia in two rows. Only two specimens,
Nos. 75 and 78, showed signs of sorophore division.
stomatal arrangement was double in three specimens,
Nos. 78, 82 and 83. Density of stomata averaged
14 to 25 per 3 mm. of frond.
The sorophore width of 1 mm. and frond diameter of 0.5 mm. were again constant.
Frond densities were not obtained for all specimens but the range for those that were was 14 to 30 fronds per 1 cm. of rhizome.
Frond scales were all of normal size and, with the exception of No. 76 which had only moderately obvious supports, had very obvious support cells.
Scales on the rhizome were the same light brown as those seen in the other New Zealand material.
Cell number was mainly three with some two and four celled ones. 32.
SOUTH CENTRAL - sou'rn I8LAl\JD
There were eight specimens, Nos. 2, 3, 27, 28,
29, 30, 56 and 57, from this region which extends from Jacksons Bay north to Charleston. All specimens were kept growing for a period at the University.
Features that were constant were the frond diameter of 0.75 mm. J sorophore width of 1 mm. and the arrangement of sporangia in two rows.
The frond heights averaged from 5.4 to 19.1 CIn. with extremes of 4 and 25 cm. The fertile zone length showed a similar range being an average of'
5.7 to 10.1 mm. with extremes of 4 and 14 mm. The width of the fertile zone ranged from 1 to 4 mm.
Sorophore pairs per frond averaged 6 to 10 with each sorophore bearing 9 to 12 sporangia. Half the specimens, Nos. 3, 27, 28 and 57, had some sorophore division.
stomata arrangement was single in all but Nos. 2 and 3 where on some sections of' the frond stomata. were in double rows. The stomata density averaged from
10 to 22 per 3 mm. of frond with extremes of 9 and
25 per 3 mm.
'1'he densi ties of fronds on the rhizomes were from
11 to 15 per 1 em. of rhizome. 33.
Frond scales were all normal in size and had obvious support cells. ill1izome scales were all normal in shape and most were three celled. A colour variation was evident in Nos. 2, 3, 30 and 57.
These were darker than the rest, which were the same colour as those seen on other New Zealand specimens. rrhese dark scales however were not as darlc as the ones from New Caledonia.
HOHTH CEN'l'RAIJ - SOUrrH ISL.lDJD
Nine specimens, Nos. 1, 20, 21, 22, 23. 24, 25,
26 and 84 were observed from this area. Eight, No .. 84 being the exception, were kept growing at the University.
This area extends from Charleston in the south to
Karamea in the North.
Frond diameter, 0.5 mm., sporangial arrangement in two rows and sorophore width 1 mm., were all constant features.
Frond height averaged from 7.4 to Ut.2 em. except for No. 26 where it was 22 em. The width of' the fertile zones was an average of 2 to 3.1 mm. with extremes of 2 and 4 mm. The fertile zone length averaged 5 to 11 mm. with extremes oi' 5 and 17 mm. except in No. 26 where the average was 1.5.6 mm. wi th 3LJ .•
extremes of 12 aLi 18 mm.
Frond densities on the rhizomes were 10 to
18 per 1 em. of rhizome.
Sporangia pe~' scrop:nores averaged 9 to 13 with
extremes of 8 anci 17. The number of sorophore pairs
averaged 6 to 11 except for No. 26 which had 15.
Four specimens, Kos. 1, 20, 24 and 26, had some
division of the sorophores.
Nos. 20, 21, 22 and 84 were the only specimens
with their stomata i~ single rows. In all specimens
the stomata de!1si t~' averaged 11 to 21 per 3 mm. of
frond wi th extrel:1eS of 11 and 23 per 3 mm.
Frond scales in all were normal in shape and
had obviol_lS suppo!'t cells.
Rhi zome scales wel'e 9.11 normal in shape with
No. 26 longer than the !'est. All were predominantly
three celled. Only =\i"o. 24 showed the dark colour that was evident in the previous group, the rest being the usual lighter brown.
NOR':PH-SOUTi-I ISLAl~D
This area is from Karamea north and fourteen specimens Nos. 4, 5, 6, 7, 8, 9, 10, 11, 31, 32, 33,
58 a-e, 77 and 80, were observed. All but 58a and b, 77, and 80 were kept growing at the University.
No. 58 a-e needs some additional explanation.
These specimens were collected from within an area six feet by two feet. They show remarkable variations for such a short distribution range. Height averages from 7 to 12.5 cm. and the fertile zone length from
7 to 15 mm.
A wide range in sizes is evident in all material from this region (See Plate No. 19). Some possible causes will be discussed in Chapter VI.
The average frond heights were 5 to 30 cm. with extremes of 3 and 33 cm. Fertile zone length varies similarly, averaging from 6 to 23 mm. with extremes of
3 to 31 mm. 'l'he width of the fertile zone is more stable averaging 2 to 3.5 mm.
The number or sorophore pairs averaged 7 to 19 per frond, each sorophore bearing an average of 9 to
18 sporangia. Only Nos. 7, 31 and 58 c-e did not have any sorophore division. In No. 8 this division was tripinnate in nature.
All the specimens had sporangia arranged in two rows, sorophore width 1 mIll. and frond diameter 0.5 mm.
Frond density ranged from 12 to 21 per 1 cm. of rhizome. 36.
Five specimens, Nos. 4, 8, 9, 31 and 80, had stomata double in places. The density or stomata averaged rrom 11 to 19 per 3 mm. or rrond with extremes or 9 and 22 per 3 mm.
A remarkable unirormity was evident in frond scales which were all normal in size and with obvious support cells.
Rhizome scales were all normal in shape and size, being mostly three celled. The scale colour in all but No. S8d was normal, 1. e. lightish brown; the exception was dark.
SOUTH - NORTH ISLAl'JD
There were six specimens from this area, two living, IlJ"os. 12 and 36, and Nos. 68, 69, 70, and 74 llerbaril..Uil specimens.
The usual constant characters of sorophore width,
1 mm, sporangia in two ro~s and rrond diameter, 0.5 mm., were observed"
Frond lengths averaged rrom 5.3 to 10.1 cm. The length and width of the rertile zone varied only slightly averaging 5 to 8.5 mm. and 2 to 2.9 mm. respectively.
The number of sorophore pairs averaged 7 per rrond in all specimens with each sorophore bearing 7 to 10 sporangia. All but Nos. 12 and 74 has some sorophore 37.
division.
The density of stomata averaged 15 to 18 per 3 mm. of frond and three, Nos. 36, 68 and 70, showed signs of a double row.
Frond density was not easy to determine but appeared to be 30 per 1 cm. of rhizome excepting in
Nos, 12 and 36 where it was 15 per 1 cm.
Frond scale support cells were all obvious and, with the exception of No, 69, the scales were of normal size. In No. 69 the scales were slightly smaller and finer than in the rest.
Rhizome scales were normal in size, shape and colour.
NORTH - NORTH. ISLAND.
Only three herbarium sheets were observed from this region, Nos. 71, 72 and 73.
All specimens were large with heights averaging from 20 to 33 cm. in length, and the fertile zones had a correspondingly long appearance with average lengths from 17 to 20 mm. rrhe fertile zones were 3 to 4 mm. wide.
Sorophores all showed signs of division and there were from 16 to 21 pairs per frond. Each of the sorophores bore an average of 15 sporangia. 38.
Stomata averaged 13 per 3 mm. OL Lrond and only
No. 71 had them arranged completely in single rows.
Frond density was an average OL 10 per 1 cm. OL rhizome.
Constant Leatures were sporangia in two rows,
Lrond diameter of 0.75 mm. and sorophore width of
1 mm.
Frond scales were slightly longer than normal
wi th obvious support cells e
Rhizome scales were normal in shape and colour.
There was a predominance OL two celled scales in
Nos. 71 and 72 while in No. 73 they were three celled.
The New Zealand material shows similar variations in frond length, fertile zone length, etc. to the non New Zealand material (see table of averages in
Fig, No.8). There were no variations, OL significance, noted in the nature of the frond or rhizome scales such as were seen in the New Caledonia group. Fig. ~'To. 9 sho\vs several characters graphed and at no point is there a signiLicant break that could be used to delimit the species and the variety. As is seen in Fig. ~o. 7 all the characters become smaller with frond length except stomata number 'iv-hich increases. 39.
rhe frond length variation etc. would seem, more and more, to be related to environmental conditions. s is endorsed by looking at the variations seen material from Stockton and Okarito or the Plate No, 19. The ecological factors will be discuss in Chapter VI..
To sum up the external morphology; for all material is a definite uniformity, within the ecologic on of size, except for the New
Caledonia whic11 has the scale difference. CHARAC'l'ERS REFERRED TO IN FIG. No. 8
1. Frond length.
2. Fertile zone length.
3. Fertile zone width.
5. Sorophore number. 6. Sporangia per sorophore. 7. Fronds per 1 em. of rhizome. 8. Stomata per 3 mm. of frond.
9. Sorophore division. s = division n ;: no division.
10. Stomata arrangement.
si - single rows.
d "" double rows. Fig. No.8.
Character l!umber ere Q 1 2 3 5 6 7 8 ./ --._-----'"-
AUCKL.AJ'JD ~ill'm ? h 6L~ 5.11 2 L.. .. ~ 5.3 10 17 1'-1 .• 6 n 81 65 5.q 7.1 2.5 6.9 12.1 16.6 s si 66 3.2 5.8 2.4 6.2 6 18.3 25 n d 67 6 6.5 3 7 5 17.5 22 n si
I 13 .2 11. 6 3.4 8.l~ 11. 2 10.6 12.3 8 st .. 9 8 1:'c 3.3 6.6 12.5 13 12.6 s d 16 8 8.8 3 8 12 18.5 15 8 si 34 7.6 6 2 7 . ~') 10 20 20 11 si 35 8 .. 5 8 . ./;::; 2.2 12 16.6 18 n si ., 1 81 13 _t~ .. 1.., 3 .'3.5 17 n si
85 9.1 9. Lr -'", 18.7 n si 86 6 6.8 2 7 8 5 35 19 .. 3 n si 87 12.5 11. 8 3-.9 1_0.3 10.5 15.5 17.1 s d es 13 J.1. ~, 3.7 9.3 11 12.5 8 n si a 607 9 .. 8 2.9 7.9 5 20 s d
ISI,Jl.lfD 7.6 7.6 2.l 7.6 10 15 20 n 3i h q 75 15 2.9 ~. ill ..; 9 15 s d q ') ''-' 76 17.7 ../ L.. 7 .. 1 9.3 11. 3 .~ si r 78 lli.2 12.2 3 • 6 l v' 30 14 vC! si 7CJ 9.7 8.1 2 S 8.8 14 16.. 7 n si 82 5.1 6.2 2 6· 3 25 n d 83 7.5 7.9 2 7 7 19.8 r~ d CE?ITrL'1.L - SOU'!'H ISU1J'W n 2 ;, 5.7 2.3 6~7 10 15 21 u d 3 5.4 6.3 1.3 7.6 9 22 s d 27 8.9 7.1 2.1 7.6 12 14.7 s si 28 12.1 .1 2. L: 803 J.l 12 15 s s 29 14- 5 3 10 12 14·.5 17 n si 30 10 9 2 6 12.5 10 n si 56 15.1 7.8 2. }..j 5 9.1 11 11-!-0 7 n si q r 57 19.1 ~. It .J 2·7 9.2 11. 2 15.2 s si ]i'ig" No. 8 cant.
Species Character number number 1 2 3 5 6 7 8 9 10
:WR'fH CEI\;TP...P.L - I 1 13.4 .. 1 2.2 11 11 13 .. 6 IB s d 20 14,.2 10 3.1 9.6 13 12 14 s si 21 6 5 2 6 9 10 21 n si 22 6 8.6 2.4 7.6 16 18 16 n si 23 B.l 8 2.2 B 9 16 18 n d 24 6 7 2.2 8 9 13 18 s d 7.4 6.8 2 6.6 11 17.3 20 n d 2 22.2 15.6 2.4 14.6 11 15 s d 84 6 9 3 7.7 10.8 18 .5 n si - SOUTE ISLM-m II 16,,5 18.1 3.1 15.1 5 s d 5 16.8 18.7 3.3 14 7 16 13 s si h '-' 11. 3 11. 8 2.3 11. 9 18.3 16 G s·i 7 5 7 2. LI 6 9 17.3 11 n si 8 7.5 12.5 2.6 12.5 HI 12 19 s d Q ~ 19.1 17 2.6 15.7 18 12 14 s d 10 10.2 9.6 2 10 15.5 17 s si 11 23.3 23 3.3 • 6 1B 18.5 12 s si 31 11 6 2 8.6 11 15 12 n d 32 10.2 .1 2.8 11. 3 10 21 18 s oi 33 8 8 2.2 7.2 9 18 11 s 58 12.5 .3 2.5 8.7 14 16.5 14.2 s si 77 4 16.8 3.5 14.3 13 12 12.5 s: si 80 30.4 16.2 2.8 16.4- 13.5 12.7 s d
5 5 2 7 10 8 n si 36 10.1 8.5 2.9 7.6 13.6 16 19 s d 68 8.3 6.9 2.3 7 r) 8.2 .5 16 s d · "- 69 7" !.J- 7.3 2.4 7.6 7. LI .5 5 s si 70 7,,6 7.2 2.4 7.2 7.2 16.8 "CJ d 74 5.3 7.1 2 7 9 40 18.11 n si - NORTH ISLAND 71 26 16.9 3.6 15.5 IG.2 11. 6 s si 72 19.3 17.1 3 1 !.J- Ib.5 14-.8 s d 73 32.6 20.1 3.6 · 3 13 s (1 34 FIQ N"~ Na ..... ZoaLIl.r'ld mnlcrl.aL,
3Z -Frond lOflgtt'o In .::mt.- ,0 -Forlllll' zone LongHI in mm. ---Su;l1'natll por J I'T1m. U .. ,, 2' ,, 22 , 20 / .& ! ,) •• / I •• , I 12 , I 10
2 15 G 31 10 32 3 Cf 30 '] 22 e 27 35 U 23 33 1& 10 3:4 4 1% U 40.
Spores were c ected from to give a rrcrDresentat coverage of the ous forms ranging from the extreme reco zed as the variety to se identified as species. In all res
from 12 spec imens were planted, (Nos. 3 1 Lt, 7, 8, 11,
16, 21,23, ,28, 31 a~'ld 36). these spores were planted out in ~ i dishes on 30uthl
Fe , in March of 1964, t in the as e at the University in TId s had no for different c tions and the resul t vvas that the gametophytes ed from overheating the
er of 1964" rrhe second. attempt, in ember
, was made by growing them on Pakihi so from
Rockville. '1'hese l,vere kept in the glas e at Ilam where is some control over temperature and humi ty. '1'11.e SDores and grew much better
those tried on the Peat. The reasons this are most probably more stable tions and the fact that soil is more to the natural habitat than ~he peat. cimens were collected peaty soils but these were of a different t to 41.
the Southland peat, and contained mainly undecomposed plant pemains. These were moved in March 1965 to
the growth room in the Physiology Section at the
University in town. This gave standard conditions of light and temperature and the gametophytes grew very well. This move was made in an attempt to stimulate the production of sex organs but these did not eventuate. It was noticed that spores placed in the growth room all tIle time germinated more rapici.ly tllBll ones III tne glass house. The dishes were moved in li'ebruary 1966 to one oi' the rooms at Ilam until they were not needed any longer.
The soil was not sterilised prior to planting spores because of a report (Britton and Taylor 1901) of ¥nycorrll.i zal flUlgus in the gametophytes Q1' s. pusilla. 'rhe gametophytes of S. pl.lsilla are very similar to those of S. fistulosa and it was assumed, rightly, that S. fistulosa may have a similar fungus, which Britton and Taylor suggest comes in from the soil and forms a symbiotic relationship with the gametophyte.
Because the soil was not sterilised this led to problems which caused minor upsets, the most obvious being the presence of various contaminants in the soil on which the gametophytes were growing. These consisted of a.
Fig. N~10. Stagos in GamGto phyte Growth a.Spor@ Garmination b.Rhizoid Growth C. Fi rs t Bra nch Formin~ 42.
interwoven protonemal threads of mosses, some stray seeds of grasses and shrubs and pro thalli of other I fern species. rl'hese were rela ti vely easy to combat merely by weeding the dishes, but the control of algal contaminants was not so easy. 'rhese algae were of two types, a unicellular species and a filamentous species. Two methods of control were tried but neither was successful. The first was removal of as much of the soil bearing the algae as possible but at no time was this totally effective, there always being some left to respread the infection over the dishes. The second method was to apply a solution of potassiwn permanganatc. 'rhis was unfortunately effective on the gametophytes as well as the alga. It was finally found. that some measure of control could be gained by allowing the surface soil to dry out. Some algae remained in the moist areas around the gametophytes but this did not seem to hamper their growth.
VEGETA'l'IVE DEVEI,oPMEliTT
Develo~ing spores initially take u~ moisture and swell slightly, at the same time becoming slightly green. The spore coat then splits and the first filament grows out (Fig. No. lOa). This may develop up to four 6e113 before the rhizoid grows out (Fig. a.
Fi g. N~ 11. Stages in Gametophyte Growth a. Fi rs t 8 ranch CompletQ b.Sphi.Hical Cell tion
b. No. lOb), also from the spore. At this stage the spore is divided off from the rhizoid and the filament.
The cell next to the spore branches and cuts off a. cell which forms a filament that grows into the soil
(Fig. Nos. 10c and lla).
Growth of the filament is by an apical cell which elongates then cuts off a small cell at its tip. rEhis small cell then elongates and repea.ts the process. The small cell formed by these divisions is approximately one Cluarter the length of the mature cells.
Branching occurs in a similar way. A bulge which appears on the side of a branching cell grows out into a. filament and when it reaches a size similar to a new apical cell it is separated from the original cell to continue growth as a ney{ filament. These branches normally arise on the distal end of a cell.
SubseCluent development f'rom the four to five cell stage is slow, the filaments growing along the surface of the sailor down into it, and all the while branching.
Branching is not confined to any particular cells, although it is more common in the basal region of the aerial filaments; the underground filaments branch at all stages (Fig. No. 12).
rEhe aerial and underground filaments are basically similar but the aerial ones contain chlorophyll Rnd 44.
are shorter than the underground filaments which have
clear contents. Some gametophytes removed for
observation were returned to the dishes very little
care was taken in seeing them properly r anted.
When were next observed the rground filaments
that were above ground contained some chlorophyll and
new cells produceti were er than their
essorS4 The reverse had also occurred with
burried aerial filaments.
On short lateral projections from und.ergr
C 3 special spherical cells are formed. These are,
as the name sts, almost spherical and they
produce two or e three rllizoids.
Some of the cells were iId'ected th a
mycorrhizal similar to the c t reported by 12ritton 'l'aylor (1901). :\To .. "CA shoVls ari
w:infeoted spherical cell and Fig. No. an i:nfeoted
one. Whether the infection is c ed in the spores or
comes the soil was not investigated. Wone of the
t on the peat d sufficient size to
have cal cells, a that may have assisted in
p 1ally determining s. Eo Ettempt to isolate
i ify the fungus was talwn, Fig.NC? 12. Gamgtophyte Well Develop Showes all phases of vagGtat iva growth. Scale·2mm.= O·014mm. Uninfectad celt from Ni?16.
Infected ceU from N913. The infection did not appear until the gametophytes were well developed, like the one in
Fig. No. 12, or a little earlier. Fig. Eo. 13b shows a hypha brancnin&; outside t.he rhizoid and could possibly mean that infection is internal. rrhis could also be a hypha returnin6 to the outside after infection from the soil. This is pUI'e conjecture and further study would be need.ed to determine whicr.L is correct.
The time gametophytes took to reach various sizes was not clearly detersined. All the spores germinated. within the first month aft.er planting after which growth deuended on the concentration of spores in the area'Q rrhose that were not craTI"iped developed. to a large size, 3 to 4 cms. in leLgth with associated branches. Fig. No. 12, from specimen No. 16, shows one which tool{ only 14 weel{s to reach that sizea Others in this culture, which were undisturbed reached.. much greater proportions. GaJ:1.etophytes which were cramped cUd not reach the size shown in Pig, No. 12 even though they were growing for the same length of time, the majori t.;y being only about half a centtmetre in length. a.
d. e. c, g. NC?1L.. Antheridia Development. a.Swo Lle n ter minaL CQ II b.PGdestal cell formed c.Cap coll A ntherozoids fo rmod d.Cell cap ruptured - contents discharged e.Antherozoid 46.
i!l'I'l'ILERIDIP..L DEVELOPMENT
It would seem most likely that the formation of
ri a is induced by vt'arnl weather
surrounding soil begins to slightly dry out. se were the conditions when development occurred in
Laboratory althou~h it is likely to be rare in some of
lac ali ties where the s cimena are f'ound e. g.
ston. P~theriQia observed d froIT. gametophytes that were in clusters. apex of either
the main or, more commonly, a two c ed lateral branch swells (Fig. No. ) is ~ut off from main filament or branch. chlorophyllous and nuclear cor:.tents then tend to curr-ulate in the centre,
a destal ccll (Fig. No. 14b) c11 retains some
the or aGts, is cut off.
The next sions are the cutting off a cell and the formation a round the chlorophyllous and nuclear contents (Fig. No. 14c). The formation of this wall takes place without cell division. It appears as a partition between clear contents and the dense c 8 and leads up to cap. This leaves the appearance a swollen outer encircl cell with a cap sitting on a s Subsequently central contents develop into a or of densely protoplasmic
each of which represents an antherozoid. e I N«?1S. Antheridia. at various staggs of dQvelopment. as they are normally borne. 47.
are normally 12-16 in number. The release of the
antherozoids was not observed but by some means the
cap is sprung open and the Bntherozoids are able to
escape (Fig. No. 14d). Tl1e antherozoids have a central
nucleus and an outer layer which ap~gears Gelatinous
(Fig. Ho. 14e).
P~theridia were seen to be borne singly or in
pairs (Fig. No. 15). If they were paired it was noted
that the two were at different stages of development with
one always much younger than the other.
No archegonial development was observed.
The development of the gametophytes vegetatively,
and the antheridia are very similar to the development
of 3. pusilla gametophytes described_ b3r Bl~i tton and
'l'aylor in 1901. This would tend to sugzest 3. remarkable
uniformi ty in gaffietoph;)rtes wi thin this section of the
genus as the two observed species are geographically widely separated. There is certainly no suggestion of any difference between the garnetophytes of S. fistulosa
"species or "variety". 48 ..
CR.Apr.rl~R IV G
CHROMOSOME NUMBgH
In view of the increased emphasis being placed on chromosome numbers in classifications, counts were made of as many specimens as possible. This resulted in twelve specimens, Nos. 1, 4, 7, 8, 9, 10, 11, 21,
22, 25, 27 and 29, being observed.
The fertile heads were fixed in aceto-alcOhol for several days then smeared in and staine~ with aceto-carmine and mounted in Gurrs Neutral Mounting
Medi um. Apart from the mounting medirull the sequence was that described by Manton (1950). The sorophores
~ere nat easy to Get at the richt stage as there is almost a simultaneous development of spores and it was necessary for the sorophores to be not completely soft nor beginning to become hard indicating that some spores had been formed.
The spore mother cells are relatively large corrrpared wi th those of' other ferns but the chromosomes do not contract very greatly when compared with many other ferns genera.
This lack of contraction and the large nu.111ber of chromosomes Bakes it difficult to get an accurate count.
Jrhe counts obtained were n = 94 (Fig. No. 1);'4 for Fig. No. 16. From specimen No.7.
49.
Nos. 7, 21, 22, 25, 27 and 29 and n = c.150
(Fig. No. l&~ for Nos. 1, 4, 8, 9, 10 and 11.
Brownlie (1963) records counts of 94 and c.270.
It is possible that the larger count is from a tapetal cell. These are very numerous and at times hard to distinguish from the spore mother cells.
No. 11, with a count of c.150, is from the same locali ty, 'fakaka, as Brol.'mlie' s reco:!:'d for the count of c.270. This does not mean his count is wrong but would tend to suggest GO. Nos. 7, 8, 9 and 10 all came from within the same locality
(Plate No. 19 is from the same area). Nos. 7 and 8 in fact were growing within ten yards of one another but have different counts.
It would seem that generally the taller specimens have a count of n := c.150 while the shorter ones have a count of 94 (Fig, No. 18). 'l'here is however no marked division between the two for any of the characters studies in Chapter II. Some shorter specimens, No.8, have high counts and taller specimens,
No. 29, have the smaller counts. A similar picture
develoDs.... if other morDholovical....L. <---.' characters are comnared_ with the chromosome n'-Ll1bers. Klekoweski and Baker
(1966) suggest that most fern taxa are capable of' Fig. No. 17. n c. 0 from specimen No.1.
50.
producing complete homoz;ygotes by self-fertilization and tho.t they do this frequently in nature. The production of these homozygotes he suggests is reI a ted to the high percentage (96~~s of 1166 species of which he had records had a count greater than 27) of polyploids in homosporous ferns compared with heterosporous ferns (90% of 50 species less than 27) where there are few polyploids' and where complete homozygote formation is highly unlikely. They go on to state that polyploidy increases gene redundancy and if dominant alleles are present then the recessive ones, of the same gene, are kept from immediate exposure to selection. These recessives may take up new functio:1s without upsetting the essential processes of the organism.
This combination of homozygosity and polyploidy could well explain the remarkable uniformity of morphological characters in S. fistulosa where there are at least two counts.
Lovis (1958) and Wagner (1963) record counts for some of the other species observed. These counts are:- s. bifida 77 -----s. rObl.lS ta 96 103 Fig. N?1B,
24 ~ ~ a1 at ~ a1 a1 Chromosome Num ber 22 20 \ 18 , 6
14 vi uE 1 2 c .- 10 ...... c Ol 8 '0 I c 6 u 4
" 9 4 2 9 1 10 2 2 2 7 25 21 7 Specimen Number 51.
Chiarugi (1960) suggested that the base nwnber for this group could be 11. He'made this assumption from the record for S~~~if'ida given l)y Lovis (1958).
Pabbri (1963 and 1965) doubted this and stated in the 1963 paper that this was not because or the i~possibility of interpreting the data but because he prefers to abstain from suppositionso In view of the
COl.l:'lts for S. f'istulosa it seems to case mOI'e doubt on the possibili ty of' the base number being 11. 52.
CH.APTER V.
SPORE MORPHOI,QGY
Spores were observed from all material but
Nos. 2, 10, 12, 35, 56-58 and 90-100. These were
stained and mounted in the same way as detailed by
Harris (1955).
Included in the group Nos. 90-100 are
S. bifida, S. robusta and g. pusilla. The spores from
these three are described by Selling (19L~4) and they
are distinct from S. fistulosa. The difference is very
slight in S. Y'obusta l)ut it is still sufficient to
make it distinguishable from the smooth nature of the
exos~ore of S. fistulosa.
S. malaccana, No. 48, was distinct from the rest of the material by having a completely smooth exospore.
All the other specimens have some markings or sculpturing but S. malaccana has none.
The external a:ppearance of all the s. fistulo~
~aterial was the same regardless of where it was collected. The exospore gave the alJpearance of being fine grained. This would tend to suggest that it was covered with fine projections but this was not apparent 53.
on close observation when the exospore appeared to be smooth.
The size of the spores diA not vary greatly.
Ten spores were measured from each specimen and they all measured between 0.109 ± 0.009 mm. by
0.073 ± 0.009 mm. These measurements were for the overall length and width of' the spores.
On these grounds I Vlould support Selling (1944) and say that there is, by s90re morphology, only need for the one snecies and no variety. 54.
CHAPTEH VI.
DISTRIBUTION AND ECOLOGICAL FACrrORS.
DISTRIBU'l'ION.
From information in various publications
(Godley 1963, l,ooser 194L~), from herbarium sheets and from my own collections I have made up the distribution maps (Fig. Nos. 19, 20, 21).
Going from West to East on a global scale the distribution is, West Australie.., Borneo, Hew Guinea,
Tasmania, ~ew Caledonia, New Zealand, Chile and the
Palkland Islands. The species has also been reported from Madagascar but this, as is stated in Chapter I, is incorrest.
Within most of the areas the distribution anneal'S.J.. _•. to be limited. In Borneo it is reported from one place only - Iva. IUnabalu. The specimens from West Australia were both from south of Perth. Tasmania would seem to have a general distribution in sui table habitats.
In NeVi Guinea and New Caledonia tl'le collections are from the eastern halves. This may be because these regions have been visited most by collecting expeditions.
The Chilean specimens were collected from almost coastal sites, althous-h some are from a reasonable height.
Godley (1963) reports that the Chilean species are Fig. N~ 19. Known distribution of Schizai.?3 fistuLosa seas.tat.
South America.
~ New Zealand N°!.17,37. Cl # lsee Figs.20&21.1 N°s. 61,62,63 ..---_ Tasmania /'-'" Southern Chile
N°s. 3 8,3 9, 40,54,5 5. 1>_ -Auckland Islan d N°s. 64,65. N°s. 59.60 ..--- __~ .. Campbell Isla nd N°s. 66,6 7. Falkland 1s1and5~ 55.
sparsley scattered between 39°30'8 to 49°10'8. The species is also in suitable habitats on the Falkland
I~,lands, this being the t;'lpe 10ca11ty for Schizaea aUG tralis Gaud.
Within New Zealand the distribution is localised.
Carr.pbell, Auckland and 8tewart Islands all have a general distribution in suitable areas. Chilton (1909) reports that specimens are more commorL on Auckland
Island, in open peaty places, than on the New Zealand mainland. The i'Iorth Islancl_ dis tribution is general but localised to lJountain bOES and similar areas south of Hamilton. North of this they are found in poor soils, more especially in Northland.
South Island distribution C311 be £iven 8S Golden
Bay and West Coast from Seddonville down through
Fiordland to Bluff. 'l'here are three reports from outside this area and these are Arthurs Pass,
Waimakiriri Basin and the Croixelles in the Marlborough
80UIlds.
ECOLCGICAL CONDI'l'J:mm
The specimens observec. came from a variety of habi tats from the point of clime.te exposure. One common feature to all vms the poor, slightly acidic, Fig. NC?20 South Land &. Stawart Island distribut ion & zones.
Takaka N° " OnQ kaka N° 77, Collingwood N?'5 Roc k v ill e N° s. ~4.;Z;8.#~9~.1;oi.~31.~32!7.:~,5~8~.-=_=_=-= _= _=_= -;::::::;;--=~ Bainham N°'s, 6,33.- Seddonvtle NC? 20. --______-'- North Stockton N? 21. --~--'------...... l. 5.1. Denniston N?s.22,23,24.------;:::C"
westPort.~~N~o:S~·1.32i~i2ro~n8~4=·====~::~~lCharla stan N°s. Nt;' 28. Central SJ Woodpecker Kumara N? South-Central Arthurs s N? 3. S.1. Okarito N'?5 Bruce
Caswell und N~S3, Dusky nd NOS.75.76 South 5.1. Chalky Inlot Otautau N? 82,-~-~q__ _ Bluff N?14.----~-.-..:l.",A-_ Masons Bay NO'S.34135.~ Pryses Pea~ - Stewart Island N°5.16.81,85,86,87,BS,89. BIg South Cape Nc:;' 15. .... ,- Codfish Island N? 13. 56. soil be it b or reasonably well drained pakihi. is menti Chapter II the feeling is that climate and exposure as well as titude influence
the size, expressed 1 in i , of the c imens. No specific evidence has been c lected this but, prompted by the observation ma.de by Miss E. F.
Kibblewhite (Dobbie 1952) that the variety,
her fernery, developed into perfectly normal
careful observations were made
of tions specimens were growing in.
. f' A comb lon o~ and degree of exposure would
seem to 'De
Altitudinal e is evident in the Kew
Guinea material (Fig. No.5) for one cimen.
This s~ecimen, No 47, was from an en area in sub-
ine scrub and, because of a greater e of exposure, could accOl.1...Tlt f"'Jr its being smaller"
an specimens show almost a reverse with regard to but exposur'e would seem to influence this.
No. 51, collected at the greater altitude, was on bush e more sheltered than those from the other areas. ( ie rs. cOffim.)
The Chilean spec seem to be cd latitude and altitude. • 59 and are from about
49°8 J the former at sea level and latter Fig. N? 21 North Island dist ri but ion 8. zones.
North N.1. Kaitaia N°s.71,72.
Ka ihu.------~
KourawhGro N? 73.-----'<-'1f- ~-Mt.Mo8hau.
Ta uhGi. - ______+-_
Hami it on. ------~ N~70. Oh in em ut u.------_-r::::-___~ Waiotapu. --~~--_____~ ____
Ah ukawakawa Swamp N'?69. ----1-_ ----;C---k: Mt.Ruapehu N~12. N°s 6u8~,7~4~.~------~~------".--- Raporoa Bog. Waimarino. -
Mt.Quain N? 36.------7 So uth N.!. Nos. 61, 62 and 63 are from about 42°S, Nos. 61 and 63
being alpine and No. 62 from sea level. This Shows,
once again, the influences, on size, b;y these factors.
Within New Zealand altitude and exposure seem to
have ~uite an influence. As would be expected the
specimens from Northland are large and those from the
Auckland and Campbell Islands are small but between
these points there is considerable variation.
By considering the influences of altitude and
exposure on a latitudinal range the point can be made
cle&rer. Specimens fron: Stockton, Dell.. niston, Charleston,
Margaret's Tarn, Okarito and Bruce Bay show this
extremely well. The specimens from Stockton, Charleston
and Okarito were collected from slishtly raised. areas of
pakihi while the rest came from very wet bogs.
Stockton and Denniston are both c.2000' above sea
level and are subjected to frequent strong cold 'winds
and misty conditions. The specimens collected from here were in no way sheltered fram these conditions. The
Charleston specimens came from a lower altitude and,
although they were exposed to all weather, the climate
is mlJ.. ch milder than at Stocldon or Denniston. l'£argaret's
Tarn is the highest of the sites chosen and is sheltered 58. from winds but is subjected to snmv as la te as
December as well as being covered by snow all winter and into the spring. Okarito and Bruce Bay specimens were from close to sea level and well sheltered.
Fig. No. 22 shows the relative sizes of specimens from these areasu Equating this with altitude and exposure makes it rather evident that these factors play an important role in specimen size.
The specimens observed from Rockville showed a variation in size (Plate No. 19) that would seem to be influenced by exposure. The taller specimens were sheltered. by high manuka while the shorter ones were all exposed to the frequent colc!. south anU south-west winds that blow in the area.
Consideration of these factors, in relation to the rest of the material, enhances the concept of size being determined by altitude and exposure. Plate No. 19. Range of sizes. These were collected over a distance of 400 yards in Rockville.
THE LIBRARY UNIVERStTY OF CANTERB.URY CHRISTCHURCH, N.Z. I ) '.
I , t .J 1 Spcec i n ight in ems.
~ -'" -'" -.II. -'" N 0 N .t"- O") (jt) a N s:- en Q) a
Stockt on 2000' 'V -0 DGnni stan 1000' N ~ N N BurnoUs CG 2000' N -- __I W nni s ton s:- an~ts 30--' , CharlGs ton 200 x N Charloston 200' x N 100· """-11 Okari to )( U1 en BrucQ 50'" x U1 ...... :! 59.
CHAFfER VII
CONCLUSIONS.
To come to a satisfactory conclusion in this
thes is f~11 the findings mUG t 1) e slJmmed up and weightect according to their influence on classific-
a tiona Four lines of study were followed some
shmving uniformi ty and others variation.
The spores from all the S. fistulosa specimens
showed a urliformi ty in their size, shape and
sculpturing or markings on the exospore. This is in
accord with the findings of Selling in 1944.
'rIle gametophytes, although they only produced
antheri,iia, once again give complete uni:f0 r'llt ty.
Vegetative growth was consistent throughout t11e twelve cultures planted and antheridial development, on the three where it occurred, was also uniform.
The vegetative and anthridial development are both very similar to those in S. pusilla, as described by
Britton and Taylor (1901), suggesting that the gaILeto:.ohytes are uniform throughout this section of the genus.
Extcrl1al :norphologjT did produce variations but onl;;,' two of these could be thought to have any
"':'HE UB~\.:-\~~Y UNlVEF,,:Try Of CANTERBUK.Y 60.
significance. These two are the frond and rhizome
scales ivhere the New Caledon.ia-Borneo material shoyvs
a distinct difference from the rest of the specimens
examined. All the other morphological characters
varied; some, such as length of the fertile zone or
nUlnber o:~ stomata, seem relateci to fronCi length
(Fig. Nos. 7 and 9); others had no set pattern in
tl1eir variation e.g. sorophore division and double
rows of stomata.
The cytological inves tiga tions give two :!Oli..-"'lts but
these cannot be related to any morphological break.
It can be said that the smaller specimens have a
predominant count of n =9!~ and the J.erger ones a cOlmt
of n=c.150 but there is no clear break in size that
could warrant recognition of' two varieties.
This then leaves three characters, frond and
rhizome scales and chromosome number, to be considered
in the classification. The spores, g8.metophytes and
remaining morphological characters are either completely
uniform or vary too greatly to be of significance.
Although these latter characters vary they in no way
detract from the overall basic pattern o~ the specimens.
To give the Nev{ Caledonia-Borneo group varietal
status, for it is certainly not a separate species, on 61.
the strength of two knovm characters is not possible.
These vary slightly 'Nithln themselves and it would re~uire a much more detailed study of a greater amount of material from these areas before any definite decision could be reached.
To split the New Zealand specimens into a species and a variety on the strength of chromosome counts is also not practicable. Al though cytology shows, two races this is not endorsed, as pointed out above, by any other feature. In view of the findings of
Klekowski and Baker (1966) and a consideration of the ecological conditions in which specimens are found, it is feasible to assume that polyploidy in tll.is case does not indicate varieties within the species.
From all this I would 3~ree with Allan (1961)
that there is only the one species, S. fistulosa t and no S. fistulos8 var australis. I would not confine this to New Zealand but would state it as covering the complete geographic distribution of S.-.fistulosa.
s. fistulosa Labil1. would then have the s;ynonyms given in Chapter I and the description that is in
Allan (1961) modified slightly to give a greater range in the measurements ~uoted. A - Arnold Arboretum.
CAl'!"B - Herbarium Aus traliense, Canberra.
CAN'fY - Canterbury Museum, Christchurch.
CHR - Botany Divis ion Herbarhlffi, D. S. I. R. , Chris tclmrch.
CIJl.fTU - University of Canterbury Herbarium, Christchurch.
- l'.~r. G. ~,rownlie' s IIerbaril,J]n, ChristC.flUrch.
HO - University of Tasm2~ia Botany Department, f:lerbari u;r., 'rasmania.
ILH - Ian Lash's Heroarium, Christchurch.
- lTniverE,i t;y of California Herbarium, T~ 0 s i~"l,S:1 e s •
- National ':lerbari um oi~ I\Tew South Wales, Sydney.
L - Ri jksherbari urn, LejJten, Holland.
WA - Universit;y oi' West Australia HeroariulT'., Perth.
SPECIMENS
]\To. 1. 1LH No,. B065, Ian Lash, now GB, Yifestport, i'T. Z.
open pakihi on slight rise.
No.2. 11,:3. l'ro. B.066, Ian Lash, now em, Charleston, N. Z. ,
open nakihi on terrace edge. lUI Ho. B067, Ian IJash, now GB, Margaret's Tarn,
Arthurs Pass, N.Z., outer edge o~ cushion bog,
C. 3000' .
l'r 0 • [.L. IL~ No. B068, Ian Lash, now 0B, Jacobs Ladder,
Rockville, N.Z., pakihi sloping to west C.150'.
ILH No. B069, Ian IJCtsh, novi GB, Collingwood,
N.Z., open pakihi, C.IOO'.
Ro. 6. II,ll No. B070, Ian I,ash,~ow GB, Bain...rlam, !'T.Z.
paldhi bank s~el tered by manuka, C. 300'.
No.7. ILH No. B071, Ian I~ash, now GB, Ap"};)Qs Flat,
Rockville, IT.Z., open yakihi exposed to sout~,
C.200'.
IL~I ~\TO. r3072 , 12.~ Lash now GB, 8.S ~or No.7.
IJ,E 1'10. 3073, IO.n 1..3.811, now :'+B, ADPOS Flat,
~ockville, ~\T. Z., 8;~el tered pakihi J C. 150'.
Ho. 10. n,H No. ~307Ll., Ian I.a.sh, now GB, Roclcville, 1\T.2.,
open pakihi, C.150'.
No. 11. ILH, No. B075, Ian Lash, now GB, Takaka, N.Z"
sheltered pakihi, C,lOO'.
No. 12. ILIi No. B076, A. P. Druce, now GB, lVit. Ruaphehu,
l\i.Z., C.4500'.
No. 13. IU·I 1'[0. B077, P. ,Johns, noV! GB, Co(lf1sh Isl,:md,
Ste'uart Isln.nd,
lUi i\o. 13078, Pr'cL j~a;;,rlt8, novr DB, B111f~ 1":. Z.
peat-sphagnum bog. No. 15. II,H No. B079, P. Johns, now GB, Stewart Island N. Z.
No. 16. IIJE B(JSO, Mrs. Leaslc, novi em, Stewart Island N. Z.
No. 17. WA, D. Churchill No. Li58, now GB, Walpole,
West Australia, amoncst sed~eE in surface of
deep peat.
No. IS. L, 'N. Vink No. 16272, now GD, West Highlands,
New Guinea, alpine grasslano., 3590m.
-:'Yo. 19. L, \'V. Vink ~;-o. 16022, now OB, V/est Highlands,
New Guinea, alpine shrubbery, on peaty
soil, 3270m.
No. 20. IU:: No. :;:081, Ian Lash, no\~' OB, Seddonville, N. Z.
in sphagnur.I on vret pakihi, C. 200 ' .
1":0. 21. ILH No. B082, Ian Lash, nD~ 3D, Stockton, N.Z.
exposed pakihi, 0.2000'.
~':~o. 22. B083, Ian Las!'",", now f'''''.-AD, Dermiston,
eXIJOsed paJeihi on hill, C. lOCO' •
No. 23. n,H :No. B084, Ian I~ash, now OE, Burnetts Face,
N.Z. very wet bog in pakihi, C.2000'.
No. 2Li. II,n, Ho. B085, IaL. La3h, new G3, Dennistoll,
N.Z., bog in pakihi exposed and wet C.2000'.
ILH No. POS6, Iall IJas!l, now 013, Westport, ::.Z.
open pakihi, C.500'.
1100 26. ILH No. ;3087, Ian Lash, now C-·B as for r{o. 1.
Ho. 27. IUI l:To u B088, 1 ali IJ8.sIl , now GB, as f>or [Oro. 2. IL~l No. B089, Ian IJash, now GB, Charleston, R.Ze,
in wet moss on sandstone road cutting.
Yo. 29. ILH No. B090, IaCl IJa3h, now GB 1 Wood.pecker J3ay
N.Z., exposed well drained pakihi, C.BOO'.
1'To. 30. IIJH No. E091, Ian Lash, now GB, Kumara, N.Z.
on vertical gravel face -dry.
No. 31. ILH No. B092, Ian Lash, now GB, Lightbands Gulley
Rockville, N.Z., south east slope of pakihi,
C. 200' •
No. 32. ILH, !\Jo. B093, Ian Lash, nO'N GIl, J~ightbalJ.ds
Gulley, Rockville, ~';.Z., pakihi C.200'. l\~ o. IVl No, 13094, Ian I~ash, nOVI GB, Bainham, IL Z. ,
sheltered pakihi, C.300'.
II,E Xo. 13095, ]',H'. l.':acarthur, now em, :\~asons Bay,
Stewert Isl8.nr1, 1T. Z.
ILH, No. 13096, Mr. stacarthur, now GB, Masons Bay,
Stewart Island, N.Z. lJo. 36. ILH, No. :8097, A.P. Druce, now GB, Rirnutaka
Ra:ilge, H. Z.
rs~, No. NSW-P430, E. Pritzel No. 251,
West Australia.
Ho. 38. HS'iV, ITo. HS'N-P8266, T. and J. lNhalte, :1'fo. 221-1-9
Tasmanla, or,. but ton grass ~plains. -..;q l~o • --' -' . 1'18'.'\/, No. N8Vl-F8643, G. and C. Davis" Tasmania. J'! o. 40. WSW, No. P1812, W.R. Archer, Tasmania.
Ho. 41. NS~, R.D. Hoogland and R. Pullen, No. 5694
New Guinea, open rocky slope, 12,500'.
No. 42. C..:'l\fB, now NSW, R. Schodde, ~\:o. 1733, Papua,
alpine grasslallds - boggy ground, 10,300'.
NSW, R. Bchodde No. 20~3, Papua, bog grasslands
8800' .
No. Lt4. CJ(;'m, No. L~0561, duplicate of' No. L/.1.
No. 45. c..:'ti.m, No. 107131, R. Schodde No. 1867,
Papuc.. alpine grassland, 10,000'.
L, nm'{ C.A1:lB. No. 6699L\ jI }L. Jacobs
No. 5732, t~t. Kinabalu, 30rneo. stunted
gnarled mossy forest, 2500-2900m.
L, no~,-{ CiLJB. No. 96199, C. KalkmanS)
No. 4531, Eastern New Guinea, open places in
sub alpine scrub vegetation, C.3380m.
No. 48. T..LJ, Now CAlm Ho. 128196, P. van Royen
and He. Slewner =\;0. 8024, Eastern Nevv Guinea,
in forest, 1800m.
CAlm No. 108113, duplicate of No. LJ.3.
P.S. Green No. 1206 now GB, New Caledonia,
in bog by a stream.
.No. 51. GB :1'10. 450 (~'l".C.149) G. Brownlie, NeVI Caledonia,
forest floor in mist zone just inside forest,
3700' . reP 1Yo. 52. u .L' No. 545 (E.C.242) G. Brownlie, New Caledonia
open ridge in scrub, 2500' •
No. 53. GB No. 4w.4 (N.C.143) G. Brownlie, New Caledonia
among scrub on steep hillside, 2500'.
No. 54. HO, J. Somerville J 'l'asmania.
No. 55. RO, J. Som~rville, Tasmania.
No. 56. ILE, No. B098, Ian Lash, now GB, Okarito, N.Z.,
raised section of large pakihi, C.150'.
No. 57. ILH I~o. ~i099J Ian Lash, now GB, Bruce Bay, N. Z.,
in sphagnUlll in large bog C. 50'.
No. 58. a-e. lUI :No. BIOO, Ian Lash, now GB, Rockville, N.Z.,
on pa~ihi slope, C.IOO'.
Ho. 59. CRR ~o. 153891, E.J. Godley, No. 579, Chile,
in bog at sea level.
No. 60. CER, No. 153892, E.J. Godley, No. 754, Chile
C. 2000'. l'~o. 61. CHR No. 153889, E.,]. Godley No. 1..j.25, Chile,
amongst tussocks.
No. 62. CllR No. 153888, E.J. Godley No. 81, Chile,
swamp at sea level.
No. 63. CRR No. 153890, E.,I. Godley, .No. Lr65 , Chl1e,
in bog.
:J o. 6Ij.• CER No. 88842, ~~.fr. Hoar No. 715, Auckland
Is 1 an d , l'J. Z. Ifo. 6511 ORR l')o. 88854, N. 'I'. Maar No. 7, Auckland Island, z.
No. 66. CHR No. 117939, E. ,T. Go ell Island, N.Z. cushion bog. No. 67. CHR No. 118021, E.J. ey, ell Island,
]\I. Z., among tussocks. No. 68. CUU No. 33247, A.J. Healy, near National Park, N.Z.
:-T No. 69. CdR ':"\10 II) 86763, A.P. Druce, • Egmont area Z.
No. 70. Cli'R No. 79485, A.P. Druce, lii eau, N.Z
No. 71. CI1H No. 69863, lTV ~ E' ~ Harris, a N. Z.
,:,'1 0 ~ 72. CHR No. 2276, Carse Callee t a, l'i. Z. i'To ~ 73. ClIP No. u4579, L. B. L;oore,
No. 7L;. C;}IR. IIa t) 151107, G. G. 0 e t ro ional
Parl{ , :'!. Z.
No. 75. C~R No. 55311, H.E. Allan, t Jacket Arm,
Dusky 8ourtd, N. Z.
CI{R 1:10. 55390, R.
Dusky Soured, N. Z.
:':0. 77. no. 9573u, R. ~ason N.T. Maar, No. 4777
• Z.
~\O G 78. GIrE l\r a. 117706, I-I. All
0 ~1 o. 79. mH? No. 117707, o. u , N. Z.
:':0. 80. CHR No. ll.1.05Ot,. , D. R. C:iive:D, es, 1\J'~ Z. No. 81. CHR No. 78024, J.F. Findley, Stewart Island, N.Z.
No. 82. CRR No. 82674, I\f. T. l\f:oar No. 2461 Otautau, I-T. Z.
:no. 83. Cj~H. No. 67L88 , A. Poole, Caswell Sound., N. Z.
}1 o. 84. CAITTU No. 202~" Paparoa Range, N.Z.
N'o. 85. ci~r'.ru No. 5689, Poy Johns, Stewart Island, 1\'.Z.
No. 86. Ci0JTTJ 'No. 7260, Port Pegasus, H.Z.
No. 87. C1JiTU No. 8283, Stewart Island, N. Z.
No. 88. C.A;-;TU No. 8282, Stewart Island, N. Z.
No. 89.
No. 90. Noumea, J.P. Blanc!lon No. 350, New Caledonia. No. 9L 902706.
ITo e 92. IIo It 122695.
~,!o • 93. L~ .LJ o. 735179.
ITo. 9L~. 1,.ll., i-To" 5331t23.
Ho. 95. IJA }~~ 0 e 8053L,I.,.
1'To. 96. I,A No. /).2972.
No. 97. LA No. 122660.
Ho. 98. LA No. 122661.
:rTo 41 99.
Ho. 100. ILH No. B043 , Ian Lash, Charleston, N.Z. BIBL T OG-RP.PEY.
1\T r; ALI..JI..l1J J H. H. (1961) 1 Flora of' J. .. 0 L.J. , Vol. 1, Govt. Printer,
No Z.
ARIiOLD, C.A. (1947), 1m IntI'oduction to Paleobotany;
McGraw - Hill, New York.
BR I'l"r ON , E. G. and rrAYLOR, A. (1901), Lif'e His tory of'
Schizaea DUBilla; Bull. of the Torrey Bot.
Club, Vol. 23, No. 1.
(1963), Chromosome ·NUlnl)er in some Pacific
Ferns; Facii'. Sci., Vol. IIi, pp. ~-93-I+97.
CH:~~ES:F~l\'lr'\:"! , '1'. F. ( 1925 ), l!:anual of N. Z. Flo r a;
(~~-vt. Printer,
CHIAPJ]GI, A. (1960), rravole Cromosomiche Delle
Pteridophyt3.; Caryologia, Vol. 13, pp.27-150.
CHIJ..1TON , c. (1909) Editor, The Sub-antarctic Islands of
N. Z., Vol. II.
CIffiIS'l'El'TSEH, C. (1906)) Index Filicum; Hagerup.
CHRIS'rE:;\TSEli, C. and EOLrrrrUlJ, R.E. (1934), The Ferns of
Mt. Kinabal11; 'I'he GardeLls' Bull.
Straits Settlement, Vol. VII, p.210.
DOBBIE, H.B. (1952), N.Z. Ferns; 5th Ed., W. and T. Ltd. F.l~BBHI J F. (1963) Primo Supplemento Alle Tavole
Cromosomich Delle Pteridophyta Di .lUberto
Chiarugi; Caryologia, Vol. 16, pp.237-335. (1965), Secondo SUI)plemento
Caryologia, Vol. 18~ pp. 675 - 731.
;- GAUDICll.!\H.D - B&~UPRE, C. (1825), Rapport sur la Flore des
iles Malouines, par; Ann. Des. Sci. Nat.,
Tome 5, IJ. 98.
GODIJIT, E.J. (1963), Contributi.ons to the plant
Geography of Southern Chile; Revista
Universits.ris. (UniversiCiad Catolica de
~~IARJUS, 'N. F\ (1955), A ma:lual of' spores of N. Z.
Pteridophyta; Bull. D. S. 1. R., N. Z. Vol. 116,
pp. 1-186. lIOI~rl'Tm.i, R.E. (1959), Flora lvTalesiana; Sere II, Vol. 1.
HOOKE..R, J.D. (1867), Hnbk. of theN.Z. Flora; ILondon.
HOOKEH, J.D. and BJ't.1(ER,J.B... (1874), Synopsis Filicum;
2nd Ed., Robt. Hardwicke, London.
KLEKOWSIG, E. J. jnr. and BA..XIm, H. G. (1966), Evolutionary
s ignificaIlC e of' Polyploidy in Pteridophyta;
Science, July 15, 1966, Vol. 153, No. 3733,
pp. 305-307. LOOSER, G. (1937) J La Schizaea Chilena (Filices);
Revista Universitaria, Universidad Catolica,
Ano XXII No.1, pp. 115-121.j..
(191.1.4) The Perns of Soutll.ern Chile;
Am. Fern. ~Tol., Vol. 38, No.3, pp. 71-87 e
LOVIS, tTeD. (1958) A Chromosome count in Schi~aea;
Nature, Vol. 18, April 12, p. 1058.
(1950) Cytology and Evolution in the
Pteridophyta; Camb. Univ. Press.
PRi\NTL, K. (1881), Untersuchungen zu~ Morphologie der
Gefass IZryptogamen. 11. Heft. Die
Schizaeaceen. LipLjig. (Not seen).
RI Carl'EIt , A. (1916) J Eine neue 3chizaea G1J.G Borneo;
~:lededeelingen van's Hi jl\:s Herb&ri um,
Leiden. :':'To. 28.
SELLING, O.H. (194Lt-) J Studies in the recent and .b'ossil
Species of' Sch:l~~~~, with particular
reference to their spore characters;
Medd. fran. Goteb. Bot. Tradg. XVI.
(1947), Further Studies in Schizaea;
Svensk Botanisk Tidskrift, Bd. 41, H.4.
SIll~M, rr. R. (1915), J?erns of'S. Africa. THOMsmr, G.lvI. (1882), The ferns and Fern Allies ofN.Z.,
Dunedin.
WAGNER, W.R. jnre (1963), A Biosyshomatic survey of V.3.
Perns - prel im. Abstract; Am. Perno Jol.,
Vol. 53, No. 1. ACKl'JQ,ifIJEDGEMENTS
It is v~itl1 gratitude and thanks that the author acknowledges the guidance and constructive c ticism
gi ven by IvJ:ro G. Brownlie throughout s s I would also like to
Mr F. Johns, Mr. A.F. Druce, Mrs. Leask,
~rs. Willa, Mr. Macarthur Frofessor Bayljs for live material.
Farents for supplying soil from Hockv e at various stages.
University of Ta a not. t., HyksherbariuTl1,
Holland, National HerbarillTIl e,f E. S. W. , [ulstraliense, erra, Botany Division D.S.l.R. ,
erbury eU.lTI, and Uni vel'S i t.:r of GalLeo , s Angles Herbaria for forwarding al for study.
Dr. E. cT. Godley for use of selected papers.
P. T. R. acLean aro for translation of papers,
and !ny for proof reading and encouragement.