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Volume 85, 1957-58
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A Descriptive Review of the Phaeophyceae of New Zealand

[Received by the Editor, March 28, 1957.]

Abstract

This paper presents a system of keys for the taxonomic and ecological classification of the New Zealand Phaeophyceae.

This review is intended primarily for the determination of specimens, and it is hoped that it will be of use as a short-cut to identification for the busy collector. A part of the research necessary for the compilation of this information was carried out under a special grant from the Research Fund Committee of the University of New Zealand, to whom the writer offers his grateful acknowledgments.

In the distribution notes, the word “south” signifies south of Cook Strait, including its Wellington shores, but its boundaries may in some cases be extended to include the area south of Lat. 39 degrees (the southern half of the North Island, from Waitara to Mahia Peninsula), when the word “southern” is used. “Northern” signifies north of Lat. 39 degrees.

In giving the ecological details a combination of letters has been used in two groups separated by a colon; the first group represents the tide-levels, thus: Hwnt-Lwst signifies “high water neap tides to low water spring tides”, and MT-LW signifies “mid tide to low water”. The second group of letters describes the type of shore, thus: EHRB-SR signifies “exposed horizontal rocky beach to sheltered rocks”, and ERF and P signifies “exposed rocky face and pools”.

First Combination Second Combination
L low E exposed
H high S sheltered
M mid M muddy
N neap R rocky or rocks
S spring B beach
W water F face, vertical
T tide P pools
H horizontal; gently shelving beach to and below LW.

After some of the names in Part I appears a number in square brackets which refers the student to Part II, where are given brief anatomical details for the more accurate determination of specimens, especially where some difficulty might arise. Where such details are felt to be superfluous they have not been given.

For one plant no name is as yet provided, but it should be recognized by the description given.

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1. Thalli very small, minute, or microscopic, massed together into low, dense, expanded colonies, individual filaments either visible as a fur or agglutinated into a solid disc or layer* 2
Thalli microscopic, solitary, filaments bearing isolated grape-like nodes; on Sporochnus; Stewart Island; summer; sublittoral Myriotrichia adriatica [10]

[Footnote] * Except in the case of Aglaozonia, which is of cellular structure.

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Thalli forming tiny, solid, glabrous or hairy balls 3
Thalli larger, constituting definite individual plants of various growth-forms, but without leaves or their equivalent, or air-vesicles 4
Thalli well-branched, usually bearing leaves and vesicles 11
2. Thalli forming (1) minute stain-like spots, (2) small, sucker-like discs, (3) crusts, (4) velvety patches of definite shape, (5) irregular mats, low and turf-like:
 (1) Spots isolated or massed, sometimes very indistinct:
Epiphytic
 (a) on Pachymenia himantophora; northern; May; LWST: ERB Mikrosyphar pachymeniae [1]
 (b) on Ulva, Hormosira, Ecklonia, Carpophyllum, Marginariella, etc.; common; summer and autumn Myrionema [2]
 (2) Epitithic
Circular discs rich dark-brown, flattish, solid, firm, carnose, about an inch across, radially corrugated, closely adhering to, and difficult to remove from, the substratum, slimy and inward-curling when removed; common; mostly summer; LWNT: EHR. Petrospongium berkeleyi [16]
 (3) Eplithic
When exposed and dry, blackish; when wet (a) mostly dark-coloured and brittle, (b) soft ochre-yellow and mushy, (c) reddish-brown and carnose:
 (a) Encrustations hard, woody or leathery, in brown, blackish or rusty circular expansions, rarely of irregular shape, often concentrically and radially marked or corrugated; common; H-LW: E-S Ralfsia [4]
 (b) Crusts very smooth, slippery, dull-coloured and matt, elliptical or circular, frequently coalescing, often of large size, extremely lubricous and mushy when scraped off substratum of clean, smooth rock; common locally; LWST: ERB Hapalospongidion saxigenum [3]
 (c) Irregular masses, usually not extensive, composed of scarcely distinguishable superimposed layers of small, flat, ribbon-like, somewhat forked, overlapping, prostrate fronds attached by basal rhizoids, resembling the decumbent juvenile thallus of Glossophora, but somewhat more cainose and redder in colour; often on colonies of sand-tube worm sheltered from sun by overhanging ledges; common locally; summer-autumn at least; LWST:ERF Aglaozonia [20]
 (4) Epiphytic
Velvety or spongy patches of distinctly filamentous structure, (a) circular or broadly-elliptical on surface of host, (b) small, narrow-elliptical, almost linear, on margins of host
 (a) (i) on Durvillea, (ii) on Xiphophora, (iii) on Hormosira:
 (i) Patches conspicuous, fundamentally elliptical but often coalescing, identical in size and shape on both surfaces of host frond (Durvillea antarctica); not on Durvillea willana; South Island; LWNT: ERF and ledges in surf and ER offshore Hapalospongidion durvilleae [3]
 (ii) Patches small, circular or oval, often coalescing, distinctly concentrically ruffled, velvety, with a visible pile, much darker when submerged than when dry; from Wellington south; most luxuriant from autumn to winter; LWNT: E ledges and P Herponema maculaeforme [5]
 (iii) Dark spongy masses capping or covering distended internodes of host; if northern, HWNT: P, winter; if southern, LWNT:HRB, February Herponerma pulvinatum [5]
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 (b) Primary tufts minute, coalescing and forming dense, putty-like sub-elliptical, linear lines along the margins of the upper parts of Carpophyllum maschalocarpum, less often on C. elongatum; North Island; most of year; LWST: S-ER . Sphacelaria pulvinata [11]
 (5) Mat-forming algae; mats of irregular shape and size, some almost reduced to close-set clusters of small, detached tufts, (a) minute, (b) under 1 inch high, (c) under 2 inches high:
 (a) Epiphytic
On Marginariella urvilleana, olive-green matt-surfaced fur covering portions of surface of blades; Stewart Island; June; sublittoral [6]
 (b) (i) soft, hairy, (ii) stiffish, somewhat bristly; with a large, dark, apical cell; epiphytic or epilithic:
Epiphytic
 (i) On most large algae, especially on Carpophyllum, Ecklonia, Lessonia, Marginariella, Durvillea, Desmarestia, Pachymenia, etc., smothering the decaying tips or blades of host; mostly summer; LWNT and below: S-EHRB Ectocarpus [7]
 (ii) On Xiphophora, Cystophora, Cladostephus, Halopteris hordacea, Pterocladia capillacea; forming patches on surface near tip of Xiphophora, otherwise smothering terete axes of other hosts; mostly southern; spasmodic, all seasons; LWNT:HRB: P Sphacelaria [11]
Epilithic
 (i) Soft, dark-olive, on upper surface of boulders, forming a distinctive turf-community; Little Barrier Island, Ti Titoki Flat; October; LWNT: HRB Ectocarpus dellowiana [7]
 (ii) Harsh, (x) with filaments erect, (xx) with filaments prostrate:
 (x) Olive turf at vertical base of low rock washed by sand; Ahipara and Houhora, Northland, Stewart Island; all seasons; LWNT: EHRB Sphacelaria tribuloides [11]
 (xx) Forming a dense, mud-accumulating, felt-like covering on mud-embedded rocks at bottom of a creek-covered by salt water at half tide; Stewart Island, Paterson's Inlet, Cunning Cove; best in winter; MT: SM Sphacelaria limicola [11]
 (c) Epilithic
 (i) Soft, Ectocarpus-like, (ii) stiff, flat, feathery
 (i) Only distinguishable from Ectocarpus under the microscope; dull-brownish, forming somewhat woolly, dense carpets accumulating sand, lining floor of very shallow hollows in rock, or fringing deeper pools in a continuous line at water-level; rarely epiphytic on Hormosira banksii var. sieberi; HWNT: P, sometimes LW on EHR Pylaiella novae-zealandiae [8]
 (ii) Thalli dark-olive, stiff, flat, feather-like, overlapping, accumulating sand, capping clean, low boulders; Pihama, Taranaki; all year, best in summer; LWST: EHB Halopteris platycena
3. Small, solid, globular or hemispherical balls or cushions,
 (1) smooth, firm; (2) hairy, soft.
Epiphytic
 (1) (a) on Gigartina lanceata; thallus resembling tiny peas, brown, glistening, kidney-shaped when removed; Stewart Island; summer; ERB sublittoral Leathesia novae-zelandiae [14]
 (b) On Cystophora; thallus resembling tiny beads or drops of “solution” (when fresh), brown,
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clustered; common locally; southern; summer; LWNT: EHRB and P Corynophlaea cystophorae [15]
 (2) on Xiphophora; thallus brown, pea-sized, distinctly hairy; throughout N.Z: summer; LWNT: EHRB and P Elachsta australis [13]
4. Thallus branched, forming tresses, sometimes bushy, without obvious, percurrent, axis 5
Thallus branched, more or less terete, with a percurrent axis 6
Thallus more or less plump, soft, and gelatinous, somewhat vermiform or filiform, solid, flaccid, lubricous, poorly or richly flexuously branched, often clothed with a visible peripheral coat of short, horizontal, hair-like filaments 7
Thallus obviously hollow, usually simple, of various forms 8
Thallus flat, membranous or cartilaginous, but not thick and leathery, or with a massive holdfast 9
Thallus flat, coarse, or thick and leathery, variously divided, usually with a massive holdfast 10
5. Tresses (1) soft and hairy, without a conspicuous apical cell, (2) somewhat wiry and bristly, with or without a conspicuous apical cell
 (1) Tresses of hair-like fineness, composed of free, branched, more or less monosiphonous, articulated filaments, not terminating in a conspicuous, dark-coloured apical cell, all unidentifiable to the naked eye, all both epiphytic and epilithic; (a) filaments monosiphonous throughout, (b) filaments mostly monosiphonous, with here and there a longitudinal wall, (c) filaments mostly polysiphonous:
 (a) Sporangia ovoid or cone-shaped, placed laterally on the branches, sessile or on short pedicels, rarely opposite. Mostly summer; LWNT: HRB and P Ectocarpus [7]
 (b) Sporangia intercalary, catenate, in short or long series; southern; mostly summer; LWNT: S-ERB Pylaiella littoralis [8]
 (c) Filaments at first monosiphonous, later polysiphonous; sporangia and branches opposite, frequently in paired series of either; south to antarctic, Lyttelton, Gladstone Pier piles, September; Campbell Island. Tucker Cove (intertidal), April: in N.Z. just below LWST Geminocarpus geminatus [9]
 (2) Epilithic
Tresses coarser (a) with a dark-coloured, broad apical cell, (b) with apical pencil (often denuded) consisting of a minute tuft of extremely fine hairs:
 (a) Harsh thallus (i) bristly-looking with short lateral whorls, (ii) shaggy with black axes, (iii) shaggy without black axes:
 (i) Thallus dichotomously branched, clothed with dense whorls of very short, bristly, incurled. pronged branchlets; southern, common; LWNT: EHRB Cladostephus
 (ii) Thallus bushy, tufted, coarse-haired through-out, rachides denuded when fertile, bearing small globular balls of incurling ramuli; southern; summer; LWNT: EHRB Ptilopogon botryocladus
 (iii) Thallus shaggy, finer or coarser, bushy, tufted, without black, wiry rachides, but lower axes clothed with foxy-brown stupa of descending rhizoids; common, one northern, others southern; throughout the year; LWNT to sublittoral; EHRB Haloptens
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 (b) Thallus horse-hair-like, very dark-brown or gingery, 18 inches high, sporangia clustered into minute, bullrush-like receptacles along the filaments; northern; all the year round; sublittoral: ERB Perithalia capillaris
6. Terete rachides (1) fine and flexuous, (2) coarser.
Epilithic
 (1) Thallus filiform, terete, or flattened in part, flexuous, more or less distichously branched along a percurrent axis which may be (a) wiry, or (b) flaccid:
 (a) Plants (1) delicate, branching alternate or pinnate, (ii) coarse, branching whorled.
 (i) Thallus pyramidal in outline, branches close-set, more or less opposite, all branches and receptacles terminating in a dense pencil of fragile hairs (vide Perithalia and Carpomitra), receptacles small, ellipsoid or spherical terminating lateral, generally short, branchlets; from Auckland south; mostly southern; summer; sublittoral: E Sporochnus
 (ii) Thallus branched on all sides, branches with whorls of branchlets; receptacles cylindrical to oval; Three Kings Island only Perisporochnus regalis
 (b) Limp thallus (i) dotted transversely with lines of minute sporangial sori, (ii) no lines of sporangia, plant turns green on exposure:
 (i) Thallus hollow throughout, sometimes distended with sand, branches long, lax, patent, in twos or threes, tapering to an apical cell capped by a long hair (frequently eroded), branches monosiphonous at tip, soon widening downwards with increasing longitudinal septation, and becoming cellular below; Portobello, Dunedin; Stewart Island: summer; LWST: SHB, somewhat muddy Striaria attenuata [12]
 (ii)Thallus solid, soft, oppositely pinnate, axis linear, branches terete, the ultimate capillary; subant-arctic; sublittoral; summer Desmarestia willii
 (2) Thallus shrubby, erect, stiffish, promiscuously branched, (a) epiphytic, (b) epilithic:
 (a) Epiphytic
On Hormosira, thallus small, several inches high, branches irregular, short, narrow-fusiform, patent, arising in the conceptacles of the axis, branchlets similar, shorter; conceptacles scattered over entire thallus; HWNT: P: LWNT; summer, autumn; common spotwise Notheia anomala
 (b) Epilithic
Thallus bushy, larger, coarser or finer, branches lax, more or less flattened, hollow in one sp., often clothed with flattened, tapering branchlets; sporangia embedded in cortical layer; MT-LWNT according to species; common, one sp. south; S-EHMB Scytothamnus
7. (1). With a fur of fine, free, horizontal, peripheral assimilatory filaments, (2) with peripheral filaments immersed in gelatine and visible only microscopically; epiphytic and epilithic
 (1) (a) epiphytic, (b) epilithic:
 (a) Epiphytic
On Sargassum spinultgerum; thallus small, up to 5 inches high, light-brown, almost filiform, slightly patently branched; very limp and difficult to distinguish on its host unless floated in. water, when fur also becomes visible; northern: Russell, Whanga-paraoa, Waiheke Island; summer; sublittoral on reefs Nemacystus novae-zelandiae
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 (b) Epilithic: (i) from MT-LWNT, (ii) sublittoral:
 (i) Thallus simple or nearly so, fat-cylindrical, rat-tail-like, very dark brown, up to 3 feet long, peripheral filaments conspicuous only in juvenile plants; common spotwise; best in spring and summer; MT-LWNT: HRB Myriogloia
 (ii) Thallus yellowish, brownish, greenish, slippery, up to 24 inches high, more or less abundantly and laxly branched, the long branches bearing short, erratically-divergent branchlets, axes narrow and dark-coloured, surrounded by a conspicuous fur of fine, free, extremely lubricous, yellow or greenish hairs; common spotwise; summer; sublittoral: EHRB Papenfussiella
 (2) (a) epiphytic, (b) epilithic:
 (a) Epiphytic
On Colpomenia (?) or Leathesia (?); thallus up to 4 inches high, axis comparatively long, bearing many short, simple branches; subantarctic; summer; LWNT: P; sublittoral Caepidium antarcticum
 (b) Epilithic: (i) on lowest littoral, (ii) sublittoral: (i) Thallus blackish, up to 8 inches high, firm, slippery, when emerged lying prone on surface, branches erratically patent or horizontal, apices frequently forked, sometimes curling; south; summer and autumn; LWNT: mudflats; sometimes epiphytic Chordaria dictyosiphon [18]
 (ii) Thallus brown, up to 18 inches high, fat-cylindrical, extremely lubricous, very laxly branched, apices obtuse; spotwise, both islands; summer; sublittoral: ERB Tinocladia novae-zelandiae [17]
8. Thallus (1) globular, (2) clavate, (3) cylindrical,
(4) moniliform:
 (1) Spherical, cerebriform, or tuber-like balls or sacs, exceptionally rupturing and forming an expanded net: (a) moderate-sized, somewhat squat, (b) very squat, (c) small, warted vesicles
 (a) Sac bullate and fully-distended when juvenile (tiny greenish or yellowish beads), later becoming somewhat squat and more or less convoluted, (i) remaining entire at all stages of growth, (ii) becoming fenestrate and expanded; epiphytic or epilithic:
Epiphytic on most brown algae:
 (i) Sac smooth to matt, somewhat papery and thin-skinned, greenish to yellowish, dull-brownish or greyish when mature; littoral P at all levels and sublittoral Colpomenia sinuosa [26]
Epilithic
 (ii) Sac smooth and entire, resembling Colpomenia until after spores are shed, then becoming fenestrate and breaking open into a net or lattice; subtropical; Kermadec Islands; P Hydroclathrus clathratus [27]
Epiphytic on Corallina:
 (b) Sac subglobular, very squat, much convoluted, substance thickish, fleshy, brittle, smooth, honey-yellow to rich-brown; common; mostly summer; LWNT: HR Leathesia difformis [14]
Epilithic
 (c) Sac very small, inflated, lobed or warted, “Colpomenia-like” with Leathesia-like structure, sac bearing a Chordaria-like erect thallus; requires further investigation; subantarctic, Campbell Island; summer; LWNT: P and sublittoral Caepidium antarcticum
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 (a) Thallus wide- or narrow-top-shaped, much distended with sea-water, ½–4 inches high, stipe short or longer, narrow, sometimes bearing an additional frond; mostly south; largest in summer; MT-LWST sometimes in HWP: HMSB Adenocystis utricularis [25]
 (b) Small, narrow, long-stiped; a very doubtful genus, possibly an alternate generation of A. utricularis; subantarctic; H-LWST and deeper: P Utriculidium durvillei
 (3) Thallus more or less long-tubular, narrow or broad, filled with seawater or gelose, (a) simple, (b) branched:
 (a) filled with sea-water:
 (i) Cylindrical or narrow-linear, (ii) intestiniform:
 (i) Thallus 4–16 inches high, constricted here and there, tapering downwards gradually into a fine stipe; common seasonally; August in the Far North, best in summer in the south; H-LW and beyond: P Scytosiphon lomentaria [21]
 (ii) Thallus under 7 inches high, light ochre-coloured, sac-like, baggy, sausage-shaped, tapering suddenly to a short stipe, thin-skinned; Stewart Island; summer and autumn (in drift) Asperococcus bullosus [24]
 (b) Filled with gelose:
Thallus under 12 inches high, greenish-yellow to dark-brown, axis erect, cylindrical, obtuse above, attenuated below, branches few, irregular, similar to main axis, sporangia in scattered conceptacles; common; best in summer; MT-LWST and below: EHRB Splachnidium rugosum
 (4) Thallus resembling dichotomously-branched chains of beads, internodes inflated and filled with sea-water, nodes stipe-like, conceptacles scattered over surface; common; all year; HWNT: P; normal from MT-LWNT Hormosira banksii
9. Thallus (1) simple, leaf-like, (2) ribbon- or strap-like throughout, variously divided, (3) cuneiform, palmate, flabellate, to suborbicular, more or less deeply incised:
 (1) Thallus small, consisting of a flat, expanded, leaf-like blade, narrow stipe, and minute, disc-like holdfast, (a) epiphytic, (b) epilithic:
 (a) Epiphytic
On Carpophyllum, Macrocystis, Marginariella urvilleana, drift logs, launch hull; rarely on rock or clay; thallus very variable in size and shape, from linear to broad-oblong, usually thin and delicate, light to dark-olive, turning green on exposure; south; best in summer; LWST and beyond: S-EHRB and P Punctaria latifolia [23]
 (b) Epilithic
Thallus smaller, resembling a ribbon-like Punctaria but more cartilaginous, dark-coloured, and not turning green on exposure; common locally; spasmodic all the year round; HW pools on elevated platform or in runnels and P to LW and beyond: ERB Ilea fascia [22]
 (2) Thallus wider or narrower, strap-like to linear, flat and flexible, substance membranous to cartilaginous, dichotomously or oppositely branched, (a) costate, (b) ecostate:
 (a) (i) dichotomously, (ii) oppositely branched:
 (i) (x) Kermadec Is. only, (xx) all over the N.Z. mainland and neighbouring islands:
 (x) Thallus about 6 inches high, linear, axils of dichotomies acute, apices attenuate, veins distinct, running obliquely out from the costa; subtropical, Kermadec Islands; sublittoral Dictyopteris kermadecensis [36]
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 (xx) Thallus up to 12 inches high, subdichotomous, texture firm and stiff, all branches and receptacles terminating in a microscopic tuft of delicate hairs, often denuded; receptacles mitriform, terminal; common; all year round; sublittoral: ERB Carpomitra costata
 (ii) Thallus up to 4 feet long, rich dark-brown, flat, smooth, highly glistening, membranous to cartilaginous, decompound, axis broad, strap-like, broadly costate, branches similar, distichous, opposite, but narrow and cylindrical at point of insertion, ultimate pinnae delicate, ciliate; plant turns verdigris-green and limp on exposure; south; sublittoral in channels and pools; ERB Desmarestia firma
 (b) Dichotomous throughout, all branches terminating in (i) a large, lenticular apical cell (not visible to naked eye), (ii) a comb-like row of small, branched hairs:
 (i) (x) N.Z. mainland species free from superficial folioles,* (xx) surfaces beset with many tongue-like processes:
 (x) Thallus smallish, ribbon-like to linear, membranous, sometimes intricately entangled; common locally; summer; LWNT and P: E-SHRB and in shallow pools in low, flat clay Dictyota [28]
 (xx) Thallus of moderate size, resembling Dictyota but more cartilaginous, and with longer dichotomies, both surfaces covered with small ligulate processes; common; best in summer; LWNT and P: EHRB and channels Glossophora kunthii [29]
 (ii) Thallus up to 12 inches high, dark-brown, narrow-linear, segments of dichotomies subcunei-fohn, tips terminating in numerous fine, short, thread-like, forked filaments giving the appearance of being shredded; south; summer; sublittoral and P: SHB Cutleria multifida [19]
 (3) Thallus complanate, membranous to cartilaginous or subcoriaceous, more or less wedge-shaped, or kidney-shaped, at least when juvenile, base stupose, undersurface smooth or covered wholly or in part with a soft, brown, woolly stupa, or with a fine, white, powdery bloom; growth-zone marked by a more or less conspicuous distal marginal series of dark-coloured peripheral initials; concentric lines of development present in all but one; thallus (a) wedge-shaped, (b) reniform-orbicular, (c) fanshaped in outline, but much falsely-branched and divided into long, narrow, subcostate laciniae with more or less flabellate tips.
Epilithic
 (a) Thallus (i) smallish, with pronounced lines of development, (ii) large, concentric lines wanting:
 (i) Thallus 6 inches high, hight olive-green, of delicate substance, deeply and unevenly incised, tips conspicuously truncate; subtropical, Kermadec Island; sublittoral Taonia australasica [31]
 (ii) Thallus up to 80 inches high, greenish-olive to dark-brown, texture firm or somewhat harsh, broadly cuneate at first then more or less deeply incised and becoming flabellate-palmate, the lacmiae somewhat subdichotomous, up to 2 inches wide, distally attenuated, with round apices, concentric lines wanting; from Whangarei Heads

[Footnote] * One Kermadec Island species has surface beset with small, curved folioles.

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southwards, largest specimens at Stewart Island; September to February; sublittoral: EHRB Spatoglossum chapmanii [37]
 (b) (i) Thallus small, substance firmer, stupa covering lower surface completely, (iii) thallus somewhat larger with inrolled terminal margin, under-surface lightly calcified, (iv) thallus scarcely larger, almost black, hard, stiff, stupa extending upwards somewhat, radially from base and in the axils of sinuses:
 (i) Thallus up to 2 inches high, olive-brown, delicately membranous, reniform when young, orbicular-flabellate later, irregularly cleft downwards, finely radially striate and concentrically lined; Russell, Brampton Roof, sublittoral; September Distromium skottsbergii [30]
 (ii) Thallus up to 2 inches high, dark-brown to almost black, firm and somewhat brittle, orbicular, flabellate, under-surface completely covered with a foxy velvety stupa, upper margin fimbricated with very narrow, even and short fimbriae; west coast of North Island, and southern; littoral plants in summer, deep-water in winter; LWNT: EHRB Microzonia velutina [34]
 (iii) Thallus up to 4 inches high, olive above, darker below, with a darker line at the inscrolled upper margin, substance delicate, stupose at base only, the whole under surface lightly powdered with a white bloom, upper surface zoned with up to 10 concentric bands of minute hairs; subtropical, Kermadec Islands; sublittoral and P Padina fraseri [32]
 (iv) Thallus up to 4 inches high, very dark brown, black when dry, stiff, subcoriaceous, broadly flabellate-orbicular, divided irregularly into lobes each crowned with a wide, broadly-rounded fan-like tip, stupa extending upwards radially from base; subtropical, Kermadec Island; sublittoral and littoral pools Pocockiella nigrescens [35]
 (c) Thallus up to 12 inches high, pale olive-green to rufous-brown, stipe stupose—in extreme cases long, cylindrical, woolly; thallus in juvenile specimens more or less orbicular, but soon dividing and splitting in one plane into many broader or narrower branched segments with pointed, forked, obtuse or cuneate tips, branches slightly to markedly falsely costate through ascending lines of stupa; common locally; all the year; LWNT and P to sublittoral: EHRB Zonaria angustata [33]
10. Thallus large to massive, with a flattened lamina without leaves, vesicles, or receptacles proper, blade variously divided, (a) flabellate-palmate, (b) pinnate, (c) dichotomous:
 (a) Thallus up to 40 feet long, attached by a large, solid, conical disc, stipe short or long, cylindrical, simple or prcliferated, blade thick, leather-like, broadly flabellate, divided more or less deeply into wider or narrower segments, round in section or flat; fertile frond bearing densely-scattered, minute conceptacles, rare in the north: on exposed islands and capes only; common in the south in swift currents and surge; perennial; LWNT and coastal rocks Durvillea
 (b) Thallus 1–6 feet high, attached by dichotomously branched hapteres, stipe extremely short to very long, solid or fistulose in part, blades more or less broad, usually long, margins pinnated with thinner, equally flat, strap-like or contorted segments arising originally at the transition-place be-
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tween blade and stipe, or (exceptionally) from any marginal tooth; fertile areas appearing as milky sori in longitudinal or oval patches on blade or segments; common locally; perennial; LWST: P: S-ER or vertical faces, in one case on a sandy substratum Ecklonia
 (c) Thallus (i) coarse, but not thick and leathery, terminating in somewhat thin-textured, inch-wide, paired blades, (ii) thallus thick and leathery, terminating in narrow, dichotomo-flabellate segments:
 (i) Thallus up to 6 feet high, attached by extremely tough, dichotomously-branched, woody, buttressed hapteres, branching dichotomous through repeated splitting of a primary laminarian frond, each splitting giving rise to a pair of stiped segments similar in appearance to those of Ecklonia; sori as in Ecklonia; common spotwise; perennial; usually sublittoral in deep, exposed situations with swiftly-moving water Lessonia variegata
 (ii) Thallus 1–3 feet high, attached by a small, but tressed holdfast, branching dichotomous and somewhat falsely pinnate, ultimate branchlets narrower, more soft and flexuous, and regularly dichotomous, forming the fertile areas (receptacles) of the plant; conceptacles arranged in 1-several rows in the fertile pinnules; common on many shores; LWNT: E ledges and forming the upper fringe of the coastal zonation Xiphophora chondrophylla
11. Thallus with flat or terete axis, which may be truncate, percurrent, or dichotomous. bearing branches, receptacles, and usually “leaves,” branching, unilateral, bilateral, or radial; vesicles (1) present, (2) wanting:
 (1) Vesicles present; axis (a) flat, (b) terete:
 (a) Axis flat, vesicles (i) unilateral, (ii) axillary, (iii) in place of ramuli of the last order:
 (i) Vesicles unilateral, marginal, on adaxial side of flat branches (“leaves” being absent) in series with subsimple marginal receptacles; south; perennial; LWST and below: E Marginariella
 (ii) Vesicles in axillary-branch-system (leaf, vesicle, and (when fertile) receptacular ramulus), branching bilateral from margins of axis, receptacles small, fasciculate; all sp. in the Far North, fewer in the south; all types of coast and harbours; LWNT and P to sublittoral; perennial Carpophyllum
 (iii) Vesicles rising from ramuli of the last order, branches not marginal (except in one), all others emerging from the flat faces of the rachides; receptacles representing modified ramuli, not specialized subsequent branchlets; common throughout N.Z. in open situations; mostly south; perennial; LWNT and P, and sublittoral: EHRB and shallows Cystophora
 (b) Axis terete, (i) first dichotomously, then unilaterally branched, (ii) percurrent-truncate, and radially branched:
 (i) Vesicles large, pear-shaped, unilaterally placed along the branches representing the inflated petioles of long, broad, lanceolate leaves, the whole leaf-system arising from the unilateral splitting of a large, sickle-shaped terminal leaf; sporangia in sori in specialized small leaves, sporophylls, at base of plant; south; perennial; deeply sublittoral on reefs along the coasts, and in harbours Macrocystis pyrifera
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 (ii) Vesicles in axillary-branch-system, as in Carpophyllum, main axis truncate, branches longer, leaves, both primary (basal) and secondary (upper) midribbed and dotted with cryptostomata; receptacles small, fascicled, in the axils of terminal branches; common locally; perennial; sublittoral and in deep lagoons: EHRB Sargassum
 (2) Vesicles absent, main axis percurrent, terete, with radial branching, leaves oak-like or small, laurel-like; receptacles smaller modified leaves dotted with conceptacles; oak-leaved species common, the other rare and at Chatham Island only; perennial; sublittoral: ERB Landsburgia

Basic Structural Details of Certain Genera

The following broad details of vegetative structure and reproduction are given as a means of verifying (where necessary) identifications resulting from the use of the key. Specimens determinable by their habit alone are excluded, but where confusion might occur, as between Ilea and Punctaria, or Distromium, Zonaria, and Microzonia, or juvenile Asperococcus and Adenocystis a description of the transverse section of the thallus is given.

1.

Mikrosyphar (sometimes included in Streblonema). Thallus wholly creeping, monosiphonous, articulate, branched; erect filiments wanting; sporangia scarcely differentiated from the normal cells, protruding.

2.

Myrionema, Thallus consisting of a single-layered basal disc of prostrate, radiating, monosiphonous, articulate, branched filaments, most articulations of which giving rise to short, erect, free, simple filaments, or sporangia.

3.

Hapalospongidion. Thallus consisting of a distromatic (scarcely distinguishable) horizontal basal layer, each articulation giving rise to a long, thin, simple, almost colourless monosiphonous, articulate, erect filament or sporangium, terminal on a short filament.

4.

Ralfsia. Thallus consisting of a rusty-coloured, thick or thin basal layer of horizontal radiating filaments from which obliquely ascend erect, somewhat forked, parallel filaments, closely compacted in gelatine; sporangia in scattered sori on upper surface, with free paraphyses.

5.

Herponema. Thallus consisting of a basal layer of dark-coloured, branched, radiating filaments from which sparsely-branched, densely packed, free, erect, monosiphonous, articulate filaments arise, bearing very numerous dark-coloured, ovoid unilocular sporangia, or large, ectocarpoid (siliquose or cone-shaped), plurilocular organs.

6.

Thallus consisting of a dense basal portion from which erect, parallel, sparsely-branched filaments ascend, the lower portion of axes closely agglutinated, chromatophores ringed bands (at least when dry), unilocular sporangia ovoid, plurilocular ectacarpoid, very erect and closely adpressed to filament.

7.

Ectocarpus. Thallus consisting of a basal disc, generally of creeping rhizoids, or penetrating filaments, and a much larger erect portion of branched, monosiphonous, articulate filaments without longitudinal septation, but occasional cortication on lower parts of upright axis; colourless hairs present in some species; reproduction by uni- or plurilocular organs, the former ovoid, the latter siliquose, conical, or cylindrical, borne laterally, rarely terminally, usually on short pedicels, on the branches.

8.

Pylaiella. Thallus consisting of a basal attachment of creeping rhizoids and an erect portion of branched monosiphonous, articulate filaments showing occasional longitudinal septation; hairs present in some species; reproductive organs, uni- and plurilocular, intercalary, catenate (taking the place of normal vegetative cells) in the filaments.

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9.

Geminocarpus. Thallus consisting of a rhizoidal holdfast and an erect Ectocarpus-like portion, monosiphonous fundamentally, but soon developing repeated longitudinal septation, branches and reproductive organs paired often on successive cells (catenate); hairs wanting; reproductive organs Ectocarpus-like, the plurilocular somewhat grotesque, the uppermost pair of a series longest, horn-like.

10.

Myriotrichia. Holdfast rhizoidal, erect filament long, bearing a few very short widely-spaced branchlets, the whole thallus articulate, monosiphonous, except at intervals where longitudinal septation occurs, giving rise to nodes where whorls of hairs and sporangia cluster.

11.

Sphacelaria. Holdfast rhizoidal or of penetrating filaments, erect filaments rather stiff, branched (except in one*, terminating in a large, black cell, followed below by a short monosiphonous part, gradually becoming polysiphonous downwards by 1–6 longitudinal walls, and cells either not divided transversely or 1–3 times; hairs present or wanting; reproductive organs borne as in Ectocarpus, unilocular ovoid, plurilocular cylindrical; vegetative reproduction in some by means of propagula or characteristic (often horned) buds.

12.

Striaria. Holdfast a small disc and rhizoids; erect thallus long, limp, hollow, branched distally, opposite or in whorls of 3, attenuating to both extremities, upwards to an apical cell capped by a long hair, terminal portion at first monosiphonous, articulated, then gradually becoming polysiphonous downwards by development of from 1-many longitudinal walls; transverse section fistulose, with, lining the cavity, a medulla of 1–2 rows of large, colourless, oval cells, followed by an assimilatory layer of small, rectangular, coloured cells; reproduction by unilocular sporangia only, spherical or ovoid, projecting, accompanied by hairs and unicellular paraphyses in small, punctate sori arranged in transverse bands across the thallus.

13.

Elachista. Structure: a hemispherical cushion arising from a basal monostromatic disc, cushion forming a solid, lubricous mass (medulla) of closely-agglutinated, erect, branched, radiating, colourless filaments, finally branching corymbosely and forming a pilose, peripheral zone (cortex) of free, short, coloured, simple, closely compact, claviform paraphyses, with them arising also numerous extremely long and broad, coloured assimilatorv filaments and unilocular and plurilocular reproductive organs; hairs wanting.

14.

Leathesia. Structure: a very large medulla of large, colourless, cylindrical filaments, ramifying, and becoming broader and shorter upwards, finally breaking up gradually into a subcortex of small, densely-packed spherical cells which bear, distally, a narrow peripheral level-topped layer (cortex) of (1) very short, simple, erect, clavate, coloured, assimilatory filaments (paraphyses) of 2–6 cells, the terminal large and globular; (2) narrow, long, colourless hairs; and (3) unilocular or plurilocular reproductive organs, or both together, all embedded in gelatine.

15.

Corynophlaea. Thallus in transverse section similar to that of Leathesia, but smaller, with a medulla of narrower cells, a wider cortex, and generally much longer, less clavate paraphyses of up to 20 or more cells.

16.

Petrospongium. Structure: a medulla of long-cylindrical, long-articulated filaments, giving off, below, long rhizoids similar to the filaments, and above, dividing dichotomously, the cells decreasing in length and width, becoming oval, the ultimate divisions forming a dark-coloured, corymbose, peripheral layer (cortex) of cylindrical (not claviform) filaments (paraphyses), at the bases of which hairs and cylindrical or difform unilocular sporangia arise on 1–5-celled pedicels, all embedded in gelatine.

[Footnote] * Sphacelaria pulvinata, besides lack of branching, has a lighter coloured apical cell than usual, and shows very little longitudinal septation.

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17.

Tinocladia. Structure: a medulla composed of a compact bundle of colourless filaments (axial threads) from which radiate outwards many branched laterals forming a transitional region (or subcortex) and terminating distally in a cortex composed of tufts of simple, coloured, curved paraphyses, hairs, and unilocular sporangia, all embedded in mucilage.

18.

Chordaria. Structure: a large medulla of firmly-agglutinated axial threads, no subcortex, cortex of (1) short, 2–5-celled, cylindro-clavate paraphyses of even height, with (2) unilocular sporangia interspersed, and (3) plurilocular sporangia metamorphosed from the top half of the paraphyses (divided into 4–6 loculi) and (5) colourless hairs, the whole embedded in gelatine.

19.

Cutleria. Structure: medullary layer of large, swollen, colourless cells, subcortical zone of smaller, isodiametric cells, and a cortex of a single row of small, coloured, photosynthetic cells; reproduction sexual, oogonia and antheridia borne on branched paranemata in superficial sori on separate individuals; hairs in tufts.

20.

Aglaozonia. Structure: cross section similar to that of Cutleria, of which it is the sporophytic generation, but lacking the layer of subcortical cells on the under surface of the crust from which rhizoids descend; unilocular sporangia in dense sori, without paraphyses, on upper surface.

21.

Scytosiphon. Structure: a medulla of large, colourless cells, becoming progressively smaller outwards, followed by a cortex of 1–2 rows of minute coloured photosynthetic cells; hairs present; plurilocular organs arranged in a palisade-like layer forming a continuous sorus interspersed with large, coloured, unicellular paraphyses.

22.

Ilea. Structure: similar to that of Scytosiphon; plurilocular reproductive organs, only, arranged in palisade-like tiers without unicellular paraphyses.

23.

Punctaria. Structure: a medulla of up to 7 layers of large, colourless, cuboidal cells, with a scarcely smaller-celled, coloured cortical layer of photosynthetic cells; hairs tufted; reproduction by uni- and plurilocular sporangia scattered over the surface singly or in punctiform sori, immersed or nearly so; paraphyses wanting.

24.

Asperococcus. Structure an inner stratum of large, colourless cells decreasing into a 1-layered, small-celled cortex; hairs in tufts; unilocular sporangia globose, projecting, with paraphyses in small groups; plurilocular sporangia conical, without paraphyses, scattered more or less singly, both types on same plant.

25.

Adenocystis. Structure: an internal larger-celled tissue of 3–5 layers of almost equal-sized colourless cells, with a cortical zone of 1–8 rows of very small superimposed cells and, when fertile, an external, compact layer of minute, short, erect, articulated paraphyses, forming the sorus, among which pyriform unilocular sporangia are borne; hairs tufted in cryptostomata.

26.

Colpomenia. Structure: a medulla of 5 rows of large, rounded, colourless cells, decreasing progressively in size towards a cortex of 1-several rows of small, cuboidal, coloured cells; plurilocular organs scattered in small sori developed around clusters of hairs within cryptostomata, and interspersed with unicellular paraphyses.

27.

Hydroclathrus. Exactly of the same structure as Colpomenia; fertile only before surface ruptures and becomes fenestrate.

28.

Dictyota. Structure: a monostromatic medulla of very large, colourless, rectangular cells, and an assimilatory layer on each surface of minute rectangular, photosynthetic cells.

29.

Glossophora. Structure: a monostromatic medulla of large, colourless, rectangular cells, followed by 1-several rows of smaller subcortical cells, and an assimilatory layer of minute, rectangular, photosynthetic cells; both surfaces covered with small, scattered ligulae (lingulae).

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30.

Distromium. Structure distromatic: a double layer of large, equal, rectangular cells regularly arranged in transverse and longitudinal rows; small cortical, photosynthetic cells wanting.

31.

Taonia. Structure: 4 rows of large, rectangular, colourless, medullary cells, and an assimilatory layer of smaller, rectangular, coloured photosynthetic cells.

32.

Padina. Structure: 4 rows of large, rectangular medullary cells bounded on each side by an assimilatory layer of scarcely smaller photosynthetic cells.

33.

Zonaria. Structure: 6–10 superimposed rows of large, equal, rectangular medullary cells arranged in regular transverse and longitudinal rows, bounded on each surface by an assimilatory layer of small, rectangular, twinned, photo-synthetic cells.

34.

Microzonia. Structure: 2–8 superimposed rows of large, equal, rectangular medullary cells, bounded by an assimilatory layer of small, rectangular, twinned, photosynthetic cells.

35.

Pocockiella. Structure: a 1-layered inner medulla (sometimes wanting) of extremely large, colourless, rectangular cells, followed by 2–6 superimposed rows of smaller, equal, colourless, rectangular cells, and an assimilatory layer of small, rectangular, twinned assimilatory cells, all in strict transverse and longitudinal rows.

36.

Dictyopteris. Structure: 4–10 rows of large, rectangular, colourless cells at the midrib, diminishing in number to 2–3 rows at the margins, and an assimilatory layer of small, cuboidal, photosynthetic cells bounding each surface.

37.

Spatoglossum. Structure: 2–3 rows of large, somewhat rhomboidal, colourless medullary cells, 1–2 rows of somewhat irregular subcortical cells, usually more rows on one surface than the other, and an assimilatory layer of small photo-synthetic cells.

I would like to express my thanks to Prof. V. J. Chapman for his help in revision and correction of the proofs.

Victor W. Lindauer,
Russell

,
Bay of Islands.