Flora of New Zealand Series

Fls in axillary or terminal racemes or spikes, infls sts compound. Calyx us. deeply and almost equally 4-lobed, the fifth lobe when present us. smaller. Corolla short- or long-tubed, with 4 subequal spreading lobes. Stamens 2, anthers held above tube. Style long, stigma capitate. Capsule dehiscing by the sagittal splitting of the septum and each carpel opening by distal median suture through the septal wall and in varying degrees also through the locule wall; septum us. across widest diam. and capsule ± dorsally compressed; seeds us. flattened and smooth. Ever-green shrubs or small trees with lvs opp., us. entire though often toothed in seedling, rarely showing veins other than midrib. Perhaps c. 100 spp., mostly endemic in N.Z. but two shared with S. America and one of them extending to Falkland Is; a few spp. in Tasmania, south-east Australia and New Guinea. Type sp. H. magellanica = H. elliptica.

SYNOPSIS

• • Capsule compressed dorsally; septum across widest diam.
  • A. "Subdistichae". 
  • • Infls lateral. Lf-bud with narrow pointed sinus. Bracts sts, but not us. opp. Rather closely branched, us. small, shrubs with smallish lvs tending to distichous arrangement
    • (a)
    • • Infls corymbose; bracts small; pedicels sts long:
      • 1. diosmifolia, 2. divaricata
    • (b)
    • • Lowest infls often tripartite; pedicels short or absent:
      • 3. insularis, 4. colensoi, 5. rupicola, 6. rigidula, 7. venustula
    • (c)
    • • Infls simple (very occ. tripartite); pedicels short, bracts small:
      • 8. brachysiphon, 9. cockayniana, 10. canterburiensis, 11. vernicosa
  • B. "Apertae". 
  • • Infls lateral. Lf-bud with broad, ± square sinus. Bracts not opp., small. Fls pedicellate. Many-branched shrubs, lvs medium to large:
    • 12. elliptica, 13. speciosa, 14. townsonii, 15. gracillima, 16. salicifolia, 17. corriganii, 18. pubescens
  • C. "Occlusae". 
  • • Infls lateral. Lf-bud without sinus. Bracts not opp., mostly small. Fls pedicellate. Many-branched shrubs, occ. small trees
    • (a)
    • • Lvs mostly > 3 × 1 cm.:
      • 19. ligustrifolia, 20. acutiflora, 21. stricta, 22. obtusata, 23. macrocarpa, 24. breviracemosa, 25. bollonsii, 26. dieffenbachii, 27. barkeri, 28. chathamica
    • (b)
    • • Lvs either < 8 mm. wide or < 3 cm. long:
      • 29. evenosa, 30. truncatula, 31. parviflora, 32. subalpina, 33. fruticeti, 34. urvilleana, 35. traversii, 36. strictissima, 37. rakaiensis, 38. glaucophylla, 39. topiaria
  • D. "Subcarnosae". 
  • • Infls lateral. Lf-bud without sinus. Bracts often opp. Fls mostly sessile or only shortly pedicellate, giving compact infl. Capsule often turgid. Mostly ± decumbent rather woody shrubs with lvs ± fleshy and/or glaucous
    • (a)
    • • Fls shortly pedicellate; bracts not opp., mostly < calyx-lobes:
      • 40. decumbens, 41. recurva, 42. albicans, 43. matthewsii
    • (b)
    • • Fls sessile; lowest bracts opp., = calyx-lobes:
      • 44. amplexicaulis, 45. allanii, 46. gibbsii, 47. pinguifolia, 48. buchananii, 49. pimeleoides
  • E. "Buxifoliatae". 
  • • Infls lateral or terminal or both. Lf-bud with ± heart-shaped sinus. Bracts opp., the lowest large and ± lflike in texture. Fls sessile. Small shrubs with small stiff lvs and strict us. erect twigs:
    • 50. odora, 51. pauciramosa, 52. pauciflora
  • F. "Flagriformes". 
  • • Infls terminal, simple. Lf-bases connate. Bracts opp., often slightly > lvs. Fls sessile. Shrubs, us. low-growing, with twigs of whipcord form
    • (a)
    • • Nodal joint well-marked; anterior calyx-lobes free except near base:
      • 53. tetragona, 54. subsimilis, 55. coarctata, 56. hectori, 57. laingii, 58. propinqua, 59. lycopodioides, 60. poppelwellii, 61. imbricata
    • (b)
    • • Nodal joint obscure, anterior calyx-lobes almost completely fused:
      • 62. salicornioides, 63. annulata, 64. armstrongii, 65. ochracea
    • (c)
    • • Capsule laterally compressed and ± didymous, very small:
      • 66. cupressoides
  • G. "Connatae". 
  • • Infls terminal, sts lateral also, forming compact head. Lf-bases connate. Bracts opp. and ± lflike. Fls almost sessile. Low-growing to decumbent shrubs with ascending tips, lvs us. imbricate:
    • 67. petriei, 68. ramosissima, 69. epacridea, 70. haastii, 71. benthamii
  • H. "Paniculatae". 
  • • Infls terminal and us. compound. Lvs all petiolate, members of a pair diverging early in bud. Fls mostly sessile. Capsule turgid, ± didymous. Seed narrow, spindle-shaped. Low-growing half-woody shrubs, lvs regularly toothed:
    • 72. raoulii, 73. lavaudiana, 74. hulkeana
• • Capsule compressed laterally, especially towards tip; septum across narrow diam. Infls lateral, few-fld. Foliage almost black when dry
  • I. "Grandiflorae". 
  • • Lvs large, toothed, petiolate, members of a pair diverging early in bud. Fls very large. Short woody shrub with lfy twigs:
    • 75. macrantha
  • J. "Semiflagriformes". 
  • • Lvs small, margins ciliate, bases connate. Fls small, unisexual, plants us. dioecious. Much-branched very low shrubs from stout woody base, twigs of semi-whipcord form with close-set lvs almost or quite hiding very slender stem:
    • 76. ciliolata, 77. tetrasticha, 78. cheesemanii, 79. tumida

The name koromiko (korokio in the Far North) is commonly used for the larger willow-lvd spp. In the high country the name "boxwood" us. refers to some small-lvd Hebe.

Cheeseman (Man. N.Z. Fl. 1925, 778 et seq.) treated under Veronica three "Divisions", Hebe, Pygmea and Euveronica. Pennell (Rhodora 23, 1921, 2-5) regarded Hebe as a genus distinct from Veronica for the following reasons: septicidal dehiscence of capsule; shrubby or arboreal habit; fls in specialized axillary racemes; austral distribution; "exceptionally baffling tendency to form local races". Later work, mostly by Frankel, has shown that basic chromosome numbers, all × = 20 or 21, differ from those within Veronica. The genus Hebe has been generally adopted in N.Z. and many of the necessary name changes were made by Cockayne and Allan (T.N.Z.I. 57, 1926, 11-47).

The species exhibit great diversity and not all fit the diagnosis of Pennell. In the capsule the septum always splits sagittally but there is us. also some loculicidal splitting which sts extends almost to the base, especially in those spp. where the septum is narrow. In the type sp. H. elliptica, and a great many others, "the stem continues indefinitely as a vegetative axis, its leaves opposite throughout, while the inflorescence is localized in specialized axillary racemes", as described by Pennell, and here fls are irregularly arranged and pedicellate, and bracts are small. Such specialized infls are connected through many intermediates with the relatively simple ones seen in whipcords and some other spp., where the stem terminates in a simple spike with short internodes, the fls are opp. and sessile, and the bracts are ± lflike. In the most complex infls, lvs subtending individual spikes or racemes are reduced to bractlike structures; two types occur; (a) as a result of branching and elongation of a terminal infl., giving in the extreme case the large lax panicle of H. hulkeana; (b) as a result of branching of a lateral infl., the greatest complexity being where there is repeated branching, sts of the second order, giving a many-fld lateral corymb. Plants grown in good conditions in gardens sts tend to develop unusually complex infls.

H. hulkeana and the closely-related H. lavaudiana and H. raoulii stand rather apart from Hebe not only in their panicled infls but also in hardly shrubby habit, regularly toothed and us. Long-petioled lvs, open corolla with very short tube, turgid capsule and particularly in narrow slightly wrinkled spindle-shaped seeds with only a trace of wing.

Also difficult to accommodate in Hebe are the five spp. in which the capsules are strongly laterally compressed. In having the septum across the narrowest diam. they resemble Pygmea, Parahebe, and Veronica, but the septum is long, not short as in those genera, the fls do not agree with those of Pygmea and the growth forms and woody habit do not accord well with anything in Parahebe or Veronica. The four spp. which Frankel and Hair called "semi-whipcords" would have good claims to separate generic rank were it not that H. macrantha with similarly shaped capsule is quite Hebe -like in appearance, apart from its exceptionally large fls.

The "whipcord" spp. (subgenus Pseudo-veronica of J. B. Armstrong 1881), though quite closely related to H. odora and its allies, form a well-defined group characterized by a peculiar habit and sharing a number of other features. For practical convenience the key is placed near the figures and descriptions.

In all the more typical spp. of Hebe the two lvs of an opp. pair do not diverge from one another until they are almost full-grown and the large closed lf-buds give a characteristic appearance to many plants. In some spp. the lf-margins remain in close contact throughout their length, but in those where a true petiole develops early the two lf-bases are separated in the bud by a distinct gap or sinus and this is a generally useful diagnostic character; it was noted by Petrie (T.N.Z.I. 48, 1916, 189) in distinguishing V. evenosa from V. laevis. In only a few spp. do the lf-tips break apart in the early bud.

The scars left when lvs fall are very persistent and, especially where internodes are short, the resulting stem pattern is often characteristic of the sp., though not easily described. Under binocular microscope lf-margins sts display disinctive features, and in some spp. stomata are strikingly obvious. Hydathodes are sts visible to the naked eye as small thickened us. colourless patches of cells, either at the lf-apex or forming slight widely spaced projections on margins of longer lvs. Where the ovary is pubescent the style us. bears spreading hairs also, and these are sts more easily seen.

In the mountains flowering is us. confined to a few spring-summer months, but in some lowland spp. fls appear over a long period, extending either from late summer into winter or from winter into late spring or early summer. Capsules develop quickly and seed is shed not long after corollas fall, but old capsules often remain on the bush throughout the non-flowering period. Approximate flowering times are given, based on available specimens.

Hooker (Handbk N.Z. Fl. 1864, 204) noted that many spp. "probably are, if not bisexual, still partially so, the two sexes presenting differences in the size of the stamens and calyx and capsules, a point worthy of the close attention of the colonist". Male sterility has since been recorded in many spp. (Frankel J. Gen. 40, 1940, 171-184). One of the peculiarities of the semi-whipcords is that the fls are apparently regularly unisexual, and the plants are almost completely dioecious.

Various schemes for subdividing Hebe have been proposed (e.g. Bentham in DC. Prodr. 10, 1846, 459; J. B. Armstrong in T.N.Z.I. 13, 1881, 349; Cheeseman in Man. N.Z. Fl. 1906, 492) but where sectional or subsectional names were given no types were indicated. A somewhat novel arrangement is adopted here, but no attempt is made to correlate the subdivisions with those previously named and no precise rank is suggested for the ten groups recognized; for easy reference informal Latin names are given, each indicating a feature that is characteristic of but not necessarily exclusive to the group in question. These groups are represented very unequally in different parts of the country. All the lowland North Id spp. and those of Kermadec, Three Kings and Chatham Is are included in A, B and C, and a few of the North Id mountain spp. fall into E and F. The other five groups are not found in North Id but all ten occur in South Id; G, I and J contain only mountain spp., while D and H have a few lowland representatives, most of them rupestral. B and E each take in at least one sp. extending to Stewart Id and the Subantarctic Is, F has one Stewart Id sp., and one sp. endemic to Auckland and Campbell Is is placed in G.

Since the second edition of Cheeseman's Manual some 26 new taxa have been proposed in N.Z. Hebe. At least half of these, mostly from the original collections, are represented by plants growing at Otari Open Air Plant Museum, where also living plants of most of the older spp. (many of them from type localities) have been readily accessible for study and comparison. A. P. Druce has transplanted a fine collection of North Id spp. from the wild into the garden at Soil Bureau, Taita, and has made available information about their ecological and geographical relationships. Special help has been obtained from Kew in the shape of photographs of type specimens taken by Miss N. T. Burbidge of Canberra, and illuminating notes about minutiae prepared by Dr. R. Melville. Even with these advantages and with the excellent basis of the Manual, Cheeseman's 1925 statement describes very exactly our experience and modest claim for the present revision; he wrote: "The arrangement and limitation of the species, and the preparation of the necessary diagnoses, has proved to be a most difficult and perplexing task, and I am far from satisfied with the result. But, imperfect though it may be, it represents many months' assiduous study, and the examination of some thousands of specimens, and is, at any rate, an honest effort to clear away some of the difficulties which impede the study of the genus".

Fig. 35. HEBE. Capsules, calyces and bracts: lateral views on left and below, anterior views on right and above. © All rights reserved. [Image: 4X1J]

FIG. 36. HEBE. Leaf-buds: above, base from midrib view; below, base from margin view, those to left of line with sinus. © All rights reserved. [Image: 4X1K]

FIG. 37. HEBE. Branchlets of whipcord species: drawn from type material except H. tetragona and H. cupressoides; shown in dry condition except H. salicornioides. © All rights reserved. [Image: 4X1L]