l9s9l LOTHIAN: FurfherNotes Concerning The Cenfral AusfralianCabbage Palm - LivistonaMariae

T. R. N. Lorur.Llv, Director, Botanic GartXen, Adelaide, South Australia.

In this journal (Principes 2:92-94, lings of this palm were seen. These oc- I95B) some general comments were curred in various places: in densecolo- given earlier concerning the habits and nies under palms, as rows of seedlings habitat of Liuistona Maria,e. Since these which have grown from seed washed were written" a visit has been made to into rock crevices; or as individual seed- Palm Valley as part of an overall bo- lings in isolatedsmall pocketsof soil in tanical reconnaissanceof the northwest the rock mass. of South Australia and southern central The age to which palms grow before portions of Central Australia. Whilst dying is not known. A careful investiga- in Palm Valley, the opportunity was tion was made in Palm Valley, but very taken to make further observations and few fallen trunks were noted. Most cer- to collect a small quantity of seed.This tainly none had fallen in recent years. seedwas obtained with a view to carry- It is likely, however, that trees live to a ing out a few germination tests; not considerable age, becausepalms of 50 only to discover the of seedwhich or 60 feet high had only commenced to was viable, but to enquire into the stages develop the root mound which most of germination. palms produce as they reach maturity. Gencral Obseroations In half a doz,encases only were these The natural soil of Palm Valley is, as root mounds of any consequence(Fig. one would expect, river-washed sand 28) . In these instances they were 4-6 and gravel. About 6-9 inches below the feet through at the base and about 5 feet surface, heavier material is found, and high. Other than this, the trunks ap- stones often 4-5 inches in diameter oc- peared to develop straight from the soil cur. The surface of the soil is covered level. with decaying organic matter. In the The old fronds initially remain on the palm thickets (Fig. 30) this debris has trunks, bend downwards, and become built up to a considerablethiclcness. It appressed around the trunk like an is often several feet above the general apron, where they die. As in Washing- level of the river bed, and unlessthe riv- tonia, these old fronds stay in position er is in very high flood, very little of until blown off by strqng winds, but in this matter is lost by water. the centre of the thickets where wind The bed of the Finke River, which velocity is greatly reduced, the fronds forms the gorge and habitat whereLiais- remain adhering to the trunk. However, tona grows, is approximately two miles as the palm grows and the crown and long, an'd up to 200 yds. wide. There trunk are taken above the protective are, however, one or two outliers still stand, the dead leaves either fall or are within the general gorge area, but quite blown off, leaving the trunk bare (Fig. separate,and forming "islands." 26). In the field a Iarse number of seed- One feature rarely noted amongst PRINCIPES IVol. :l standsof palms was the number of curv- though interferenceby other plantscan- ing, twisted, and crooked trunks. From not be overlooked. Reference to the observationsit was not possible to de- plate (Fig. 52) in my previous article cide what had causedthe trunksto grow will show that the tall palm is greatly in this position (Fig. 30, right). al- arched; but as the photo shows lFig.

26. Palms shorv the retained apron oi old fronds in the centre of a thicket, but bare trrrnks above and to right. les9l LOTHIAN:LIVISTONA 55 "S." 29 ) the trunks are in the forrn o{ an probably in a soil deficient in organic The height of these palms is approxi- matter. mately 60 feet. The rate of growth following the pro- Seetllings in the tr'ield duction of the initial leaf is not known, but it undoubtedlydepends on the water The initial seedlingleaf is single, and supply. Rain in this region is by no except for some broadening towards the means re$ular, although there is prob- apex, the next three to six leaves are ably a certain amount of underground undivided. This broadeningof the la- water which the roots o{ these mina proceeds, then the leaf divides at can draw on freely. When the seedling the tip, and in the lOth or l2th leaf a has developeda trunk approximatelf palmate frond develops. The young the diameter of the mature palm, the leaves have weak spines along their trunk commencesto elongate. This may edges but these quickly disappear. As take at least l0 to 12 yearsin the fielj. the subsequent leaves occur, the spines At this stage the petioles may still be beconrernore rigid and stronger along coloured,but the lamina commencesto their petioles, until those along the changefrom red to glaucouscolour, and stalks of the initial palmate fronds are when the trunk is about 6 feet hish nor- quite strong and sharp and are small mal greenleaves are produced. replicas of those to be seen along the Undoubtedly the rate of growth also mature leaf stalks. determines not only the age but the The leaves of the seedlings growing height at which this palm reachesma- amongstthe mature palms are green and turity (flowers). Numerous specimens never assume a reddish hue. It was were examined,but none had uroduced noted, however, that in those seedlings flowers if the trunk was under l5 feet. growing away from the rnain colony However, palms of 25-30 feet showed of palms the leaves were a deeper col- old flower headspresent and it is likely our, and in one or two instancesthey that somewhereabout this height they had already commencedto turn reddish. commenceto flower. It would appear that the young plants It was previonsly stated that flower- "probably assumethe reddish tinge to dull red and ing is annual." In the main almostcinnamon red only when growing this statement remains substantially in fully exposed sites, and more espe- true, becauseat this height the palms cially when growing as individual seed- are likely to be drawing freely on under- lings well separatedfrom the main palm ground water supplies and are therefore (such colony as in rock crevices or no longer dependenton seasonalrains. pockets). On the other hand, without Prolonged droughts only would curtail exception, the seedlingsgrowing in ex- regular growth and flowering. posedsites had red leaves,and it was not A large number of seedlingswas ex- until theseplants commencedto produce cavated in an endeavour to trace their a trunk that the colour of the leaves root growth and root developmentdur- changedfrom red to a glaucouscolour. ing the young years. Following germi- It would appear, therefore, that to nation, roots descend rapidly. The juvenile produce red or young foliage crown was at least 4-6 inches below the under cultivation it is essentialto grow soil level and the roots of seedlinsshav- seedlingsin a hot fully exposedsite, and ing three or four leavesextendej to at 56 PRINCIPES [Vol. 3 least lB inches deep. By this stage Five weekslater (5th September) no there was also considerablelateral root seedlingshad occurred in groups A and spread. As can be seenfrom the accom- E. In group B there were two seedlings, panying photograph (Fig. 32), the de- and in groups C and D one seedlingin velopmentof the seedlingsfollowing the each had appeared above the ground. production of the shoot and fourth leaf By 27rh September,the following ger- is fairly regular, but the trunk develop- minatibns had taken place: Group A- ment is extremelyslow, one seedlingcommencing to show above the soil ; Group B - two seedlings; Germinntion ol Seed Group C-one seedling; Group D- From the seedcollected in the field, a eight seedlings, five several inches high quantity was sown under controlled con- and three just breaking the soil level; ditions at the Adelaide Botanic Garden. Croup E-no germination. Before sowing the seed was sorted into On 27th October, the following was five groups. (SeeFig. 2?.) the progressof the five batchesof seed: Group A-two seedlingswere growing, Group A: The thin but fleshy exocarp one freely with an upright leaf, the of brownish-plum colour was entire. other slowly with an abnormal leaf This was presentseason's seed. (curled and prostrate) ; Group B-two Group B: Exocarp covering was seedlings produced, one growing slightly chipped, brownish'black in strongly, the other very weak; Group C colour, but at least 75 per cent of the - two seedlings which subsequently individual seed was covered. Prob' died; Group D-nine seedlings well ably present season'sseed. above the soil level, all growing very strongly; Group E-no germination. Group C: The covering had started to On l9th November,approximately 15 rot away, was comparatively thin, weeks after sowing the seed. the pots grey coloured, and in some instances were removed from the propagating pit, covered less than approximately 50 eachwas emptied, and a checkwas made per cent of the seed. This could be on the number of seedswhich had ger- seedof previousyears (seasons). minated, the condition of the seedlings Group D; The covering was entirely re- and of the remaining seeds. The results moved, and representedfresh seed. were as follows: Group /: This comprised obviously old Group l-Four seedlingsproduced, of seed. The fruit was in some instances which two seedshad germinated re- deeply pitted or indented, black in cently and were just pushing through colour, and when cut in half before the surface of the soil. The other two sowing, it appearedeither dead or al- were well established. The seedling most so, with the curled and prostrate leaf had continued to gror,vbut at a rate slower sown on lst The five lots of seedwere than the other. The length o{ leaf in August in our usual seed soil mixture the vigorous seedling was l2 inches. (based on John Innes Standard) in Within the seedsoil, five further seeds 4-inch pots which were plunged into had recently germinated. One seed bottom heat of 80' F. Ten seedsof each failed to germinate, but was still type were sown to a pot. viable on dissection. lesel LOTHIAN: LIVISTONA JI

@dfu E 27. The various groups of seeds which were used in germination tests, Note the cracked appear. ance of Group E. Croup B-o{ the ten seeds, four had vigorous primary leaf up to 14 inches germinated. Two of the seedlingshad long, including three seedlingswhich died, the two remaining were growing had commencedto produce a second strongly. Within the seed soil two leaf. One seedling had subsequently further seeds had germinated into died. The primary leaveswere green, strongly developing seedlings. The somewhatchannelled, and along their primary root was well developedand edges were fine but distinct bristles. the shoot (plumule) was commencing Croup E-No germination. After care- grow. to Four viable seedsremained. fully washing the seedsoil, five rotten Croup C-A total of six seeds germi- seeds only were recovered, the re- nated and one of thesehad produced mainder had disintegrated. All were a sturdy primary leaf well above the probably dead before sowing. level of the seedsoil. Of the remain- The above results are rather inter- ing five seedlings, four had com- esting (although the number of seeds menced to produce shoot growth used was limited). It would appear above the ground, but subsequently that the fleshy fruit covering may con- died, and one had just broken the soil tain an inhibiting agent such as is level. The four remaining seeds were known to exist in the fleshy covering of rotten and infertile, and quite were many fruits (e.g. Rosaceae)which pre- likely dead before sowing. vents the seed from germinating until Group D-All ten seeds germinated. this has rotted. It wilt be noted ih"t ut Nine seedlingshad each produced a this cover rotted, so germination occur- "root 28. These palms rarely produce mounds," and it would appear that external cattses contribute to their develoPment. "straight" 29. A view looking rup the trtrnk of the palms shown in I'igure 30.

30. Grove of Liaistona Mar- iac showing seedlings (cen- tre) and variorrs stages of growth. Note twisted trunks on right. Tall palm in centre is approximately 55-60 feet high. 60 PRINCIPES lVol. 3 red. The five seeds of Group A which made to some of the seedlings dying had germinated by the middle of No- subsequent to germination. Two rea- vember but had not yet produced shoot sons can be advancedfor this happen- growth above ground level, showed ing. The first is that the bottom heat was maintained strong deterioration of the seed cover. well at B0o and on some days varied between85o and 90'F. As It would appear essential before sow- this temperature is too high for the well- ing seedsoI Liuistona (and possibly of being of the seedlings,the pots should other palms as well) that this soft suc- be removed to slightly cooler tempera- culent covering should be removed, and tures following germination. further, that fresh seedsonly should be The second reason, either coupled sown. This appearsto be the explana- with the above or independent,is that tion for the rather contrary results ob- for a very short period the bottom lay- tained in Group C, of which the four ers of the seed soil dried out. The ef- seedswhich were rotten (and probably fect of this can be clearly seen in Fig. dead) were in actual fact old seed, 3IAa. (Fig. 31Ab is a crosssection o{ a while the fresh seed germinated. fertile seed,showing the flesh still firm Deuelopmefi ot' Seed,lings and shiny whereas the remains of the In the above results. reference is germinated seed were mealy.) ^@ ,i.,,iwWwffi[@ i , wil'tl \\ )j \\

D

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3t. Germination oI Liuistona Mariae. A, cross section of seeds; Aa, seed germinated but sub- sequently died-note mealy appearance of endosperm; Ab, viable but ungerminated seed: B, stages of germination-note that the plumule commences growth below the level of the seed: C, greatly enlarged and somewhat later stage oI Bd, showing development of leaf sheath: D, various stages lesel LOTHIAN:LMSTONA 6l

SO|L LEVEL

of growth subsequent to the production of the lea{ sheath-note hardened leaf tip_ at- B. weeks afte"r germination: E, seedlings at 15 weeks a{ter germination showing stages of second leaf, destruc' tion of endocarp and (d) an abnormal leaf. PRINCIPES [\'o1.3

32. A range o{ seedlingsdug rrp in the field. Tl.re ages of the individual plants range from one season(left) to probably 7-10 (right).

The first developmentof the seedling in a descendinghabit, are being freely occurs as a thin cotyledon which for produced. the lower three-quartersof its length In Fig. 3lDa-c the subsequentde. very quickly thickensto almost double velopment of the young seedling is the initial diameter. At approximately clearly shown. It will be seenin 3IDc I inch below the seedthe primary shoot and to a lesserextent in 3lDb that the commencedto develop where this coty- tip of the leaf has now commencedto ledon is thickest. The staeesare clearlv wither and at this staee the hard rein- seenin Fig. 3l Ba-d. forcedtip is disappeaiing.Fig. 3lEa-c In Fig. 3lC the young shoot is now shows the subsequent development of ready to break through the soil, but it the seedling. Ground level has been will be noted that the apex of the leaf marked and it will also be noticed that has developeda very thick sharp but the tip is ready to break off. The pri- stout point. This is extremelysharp. If mary root has almost completely dis- the finger is pressedon it the skin can appeared,and a fibrous root system is be punctured. It will be noted also at now well developed. this stage that the primary root has It was previously stated that "con- commencedto shrivel and the first of tractile roots may be responsible for the true roots, developing laterally but drawing the crown of the below l959l I.OTHIAN: LIVISTONA ground level." From the current in- are washed into crevices or holes could vestigations it is obvious that the crorvn be deeply buried and, while the seeds is already below ground level, but it is germinate, the depth of material above not clear lvhether in actual fact con- may be too grtat to allow the seedling tractile roots do pull the seedlings to reach the soil surface before the down further irrto the s

PLANTINGS OF LIVISTONA MARIAE IN AMERICA

The following is an extract from a I was also able to compare seed samples letter from Mr. Nat J. De Leon. Miami. which che,ck out. Incidentally, the orig' F'lorida, who writes: inal date in the inventory records is De' "Further Mont' information concerning' cember 11, 1931. The trees at the the same seed plantings of Liuistona Mariae:-I have gomery collection from about five trees' since combed the area, the three major lot are represented by smaller and average 130 collections, and have pored through all These are all to These trees are planted on the records. The plants at Chapman 40 feet. higher land not receive the con- Field and the Montgomery collection and do introduction. In stant moisture found at Chapman Field. are from the same seed "In the U. S. Plant Introduction Inventory further checking trees and rec- records is the following listing: P. I. ords, I can conclude that all members 'Seeds No. 95077, presented by Council of this genus do very well with us, and for Scientific and Industrial Research. are fast growers when once planted out. Canberra; collected at Palm Valley, This includes those from more tropical Central Australia.' There can be no areas, as we have at least four species doubt that the palms correspond to f rom the Philippines that grow like weeds." this introduction. At Chapman Field I This information is most interesting. found some 12 trees. These are all The very rapid growth which these planted along a stream that was man- plants have made rather astounds me. made during the early days of that If. as stated. the initial introduction was station. The stream is shallow, but made in Decemher 19i31.the srowth has there is always enough water in it to been most remarkable. I can only con- keep the surrounding ground moist. clude that where a constant supply of The trees average in height from 4,0 to water is available, together with mild 60 feet. Only the tallest of these fruited to warm conditions, grolvth is con- this year, and bore quite a heavy crop. tinuous rather tharr seasonal.