6. Breeding systems and h y b rid iz atio n
6.5. Hybridization in nature and cu ltu re
Cockayne et Allan 1934, M oore in Allan 1961 who cites H ooker (1854), M etcalf 1987, Rooney 1987, Chalk 1988, A.P. Druce pers. comm., P. Gamock- Jones pers. comm.)
Hybridization in nature is limited because distri
bution o f most taxa is localized, but fertile and sterile hybrids are to be found in many habitats. The chromosome number varies from monoploid (n=20, n= 21) to diploid (n=40, n=42), triploid (n=59, n=60, n= 61, n=62, n=63) groups (Appendix 1 and Chapter 4). Cockayne and Allan (1934) list for example 43 wild hybrids, while M oore (Allan 1961) lists 13 wild hybrids and 15 horticultural forms.
Man has been involved in breeding since the first species came to Europe in 1776 and now more
cultivars than species are present in Europe (mainly Britain) while the opposite is the case in New Zealand (Chalk 1988). An updated "Hebe Interna
tional Check List o f Cultivars" is in preparation (Chalk 1988, L.J. M etcalf pers. comm.) and about 500 cultivar names are being examined for validity (L.J. M etcalf pers. comm.). H ebe species and culti
vars make very nice evergreen shrubs in gardens and parks, and I will assert that the potential for further use as ornamentals in temperate to subtropi
cal climates is almost unlimited.
Different ways to establish new cultivars are:
1) a breeding program,
2) collection and selection o f types from the wild, 3) casual crosses happening within a collection.
Breeding programs are not very common. Hebe breeding at Auckland Regional Botanic Gardens is an example. The first crossing was m ade in 1979 (Hobbs 1986), and the program started in 1982. The main object is “to produce attractive cultivars which perform well in Auckland gardens. Pest and disease resistance is a particularly important requirem ent”
(Hobbs 1988).
The first crosses included//, speciosa crossed with some o f the more disease resistant varieties in the Auckland collection. Out of 232 seedlings raised, 5 proved improvements at various positions within the Gardens and were selected and named in 1988:
H. ‘ W iri Joy‘, rose pink flowers on a plant with assemblance to H. ‘Inspiration1,
H. ‘W iri Splash1, m auve flowers, attractive goldengreen foliage
H. ‘Wiri Jew el1, magenta flowers, texture and undulating margins o f foliage and pointed tips H. 1 Wiri Spears1, long spear shaped spike, mauve flowers, quick growing
H. ‘ Wiri Grace ‘, mauve flowers, compact growth, with H. stricta, H. speciosa and H. bollonsii in its parentage
The large variation within Hebe species led some plantspeople to select and name cultivars from wild grown species. An exam ple is the British nursery
man Graham Hutchins, County Park Nursery. During expeditions into the native bush o f New Zealand he collected and selected forms o f different species of H ebe and other plant species. The forms are grown on at H utchins’ nursery near London, England. If the forms maintain their differences in shape and behaviour, they are given cultivar names. Crossings are made in the nursery as well, and a number of cultivars has been released (G. H utchins pers.
comm.).
M any people find seedlings in their garden.
Seedlings that look a bit different from their possib
le parents; and an unknown number o f these seed
lings are named and find their way into com mercial production both in Europe and New Zealand (L.J.
M etcalf pers. com m., pers. obs.).
6.6. Discussion
M onoploids, n=20 and n=21, have evolved to dip
loids and triploids. A t present new forms, either species (stable from seed) or hybrids (mostly un
stable from seed) occur both in nature and in culti
vation. From which origin the monoploids arrived is not known, but suggestions have been made (Chap
ter 3). Evidence o f phylogeny would give important information for understanding the characteristics of the Hebe genus.
Fertility is high, and plants have a high propor
tion o f gender dimorphism. M ost species are self
compatible but mechanisms which avoid self-polli
nation have evolved and flies and native bees seem to be the most im portant pollinators in alpine and subalpine habitats. These three recognized features must correspond in their functions. Firstly, it does not seem logical that plants are self-fertile and have evolved dimorphism at the same time. But because the populations o f plants often are local, the flow ering time short and the weather cool and humid (for example in a wet subalpine habitat in the South Island), the plants must be advantaged by developing mechanisms which secure the highest degree of cross-pollination. Then, if the climate conditions are poor, the flower longevity increases and polli
nation is delayed until the weather improves and pollinators are available. Therefore, I hypothesize that the ability to be self-pollinated is only used if cross-pollination can not be carried out.
How pollination takes place in Hebe taxa gro
wing in montane and lowland habitats has not been studied but birds m ight be involved. Flower colours also indicate pollination by flies and native bees in higher altitudes (white and pale colours). Other pollinators m ight be attracted at low altitudes (blue, purple and red colours). The bright coloured species have long tubed corollas which in other plant spe
cies are found to be more likely to have bird- pollination. Further studies are required.
Patterns of inheritance in the genus Hebe have not been studied, but casual hybridization in nature and cultivation happens frequently. The genus is still
under development, adaptations to climates and ecological system s are im proved by natural selec
tion of genotypes. If breeding systems becam e known, the hybridization within the genus would be understood and would be valuable for controlled
Thank you to Professor Arne Skytt Andersen and Mr. Ole Voigt Christensen for the support of carrying out one year o f my Ph.D. study in New Zealand where Hebe can be investigated in its own envi
ronment.
References
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Gamock-Jones, P. J. 1976. H ebe rapensis (F. Brown) G am ock-Jones comb. nov. and its relationships.
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G am ock-Jones, P.J. & Molloy, P.J. 1982. Poly
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Hong De-yuan. 1984. Taxonom y and evolution of the Veroniceae (Scrophulariacae) with special reference to palynology. Opera Botanica 75: 5- 61.
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Laing, R.M. and Blackwell, E.W. 1928. Plants of N ew Zealand. W hitcombe and Tombs Limited, Christchurch. 499 pp.
Lee, J., Brooks, R.R., Reeves, R.D. & Boswell, C.R.
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M acaulay, J.U. & Beavis, E.J. 1983. Senior Atlas for New Zealand. Collins (NZ) Ltd. & Long
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Moore, L.B. 1967. How to look at//e ö e.T u a ta ra 15:
10-15.
M oore, L.B. 1973. Protogyny in Hebe “Panicuala- tae” . New Zealand Journal o f Botany 11: 173- 176.
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Poole, A.L. 1987. Southern Beeches. New Z ealand Department o f Scientific and Industrial R ese
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383-437.
Sakai, A. & W ardle, P. 1978. Freezing resistance o f New Zealand trees and shrubs. New Zealand Journal o f Ecology 1:51-61.
Saunders, E.R. 1934. A study from the viewpoint o f certain floral charaters. Linnean Society’s Jour
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Scott, D. 1977. Plant ecology above timber line on Mt Ruapehu, North Island, New Zealand. 1. Site factorsand plant frequency. New Zeal and Journal of Botany 15: 255-294.
Simpson, M.J.A. 1976. Seeds, seed ripening, germ i
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Thomson, G.M. 1927. T he pollination o f New Zealand flowers by birds and insect. T ransac
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Wardle, P. 1963A. E volution and distribution o f the New Zealand flora, as affected by quaternary climates. New Zealand Journal of Botany 1(1):
3-17.
Wardle, P. 1963B. Grow th habits of New Zealand subalpine shrubs and trees. New Zealand Jour
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A p p en d ix 1
Habitat and distribution of New Zealand Hebe taxa
Characteristics and specifications on New Zealand Hebe species and varieties: Names (Allan 1961), tagnames for unnam ed species (A. P. Druce pers. comm.), botanical sections (M oore in Allan 1961 and A.
P. Druce pers. com m.), chromosom e numbers (Hair 1967, A. P. Druce pers. com m .), original habitat (A. P.
Druce pers. comm, and Eagle 1986) and distribution (A. P. Druce pers. comm, and Eagle 1986).
H abitat Hydro- G row th-
Plant-N am e altitudinal zone logy Landform form height (m)
A . Subdistichae, leaf bud with narrow pointed sinus; dorsally com pressed capsule; inflorescences lateral, shrubs with sm allish leaves tending to distichous arrangem ent.
Chromosome number: n = 20
|jÜ | H. diosm ifolia (spring flow.) m ontane-low land wet forest m argin, cliff bushy shrub 2 (A. Cunn.) Ckn. et Allan
[ :‘ j H. insularis m ontane-low land wet m aritim e cliff e rect or sprawling 1
(Cheesem .) Ckn. et Allan shrub
H l H. colensoi m ontane-low land dry cliff spreading bushy 1
(Hook, f.) Ckn., incl. H. c. var. colensoi and H. c. var. hillii (Col.) L.B. M oore shrub
I J I I H. rupicola subalpine-low land dry cliff, rock e rect shrub 1.5
(Cheesem .) Ckn. et Allan
3 H . rigidula subalpine-low land dry forest m argin, rock sm all shrub 0.6
(Cheesem .) Ckn. et Allan, incl. form (II) (A. Eagle)
[ I H. rigidula form (I) (A. Eagle) m ontane-low land w et calcareous cliff sm all shrub 0.6 tagnam e “H. L ady” A.P. Druce
H H . sp. (q) Eagle, tagnam e alpine-subalpine dry tussockland,rock dom e-shaped shrub 1.5
“H. aff. rigidula” A.P. Druce
1 1 H . canterburiensis alpine-subalpine w et tussockland straggling or 1
(J.B. Arm st.) L.B. M oore rounded shrub
Chromosome number: n = 21
I “ 1 H. vernicosa subalpine-low land w et-dry forest spreading shrub 1
(H ook, f.) Ckn. et Allan
Chromosome number: n = 40
1 1 H. divaricata m ontane-low land w et forest margin e rect shrub 3
(Cheesem .) Ckn. et Allan
U H H. diosm ifolia (sum m er flow.) m ontane-low land w et forest margin e rect tall shrub 6 (A. Cunn.) Ckn et Allan
|jf j| H. carnosula (Hook, f.) Ckn. alpine-subalpine dry rock shrub 0.5
Chromosome number: n = 60
PUjjl H. venustula (Col.) L.B. M oore alpine-low land w et-dry tu sso ck lan d ,ro ck e rect bushy shrub 1.5 (syn. H . brachysiphon Sum m erhayes)
|5jj| H . cockayniana alpine w et tussockland, rock erect shrub 1 (Cheesem .) Ckn. et Allan
Chromosome number: n = ?
1 1 H. dilatata Sim pson et Thomson alpine w et rock p rostrate spreading ?
y y and H . craw ii M. H eads shrub
II
A. Subdistichae
III
N am e
H abitat
Hydro-altitudinal zone logy Landform
G row th-form
Plant-height (m) B. A pertae, leaf bud with broad, + square sinus; dorsally com pressed capsule; inflorescences lateral; m any-branched shrubs;
leaves m edium -large.
Chromosome number: n = 20
1 1 H. elliptica (Forst, f.) Pennell incl. H . e. var. elliptica and H.
m ontane-low land e. var. crassifolia
w et m aritime cliff Ckn. et Allan
bushy shrub 2
|f8j|
H. speciosa(A. C unn.) Ckn. et Allan
m ontane-low land wet maritim e cliff rounded bushy shrub
2
I ; 1 H . townsonii m ontane-low land wet calcareous cliff upright shrub 2.5
(C heesem .) Ckn. et Allan
1 j H. p ubescens m ontane-low land w et (Banks et Sol. ex Benth.) Ckn et Allan
incl. form (I), tagnam e H . p. var. “B arrier” A. P. Druce
forest margin m any branched shrub
2
P 1 H. salicifolia m ontane-low land wet-dry forets m argin, cliff erect shrub 5 (Forst, f.) Pennell
1 H. sp. (v) Eagle m ontane-low land wet cliff shrub 2
tagnam e “H. m okohinau” A. P. Druce
Chromosome number: n = 40
I" I H. gracillim a m ontane-low land wet-dry forest margin spreading shrub 2 (K irk) Ckn. et Allan
H. corriganii Carse (see O cclusae)
Chromosome number: n = ?
IU I H. sp., tagnam e “H. U nuw hao” m ontane-low land w et rock erect multibranched ?
A. P. D ruce shrub
IV
N am e
flow ers pedicellate; m any-branched shrubs, occasionally sm all trees.
Chromosome number: n = 20
□
H . ligustrifolia m ontane-low land wet forest m argin shrub 3
(A. Cunn.) Ckn. et Allan
H . acutiflora Ckn. m ontane-low land wet forest m argin erect shrub 1
H . stricta var. stricta subalpine-low land w et forest m argin shrub 1
(Benth.) L. B. M oore m ontane-low land wet-dry m aritim a cliff shrub 2 var. m acroura
(Benth.) L. B. M oore
var. atkinsonii subalpine-low land wet-dry forest m argin,scrub, tall robust shrub 3
(Ckn.) L.B. M oore rock
form (I) A. Eagle, m ontane-low land wet cliff shrub 1
tagnam e “H. angustissim a” A.P. Druce
H . obtusata m ontane-low land w et maritim e cliff spreading prostrate 0.5
(Cheesem .) Ckn. et Allan shrub
H. bollonsii m ontane-low land w et cliff? erect shrub 1
(Ckn.) Ckn. et Allan
H . dieffenbachii m ontane-low land wet maritim e cliff low spreading shrub 1 (Benth.) Ckn. et A llan
H . barkeri m ontane-low land wet forest large shrub to 7
(Ckn.) Ckn. small tree
H . chatham ica m ontane-low land wet m aritim e cliff sm all trailing 1
(Buchan.) Ckn. et Allan shrub
H. traversii subalpine-low land dry forest m argin, scrub, com pact ball-like 2
(Hook, f.) Ckn. et Allan cliff shrub
H . treadwellii Ckn. et Allan alpine-subalpine wet rock sm all low shrub 0.5 incl. H. brockiei Sim pson et Thomson
H . sp. (x) A. Eagle, m ontane-low land wet cliff shrub 2.5
tagnam e “H. B artlett” A. P. Druce
H. parviflora var. angustifolia* m ontane-low land w et cliff lowgrowing or 2
(Hook, f.) L.B. M oore bushing shrub
(syn. Veronica squalida Kirk), tagnam e “H. squalida” A. P. Druce H. sp (h),
(syn. Veronica x bishopiane) m ontane-low land w et cliff H . sp (m ), tagname
"H. W hangarei" A.P. Druce m ontane-low land w et forest, cliff?
Chromosome number: n = 40
I 1 II m acrocarpa var. m acrocarpa, m ontane-low land w et forest
C. Occlusae
CD
*) The two varieties are regarded as tw o distinct species by A. P. Druce (pers. com m .)
**) This species is considered to be a variety o f H. m acrocarpa by A. P. Druce (pers. com m .)
***) These tw o varieties o f H. s tr id a are regarded as belonging to a separate species by A. P. Druce (pers. com m .)
N am e
Chromosome number: n = 40 continued
m H . stricta var. lata** alpine-low land wet
. m acrocarpa var. brevifolia m ontane-low land wet (Cheesem .) L.B. M oore*****, tagnam e “ H. brevifolia” A. P.
Chromosome number: n = 60
[fill H . m acrocarpa var. latisepala m ontane-low land wet (K irk) Ckn. et Allan
m H . m atthew sii alpine-subalpine wet
(Cheesem .) Ckn.******
C. Occlusae
****) There are num erous form s/geographic races o f H. glaucophylla in NW N elson, South Island, som e alm ost indistin
guishable from H. albicans except by chrom osom e num ber. These tw o species should belong to same botanical section (A. P. Druce pers. com m .)
*****) These variety is suggested to be given specific status (D ruce et al. 1979)
******) j-i m atthewsii is re-discovered by A. P. Druce 1989, from not being seen in nature since first discovery in 1906 by Cheesem an (A. P. D ruce pers. com m ., A llan 1961). The species is suggested by A. P. Druce to belong to ' 'O cclusae' ’ not “Subcarnosae” , because it has no sinus (pers. com m .)
Nam e
Habitat altitudinal zone
Hydro
logy Landform
Growth-form
Plant-height (m) D. Subcarnosae, leaf bud with out sinus; dorsally com pressed capsule; inflorescences lateral; bracts often opposite; flow ers mostly +decum bent rather w oody shrubs with leaves +fleshy and/or glaucous.
Chromosome number: n = 20
H I H . decum bens alpine-subalpine dry cliff, rock prostrate shrub 1
(J.B. Armst.) Ckn. et Allan
g H. albicans (Petrie) Ckn.* subalpine-low land wet cliff, rock, bog spreading shrub 1
□
H . recurva Sim pson et Thomson* m ontane-low land wet cliff, rock shrub 1g H. am plexicaulis alpine-subalpine dry cliff, rock
(J.B. Armst.) Ckn. et Allan
(incl. H. a. var. erecta Ckn. et Allan and H. allanii Ckn.)
■
H. gibbsii (Kirk) Ckn. et Allan alpine-low land dry rock spreading shrub 0.3m H. sp. alpine dry rock shrub ?
tagnam e "H. aff. pinguifolia" A.P. Druce
□
H. pim eleoides var. pim eleoides subalpine-low land dry rock small shrub 0.3 (Hook, f.) Ckn. et A llan (syn. H. p. var. m inor (H ook, f.) Ckn. e t Allan)□
H. pim eleoides var. rupestris** m ontane-low land dry rock straggling shrub 0.5 Ckn. et AllanH
H. pareora m ontane-low land dry calcareous cliff large shrub 1.5G am ock-Jones et M olloy
■
H. biggarii Ckn. subalpine dry cliff, rock sm all shrub 0.3Chromosome number: n = 40
a H. pinguifolia alpine-subalpine dry rock low or erect shrub 1
(H ook, f.) Ckn. et Allan
□
H. buchananii alpine-subalpine dry rock, cliff sprawling shrub 0.2(Hook, f.) Ckn. et Allan
□
H. sp. m ontane-low land dry rock small shrub 0.2tagnam e "H. aff. pim eleoides” A.P. Druce
*) regarded as one species with two varieties, H. albicans var. recurva and H. albicans var. albicans by A. P. Druce (pers.
com m .) (D ruce et al. 1987)
**) regarded as a species distinct from H. pim eleoides by A. P. Druce (pers. com m .) H . pim elioides var. glauca-caerulea is based on a hybrid (A. P. D ruce pers. comm.)
D. Subcarnosae
N am e
Habitat
Hydro-altitudinal zone logy Landform
Growth-form
Plant-height (m) E. B u xifo lia ta e, leaf bud with -(-heart-shaped sinus; dorsally com pressed capsule; inflorescences lateral or terminal o r both; bracts opposite, the lowest large and + leaflike in texture; flow ers sessile; small shrubs with sm all stiff leaves and strict usually erect twigs.
Chromosome number: n = 21
I I H. odora (H ook f.) Ckn.* alpine-subalpine wet I I /■/. pauciram osa var. pauciram osa alpine-subalpine wet
(Ckn. et Allan) L. B. Moore
I ?| H. pauciflora alpine-subalpine wet
Sim pson et Thom son
Chromosome number: n = 42
IU I H. sp. (u) form (I) A. Eagle alpine-subalpine tagnam e “H . anom ala" A. P. Druce
Chromosome number: n = 59
I I H. pauciram osa var. masonae alpine-subalpine wet L. B. M oore (Leonohebe m asoniae M. Heads)
Chromosome number: n = 63
I - v[ H. sp. (i) A. Eagle subalpine wet (Leonohebe m ooreae M. Heads)
bog, tussock, scrub shrub shrub bog, tussock
w et bog?, rock?
wet-dry scrub, tussock
b o g ,tu sso ck
bog, tussock
sm all shrub
shrub
robust shrub
1.5 0.5
0.2
1.5
shrub 1.5
*) This species is stated by H air (1967) to probably include more than one good species. H air recorded the chrom osom e num bers n = 2 1,42 and 63. Further a variety, H. odora var. prostrata, was recorded to have the chrom osom e num ber 2n=84.
E. Buxifoliatae
N am e
Habitat
Hydro-altitudinal zone logy Landform
Growth-form
Plant-height (m) F . F lagriform es, leaf bases connate; dorsally or laterally com pressed capsules; inflorescences terminal, simple; bracts opposite, often slightly > leaves; flow ers sessile; shrubs, usually low growing, with tw igs o f w hipcord form.
Chromosome number: n = 20
B l H. tetragona alpine-subalpine
(H ook.) Ckn. et Allan
Ifljfjl H. subsim ilis (Col.) M.B. A shw in alpine-subalpine var. subsimilis*
P 1 H. subsim ilis var. astonii alpine-subalpine (Petrie) M . B. Ashwin*
[5j|
H . coarctata alpine-subalpine(C heesem .) Ckn. et Allan*
|Hf
H. hectori alpine-subalpine(H ook, f.) Ckn. et Allan var. hectori* incl. H. subulata Simpson I Q H. hectori var. demissa alpine or
(Sim pson) M. B. Ashwin* subalpine?
I I H. laingii (Ckn.) Ckn. et A llan* alpine-subalpine
IUI
H. propinqua alpine-subalpine(Cheesem .) Ckn. et Allan
lüüil H . lycopodioides alpine-subalpine (Hook, f.) Ckn. et Allan var. lycopodioides . lycopodioides var. patula alpine-subalpine
Sim pson et Thomson**
wet tussock stout erect shrub 1
wet tussock shrub 1
wet tussock small shrub 0.3
wet tussock spreading shrub 1
wet tussock, bog small, 0.8
lata Simpson erect shrub 0.8
wet ?
wet tussock, bog? small shrub 0.3
wet-dry tussock shrub 1
dry tussock stout shrub 1
dry tussock decum bent shrub 0.2
dry tussock slender shrub 0.2
wet ? m any branched shr. 0.6
wet-dry bog erect shrub 1
dry tussock sm all shrub 0.2
dry scrub rounded shrub 2
dry scrub spreading shrub 1
wet tusso ck ,ro ck spreading shrub 0.7
*) T hese six species and varieties are considered to be part o f the wide spread species H. tetragona by A. P. Druce (D ruce et al. 1987)
**) This variety and species are considered to be part o f the species H. lycopodioides by A. P. D ruce (pers. comm.)
**) This variety and species are considered to be part o f the species H. lycopodioides by A. P. D ruce (pers. comm.)