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Austral Hepaticae. 32. A Revision of th

Lepidozia (Hepaticae) for New Zealand

John J. Eitgel

Rudolf M. Schuster

Angus! 31. 2001

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FIELDIANA

HISTORY

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Botany **n

NEW SERIES, NO. 42

Austral Hepaticae. 32. A Revision of the

Genus Lepidozia (Hepaticae) for New Zealand

John J. Engel

Department of Botany

Field Museum of Natural History

1400 South Lake Shore Drive

Chicago, Illinois 60605-2496, U.S.A.

Rudolf M. Schuster

Cryptogamic Laboratory

Hadley, Massachusetts, U.S.A.

Research Associate

Field Museum of Natural History

1400 South Lake Shore Drive

Chicago, Illinois 60605-2496, U.S.A.

Accepted July 18, 2000

Published August 31, 2001

Publication 1513

PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY

© 2001 Field Museum of Natural History

ISSN 0015-0746

PRINTED IN THE UNITED STATES OF AMERICA

Table of Contents

ABSTRACT 1

INTRODUCTION 1

LEPIDOZIA (DuM.) DUM 4

Affinities and Intrageneric Classification

Taxonomic Characters and Morphology .

Conspectus of Subgenera of Lepidozia . .

Conspectus of New Zealand Lepidozia

Taxa

Key to New Zealand Species of Lepido-

zia

5

14

18

19

20

ACKNOWLEDGMENTS 104

LITERATURE CITED 104

INDEX TO TAXA . 106

List of Figures

1. Telaranea remotifolia Hodgs 6

2. Telaranea mooreana (Steph.) Schust 7

3. Telaranea centipes (Tayl.) Schust 9

4. Telaranea centipes (Tayl.) Schust 10

5. Telaranea tuberifera Engel & Schust. ... 11

6. Lepidozia pendulina (Hook.) Lindenb.,

L. concinna Col., and L. microphylla

(Hook.) Lindenb 12

7. Lepidozia spinosissima (Hook. f. &

Tayl.) G. L. & N 24

8. Lepidozia spinosissima (Hook. f. &

Tayl.) G. L. & N 25

9. Lepidozia microphylla (Hook.) Lin-

denb 29

10. Lepidozia pendulina (Hook.) Lindenb.

33

11. Lepidozia setigera Steph 37

12. Lepidozia setigera Steph 38

13. Lepidozia procera Mitt 42

14. Lepidozia obtusiloba Steph 44

15. Lepidozia obtusiloba Steph 46

16. Lepidozia obtusiloba Steph 47

17. Lepidozia ornata Engel 50

18. Lepidozia concinna Col 54

19. Lepidozia concinna Col 55

20. Lepidozia laevifolia (Hook. f. & Tayl.)

G. L. & N 58

21. Lepidozia laevifolia (Hook. f. & Tayl.)

G. L. &N 60

22. Lepidozia laevifolia var. alpina Schust.

& Engel 63

23. Lepidozia fugax Engel 64

24. Lepidozia novae -zelandiae Steph 68

25. Lepidozia novae-zelandiae var. heteros-

tipa Schust 69

26. Lepidozia novae-zelandiae var. minima

Schust., L. spp 72

27. Lepidozia acantha Engel 73

28. Lepidozia elobata Schust 75

29. Lepidozia pumila Engel 77

30. Lepidozia bidens Engel 80

31. Lepidozia kirkii Steph 83

32. Lepidozia hirta Steph 86

33. Lepidozia ulothrix (Schwagr.) Lindenb.

89

34. Lepidozia ulothrix (Schwagr.) Lindenb.

90

35. Lepidozia glaucophylla (Hook. f. &

Tayl.) G. L. & N 93

36. Lepidozia bisbifida Steph 96

37. Lepidozia digitata Herz 99

38. Lepidozia digitata Herz 100

39. Lepidozia glaucescens Engel 102

111

Austral Hepaticae. 32. A Revision of the Genus

Lepidozia (Hepaticae) for New Zealand

John J. Engel and Rudolf M. Schuster

Abstract

Twenty-two species of Lepidozia are recog-

nized in New Zealand, and these are placed in five

subgenera. The following new taxa are proposed:

Lepidozia subg. Notholepidozia Schust., L. acan-

tha Engel, L. bidens Engel, L. elobata Schust., L.

fugax Engel, L. glaucescens Engel, L. laevifolia

(Hook. f. & Tayl.) G. L. & N. var. acutiloba En-

gel, L. laevifolia var. alpina Schust. & Engel, L.

novae-zelandiae Steph. var. heterostipa Schust.,

L. novae-zelandiae var. minima Schust., L. obtu-

siloba Steph. var. parvula Engel, L. ornata Engel

and L. pumila Engel.

A key to the Lepidozia species in New Zealand

is included.

Introduction

The genus Lepidozia was proposed by Dumor-

tier in 1835 (p. 19), based on an earlier taxon,

Pleuroschisma sect. Lepidozia (Dumortier, 1831,

p. 69). Basically, the latter concept included taxa

now placed into Bazzania and allies (the Pleuros-

chisma element) and a series of eight taxa that

constitute Lepidozia. In subsequent years a wide

range of other taxa were assigned to Lepidozia but

failed to fit into the genus as we here define it.

Spruce (1885, p. 358-360) divided the genus into

two subgenera, Eu-Lepidozia ( = Lepidozia s. str.)

and Micro-Lepidozia. The latter was divided into

two unnamed groups with four species, of which

the first three constitute Microlepidozia s. str., to-

day placed within Kurzia. v. Mart. The fourth spe-

cies (L. chaetophylla Spruce) represents the au-

tonomous genus Telaranea.

In the 19th century and into the early years of

the 20th century, various taxa, such as the south-

ern South American L. oligophylla and the tem-

perate Australasian L. capilligera (cf. Gottsche et

al., 1845, pp. 201, 204), were included under

"Lepidozia." These taxa were segregated from

Lepidozia by Fulford and Taylor (1959) as Neo-

lepidozia, with Lepidozia (Jungermannia) capil-

ligera as the type. Taxa falling into this last ele-

ment were regarded (Schuster, 1963, 1969) as

constituting a subgenus of Telaranea. Finally, an

isolated element, Lepidozia integristipula Steph.

(Stephani, 1922, p. 331), was segregated by

Hodgson (1962b) and placed into the autonomous

genus Drucella. Thus, at various times, taxa that

today fit into Bazzania, Telaranea (incl. Neolepi-

dozia), Kurzia (incl. as a subgenus of Microlepi-

dozia), and Drucella were assigned to Lepidozia.

This confusion continued into the relatively re-

cent account of the New Zealand species of the

genus Lepidozia by Hodgson (1956). To date, this

is the only revision of the New Zealand taxa.

Hodgson assigned some 33 species to the genus,

but of these 33 species, 22 presently belong to

other genera (Table 1).

Thus, in the only prior revision of the New Zea-

land taxa of Lepidozia, the 33 species assigned to

that genus belong in some five genera. Hodgson

(1964, p. 70) assigned Telaranea herzogii to a

sixth genus, Arachniopsis, which does not occur

in New Zealand. Hodgson (1962) also segregated

Lepidozia glaucophylla, which she had assigned

in 1956 to Lepidozia s. str. (her subg. Lepidozia),

to a new genus, Lepidoziopsis; this was trans-

ferred (Schuster, 1973) to Lepidozia subg. Glau-

colepidozia.

From this brief account, it is evident that the

New Zealand taxa of Lepidozia have been widely

confused with other genera. Rather than 1 1 spe-

FIELDIANA: BOTANY, N.S., NO. 42, AUGUST 31, 2001, PP. 1-107

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FIELDIANA: BOTANY

cies of "true" Lepidozia, as in Hodgson (1956),

we now recognize some 22 species, of which sev-

en are newly described here. The genus s. str. is

thus a large and complex group in New Zealand,

and a definition of Lepidozia s. str. emerges from

the following conspectus. In the past, Lepidozia

was confused chiefly with Telaranea subg. Neo-

lepidozia; this problem is addressed under Affin-

ities and Intrageneric Classification (p. 5).

After exclusion of the foreign elements, the ge-

nus remains forbiddingly complex, even after the

exclusion of the more isolated elements. Taxo-

nomic problems in the genus s. str. are discussed

on pp. 5-8.

Conspectus of Generic and Subgeneric

Elements Previously Confused with Lepidozia

1. Vegetative branches all or mostly Frullania

type, lateral-terminal and/or ventral-intercalary

(exc. Pseudocephalozia); stem with several to

many cortical cells intervening between suc-

cessive lateral leaves; oil-bodies present (ves-

tigial or lacking from most cells in some Kur-

zia spp.) 2

2. Leaves clearly incubous to transversely in-

serted; underleaves never present (vestigial

in some Telaranea spp.); terminal branches

abundant and lateral-intercalary leafy

branches usually lacking 3

3. Terminal branches of the Microlepidozia

type (replacing dorsal half of lateral leaf)

lacking, the lateral branches all Frullan-

ia type (replacing ventral half of a lateral

leaf); oil-bodies present in all leaf cells,

conspicuous, granular to botryoidal (exc.

L. reptans); leaves ± incubously insert-

ed; plants usually whitish or light to

clear green, rarely brownish 4

4. Stem without a conspicuous hyalod-

erm, the cortex firm- to thick-walled,

the cells not conspicuously enlarged

in diameter, mostly in numerous

(>24) rows; leaf cells almost always

firm, mostly thick-walled, usually rel-

atively small in proportion to leaf,

never tiered within the lamina; spores

(as far as known) with granular exine;

$ bracts usually shallowly lobed or

lobulate . . . Lepidozia (Dum.) Dum.

4. Stem with a conspicuous hyaloderm

(exc. Telaranea subg. Tricholepido-

zia), formed of very large, soft-tex-

tured, leptodermous cells, the cortex

of a limited number of cells (except

Telaranea subg. Tricholepidozia),

usually in 6-24 rows; leaf cells thin-

walled, usually ± pellucid, large in

proportion to leaf, ± tiered within

lamina (or lamina lacking); spores

delicately areolate; $ bracts very

deeply (mostly 4-) lobed

Telaranea Spruce ex Schiffn.

3. Terminal branches of both the Microle-

pidozia and Frullania types; oil-bodies

of some or most leaf cells lacking or few

(1-2, rarely 3-5) and homogeneous to

few-segmented, less often granular;

leaves usually transversely inserted

(very deeply 3-4(5-6)-lobed); plants

usually ± fuscous (exc. Leucolepidozia,

which is nitid and whitish green). Stem

with ± firm-walled cortical cells, usually

in (8)9-14 rows (exc. subg. Dendrole-

pidozia) Kurzia v. Mart.

2. Leaves somewhat to clearly succubously in-

serted, only the dorsal end of the insertion

line transverse; underleaves vestigial, a few

isodiametric basal cells giving rise to 2-4

lobes formed of 1(2) fingerlike cells, each

ending in a slime papilla; terminal branches

rare or lacking in most phases, but with lat-

eral-intercalary branches frequent. Leaves

soft-textured and mostly 4-lobed

Pseudocephalozia Schust.1

1. Vegetative leafy branches thecal, lateral-inter-

calary (from axils of the incubous, asymmetri-

cally 3-lobed lateral leaves); stem with 1-2

rows of strongly elongated, sublinear cortical

cells intervening between successive lateral

leaves; oil-bodies lacking (numerous lipid drop-

lets may be present) Drucella Hodgs.2

[D. integristipula (Steph.) Hodgs.]

Of the five genera in the preceding conspectus,

genera 1-3 belong in the subfamily Lepidozioi-

deae (sensu Schuster, 1973, 1984), while genus 4,

Pseudocephalozia, is an enigmatic genus that

probably falls close to the point of origin of the

Zoopsidoideae, a subfamily unusual in the Lepi-

1 Lepidozia leptodictyon Herz. (Trans. Roy. Soc. New

Zealand 68: 45. 1938) is placed as a synonym of Pseu-

docephalozia lepidozioides Schust. by Engel and Schus-

ter (1988). Pseudocephalozia lepidozioides is treated in

detail in Schuster and Engel (1974, p. 677, figs. 5-7).

2 For this genus, see Schuster (1980, p. 411, figs. 17-

18) and Boesen (1982).

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

doziaceae in developing succubous to longitudi-

nally inserted leaves, although possibly also bear-

ing "distant affinities to the Lembidoideae

Schust." (cf. Schuster & Engel, 1974). The last

genus, Drucella, is so isolated that it is split from

other Lepidozioideae by Boesen (1982) as a tribe

Drucelleae Boesen, elevated by Schuster (1984)

to the status of an autonomous subfamily, the

Drucelloideae (Boesen) Schust. After removal of

these two last genera, a complex remains formed

by Lepidozia, Telaranea, and Kurzia. These gen-

era are clearly allied, but Kurzia seems to us to

bear relatively remote affinities to the other two

genera. It is distinct in (a) retaining the capacity

to form Microlepidozia-type terminal branches;

(b) a tendency for most taxa to lose oil-bodies

from all or most leaf cells; (c) the transverse

leaves, ranging in some taxa to clearly succu-

bously inserted and oriented.

Lepidozia and Telaranea, however, are so

closely allied that until relatively recently they

were considered identical, or at least the Neole-

pidozia element was placed as a group "Sym-

metricae" in Lepidozia, while the subg. Telaranea

element was recogized as an autonomous genus.

The present segregation into two genera, with the

"Symmetricae" element placed within Telaranea,

was introduced in Schuster (1963).

Lepidozia (Dum.) Dum.

Pleuroschisma sect. Lepidozia Dum., Syll. Jungerm.

Eur. Indig. 69. 1831.

Lepidozia (Dum.) Dum., Receuil Observ. Jungerm.

19. 1835, nom. cons.

Herpetiwn sect. Lepidozia Nees, Naturg. Eur. Leberm.

3: 31. 1838.

Lepidoziopsis Hodgs., Rec. Domin. Mus. 4: 105.

1962.

Plants anisophyllous, creeping to loosely pros-

trate to, more often, somewhat ascending, rarely

stiffly erect, the leafy shoots mostly loosely at-

tached to substrate, firm, yellowish or whitish to

olive-green, rarely brownish in life, rarely glau-

cous, usually relatively opaque and chlorophyl-

lose, vigorous to medium-sized, usually 0.5-

2.5(4) mm wide. Branching irregular to regular,

copiously 1- to 2- or 2-3-pinnate, of Frullania

type; Acromastigum type present in Dendrolepi-

dozia and one species of sect. Notholepidozia; Mi-

crolepidozia-type branches and lateral-intercalary

branches lacking; ventral-intercalary microphyl-

lous or leafy branches in some taxa; lower lateral

terminal branches partly or mostly attenuate at the

apices, often becoming rhizoid-bearing; half-leaf

associated with Frullania-lype branches usually

bifid; initial branch underleaf of primary branches

usually bifid, less often unlobed or trifid to quad-

rifid. Stem firm to rigid, terete to elliptical in cross

section, with (18)20-32 or more rows of firm-

walled cortical cells, typically little or at most

moderately larger in diameter than those of ad-

joining medulla, not forming a discrete hyalo-

derm;3 subepidermal layers occasionally present,

composed of thick-walled cells; medullary cells

equal to or slightly smaller than cortical, firm-

walled (the outer sometimes strongly so). Rhi-

zoids of normal-leaved axes usually rare or ab-

sent, except at shoot bases, arising from underleaf

bases; rhizoids on flagelliform shoots or shoot

apices from all three merophyte rows. Leaves al-

most without exception 4-lobed or 4-lobulate on

main axes, incubous to subtransverse, the inser-

tion extending to stem midline dorsally, the leaves

typically asymmetrical, the dorsal half (and dorsal

1-2 lobes) larger than the ventral, the lobes usu-

ally many-celled (at bases 3-4 or more cells broad

in most taxa), the cells similar to those of lamina,

the lobes usually sharp, often deflexed or inflexed

in most taxa; lamina margins entire or toothed to

spinose-dentate or ciliate. Cells not tiered within

the lamina, firm-walled, the lumina often guttulate

or rounded, the trigones concave-sided and mostly

obscurely defined; median cells mostly 18-25(40)

//.m wide in median portions of lamina, cuticle

smooth or clearly papillose. Oil-bodies present in

all leaf cells, colorless, usually (2)5-16 per cell,

relatively large, usually granular-botryoidal, less

often subhomogeneous. No asexual reproduction4

or (L. fugax, occas. in L. digitata) reproduction

via fragmenting or caducous leaf lobes. Under-

leaves usually 0.5 or less the lateral leaves in area,

transversely inserted, symmetrically 4-lobed on

main axes.

Dio- or rarely (L. reptans) autoecious. Game-

tangial branches all short (<? rarely ± elongated,

leafy), devoid of normal leaves and underleaves,

ventral-intercalary in origin (the androecia less

3 Some taxa, such as L. microphylla, have two or more

strata of smaller cells lying within a rather distinct hy-

aloderm (Fig. 9 : 14). Even though stem anatomy, in

general, serves to separate Lepidozia from Telaranea,

the distinction is bridged by several exceptions.

4Degenkolbe (1938, p. 87) reported gemmae in the

generic type, L. reptans (L.) Dum. Over four decades of

study of masses of material of this species have not re-

vealed any gemmiparous population of this or any other

true Lepidozia species.

FIELDIANA: BOTANY

frequently terminal on primary and secondary

branches). Androecia typically spicate, smaller

than vegetative shoots, the bracts concave, ± im-

bricate, lobed like (but usually more shallowly

than) vegetative leaves; antheridia 1-2 per bract,

the stalk 2-seriate; bracteoles small, lacking an-

theridia. 9 Branches with several progressively

larger gyres of identical bracts + bracteoles, the

inner 1-2 gyres of bracts + bracteoles erect, con-

cave, clearly sheathing the perianth, the innermost

bracts mostly oval or ellipsoidal in outline, 4(6)-

lobed to lobulate at the apex, but occasionally

bracts 2-4 dentate-lobulate. Perianth fusiform or

ellipsoidal-fusiform, bluntly 3-gonous in at least

the narrowed distal sectors, the mouth usually

crenulate to crenate-denticulate.

Seta with (8)12-16 large epidermal + 16 or

more rows of much smaller internal cells. Capsule

ellipsoidal to oblong, the wall 3-5(6)-layered (in

studied taxa); epidermal cells with ± oblong cells,

often imperfectly tiered, the primary longitudinal

and all transverse walls normally lacking pig-

mented thickenings, the secondary (longitudinal)

walls with conspicuous radial (nodular) thicken-

ings, often ± coalescent; inner cells narrower, nar-

rowly oblong, with radial (nodular) thickenings,

± connected by weak, often vestigial, tangential

bands. Spores papillose-vermiculate, less than

1.5X the diameter of the 2-spiral elaters.

TYPE — Lepidozia reptans (L.) Dum. (The type

species is unfortunately not representative of the

genus; cf. remarks under subgenus Notholepido-

zia, p. 31.)

Lepidozia is a large, taxonomically complex

genus that is found in almost all areas except the

Arctic and true Antarctic regions, arid continental

regions, and some isolated lowland islands. Al-

though Stephani (1922) included some 297 taxa

in the genus, and a few have been described sub-

sequently, many of the taxa he referred here have

now been assigned to other genera (Telaranea,

incl. Neolepidozia; Kurzia, incl. Microlepidozia;

Drucelld). Even after exclusion of these elements,

the genus remains large and unwieldy, although

many described taxa are synonyms. On that ac-

count, the genus was divided into five subgenera

in Schuster (1973, pp. 385-388); two other sub-

genera, Notholepidozia Schust. and Austrolepi-

dozia Schust., are accepted here. Of these six sub-

genera, only Chaetolepidozia Schust. and Clado-

lepidozia Schust. do not occur regionally.

The center of diversity of the genus is the re-

gion of former Gondwanaland: present-day South

America (with 30 species recognized by Fulford,

1966), New Zealand (we recognize 22 species),

ranging to New Caledonia, New Guinea, and the

Indomalayan region. Africa is much poorer in

species.5 By contrast, Holarctic regions bear only

an impoverished representation of species: All of

eastern North America has a single species (L.

reptans), and Europe has only three species (L.

cupressina (Sw.) Lindenb., L. pearsonii Spruce,

and L. reptans). The holarctic taxa belong to the

single subgenus Lepidozia. By contrast, in New

Zealand we find representatives of five of the sev-

en subgenera, while the other two subgenera oc-

cur in New Guinea (largely Gondwanalandic in

origin) westward into Indomalaya.

Affinities and Intrageneric Classification

As the conspectus clearly suggests, Lepidozia

is most immediately allied to Telaranea. In ad-

dition to the critera in the conspectus, a series of

additional features and, more important, a suite of

phylogenetically relevant "tendencies" separate

the two genera. A discussion of these follows.

1 . Leaf lobe number is relatively stable in Lep-

idozia, with main axes usually with 4-lobed

leaves, although sporadic 3- and 5-lobed leaves

occasionally occur in some taxa; in Telaranea,

rampant variation from 2- to 3-4- to 5-6- or even

8-13-lobed leaves occurs (Figs. 1 : 1-6; 2 : 1-3;

3 : 1, 2; 5 : 6, 7).

2. The leaf tends to be conspicuously asym-

metrical in Lepidozia, with a longer dorsal lobe

(and margin) than ventral lobe (and margin; Figs.

9:3; 10 : 3, 4), while in Telaranea the leaves are

typically symmetrical or almost so (Fig. 5 : 5-7).

However, obvious exceptions occur; thus, in T.

inaequalis Schust., the bifid leaves have a con-

spicuously shorter ventral lobe (cf. Schuster,

1969, fig. 88 : 5), and in the very different T.

mooreana (Steph.) Schust., the dorsal portion of

the disc tends to be higher than the ventral (Fig.

2 : 3); the same is true of such typical taxa of

Telaranea subg. Neolepidozia as T. remotifolia

(Hodgs.) Hodgs., in which the leaf shape is de-

cidedly Lepidozia-\ike (cf. Fig. 1 : 1-6).

5 However, the closely allied genus Sprucella Steph.,

with two endemic species, occurs in tropical Africa. This

is so closely allied to Lepidozia that some workers in-

clude the two species within Lepidozia. It differs from

Lepidozia s. str. in the 2-3-lobed lateral leaves, strongly

reduced underleaves, and the leaf areolation (cf. Schus-

ter, 2000).

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

FIG. 1 . Telaranea remotifolia Hodgs. 1-6. Leaves, cellular detail. 7. Dorsal lobe of leaf. 8. Main shoot (fbu —

first branch underleaf), medullary strand shown with stipple; ventral view. 9. Underleaves, cellular detail. 10. Stem,

cross section. 11. Antheridial stalk. 12, 13. First branch underleaves (both at same scale). (Figs. 1, 5-6, 10, 12, 13

from Engel 18921, New Zealand, South Is., Otago Prov., Mt. Aspiring Natl. Park, Blue River; 2-4, 7-9 from type;

1 1 from Child 5366, New Zealand, South Is., Westland Prov., 5 km N of Runanga.)

FIELDIANA: BOTANY

FIG. 2. Telaranea mooreana (Steph.) Schust. 1. Sector of main shoot, dorsal view (FB = Frullania-type branch;

HL = half-leaf). 2, 3. Leaves. 4-7. Leaf lobes. 8. Terminal cells of leaf lobes. 9. Sector of main shoot, ventral view

(FB = Frullania-type branch; FBU = first branch underleaf)- 10. Apex of innermost 9 bract, with (at arrows) 2

lobules shown in detail. 11. Stem, cross section. (Figs. 1, 6, 9 from Engel 21771, New Zealand, South Is., Westland

Prov., ca. 4 km N of Haast River; 2, 7 from type of Lepidozia pulcherrima; 3, 4, 8 from type of Lepidozia mooreana;

10, 1 1 from Engel 16407, Tasmania, road between Zeehan and Trial Harbour; 5 from Engel 79927, Tasmania, Tahune

Forest Reserve, along Huon River.)

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA 1

3. The stem in Lepidozia generally lacks ob-

vious differentiation of a hyaloderm (Figs. 8:9;

9 : 14), although cortical cells in some taxa, in-

cluding the generic type, tend to be somewhat

larger than the medullary (cf. fig. 85 : 8 in Schus-

ter, I.e., and Fig. 6:13 in this paper). In many

taxa the cortical cells and/or several strata of in-

tracortical cells may be strikingly thick-walled, as

in L. spinosissima and L. microphylla (Figs. 8 : 7,

9; 9 : 14), and stems are quite rigid. In general,

cortical cells are at least slightly firm-walled.6 In

Telaranea, in contrast, at least dorsal and dorso-

lateral cortical cells are much larger than medul-

lary cells (Figs. 1 : 10; 5 : 4), even though this is

hardly distinct in subg. Tricholepidozia (Fig. 2 :

11 of T. mooreana). In Telaranea there is never

a clearly defined zone of thick-walled cells lying

within the hyaloderm (Figs. 2 : 1 1 ; 5 : 4), whereas

such a region of thick-walled cells normally is

discernible in Lepidozia (Figs. 6 : 13; 8 : 9; 9 : 14;

10: 15).

4. Leaves in Lepidozia are usually much less

deeply divided (usually for less than 0.55 their

length) than in Telaranea, although some taxa of

the latter (e.g., T. centipes; cf. Figs. 3 : 1, 7; 4 :

2) may have less deeply lobed leaves. Even in this

last species, lobes are only inserted on 2 basal

cells and are uniseriate distally, a tendency ex-

pressed more fully in T. nematodes and T. moo-

reana (cf. fig. 90 : 10-12 in Schuster, I.e., and Fig.

2 : 2, 3, this paper). By contrast, in Lepidozia, ex-

cept in the highly derived subg. Cladolepidozia,

leaf lobes tend to be much broader, typically tri-

angular, and fail to end in ciliiform apices, and

they are never ciliiform throughout; this is true of

the generitype, L. reptans (cf. fig. 86 : 4 in Schus-

ter, I.e.).

5. Leaf cells in Lepidozia are almost without

exception perceptibly firm-walled and, in the lam-

ina and usually into the lobes, are irregularly ori-

ented, irregularly polygonal, and never consis-

tently elongated (Figs. 8:8; 12 : 1; 13 : 9); this is

true even of weaker taxa such as the type, L. rep-

tans (cf. fig. 86 : 4, 6 in Schuster, I.e.). In Telar-

anea, leaf cells are usually consistently thin-

walled and in the lamina and/or lobe bases tend

to be oriented in longitudinal lines and are mostly

perceptibly tiered, as, for example, in T. centipes

(Figs. 3 : 2, 3, 7; 4 : 2) and T. tuberifera (Fig. 5 :

5,7).

The cells in Lepidozia are not only usually

somewhat firm-walled (L. reptans, the generic

type may have them minimally incrassate; cf. fig.

86 : 6 in Schuster, I.e.), but they average much

smaller than in Telaranea species and seem more

chlorophyllose. In Telaranea the relatively large

and thin-walled cells seem more subhyaline in

most of the taxa. As a consequence, Lepidozia

species are usually firmer than those of Telaranea.

The cells in the lamina of Telaranea are often

more rectangulate (Fig. 2 : 3) and are often, if im-

perfect, tiered; when not clearly tiered they are

often clearly 5-6-angled (Figs. 1 : 1-5; 3 : 1; 4 :

2). They are always clearly angulate. Those of

Lepidozia are much less rectangular or angulate,

are only sporadically elongated, and are never

clearly tiered (Figs. 6 : 1-3, 5-6, 10; 12:1). In

practice, the cell differences between the two gen-

era are readily perceived.

6. The 9 bracts in Lepidozia, with very isolat-

ed exceptions, have a very high lamina (Figs. 7 :

5; 15 : 2), often longer than broad, and have re-

duced lobes; often the bracts are merely lobulate

at their apices, as in the type, L. reptans (fig. 87 :

6-7 in Schuster, I.e.). In Telaranea the bracts,

like the leaves, tend to be deeply lobed and some-

times bear accessory lobes and/or cilia or lacinia,

as in T. nematodes (cf. fig. 89 : 9 in Schuster,

I.e.).7

7. Spores in Lepidozia species always appear

to be papillose or bear irregularly coalescent pa-

pillae, forming short, vermiculate markings (as in

Lepidozia fugax, p. 66; L. hirta, p. 87; L. kirkii,

p. 84; L. microphylla, p. 30; L. obtusiloba, p. 45;

L. pumilla, p. 79; and L. setigera, p. 39). In Te-

laranea the spores are always delicately areolate

(as in T. nematodes; cf. fig. 90 : 4 in Schuster,

I.e.). The spore differences appear to be absolute:

even in taxa like L. kirkii, in which spores may

show a marked tendency for the papillae to coa-

lesce to form vermiculate "ridges," these ridges

may fork, but never delimit areolae.

In view of the considerable intrageneric vari-

ability in criteria 1-6, the differences in spore

"ornamentation" seem to be of major significance

in separating the two genera. Unfortunately, they

have not been observed in more than a dozen

taxa.

6 In L. microphylla, cortical cells are both larger and

thinner- walled than the much smaller, thick- walled, 1-3

strata of cells lying immediately within (Fig. 9 : 14).

7 The difference is far from absolute. Thus T. centipes

has shallowly lobed or lobulate bracts (Fig. 4 : 10, 12),

much like the typical condition in Lepidozia. Also, in

Lepidozia setigera 9 bracts are deeply lobed or laciniate

(Fig. 12:3).

FIELDIANA: BOTANY

FIG. 3. Telaranea centipes (Tayl.) Schust. 1. Leaf of main shoot. 2, 3. Leaves of main shoot. 4. Plant with 3

androecia-bearing Frullania-type branches. 5. Male bract. 6. Stem, cross section. 7. Opposing leaf pair of main shoot,

dorsal view. 8. Antheridium. 9. Androecial branch. 10. Median lobe of leaf. (Figs. 1 , 4-9 from Engel 15593, Tasmania,

track from Ferndene to Mt. Dial; 2, 3, 10 from Engel 15191, Tasmania, Lillydale Falls.)

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

FIG. 4. Telaranea centipes (Tayl.) Schust. 1. First branch underleaves. 2. Portion of main shoot with branch,

ventral view (BUL = first branch underleaf; FB = Frullania-type branch; HL = half-leaf, shown in outline). 3, 5-

7. Underleaves from leading shoots, stem cells stippled (note insertion of 3 young rhizoids in Fig. 5 and of mature

rhizoids in Fig. 7). 4. Half-leaf (sc = subtending stem cell; be = subtending branch cell). 8. Portion of perianth

mouth showing cuticular detail. 9. Perianth, cross section through median third. 10. Perianth bearing shoot. 11. Female

bract (left) and bracteole (right) of innermost series. (Figs. 1-7 from Engel 15593, Tasmania, track from Ferndene to

Mt. Dial; 8-1 1 from Engel 15191, Tasmania, Lillydale Falls.)

10

FIELDIANA: BOTANY

14

FIG. 5. Telaranea tuberifera Engel & Schust. 1. Plant, dorsal view. 2. Germinating tuber; note scattered leaf

rudiments and previous point of attachment of tuber to stolon at opposing end of tuber. 3. Portion of tuber surface

showing a leaf rudiment. 4. Stem, cross section. 5-7. Leaves. 8. Underleaf; note rhizoid position. 9. Median discs

cells showing cuticular detail in part. 10. Old, basal portion of shoot showing stoloniferous branches and a tuber. 11.

First branch underleaves. 12. Tuber, cross section. 13. Portion of main shoot; note ragged leaf apices. 14. Median

lobe of leaf. (All from type of T. tuberifera.)

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

11

FIG. 6. Lepidozia pendulina (Hook.) Lindenb. (1-3 and 8-9; Schuster 67-238); Lepidozia concinna Col. (4-7;

Schuster 67-237d); Lepidozia microphylla (Hook.) Lindenb. (10-13; Schuster 67-237 c, Westland). 1. Median cells

with oil-bodies (X432). 2. Basal leaf cells (X370; 40 /urn scale). 3. Underleaf (top) and stem leaf (X20). 4. Two

leaves and, top, underleaf; to scale of Fig. 3 (X20). 5. Median cells with oil-bodies and (upper right) chloroplasts

(X325; 40 /zm scale). 6. Basal cells with oil-bodies (X272; 60 /mi scale). 7. Two underleaves + three leaves (X15.7;

1 mm scale). 8. Three leaves (X15.7; 1 mm scale). 9. Three underleaves (X15.7; 1 mm scale). 10. Median cells with

oil-bodies + chloroplasts (X650). 11. Underleaves (X40.5; 400 /irn scale). 12. Leaves (X40.5; 400 /mi scale). 13.

Stem cross section.

INTRAGENERIC CLASSIFICATION — The intragener-

ic classification and phylogeny of the genus Lep-

idozia remain poorly understood. Study of a wide

array of taxa has led Schuster (2000, pp. 183-207)

to divide the genus into six subgenera, each with

1-5 sections. What follows is tentative, in part

because the sporophyte generation of so few taxa

is known. Nonetheless, it is clear that the genus

consists of an array of relatively well-defined

groups (subgenera, sections). The aspect of, for

example, Lepidozia reptans and L. (Chaetolepi-

dozid) pinnaticruris (cf. fig. 64, Schuster, 2000)

and L. (Cladolepidozid) bursifera (cf. fig. 66A,

Schuster, 2000) is so different that initially, one

might consider placing them in different genera.

Fulford (1966, p. 181) states that "unlike Bazza-

nia," Lepidozia "does not consist of obvious,

well-marked groups or sections." We are im-

pressed, on the contrary, with the relatively

marked homogeneity of Eazzania (and cannot rec-

ognize subgenera within it) versus the perceptible

heterogeneity of Lepidozia. Students of the ho-

12

FIELDIANA: BOTANY

larctic flora tend to conceptualize Lepidozia by L.

reptans; as we show elsewhere (p. 31), this is

highly atypical of the genus and should not serve

as a "model" for the genus.

We assume that erect growth, branching mal-

leability, weakly asymmetrical leaves, and trans-

verse leaf orientation are all signs of "primitivi-

ty"; on these bases, subg. Dendrolepidozia is

primitive. With the exception of only one other

occurrence, it is the only element in the genus in

which, at least sporadically, terminal branches oc-

cur from all three merophyte rows. In Schuster

(1980), it is proposed that the most generalized

element within the allied genus Kurzia is formed

by K. mollis (Steph.) Engel and Schust. and K.

setiformis (De Not.) Engel & Schust., taxa in

which branching is "highly plastic"; in the first

of these taxa also we have seen occasional ven-

tral-terminal (Acromastigum-type) branches. In

these species also, leaves are very deeply quad-

rifid, with a low disc and tapered, narrow lobes,

exactly as in Lepidozia (Dendrolepidozia) spinos-

issima. In both of these taxa the leaves are slightly

asymmetrical, with longer dorsal margins (and

lobes) (compare Figs. 7 : 9 and 8 : 3, 4, 10 in this

paper with fig. 1 : 8 in Schuster, 1980). Both taxa

have leaves that are almost transversely inserted;

leaves and underleaves tend to be similar; the lat-

ter tend to be more than two-thirds the area of the

former. The erect growth of Dendrolepidozia also

may be considered a primitive feature. Interest-

ingly, Acromastigum-type branching occurs again

only in L. pumila (Fig. 29 : 3), a species of the

relatively derived sect. Notholepidozia. The pres-

ence of ventral-terminal branching in L. pumila is

all the more remarkable because it occurs in a

plant with a closely prostrate habit.

TAXONOMIC SEQUENCE ADOPTED — We start our

treatment with Dendrolepidozia because of the

features emphasized above. In the erect, almost

"woody" axes, Dendrolepidozia seems allied

with Mastigolepidozia (the L. microphylla ele-

ment), and we treat this group next.8 Both of these

groups show copious branching, and they are reg-

ularly 2-3-pinnate. By contrast subg Notholepi-

dozia consists basically of loosely prostrate plants

(becoming suberect only when crowded), with a

stem usually showing slight cortication. In this

8 There is also a clear similarity (and presumable af-

finity) in the "woody" main stem, the transversely ori-

ented leaves, and the virtual isophylly to the otherwise

highly derivative subg. Cladolepidozia (compare figs.

65, 66, and 66A in Schuster, 2000).

subgenus anisophylly is usually marked; the

leaves are usually strongly incubous, often with

deflexed apices. Branching in subg. Notholepido-

zia is generally once-pinnate, with branches typ-

ically of limited length (unless they end in flagel-

la; in this case, they often bear secondary branch-

es). This is clearly seen in the generic type, L.

reptans (cf. fig. 86 : 7, 9, in Schuster, 1969).

Subgenus Notholepidozia also shows a series of

derived tendencies: Some taxa develop strong

dentition of the leaves; others develop strikingly

ampliate dorsal leaf bases; some develop cuticular

ornamentation. Subgenus Notholepidozia is there-

fore treated as third in a linear arrangement, al-

though we do not wish to imply that Mastigole-

pidozia "gave rise" to subg. Notholepidozia.

Subgenus Glaucolepidozia is treated last in our

arrangement, for several reasons. It is highly de-

rived in the peculiar cutinization, which gives the

plants a striking opaque, whitish aspect not seen

in any other taxa of Lepidozia. In other respects,

it is relatively close to subg. Lepidozia, for ex-

ample, in its creeping growth, its leaf and under-

leaf form, and its ramification. Subgenus Glau-

colepidozia is derived in its leaf cells. These are

mostly equally firm-walled, very chlorophyllose,

and have few, very small, mostly few-segmented

oil-bodies, always smaller than the chloroplasts

(Figs. 36 : 9, 10; 38 : 8-10).

Subgenus Austrolepidozia, with only L. uloth-

rix, also represents a highly derivative element. It

has ciliate to spinose-ciliate leaf margins, much as

in Chaetolepidozia; as in that subgenus, cilia are

formed of strongly elongated, thick-walled cells.

In Austrolepidozia, cilia are usually alternate,

more sporadically opposed; in Chaetolepidozia,

each lobe typically bears two to three pairs of op-

posed cilia. Austrolepidozia approaches subg.

Lepidozia in the strongly asymmetrical leaves that

are relatively shallowly lobed (Figs. 33 : 3-5; 34 :

1,2).

The two subgenera not regionally represented,

Cladolepidozia and Chaetolepidozia, would nat-

urally fall last in any linear arrangement; both are

highly derived groups. In Cladolepidozia, leaf

lobing is greatly reduced, and the vestigial lobes

are represented by mere cilia (as in L. cladorhiza

(Reinw. et al.) G. L. & N; see Schuster, 2000, figs.

65 and 66, and Piippo, 1984, fig. 6 : a-g) or

coarse, serrate teeth (as in L. ferdinandi-muelleri

Steph.; see Piippo, 1984, fig. 6 : h-n; Mizutani,

1968, fig. VI). Some taxa in this subgenus have

ampliate leaf- and underleaf-bases, converted into

water sacs (see Grolle, 1967, fig. 2, and Schuster,

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

13

2000, fig. 66A of L. burst/era Hatt. & Grolle). In

Chaetolepidozia the deeply lobed leaves and un-

derleaves are armed with several pairs of spines-

cent teeth and/or cilia (see Schuster, 2000, fig.

64), much as is typical of, for example, Temnoma

and Trichocolea (as in L. lacerifolia Steph.; see

Piippo, 1984, fig. 8 : a-b). This last subgenus is

known only from gametophytic plants, and in-

deed, discovery of sporophytic material may ne-

cessitate removal of the subgenus from Lepidozia

and its elevation to generic status. Both Cladole-

pidozia and Chaetolepidozia have almost

"woody" main stems, 20-24 cells high, with 3-

5 strata of thick-walled cells peripherally, forming

a rigid cortex (figs. 64 : 9; 66A : 13, Schuster,

2000). Such rigid stems are approached, in our

taxa, only by L. pendulina (Fig. 10 : 15) and L.

microphylla (Fig. 9 :14).

After much deliberation, we have excluded

subg. Lepidozia in our treatment. That subgenus

is exclusively Laurasian; it has consistently tiny,

homogeneous, smooth (rarely with an "appen-

dix" cut-off) and numerous oil-bodies per cell (p.

16). In all Australasian taxa we have studied, oil-

bodies are fewer per cell, relatively large, and

consistently coarsely botryoidal (Fig. 6:1, 2, 5,

6, 10); cf. p. 16.

The bulk of taxa thus belong in subg. Notho-

lepidozia. This remains relatively heterogeneous

and, after a suite of sporophytes has been ade-

quately studied, may possibly be subject to sub-

division.

The perimeters of the subgenus emerge from

the conspectus (p. 18) of subgenera, modified

from Schuster (1973, 2000).

Taxonomic Characters and Morphology

BRANCHING, FORM, AND ORIENTATION OF BASAL

BRANCH UNDERLEAVES — In general, the first un-

derleaf formed on a Frullania-type branch differs

substantially from succeeding branch underleaves.

Usually the initial branch underleaf is so oriented

that it is not "in line" with succeeding branch

underleaves; it is often displaced so that it is more

nearly in line with lateral stem leaves. In most

taxa the initial branch underleaf is substantially

smaller (and often has fewer lobes) than stem and

branch underleaves distad of the basal one.

In L. kirkii (Fig. 31:3, 4), however, the basal

branch underleaf is nearly identical to succeeding

branch underleaves, aside from usually being 3-

rather than 4-lobed; it is oriented in line with suc-

ceeding branch underleaves.

In L. spinosissima the basal branch underleaf is

normally 2-3 lobed and narrower than successive

4-partite underleaves (Fig. 7 : 1 1); it is oriented so

that it is not quite in line with successive under-

leaves (Fig. 8 : 1,2), and the branch superficially

appears almost axillary to it. Such displacement

of the initial branch underleaf is typical of most

taxa. Thus in L. acantha and L. elobata the first

branch underleaf is displaced and is 1 -segmented,

versus the 4-segmented succeeding branch under-

leaves (Fig. 27 : 3, 5).

In related taxa, the first branch underleaf may

be a useful character. The consistently monocru-

rous condition in L. acantha and L. elobata is

unique in sect. Notholepidozia. Moreover, the

shape of the first branch underleaf is different be-

tween these species: subulate in L. acantha (Fig.

27 : 3, 5), versus ovate-lanceolate in L. elobata

(Fig. 28 : 1, 2). However, the first branch under-

leaf of related taxa, clearly of the same species

complex, may be variable and therefore of little

taxonomic utility. Thus, in L. novae-zelandiae the

initial branch underleaves range from (1)2 to 3(4)-

lobed, and may be 1-2-fid even on a single stem

(Fig. 26 : 7). In the allied L. pumila, initial branch

underleaves are (l)2-4-fid and may be 2-4-lobed

even within the same population (Fig. 29 : 2, 3,

11, the type).9

STEM ANATOMY — The generic type, L. reptans,

has ca. 20-24 rows of cortical cells that are slight-

ly to moderately larger than the medullary cells;

the medullary have walls scarcely more firm-

walled than the cortical (fig. 87 : 8 in Schuster,

1969). A cortex is hardly differentiated.

In New Zealand taxa there is wide variation in

(a) degree of differentiation of a hyaloderm and

(b) degree of rigidity of the medulla, depending

on how strongly thick-walled the outer and some-

times the inner strata of medullary cells are. The

following general types exist:

1. In the most vigorous taxa, such as L. pen-

dulina (Fig. 10 : 15), a hyloderm hardly exists;

some 5—8 outer strata are thick-walled, and the

outermost layer is not differentiated as a hyalo-

derm. The central medulla consists of thin-walled

cells.

9 A cautionary note: Initial branch underleaves and

their number of lobes may reflect vigor of the shoot.

Thus, in L. concinna the lower, weaker shoot sectors

may have 3-fid basal branch underleaves; the vigorous

distal shoot sectors have them (3)4-fid (Fig. 19 : 1, 1A).

14

FIELDIANA: BOTANY

2. In L. microphylla the main axis is rigid, as

in L. pendulina, and is ca. 22-26 cells in diameter

(Fig. 9 : 14). Here the peripheral stratum is dif-

ferentiated as a recognizable hyaloderm, and 2-4

strata within are of smaller cells, with rather firm

walls.

3. In L. spinosissima (Fig. 8:7, 9) the main

axis is much as in L. microphylla: a weakly dif-

ferentiated hyaloderm, with several strata of

smaller thick-walled cells within that grade into a

larger-celled medulla.

These taxa and L. kirkii (Fig. 31 : 10) agree in

having rather massive stems, with numerous rows

of cortical cells.

4. Within the more reduced taxa, such as L.

acantha (Fig. 27 : 2) and L. novae-zelandiae (Fig.

26 : 13), we see progressive reduction in axial

complexity. An ultimate point is reached in L. no-

vae-zelandiae var. minima, with the stem only ca.

6-7 cells in diameter (Fig. 26 : 13). Stems of var.

minima have slightly larger cortical cells that are

strongly thick-walled, much as the cells within,

which average rather smaller in diameter.

In general, there is a correlation between vigor,

erect growth, and evolution of a relatively

"woody" stem, as in L. microphylla. Smaller

taxa, especially if not developing a stiffly erect

main axis, tend to have weaker stems, with less

histological differentiation. Thus in L. acantha

(Fig. 27 : 2) and the allied L. laevifolia (Fig. 22 :

12), stems with a fundamentally similar anatomy

are found. The cortical and medullary cells in L.

laevifolia are considerably firmer-walled than in

L. acantha.

In virtually all the cases cited there is a weakly

to at most moderately differentiated hyaloderm of

larger, usually more leptodermous cells, contrast-

ed to a medulla in which at least peripheral strata

are firm to thick-walled. In subg. Glaucolepido-

zia, however, the stem is formed of thin-walled or

virtually thin-walled cells throughout, and the cor-

tical cells are hardly differentiated (at least in

cross section) from the medullary cells (Figs. 36 :

11; 37:7; 38:7).

We have illustrated the axial anatomy of a

range of species but have de-emphasized axial

anatomy in the taxonomic treatment because,

aside from the four general types (1-4, above),

the degree of histological differentiation in the

stem reflects, in part, overall gametophyte size.

Thus, vigorous main axes in L. microphylla (Fig.

9:14) have stems ca. 22-26 cells in diameter,

versus only ca. 12 cells high on axes of weaker

plants (Fig. 6: 13).

VENTRAL MEROPHYTE WIDTH — In the generic

type, L. reptans, ventral merophytes are ca. 5 cell

rows broad (Schuster, 1969).

Austral taxa exhibit a broad range of merophyte

width, a character best observed in material

cleared in Hoyer's fluid. Some taxa have many

cell rows (e.g., L. microphylla, L. pendulina), and

merophyte width data are not cited under such

species. In the smaller taxa, however, merophyte

width is typically fixed and may be useful in spe-

cies discrimination.

Thus, in the complex centering on L. novae-

zelandiae, L. acantha, et al. (sect. Notholepido-

zia), ventral merophytes may be only 4-5 cell

rows broad (L. novae-zelandiae; Figs. 24 : 14;

26 : 13), ranging to 5-6 rows (L. pumila. Fig. 29 :

10), to 7 in L. fugax (Fig. 23 : 10), to as many as

8 rows in L. laevifolia (Fig. 20 : 11). Ventral mer-

ophyte width is used to differentiate between spe-

cies pairs within the key. For example, L. bidens

has ventral merophytes 8 cells across (Fig. 30 :

10) versus 5-6 cells in L. pumila; the character

also is used in the couplet distinguishing L. fugax

from L. novae-zelandiae. Lepidozia concinna,

also of sect. Notholepidozia, has ventral mero-

phytes 10 cells broad (Fig. 18 : 11).

All species within subg. Glaucolepidozia differ

at least in the range of ventral merophyte width.

The widths range from 3-4 cells across (L. digi-

tata}, to 4-6 (L. glaucescens), to 8-9 (L. glau-

cophylla), to 9-10 (L. bisbifida). In the key, the

widths easily separate L. bisbifida from L. digi-

tata.

LEAVES — Leaves of Lepidozia may be subsym-

metrical or weakly asymmetrical, as in L. spinos-

issima (Fig. 7 : 9) or L. acantha (Fig. 27 : 6), but

typically they are moderately to strongly asym-

metrical. Such leaves have the dorsal margin of

the disc longer than the ventral margin. The lon-

ger margin is often accompanied by various de-

grees of dilation of the dorsal sector of the leaf;

examples are L. setigera (Fig. 1 1 : 3, 4) and L.

procera (Fig. 13:5, 7).

Leaves in Lepidozia typically are 4-lobed, al-

though in some taxa sporadic 3- and 5-lobed

leaves occur. The sinuses usually descend for less

than 0.55 their length, although exceptions are

present, for example, in L. spinosissima, where

leaves are divided to 0.7-0.9 (Fig. 7 : 9). The dis-

tance from the dorsal sinus base to the leaf inser-

tion is at times much greater than from the ventral

sinus to the insertion. This occasionally results in

the dorsalmost two lobes being conspicuously

paired (e.g., L. bidens. Fig. 30 : 2, 3). The differ-

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

15

ences in distance between sinus base and leaf base

often correlate with the degree of leaf asymmetry.

Thus, the conspicuously shallow dorsal sinus of

L. procera accompanies a strongly asymmetrical

leaf (Fig. 13:5, 7). The dorsal sinus in L. obtu-

siloba is exceptionally shallow, and in this species

it may be reduced to a mere notch (Fig. 14 : 1).

Leaves of New Zealand Lepidozia species are

typically devoid of teeth, although some species

may at times have the dorsal margin of the disc

armed with a few distant, irregular, few-celled

teeth (e.g., Fig. 35 : 7, 9, of L. glaucophylla).

However, conspicuous leaf margin ornamentation

occurs in two groups. Subgenus Austrolepidozia,

with L. ulothrix, has leaf lobes with up to 10-12

cilia, usually oriented in opposed pairs (Fig. 34 :

2, 6). Also, the dorsal margin and the confluent

margin of the dorsal lobe have up to 12-19 spi-

nose teeth or cilia. Members of sect. Kirkii have

the dorsal margin of the disc and the dorsal 1-2

lobes coarsely and variably toothed (Figs. 31 : 1;

32 : 2, 4, 10).

LEAF CELLS, CUTICLE, AND OIL-BODIES — Leaf

cells in Lepidozia are typically firm-walled; and,

at times, well-developed trigones may be present

(as, for example, in L. concinna (Fig. 18:6) and

L. ornata (Fig. 17 : 12). However, some of our

New Zealand species may have uniformly thin-

walled cells, and it is not altogether surprising that

species with such cells are found in sheltered,

deeply shaded loci such as cavelike recesses at

cliff bases or in pockets at tree bases. Lepidozia

digitata is an example (Fig. 38 : 9, 10). Lepidozia

reptans, the generitype, has cells minimally in-

crassate (cf. fig. 86 : 6 in Schuster, 1969). Leaf

cells in Lepidozia are irregularly arranged, irreg-

ularly polygonal in shape, and never consistently

elongated.

The ornamentation of the cuticle is a significant

character within Lepidozia. Plants of subgenus

Glaucolepidozia have a dull, water-repellent sur-

face due to the presence of a waxy cuticle. The

leaves and stems of such plants are opaque, whit-

ish, and glaucous, sometimes becoming pale

brownish with age. Some Lepidozia species have

papillose lobes but a smooth disc (Fig. 23 : 4, L.

fugax), while others have both lobes and disc pa-

pillose (Fig. 27 : 7, 8, L. acantha). The character

is particularly useful in distinguishing the smaller

species of sect. Notholepidozia. For example, dis-

tinctly papillose leaves occur in L, laevifolia (Fig.

20 : 8-10) and L. novae -zelandiae (Fig. 24 : 6),

whereas leaves are smooth (or at most faintly and

indistinctly papillose) in L. pumila and L. bidens.

Care is required in the use of this character, be-

cause ornamentation at times is subtle and rather

difficult to observe. For this reason, it is recom-

mended that several leaves, particularly from sev-

eral different shoots, be routinely checked for

presence versus absence of cuticle features.

We have studied the oil-bodies of some 5-6

taxa of subg. Notholepidozia. They are consis-

tently relatively few per median cell (usually 2-

7(8) in L. pendulina and 1-5 per cell in L. con-

cinna} and are always distinctly, if not coarsely,

botryoidal. In subg. Mastigolepidozia (L. micro-

phylla) they are somewhat less coarsely botryoi-

dal, average rather larger than in Notholepidozia

species studied, and occur 2-5 per median cell

(Fig. 6 : 10). Only in subg. Glaucolepidozia do we

see very different, often vestigial oil-bodies (cf.

p. 92 and Figs. 36 : 9; 38 : 10).

By contrast, all workers find that the generic

type, L. reptans, has homogeneous oil-bodies

(Hattori, 1953; Schuster, 1969, fig. 86 : 6; Furuki

& Higuchi, 1997). Similar oil-bodies are reported

for the other two Asiatic taxa, L. robusta and L.

subtransversa (Furuki & Higuchi, I.e.). In these

three taxa, oil-bodies are more numerous: 5-16

per cell in L. reptans (Schuster, 1969; Furuki &

Higuchi, I.e.) and 5-15 or 5—20 per cell in L. ro-

busta and L. subtransversa (Furuki & Higuchi,

I.e.). In none of the holarctic taxa studied did we

ever see 2—4 or 2-5 oil-bodies per cell, typical of

the New Zealand species studied (cf. Fig. 6 : 1,5,

10). Only in basal cells (as of L. pendulina; Fig.

6 : 2) may there be more numerous oil-bodies, but

in other taxa (e.g., L. concinna), there may be

only 4-8 per basal cell (Fig. 6 : 6). The taxonomic

implications of these observations need to be but-

tressed by in-depth studies of a wider range of

taxa. They strongly suggest that the internal clas-

sification of Lepidozia s. lat. needs intensive

study.

ASEXUAL REPRODUCTION — Asexual reproduc-

tion is sporadically present in our Lepidozia spe-

cies and occurs by fragmenting or caducous leaf

lobes, whose bases persist as truncate stubs. Cad-

ucuous leaf tips are found in L. fugax (Fig. 23 :1,

3, 5) and give the plants a distinctive ragged ap-

pearance (for details, see Differentiation section

under that species, p. 66). Lepidozia fugax is the

only regional representative of subg. Notholepi-

dozia with asexual reproduction. At least the

weaker phases of L. digitata (subg. Glaucolepi-

dozia) may have fragmenting or irregularly ca-

ducous leaf lobes (Fig. 38 : 1, 3, arrows), and

16

FIELDIANA: BOTANY

even underleaf lobes may be caducous. Dissemi-

nation presumably is by such leaf fragments.

ANDROECIA — Although the generitype (L. rep-

tans) is autoecious, the vast majority of species

within the genus are dioecious. Androecia occur

on short, spicate, often cernuous, determinate,

ventral-intercalary branches devoid of normal

leaves and underleaves (Fig. 7 : 2) and are typi-

cally found in the median or basal sectors of a

shoot. (Figure 7 : 1 shows androecia on the pri-

mary and secondary branches of the median sec-

tor of the plant.) In some species, androecia also

occur terminal on primary and secondary branch-

es. Regardless of position, the androecia are in-

conspicuous, and those of the main shoot may not

exceed the tips of the underleaf lobes, as in L.

ulothrix.

The androecia are rather uniform and do not

yield characters of taxonomic significance at any

level. Bracts are bifid to at most 0.4, with acute

to acuminate lobes. The dorsal margin is entire or

may have a tooth, and several slime papillae may

be present, with or without the tooth. The dorsal

margin in several unrelated species (L. ornata, L.

ulothrix, L. bisbifidd) is dilated, forming a mod-

erately rounded projection (Fig. 34 : 12, L. uloth-

rix).

GYNOECIA — The abbreviated gynoecial branch-

es in Lepidozia have several progressively larger

gyres of bracts and bracteoles (Fig. 16 : 1, L. ob-

tusilobd). The lowermost gyre typically consists

of inconspicuous scalelike bracts and bracteoles,

while the inner 1-2 gyres are erect, concave, and

sheath the lower part of the perianth (Fig. 7 : 4,

L. spinosissima). In general, in Lepidozia the 9

bracts are strongly differentiated from leaves in

both form and size and are subisophyllous to is-

ophyllous (Fig. 7 : 5, L. spinosissima; Fig. 15:2,

L. obtusilobd). The generic type, L. reptans, has

very large bracts and bracteoles, with the apices

usually 4-lobulate (Schuster, 1969, fig. 87 : 6, 7).

The $ bracts and bracteoles in most of our taxa

are shallowly 4- (exceptionally 5- or 6-) lobate to

lobulate, as in, for example, L. spinosissima (Fig.

7 : 4, 5). They are particularly shallowly lobed in

L. obtusiloba (Fig. 15 : 2, 3), and in this species

the bracts may occasionally have obsolete lobes,

with each represented solely by a slime papilla

mounted on 2 laterally juxtaposed cells that pro-

ject slightly more than the neighboring cells (Fig.

15 : 2, bract at left, 4). In Lepidozia laevifolia the

bract summit may have 4 small teeth that are

hardly differentiated from the otherwise crenate-

denticulate apex (Fig. 21 : 2).

There are, however, two notable deviations

from this general pattern. First, in L. setigera the

bracts and bracteoles are 4-partite for ca. 0.4-0.5

their length and are much less elongated than in

the preceding taxa cited (Fig. 12:3). Second,

gynoecial bracts may have only 2 teeth or lobules,

as in L. pumila, L. bidens, and L. hirta, but in all

of these species, 4 dentate-lobulate bracts are also

present. Bracts with 2 short lobes at times are pre-

sent in L. laevifolia (Fig. 21 : 3).

Characters involving bract lobe and lamina

margins are not of taxonomic significance, al-

though differences exist between some species.

The lobes typically are composed of ± regularly

rectangular and (particularly toward the lobe mar-

gins) irregularly sinuate-rhomboidal cells. The

apical end of the lobe marginal cells is frequently

thickened and variously diverging, often only fee-

bly so, as in L. obtusiloba (Fig. 15:3). In other

instances the cells markedly diverge and result in

a crenate to denticulate lobe margin (Fig. 7 : 6, L.

spinosissima).

Bract lamina margins are bordered by cells var-

iable in shape and orientation, often thinner-

walled, typically narrowly elongated, and often ±

sinuate-rhomboidal in shape. The apical or free

end of the marginal cells in most species is vari-

ously divergent and forms a short projection or a

tooth, and in such cases an irregularly crenulate

to crenate-denticulate margin results. In some spe-

cies, ornamention is primarily in the distal half of

the lamina margin (Fig. 7 : 5, L. spinosissima); in

others, the margins are ornamented to the base (L.

laevifolia, Fig. 21 : 5). The apical end of the mar-

ginal cells is exceptionally nonprotuberant, a con-

dition that results in an esentially smooth margin,

as in L. obtusiloba (Fig. 16:5), although the mar-

gins of this species may occasionally be crenu-

late-denticulate by variously projecting cells (Fig.

16: 1).

Perianths are fusiform or ellipsoidal-fusiform in

shape (Figs. 7:4; 15 : 1) and are terete in the me-

dian and basal sectors (Figs. 7 : 7a; 15 : 9); the

cylindrical sector of the perianth is (2)3-stratose

(Fig. 15 : 8b, m). Perianths become narrowed and

bluntly 3-gonous distally (Fig. 7 : 7b, c) and deep-

ly plicate for varying distances just below the

mouth. The perianth also becomes unistratose dis-

tally (Fig. 15 : 8d, d'). The perianth mouth in most

of our species is shallowly 3-lobed, with the lobe

margins composed of irregularly sinuate-rhom-

boidal cells (Fig. 15:6). These mouth cells are

laterally free for varying lengths at the apical end,

and only rarely are completely free laterally. This

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

17

condition results in the prevailing crenate-dentic-

ulate mouth. Examples of exceptions to this con-

dition follow. The perianth mouth in L. spinosis-

sima is shallowly lobulate (not divided into 3

lobes), and the teeth of the mouth are composed

of a single cell or a uniseriate row of at most 3

cells (Fig. 7 : 8), with the teeth thus being more

protracted than in most species. The perianth of

L. laevifolia is shallowly 3-lobed, and the mouth

crenate-denticulate, with mouth cells laterally free

for varying lengths at the apical end (Fig. 21 : 6,

7). The shallowly 3-lobed perianth of L. obtusi-

loba has mouth cells laterally fused to the summit

or nearly so, and the mouth is thus merely regu-

larly crenulate (Fig. 15 : 6, 7).

The calyptra extends ca. 0.5 the perianth length,

and no evidence is present of stem calyptra de-

velopment (the unfertilized archegonia remain at

the extreme base of the calyptra).

SETA ANATOMY — In Schuster (1969, p. 17), the

seta of Lepidozia is stated to be "where known

of 12 or more rows of epidermal cells'" (author's

emphasis). The generic type, L. reptans, has 12

epidermal cell rows.

However, the seta has 8 epidermal cell rows in

L. laevifolia (Fig. 20 : 12), L. fugax, and L. pum-

ila, and 9 epidermal cell rows in L. hirta (Fig.

32 : 9). Several relatively vigorous species have a

greater number of epidermal cell rows. Thus, L.

setigera has 11-13 rows (Fig. 12 : 11), and L. ob-

tusiloba (Fig. 16 : 2) and L. pendulina each have

16 rows. There seems to be at least a weak cor-

relation between gametophytic vigor and seta

anatomy. The character is used in the key to help

separate L. obtusiloba, with 16 rows, from the

taxa with 8 rows mentioned above. Seta differ-

ences also distinguish L. kirkii (10-14 rows) from

L. hirta (9 rows).

CAPSULES — The capsule in Lepidozia is ellip-

soidal to oblong in profile (Fig. 15 : 10, L. obtu-

siloba), and, like the seta, there seems to be a

weak correlation between gametophytic vigor and

the thickness of the capsule wall. Thus, smaller

species, such as L. fugax and L. pumila, have 3-

stratose capsule walls, whereas larger species,

such as L. microphylla and L. obtusiloba (Fig. 16 :

3), have 4-5-stratose capsules, and the vigorous

L. setigera has a 5-6-layered capsule wall. The

correlation is not absolute, since L. laevifolia, a

small species, has both 3-4- and 4-5-stratose

valves (Fig. 21 : 8, 9), and L. kirkii, a relatively

small species, has 5-stratose capsule walls. The

character is used in the key to species to aid in

the separation of L. obtusiloba, with thicker cap-

sules, from L. fugax and L. pumila. In all Lepi-

dozia species we have studied, the epidermal cells

have a "two-phase" ontogeny, as defined in

Schuster (1966). This is true of the generic type

(fig. 87 : 3, 4, Schuster, 1969) and of studied New

Zealand taxa, for example, L. obtusiloba (Fig. 16 :

4) and L. laevifolia (Fig. 21 : 10). Aside from dif-

ferences in degree to which the "nodular" thick-

enings are developed and whether they tend to

coalesce (Fig. 16:4) or are hardly so (Fig. 21 :

10), the genus seems singularly uniform with re-

gard to capsule wall microanatomy.

SPORES — The spores in Lepidozia species bear

coarse, sharply defined papillae and short, ver-

miculate markings (Fig. 16 : 6, L. obtusiloba).

The markings or ridges may fork, as in L. pumila,

L. kirkii, and L. hirta, but do not delimit areolae.

Conspectus of Subgenera of Lepidozia

1. Leaves clearly, usually conspicuously (3)4(5)-lobed, devoid of water sacs; underleaves various, never

with water sacs; leaves subtransverse to clearly incubous, normally conspicuously asymmetrical, not

strongly narrowed to the lobe apex 2

2. Leaves and underleaves with lobes entire-margined or variously toothed, the teeth, if present,

usually broad-based, never in opposing pairs 3

3. Leaves clearly incubously inserted, obliquely or virtually horizontally oriented, ± clearly asym-

metrical, quadrilobed for under 0.6 their length; Acromastigum-type branches uniformly lacking

(exc. in 1 species of sect. Notholepidozia) 4

4. Plants devoid of a waxy cuticular covering, not wholly opaque, green to yellow-green or

(rarely) brownish in life; cells quite distinct and firm in moist preparations; oil-bodies con-

spicuous, homogeneous or ± botryoidal, persistent, (2)4-10 or more per cell 5

5. Plants with leaves ± imbricate, much broader than stem (very incubously inserted and

oriented); branching primarily 1 -pinnate (the pinnae simple or rarely with more than one

secondary branch); stem with cortication weakly developed 6

18

FIELDIANA: BOTANY

6. Oil-bodies essentially homogeneous, smooth (sporadically some few septate), usually

(8)10-16(23) per median cell [Subg. Lepidozia]

6. Oil-bodies finely to coarsely botryoidal, usually 2-8 per median cell (in basal cells

to 4-18 per cell) Subg. Notholepidozia Schust.

5. Plants with leaves scalelike (especially on branches), very remote, their width not ex-

ceeding that of the stem; branching regularly 2-3-pinnate; stem strongly corticated, stiffly

erect but branches drooping. (Oil-bodies usually 2-5 per cell, rather finely botryoidal.)

Subg. Mastigolepidozia Schust.

4. Plants with a waxy cuticle, whitish, opaque, glaucous, the cells obscured because of the

cutinized surface; cells thin-walled; oil-bodies tiny, few-segmented, often ephemeral and

obscure, usually (1)2-4(5) per cell. Plants flatly creeping, with subcontiguous to contiguous

leaves Subg. Glaucolepidozia Schust.

3. Leaves subtransversely inserted and oriented, subsymmetrical, 0.7-0.9 quadrifid (the lobes nar-

rowly lanceolate, longly tapered); Acromastigum-type branches sporadic. Plants erect, dendritic,

stiff, with cernuous branches, nitid Subg. Dendrolepidozia Schust.

2. Leaves and underleaves with lobes armed with (2)3-4 or more pairs of sporadically to mostly

opposed spinose or ciliiform teeth (the lower tooth sometimes elaborated as a secondary lobe).

Leaves large, clearly incubuous, concave, imbricate; cuticle striolate 7

7. Dorsal sinus of leaf descending less than 0.45 the leaf length, the disc high; leaf asymmetrically

ovate, length equal to or greater then lamina width; leaf lobes with 1-3(4-5) cilia, which are

mostly sporadically opposed; basal branch underleaf (BUL1) 3-5(8) ciliate and/or lobed; stem

half-leaf, associated with branch, less than 0.5 bilobed, lobes usually with cilia and/or teeth

alternate Subg. Austrolepidozia Schust.

7. Dorsal sinus of leaf descending 0.55-0.65 leaf length, the disc low; leaf (flattened) asymmet-

rically obtrapezoidal, length inferior to disc width; leaf lobes with 7-16 cilia, mostly in opposed

pairs; basal branch underleaf deeply 2-3 fid, with copious paired cilia; stem half-leaf to 0.65-

0.8 trifid, with copious cilia, mostly in pairs Subg. Chaetolepidozia Schust.

1 . Leaves not lobed at apex, the leaves merely ciliolate or ciliate at the narrowed apex, with or without

an elaborated water sac at the base; underleaves ovate, broadest at base, with or without basal water

sacs; leaves transverse or feebly incubous only, erect and sheathing, ovate-reniform to very broadly

ovate, subsymmetrical, strongly and ± abruptly narrowed to the triangular apex, which bears merely

4 minute, uniseriate cilia (= the lobes). Cells finely papillose. New Guinea to Indomalaya

Subg. Cladolepidozia Schust.

Conspectus of New Zealand Lepidozia Taxa

Subg. Dendrolepidozia Schust. (p. 23)

L. spinosissima (Hook. f. & Tayl.) G. L. &

N. (p. 23)

Subg. Mastigolepidozia Schust. (p. 27)

Sect. Microphyllae (Lindenb.) Grolle (p. 28)

L. microphylla (Hook.) Lindenb. (p. 28)

Subg. Notholepidozia Schust., subg. nov. (p. 31)

Sect. Pendulinae Schust. (p. 32)

L. pendulina (Hook.) Lindenb. (p. 32)

Sect. Setigerae Schust. (p. 36)

L. setigera Steph. (p. 36)

Sect. Notholepidozia (p. 40)

L. procera Mitt. (p. 41)

L. obtusiloba Steph. (p. 43)

var. obtusiloba (p. 48)

var. parvula Engel, var. nov. (p. 48)

L. ornata Engel, sp. nov. (p. 49)

L. concinna Col. (p. 52)

L. laevifolia (Hook. f. & Tayl.) G. L. & N.

(p. 56)

var. laevifolia (p. 59)

var. acutiloba Engel, var. nov. (p. 61)

var. alpina Schust. & Engel, var. nov. (p.

62)

L. fugax Engel, sp. nov. (p. 63)

L. novae-zelandiae Steph. (p. 66)

var. novae-zelandiae (p. 70)

var. heterostipa Schust., var. nov. (p. 70)

var. minima Schust., var. nov. (p. 71)

L. acantha Engel, sp. nov. (p. 71)

L. elobata Schust., sp. nov. (p. 74)

L. pumila Engel, sp. nov. (p. 76)

L. bidens Engel, sp. nov. (p. 79)

Sect. Kirkii Schust. (p. 82)

L. kirkii Steph. (p. 82)

L. hirta Steph. (p. 85)

Subg. Austrolepidozia Schust. (p. 87)

L. ulothrix (Schwagr.) Lindenb. (p. 88)

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

19

Subg. Glaucolepidozia Schust. (p. 92) L. bisbifida Steph. (p. 95)

L. glaucophylla (Hook. f. & Tayl.) G. L. & L. digitata Herz. (p. 98)

N. (p. 92) L. glaucescens Engel, sp. nov. (p. 101)

Artificial Key to New Zealand Species of Lepidozia

1. Branching irregularly to regularly once-pinnate, secondary branches rarely or sporadically devel-

oped, if present only 1(2) per primary branch (branches sometimes developing into new leafy

shoots); leaves imbricate to contiguous, the insertion weakly to distinctly incubous 2

2. Plants opaque and ivory-white, becoming pale brownish with age; leaves and stems with a dull,

waxy, water-repellent surface (at least on new growth) 3

3. Dorsal margin of leaf moderately to distinctly ampliate; cells of disc and lobes not papillose

(but with a fine, granular coating); leaves broadly inserted and clasping the stem, distinctly

concave, with involute lobes and margins; leaf insertion strongly incubous to horizontal

4

4. Disc cells with evenly thickened walls, trigones never present; dorsal margin of leaf

strongly ampliate, distinctly auriculate at the insertion, the dorsal margin sinuate to

distantly toothed; leaf lobes subcaudate, ending in a uniseriate row of 3-5 cells. Rare

(South Is., Paparoa Mts.) L. glaucophylla (p. 92)

4. Disc cells lax, nearly thin-walled, trigones lacking to medium; dorsal margin of leaf

moderately ampliate, ± clearly cordate at the insertion, the dorsal margin normally

entire; leaf lobes acute to apiculate, ending in a single cell or a uniseriate row of 2-

3(4) cells 5

5. Plants medium, to 1.5 cm wide (including branches); leaves 0.8-1.3 mm long and

broad, the disc 15-22 cells high at dorsal sinus, the median lobes 6-10 cells wide

at base; underleaves inserted on 9-10 rows of stem cells, the disc 5-9 cells high;

first branch underleaf 2- to 4-lobed, transversely inserted on the stem, often some-

what below the branch. Typically higher-elevation forests to subalpine to alpine

L. bisbifida (p. 95)

5. Plants small, to 0.6 cm wide (including branches); leaves at most 0.55 mm long

and broad, the disc 7-13 cells high at dorsal sinus, the median lobes 3-5 cells

wide at base; underleaves inserted on 3-4 rows of stem cells, the disc 2-4 cells

high; first branch underleaf 2-lobed (or rarely undivided), obliquely inserted at

branch base. Forest plant; 950-1390 m L. digitata (p. 98)

3. Dorsal margin of leaf ± straight; cells of disc and lobes striate-papillose, the papillae

obscured by the continuous waxy coating; leaves narrowly inserted, the disc flat, with

spreading lobes; leaf insertion weakly to distinctly incubous. Disc and lobe cells distinctly

thick-walled; first branch underleaf undivided or 2-lobed (rarely 4-lobed). Chatham Is. +

South Is. (150-300 m) + North Is. (Waipoua area, 340-540 m)

L. glaucescens (p. 101)

2. Plants green, yellow-green, brownish to dark brown, but without a dull, waxy, water-repellent

surface 6

6. Dorsal margin of leaves serrate or dentate-laciniate or ciliate 7

7. Distal sector of leaf lobes ciliiform, ending in a uniseriate row of (5)8-15 distinctly

elongated cells (3-4:1); lobe margins ciliate, often with paired cilia; leaves 2-2.2 mm

wide X 1-1.2 mm long L. ulothrix (p. 88)

7. Distal sector of leaf lobes caudate to acute, the uniseriate row of 0-6 cells that are ±

isodiametric or at most 2:1; lobe margins entire or with a few short teeth or spines;

leaves at most 1.4 mm wide X 0.95 mm long 8

8. Dorsal margin of disc and contiguous margin of lobe irregularly dentate to coarsely

serrate; leaf lobes acute to short- acuminate; underleaves divided to at most 0.5

(the disc 12-16 cells high), the lobe margins entire or sparingly toothed; seta of

10-14 epidermal cell rows L. kirkii (p. 82)

20 FIELDIANA: BOTANY

8. Dorsal margin of disc and confluent margin of lobe ± regularly spinose-dentate,

with discrete, often curved, acuminate spines; leaf lobes caudate; underleaves di-

vided to 0.55-0.8 (the disc 6-9 cells high), the lobe margins laciniate-multifid

(rarely entire); seta of 9 epidermal cell rows L. hirta (p. 85)

6. Dorsal margin of leaves entire (rarely with a basal tooth) 9

9. Leaf lobes setaceous, terminating in a uniseriate row of 6-10(12) elongated capillary

cells. Leaf insertion distinctly incubous; trigones large and bulging to knotlike; under-

leaves distinctly connate on one side L. setigera (p. 36)

9. Leaf lobes never setaceous, the lobes acute or terminating in a short uniseriate row of

up to 2-5 cells 10

10. Leaves appressed to stem, distinctly ventrally secund, ± equally and shallowly

(0.25-0.4) lobed, the two median lobes wider than the outer lobes. Slender, wiry

plants with strongly ventrally secund branches L. procera (p. 41)

10. Leaves spreading, not appressed to stem, the two median lobes not clearly wider

than outer lobes. Leaves often asymmetrically lobed, the dorsal lobes often ±

united in a perceptible pair, the ventral lobes usually shorter and often somewhat

divergent from the dorsal pair 11

1 1 . Leaves and underleaves variously appendaged, with accessory lobes and often

lamellae; underleaves plicate, the lobes ventrally sulcate, the sinus bases

strongly reflexed. Plants brown; branches spreading. Subalpine to alpine spe-

cies L. ornata (p. 49)

11. Leaves and underleaves not appendaged (rarely with marginal teeth); under-

leaves plane 12

12. Plants with brown pigments, typically erect, with distinctly ventrally se-

cund branches (occasionally greenish with spreading branches in var. par-

vuld). Disc cells often distinctly thick-walled, brownish; leaf and underleaf

lobes often bluntly rounded. Seta with 16 rows of epidermal cell rows;

capsule wall 4-5 stratose. Montane to alpine species

L. obtusiloba (p. 43)

12. Plants green, typically prostrate to procumbent, with spreading or ± de-

curved branches. Seta (where known) with 8 rows of epidermal cell rows;

capsule wall (where known) 3-stratose 13

13. Median cells of leaf disc large (26-41 /xm wide); plants medium-

sized; underleaves inserted on 10 rows of stem cells; leaves 0.8-1

mm wide X 0.6-0.9 mm long. Leaves variable in shape, the ventral

lobes typically not much shorter than the dorsal (the branch leaves ±

equally 4-lobed). Cuticle of leaf disc smooth; median disc cells typ-

ically with trigones straight to large and bulging

L. concinna (p. 52)

13. Median cells of leaf disc smaller: 25 /xm wide or less (exc. sometimes

in L. bidens); plants small to minute; underleaves inserted on 9 or

fewer rows of stem cells; leaves usually less than 0.5(0.6) mm in

longest dimension 14

14. First branch underleaf consistently unlobed. South Is. (Westland

Prov., 990 m) + North Is. (Rotorua area) 15

15. Leaves deeply (0.5-0.6) quadrifid, the lobes lanceolate; dor-

sal sinus descending at least 0.5 leaf length; lobes ending in

± elongated cells, the lobe cells mostly rectangulate; first

branch underleaf subulate; stem half-leaf usually 0.45-0.55

bifid; branch leaves mostly 3-lobed . . . L acantha (p. 71)

15. Leaves less than 0.5 quadrifid, the lobes triangular, less than

3X as long as broad; dorsal sinus usually descending to 0.2-

0.3; lobes ending in nonelongated cells, the lobe cells mostly

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA 21

subquadrate; first branch underleaf ovate-lanceolate; stem

half-leaf under 0.25 bifid; branch leaves 4-lobed

L. elobata (p. 74)

14. First branch underleaf of mature shoots 2-4-lobed or mixed 1—

2-lobed. Leaves mostly bisbifid: median sinus deeper, dorsal sinus

usually shallow, usually strongly asymmetrical, the lobes always

strongly unequal 16

16. Leaves ± conspicuously papillose (at least on the lobes) or

if indistinctly papillose (suboptimal populations of L. lae-

vifolid) then the dorsal margin of disc ampliate and the outer

wall of the marginal cells of disc and lobes not thickened

17

17. Marginal cells of disc and lobes with free wall not dif-

ferentially thickened; dorsal margin of disc ampliate, the

margin cordate to auriculate at the insertion; leaves usu-

ally distinctly concave, wet or dry. Leaves coarsely pa-

pillose, the disc closely and distinctly striate-papillose;

lobes with uniseriate row (when present) of nonelongated

cells. Common; usually not below 300 m

L. laevifolia (p. 56)

17. Marginal cells of disc and lobes with a strongly thickened

free wall (the wall often bulging into the cell lumen); dor-

sal margin of disc ± straight to moderately ampliate prox-

imally, cordate at insertion, not clearly auriculate; leaves

plane or only moderately concave, the lobes spreading or

incurved. Rare or infrequent species 18

18. Lobe tips caducous, the cells turgid, barrel-shaped,

the septa constricted; lobes with uniseriate row of 2—

4(5) cells, the cells ± isodiametric, the terminal cell

rounded at the tip; median disc cells 17-24 /nm

wide; underleaves inserted on 7 rows of stem cells.

Margins of disc and lobes occasionally toothed.

South Is., 560-900 m L. fugax (p. 63)

18. Lobe tips not caducous, the cells with straight or

concave walls, the septa dilated; lobes terminating

in a single cell or at most 2 superposed cells, the

cells elongated (to 2:1), the terminal cell ± tapering

to a point; median disc cells 11-16 /urn wide; un-

derleaves inserted on 4-6 rows of stem cells. South

Is. + North Is.; forest species

L. novae-zelandiae (p. 66)

16. Leaves smooth (rarely finely and indistinctly papillose); outer

wall of marginal cells of disc and lobes distinctly thickened

19

19. Plants medium-sized (the main shoots, 700-900 /um

wide, leaf tip to leaf tip); underleaves inserted on 8 rows

of stem cells; leaf disc 8-10 cells high at dorsal sinus,

the cells ± longitudinally elongated; leaves with dorsal

lobes ending in a uniseriate row of elongated (up to 2:

1) cells. Plants of forests (sea level to 330 m on South

Is.; to 920 m on North Is.) L. bidens (p. 79)

19. Plants minute to small (the shoots 365-490 /x,m wide,

leaf tip to leaf tip); underleaves inserted on 5-6 rows of

stem cells; leaf disc 9-12 cells high at dorsal sinus, the

22 FIELDIANA: BOTANY

cells quadrate or slightly longer than wide; dorsal lobes

with cells of uniseriate row ± isodiametric or at most

slightly longer than wide. Plants of forests + subalpine

zone (570-1370 m); South Is. + North Is

L. pumila (p. 76)

1 . Branching regularly and consistently twice-pinnate, the branches slender and drooping; leaves typ-

ically distant, vertically oriented, the insertion transverse. Plants erect, with rigid, woody stems . .

20

20. Leaves inconspicuous, scalelike, narrower than stem, strongly erect and commonly ± appressed

to stem, divided to 0.4-0.5 (median sinus) L. microphylla (p. 28)

20. Leaves conspicuous, wider than stem, stiffly patent or erect-patent from a spreading base, di-

vided to 0.65-0.9 21

21. Leaves ± symmetrical, lobes 6-8 cells wide at base, ending in a uniseriate row of 7-12

cells; dorsal margin of disc straight to slightly concave; underleaf disc 3-6 cells high, the

lobes with a uniseriate row of 7-16 cells L. spinosissima (p. 23)

21. Leaves asymmetrical, lobes 14-24 cells wide at base, with a uniseriate row of 2-4(6) cells;

dorsal margin of disc broadly ampliate; underleaf disc 14-24 or more cells high, the lobes

with a uniseriate row of 1-3(4) cells L. pendulina (p. 32)

Subg. Dendrolepidozia Schust.

Lepidozia subg. Dendrolepidozia Schust., J. Hattori

Bot. Lab. 36: 386. 1973 (1972).

Plants usually stiffly erect, with a rigid, almost

woody main axis with a thick-walled, polystratose

cortex grading into the ± firm-walled medulla;

plants usually a clear light to deep green, without

brownish pigmentation of any sort. Shoot system

usually repeatedly, copiously pinnate, sometimes

with Acromastigum-type branches, with a regular,

progressive diminution in vigor of both axes and

appendages of the branches (the branch leaves

typically smaller, more remote, often fewer-

lobed), at least the ultimate branches often cer-

nuous or drooping, often microphyllous and be-

coming flagelliform. Leaves usually remote, often

strongly so, somewhat vertically oriented and

only weakly to moderately incubously inserted,

the stems extensively exposed both dorsally and

ventrally; lateral leaves usually weakly asymmet-

rical, varying from handlike and deeply 4-5-part-

ed to smaller, handlike to even scalelike, with ab-

breviated lobes. Underleaves similar in form to

leaves, but somewhat smaller, 3-4-lobed with

equal lobes, usually without rhizoids at their api-

ces (rhizoids usually confined to tapered flagelli-

form axes or to lower portions of main stems).

TYPE — Lepidozia spinosissima Mitt.

Dendrolepidozia occurs usually in exceedingly

deeply shaded sites in lowland forests and, as the

name implies, the plants are strongly dendritic.

Although leaf insertion is incubous, leaves are

never incubously shingled. They are remote (on

branches often exceedingly remote), and photo-

synthesis is, to a large extent, a function of the

strongly chlorophyllose stems. The regular dimi-

nution in leaf size and complexity is marked (Fig.

8 : 3, 4, 10). Although terminal branches are usu-

ally of the Frullania type (Fig. 8 : 1, 2), in L. spi-

nosissima, Acromastigum-type ventral terminal

branches occur, with the half underleaf bifid rather

than quadrifid (Schuster, 1982, fig. 1:2 and pp.

23-24).

In the freely 2-3-pinnate branching L. spinos-

issima is similar to L. pendulina, which we place

within subg. Notholepidozia, in section Penduli-

nae. The two taxa also agree in the nearly vertical

leaf orientation, the massive stems, and the mostly

handlike leaves. Lepidozia pendulina, however,

lacks Acromastigum-type branching (Fig. 10: 1,

2), and has relatively strongly asymmetrical

leaves with ampliate dorsal bases (Fig. 10:4) ver-

sus the weakly asymmetrical leaves with subequal

lobes of L. spinosissima (Fig. 8 : 3, 4, 10). In L.

pendulina stem half-leaves associated with ter-

minal branches are nearly always bilobed (Fig.

10: 2, 8, 12), while in L. spinosissima sporadic

unlobed half-leaves may occur (Fig. 8 : 2). Basal

branch underleaves in L. pendulina are (3)4-lobed

(Fig. 10 : 1) versus 2-3(4)-lobed in L. spinosissi-

ma (Figs. 7:11; 8:1, 11). On balance, the

strongly asymmetrical leaves of L. pendulina sug-

gest that this species belongs in subg. Notholepi-

dozia rather than in Dendrolepidozia.

Lepidozia spinosissima (Hook. f. & Tayl.) Mitt.

Figures 7 and 8

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

23

FIG. 7. Lepidozia spinosissima (Hook. f. & Tayl.) Mitt. 1. Plant showing androecia on primary and secondary

branches, the main axis with 9 androecia from ventral face of stem between arrows a and b, the plant also with a

number of androecia (not shown) between the plant base and those drawn (these basal <? not included appear older,

at times broken and presumably from a previous year's growth). 2. Main shoot with androecium, ventral view (the

leaf at lower left not shown); note also terminal branch and position of first branch underleaf. 3. Antheridium. 4.

24

FIELDIANA: BOTANY

FIG. 8. Lepidozia spinosissima (Hook. f. & Tayl.) G. L. & N. 1. Sector of upper part of main stem, ventral aspect,

with two Frullania-type branches; at arrows, first branch underleaves (BUL) (X23; 1 mm scale). 2. Sector of main

axis, with primary and secondary branches (at upper right, continuation of the attenuate branch drawn to right; note

dorsal half-leaves (HL) at arrows, and the primary branch underleaves (UL) (X23; 1 mm scale). 3. Stem leaf and

underleaves (X23; 1 mm scale). 4. Stem leaf (X23; 1 mm scale). 5. Stem underleaf (X23; 1 mm scale). 6. Leaf lobe

(XI 10; 120 /Jim scale). 7. Stem cross section (X127; 150 /Am scale). 8. Lobe base cells with oil-bodies and (upper

right) chloroplasts (x ca. 850). 9. Main stem cross-section (X180; 100 /Am scale). 10. Leaf (X20; 600 /im scale). 11.

Underleaf (X20; 600 /Am scale). 12. Lobe apex (XI 10; 120 /im scale). (Figs. 1-2 from Schuster 53380b; 3-9 from

Schuster 67-244a; 10-12, ex type.)

Portion of plant with perianth, ventral view; note primary and secondary terminal branches and the position of the

first branch underleaves. 5. Innermost bracts and bracteole. 6. Bract lobes. 7. Perianth, cross sections through median

sector (a), distal 0.2 (b) and near apex (c). 8. Portion of perianth mouth. 9. Two leaves and (at right) an underleaf

of main axis (DS = dorsal sinus). 10. Half-leaf. 11. First branch underleaves. (Figs. 1-3 from Braggins 98/31 1, New

Zealand, North Is., North Auckland Prov., E edge of Waipoua Forest, Mataraurau Plateau; 4-8 from Cameron 3797a,

New Zealand, South Auckland Prov., Coromandel Ranges, SW side of Maumaupaki, Cameron 3797a; 9-11, from

Engel 21 109, New Zealand, North Is., North Auckland Prov., SE corner of Waipoua Forest, just N of Tutamoe.)

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

25

Sendtnera spinosissima Hook. f. & Tayl. in Taylor,

London J. Bot. 5: 373. 1846. Lepidozia spinosis-

sima (Hook. f. & Tayl.) Mitt, in Hooker, Bot. An-

tarc. Voy. 2: 146. 1854. Herbertia spinosissima

(Hook. f. & Tayl.) Trev., Mem. 1st. Lomb. Sci. Lett.

III. 4: 397. 1877. Original material: New Zealand,

Edgerley (non vidi).

Lepidozia longifissa Steph., Spec. Hep. 6: 364. 1922.

Original material: New Zealand, Ziirn (non vidi).

Plants erect, rigid, with somewhat ventrally se-

cund spinescent branches, light green to yellow

brown, highly nitid when dry, the shoots to 3.2

cm wide, including branches. Branching mostly of

Frullania type, consistently bipinnate (rarely 3-

pinnate), the primary branches each usually with

1-2 strong secondary branches arising near the

base of the branch and positioned on the side of

the branch toward the shoot apex, the secondary

branches equal to or longer than primary branch-

es; branches whiplike, occasionally becoming fla-

gelliform, the leaves like those of main shoot ex-

cept smaller; branch half-leaf 2-lobed or sporad-

ically unlobed; first branch underleaf 2-3(4)-

lobed, inserted on ventral-lateral side of main axis

in similar plane to leaves of the main shoot, the

branch appearing to originate in its axil. Primary

branches with morphological ventral side oriented

toward substrate. Acromastigum-type branches

(?)occasional. Ventral-intercalary branching oc-

casional. Stems rigid, ca. 16 cells in diameter, the

cortical cells in 1 layer of somewhat elongated (at

most 2-3 X longer than wide and best viewed in

longitudinal section) thick-walled cells; medullary

cells longitudinally elongated, thick-walled, grad-

ually becoming larger in diameter toward stem

middle. Leaves rigid, plane or feebly concave, ap-

proximate, 0.7-1 mm long at longest point, 0.8-

1.3(1.5) mm wide at widest point, stiffly patent,

the insertion narrow, transverse; leaves slightly

asymmetrical, subequally deeply 4(5)-lobed, di-

vided to ca. 0.7-0.9, the lobes widely divergent,

the distance from dorsal sinus base to insertion

not much greater than that from ventral sinus to

insertion, the sinuses gradually becoming deeper

ventrally. Lobes narrowly and gradually attenuate,

not noticeably in pairs, entire, 6-8(10) cells wide

at base, terminating in a uniseriate row of (6)7-

12 cells; cells of uniseriate row thick- walled, ±

isodiametric to slightly longer than wide. Disc

moderately asymmetrical, 7-9 cells high at dorsal

sinus, 3-6 cells high at ventral sinus, the margins

entire, the dorsal margin straight to slightly con-

cave, the ventral shorter than the dorsal. Marginal

cells of disc and lobes ± isodiametric, forming an

indistinct border. Cells of disc middle thick-

walled, trigones medium and straight-sided, oc-

casionally with intermediate thickenings, isodia-

metric to longitudinally elongate along the lobe

axis (the leaves thus subvittate), (15)18-27(32) X

26-35 /im; cuticle smooth. Oil-bodies colorless,

4-10(12) per cell, relatively large (3-5 X chloro-

plasts in size), coarsely botryoidal, ovoid to ellip-

soidal, sporadically subspherical. Underleaves

widely spreading, symmetrically quadrifid to ca.

0.7—0.9, the lobes ± parallel to slightly divergent,

filiform-attenuate, usually entire, rarely with a cil-

ium toward base, terminating in a uniseriate row

of 7— 16 cells; disc 3-6 cells high at median sinus;

disc margins entire. Underleaves indistinctly bor-

dered like the leaves.

Androecia on inconspicuous, short, determi-

nate, tightly spicate, cernuous, ventral-intercalary

branches from main stem as well as primary and

secondary branches, or terminating on such

branches; bracts ventricose-cucullate, 2-lobed to

ca. 0.3-0.5, the lobes apiculate to short acuminate,

the lateral margin entire or on each with a tooth;

antherida 1(2) per bract, the stalk 8-13 cells long,

biseriate. Gynoecia rarely produced, on abbrevi-

ated ventral-intercalary branches issuing from

main stem, with 1-2 ± mature gynoecia produced

per shoot; bracts of innermost series much larger

than leaves, erect and sheathing the perianth, the

bracts deeply concave, narrowly ovate to subel-

liptical, shallowly and irregularly 3-4-lobed, the

lobes acute to acuminate, composed of ± regu-

larly rectangular and subrhomboidal cells, the api-

cal end of the marginal cells often diverging and

forming a crenulation or tooth, the lobe margins

at times with a tooth composed of a uniseriate row

of 2-3 cells, the lobe tips composed of a uniser-

iate row of 2 cells or a single cell or several lat-

erally juxtaposed cells; lamina margins composed

of narrowly elongate cells somewhat irregular in

shape, the cells of median and basal portion of

margin not projecting, the margin distally with

cells often divergent and forming a crenulation

and (sporadically) the margin with a few small

teeth; bracteole similar in size and form. Perianth

long and prominent, slenderly cylindrical-fusi-

form, terete below, obscurely trigonous above,

distinctly and deeply 3-plicate toward mouth, the

perianth narrowing toward the contracted, shal-

lowly lobulate mouth, the mouth crenulate-dentic-

ulate, the teeth composed of a single cell or a

uniseriate row of at most 3, thin to slightly thick-

ened cells, the cells of the mouth finely striate

papillose; perianth cells ± regularly long-subrect-

26

FIELDIANA: BOTANY

angular below mouth, the cells much thicker than

those of the mouth and with a smooth cuticle;

perianth (2)3-stratose near base.

Sporophyte not seen.

DIFFERENTIATION — Lepidozia spinosissima is

one of the most distinctive New Zealand species

of the genus. The erect, wiry, highly nitid (when

dry) plants, with stiffly spreading, ventrally se-

cund, spinescent branches, and the distant, rigid,

stiffly patent, transversely inserted, deeply and

symmetrically divided leaves are immediately rec-

ognizable.

DISTRIBUTION-ECOLOGY — Endemic to New

Zealand (South Is., North Is.). A species of dense,

wet, rich, low- to middle-elevation forests (typi-

cally sea level to 365 m). It typically occurs over

decaying organic matter, such as on the forest

floor among leaf litter and to a lesser extent on

old logs. When terricolous, it occurs in particu-

larly humid niches, such as damp hollows, stream

banks, moist, shaded, humus-covered banks, and

boggy areas. It also may occur as a component of

dense bryophyte cover of vertical, dripping cliffs,

such as those along the sea-level track to Bowen

Falls at Milford Sound (Engel 22010), and may

extend as high as 940 m on Mt. Te Aroha (North

Is.), where it occurs on vertical cliff faces in a

forest dominated by Nothofagus menziesii and

Griselinia littoralis (Engel 22130}.

SELECTED SPECIMENS SEEN — NEW ZEALAND.

SOUTH ISLAND. SOUTHLAND PROV.: Doubtful

Sound, Simpson s. n. — c. 6 (F); Fiordland Natl. Park,

Milford Sound, track to Bowen Falls, sea level, Engel

22012 (F); ibid., Wilmot Pass, between Lake Manapouri

and head of Doubtful Sound, Barnard NZ-49 (F); ibid.,

Stuart Mts., W shore of Lake Thomson N of stream

draining from Lake Wade, Fife 7622 (F). OTAGO

PROV.: Fiordland Natl. Park, near McKerrow Hut, head

of Lake McKerrow, Hatcher 1434 (F); just S of Haast

Pass, 540-550 m, Schuster 49648b (F). WESTLAND

PROV.: Cascade Road, Cascade ultramafic moraine, W

of Martyr Saddle, SSW of Jackson Bay, ca. 30 m, Engel

21777 (F); Jackson Bay, between confluence of Jackson

River and Arawata River and Lake Ellery, off Jackson

River Road, ca. 30 m, Child H4255 as L. microphylla

(F); Lake Ellery, ca. 30 m, Child H4573 (F); Westland

Natl. Park, along Gillespies Cook River Road, between

Tornado Creek and Whelan Creek, Engel 6595 A (F);

near Hercules Creek on Route 6 in Mount Hercules Sce-

nic Reserve, Engel 6564A (F); Arthur's Pass Natl. Park,

Otira River gorge, W of Arthur's Pass, ca. 300-365 m,

Schuster 48488c (F); ca. 1 2 mi N of Greymouth, Moore

85 as Lepidozia gottscheana (cm). NELSON PROV:

Southern end of Heaphy Track, between summit of sad-

dle beside Kohaihai Bluff and footbridge over Kohaihai

River, less than 120 m, Fife 7084 (F); Paparoa Natl. Park,

Bullock Creek Road, along Bullock Creek, NE of Pun-

akaiki, ca. 25 m, Engel 21591 (F); N bank of Fox River,

ca. 1.5 km E of Route 6, less than 30 m, Fife 4968 (F);

Stony Creek, E of Westport, ca. 150 m, Child H3549 as

L. microphylla (F); valley of the Oparawa River, on track

between logging road bridge at mouth of Narya Creek

and Moria Gate Limestone Arch, 215 m, Fife 7049.

NORTH ISLAND. WELLINGTON PROV: Akatarawa,

820 m, Braggins 84/412A—C. per. (AKU); ibid., Waterfall

Creek near Bullstream, E of Akatarawa summit, 335 m,

Braggins 84/367A (AKU); Tararua Range, Puffer to

Smiths Creek track, Moss s.n. (AKU). TARANAKI

PROV: Pukeiti Bush, near New Plymouth, Hatcher 404

(F). SOUTH AUCKLAND PROV: Mt. Te Aroha, ca. 3

km E of Te Aroha, 900-940 m, Engel 22130 (F); Cor-

omandel State Forest Park, ridge between Webb Creek

Track and Billy Goat Track, 510-540 m, Engel 22330

(F); ibid., summit of Table Mt., 835 m, Engel 22384 (F);

Coromandel Ranges, Kauaeranga Valley, Waiora Stream

Valley, ca. 200 m, Cameron 2852 (AKU); Coromandel

Ranges, SW side of Maumaupaki (Camels Back), ca.

800 m, Cameron 3797a (AKU). NORTH AUCKLAND

PROV: NE Waitakere Ranges, Spraggs Bush, ca. 360

m, Engel 22191 (F); West Auckland Waitakere Ranges,

Fairy Falls Track, ca. 320 m, Braggins 92/48c (AKU);

Kaihu Forest, headwaters of Opouteke Stream, Belling-

ham (AKU); SE corner of Waipoua Forest, just N of Tu-

tamoe, 540 m, Engel 21109 (F); Mangamuka Gorge, 245

m, Braggins 84/312 (AKU).

Subg. Mastigolepidozia Schust.

Lepidozia subg. Mastigolepidozia Schust., Beih. Nova

Hedwigia 118: 187. 2000.

Plants stiffly erect, at maturity 2-3-pinnate usu-

ally, with drooping, progressively and conspicu-

ously smaller, often distally attenuate branches;

vegetative branching uniformly of Frullania type,

remote. Stem of main axes woody, rigid, 22-25

cells in diameter, with a very weakly distinct hy-

aloderm; 2-3(4) intracortical strata smaller-celled,

firm-walled, grading into the somewhat larger-

celled, less firm-walled medulla. Leaves of stems

and branches exceedingly remote, scalelike,

broad-based, only weakly incubous, suberect, ap-

pressed or weakly spreading from stem; leaves of

main axes asymmetrical (ventral lobe smaller,

shorter), usually less than 0.45 quadrilobed, the

lobes and disc edentate; leaves of branches pro-

gressively smaller, 3-4-lobed, narrower and less

asymmetrical. Leaf cells firm-walled, with 2-5

rather large, finely botryoidal oil-bodies. Under-

leaves of main stems exceedingly remote, mostly

broader than long, quadrilobed.

TYPE — Lepidozia microphylla (Hook.) Lindenb.

Including perhaps no more than 4-5 species of

almost exclusively austral range. Only L. porto-

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

27

ricensis Fulf., which we have not studied, occurs

in the Antilles.

It is uncertain whether the few species belong

in one or two sections. The only regional species

belongs in sect. Microphyllae. Our single species

is similar to L. subdichotoma Spruce (Trans. Proc.

Bot. Soc. Edinburgh 15: 361. 1885) of Ecuador

in that, even though primary branches lie mostly

at an acute angle to the main axis (Fig. 9:1), the

primary branches typically are almost dichoto-

mously furcate and both secondary and tertiary

branches are greatly reduced in vigor compared

to the main axis. The pseudodichotomous branch-

ing of primary branches is highly distinctive. Peri-

anths in this group are very large for the overall

plant size.

Sect. Microphyllae (Lindenb.) Grolle

Sect. Microphyllae (Lindenb.) Grolle, Feddes Repert.

87: 269. 1976.

Lepidozia I. Microphyllae Lindenb. in G. L. & N.,

Syn. Hep. 201. 1845. Lectotype (fide Grolle, 1976):

Lepidozia microphylla (Hook.) Lindenb.

Lepidozia microphylla (Hook.) Lindenb. Figures

6: 10-13; 9

Jungermannia microphylla Hook., Musci Exot. 1 : pi.

80, f. 1-6. 1818. Mastigophora microphylla

(Hook.) Mont, in d'Orbigny, Voy. Amer. Mer. 7,

Bot. 2: 73. 1839. Lepidozia microphylla (Hook.)

Lindenb. in G. L. & N., Syn Hep. 202. 1845. Orig-

inal material: New Zealand, South Is., Dusky Bay,

Menzies (non vidi).

Lepidozia multipinna Steph., Spec. Hep. 3: 591. 1909.

Original material: New Zealand, Dall (non vidi).

Plants erect, rigid, distinctly wirelike, delicate

in appearance, with drooping, ventrally secund

branches, the branch axils (in field) with promi-

nent, glistening droplets of water, the plants pale

green (± strawlike color or whitish in herb); the

shoots to 4.5 cm wide, including branches.

Branching mostly of Frullania type, consistently

bipinnate (occasionally 3-pinnate), the primary

branches each usually with 2-4 strong secondary

branches arising near the base of the branch and

positioned on the side of the branch toward the

shoot apex, the secondary branches equal to or

longer than primary branches; branches whiplike,

filiform-attenuate, the leaves scalelike, obliquely

spreading, approximate; branch half-leaf often ca-

ducous; first branch underleaf 2-4-lobed, inserted

on main axis in similar plane to leaves of main

shoot, the branch appearing to originate in its axil;

primary branches with morphological ventral side

oriented toward substrate. Ventral-intercalary

branching occasional. Stems fleshy, rather brittle,

ca. 24 cells in diameter, the cortical cells in a dis-

tinct layer of isodiametric to slightly elongated

cells (best viewed in longitudinal section), larger

in diameter than subepidermal cells; medullary

cells thick-walled, the outer several rows with

walls thicker than the median core cells. Leaves

inconspicuous, scalelike, narrower than the stem,

slightly concave, markedly distant, (340) 385-

420(450) /Jim long at longest point, (340) 380-

450(490) fjim wide at widest point, strongly erect,

commonly ± appressed to stem, occasionally fee-

bly spreading, the insertion transverse to weakly

incubous; leaves asymmetrical, unequally 4-

lobed, divided to ca. 0.4-0.5 (median sinus), the

distance from dorsal sinus base to insertion great-

er than that from ventral sinus to insertion, the

sinuses gradually becoming deeper ventrally.

Lobes acuminate, not noticeably in pairs, entire,

4-5 cells wide at base, terminating in a uniseriate

row of 2-4(5) cells; cells of uniseriate row ± iso-

diametric, thick-walled; cuticle as in disc. Disc

asymmetrical, 7-9 cells high at dorsal sinus, 3-5

cells high at ventral sinus, the margins entire, the

dorsal margin straight to slightly curved, the ven-

tral shorter than the dorsal. Marginal cells of disc

± isodiametric, forming an indistinct border. Cells

of disc middle thick-walled, ± isodiametric, 24-

31 /Jim wide and long; cuticle faintly striate-pa-

pillose. Oil-bodies 2-5 per median cell, finely bot-

ryoidal and almost granular in aspect (the spher-

ules small), the oil-bodies relatively large for cell

size, 3.5 X 6.5 to, more usually, 4.5-5 X 9-1 1

up to 5 X 7-8 fjLm. Underleaves minute, widely

spreading, symmetrically quadrifid to ca. 0.3-0.4,

the lobes abbreviated, only 3-5 cells long; disc 3-

4(5) cells high at median sinus; disc margins en-

tire.

Plants dioecious. Androecia10 on inconspicu-

ous, short, determinate, tightly spicate, straight,

ventral-intercalary branches from primary or sec-

ondary branches; bracts ventricose-cucullate, 2-

lobed to ca. 0.3-0.4, the lobes acute to apiculate

to short acuminate, the lateral margins entire or

on each side with a small to large tooth, the mar-

gins often also with a few slime papillae; anther-

idia 1 per bract, the stalk biseriate. Gynoecia on

abbreviated ventral-intercalary branches issuing

from main stem; bracts of innermost series much

10 Some populations have nematode galls at the tips

of flagelliform branches; these should not be confused

with androecia (see comments under L. pendulina).

28

FIELDIANA: BOTANY

FIG. 9. Lepidozia microphylla (Hook.) Lindenb. 1. Part of erect, leafy axis, ventral aspect; at asterisks sector of

main axis; at left, part of the branching system (BL1 and BL2), near base, first two lateral primary branch leaves;

distally, BUL2 = primary appendages, underleaves, of secondary branches and BL2 = primary lateral leaf of secondary

branch; at HL portions of the half-leaves associated with Frullania-type branches visible; on right-hand side, primary

branch underleaf (BUL1) and primary branch leaf (BL1), as well as secondary branch primary leaf (BL2) labeled;

above primary appendage, an underleaf (BUL3) of tertiary branch labeled (X14.5; 2 mm scale). 2. Small sector of

main axis and bases of primary and secondary branches, dorsal aspect; stem half-leaf = HL1, half-leaf of primary

branch = HL2, first branch underleaf of primary branch = BUL1, of secondary branch, BUL2 (X29; 1 mm scale). 3-

4. Stem leaves (X50; 400 /xm scale). 5. Stem half-leaf, associated with branch (X50; 400 /urn scale). 6. Stem

underleaves (X50; 400 /Ltm scale). 7-8. Leaves from primary branches (X50; 400- /Am scale). 9. Underleaf, primary

branch (X50; 400 /tm scale). 10. Leaf from secondary branch (above) and first branch underleaf from secondary

branch (below) (X50; 400 /xm scale). 11. Underleaves from secondary branches (X50; 400 /un scale). 12. Lobe of

stem leaf (X230; 100 /xm scale). 13. Dorsal cortical cells of stem (X230; 100 /tm scale). 14. Stem cross section

(X142; 200 /xm scale). (All from Schuster 85-2719.}

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

29

larger than leaves, erect and sheathing the peri-

anth, the bracts deeply concave to canaliculate,

narrowly ovate; apices with 4 short, irregular

lobes composed of irregularly sinuate-rhomboidal

cells that at their apical end are laterally free for

varying lengths, the lobes crenate-denticulate to

ciliate; lamina composed of ± regularly subrect-

angular cells, the submarginal cells rhomboidal,

the margin bordered by elongate, thin-walled, sin-

uate-rhomboidal cells that at their apical end oc-

casionally diverge to form a projection, the mar-

gin irregularly crenate-denticulate to the base, the

teeth at most of 3 superposed cells; bracteole sim-

ilar in size and form. Perianth long and prominent,

slenderly cylindrical-fusiform, terete below, ob-

scurely trigonous above, distinctly 3-plicate to-

ward mouth, the perianth gradually narrowing to-

ward the contracted, shallowly 3-lobed mouth, the

lobes composed of irregularly sinuate-rhomboidal

cells that at the apical end are laterally free for

varying lengths, the lobes crenate-denticulate to

ciliate; perianth cells ± regularly subrectangular

below mouth; perianth 2-3 stratose near base.

Capsule oblong, the wall 54-60 /um thick, of

4-5 layers; outer layer of cells with two-phase

development, the longitudinal walls with sinuous,

sheetlike thickenings and several small nodules

alternating with walls that are devoid of thicken-

ings (or are sporadically locally thickened), the

transverse walls usually devoid of thickenings or

sporadically with an isolated nodule; innermost

layer of cells with ± tiered, narrowly rectangular,

with semiannular bands common, rather wide,

close and numerous, usually complete, rarely

forked.

Spores 10.1-11.9 /am, exine pale brown, thin,

with low, rather coarse, close papillae and short-

vermiculate markings. Elaters rigid, nontortuous,

11-14.9 /um wide, only slightly tapering toward

tips, bispiral, the spirals 3.4-4.8 /um wide.

DIFFERENTIATION — Lepidozia microphylla is im-

mediately recognizable by the erect, fleshy, pale

yellow green shoots with rigid, distinctly wirelike,

drooping, ventrally secund branches. The shoots

are rather brittle when dry. The leaves are incon-

spicuous, distant and scalelike, nearly appressed

to and narrower than the stem, and almost trans-

versely inserted (Fig. 9 : 1, 2). A distinctive field

character of the species is the presence of tiny,

glistening water droplets in the branch axils.

A notable feature of this species is the "low"

insertion of the basal branch underleaf of primary

branches: this structure, usually trifid, superficial-

ly appears to be inserted on the stem rather than

the branch base (Fig. 9 : 1, 2).

Although L. microphylla and its immediate al-

lies of the Neotropics are very distinct plants veg-

etatively, the gynoecium — in particular the $

bracts — is very similar to what is seen in ordinary

taxa of subg. Lepidozia and Notholepidozia.

Lepidozia microphylla is apt to be confused in

the field with L. spinosissima owing to the stiffly

erect growth, the copious branching, and the pale

yellowish color. Both taxa tend to occur in large,

often pure swards, are ± similar in size, with the

well-separated individual plants having a dendritic

aspect.

The leaf of L. microphylla is similar to that of

the otherwise very different L. pendulina in being

strongly asymetrically 4-lobed (Fig. 9 : 3, 4). The

leaf, however, is much smaller, averaging 385-

415 jam broad X 385-410 /am long. The ventral

lobe is shaped much like the others and is not set

off as a narrow "finger." The distal portions of

the leaf are distinctly smaller-celled; cells are ca.

(20)23-25 X 24-32 /um versus 33-37 X 44-52

/urn in a small basal region. Median cells (Fig. 6 :

10) have a conspicuous middle lamella, are evenly

thick-walled, and have mostly 2-5 oil-bodies

each. The oil-bodies are clearly less coarsely bot-

ryoidal — almost granular-botryoidal — than in L.

pendulina and L. concinna (compare Fig. 6 : 1,5)

and, in median cells, range from 3.5 X 6.5 to 4.5-

5 X 9-1 1 /urn, or 5 X 7 /am.

Underleaves are distinctive in being broader

than long, rather shallowly 4-lobed, with narrow

lobes, unusual in the relatively conspicuous and

persistent apical slime papilla (Fig. 6 : 11).

The stem (Fig. 9:14) has a rather well-defined

hyaloderm, surrounding a layer 1-2(3) strata thick

of somewhat smaller, very thick-walled cells that

gradually grade into a firm-walled medulla.

DISTRIBUTION-ECOLOGY — Endemic to New

Zealand (South Is., North Is.). A species of dense,

wet, rich, lower-elevation forests (sea level to 340

m). It typically occurs on the forest floor, where

it may form extensive, dense carpets. It is a lo-

cally common floor plant at some sites, such as

rich mixed Nothofagus forests or kakikatea

swamps, and often occurs in boggy, seepage nich-

es. To a lesser extent it may also be found on old

logs or tree bases (such as of kauri).

SELECTED SPECIMENS SEEN — NEW ZEALAND:

SOUTH ISLAND: SOUTHLAND PROV.: Lake Hau-

roko, ca. 215 m, Child 1538 (F); Doubtful Sound, Simp-

son s. n. (F); Fiordland Natl. Park, Tutoko River, W of

30

FIELDIANA: BOTANY

Milford Sound, 50 m, Engel 18820 (F). WESTLAND

PROV: Cascade Road, just W of Jackson River, ca. 8-

12 km SW of confluence of Jackson and Arawata Riv-

ers, 25-90 m, Engel 22970 (F); Lake Ellery, Child

H4571 (F); between confluence of Jackson River and

Arawata River and Lake Ellery, off Jackson River Road,

sea level, Engel 17913 — c. S + sporo. (F); Haast, 3 mi.

N of bridge, ca. 15 m, Child H702 (F); Lake Matheson,

sea level, Child H371 (F); Westland Natl. Park, along

Gillespies Cook River Road, between Tornado Creek

and Whelan Creek, Engel 6578A (F); Gillespies Beach

Road, 12 km W of Fox Glacier, ca. 150 m. Child

H4849 — c. 6 + sporo. (F); Mahinapua, S of Hokitika,

ca. 60 m. Child H3668 (F). NELSON/WESTLAND

PROV. BOUNDARY: Porarari River Gorge, junction of

Porarari River Gorge, junction of Porarari River Track and

the Inland Pack Track, ca. 30 m, Fife 4850 (F). NELSON

PROV.: Little Wanganui, W coast, ca. 15 m, Child 416

(F); valley of the Oparara River, 0.5 km S of Moria Gate

Limestone Arch, 226 m, Fife 7073 (F); Paparoa Natl.

Park, Inland Pack Track, SW of terminus of Bullock

Creek Road, NE of Punakaiki, ca. 35 m, Engel 21675 (F);

swamp at N end of Lake Hanlan and gorge of stream

draining lake and entering Tidal Creek between lake and

State Highway 67, 30-90 m, Fife 7142 (F). NORTH IS-

LAND: NORTH AUCKLAND PROV: Waipoua Forest,

track to Te Matua Ngahere, ca. 340 m, Engel 22561 (F);

SE corner of Waipoua Forest, just N of Tutamoe, Engel

21086 (F); same loc., Braggins 94/246 — c. sporo. (AKU);

Omahuta Forest Kauri Sanctuary, E of Mangamuka

Bridge, 260 m, Engel 21001— c. per. (F).

Subg. Notholepidozia Schust., subg. nov.

Subgeneri Lepidoziae similis sed corporibus

oleosis distincte vel grosse botryoideis, inflores-

centiis uniformiter dioicis, axe e cellulis intracor-

ticeis rigidis atque parietibus firmis vel percrassis

composite differt.

TYPE — Lepidozia novae -zelandiae Steph.

Plants typically creeping to ascending, often

loosely procumbent, occasionally erect, light yel-

lowish or even whitish green, ranging to olive to

brownish, but never glaucous, the cells lacking a

waxy cuticle. Branching l(2)-pinnate (consistent-

ly bipinnate only in L. pendulind), often regularly

so, the branches often running out into flagellae;

lateral branching always Frullania type; ventral-

intercalary vegetative branches usually few or

none, but 6 and 9 branches normally from axils

of underleaves of leading stems or primary

branches. Stems not woody (exc. L. setigera),

mostly with a 1-stratose cortex of larger cells (at

times verging on an indistinct hyaloderm) which

have only moderately thickened or almost thin

walls; medullary cells smaller, often thick-walled.

Leaves remote to imbricate, typically strongly in-

cubously inserted and oriented (exc. L. penduli-

na), often handlike, distally decurved, strongly

(infrequently weakly) asymmetrically (3)4(5)-

lobed; leaves with both disc and lobes usually

asymmetrical, the dorsal sectors larger and longer,

the dorsal margin longer, usually more convex

(sometimes strongly ampliate); leaves usually

0.3-0.5 lobed, with narrowly triangular to longly

acuminate lobes, the lobe (and disc) margins en-

tire or toothed with coarse, broad-based, teeth

which are never opposed in pairs. Cells of leaves

usually firm-walled, often quite thick-walled and

with lumina rounded at the angles, less often with

distinct trigones. Oil-bodies in distal and median

cells usually 2-5 per cell, finely to coarsely bot-

ryoidal. Underleaves smaller than lateral leaves,

usually 3-4-lobed.

Subgenus Notholepidozia is very well repre-

sented in New Zealand and is strongly polytypic

there. Species occur from sea level well into the

alpine region, where they occur mostly on soil in

clefts between ledges or crevices in cliff faces, but

also sometimes in moory areas or in snow-tussock

grasslands. The species usually grow procumbent

when not crowded and almost never occur stiffly

erect as do the taxa of subg. Dendrolepidozia.

Most of the species are easily known by their pre-

dominantly 4-lobed, almost handlike, asymmetri-

cal leaves, with larger dorsal lobes.

We have, with some reluctance, concluded that

the Austral taxa here placed into Notholepidozia

must be segregated as an autonomous subgenus

from subg. Lepidozia, as typified by L. reptans.

All of the taxa of Notholepidozia that we have

seen have distinctly botryoidal oil-bodies (Figs.

31 : 11, L. kirkii; 6 : 1, 2, L. pendulina; 6 : 5, 6,

L. concinna), while L. reptans and two other

Northern Hemisphere species have small, homo-

geneous oil-bodies (cf. fig. 86 : 6, Schuster, 1969).

Lepidozia reptans has a relatively simple axial

anatomy, with a leptodermous medulla; it is au-

toecious. All of the Austral species we have stud-

ied from living plants consistently have botryoidal

oil-bodies, have a stem with a firm-walled or rigid

medulla, and are unisexual.

Lepidozia reptans in many respects is not typ-

ical of the genus; it is a pity that it is the generic

type because it is a thoroughly "strange" element

within Lepidozia. Oil-bodies in L. reptans are

small, smooth, and numerous, occurring (8)10-

16(25) per cell versus the botryoidal oil-bodies,

often larger, found usually (1)2-8 per median cell

in Notholepidozia. In the botryoidal oil-bodies

Notholepidozia is similar to the two Austral sub-

genera, Dendrolepidozia (cf. Fig. 8 : 8, L. spinos-

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

31

issima) and Austrolepidozia (cf. Fig. 6 : 10, L. mi-

crophylld). This suggests that botryoidal oil-bod-

ies are part of the original genetic "equipment"

of the genus as a whole and that L. rep tans rep-

resents, geographically and cytologically, an iso-

lated extreme within the genus as a whole.

Lepidozia reptans is, as far as known, wholly

Laurasian in range. Early reports (from Java, Juan

Fernandez, Hawaii, South America; Frye & Clark,

1937_47) are all erroneous (Schuster, 1969, p.

25). The report of the species from as far south

as Colombia and Venezuela (Fulford, 1966) is al-

most surely incorrect; pi. 40 : 5 in Fulford (I.e.)

illustrates a plant with a high disc and short lobes,

whereas L. reptans s. str. has a lower and more

asymmetrical disc (cf. fig. 86 : 3, 4, Schuster I.e.).

Sect. Pendulinae Schust.

Lepidozia sect. Pendulinae Schust., Beih. Nova Hed-

wigia 118: 197. 2000.

Plants firm, erect, vigorous, 2(3)-pinnate, the

branches drooping, ventrally secund, freely be-

coming flagelliform. Basal branch underleaf in-

serted at branch base, usually deeply quadrifid.

Stem woody, with up to (5)6-7(8) thick-walled

strata of relatively small cells, the outer layer not

or hardly distinguishable from the inner ones; me-

dulla larger-celled, the cells rather thin-walled.

Leaves asymmetrical, almost transversely insert-

ed, concave, quadrifid, the two dorsal lobes larger,

separated by a shallower sinus; ventral lobes nar-

rower, lanceolate, separated from the dorsal two

by a deeper sinus.

TYPE — Lepidozia pendulina (Hook.) Lindenb.

Including a single regional species. When well-

developed, the freely bipinnate, sporadically tri-

pinnate, tapered, and ventrally secund branching

separates the species from any other regional tax-

on of subg. Notholepidozia. Branching is sugges-

tive of Dendrolepidozia, but the total lack of Ac-

romastigum-type branches separates L. pendulina

from L. (D.) spinoss issima. On well-developed

plants, branches, apices of uppermost branches,

and the shoot tip are diagnostically ventrally

strongly secund or even coiled, unlike in L. spi-

nosissima. The sporadic ventral branches in L.

pendulina are all intercalary and axillary.

The strongly asymmetrical leaves of L. pendu-

lina consist of two similar dorsal lobes that are

separated by a relatively shallow sinus; the dorsal

pair of lobes is separated by a deeper sinus from

the two narrow, lanceolate, longly tapered ventral

lobes. Ventral lobes typically are separated from

each other by a very deep sinus that often de-

scends for 0.75-0.85 the leaf length (cf. Fig. 10 :

3,4). Cells in L. pendulina at leaf bases may have

as many as 16-20 oil-bodies (Fig. 6:2).

Lepidozia pendulina (Hook.) Lindenb. Figures 6 :

1-3, 8, 9; 10

Jungermannia pendulina Hook., Musci Exot. 1 : pi.

60, f. 1-5. 1818. Lepidozia cupressina var. T pen-

dulina (Hook.) Lehm. & Lindenb. in Lehmann,

Nov. Min. Cogn. Stirp. Pug. 4: 39. 1832. Lepidozia

pendulina (Hook.) Lindenb. in G. L. & N., Syn.

Hep. 208. 1845. Mastigophora pendulina (Hook.)

Trev., Mem. 1st. Lomb. Sci. Lett. III. 4: 416. 1877.

Original material: New Zealand, South Is., Dusky

Bay, 1791, Menzies (non vidi).

Lepidozia gigantea Steph., Spec. Hep. 3: 600. 1909.

Original material: New Zealand, Doll (non vidi).

Plants erect, rigid, with ventrally secund

branches, pale green, the shoots robust, to 6 cm

wide, including branches. Branching mostly of

Frullania type, consistently bipinnate (rarely 3-

pinnate), the primary branches each usually with

1-2 strong secondary branches arising near the

base of the branch and positioned on the side of

the branch toward the shoot apex, the opposing

side of the primary branch with 0-2(3) branches,

the secondary branches equal to or longer than

primary branches; branches whiplike, often be-

coming flagelliform, the leaves like those of main

shoot except smaller; branch half-leaf 2-lobed or

sporadically unlobed; first branch underleaf (3)4-

lobed, inserted on ventral-lateral side of main axis

in a plane similar to leaves of main shoot, the

branch appearing to originate in its axil; primary

branches in suberect to erect plants with morpho-

logical ventral side oriented toward substrate.

Ventral-intercalary branching rare and sporadic.

Stems rigid, ca. 30 cells in diameter, the cortical

cells in 1 layer of isodiametric, firm-walled cells

(best viewed in longitudinal section), internal to

the epidermal layer are 2-3 to 4-6 rows of thick-

walled, longitudinally elongated cells; central por-

tion of stem made up of cells larger in diameter

and with thin walls. Leaves rigid, slightly con-

cave, distant, 1.1-1.4 mm long at longest point,

(1.7)1.9-2.5 mm wide at widest point, stiffly pat-

ent, the insertion transverse; leaves asymmetrical,

unequally 4(6)-lobed, the leaves divided to ca.

0.65-0.75 (median sinus), the distance from dor-

sal sinus base to insertion much greater than that

from ventral sinus to insertion. Lobes broadly at-

32

FIELDIANA: BOTANY

FIG. 10. Lepidozia pendulina (Hook.) Lindenb. 1. Part of plant, ventral view; main axis at asterisks; at BUL1 the

primary appendages, underleaves. of primary branches; at BL1 primary lateral leaves of branches; at BUL2 primary

underleaf of secondary branch, and distal to it (at 3) succeeding branch underleaf; at HL, stem half-leaves; the

secondary branch at upper right with sector removed (arrows), the branch so curved that underleaves (UL) are not

perfectly aligned; at asterisks, the distal sectors join together (X12; 2.5 mm scale). 2. Small sector of main axis,

dorsal aspect; two Frullania-type branch bases (FB), with the bifid half-leaves (HL) and primary branch appendages

(BUL1) labeled. Note the juxtaposition (upper branch base) of half-leaf and primary branch underleaf, so that the

branch superficially appears to arise in an axillary position. At SUL the stem underleaf visible, in part; note the

subtransverse insertion of dorsal halves of leaves (X15.5; 1 mm scale). 3. Stem leaf (X 17; 1.5 mm scale). 4. Stem

leaves (X14; 2 mm scale). 5. Stem underleaves (X14; 2 mm scale). 6. Leaves of primary branches (X17; 1.5-mm

scale). 7. Underleaves, primary branch (X17; 1.5 mm scale). 8. Stem half-leaf, primary branch (X17; 1.5-mm scale).

9. Initial branch underleaves from primary branches (BUL1 in Figs. 1-2), showing variation in lobe number (X14; 2

mm scale). 10-11. Appendages from secondary branches; above, two underleaves; below, two leaves (X17; 1.5 mm

scale). 12. Half-leaf of stem (X14; 2 mm scale). 13. Dorsal cortical cells (X160; 150 /urn scale). 14. Cells of leaf

lobe (x 160; 150 /*m scale). 15. Sector of stem cross section (X 160; 150 /im scale). (All from Schuster 67-400, New

Zealand, Fiordland, Falls Creek.)

ENGEL & SCHUSTER: AUSTRAL HEPATIC AE. LEPIDOZIA

33

tenuate (the dorsal lobes acute, the ventral lobe

more narrowly attenuate and curved), widely di-

vergent, the dorsal pair of lobes partially united,

the others distinct and spreading, the ventral lobe

often widely divergent as a "claw," the dorsal

lobes entire, (10)14-24 cells wide at base, the

lobes terminating in a uniseriate row of 2-4(6)

cells (6-7 in ventralmost lobe); cells of uniseriate

row ± isodiametric to slightly longer than wide,

thick-walled. Disc distinctly asymmetrical, 20-30

cells high at dorsal sinus, 12-18 cells high at ven-

tral sinus, the margins entire, the dorsal margin

broadly ampliate, the ventral much shorter than

the dorsal, slightly curved. Cells of disc middle

thick-walled, trigones medium and straight-sided,

occasionally with intermediate thickenings, ± iso-

diametric to elongate, 18-26(29) X 24-33(42)

fim, in ± regular longitudinal files; median basal

cells enlarged, in 1-2 rows, with walls often nod-

ulose; cuticle smooth to indistinctly striate. Oil-

bodies in median cells (2)3-8, locally to 10-14

per cell, rather coarsely botryoidal, spherical to

ovoid, 4 X 4-5 /tm to 4-4.5 X 5-7 /xm, sporad-

ically 3.6 X 4.5 yam; basal cells locally large and

40-42 X 50-59 /xm and with 16-20 oil-bodies

each, coarsely segmented, 5-7 ^m to 5.5 X 10

/jim. Underleaves often weakly to distinctly pli-

cate, widely spreading, symmetrically 4(6)-fid to

ca. 0.65-0.7 (median sinus), the sinus bases some-

what reflexed, the lobes ± parallel, somewhat

ventrally sulcate, narrowly and gradually attenu-

ate, entire, terminating in a uniseriate row of 1-

3(4) cells; disc 14-24(32) cells high at median

sinus, the margins reflexed, entire or sporadically

with a tooth.

Plants dioecious. Androecia" on inconspicu-

ous, short, determinate, tightly spicate, cernuous

ventral-intercalary branches from main shoot and

primary + secondary branches (in flagelliform or

leafy sectors); bracts ventricose-cucullate, 2-lobed

to ca. 0.3, the lobes acute to short acuminate, the

lateral margin with several slime papillae; anther-

idia 2 per bract, the stalk biseriate. Gynoecia only

sporadicaly produced, on abbreviated ventral-in-

11 Some populations have nematode galls at the tips

of primary and secondary branches. These are conspic-

uous, budlike, and spherical, with several nematodes in

each swelling. The nematodes considerably alter the

morphology of the leaves to structures that are in ca. 2-

3 pairs, cucullate, 4-lobed to ca. 0.2-0.3 and with lobes

acute to short acuminate. The budlike swellings should

not be confused with androecia. Nematode galls occur

in a variety of Hepaticae but heretofore are unknown for

Lepidoziaceae (cf. also Schuster, 1966).

tercalary branches issuing from main stem; bracts

of innermost series much larger than leaves, erect

and sheathing the perianth, the bracts deeply con-

cave, broad ovate-subrectangular; apices with 4-

5 short, ± regular lobes composed of ± regularly

rectangular and (especially at the lobe extremities)

irregularly sinuate-rhomboidal cells, the apical

end of the marginal cells often thickened and fee-

bly diverging and forming a crenulation, the lobes

thus finely crenulate; lamina composed of ± reg-

ularly subrectangular to prosenchymatous cells,

the margin bordered by thin-walled cells of vari-

able shape and orientation, the cells often ± sin-

uate-rhomboidal, the apical or free end of margin-

al cells variously divergent and forming a short

projection or a tooth, the margin irregular and

crenulate to crenate-denticulate, the margins oc-

casionally with a ciliiform-laciniiform process;

bracteole similar in size and form. Perianth long

and prominent, slenderly cylindrical-fusiform,

slightly curved, terete below, obscurely trigonous

above, distinctly and deeply 3-plicate toward

mouth, the perianth gradually narrowing toward

the strongly contracted, shallowly 3-lobed mouth,

the lobe margins composed of irregularly sinuate-

rhomboidal cells that, at the apical end, project

for varying lengths; mouth cells laterally free for

varying lengths and only rarely completely free

laterally, the mouth thus crenate-denticulate; peri-

anth cells ± regularly subrectangular to feebly

prosenchymatous below mouth, and here the cu-

ticle is obscurely striate; perianth 3-4 stratose

near base.

Seta with 16 rows of outer cells surrounding an

inner core of numerous much smaller cells. Ma-

ture capsules not seen.

DIFFERENTIATION — Well-developed plants of

this species are not likely to be confused with any

other New Zealand member of the genus on gen-

eral aspect alone. However, small, poorly devel-

oped populations are often difficult to recognize.

Such plants are compact, with secondary branches

weakly developed or absent, and have imbricate

leaves. In these cases the broad, transversely in-

serted, entire-margined leaves (at least near the

base of best-developed shoots), the narrow me-

dian disc cells, and the large underleaves with ±

parallel, ventrally sulcate lobes will generally dis-

tinguish this species. Absence of pigments and the

lack of underleaf armature will aid in distinguish-

ing the species from L. obtusiloba. Lepidozia con-

cinna is sometimes suggestive of a diminutive L.

pendulina in the shape of its leaves and under-

34

FIELDIANA: BOTANY

leaves (as, for example, in the type of L. latiloba

Col.) but typically has at least a moderately in-

cubous leaf insertion, much larger median disc

cells, and dorsal lobes that are at most 9 cells wide

at the base.

The leaf areolation of L. pendulina is distinc-

tive: The leaf cells are thick-walled, arranged in

± regular longitudinal files, and appear small and

narrow in proportion to the size of the leaf, giving

the leaf a densely cellular appearance, even under

the dissecting microscope.

Vigorous plants of L. pendulina may have very

rigid main stems, with 6-8 outer cell layers form-

ing, collectively, a rigid cortex (Fig. 10 : 15). Al-

though the main shoots are moderately aniso-

phyllous, at least the secondary branches may be-

come virtually isophyllous (Fig. 10 : 1).

DISTRIBUTION-ECOLOGY — New Zealand (South

Is., North Is.), Tasmania, Australia (Victoria).

The species occurs in dense, very wet forests

over a broad altitudinal gradient but is more typ-

ical of middle- to upper-elevation sites. For ex-

ample, it may occur at the upper limits of Noth-

ofagus menziesii forests (stations at ca. 900 m on

South Is. and ca. 1180 m on North Is.), as well

as in subalpine Nothofagus solandri forests with

Phyllocladus alpinus, Senecio bennettii, Pseudo-

panax spp., Dracophyllum traversii, and Copros-

ma (915 m, Fife 8005). In the southern sectors of

South Is, it may occur in forests at lower eleva-

tions, including at sea level. For example, it is

found on the sides of bryophyte-covered logs in

the rich mixed Nothofagus forest with tree fern

understory along the Tutoko River at 50 m in

Fiordland.

The species typically occurs over organic sub-

strates such as the forest floor (particularly in

damp leaf litter), where it may form extensive, at

times pure, carpets; it may be associated with Tri-

chocolea mollissima, Plagiochila suborbiculata,

P. gigantea, Schistochila nobilis, and Bazzania ni-

tida, among others. It also occurs on damp, bryo-

phyte-covered banks and wet, bryophyte-covered,

rotted logs. The species often is a component of

the flora of sites where the floor, boulders, and

trees are densely covered with bryophytes. At the

upper forest limits smaller phases of the species

may form extensive pure populations on rotted,

decorticated wood (Moraine Creek Track, Engel

23218) or on old tree stumps (Punekiri Bluff, En-

gel 23321).

SELECTED SPECIMENS SEEN— NEW ZEALAND.

SOUTH ISLAND. SOUTHLAND PROV.: Lake Hau-

roko, ca. 215-305 m, Child HI 547, HI 59 1 (F); Fiord-

land Natl. Park, Expectation Stream, Zotov s. n. (F);

ibid., Stillwater River, Zotov s. n. (F); ibid., Stuart Mts.,

W shore of Lake Thomson N of stream draining from

Lake Wade, 300 m, Fife 7722 (F); immediately N of Ten

Mile Bush, near W shore of Lake Te Anau and ca. 20

km N of town of Te Anau, 200 m, Engel 23203 (F);

Fiordland Natl. Park, off track along East Branch of

Eglinton River, SE of Mt. Eglinton and NNE of Te

Anau, 440 m, Engel 18773— c. S (F); ibid., Cascade

Creek, near Lake Gunn, tributary of Eglinton River, ca.

365 m, Child H2591 (F); ibid., Moraine Creek Track,

area N of Moraine Creek, W of Hollyford River, 610m,

Engel 23218 (F); ibid., Moraine creek, Hollyford Valley,

ca. 120-200 m, Schuster 48111 (F); ibid., Tutoko River,

W of Milford Sound, 50 m, Engel 18813 (F). OTAGO

PROV.: Fiordland Natl. Park, track along Fails Creek,

near junction of Hollyford River road and road to Mil-

ford Sound, Schuster 67^t06a (F); Route Burn Valley,

ca. 520 m, Child 2460 (F); Sylvan Lake, Dart River, ca.

365 m, Child HI 167 (F); Beans Burn, tributary of Dart

River, 455 m, Child 1267 (F); Fiordland Natl. Park, trail

between Gunn's Hut and Hidden Falls, ca. 30 mi SE of

Lake McKerrow, Hatcher 696 (F); ibid., head of Lake

McKerrow, Hatcher 1338 (F); ibid., N of McKerrow

River, Martins Bay, Hatcher 749 (F); ibid., Martins Bay,

Hatcher 1375A (F). OTAGO/WESTLAND PROV.

BOUNDARY: Summit area of Haast Pass, 570 m, Engel

17963 — c. S (F); Haast Pass watershed, ca. 455 m, Child

s.n. 48 (F). WESTLAND PROV: Cascade Readjust NE

of Martyr Saddle and immediately N of Jackson River,

ca. 20 m, Engel 21835A (F); ibid., just W of Jackson

River, ca. 8-12 km SW of confluence of Jackson and

Arawata Rivers, 25-90 m, Engel 22994 (F); S facing

upper reaches of Haast River, ca. 0.6 miles N of Haast

Pass, ca. 520 m, Schuster 59616a (F); Mt. Aspiring Natl.

Park, Cross Creek, 1.1 km N of Haast Pass, 540 m,

Engel 21881 (F); ibid., Blue River near confluence with

Makarora River, NNE of Makarora, 310m, Engel 18912

(F); ibid., off track to Mt. Brewster, below and W of Mt.

Armstrong, SW of Mt. Brewster, ca. 900 m, Engel 17842

(F); Lake Ellery, ca. 60 m. Child H4574 (F); between

confluence of Jackson River and Arawata River and

Lake Ellery, off Jackson River Road, sea level, Engel

17915 (F); near Fox Glacier, Mrs. Knight (F); N side of

Waiho River, within 0.8 mi. of jet. with Gallery River,

ca. 2-3.5 mi. below Franz Josef Glacier, 75-250 m,

Schuster 59729a (F); Arthur's Pass Natl. Park, N of Kel-

lys Creek near Hwy 73, above campground, 420-475

m, Engel 18353 — c. 6" (F); Otira, track to Mt. Barron,

Whitehouse 29641 (F); Camp Creek, W of Alexander

Range, 510-1040 m, Reif C63D, C216C (F). WEST-

LAND/CANTERBURY PROV. BOUNDARY: Arthur's

Pass Natl. Park, Bealey Valley Track, ca. 875-900 m,

Engel 22854 (F). CANTERBURY PROV: Arthur's Pass

Natl. Park, near Bealey Glacier Vista, Engel 6839 (F);

ibid., S bank of upper reaches of Bealey River near Be-

aley Track (to Bealey Glacier), within 1.5 km NW of

Jack's Hut, 875-950 m, Fife 6145 (F); Margaret's Tarn

(Bealey Glacier Track), 915 m, Fife 8005 (F); Arthur's

Pass vicinity, 1/4 mile W of Bealey Valley, Whitehouse

29585 (F). NELSONAVESTLAND PROV. BOUND-

ARY: Porarari River Track. 4 km from road, at junction

with Bullock Creek Track, ca. 150 m, Child H4978 (F).

NELSON: Mt. Sewell. Paparoa Range, ca. 915 m, Child

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

35

H2783 (F); Paparoa Mts., cirque on E flank of Mt. Priest-

ly, 960 m, Fife 5573 (F). NORTH ISLAND: WELLING-

TON PROV.: Erua (South of National Park), Erua Forest

2 km from State Highway 4, Braggins 92/129 (F); Ton-

gariro Natl. Park, ca. 8 km from Ohakune on Ohakune

Mt. Road, ca. 950 m, Engel 21334— c. <S (F); Tongariro

Forest, Okupata Caves site, ca. 600 m, Braggins 92/136

(F). NEAR GISBORNE/SOUTH AUCKLAND PROV.

BOUNDARY: Urewera Natl. Park, track to Whakataka

Hut from Lake Waikaremoana, N of western extremity

of Lake Waikaremoana, 1000-1 100 m, Engel 23311 (F).

GISBORNE PROV.: Urewera Natl. Park, Panekiri

Range, summit area of Pukenui in vicinity of Punekiri

Bluff, 1180 m, Engel 23321 (F); ibid., Huiarau Range,

summit area of Te Rangaakapua, 1265-1320 m, Engel

23409 (F).

Sect. Setigerae Schust.

Lepidozia sect. Setigerae Schust., Beih. Nova Hed-

wigia 118: 193. 2000.

Plants closely 1-, sporadically l(2)-pinnate,

erect or suberect, pale yellowish or whitish green,

vigorous. Stems rigid, with a hyaloderm moder-

ately distinct, the 2-3 cell layers within firm-

walled, grading into the medulla, which is less

strongly thick-walled (in section often strongly

thickened only at the angles). Leaves strongly in-

cubous, imbricate, strongly asymmetric, 4(5)

lobed (dorsal two lobes shorter, separated by a

shallower sinus than ventral 2-3 lobes), with

strongly ampliate dorsal margin, and a short, non-

ampliate ventral margin; lobes acuminate, ending

in up to 6-10(12) superposed, elongated cells.

Underleaves often connate on one side, cuneate-

based, obtrapezoidal, 4-6-lobed, the lobes acu-

minate and setigerous like the leaf lobes.

Gynoecial bracts in 3-4 progressively larger se-

ries, at least 0.5 quadrifid into narrow, erect, grad-

ually tapered lobes that end in ca. 9-13 super-

posed single cells.

TYPE — Lepidozia setigera Steph.

Monotypic. Without close allies. In dorsal as-

pect the leaves appear conspicuously imbricate on

account of the broadly overlapping ampliate dor-

sal margin of the disc (Fig. 11 : 1). On main

shoots the stem is wholly hidden in dorsal aspect.

Both leaves and underleaves may show elabora-

tion of a strong accessory lacinium, so that seem-

ingly 5-lobed leaves (Fig. 11 : 3, 4) and 5-6-lobed

underleaves (Fig. 11 : 5, 6) frequently occur. Bas-

al branch underleaves are oriented in line with

stem lateral leaves and the branches superficially

seem to originate in their axils (Fig. 11 : 2). The

first branch underleaves are large, often trifid, at

times with one lobe seemingly again bifid, or

bearing a strong cilium (Fig. 11 : 7).

Lepidozia setigera is a strikingly isolated spe-

cies. Unlike ordinary species of the genus, which

have shallowly lobulate bract apices, it has the

bracts basically deeply incised, for at least 0.5

their length, into narrowly lanceolate-caudate

lobes that end in setae formed of 9-13 cells. (Fig.

12: 12).

Lepidozia setigera Steph. Figures 11 and 12.

Lepidozia setigera Steph., Spec. Hep. 3: 599. 1909.

Original material: New Zealand, South Is., Kirk

(non vidi).

Plants suberect, somewhat flexuous, with ven-

trally secund branches, pale green, the shoots ro-

bust, to 3 cm wide, including branches. Branching

mostly of Frullania type, closely and regularly

pinnate, the primary branches often very long, be-

coming whiplike and flagelliform, the primary

branches occasionally developing into new lead-

ing shoots; secondary branches occasionally pre-

sent, 1(2) per primary branch; branch half-leaf ±

symmetrical, broadly ovate, 2-lobed to ca. 0.5, en-

tire; first branch underleaf 3-4-lobed, with a few

marginal cilia, inserted on lateral side of stem im-

mediately below branch base and aligned with

leaves of main shoot, the branch appearing to

originate in its axil; second and subsequent branch

underleaves ventral-lateral in position and gradu-

ally becoming morphologically ventral. Ventral-

intercalary branching sporadic, stoloniform or

leafy and developing into a new leading shoot.

Stems stiff and woody, ca. 24 cells in diameter,

the cortical cells in 1 layer of thin-walled cells

distinctly larger than medullary cells (up to 2X

larger); medullary cells moderately thickened, the

outer several rows with walls thicker than the me-

dian core cells. Leaves rigid, slightly concave, the

lobes in same plane as lamina and not incurved,

strongly imbricate and completely hiding stem in

dorsal view, 1.5-1.7 mm long at longest point, 2-

2.4 mm wide at widest point, patent, the insertion

distinctly incubous and somewhat recurved at dor-

sal end; leaves distinctly asymmetrical, unequally

4(5)-lobed, the leaves divided to ca. 0.5-0.6 (me-

dian sinus), the distance from dorsal sinus base to

insertion greater than that from ventral sinus to

insertion, the sinuses ± U-shaped, gradually be-

coming deeper ventrally. Lobes subulate, the

lobes somewhat divergent, not noticeably in pairs,

the ventral lobe often spreading as a "claw," the

36

FIELDIANA: BOTANY

030002!

RT;.

FIG. 1 1. Lepidozia setigera Steph. 1. Shoot sector, main axis, with base of lateral branch at left, dorsal view. At

HL the stem half-leaf; at UL1 the primary branch underleaf, trifid in form; at BL the branch leaf (X20; 1.5 mm scale,

adjacent). 2. Shoot sector, ventral aspect. At SU, stem underleaf; at BUU, the trifid primary branch underleaf, the

branch superficially axillary in it; at HL, the stem half-leaf (X17.5; 1.5 mm scale, adjacent). 3-5. Two stem leaves

and an underleaf (X17.5; lower 1.5 mm scale). 6. Branch underleaf (X17.5; lower 1.5 mm scale). 7. First branch

underleaf (X17.5; lower 1.5-mm scale). 8. Half-leaf associated with Frullania-type branch (X17.5; lower 1.5 mm

scale). 9, 11. Underleaf lobes (X90; 200 /Am scale). 10. Dorsal lobe at sinus, stem leaf; note small cells in sinus

(X90; 200- /im scale). 12, 13. Basal and median leaf cells, respectively (X90; 200 /tm scale). 14. Part of stem cross

section (X198; 150 /xm scale). (All from Schuster 67-400, New Zealand.)

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

37

10

FIG. 12. Lepidozia setigera Steph. 1. Median leaf cells (X220; 100 ^tm scale at right). 2. Foot and seta base,

excised sporophyte (X25; 700 /j.m scale). 3. 9 Bracts + bracteole, innermost series (X17.5; left 1.5 mm scale). 4.

9 Bract, second series (X17.5; left 1.5 mm scale). 5. 9 Bract, outermost series (X17.5; left 1.5 mm scale). 6.

Perianth-bearing ventral-intercalary branch (X22; lower 1.5 mm scale). 7. Stem leaf (X17.5; left 1.5 mm scale). 8.

Stem underleaf (X17.5; left 1.5 mm scale). 9. Stem half-leaf (X17.5; left 1.5 mm scale). 10. Stem half-leaf (X 17.5;

left 1.5 mm scale). 11. Cross section, immature seta (XI 10; 200 yum scale). 12. Apex of lobe of 9 bract (X185; 150

/mi scale). (All from Schuster 95-750, New Zealand, North Is.)

38

FIELDIANA: BOTANY

lobes entire, the dorsal lobes 8-1 1 cells wide at

base, terminating in a uniseriate row of 6-10(12)

cells (the ventral often somewhat longer); cells of

uniseriate row, often hyaline, tapering, with ±

swollen dilated septa, 2-5:1, thick-walled, becom-

ing longer and narrower toward the tip and cap-

illary, the terminal cells finely tapering, 8-10:1,

the cuticle finely striate-papillose. Disc distinctly

asymmetrical primarily because of strong dilation

of dorsal sector, the dilated portion up to ca. 0.5

the disc area, the disc ( 1 8)20-25 cells high at dor-

sal sinus, 12-16 cells high at ventral sinus; dorsal

margin distinctly and broadly ampliate, entire;

ventral margin much shorter than the dorsal, en-

tire or sometimes with a process. Cells of disc

middle thin-walled, trigones large and bulging to

knotlike, 26-33 X 34-52 /xm; median basal cells

larger; cuticle smooth to indistinctly striate-papil-

lose. Underleaves often distinctly connate on one

side (the connation to 7-8 cells high) spreading,

concave, imbricate, symmetrically 4-fid to ca.

0.5-0.6 (median sinus), the sinus bases somewhat

reflexed, the lobes plane or reflexed, diverging,

filiform-attenuate from a narrow base and ulti-

mately with a long-ciliiform tip, entire or with a

ciliiform spine, the lobes appearing to terminate

in a blunt tip, the lobe continuing as a subapical,

often abaxially oriented, ciliiform process with a

uniseriate row of 8-13 cells; disc 18-22 cells high

at median sinus, the disc margins somewhat re-

flexed, on each side often with a ciliiform process

distally, the underleaves then appearing 6-lobed.

Androecia not seen. Gynoecia rarely produced,

on abbreviated ventral-intercalary branches issu-

ing from main stem, solitary or with up to 6 ma-

ture gynoecia produced per shoot, the base of gy-

noecium bulbous and rhizoidous; bracts of inner-

most series much larger than leaves, erect and

sheathing the perianth, the bracts deeply concave,

broadly ovate to suborbicular from a narrow base,

4(6)-lobed to ca. 0.5-0.6, the lobes caudate, the

setaceous tips, composed of a uniseriate row of

6-10 thick-walled, elongate cells, the cells im-

mediately basal to the uniseriate row elongate and

at times subsigmoid, the lobes entire or armed

with a few short to long basal cilia; lamina mar-

gins armed with spinose teeth and straight to con-

torted cilia; bracteole similar in size and form;

gynoecium with a few scales within or immedi-

ately exterior to innermost bracts, the scales

armed with contorted cilia. Perianth long and

prominent, slenderly cylindrical-fusiform, terete

below, obscurely trigonous in distal 0.25-0.4, dis-

tinctly and deeply triplicate toward mouth, the

perianth gradually narrowing toward the strongly

contracted, contorted mouth, the contracted sector

membraneous and, with emergence of the capsule,

splitting into 4-6 narrowly long-attenuate lobes,

the contracted sector composed of elongate, nar-

row, thick-walled, slightly sinuate cells; mouth

cells 4-9: 1, laterally fused for varying lengths and

thickened at the summit, the mouth thus rather

regularly crenulate to crenate-denticulate; perianth

cells ± regularly subrectangular below contracted

sector and here the cuticle is inconspicuously

striolate; perianth 3-5 stratose near base.

Seta with 12-13 rows of large outer cells sur-

rounding an inner core of ca. 40-50 small cells.

Capsule wall 52-54 pm thick, of 5-6 layers; outer

layer of cells with two-phase development, the

longitudinal walls very thin, with sinuous, sheet-

like swellings (very narrow for valve size) and a

few nodulelike swellings alternating with walls

that are devoid of thickenings, the transverse

walls usually devoid of thickenings or sporadi-

cally have an isolated sheetlike thickening; inner-

most and each of intermediate layers subequal in

thickness, the innermost layer of cells ± tiered,

narrowly rectangular, semiannular bands com-

mon, narrow, close and numerous, sporadically

incomplete, rarely forked.

Spores 11-12.5 /^m, exine brown, thin, with

dense, close papillae and short vermiculate mark-

ings. Elaters rigid, nontortuous, 15.8-16.8 yu,m

wide, slightly tapering toward tips, bispiral, the

spirals 4.8-5.3 /xm wide.

DIFFERENTIATION — This is one of the easiest of

our species to identify. The lobes of both leaves

and underleaves are setaceous at their apices, ter-

minating in a uniseriate row of greatly elongated

cells (Fig. 11 : 9). Unlike L. ulothrix, which also

has lobes terminating in a uniseriate row of elon-

gated cells, this species has disc and lobe margins

that are uniformly entire (Fig. 1 1 :3, 4). Also no-

table is the broad and marked ampliation of the

dorsal sector of the leaf, the dilated portion up to

half the disc area.

Lepidozia setigera has distinctive gynoecia.

There are 3-4 gyres of gradually larger, imbricate

bracts. Those of the outer series are about 0.6-0.7

quadrifid, with tapered, acuminate, distally setig-

erous lobes that are entire or occasionally bear a

single basal cilium (Fig. 12:4, 5). Disk margins

are sparingly short ciliate, often with recurved cil-

ia. Innermost bracts are much larger (Fig. 12:3)

but similarly quadrifid, with acuminate-caudate

long, tapered lobes ending in 9-13 or more firm-

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

39

walled, elongated single cells (Fig. 12 : 12). Disk

margins are sparingly ciliate, the short cilia often

reflexed. Perianths are 2-3-layered to at least the

basal third but unistratose above; the narrow

mouth is crenulate to crenate-denticulate.

DISTRIBUTION-ECOLOGY — Endemic to New

Zealand (South Is., North Is.). The species occurs

sporadically in lower- to middle-elevation forests

(60-150 m on South Is., 260-340 m on North Is.),

where it is terricolous or found on old, humus-

covered logs, at times mixed with Bazzania ad-

nexa. On North Is. it chiefly occurs in kauri for-

ests. It is rather common in the Waipoua Forest

Reserve on relatively well-insulated peaty ground

under, for example, old Leptospermum or on very

rotted, bryophyte-covered shaded logs in Agathis

forest containing Dacrydium and other podocarps.

SPECIMENS SEEN— NEW ZEALAND. SOUTH IS-

LAND. WESTLAND PROV.: Ahaura, ENE of Grey-

mouth, ca. 150 m, Child H3770 (F); Mahinapua, S of

Hokitika, ca. 60 m, Child H3666 (F); 2 km N of White

Horse Creek, ca. 305 m, Child H5430 (F). NORTH IS-

LAND. NORTH AUCKLAND PROV.: Waipoua, track

to Mahuta, ca. 305 m, Child H2151 (F); Waipoua Forest,

Track to Te Matua Ngahere, ca. 340 m, Braggins 92/

149 (F — c. sporo.); same loc., Engel 22563 (F); ibid.,

Yakas Track, Braggins 94/214 (AKU); Omahuta Forest

Kauri Sanctuary, E of Mangamuka Bridge, 260 m, Engel

20993 (F), Schuster 95-750— c.. per.

Sect. Notholepidozia

Notholepidozia, as here circumscribed, may

still be heterogeneous. Sporophyte and gynoecium

features of L. obtusiloba are different from other

members of the section. That species has a seta

consisting of 18 outer rows (Fig. 16 : 2), whereas

other members of the section for which the seta

is known have only 8 outer rows (Fig. 20 : 12, L.

laevifolid). The capsule wall of L. obtusiloba is

4-5-stratose (Fig. 16:3), whereas other members

of the section have a 3-stratose capsule, where

known. There appears to be at least a weak cor-

relation between gametophytic vigor and seta and

capsule anatomy (see Taxonomic Characters and

Morphology, p. 18). This correlation should be

tested by the study of sporophytes of those species

heretofore known to lack them. Further subdivi-

sion of sect. Notholepidozia may be necessary af-

ter obtaining sporophyte data of L. ornata (a rel-

atively large species), L. concinna (a median-

sized species) and those of more of the smaller

taxa (L. novae-zelandiae and L. bidens}. The peri-

anth mouth of L. obtusiloba also differs. Mouth

cells of this species are laterally fused to the sum-

mit or nearly so, and the mouth is thus merely

regularly crenulate (Fig. 15 : 6, 7), versus mouth

cells that are laterally free for varying lengths at

the apical end, resulting in a crenate-denticulate

mouth in all other species of the section for which

perianths are known.

Section Notholepidozia has undergone exten-

sive, in part seemingly recent, speciation in New

Zealand. Species limits, therefore, are at times dif-

ficult to draw. Nonetheless, several characters are

of particular taxonomic value within the section.

The width of the ventral merophytes (on well-de-

veloped main stems) is often helpful, particularly

among the complex of species that includes L.

novae-zelandiae (4-6 cells wide; Fig. 24 : 14), L.

laevifolia (6-8 cells wide; Fig. 20 : 11), L. bidens

(8 cells wide; Fig. 30 : 10) and L. concinna

(10 cells wide; Fig. 18: 11) (see discussion on

p. 15).

The form of the basal branch underleaf is some-

times useful and seems fixed in both L. acantha

and L. elobata, where it is nearly uniformly un-

lobed (cf. Figs. 27 : 3, 5; 28 : 1, 2). The remainder

of our species of sect. Notholepidozia have the

basal branch underleaf 2-3- or 3-4- or, in several

cases, 2-4-lobed. Moreover, in several species

(e.g., L. laevifolia, L. pumila) unlobed basal

branch underleaves also may be present, though

exceptionally. The plasticity of the character in

the section renders it of minimal taxonomic use,

aside from the unlobed species (see also the dis-

cussion on p. 14).

The degree of leaf cell papillosity varies con-

siderably within the section from weak and deli-

cate to rather coarse, and may be subject to some

environmental influence. Nevertheless, the char-

acter is valuable within the section (see discussion

on p. 16).

Leaf shape, in general, shows sufficient intra-

specific variation that it is difficult to use taxo-

nomically. In general, the two dorsal lobes form

a pair, separated by a shallower sinus than that

which separates them from the two ventral lobes.

The many figures illustrate the variation seen; it

may be as great intraspecifically as between spe-

cies. Some taxa have a relatively shallow dorsal

sinus (Fig. 28: 3, 4, 7-9, L. elobata), usually less

than 0.3 the leaf length; others have it deeper and

narrower (Fig. 25 : 1, 2, 5, 11, L. novae-zelan-

diae), but intrapopulational variation is sufficient-

ly extensive as to largely bridge such differences.

Species differences are occasionally difficult to

draw in this section, in part because of intrapo-

pulational malleability. It is therefore essential

40

FIELDIANA: BOTANY

that several shoots be studied for assessment of

taxonomic characters.

Lepidozia procera Mitt. Figure 13.

Lepidozia procera Mitt, in Hooker, Bot. Antarc. Voy.

3: 231. pi. 180, f. 1. 1859. Original material: Tas-

mania, without specific locality, Gunn; Mt. Wel-

lington, Oldfield (nan vidi).

Lepidozia breviloba Steph., Spec. Hep. 3: 596. 1909.

Original material: New Zealand. Great Barrier Is.,

Kirk (non vidi).

Plants erect, slender and wiry, with strongly

ventrally secund branches that are often oriented

parallel to one another, pale green, the shoots to

1.5 cm wide (stem to branch extremities). Branch-

ing nearly exclusively of Frullania type, closely

and regularly pinnate, the primary branches be-

coming distinctly whiplike, long-flagelliform, mi-

crophyllous and rhizoidous, the nonmicrophyllous

leaves ± symmetrically 4-lobed, the primary

branches occasionally developing into new lead-

ing shoots; secondry branches rare, 1-2 per pri-

mary branch; branch half-leaf ± symmetrical,

narrowly ovate, shallowly bilobed; first branch

underleaf (2)3-4-lobed, inserted on ventral-lateral

side of junction of main axis and branch and

aligned with branch underleaves. Ventral-interca-

lary branching sporadic, leafy. Stems stiff, 14-16

cells in diameter, the cortical cells in 1 layer of

somewhat thick-walled cells somewhat larger than

the medullary cells (the cortical cells well-defined

in longisection); 1-2 subepidermal layers with

walls thicker than both epidermis and internal

medullary cells, the medullary cells toward stem

middle thick-walled and with trigonelike thick-

enings. Leaves appressed to stem at least in basal

sector, somewhat concave, contiguous to weakly

imbricate, 0.35-0.5 mm long and wide at longest

and widest points, suberect, ventrally secund par-

ticularly when dry, the insertion weakly to dis-

tinctly incubous; leaves slightly to rather mark-

edly asymmetrical, ± equally or, more often, un-

equally 4-lobed, divided to ca. 0.3-0.45 (median

sinus), the distance from dorsal sinus base to in-

sertion greater than that from ventral sinus to in-

sertion (often markedly so). Lobes parallel to

slightly divergent, short-attenuate to acute, entire,

the 2 median lobes wider and often slightly longer

than the outer lobes, 5-7 cells wide at base; lobes

terminating in a single cell or a uniseriate row of

2-3(4) cells, the cells of uniseriate row ± isodi-

ametric, thick-walled and with the septa distinctly

thickened and swollen; cuticle of lobes smooth to

indistinctly and obscurely papillose. Disc moder-

ately to distinctly asymmetrical, subquadrate to

deltoid, 17-26 cells high at dorsal sinus, 10-16

cells high at ventral sinus, the margins entire but

the septa of marginal cells distinctly thickened

and swollen, the dorsal margin slightly to mod-

erately ampliate and broadly curved, the ventral

straight or slightly curved (often with a slight di-

lation at base), shorter than the dorsal. Cells of

disc middle evenly thick- walled, 10-13 X 13-19

/urn, in ± regular longitudinal files; median basal

cells larger (up to 2X diameter of cells of second

row), in one row; cuticle of disc smooth. Under-

leaves often narrowly connate on one side, small,

ca. 0.7-0.9X the stem width, ± appressed to

slightly spreading, quadrate, symmetrically quad-

rifid to ca. 0.3-0.4, the lobes abbreviated, only

(4)5-6(8) cells long; disc (5)7-9 cells high at me-

dian sinus; disc margins entire.

Androecia and gynoecia not seen.

DIFFERENTIATION — The superficial aspect of

plants of this species is that of a miniature L. spi-

nosissima, both taxa being slender and wiry, with

the branches having a spinescent appearance (in

the case of L. procera due to the ventrally secund

leaves; Fig. 13 : 1,4). The primary branches of L.

procera are noteworthy in being strongly ventral-

ly secund and often oriented parallel to one an-

other (Fig. 13 : 1, 14).

The leaves are closely appressed to the stem

(Fig. 13:1,4) and, coupled with being ventrally

secund, lend the shoots and branches a markedly

smooth aspect dorsally. The leaves are shallowly

lobed, the two median lobes larger and somewhat

longer than the outer lobes (Fig. 13 : 5-7). The

orientation of leaf lobes is thus distinctive among

New Zealand species (with the exception of L.

spinosissima), which for the most part have a ten-

dency for the dorsal lobes to be paired to united

and the ventral pair of lobes smaller and some-

what divergent. Also distinctive is the broadly

curved dorsal margin of the leaf, which is longer

than the slightly curved or straight ventral margin;

the difference in margin lengths is related to the

(often marked) greater distance between dorsal si-

nus base to the leaf insertion than from ventral

sinus base to insertion (Fig. 13 : 5-7). The leaf

areolation is distinctive. Disc cells are evenly very

thick-walled, and the cells are in longitudinal files

(Fig. 13:9). The septa of marginal cells, espe-

cially those of the dorsal margin, are distinctly

thickened and swollen (Fig. 13: 11). This char-

acter is not always fully expressed in suboptimal

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

41

n'L-1 )/ }LJ,U( l.l

FIG. 13. Lepidozia procera Mitt. 1. Portion of main shoot with 2 Frullania-type branches (HL — half-leaf); lateral

view. 2. Portion of main shoot, ventral view. 3. Sector of main shoot and Frullania-type branch, ventral view (BUL

= first branch underleaf)- 4. Sector of main shoot, lateral view. 5. Leaves. 6. Leaf and within + at left, underleaves.

7. Four leaves and, at left, underleaves. 8. Dorsalmost 2 lobes of leaf (ds = dorsal sinus, ms = median sinus). 9.

Median disc cells. 10. Median-basal cells of leaf showing row of enlarged cells. 11. Cells of dorsal margin of disc.

12. Distal sector of underleaf. 13. Half-leaf (right) and first branch underleaves. 14. Outline of plant at low magni-

42

FIELDIANA: BOTANY

populations, and several leaves should be careful-

ly examined. Cells of the uniseriate row in the leaf

lobes are thick-walled, with the septa distinctly

thickened and swollen (Fig. 13:8).

In ventral aspect the plants are reminscent of L.

microphylla in having underleaves markedly

small for the shoot size; those of L. procera are

only 0.7-0.9 X the diameter of the stem (Fig. 13 :

2, 3). Also, the underleaves often are narrowly

connate on one side (Fig. 13:2), an unusual fea-

ture among New Zealand representatives of the

genus, or the underleaves may be free (Fig. 13 :

3).

DISTRIBUTION-ECOLOGY — New Zealand (South

Is., North Is.), Tasmania, Australia (Victoria,

Queensland). Within New Zealand the species

sporadically occurs terricolous or saxicolous in

low-elevation (at most 200 m) forests on South

Island, and is known from only one station (290

m) on North Island. For example, it occurs on

mossy banks in Nothofagus forests in Fiordland

(Hatcher 704a), as well as over rock in a creek

bed within a podocarp-dominated forest (Lake

Gault, Engel 21722). On North Island the species

was found on a layer of soil over a huge stream-

bed boulder in a shaded, humid, steep-sided

stream valley within a forest mainly of kauri, but

with some Weimannia silvicola (Waipoua Forest,

Engel 22678).

SELECTED SPECIMENS SEEN — NEW ZEALAND.

SOUTH ISLAND. OTAGO PROV.: Fiordland Natl.

Park, head of Lake McKerrow, near McKerrow Hut,

Hatcher 1476 (F); trail between Gunn's Hut and Hidden

Falls, ca. 30 miles SE of Lake McKerrow, Hatcher 704a

(F). WESTLAND PROV.: Jackson Bay, between conflu-

ence of Jackson River and Arawata River and Lake El-

lery, off Jackson River Road, ca. 30 m, Child H4253 as

L microphylla (F); Westland Natl. Park, track to Lake

Gault, NE of Lake Matheson, NW of town of Fox Gla-

cier, ca. 100-200 m, Engel 21722 (F). NELSON PROV.:

Valley of the Little Wanganui River, ca. 0.5 km S of

Scobie Clearing, 120 m, Fife 7116 as L laevifolia (F).

NORTH ISLAND. NORTH AUCKLAND PROV: Wai-

poua Forest, Waikohatu Stream at Waikohatu Kauri

Bridge, 290 m, Engel 22678 (F).

Lepidozia obtusiloba Steph. Figures 14-16.

Lepidozia obtusiloba Steph., Spec. Hep. 3: 598. 1909.

Original material: New Zealand, South Is., Beckett,

Helms.

Lepidozia pan'itexta Steph., Spec. Hep. 3: 598. 1909,

syn. nov. Lectotype (nov.): New Zealand, Jacksons,

Goebel (G! — c. sporo.).

Plants typically erect, rather flexuous, with lax-

ly spreading (van parvula) to strongly ventrally

secund branches that are often parallel to one an-

other, brown to yellow-brown, the shoots small to

medium, to 1 cm wide, including branches.

Branching nearly exclusively of Frullania type,

rather short, closely and regularly l(2)-pinnate,

the primary branches sporadically becoming

whiplike, flagelliform and microphyllous, the

nonmicrophyllous leaves ± symmetrically 4-

lobed; secondary branches sporadic, 1(3) per pri-

mary branch; branch half-leaf symmetrical, cor-

date, 2-lobed to ca. 0.2-0.4; first branch underleaf

2-3(4)-lobed, inserted on ventral-lateral side of

juncture of main axis and branch and aligned with

underleaves of branch or leaves of main shoot.

Ventral-intercalary branching occasional, leafy or

long stoloniform. Stems rigid. Leaves rigid, brit-

tle, somewhat concave, imbricate and nearly or

completely obscuring stem in dorsal view, 0.65-

1 mm long at longest point, 0.5-1.2 mm wide at

widest point (0.5-0.7 mm wide X 0.7 mm long

in var. parvula), spreading, the insertion narrow

to broad, and weakly incubous; ventral extremity

of leaf at times extending up to stem midline,

leaves distinctly asymmetrical, unequally 4(5)-

lobed, the leaves divided to ca. 0.45-0.65 (median

sinus), the distance from dorsal sinus base to in-

sertion much greater than that from ventral sinus

to insertion, the dorsal sinus at times reduced to

a mere notch. Lobes acute to subapiculate, the 2

dorsal lobes paired, the 2 ventral widely divergent

from the united dorsal lobes; dorsal lobes termi-

nating in several laterally juxtaposed cells or a

single cell or a uniseriate row of 2-3(4) cells, the

cells of uniseriate row ± isodiametric, thick-

walled; dorsal lobes entire, 6-8(10) cells wide at

base. Disc distinctly asymmetrical, 17-21 cells

high at dorsal sinus, 6-10 cells high at ventral

sinus, the margins entire, the dorsal margin broad-

ly ampliate, cordate at the base, the ventral much

shorter than the dorsal, subcordate. Cells of ven-

tral margin and lobes ± isodiametric, forming

border. Cells of disc middle thick-walled, often

distinctly so, trigones sometimes present, medium

fication, lateral view. (Figs. 1, 2, 7, 11 from Engel 21722, New Zealand, South Is., Westland Prov., Westland Natl.

Park, track to Lake Gault; remainder from Hatcher 1476, New Zealand, South Is., Otago Prov., Fiordland Natl. Park,

head of Lake McKerrow.)

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

43

FIG. 14. Lepidozia obtusiloba Steph. 1. Leaves. 2. Apical sectors of 4 lobes from same leaf (vl = ventral lobe;

dl = dorsal lobe). 3. Base of median sinus of leaf showing border of smaller cells. 4. Median disc cells. 5. Sector of

ventral margin of leaf disc showing border of smaller cells. 6. Sector of main shoot with 2 Frullania-type branches,

ventral view (BUL = first branch underleaf). 7. Half-leaves and, below, 2 first branch underleaves. 8. Outline of

shoot at low magnification, lateral view. 9. Underleaves. 10. Apical sectors of 4 lobes from same underleaf. 11.

Sector of main shoot, lateral view. (All from Engel 17843, New Zealand, South Is., Westland Prov., Mt. Aspiring

Natl. Park, off track to Mt. Brewster.)

and straight-sided to large and bulging, 19-28 X

24-35(42) /Am; median basal cells enlarged, in a

single row; cuticle smooth to indistinctly striate to

distinctly papillose. Underleaves widely spreading

to squarrose, symmetrically 4-fid to ca. 0.4-0.55

(median sinus), the lobes plane to slightly ven-

trally sulcate, the sinus bases plane to slightly re-

flexed, the lobes ± parallel to somewhat spread-

44

FIELDIANA: BOTANY

ing, short-attenuate, entire, blunt at the tip and

terminating in several laterally juxtaposed cells,

or with a uniseriate row of 2-3 cells; disc 7-10

cells high at median sinus, the margins parallel,

plane, subcordate at base, entire or with a blunt

tooth.

Plants dioecious. Androecia on inconspicuous,

short, determinate, tightly spicate, often cernuous

ventral-intercalary branches from main shoot;

bracts ventricose-cucullate, 2-3-lobed to ca. 0.2-

0.3, the lobes acute, often apiculate; disc margins

basically entire; antheridia 1-2 per bract, the stalk

biseriate. Gynoecia on abbreviated ventral-inter-

calary branches issuing from main stem, the base

of gynoecium swollen, bulbous and rhizoidous;

bracts of innermost series much larger than

leaves, erect and sheathing the perianth, the bracts

deeply concave, broadly ovate to suborbicular;

apices with 4 abbreviated lobes composed of ±

regularly rectangular and subrhomboidal cells, the

apical end of the marginal cells only sporadically

diverging and forming a tooth, the lobes thus ba-

sically smooth and only exceptionally with a cren-

ulation, the lobe summit composed of 2 or more

laterally juxtaposed cells, with each lobe termi-

nating in a slime papilla, the lobes at times ob-

solete, with each represented solely by a slime

papilla mounted on 2 laterally juxtaposed cells

that project slightly more than neighboring cells;

lamina composed of ± regularly subrectangular

cells, the margin bordered by thin-walled cells of

variable shape and orientation, some hardly lon-

ger than wide, others ± rhomboidal, the apical or

free end of marginal cells usually only sporadi-

cally divergent and forming a crenulation, the

margins basically smooth except for sporadic

crenulations or a several-celled tooth, or, occa-

sionally, the margin crenulate-denticulate by var-

iously projecting cells; bracteole similar in size

and form. Perianth long and prominent, slenderly

cylindrical-fusiform, terete below, obscurely tri-

gonous above, distinctly and deeply 3-plicate to-

ward mouth, the perianth gradually narrowing to-

ward the strongly contracted, shallowly 3-lobed

mouth, the lobes composed of feebly tiered, often

long, narrow, subrectangular to weakly sinuate-

rhomboidal, slightly thick-walled cells; mouth

cells laterally fused to the summit or nearly so,

the mouth thus rather regularly crenulate; perianth

cells ± regularly subrectangular below mouth,

and here the cuticle is inconspicuously striolate;

perianth 2-3 stratose near base.

Seta with 16 rows of outer cells surrounding an

inner core of ca. 60 much smaller cells. Capsule

long elliptical, the wall 46-48 /urn thick, of 4-5

layers; outer layer of cells with two-phase devel-

opment, the longitudinal walls with sinuous,

sheetlike thickenings and several large nodules

and spines alternating with walls that are devoid

of thickenings, the transverse walls usually devoid

of thickenings or sporadically have an isolated

nodule; innermost layer of cells ± tiered, narrow-

ly rectangular, semiannular bands common, close

and numerous, sometimes incomplete, sporadical-

ly forked.

Spores 13.5-14.9 /im, exine brown, thin, with

dense, sharply defined, close papillae and short

vermiculate markings. Elaters rigid, nontortuous,

9.6-10.1 /Am wide, slightly tapering toward tips,

bispiral, the spirals 3.4 /am wide.

DIFFERENTIATION — The typical form of the spe-

cies is distinctive because of brownish pigmen-

tation and erect shoots with distinctly ventrally

secund branches (Fig. 14:8). The species is allied

to L. ornata but differs in the lack of accessory

ornamentation, the broadly acute, united dorsal

lobes with a shallow dorsal sinus (in L. ornata the

lobes are narrowly acute to attenuate and narrow-

ing to a sharp apex), and the essentially plane and

unornamented underleaves. Also, the depth of the

median sinus tends not to be as great as in L.

ornata, and the ventral lobes tend to be broader

than in that species. The underleaf lobes are often

bluntly rounded at the summit (Fig. 14 : 10), giv-

ing rise to the name.

Lepidozia obtusiloba has the dorsal leaf bases

optimally ampliate (Fig. 14 : 1), and the stem half-

leaves tend to have both margins broadly ampliate

at the base (Fig. 14 : 7); it has, apparently consis-

tently, a shallow dorsal sinus, descending only

0.2-0.25 the leaf length. Basal branch underleaves

are usually 2-3-fid. In leaf form closely approach-

ing L. laevifolia var. alpina.

Hodgson (in Herzog, 1938) refers Lepidozia

parvitexta Steph. to L. breviloba Steph., a syno-

nym of L. procera; however, the lectotype of L.

parvitexta, labelled "original" by Stephani, is

typical L. obtusiloba, as is the syntype, leg. Nay-

lor Beckett. Both are abundantly fruiting. Ste-

phani (1909) described the species as "brunneo-

la"; the branches of the type are characteristically

ventrally secund, and the underleaves widely

spreading and divided to ca. 0.5. Although L. par-

vitexta and L. obtusiloba were published simul-

taneously, the latter is the name in current use for

this taxon.

Allison and Child (1975) called attention to the

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

45

FIG. 15. Lepidozia obtusiloba Steph. 1. Portion of main shoot with mature gynoecium, lateral view. Base of

terminal branch shown, but with initial leaves and underleaves removed from branch for clarity (hi = half-leaf). 2.

Innermost bracts and bracteole. 3. Apex of innermost bract. 4. Portion of apex of innermost 9 bract showing obsolete

median pair of teeth, each represented by a slime papilla. 5. Portion of lateral margin of innermost bract showing

nearly entire condition. 6. Perianth mouth, the summit at upper left. 7. Portion of perianth mouth. 8. Perianth, cross

sections (top to bottom) through basal (= b), median (= m) and distal (= d) sectors; d* drawn at higher magnification.

46

FIELDIANA: BOTANY

FIG. 16. Lepidozia obtusiloba Steph. 1. Portion of lateral margin of innermost 9 bract showing crenulate-dentic-

ulate condition. 2. Seta, cross section. 3. Capsule wall, cross section. 4. Capsule wall, outer layer. 5. Capsule wall,

inner layer. 6. Spore (X1880). 7. Elater (X233). (All from Engel 17565, New Zealand, South Is., Otago Prov., S side

of Mt. Cargill.)

9. Perianth, cross sections through median (= m) and distal portions (= d). 10. Capsule profile. (All from Engel

17565, New Zealand, South Is., Otago Prov., S side of Mt. Cargill.)

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

47

presence of 3-lobed branch leaves in this species.

We have not been able to verify this character.

The species is separable into two varieties, as

follows:

Key to the varieties of L. obtusiloba

1. Plants medium (to 1 cm wide, including

branches), erect, dark brown; branches strong-

ly ventrally secund; leaves rather broadly in-

serted, (0.7)0.8-1.2 mm wide X 0.65-1 mm

long var. obtusiloba

1. Plants small (similar to L. concinna in size or

smaller), procumbent, brown or yellow-brown

to greenish; branches spreading and not dis-

tinctly secund; leaves narrowly inserted, easily

detached, only 0.5-0.7 mm wide X 0.7 mm

long var. parvula

Lepidozia obtusiloba Steph. var. obtusiloba

Plants medium (to 1 cm wide, including

branches), erect, dark brown; branches strongly

ventrally secund; leaves rather broadly inserted,

(0.7)0.8-1.2 mm wide X 0.65-1 mm long.

DISTRIBUTION-ECOLOGY— Auckland Is., New

Zealand (South Is.), Australia (Victoria).

A montane variety typically present in the up-

per reaches of forests of, for example, Nothofagus

menziesii or Nothofagus solandri. It is often abun-

dant, and then appears as the most common he-

patic on the forest floor, rotten logs, and tree

trunks. Variety obtusiloba may extend to the sub-

alpine and alpine zones in protected niches on the

faces of boulders or cliffs.

SELECTED SPECIMENS SEEN— AUCKLAND IS.: E end

of Hooker Hills, 396 m, Johnson 7/32 (WELT). NEW

ZEALAND. SOUTH ISLAND. SOUTHLAND PROV.:

Takahe Valley, W of W side of Lake Te Anau, Mark (F);

Fiordland Natl. Park, Gertrude Valley, off track to Ger-

trude Saddle, NE of Homer Tunnel, 1880-1900 m, En-

gel 18864 (F); ibid., S end of Lake Marion, W of Hol-

lyford River, 695 m, Engel 23183 (F). SOUTHLAND/

OTAGO PROV. BOUNDARY: Key Summit, NNE of

Lake Gunn, ca. 915 m, Child H2585 (F). OTAGO

PROV: Ajax Swamp, ca. 1 km N of Ajax Hill, Catlins

River area, ca. 610 m, Child H5523 (F); Maungatua, W

of Mosgiel, ca. 825 m, Child 139 (F); S side of Mt.

Cargill, just below summit, N of Dunedin, ca. 670 m,

Engel 17565 — c. sporo. (F); unnamed peak immediately

NE of Mt. Cargill, N of Dunedin, ca. 550 m, Engel

17585 (F); Blue Mts., 915 m, Child 389 (F); Fiordland

Natl. Park, trail between Gunn's Hut and Hidden Falls,

ca. 30 mi SE of Lake McKerrow, Hatcher 607 (F); ibid.,

head of Lake McKerrow, Hatcher 725 (F); Paradise, Dart

Valley, N of N end of Lake Wakatipu, ca. 610 m, Child

H107 (F); Mt. Aspiring Natl. Park, Blue Valley Track,

above Blue River just N of confluence with Makaroa

River, 430-480 m, Engel 21916 (F); ibid., ridge below

and W of Mt. Shrimpton, 1250 m, Engel 17856 — c. spo-

ro. (F); tributary of Siberia Stream, opposite Siberia Hut,

WNW of Makarora, ca. 760 m, Child H2990 (F). OTA-

GOAVESTLAND PROV. BOUNDARY: Haast Pass, ca.

610 m, Child H4504—C. S + per. (F). WESTLAND

PROV: Mt. Aspiring Natl. Park, off track to Mt. Brew-

ster, below and W of Mt. Armstrong, SW of Mt. Brew-

ster, ca. 900 m, Engel 17843— c. per. (F); Westland Natl.

Park, Franz Josef Glacier Valley, Roberts Point, SW of

Mt. Gunn, ca. 620-670 m, Engel 18113 — c. per. (F);

upper Otira Valley, ca. 1220 m, Child H5266 (F); Camp

Creek, W of Alexander Range, 1040 m, Reif C259F—

c. sporo. (F). WESTLAND/CANTERBURY PROV.

BOUNDARY: Arthur's Pass, Beckett 725, syntype of L.

parvitexta (G — c. sporo.); Arthur's Pass Natl. Park, Ar-

thur's Pass, near Temple Basin Ski area, Engel 6456 (F);

ibid., Bealey Valley Track, ca. 875-900 m, Engel 22865

(F). CANTERBURY PROV: Cass, Woolshed Hill, Visch

s. n. (F); Arthur's Pass Natl. Park, Scotts Track to Ava-

lanche Peak, W of town of Arthur's Pass, 950 m, Engel

22083 (F); ibid., immediately below Punchbowl Falls,

Engel 6874A (F); Bealey Spur, within 3 km W of begin-

ning of track to Bealey Spur Hut, ca. 825 m, Fife 6717

(F); Arthur's Pass Natl. Park, Bealey River, off Bealey

Valley Track, 830-850 m., Engel 18508 (F); Spencer

Mts., opposite Ada Hut, ca. 915 m, Child 4611 (F). NEL-

SON/CANTERBURY PROV. BOUNDARY: Spencer

Range, Mt. Gloriana, ca. 1675 m, Child H3965A (F).

NELSON PROV: Nelson Lakes Natl. Park, Pinchgut

Track, W of southern sector of Lake Rotoiti, SSW of St.

Arnaud, ca. 1280-1390 m, Engel 21401— c. per. (F); St.

Arnaud Range, Rainbow Skifield just below ski-tow

area, E of S end of Lake Rotoiti, 1210 m, Engel 22805

(F); Flora Hut, NW Nelson S. F. Park, ca. 915 m, Child

H4770 (F); Abel Tasman Natl. Park, Mt. Evans, ca. 1065

m, Child H4703A (F). MARLBOROUGH NEAR

BOUNDRY WITH NELSON PROV: Mt. Richmond

Forest Park, Red Hills, track to Maitland Hut, NE of St.

Arnaud, ca. 700-920 m, Engel 21445 (F). MARLBOR-

OUGH PROV: Richmond Range, Mt. Richmond, ca.

1220 m, Child H4335 (F); Mt. Fishtail, ca. 1220 m,

Child H3240 (F); Mt. Robertson, SES of Picton, ca. 1035

m, Child H4392 (F).

Lepidozia obtusiloba var. parvula Engel, var. nov.

Plantae parvae, amplitudini L. concinnae similes vel

minores plerumque procumbentes brunneae flavobrun-

neae vel virescentes; rami patentes subsecundi.

HOLOTYPE — New Zealand, South Is., Westland

Prov., Arthur's Pass Natl. Park, Kelly Range, vi-

cinity of Carroll Hut, above Kellys Creek, N of

Otira, 1150 m, Engel 18382 (F); isotype: (CHR).

Plants small (similar to L. concinna in size or

smaller), procumbent to (exceptionally) suberect,

brown to yellow-brown to greenish; branches

48

FIELDIANA: BOTANY

spreading and not distinctly secund; leaves nar-

rowly inserted, easily detached, 0.5-0.7 mm wide

X 0.7 mm long.

DIFFERENTIATION — A small extreme of the spe-

cies, L. obtusiloba var. parvula typically occurs in

rock crevices or on dripping rock faces in alpine

situations, but it is also encountered, although

sporadically, at median or lower elevations. Plants

assigned to the variety have an appearance quite

distinct from that of the more typical, well-devel-

oped plants of the species, being typically pro-

cumbent, with branches spreading and not dis-

tinctly secund. The leaves are smaller than in var.

obtusiloba, more narrowly inserted and easily de-

tached, and have disc cells often more strongly

thick-walled. The color varies from brown to pale

yellowish brown to greenish. They share with the

typical form a brownish pigmentation, similarly

shaped (though smaller) leaves, and shallowly

lobed, widely spreading to squarrose underleaves.

The variety may be distinguished from L. con-

cinna by the presence of brown pigments or

(when greenish) by the shorter, broader underleaf

lobes, the distinctly asymmetrical leaves with a

shallow dorsal sinus (which may be reduced to a

mere notch), and the smaller, thick-walled median

disc cells. It differs from L. laevifolia by the larger

shoot size, the presence of brown pigments, and

the imbricate, strongly asymmetrical leaves that

are not distinctly cup-shaped when dry.

DISTRIBUTION-ECOLOGY — New Zealand (South

Is., North Is.). Lepidozia obtusiloba var. parvula

typically occurs in rock crevices or on damp to

dripping, often shaded, rock faces in alpine or

subalpine sites (in the latter case, particularly un-

der shrub cover, and here sporadically terrestrial).

It also may occur in very wet niches, such as at

the edges of tarns between tussock and covered

by a layer of old tussock blades (Mt. Robert Ski-

field, Engel 22834). On vertical banks of alpine

rills, it may form very tight, compact populations

with little more than the tips of each shoot ex-

posed (Old Man Range, Engel 23256). The vari-

ety also (sporadically) occurs in upper-elevation

forests of Nothofagus menziesii or N. solandri var.

cliffortioides and only rarely at median or lower

elevations (examples: on old logs at 460 m in the

Mt. Cargill area and at 135 m in the Cascade ul-

tramafic moraine area in southern Westland). At

the Cascade Road site var. parvula occurs on the

floor in an area consisting of ultramafic rocks and

outcrops with rather open vegetation consisting

mainly of Gleichenia, Lycopodium, Juncus, the li-

chen Cladina, and scattered Leptospermum.

SELECTED SPECIMENS SEEN— NEW ZEALAND.

SOUTH ISLAND. OTAGO PROV: Ajax Swamp, ca. 1

km N of Ajax Hill, Catlins River area, ca. 550 m, Child

H5546 (F); Maungatua, W of Mosgiel, ca. 760 m. Child

2843 (F); Mt. Cargill, N of Dunedin, ca. 455-700 m,

Child 962, 2685 as L. Iconcinna (F); Kakanui Range,

ca. 1525 m. Child H2836 (F); Rock & Pillar, ca. 1315

m, Child 488 (F); Old Man Range, Symes Road, 1370-

1570 m, Engel 23256 (F); Jordan River, head of Lake

Wakatipu, vicinity of Paradise, ca. 455 m, Child H12I8

as L. laevifolia (F); Mt. Aspiring Natl. Park, below and

W of Mt. Shrimpton, ca. 1220 m, Child H4803 (F).

WESTLAND PROV.: Cascade Road, Cascade ultramaf-

ic moraine, W of Martyr Saddle, SSW of Jackson Bay,

135 m, Engel 23002; Mt. Aspiring Natl. Park, Cross

Creek, 1.1 km N of Haast Pass, 540 m, Engel 21879 (F);

Mt. Brewster, ca. 1435 m, Schuster 67-459a (F); upper

Otira Valley, ca. 1220 m. Child H5257 (F); Arthur's Pass

Natl. Park, Kelly Range, vicinity of Carroll Hut, above

Kellys Creek, N of Otira, 1150 m, Engel 18382 (F);

ibid., Otira River gorge, W of Arthur's Pass, ca. 305-

365 m, Schuster 48493 (F); Arthur's Pass, Otira side, 760

m, Child 336 (F); Camp Creek. W of Alexander Range,

800-1040 m. Reif CI50B, C259J (F). WESTLAND/

CANTERBURY PROV. BOUNDARY: Arthur's Pass

Natl. Park, Arthur's Pass, near Temple Basin Ski area,

Engel 6454C (F); ibid., Otira Valley Track, along Otira

River, NE of Mt. Rolleston, 1210-1310 m. Engel 22896

(F). CANTERBURY PROV: Mt. Cook Natl. Park. Sealy

Range, below Sealy Lakes, ca. 1280 m, Schuster 4971 1

(F); near summit of Mt. Tourlesse, Hatcher 1216 (F);

Arthur's Pass, ca. 915 m. Child H 2078 (F). NELSON/

CANTERBURY PROV BOUNDARY: Spencer Range.

Mt. Gloriana, ca. 1525-1830 m, Child H3967, H3983

(F). NELSON PROV: Nelson Lakes Natl. Park, E slope

of Robert Ridge in vicinity of Mt. Robert Skifield, W

end of Lake Rotoiti, 1400-1480 m. Engel 22834 (F); St.

Arnaud Range, Rainbow Skifield, E of S end of Lake

Rotoiti, 1360-1480 m, Engel 22759 (F). MARLBOR-

OUGH PROV: Richmond Range, Mt. Richmond, ca.

1615 m, Child H43J6 (F): Mt. Fishtail, ca. 1370 m.

Child H4340 (F). NORTH ISLAND. WELLINGTON

PROV: Tongariro Natl. Park, Soda Springs, Mangate-

popo Stream, 1350 m. Engel 22481 (F); ibid., Taranaki

Falls Track, E of Whakapapa Village, 1240 m. Engel

22469 (F). GISBORNE/SOUTH AUCKLAND PROV

BOUNDARY: Urewera Natl. Park, track to Whakataka

Hut from Lake Waikaremoana, N of western extremity

of Lake Waikaremoana, 1000-1 100 m, Engel 23308 (F).

GISBORNE PROV: Urewera Natl. Park, Huiarau

Range, summit area of Te Rangaakapua, 1265-1320 m.

Engel 23378 (F).

Lepidozia ornata Engel, sp. nov. Figure 17.

Plantae brunneae; rami patentes; folia amphigastri-

aque et lobis accessoriis et additamentis paraphyllaceis

et lamellis varie ornata.

HOLOTYPE — New Zealand, South Is., Westland

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

49

FIG. 17. Lepidozla ornata Engel. 1. Sector of main shoot with Frullania-type branch, ventral view. 2, 3. Leaves.

4. Apical sectors of 5 lobes from same leaf (dl = dorsal lobe; vl = ventral lobe). 5. Second dorsalmost leaf lobe. 6,

7. Underleaves. 8. Apical sectors of lateral (L) and median (M) lobes of same underleaf. 9. Apical sectors of median

(M) and lateral (L) lobes of same underleaf. 10. Sector of main shoot, dorsal view. 11. Half-leaf. 12. Median disc

cells of leaf. 13. Outline of shoot at low magnification. (Figs. 1, 3-7, 10, 12, 13 from type; 2, 8, 9, 11 from Engel

18799, New Zealand, South Is., Fiordland Natl. Park, Mistake Creek.)

50

FIELDIANA: BOTANY

Prov., Mt. Aspiring Natl. Park, below and W of

Mt. Armstrong, SSW of Mt. Brewster, ca. 1250-

1450 m, Engel 17789 (F); isotype: (CHR).

Plants suberect, stiff and wiry, rather flexuous,

with spreading to weakly ventrally secund branch-

es, warm brownish, the shoots medium, to 2.4 cm

wide, including branches. Branching nearly exclu-

sively of Frullania type, rather short, the pattern

variable: distantly and irregularly to regularly

l(2)-pinnate, the primary branches sometimes be-

coming whiplike, flagelliform, and microphyllous,

the nonmicrophyllous leaves asymmetrically to

subsymmetrically 4-lobed, the primary branches

sometimes developing into new leading shoots;

secondary branches occasional, 1-2 per primary

branch; branch half-leaf subsymmetrical, cordate,

2-lobed to ca. 0.3, the base variously armed with

teeth and accessory appendages; first branch un-

derleaf 2— 4-lobed, inserted on main axis or ven-

tral-lateral side of juncture of main axis and

branch and aligned with underleaves of branch.

Ventral-intercalary branching not seen. Stems

somewhat flexuous. Leaves rigid, brittle, concave,

imbricate and completely obscuring stem in dorsal

view or nearly so, 0.9-1.2 mm long at longest

point, 1-1.6 mm wide at widest point, spreading,

the insertion weakly to moderately incubous, re-

curved at dorsal end; ventral extremity of leaf at

times extending to stem midline, the ventral lobe

often aligned with lateral underleaf lobes, ± de-

current on the stem or the lobe with a decurrent

lobuliform flap. Leaves distinctly asymmetrical,

unequally 4(5)-lobed and variously appendaged,

the leaves divided to ca. 0.5-0.85 (median sinus),

the distance from dorsal sinus base to insertion

much greater than that from the median and ven-

tral sinuses to insertion. Lobes narrowly acute to

attenuate to acuminate, the dorsal pair of lobes

paired, the 2 ventral widely divergent from the

united dorsal lobes (when flattened), the ventral 1

or 2 lobes often ventrally sulcate, subfalcate,

sometimes with a tooth, the lobes terminating in

several laterally juxtaposed cells, or a single cell

or a uniseriate row of 2-3 cells, the cells of un-

iseriate row ± isodiametric, thick-walled; dorsal

lobes entire, usually narrowing to a sharp apex,

8-14 cells wide at base; ventral and median si-

nuses reflexed, with accessory armature often pre-

sent in the sinus base. Disc distinctly asymmetri-

cal, 20-25(31) cells high at dorsal sinus, 8-10

cells high at ventral sinus, the dorsal margin

broadly ampliate, often distinctly auriculate at the

base, sporadically with a tooth or spine near the

base, the ventral margin much shorter than the

dorsal, subcordate to subauriculate, typically with

accessory armature. Accessory lamellae some-

times present on the abaxial face of disc, descend-

ing from sinus bases. Flattened paraphyllia-like,

irregularly dentate appendages often present on

base of disc. Marginal cells of disc and lobes ±

isodiametric, forming an indistinct border. Cells

of disc middle with moderately thickened walls,

with intermediate thickenings and medium and

straight-sided to large and bulging trigones, 19-

27 X 28-40 /mi; median basal cells larger, in a

single, sometimes indistinct row; cuticle smooth

to finely striate-papillose. Underleaves plicate,

widely spreading, symmetrically 4-fid to ca. 0.5-

0.6 (median sinus), the lobes typically ventrally

sulcate, with strongly reflexed sinus bases, acces-

sory lamellae, and flattened paraphyllia similar to

those of the leaves; the lobes ± parallel to slightly

diverging, short attenuate, the margins entire to

irregularly crenulate, often bluntly rounded at the

tip or terminating in a uniseriate row of 2(3) cells;

disc 8-10 cells high at median sinus, the margins

reflexed, subauriculate at base, decurrent, often

toothed, rarely with underleaf margin confluent

with the ventral leaf margin, or adnate to face of

leaf disc.

Plants dioecious. Androecia determinate, on

tightly spicate, often cernuous ventral-intercalary

branches from main shoot and primary branches

(in leafy sectors); bracts ventricose-cucullate, 2-

lobed to ca. 0.3, the lobes broad acute, at times

apiculate; dorsal margin of disc dilated and broad-

ly and evenly rounded, the dilated portion with

several slime papillae and sometimes a tooth, the

disc otherwise entire; antheridia 1 per bract, the

stalk biseriate. Gynoecia unknown.

DIFFERENTIATION — The combination of brown

plants with ornamented leaves and underleaves

will immediately distinguish this plant. Leaves

and underleaves are variously appendaged with

accessory lobes, paraphyllia-like appendages, and

lamellae (Fig. 17 : 1-3, 6, 7, 10). The decurrent

ventral extremity of the leaf often extends to the

ventral stem midline, and at times is ornamented

with accessory lobes (Fig. 17 : 1). The ventral and

median sinus bases of leaves and sinus bases of

underleaves are frequently broadly dilated and the

reflexed margins of the lobes often extend down-

ward as irregular lamellae on the abaxial face of

the disc. The flattened paraphyllia-like append-

ages, often at the base of the disc, are present in

no other New Zealand species of Lepidozia and

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

51

are reminiscent of the disc armature of Pachys-

chistochila berggrenii Engel & Schust., also of

New Zealand.

See comments under L. obtusiloba regarding

similarities.

DISTRIBUTION-ECOLOGY — New Zealand (South

Is., North Is.). This species is nearly always sub-

alpine to alpine and occurs under snow tussock

(Chionochlod) cover (often on damp ground be-

tween tussock bases), particularly on damp slopes

or streambanks (?perhaps requiring adequate

drainage). Also present over leaf litter at the edges

of tarns, in crevices of cliff faces, and in pockets

of banks. The sole record of the species in South

Island forests is from the upper reaches of an open

Nothofagus menziesii forest at 610 m at Moraine

Creek, where it forms thick, deep masses on the

sides of bryophyte mounds.

SELECTED SPECIMENS SEEN — NEW ZEALAND.

SOUTH ISLAND. SOUTHLAND PROV.: Fiordland

Natl. Park, below Mt. Burns, E of Borland Saddle, S of

South Branch of Borland Burn, W of Monowai, 1225-

1320 m, Engel 18614 (F); ibid., Mt. Burns, ca. 1370-

1525 m, Child H5094, 5206 (F); ibid., Gertrude Valley,

near track entrance to Gertrude Saddle, E of Homer Tun-

nel, 1740 m, Engel 21948 (F); ibid., Central Earl Mts.,

Mistake Creek, between Triangle Peak and Melita Peak,

NE of N end of Lake Te Anau, 740-800 m, Engel 18799

(F); ibid., Moraine Creek Track, area N of Moraine

Creek, W of Hollyford River, 610 m, Engel 23212 (F).

OTAGO PROV.: Blue Mts., Tapanui, 915m Child 382

(F). WESTLAND PROV.: Mt. Aspiring Natl. Park, be-

low and W of Mt. Shrimpton, 1370-1470 m, Engel

17863 (F); Westland Natl. Park, track to Alex Knob, off

track to Louisa Peak, 1170 m, Engel 18974 — c. <3 (F);

upper Otira Valley, ca. 1220 m, Child H5258—C. 6 (F);

Arthur's Pass Natl. Park, Kelly Range, off track to Car-

roll Hut, above Kelly s Creek, N of Otira, 1040-1 110m,

Engel 18407 (F). WESTL AND/CANTERBURY PROV.

BOUNDARY: Arthur's Pass Natl. Park, Otira Valley

Track, along Otira River, NE of Mt. Rolleston, 1060 m,

Engel 22915 (F). CANTERBURY PROV: Mt. Cook

Natl. Park, Stocking Stream, SW facing cliffs above

Hooker Valley, 1280-1330 m, Engel 18249 (F). NORTH

ISLAND. WELLINGTON PROV: Tongariro Natl. Park,

Mangatepopo Stream below Soda Springs, ENE of Na-

tional Park, ca. 1200 m, Engel 21373 (F).

Lepidozia concinna Col. Figures 6 : 4-7; 18, 19.

Lepidozia concinna Col., Trans. & Proc. New Zealand

Inst. 18: 244. 1886. Original material: New Zea-

land, Waipawa Co., near Norsewood, 1885, Col-

enso a. 1417 (BM!, WELT!).

Lepidozia latiloba Col., Trans. & Proc. New Zealand

Inst. 19: 287. 1887 (1886). Original material: New

Zealand, Waipawa Co., near Norsewood, 1886,

Colenso a. 1421 (BM!, WELT!).

Lepidozia colensoana Steph., Spec. Hep. 3: 597.

1909, syn. nov. Original material: New Zealand,

Lake Waikare, Colenso 1021 (G! — c. per. + seta).

Plants procumbent, the stems interwoven, rath-

er flexuous, with spreading branches, green, high-

ly nitid, the shoots medium, to 1.7 cm wide, in-

cluding branches. Branching nearly exclusively of

Frullania type, often long, closely and regularly

pinnate, the primary branches often becoming

whiplike, flagelliform, microphyllous and rooting

in the substrate, the nonmicrophyllous leaves ±

symmetrically 4-lobed; primary branches often

developing into new leading shoots; secondary

branches occasional, 1-3 per primary branch;

branch half-leaf symmetrical, cordate, 2-lobed to

ca. 0.3; first branch underleaf 2-4-lobed, inserted

on ventral to ventral-lateral side of branch and

aligned with underleaves of branch. Ventral-inter-

calary branching occasional, leafy. Stems flexu-

ous. Leaves when dry curved ventrally, the shoots

(particularly branches) then appearing braided, the

main shoot leaves concave, loosely imbricate and

not obscuring stem in dorsal view, 0.6-0.85 mm

long at longest point, (0.7)0.8-1 mm wide at wid-

est point, spreading, the insertion weakly to

strongly incubous; leaves variable in shape on the

same stem, subsymmetrical to weakly asymmet-

rical (except for the ampliate dorsal base), sub-

equally 4(5)-lobed, the leaves divided to ca. 0.4-

0.6 (median sinus), the distance from dorsal sinus

base to insertion greater than that from ventral

sinus to insertion, the dorsal sinus shallow or the

sinuses gradually becoming deeper ventrally; dor-

sal sinus U-shaped, at least at base. Lobes acute

to apiculate, not noticeably in pairs (except on

asymmetrical leaves), entire, terminating in a un-

iseriate row of 2-3(6) cells, the cells of uniseriate

row ± isodiametric, thick-walled; dorsal lobes 5-

7(9) cells wide at base; cuticle of lobes smooth or

finely striate-papillose. Disc subsymmetrical, 12—

19 cells high at dorsal sinus, 9-1 1 cells high at

ventral sinus, the margins entire, the dorsal mar-

gin ampliate but not strongly so, ± cordate at the

base, the ventral not much shorter than the dorsal.

Cells in a well-defined, large median field, ± iso-

diametric to occasionally oblong-hexagonal,

thick-walled, trigones medium and straight-sided

to large and bulging (rarely weak and concave-

sided), (24)26-41 fan wide X (32)34-45(49) /un

long; cells of the ampliate portion smaller; median

basal cells not sharply differentiated; cuticle of

disc smooth. Oil-bodies in median subapical cells

(1)2-5(6) per cell, coarsely botryoidal, small,

spherical to ovoid, 3.6-4 X 3.6-5 yttrn up to 5 X

FIELDIANA: BOTANY

1 1 /im; basal cells with 4-8 oil-bodies of similar

size (4X5 /Am to 4.5-5 X 6-9 /im). Underleaves

inserted on 10 rows of stem cells, widely spread-

ing, ca. 1-1. 2x stem width, narrowed to insertion,

symmetrically 4-fid to ca. 0.45-0.55 (median si-

nus), the lobes incurved, plane, somewhat spread-

ing, slenderly attenuate, entire, terminating in a

single cell or uniseriate row of 2-3(5) cells; disc

7-9 cells high at median sinus, the margins plane,

broadly curved, entire.

Androecia not seen. Perianth (fide Stephani,

1909) broadly oblong, 4-6 cells thick at base, the

mouth constricted and entire; capsule broadly

oval; spores 18 /urn. reddish brown; elaters bis-

piral. laxly twisted.

DIFFERENTIATION — The leaves of Lepidozia

concinna are variable in shape on the same shoot

but typically are subsymmetrically to weakly

asymmetrically lobed, with the lobes not notably

(and never consistently) in pairs (Fig. 18:3, 4).

The underleaves are somewhat wider than the

stem, narrowed to the insertion (Fig. 18 : 1, 2),

inserted on 10 rows of stem cells (Fig. 18 : 11),

and with the lobes slenderly attenuate and often

broadly incurved (Figs. 18: 1, 2; 19: 1). The

plants are uniformly green, lacking secondary

pigmentation, which will immediately distinguish

this species from the L. obtusiloba complex. A

notable feature of this species is the well-defined

median field of large disc cells (to 40 /Am wide);

the cells are in large part ± isodiametric and typ-

ically have medium to large and bulging trigones

(Fig. 18:6, 7). A few basal cells may be quite

large, 25-35(42) X 35-48(56) /Am versus 22 X

25-32(36) /Am in the lobes (compare Fig. 6 : 5,

6). Lepidozia concinna is usually readily recog-

nized because of the deeply lobed underleaves,

with attenuated lobes, often 3-4 cells broad at the

base (Figs. 18:8, 11; 19:5, 6).

Mature sporophytes of L. colensoana are de-

scribed by Stephani (1909). The type of L. col-

ensoana bears a perianth with an exerted seta, but

no capsules were seen; the material is too scanty

to bear examination of fertile structures. This is

the only fruiting material of L. concinna known.

This species is rather similar in leaf form to L.

pendulina (compare Fig. 10 : 3, 4) but has much

smaller appendages; leaves average ca. 615-700

/Am broad X 555-620 /Am long versus 1640 /Am

broad X 1 300 /Am long for L. pendulina. The un-

derleaves of L. pendulina differ from those of L.

concinna in being (often) weakly to distinctly pli-

cate, with lamina margins less curved and much

less narrowed toward the base (cf. Fig. 10 : 1) and

with lobes ventrally sulcate.

In L. concinna median cells are ca. 26-4 IX

34-45 /Am; they each have (1)2-5(6) oil-bodies

each, small and coarsely botryoidal, ranging from

3-4 /Am to 4 x 5 to 3 X 6 up to 5-5.5 X 10-1 1

/tm. Basal cells are ca. 25-35 X 33-46 /Am and

have 4-8 oil-bodies each; they are similar to those

of median cells and are scarcely larger (4 X 4-7

to 4.5 X 6 to 5 X 9 /Am), similarly botryoidal.

Plants of Schuster 67-237a (Fig. 6 : 4-7) have

rather similar leaves, as regards outline and lob-

ing, as does L. pendulina (Fig. 10:3); both have

dorsal leaf bases ampliate, but not strongly so;

both have the two dorsal lobes both set off as a

unit from the ventral lobes. Both have a narrow,

very sharp ventral lobes. There are clear differ-

ences in oil-body numbers. In L. concinna cells

of the leaf middle and distal sectors have (1)2-

5(6) oil-bodies each (Fig. 6 : 5) and the basal cells

have mostly 4-8 oil-bodies each (Fig 6 : 6). In L.

pendulina the upper laminar cells have (2)3-8(14)

oil-bodies (Fig 6:1), and the basal cells bear

mostly 16-20 oil-bodies each (Fig. 6 : 2). Basal

cells in L. concinna are mostly 26-35 X 33-48

/im (Fig. 6 : 6); in L. pendulina at least a small

area has basal cells to 40-43 X 55-60 /Am (Fig.

6:2).

For additional comments, see under L. pendu-

lina and L. obtusiloba var. pawula.

NOTES — The type collection of L. concinna

consists of rather weak, lax, suboptimal shoots

with leaves moderately asymmetrical. However,

some sectors of shoots have leaves more typical

of the species, and show weakly asymmetrical

leaves with the distinct field of large median disc

cells characteristic of this species.

DISTRIBUTION-ECOLOGY — New Zealand (South

Is., North Is.), Tasmania. A forest plant, often in

extensive patches and becoming abundant, occur-

ring on old, rotten logs or, less often, over soil;

the species occurs along a broad elevational gra-

dient (150-920 m).

SELECTED SPECIMENS SEEN— NEW ZEALAND.

SOUTH ISLAND. SOUTHLAND: Lake Hauroko, ca.

215 m, Child HI 542 (F); Waikaia. Heriot Rd. ca. 610

m, Child H3920 (F); N of Te Anau, Hatcher 1515 (F);

immediately N of Ten Mile Bush, near W shore of Lake

Te Anau and ca. 20 km N of town of Te Anau, 200 m.

Engel 23201 (F); around Smithy Creek on road from Te

Anau to Milford Sound, ca. 15 km S of Lake Fergus,

Raven 26644A (F). OTAGO PROV.: Blue Mts.. Black

Gully, near Tapanui. 610m Child 20 (F); Mt. Cargill, N

of Dunedin, ca. 455 m. Child 2308 (F); Morrisons

Creek, N of Dunedin, ca. 305 m. Child 1318 (F); Jordan

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

53

FIG. 18. Lepidozia concinna Col. 1. Portion of main shoot with Frullania-type branch, ventral view. Note position

of first branch underleaf. 2. Portion of main shoot with two Frullania-type branches, ventral view. 3. Leaves. 4. Four

leaves and (lower right) 2 underleaves at same scale. 5. Dorsal lobe of leaf. 6, 7. Median disc cells of leaf. 8.

Underleaves (drawn at same scale as Fig. 4). 9. Apical sectors of 2 median lobes of same underleaf. 10. Primary

branch leaves (PEL), half-leaf (HL), and first branch underleaves (BUL). 11. Portion of stem with underleaf. Note

54

FIELDIANA: BOTANY

FIG. 19. Lepidozia concinna Col. 1. Part of main stem, ventral aspect, with a sector (below) omitted. Note five

Frullania-lype branches (F), with 3-4-fid first branch underleaves (UL1) (X34; 1 mm scale). 2, 3. Two leaves (X47:

600 /j.m scale). 4. Three underleaves (X47; 600 fj.m scale). 5, 6. Lobes of underleaves (X 150; 150 /im scale). 7. Leaf

(X47; 600 iim scale). 8. Stem half-leaf (X47; 600 /im scale). 9. Stem underleaf (X47; 600 /im scale). 10. Branch

underleaf (X47; 600 fjun scale). 11. Stem cross section (x 150; 150 /*m scale at right). 12-14. Cells of leaf lobe (12),

middle (13) and leaf base (14) (X265; 100 /*m scale). 15. Leaf (X265; 600 /xm scale). (All from Schuster 95-1435,

New Zealand.)

ventral merophyte width of 10 cells. 12. Outline of plant at low magnification. (Figs. 1, 4, 7, 11 from Engel 23201,

New Zealand, South Is., Southland Prov., immediately N of Ten Mile Bush, near W shore of Lake Te Anau; remainder

from Child 4373, New Zealand, South Is., Marlborough Prov., Red Hills.)

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

55

River, head of Lake Wakatipu, vicinity of Paradise, ca.

365-455 m, Child 2421, 2467B (F); Paradise, Dart Val-

ley, N of N end Lake Wakatipu, ca. 365 m, Child 1197

(F). OTAGO/WESTLAND PROV. BOUNDARY: Haast

Pass, Matthews s. n. (Allison H3158) (F); Haast Pass,

ca. 610 m, Child H4506 (F). WESTLAND PROV.:

Camp Creek, W of Alexander Range, 210 m, ReifCllZ

(F); Ahaura, ENE of Greymouth, ca. 150 m, Child

H3745 (F). MARLBOROUGH NEAR BOUNDRY

WITH NELSON PROV: Mt. Richmond Forest Park,

Red Hills, track to Maitland Hut, NE of St. Arnaud, ca.

700-920 m, Engel 21438 (F). MARLBOROUGH

PROV: Red Hills, W of hut ca. 915 m, Child H4373

(F). NORTH ISLAND. WELLINGTON PROV: Tonga-

riro Natl. Park, ca. 7 km from Ohakune on Ohakune Mt.

Road, 900 m, Engel 22702 (F). GISBORNE PROV:

Urewera Natl. Park, Waikareiti Track between track en-

trance and Lake Ruapani, N of eastern extremity of Lake

Waikaremoana, 650-920 m, Engel 20588 (F). SOUTH

AUCKLAND PROV: Waiotapu, Rotorua, Child 43 C

(F).

Lepidozia laevifolia (Hook. f. & Tayl.) G. L. &

N. Figures 20-22

Jungermannia laevifolia Hook. f. & Tayl., London J.

Bot. 3: 385. 1844 (3: 285. [sic] in errore pro 385).

Lepidozia laevifolia (Hook. f. & Tayl.) G. L. & N.,

Syn. Hep. 208. 1845. Mastigophora laevifolia

(Hook. f. & Tayl.) Trev., Cat. Herb. Crypt. Trevisan

2: 30. 1853. Lectotype (nov.): Campbell Is., Nov.

1840, Hooker (YR\).

Lepidozia minuta Col., Trans. & Proc. New Zealand

Inst. 18: 245. 1886 non L. minuta Steph., Spec.

Hep. 3: 603. 1909. Original material: New Zealand,

Waipawa Co., near Norsewood, 1885, Colenso a.

1419 (BM!, o = 0, WELT!).

Lepidozia retrusa Col., Trans. & Proc. New Zealand

Inst. 22: 455. 1890 (1889). Original material: New

Zealand, Waipawa Co., S of Dannevirke, 1889,

Colenso a. 1515 (BM! — c. sporo., G — 0, WELT! —

c. per.).

Lepidozia papillata Steph., Spec. Hep. 3: 595. 1909.

Original material: Auckland Is., Hooker (G!, "sub

Lepid. dispar").

Lepidozia asperifolia Steph., Spec. Hep. 3: 596. 1909.

Original material: New Zealand, South Is. (non

vidi).

Plants closely prostrate, flexuous, with spread-

ing branches, green, nitid when dry, the shoots

small, to 1 cm wide, including branches; excep-

tionally luxurient phases to 2.5 cm long X 0.9 cm

wide. Branching nearly exclusively of Frullania

type, rather short, normally irregularly and dis-

tantly pinnate, ± regularly pinnate in well-devel-

oped plants, the primary (and secondary) branch-

es sometimes becoming whiplike, flagelliform and

microphyllous, the nonmicrophyllous leaves ±

symmetrically 4-lobed; primary branches occa-

sionally developing into new leading shoots; sec-

ondary branches occasional, 1-4 per primary

branch; branch half-leaf subsymmetrical, cordate,

2-lobed to ca. 0.25; first branch underleaf (1)2-4-

lobed, inserted on ventral side of branch base to

the ventral-lateral side of junction of main axis

and branch, in both cases the first branch under-

leaf aligned with underleaves of branch. Ventral-

intercalary branching occasional, leafy, at times

becoming leading shoots. Stems flexuous, ca. 10

cells in diam., the cortical cells in 1 layer of thick-

walled cells somewhat larger than the thick-

walled medullary cells. Leaves when dry deeply

cup-shaped, with the tips of the lobes incurved

and not visible in dorsal view; leaves when moist

rigid, distinctly concave, contiguous, with much

of stem visible in dorsal view, 0.5—0.65 mm long

at longest point, 0.5-0.7 mm wide at widest point,

spreading, the insertion distinctly incubous; leaves

variable, subsymmetrical to distinctly asymmetri-

cal, subequally 4(6)-lobed, sometimes with a ±

shallow dorsal sinus, the leaves divided to ca.

0.4-0.65 (median sinus), the distance from dorsal

sinus base to insertion ± equal to or much greater

than that from ventral sinus to insertion. Lobes

broadly to narrowly acute, sometimes short-apic-

ulate, the two ventral occasionally somewhat di-

vergent from the dorsal lobes, the lobe margins

entire to somewhat sinuate to occasionally 1-2-

dentate, terminating in a single cell or a uniseriate

row of 2-3 nonelongated, thick-walled cells; dor-

sal lobe 5-10 cells wide at base (at times 4 cells

wide in minute phases). Disc subsymmetrical to

asymmetrical and then obliquely truncate, 12-18

cells high at dorsal sinus, 4-9 cells high at ventral

sinus; dorsal margin ampliate, cordate to auricu-

late at the insertion (rarely decurrent), entire to

sinuate, sometimes irregularly crenulate to den-

tate; ventral margin shorter than dorsal, ± cordate

at the insertion, entire or sometimes with an ac-

cessory lobe. Cells evenly thick-walled, occasion-

ally with small trigones, the cells of median por-

tion of disc 16-25(28) ^tm wide X 19-32 /tm

long, ± longitudinally elongated, typically sharply

differentiated from the small and isodiametric (8-

10 yam) cells of the lobes and ampliate sector of

the disc; marginal cells of disc and lobes lacking

a thickened outer wall; cuticle of abaxial surface

coarsely and closely papillose on lobes and am-

pliate sector (the adaxial surface typically

smooth), the disc closely striate-papillose. Under-

leaves inserted on 6-8 rows of stem cells, spread-

ing, short, broader than high, symmetrically 4-fid

to ca. 0.3-0.45 (median sinus), the lobes plane,

entire, acute to acuminate, terminating in a single

cell or a uniseriate row of 2-4 cells; disc 7-8 cells

56

FIELDIANA: BOTANY

high at median sinus, cordate at the insertion, the

margins plane, entire.

Plants dioecious. Androecia on inconspicuous,

short, determinate, tightly spicate, sometimes cer-

nuous ventral-intercalary branches from main

shoot or terminal on long flagelliform primary

branches; bracts ventricose-cucullate, 2-lobed to

ca. 0.3-0.4, the lobes acute to apiculate, the lateral

margins occasionally with several slime papillae;

antheridia 1-2 per bract, the stalk biseriate. Gy-

noecia on abbreviated ventral-intercalary branch-

es issuing from main stem; bracts of innermost

series much larger than leaves, erect and sheath-

ing the perianth, the bracts deeply concave, sub-

orbicular to broadly ovate to suboblate; apices

with 4 small teeth hardly differentiated from the

otherwise crenate-denticulate apex, or with (2)4

short, rather irregular lobes composed of ± reg-

ularly rectangular + irregularly and .feebly sinu-

ate-rhomboidal cells, the apical end of the mar-

ginal cells often feebly diverging, the lobes thus

finely and sparingly crenulate; lamina composed

of ± regularly short rectangular cells, the margin

bordered by cells of variable shape, some hardly

longer than wide, others rather long and narrow,

the apical or free end of marginal cells variously

divergent and forming a short projection or a

tooth, the margin irregularly crenate-denticulate to

the base, the teeth occasionally composed of a

few to several cells; bracteole similar in size and

form. Perianth long and prominent, slenderly cy-

lindrical-fusiform, slightly curved, terete below,

obscurely trigonous above, distinctly and deeply

3-plicate toward mouth, the perianth gradually

narrowing toward the strongly contracted, shal-

lowly 3-lobed mouth, the lobes composed of ±

tiered, long, narrow, subrectangular cells; mouth

cells rather thick-walled, at the apical end laterally

free for varying lengths, the mouth thus crenate-

denticulate; perianth cells ± regularly rather short

to long subrectangular below mouth and here the

cuticle is conspicuously striate-papillose; perianth

2-stratose near base.

Seta with 8 rows of outer cells surrounding an

inner core of 18 much smaller cells. Capsule wall

30-35 n,m thick, of 3-4 or 4-5 layers; outer layer

of cells with two-phase development, the longi-

tudinal walls with sinuous, sheetlike thickenings

alternating with those that are devoid of thicken-

ings (or are rarely locally thickened), the trans-

verse walls usually devoid of thickenings or spo-

radically have an isolated nodule; innermost layer

of cells ± tiered, narrowly rectangular, with no-

dulelike to spinelike thickenings common, with

semiannular bands weakly developed, only occa-

sionally present, often incomplete, pale.

Spores 11.5-13.4 ^m, exine light brown, with

dense, rather coarse, sharply defined, rather broad

papillae and simple or furcate vermiculate mark-

ings. Elaters ± rigid to feebly tortuous, 9.6-10.1

/Am wide, only slightly tapering toward tips, bis-

piral to tips, the spirals 3.4-3.8 /tm wide.

DIFFERENTIATION — Examination of the type of

Jungermannia laevifolia, a Campbell Is. plant, re-

veals that the species is identical with the com-

mon New Zealand plant called Lepidozia asperi-

folia by Stephani (1909, sub L. "levifolia"), who

maintained that the true L. laevifolia did not occur

in New Zealand. Hodgson (1956) and others (e.g.,

Allison & Child, 1975) use the name Lepidozia

laevifolia collectively to refer to all of the New

Zealand species of the "laevifolia" complex.

When differentiated, however, as they are here,

the name must be applied to the plant with leaves

densely papillose, and broadly ampliate dorsal

margins. The species epithet, "laevifolia," is un-

fortunate and misleading, standing in contradic-

tion to the more appropriate names "papillata"

and "asperifolia."

A note on the type of Lepidozia papillata (G!)

indicates that the specimen was extracted from a

specimen of Lepidozia dispar (= Kurzia hippu-

roides), leg. Hooker on Auckland Is., Herb. Kew.

At least one of the specimens labeled L. dispar at

BM includes strands of Lepidozia laevifolia, but

other liverwort species that are associated with the

type of L. papillata at G are not present, and this

element in the BM specimen cannot be confidently

regarded as an isotype of L. papillata.

Stephani (1892) referred L. minuta Col. and L.

retrusa Col. to L. praenitens (= Telaranea praen-

itens (Lehm. & Lindenb.) Hodg.). The types of

these two Colenso species (BM!) were annotated

by Stephani as L. praenitens', both are Lepidozia

laevifolia.

Small, poorly developed plants of L. laevifolia

and L. pumila may be difficult to distinguish sat-

isfactorily. Both species can have distinctly con-

cave leaves with the lobes incurved and hidden

when viewed in dorsal aspect. Well-developed

leaves should always be selected for examination,

but a number of leaves should also be examined

to determine the range of variation. The leaves of

L. laevifolia are nearly always coarsely papillose

(Fig. 20 : 8-10), and the papillae are typically pre-

sent only on the abaxial surface of the leaf (the

adaxial surface being almost smooth). Distinctly

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

57

50 M

FIG. 20. Lepidozia laevifolia (Hook. f. & Tayl.) G. L. & N. (Figs. 6, 7 from var. acutiloba; remainder from var.

laevifolia). 1. Sector of main shoot with Frullania-type branch, ventral view (BUL, first branch underleaf). 2. First

branch underleaves. 3. Sector of shoot, dorsal view. 4-7. Leaves and underleaves (ul), all drawn at same scale. 8.

Median disc cells showing cuticular papillae. 9. Marginal cells of ampliate sector of disc. 10. Leaf lobes showing

cuticular papillae in lobe at left. 11. Portion of stem with underleaf disc and lobe bases. Note ventral merophyte

width of 10 cells and rhizoid origin (arrows indicate sinus bases). 12. Seta, cross section. (Figs. 1-3 from Engel

58

FIELDIANA: BOTANY

papillose leaves may vary vis-a-vis the number of

papillae per cell: Some populations have very

coarse papillae that number ca. 7-8 or 12-15 per

cell (as in the type of L. laevifolia, Fig. 20 : 8).

Other populations are less coursely papillose, with

ca. 30 papillae per cell. The dorsal leaf margin in

L. laevifolia is usually ampliate and distinctly cor-

date at the insertion, the margins of the disc and

lobes are sinuate, and the cells of the ampliate

portion are small and abruptly differentiated from

the cells of the disc. Typically, in L. pumila the

cuticle is smooth, and the dorsal leaf margin is

straight and abruptly cordate at the base (Fig. 29 :

1, 4, 5); the leaf cells appear large in proportion

to the size of the leaf, giving the leaf a "cellular"

appearance. Another useful character in distin-

guishing L. pumila from L. laevifolia is the pres-

ence in the former species of a thickened outer

wall of at least some of the marginal cells of disc

and lobes (Fig. 29 : 6, 8).

Suboptimal phases of L. laevifolia at times have

mostly undivided first branch underleaves and

therefore may be confused with L. acantha. Such

populations differ from L. acantha by the non-

elongated cells of the leaf lobe uniseriate row, the

arched dorsal margin of the leaf disc, and the

presence of at least some bifid (or trifid) first

branch underleaves. Bilobed first branch under-

leaves in L. acantha are very rare. It is important

to select the best-developed shoots when exam-

ining suspected suboptimal populations.

In the dilated dorsal leaf bases similar to L. elo-

bata, which differs in the consistently unlobed

first branch underleaves (Fig. 28 : 1); the relative-

ly narrower leaves (Fig. 28 : 3, 4, 8); and the shal-

lower median sinus of the leaf (Fig. 28 : 3, 4, 7-

8).

The species is separable into three varieties, as

follows.

Key to the varieties of L. laevifolia

1 . Dorsal half of leaf more deeply bilobed (typi-

cally to 0.2 or more); disc cells moderately and

evenly thick-walled, occasionally with small

trigones 2

2. Leaf lobes narrowly acute, the two dorsal

lobes separated by a narrow, rather deep si-

nus (less than 45°); median disc cells large,

18-25(28) /x,m wide; leaf margins sinuate to

occasionally 1-2 -dentate; underleaf lobes

slenderly acuminate; leaves ± strongly

asymmetrical. Occurring from (60 m) 300

m to subalpine zone var. laevifolia

2. Leaf lobes broadly acute, the two dorsal

lobes separated by a broad, shallow sinus

(45-90°); median disc cells narrow, 16-20

/xm wide; leaf margins often sinuate, never

toothed; underleaf lobes acute to broadly

acuminate; leaves weakly asymmetrical.

Occurring in upper-elevation forests to al-

pine zone var. acutiloba

1 . Dorsal half of leaf very shallowly bilobed (to

0.15-0.2); disc cells evenly and very strongly

thick-walled, no trigones var. alpina

Lepidozia laevifolia var. laevifolia

Leaves ± strongly asymmetrical, obliquely

truncate, 4(6)-lobed, divided to 0.5-0.65 (median

sinus); leaf lobes narrowly acute, the two dorsal

lobes ± paired, separated by a narrow, rather deep

sinus (less than 45°); disc 12-15 cells high at dor-

sal sinus, 4-7 cells high at ventral sinus, the ven-

tral sinus often ± reflexed; leaf margins occasion-

ally 1-2-dentate; median disc cells 18-25(28) /tm

wide; underleaf lobes slenderly acuminate, at

times only 2-3 cells wide at base.

DISTRIBUTION-ECOLOGY — Amphipacific tem-

perate: Macquarie Is., Auckland and Campbell Is.,

New Zealand (South Is., North Is.), Tasmania,

Australia (Victoria, N.S.W.), Kerguelen Is., Mar-

ion and Prince Edward Is., Falkland Is., southern

South America, Juan Fernandez Is.

In New Zealand var. laevifolia typically ranges

from middle-elevation forests to the subalpine

zone and is able to tolerate considerable exposure.

However, it may be present in lower-elevation for-

ests, and it extends as far north as Radar Bush

(WSW of Cape Reinga), where it occurs on rot-

ted, decorticated logs in forests of Beilschmiedia-

Vitex-Hoheria and Cvathea dealbata (100 m).

18406, New Zealand, South Is., Arthur's Pass Natl. Park, Kelly Range; 4, 8-10 from type of L. laevifolia; 5 from

Engel 20585, New Zealand, North Is., Gisborne Prov., Urewera Natl. Park, Waikareiti Track; 6 from Engel 22816,

South Is., Nelson Prov., road to Rainbow Skifield in vicinity of Red Gate, just W of Wairau River; 7 from type of

var. acutiloba; 1 1 from Engel 22885, New Zealand, South Is., Westland/Canterbury Prov. Boundary, Arthur's Pass

Natl. Park, Bealey Valley Track; 12 from Child 2849, New Zealand, South Is., Otago Prov., Maungatua.)

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

59

FIG. 21. Lepidozia laevifolia (Hook. f. & Tayl.) G. L. & N. 1. Two bracts and (to right) bracteole from innermost

series (all drawn to same scale). 2, 3. Portions of apex of 2 innermost bracts. 4, 5. Complete lateral margin and basal

half of margin, respectively, of innermost bracts (drawn to same scale). 6, 7. Portions of perianth mouth (drawn to

same scale). 8, 9. Capsule, cross sections of 3-4- and 4-5-stratose walls. 10. Capsule wall, outer layer. 11. Capsule

wall, inner layer. (All from Engel 20669, New Zealand, North Is., near Gisborne/South Auckland Prov. Boundary,

Urewera Natl. Park, crest trail from Highway 38 towards Whakataka summit.)

60

FIELDIANA: BOTANY

Also present at ca. 60-80 m in the Kiwanis Re-

serve (S of Kaitaia), where it occurs on nikau in

mixed broadleaf forests dominated by Beilschmie-

dia and Vitex, with Rhopalostylis (nikau), Co-

prosma, Dacrycarpus, and Hoheria. Generally,

however, the variety is less common below 300

m. In forests it occurs on shaded, moist cliff faces

and boulders, soil banks (including banks of road-

cuts through the forest, and here at times mixed

with Breutelia elongata), rotted logs, and occa-

sionally over bark. It is rather common on the

floor and loosely on bark at the bases of Leptos-

permum in the "Frost Flats" (South Auckland

Prov., near Otupaka Stream, SW of Minginui). In

subalpine situations it occurs on soil under and in

between tussocks (at times where the tussock base

is rotted), under shrub cover, in boulder fields, on

cliff faces (where it sometimes forms sheets) and

on clayey banks. In boggy sites, it may occur

among Sphagnum, and then the shoots are often

quite straggly.

SELECTED SPECIMENS SEEN — CAMPBELL IS.: Kirk

314, syntype of L. novae -zelandiae — c. sporo. (G). NEW

ZEALAND. SOUTH ISLAND. SOUTHLAND: Fiord-

land Natl. Park, Borland Road, 14 km by road WNW

from Borland Lodge, near South Branch of Borland

Burn, 855-870 m, Engel 18687 (F); East Dome, Garvie

Range, ca. 1370 m, Child H3913 (F); Wilmot Pass Road

to Deep Cove, ca. 305 m, Child H5602 as L microphylla

(F). OTAGO PROV.: Ajax Swamp, ca. 1 km N of Ajax

Hill, Catlins River area, ca. 610 m, Child H5492 (F); Mt.

Maungatua, W of Mosgiel, ca. 500 m, Engel 17781 (F);

Maungatua, W of Mosgiel, ca. 855 m, Child HMO as L.

microphylla (F); Silverpeaks, N of Dunedin, ca. 455 m,

Child H3406 (F); Flagstaff, NW of Dunedin, ca. 455 m,

Child 1377 as L. microph\lla (F); Lammermoor Range,

ca. 610 m, Child H3843 (F); Great Moss Swamp, WSW

of Middlemarch, ca. 305 m, Child H472 (F); Kakanui

Range, ca. 1220 m. Child H2837 (F). WESTLAND

PROV.: Near Hercules Creek on Route 6 in Mt. Hercules

Scenic Reserve, Engel 6559 (F); upper Otira Valley, ca.

1220 m, Child H5342 (F); Arthur's Pass Natl. Park, Kel-

ly Range, off track to Carroll Hut, above Kellys Creek,

N of Otira, 1040-1110 m, Engel 18406 (F); Camp

Creek, W of Alexander Range, 320 m, Reif C29C (F).

WESTLAND/CANTERBURY PROV. BOUNDARY:

Arthur's Pass Natl. Park, Arthur's Pass, near Temple Ba-

sin Ski area, Engel 6473A (F); ibid., Dobson Nature

Walk, summit area of Arthur's Pass, 920 m, Engel 22025

(F); ibid., Bealey Valley Track, ca. 875-900 m, Engel

22885 (F). CANTERBURY PROV: Mt. Cook Natl.

Park, Governors Bush, SW of town of Mt. Cook, 760-

800 m, Engel 18206— c. per. (F); Arthur's Pass Natl.

Park, immediately below Punchbowl Falls, Engel 6870

(F); Mt. Gloriana, Spencer Range, 1676 m, Child H

3965B (F); Mt. Tourlesse, Hatcher 1221 (F). NELSON

PROV: Nelson Lakes Natl. Park. Pinchgut Track, W of

southern sector of Lake Rotoiti. SSW of St. Arnaud, ca.

1280-1390 m, Engel 21405 (F); ibid., NE margin of

Lake Rotoroa, W of St. Arnaud, 520 m, Engel 21505

(F). MARLBOROUGH NEAR BOUNDRY WITH

NELSON PROV: Mt. Richmond Forest Park, Red Hills,

track to Maitland Hut, NE of St. Arnaud, ca. 700-920

m, Engel 21472 (F). NORTH ISLAND. Without specific

locality, "alpine interior," Colenso a.2200 as L. praen-

itens (BM). WELLINGTON PROV: Near eastern border

of Tongariro Natl. Park along road to Tree Trunk Gorge,

ca. 0.5 km W from gorge, 750 m, Engel 21209 (F). GIS-

BORNE PROV: Urewera Natl. Park, Waikareiti Track

between track entrance and Lake Ruapani, N of eastern

extremity of Lake Waikaremoana, 650-920 m, Engel

20585 (F). NEAR GISBORNE/SOUTH AUCKLAND

PROV BOUNDARY: Urewera Natl. Park, crest trail

from highway 38 toward Whakataka summit, N of

northern extremity of Lake Waikaremoana, 930-1030

m, Engel 20669— c. sporo. (F). SOUTH AUCKLAND

PROV: "Frost Flats," on South Road between Totara

Salvage Road and Wong Road, near Otupaka Stream,

SW of Minginui, Engel 20747 (F); Whirinaki Forest

Park, Waterfall Loop track, near Whirinaki River, SSW

of Minginui. Engel 20694 (F); Rotorua thermal area,

Hatcher 90 (F); 7 km N of junction of Highways 33 and

30 on Highway 30, Kantak & Churchill 78— c. d (F).

NORTH AUCKLAND PROV: Auckland, Cheeseman

32, syntype of L novae -zelandiae (G); NE Waitakere

Ranges, Swanson University Reserve, Tram Valley

Road, 95 m, Engel 20466 — c. per. (F); same loc., Brag-

gins 99/275 (AKU); SE corner of Waipoua Forest, just

N of Tutamoe, 540 m, Engel 21118 (F); Mangamuka

Walkway, Maungataniwha Range, ESE of Kaitaia, sad-

dle on State Highway 1, 390 m, Engel 20960 (F); Ki-

wanis Reserve, junction of Okahu Stream and unnamed

stream, ca. 5 km S of Kaitaia, N edge of Herekino Forest

area, S of Quarry, ca. 60-80 m, Engel 20907 (F); Radar

Bush, WSW of Cape Reinga, S of Mt. Te Paki, ca. 100

m, Engel 20850 (F).

Lepidozia laevifolia var. acutiloba Engel, var.

nov.

Lobi foliares late acuti, sinu inter lobos dorsales duos

non profundo, lato (45-90°) instructi; cellulae medianae

disci angustae 16-20 /im latae; lobi amphigastrii acuti

vel late acuminati.

HOLOTYPE — New Zealand, South Is., Otago

Prov., W slope of Flagstaff, NW of Dunedin, 560

m, Engel 17632 (F); isotype: (CHR).

Leaves weakly asymmetrical, ± equally 4(6)-

lobed, divided to 0.35-0.5 (median sinus); leaf

lobes broadly acute, the two dorsal lobes separat-

ed by a broad, often shallow sinus (45-90°), the

disc 16-19 cells high at dorsal sinus, 6-10 cells

high at ventral sinus; leaf margins often sinuate

but rarely toothed; median disc cells small, 16-20

/im wide; underleaves shallowly lobed, the lobes

acute to broadly acuminate.

DISTRIBUTION-ECOLOGY — New Zealand (South

Is., North Is.). Typically an upper-elevation plant,

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

61

occurring in the scrub Nothofagus menziesii tran-

sition zone and extending into the subalpine and

alpine zones. It occurs on damp ground, some-

times appearing as a pioneer, at other times be-

tween tussocks or over leaf litter under shrubs;

also on damp cliff faces, rock outcrops, and in

rock crevices. In the forest zone it may be terric-

olous or corticolous, as well as occurring on rot-

ted logs, particularly where there has been soil

accumulation.

SELECTED SPECIMENS SEEN— NEW ZEALAND.

SOUTH ISLAND. SOUTHLAND PROV.: East Dome,

Garvie Range, ca. 1370 m, Child H 3910 (F). OTAGO

PROV.: Rock & Pillar, S end, near McPhees Rock,

WNW of Middlemarch, ca. 1220 m, Child H3861 (F);

Mt. Maungatua, W of Mosgiel, 760 m, Engel 17766 (F);

W slope of Flagstaff, NW of Dunedin, 560 m, Engel

17632 (F); S side of Mt. Cargill, just below summit, N

of Dunedin, ca. 670 m, Engel 17578 (F); Kakanui Range,

ridge to Kakanui Peak, ca. 1220 m, Child H2838 (F);

opposite Siberia Hut, WNW of Makarora, ca. 1065 m,

Child H2919 (F); Fiordland Natl. Park, near McKerrow

Hut, head of Lake McKerrow, Hatcher 1768 (F); ibid.,

Martin's Bay, N of McKerrow River, Hatcher 844 (F).

WESTLAND PROV.: Mt. Aspiring Natl. Park, below

and W of Mt. Armstrong, SSW of Mt. Brewster, 1250-

1450 m, Engel 17792 (F); road to Rainbow Skifield in

vicinity of Red Gate, just W of Wairau River, E of N

end of Lake Rotoiti, 700 m, Engel 22816 (F). CANTER-

BURY PROV: Mt. Cook Natl. Park, SW facing cliffs

off Wakefield Track, overlooking Hooker River, E of

town of Mt. Cook, 720 m, Engel 18177 (F); ibid., Stock-

ing Stream, SW facing cliffs above Hooker Valley,

1280-1330 m, Engel 18244 (F); Arthur's Pass Natl.

Park, immediately below Punchbowl Falls, Engel 6896

(F); ibid., near start of Punchbowl Creek Track just

above village of Arthur's Pass, Engel 6913 (F); junction

of Craigieburn River and Route 73, Engel 6922 (F).

NORTH ISLAND. WELLINGTON PROV: Tongariro

Natl. Park, track to Soda Springs, along Mangatepopo

Stream, ca. 1300 m, Engel 22499 (F); ibid., Blyth Track,

along small stream ca. 0.5 km from Ohakune Mt. Road,

ca. 1230 m, Engel 21302 (F). GISBORNE PROV:

Urewera Natl. Park, Waikareiti Track between track en-

trance and Lake Ruapani, N of eastern extremity of Lake

Waikaremoana, 650-920 m, Engel 20552 (F).

Lepidozia laevifolia var. alpina Schust. & Engel,

Dimidium folii dorsale non profunde bilobum, usque

0.15-0.2 longitudinis folii; cellulae disci parietibus ae-

qualiter atque pervalde incrassatis praeditae, trigonis

carentes.

HOLOTYPE — New Zealand, South Is., Fiordland

Natl. Park, Gertrude Cirque NE of Homer Tunnel,

below saddle on W facing exposed cliff, 1945 m,

Schuster 95-2089 (hb. Schuster).

Plants 500-550 to 760-810 /un broad with

leaves, rather remotely pinnate, the branches in

small part becoming flagelliform, rhizoidous;

leaves strongly asymetrically bifid, with each lobe

again bifid, ca. 380-420 /mi broad X 440-

480(500) fjim long; dorsal half of leaf much larger

and longer than ventral half, ca. 0.15-0.2(0.25)

bilobed, the lobes acute but not strongly so, end-

ing in 2-3 subisodiametric or isodiametric cells;

cells of disc strongly and evenly thick-walled, tri-

gones lacking, the median disc cells (18)20-26

jjim wide; cuticle papillose (at times obscurely so).

The var. alpina is similar to var. acutiloba, also

of higher elevations, in the shallow dorsalmost si-

nus and the broadly acute lobes. The var. alpina

differs from var. acutiloba in the exceptionally

shallow dorsalmost sinus, in the exceedingly

thick- walled leaf cells, and in the somewhat larger

median disc cells ([18]20— 26 /mi wide versus 16—

20 fjim in var. acutiloba). Also, in var. alpina the

dorsalmost sinus tends to be narrower than in var.

acutiloba and may even be slitlike.

Variety alpina has distinctly thick-walled cells

reminiscent of L. pumila. However, the var. alpina

has shorter and much less tapered dorsal leaf

lobes, separated by a rather shallow sinus. Also,

unlike in L. pumila, the sinus separating the two

dorsal lobes is usually much shallower than that

separating the two ventral lobes. In L. pumila the

dorsal sinus descends ca. 0.3-0.35 the leaf length,

in var. alpina only about (0.15)0.2-0.25 the leaf

length. Moreover, L. pumila lacks an auriculate

dorsal leaf base, has larger disc cells (to 33 /tm

wide vs. at most 26 /mi wide in var. alpina), and

has ventral merophytes 5-6 cells wide versus 6-

8 cells wide in var. alpina.

In leaf shape (particularly in the shallow dorsal

sinus and cordate to cordate-auriculate dorsal ba-

ses) the variety is similar to L. obtusiloba. Lepi-

dozia obtusiloba, however, has leaf and underleaf

lobes usually blunt; they are always sharp in var.

alpina (Fig. 22 : 1-8). Moreover, L. obtusiloba

has leaves with a much deeper ventral sinus than

in var. alpina.

The var. alpina, as the name implies, is an alpine

plant, known only from ca. 1950 m. It is possible

that var. alpina is a mere alpine extreme in which,

with extreme exposure, the dorsal leaf lobes are

abbreviated. Other characters of var. alpina rele-

vant to extreme exposure are the very thick-walled

leaf cells and the markedly thick cuticle.

Leaf surface ornamentation in this species is

variable. It may be distinctly papillose on some

shoots or obscurely so on others.

62

FIELDIANA: BOTANY

FIG. 22. Lepidozia laevifolia van alpina Schust. & Engel. 1, 2. Leaf and underleaves, main stem (X75; 300 /im

scale). 3. Leaf and underleaf, main stem (X75; 300 /im scale). 4. Ventral half of leaf from Fig. 1 (X183; 150 /j.m

scale). 5, 6. Apices of dorsalmost lobe (X183; 150 /im scale). 7. Leaf and underleaf (X51; 400 ^tm scale). 8. Two

large stem leaves (X51; 400 fim scale). 9. Part of main axis with bases of two branches; note trifid basal branch

underleaves (UL1) (X42; 600 /tm scale). 10. Shoot sector, ventral aspect, with bases of two Frullania-lype branches

at UL1, basal branch underleaves, at HL, part of stem half-leaf visible (X51; 500 fim scale). 11. Two leaves (X51;

400 /Am scale). 12. Stem cross section (X225; 100 /im scale). (All from type.)

DISTRIBUTION-ECOLOGY — Known only from

type plants, which occurred admixed with Her-

bertus alpinus and Gymnomitrion strictum, creep-

ing amid Hymenophyllum.

Lepidozia fugax Engel, sp. nov. Figure 23.

Plantae in aspectu laceratae; apices lobi foliaris caduci

e cellulis turgidis doliiformibus subisodiametrae con-

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

63

15

FIG. 23. Lepidozia fugax Engel. 1. Sector of main shoot, dorsal view. 2. Sector of main shoot and Frullania-type

branch, ventral view. 3. Leaves (note missing lobe tips). 4. Dorsal margin of leaf (dm) and (left to right, i.e., dorsal

to ventral) all lobes of same leaf (vl = ventral lobe; sp = slime papilla; cuticular papillae shown in part). 5. Dorsal

pair of lobes, the uniseriate sectors of each lobe having fallen away (cuticular papillae shown in part). 6. Second

dorsal lobe (complete; note constricted septa). 7. Sector of dorsal margin of disc with a blunt tooth (cuticular papillae

shown in part). 8. Median disc cells of leaf. 9. Leaf disc cross section (ab = abaxial face). 10. Portion of stem with

FIELDIANA: BOTANY

stricto-septatis instruct!; amphigastria in seriebus septem

cellularum caulinarum inserta.

HOLOTYPE — New Zealand, South Is., Otago/

Westland Prov. boundary: Mt. Aspiring Natl.

Park, summit area of Haast Pass, 570 m, Engel

18023 (F); isotype: (CHR).

Plants loosely prostrate, in pure mats, rather

stiff and wiry, with spreading branches, green,

distinctly nitid when dry, the shoots small, to 1.2

cm wide, including branches (shoots with rigor-

ous branches), shoots with short branches at times

only 3 mm wide. Branching nearly exclusively of

Frullania type, short to rather long, irregularly

l(2)-pinnate, the primary (and secondary) branch-

es sporadically to frequently becoming whiplike,

flagelliform and microphyllous, the nonmicro-

phyllous leaves asymmetrically to subsymmetri-

cally 4-lobed, the uniseriate portion submonili-

form; primary branches occasionally developing

into new leading shoots; secondary branches oc-

casional, up to 6 per primary branch; branch half-

leaf subsymmetrical, the basal sector somewhat

rounded to subcordate, 2-lobed to ca. 0.3; first

branch underleaf 2-3-lobed, inserted on ventral

side of branch base to the ventral-lateral side of

junction of main axis and branch, and in both cas-

es, the first branch underleaf ± aligned with un-

derleaves of branch. Ventral-intercalary branching

occasional, leafy. Stems rather stiff. Leaves frag-

ile and easily detached; when dry moderately con-

cave, the dorsal lobes stiffly spreading, the tips of

the ventral lobes usually incurved and not visible

in dorsal view; leaves when moist often with a

ragged appearance, slightly concave, contiguous

to imbricate, with much of stem visible in dorsal

view, 0.3-0.5 mm wide and long, distinctly

spreading, the dorsal margin forming an angle

with stem as much as ca. 90°, the insertion dis-

tinctly incubous; leaves variable in shape, sub-

symmetrical to moderately asymmetrical, sub-

equally to unequally 4(5)-lobed, the leaves divid-

ed to ca. 0.3-0.5 (median sinus), the distance from

dorsal sinus base to insertion greater than that

from ventral sinus to insertion. Lobes acuminate,

often appearing short in proportion to disc, ± par-

allel to weakly divergent, the tips often caducous

(lobes often lacking the uniseriate sector), the

lobes often becoming slender and 2 cells wide for

much of their length, the dorsal lobes 2-4 cells

wide at the base; lobes entire, terminating in a

uniseriate row of 2-4(5) at most somewhat elon-

gated (to ca. 1.3:1) cells, the cells of uniseriate

row turgid, barrel-shaped, the septa constricted,

the terminal cell rounded at the tip; cuticle of

lobes papillose. Disc moderately to distinctly

asymmetrical (the disc then obliquely truncate),

9-13 cells high at dorsal sinus, 6-7 cells high at

ventral sinus; dorsal margin moderately amplia-

te, cordate at the insertion, mostly entire to sin-

uate, sporadically with a 1-2-celled blunt or

sharp tooth; ventral margin somewhat shorter

than the dorsal, subcordate at the insertion, entire

or sometimes with a single tooth. Cells of me-

dian portion of disc with walls uniformly rather

strongly thickened, trigones lacking, the cells

17-24 /Ltm wide X 20-31 /im long, ± longitu-

dinally elongated; cells of ampliate sector ± iso-

diametric; marginal cells of disc and lobes in sur-

face view with a strongly thickened free wall

(the wall often crescentic and bulging into the

cell lumen), the disc cells in cross section with

crescentic thickenings of the abaxial wall; cuticle

of disc smooth. Underleaves inserted on 7 rows

of stem cells, strongly spreading, ca. 1.5-2X

stem width, broader than high, symmetrically 4-

fid to ca. 0.3-0.5 (median sinus), the sinuses V--

shaped, the lobes slender, acuminate, entire, the

tips often caducous, terminating in a single cell

or a uniseriate row of 2-3 cells; disc 5-7 cells

high at median sinus, the margins plane, entire.

Asexual reproduction by caducous tips of leaf

lobes.

Plants dioecious. Androecia on inconspicuous,

short, determinate, tightly spicate, cernuous ven-

tral-intercalary branches from main shoot or sub-

microphyllous primary branches; bracts ventri-

cose-cucullate, 2-lobed to ca. 0.3, the lobes acute

to apiculate, the lobe tips complete, the lateral

margins entire or with a tooth; antheridia 1 per

bract, the stalk biseriate. Gynoecia on abbreviated

ventral-intercalary branches issuing from main

stem, the gynoecium base bulbous, fleshy, poly-

stratose and rhizoidous; bracts of innermost series

much larger than leaves, erect and sheathing the

underleaf; note ventral merophyte width of 7 cells. 11. Underleaves. 12. Two half-leaves and (to right) 2 first branch

underleaves. 13-15. Outlines of three shoots at low magnification (all at same scale). (All from holotype.)

ENGEL & SCHUSTER: AUSTRAL HEPATIC AE. LEPIDOZIA

65

perianth, the bracts deeply concave, suborbicular

to suboblate; apices with 4 small, rather irregular

lobes that are often composed of a base of 2 tiers

of cells that are irregularly quadrate to short-rect-

angular to sporadically rather long and narrowly

subrectangular, the apical end of the marginal lobe

cells often feebly diverging, the lobes thus often

finely and sparingly crenulate, the lobes often ter-

minating in a uniseriate row of 2 narrowly rect-

angular cells or 2 laterally juxtaposed cells; lam-

ina composed of ± regularly quadrate to short

subrectangular cells, those of the distal sector

rather thick-walled, those of remainder of bract

(particularly those in a large median-basal field)

thin-walled, the margin bordered by cells of var-

iable shape, some hardly longer than wide, others

rather long, narrow, and at times ± sinuate-rhom-

boidal, the apical or free end of marginal cells at

times divergent and forming a short projection or

a tooth, the margin irregularly crenate-denticulate

in distal sector, the teeth unicellular or composed

of a few cells but not forming a uniseriate row of

2 or more cells, the basal sector of bract margin

entire except for slime papillae; bracteole similar

in size and form. Perianth long and prominent,

slenderly cylindrical-fusiform, slightly curved, te-

rete below, obscurely trigonous above, distinctly

and deeply 3-plicate toward mouth, the perianth

gradually narrowing toward the strongly contract-

ed, shallowly 3-lobed mouth, the lobes composed

of ± tiered, long, narrow, subrectangular cells;

mouth cells rather thick-walled, papillose, at the

apical end laterally free for varying lengths, the

mouth thus crenate-denticulate; perianth cells ±

regularly rather short-subrectangular below

mouth, and here the cuticle is conspicuously stri-

ate-papillose; perianth 2-3-stratose near base.

Seta with 8 rows of outer cells surrounding an

inner core of numerous much smaller cells. Cap-

sule oblong, the wall 26-30 /on thick, of 3 layers;

outer layer of cells with two-phase development,

the longitudinal walls with sinuous, sheetlike

thickenings and several large nodules alternating

with those that are devoid of thickenings (or are

sporadically locally thickened), the transverse

walls usually devoid of thickenings or sporadi-

cally have a sheetlike thickening or an isolated

nodule; innermost layer of cells narrowly rectan-

gular, semiannular bands common, close and nu-

merous, sometimes incomplete, rarely forked.

Spores 12.4-13 /on, exine brown, with coarse,

sharply defined, close papillae and short vermic-

ulate markings. Elaters rigid, nontortuous, 8.6-9.1

/Am wide, only slightly and gradually tapering to-

ward tips, bispiral to tips, the spirals 3.4-3.8 /mi

wide.

DIFFERENTIATION — The species is the only re-

gional member of subg. Notholepidozia with cad-

ucuous leaf tips (Fig. 23 : 1,3, 5), which give the

plants a distinctive ragged appearance. The turgid,

barrel-shaped cells of the uniseriate row and the

constricted septa are responsible for the fragility

of the lobes. The cells of the broken tips often

have walls that have become secondarily thick-

walled, rounded, and papillose, following the

breakage of the lobe tips.

As in Lepidozia novae -zelandiae, the abaxial

cell walls of the disc and lobes are strongly thick-

ened, as confirmed by leaf cross sections (Fig.

23 : 9). These thickenings may also be viewed in

a ventral aspect of the intact shoot by focusing on

the ventral margins of the lobes. In surface view,

only the outer wall of the marginal cells of disc

and lobes appears thickened (Fig. 23 : 4).

DISTRIBUTION-ECOLOGY — Known only from

two collections; the type occurred on an erect, de-

corticated, rotted, 30 foot high tree remnant in a

Nothofagus menziesii forest at 570 m (Haast

Pass). A second collection was made in the upper

limits of a Nothofagus menziesii forest (900 m),

where it occurred mixed with Telaranea cf. cor-

ticola on rotted wood at the base of a huge tree

(Mt. Brewster).

SPECIMEN SEEN— NEW ZEALAND. SOUTH IS-

LAND. WESTLAND PROV.: Mt. Aspiring Natl. Park,

off track to Mt. Brewster, below and W of Mt. Arm-

strong, SW of Mt. Brewster, ca. 900 m, Engel 17847b

(F).

Lepidozia novae-zelandiae Steph. Figures 24; 25;

26: 4-13

Lepidozia novae-zelandiae Steph., Spec. Hep. 3: 595.

1909 ("Nova Zelandiae"). Lectotype (nov.): New

Zealand, without specific locality, Colenso 767 (o!)

Plants brittle, prostrate, in loosely interwoven,

± pure mats, the shoots slender, delicately spi-

nescent, stiff and wirelike, with widely spreading

branches, green, nitid when dry, the shoots mi-

nute, to 1.5 mm wide, including branches.

Branching exclusively of Frullania type, ± dis-

tantly once-pinnate, the branches often becoming

whiplike, flagelliform and microphyllous, the

nonmicrophyllous leaves ± symmetrically 4-

lobed; primary branches occasionally developing

into new leading shoots; secondary branches spo-

66

FIELDIANA: BOTANY

radic, 1-2 per primary branch; branch half-leaf

subsymmetrical, cordate at the base, 2-lobed to ca.

0.4; first branch underleaf (1) 2-3(4)-lobed, in-

serted on ventral side of branch base to the ven-

tral—lateral side of junction of main axis and

branch, nearly aligned with underleaves of

branch. Ventral-intercalary branching not seen.

Stems flexuous. Stem epidermal cells in surface

view strongly thick-walled. Leaves when moist

(or dry) moderately concave, with the lobes some-

what incurved, contiguous to ± distant, with

much of stem visible in dorsal view, somewhat

longer than wide, 0.25-0.3 mm long at longest

point, 0.25-0.35 mm wide at widest point, spread-

ing, the insertion subtransverse to weakly incu-

bous; leaves variable, ± symmetrical to moder-

ately asymmetrical, subequally 4(5)-lobed, the

leaves divided to ca. 0.6 (median sinus), in ±

symmetrical leaves the distance from dorsal sinus

base to insertion not much greater than that from

ventral sinus to insertion. Lobes longly acute to

short-acuminate, typically somewhat apiculate, in

asymmetrical leaves the dorsal lobes ± paired, the

ventral shorter than the dorsal lobes and some-

what divergent, the dorsal lobes entire, 4-5 cells

wide at base; lobes terminating in a single cell or

in a uniseriate row of 2 cells, the cells of uniser-

iate row elongated (to 2:1), typically strongly

thick-walled, the septa somewhat to distinctly di-

lated, the terminal cell tapering to a point; cuticle

of abaxial surface of lobes coarsely papillose (the

adaxial surface typically smooth). Disc subsym-

metrical to obliquely truncate, 6-10 cells high at

dorsal sinus, 4-6 cells high at ventral sinus, the

margins entire, the dorsal margin ± straight,

abruptly cordate near the base, entire to sinuate;

ventral margin shorter than dorsal, at times dis-

tinctly so, entire or sinuate, sometimes with an

accessory lobe. Cells of disc middle often strong-

ly thick-walled, occasionally with small to medi-

um trigones, 11-16 /am wide X 14-22 />tm long;

median basal cells in 1 (locally 2) rows of en-

larged cells; marginal cells of disc and lobes with

a strongly thickened, often concave outer wall

(the wall thickening often crescentic and bulging

into the cell lumen), the septa ± dilated; cuticle

of disc finely to distinctly striate-papillose. Un-

derleaves inserted on 4-6 rows of stem cells,

spreading, symmetrically 4-fid to ca. 0.5 (median

sinus), narrowing to the insertion, the lobes plane,

slenderly acuminate, entire, terminating in a sin-

gle cell or in a uniseriate row of 2-4(5) cells; disc

4-7 cells high at median sinus, the margins plane,

entire. Cells of underleaf thicker-walled than

those of leaves, similar in thickness to epidermal

cells of stem.

Androecia and gynoecia not seen.

DIFFERENTIATION — Lepidozia novae-zelandiae

is closely allied to L. laevifolia, as evidenced by

the coarsely papillose leaves (Fig. 24 : 6), (the pa-

pillae are restricted to the abaxial leaf surface) and

by the sinuate leaf margins. It differs in the sub-

transverse to weakly incubous leaf insertion (Fig.

24 : 2), the ± straight dorsal margin (Fig. 24 : 2-

4), the strongly thickened outer walls and dilated

septa of the marginal cells of both disc and lobes

(Fig. 24 : 5-9), the elongated (to 2:1) cells of the

uniseriate portion of the lobe apices (Fig. 24 : 5-

8), and the distinctly thickened stem epidermal

cell walls (Fig. 24 : 14-15). Lepidozia pumila re-

sembles this species in the more or less straight

dorsal margin of the leaves and thickened walls

of the marginal cells, but the cuticle is typically

smooth, and the leaves are more distinctly incu-

bous.

In L. novae-zelandiae, the abaxial cell walls of

disc and lobes are typically thickened; these thick-

enings can also be seen by focusing on the ventral

margins of the lobes. Similarly thickened cell

walls are present in L. fugax (Fig. 23 : 4, 9), but

the characteristic deciduous lobe tips, and the bar-

rel-shaped cells and constricted septa of the uni-

seriate row, will immediately distinguish that spe-

cies from L. novae-zelandiae. It is notable that

this species was known to Hodgson, as evidenced

by a specimen (F!) identified as L. novae-zelan-

diae; she referred to this species (1956, p. 597

sub L. laevifolia) as "a form with many capillari-

attenuated branches."

The subtransverse insertion of the leaves and

the thick-walled, almost spinose terminal cells of

the inflexed lobe tips of the main shoot and

branches lend an aspect reminiscent of the genus

Psiloclada Mitt, of the same family. The incras-

sate walls of the stem epidermal cells are similar

to those of many Kurzia species.

Two of the syntypes of this species in Ste-

phani's herbarium belong to L. laevifolia: Camp-

bell Island, leg. Kirk, and New Zealand, Auck-

land, leg. Cheeseman. However, a syntype from

Auckland Island, leg. Kirk, is L. novae-zelandiae,

which establishes a rather wide-ranging distribu-

tion for this rare species.

VARIATION — Three extremes, evidently but not

surely conspecific, are assigned to this species.

The foregoing diagnosis and discussion apply

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

67

100 ll

FIG. 24. Lepidozia novae-zelandiae Steph. 1. Sector of main shoot, and Frullania-type branch, ventral view (BUL

= first branch underleaf). 2. Sector of main shoot (HL = half-leaf), dorsal view. 3. Leaves; below, underleaves. 4.

Leaves. 5. Dorsal leaf lobes. 6. Dorsalmost leaf lobe, the cuticle indicated in part. 7. Dorsalmost of median leaf lobes,

cuticlar detail not shown. 8. Distal sector of dorsal lobes. 9. Portion of dorsal margin of leaf disc. 10. Median cells

of leaf disc. 11. Two half-leaves. 12, 13. Underleaves, cellular detail. 14. Portion of stem with underleaf; note ventral

merophyte width of 5 cells. 15. Ventral cortical cells of stem, surface view. (Figs. 1-3, 5, 8-1 1, 13, 15 from Schuster

68

FIELDIANA: BOTANY

FIG. 25. Lepidozia novae-zelandiae van heterostipa Schust. 1, 2. Stem leaves, dorsal margin at left (X96; 150

IJim scale). 3. Dorsal lobe of leaf (X298; 80 /AID scale). 4. Dorsal lobes of leaf (X298; 80 /Lim scale). 5. Dorsal half

of leaf (X 185; 150 ^m scale). 6. Leaf (X96; 150 /urn scale). 7. Underleaves (X96; 150 fim scale). 8, 8', and 8". Four

sectors of a single stem showing 1- and 3-lobed basal branch underleaves (X85; 300 /urn scale). 9, 9', and 9". Three

sectors of one stem showing 1-, 2-, and 3-lobed basal branch underleaves (X85; 300 /xm scale). 10, 11. Leaves (X96;

150 /j.m scale). 12. Shoot sector, dorsal aspect, at HL, half-leaves, at HUL1, part of first branch underleaf visible

(X85; 300 /im scale). (All from type.)

59645b, New Zealand, South Is., Westland Prov., upper reaches of Haast River, 0.6 mi. N of Haast Pass; 4, 6, 7, 14

from Engel 23107, New Zealand, South Is., Westland Prov., Mt. Aspiring Natl. Park, Cross Creek; 12 from Hodgson

s. n., New Zealand, North Is., Gisborne Prov., Lake Waikaremoana.)

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

69

solely to var. novae-zelandiae. The three differ as

follows:

Key to the varieties of L. novae-zelandiae

1. Leaf lobes, excluding ventralmost, (3) 5-6

cells broad, to 7-9 cells long; basal branch un-

derleaf (l)2-3(4)-lobed; underleaves with disc

4-5 cells high, the lobes 4-5 cells broad; un-

derleaves inserted on 4-6 cell rows 2

2. Dorsal leaf sinus descending ca. 0.35-0.4;

ventral sinus descending usually over 0.5

the leaf length; dorsal and ventral sinuses

usually wide open, V-shaped; leaf lobe api-

ces ending in (1) 2 clearly elongated (to 2:

1) cells var. novae-zelandiae

2. Dorsal leaf sinus descending ca. 0.3-0.35 the

leaf length; ventral sinus descending 0.45-

0.5; dorsal (and often ventral) sinuses slitlike

or narrowly V-shaped; leaf lobe apices end-

ing in (1)2-3 subquadrate to short-triangular

cells (1-1.5:1) var. heterostipa

1. Leaf lobes 2-3 cells wide, 4-6 cells long; bas-

al branch underleaf 1- or 2-lobed; underleaves

with disc 2 cells high, the lobes usually few-

celled, 2 cells broad at base; underleaves (8

cells broad at base) inserted on 4 cortical cell

rows var. minima

Lepidozia novae-zelandiae Steph. var. novae-ze-

landiae

DISTRIBUTION-ECOLOGY — Aside from the type

(of unknown provenance, but North Is. because it

is a Colenso collection), this species is known

from the Haast Pass area (Westland Prov.), Ton-

gariro Natl. Park, Lake Waikaremoana, and the

Waipoua area on North Island. Haast Pass collec-

tions were from steep, humid but never wet, shad-

ed, vertical, south-facing cliffs in a Nothofagus

menziesii forest, and may occur with Plagiochlla

caducifolia, Gymnomitrion cuspidatum, Diplo-

phyllum domesticum, and Radula sp. The Tonga-

riro National Park plant occurred within a mixed

broadleaf forest (with Nothofagus menziesii com-

mon), and grew very loosely over soil on a ver-

tical bank of the Mangawhero River. In the Wai-

poua area, the species occurred on a roadside

bank under protected cover, with Blechnum, be-

neath a kauri forest.

SPECIMENS SEEN — AUCKLAND IS.: Kirk s. n., syn-

type of L. novae-zelandiae (G). NEW ZEALAND.

SOUTH ISLAND. WESTLAND PROV.: Upper reaches

of Haast River, 0.6 mi. N of Haast Pass, ca. 520 m,

Schuster 53382a, 59645b (F); Mt. Aspiring Natl. Park,

Cross Creek, 1.1 km N of Haast Pass, 540 m, Engel

21884 (F). NORTH ISLAND. WELLINGTON PROV.:

Tongariro Natl. Park, Mangawhero River near juncture

of Rimu Track and Forest Walk, just NE of Ohakune,

625 m, Engel 22721 (F). GISBORNE PROV: Lake Wai-

karemoana, 610 m, 1935, Hodgson s. n. (as L. novae-

zelandiae) (F). NORTH AUCKLAND PROV: Waipoua

River, ca. 70 m upstream from State Highway 12 bridge,

ca. 95 m, Engel 21044 (F).

Lepidozia novae-zelandiae var. heterostipa Schust.,

var. nov.

Sinus folii dorsalis usque 0.3-0.35 longitudinis folii

descendens, ventralis usque 0.45-0.5; sinus dorsalis

atque saepe ventralis rimiformis vel in forma litterae

'V; apices loborum foliarium in (1) 2-3 cellulas sub-

quadratas vel brevi-triangulares terminantes.

HOLOTYPE — New Zealand, North Is., Tongariro

Natl. Park, Tree Trunk Gorge, along west bank of

Tongariro River, 15 Feb. 1995, Schuster 95-772.

Branching with strongly variable first branch

underleaves; on one stem, primary underleaves 1-,

2-, and 3-lobed. Leaves usually strongly asymet-

rically quadrifid; dorsal lobes separated by a sinus

descending usually (0.35) 0.4-0.45 the leaf

length, the lobes narrow, usually 3-4 cells broad

at base; dorsalmost sinus (between the two dorsal

lobes) usually narrow, often slitlike. Underleaf ba-

ses conspicuously cordate.

This variety (? microspecies) of the L. elobata-

fugax-novae-zelandiae "complex" appears to be

most closely allied to typical L. novae-zelandiae.

It shares with this the basic leaf form (both taxa

tend to have both leaf bases a little cordate); the

thick-walled leaf cells, with the marginal cells

with the free walls prominently thickened; and the

variability in form of the first branch underleaf.

In typical L. novae-zelandiae, it may be 1-, 2-, 3-,

or 4-lobed; in var. heterostipa, we have seen only

1-, 2-, or 3-lobed first branch underleaves (Fig.

25 : 8, 9).

In var. novae-zelandiae, leaves average rather

deeper lobed than in var. heterostipa: the wide

open ventral sinus usually decends well over 0.5

the leaf length (in var. heterostipa the disc, as a

whole, is higher and the ventral sinus descends

ca. 0.5 the leaf length). The two ventral lobes in

typical L. novae-zelandiae are separated by a rath-

70

FIELDIANA: BOTANY

er broadly V-shaped sinus (in var. heterostipa the

ventral sinus is often slitlike, and in fact, both

dorsal and ventral sinuses may be slitlike or nar-

rowly V-shaped). The bifid leaf associated with a

terminal branch in var. novae-zelandiae is divided

for 0.35-0.45 its length (in var. heterostipa it is

0.25-0.3(0.35) bifid; in var. novae -zelandiae the

disc, to the dorsal sinus, is ca. 6-10 cells high, 8-

10 cells high in var. heterostipa. Leaf-lobe apices

in var. novae-zelandiae are more drawn out, with

the terminal cells mostly elongated and ca. 2: 1 (in

var. heterostipa lobes are less attenuated and ter-

minal cells are usually under 2: 1 [Fig. 25 : 3—5]).

The asymmetry of the leaf in var. heterostipa

is often relatively slight (Fig. 25 : 1, 10, 11), and

the ventral lobes may approach the dorsal ones in

length and form. As in typical L. novae-zelandiae,

well-developed plants have strongly cordate dor-

sal leaf bases and may have ventral bases dilated

and subcordate (Fig. 25 : 10, 1 1). Underleaves on

mature shoots have cordate-auriculate bases.

The variation in lobe number of first branch

underleaves is not size related. Thus in Figure 25 :

8, four sectors of a single main stem are shown,

each with a branch base; the basal underleaves

range successively from 1 to 2 to 1 to 3. The same

variation occurs on another shoot (Fig. 25 : 9).

Thus the apparently consistent reduction of basal

branch underleaves to the monocrural condition

in L. acantha and L. elobata appears to have a

genetic basis.

Lepidozia novae-zelandiae var. minima Schust.,

var. nov.

Var. novae-zelandiae similis, sed lobis foliaribus et

trans latitudinem per 2-3 cellulas et trans longitudinem

per 4-6 cellulas extensis, amphigastrio rami primi 1- vel

2-lobo, amphigastriis disco altitudinem per 2 cellulas ex-

tenso instructis, lobis plerumque pauci-cellularibus et

basi latitudine per 2 cellulas extensis differt.

TYPE — New Zealand, South Is., Fiordland, Falls

Creek, Schuster 48802b.

Plants rather rigid and wiry, appearing small-

leaved (for the stem diameter), the shoots with

leaves to ca. 0.7 mm broad. Branching remotely

1 -pinnate, the branches short, at right angles to

stem, remote; first basal branch underleaf 1- or 2-

lobed. Stem rigid, only ca. 6 cells in vertical di-

ameter, with lateral merophytes 4-5 cells broad,

ventral merophytes 4 cells broad (underleaves in-

serted on 4 cell rows); cortical cells rather tan-

gentially flattened, very thick-walled; medulla of

rigid, strongly thick-walled cells. Leaves only

175-200 /im broad, asymmetrically 4-fid, the me-

dian sinus descending ca. 0.55-0.6, deeper than

the dorsal sinus (dorsal pair of lobes forming a

perceptible, ± parallel-sided pair); disc ca. 4 cells

high to dorsal sinus; dorsal lobes only 2-3 cells

broad at base, to 4-6(7) cells long, ending in 2(3)

single cells which are 1.5-2 X as long as broad.

Underleaves small, the disc usually 2 cells high,

the lobes 2(3) cells broad at base, (3)4 cells long.

Figure 26 : 4-13 illustrates this very small, rig-

id plant. The stem is formed of only 12-13 cor-

tical cell rows (Fig. 26 : 13) and, for a plant of

subg. Notholepidozia, is formed of a minimal

number of cell rows; the rigid, firm-walled me-

dulla is only 4-5 cells high.

Differing from var. novae-zelandiae in the 1—

2-lobed basal branch underleaves (Fig. 26 : 1, 8),

the fewer-celled lobes of leaves (Fig. 26 : 5, 12)

and underleaves (Fig. 26 : 4), the much lower leaf

disc (Fig. 26 : 5, 12), and underleaves (Fig. 26 :

4). In this plant, the underleaf bases are only ca.

8 cells broad and inserted on 4 rows of cortical

cells (Fig. 26 : 4), whereas in var. novae-zelandiae

underleaf bases are to 10-12 cells broad and in-

serted on 4-6 rows of cortical cells (Fig. 24 : 14).

Although leaf and underleaf margins in var. min-

ima are somewhat sinuous, owing to the generally

somewhat dilated septa separating marginal cells

(Fig. 26 : 5, 12) the marginal cells do not, or only

exceptionally, exhibit the constrictions seen in var.

novae-zelandiae (compare Fig. 26 : 4, 5).

Variety minima may be confused with L. acan-

tha because of the often unlobed basal branch un-

derleaves (Fig. 26 : 8), but that species has more

lanceolate, long-tapered leaf and underleaf lobes

formed of many more cells (Fig. 27 : 7); has deep-

ly bifid stem half-leaves associated with branches

(Fig. 27 : 5); and has a much more complex stem

anatomy, with ca. 22-24 rows of only weakly

firm-walled cortical cells (Fig. 27 : 2).

Lepidozia acantha Engel, sp. nov. Figure 27

Amphigastrium primarium ramalinum regulariter non

lobatum, subulatum; folia dimidia caulina bifida pler-

umque 0.45-0.55 longitudinis folii; folia ramulina pler-

umque 3-loba. Folia quadritida. 0.5-0.6 longitudinis fo-

lii; lobi lanceolati; sinus dorsalis usque 0.5 longitudinis

folii descendens; lobi in cellulas elongatas desinentes;

cellulae loborum maximam partem rectangulatae.

HOLOTYPE — New Zealand, North Is., Whaka-

rewarewa Thermal Reserve, Sept. 1 939, Hodgson

s. n. (F); isotype: (CHR).

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

71

FIG. 26. Lepidozia subg. Lepidozia; species and subgeneric criteria. (1-3) Lepidozia spp., showing the clearly if

finely botryoideal oil-bodies. (4-13) Lepidozia novae -zelandiae van minima Schust. 1. Laminar cells with oil-bodies

and, lower right, chloroplasts; oil-bodies 5 X 5-7 to 5 X 10-15 up to 6 X 10-17 yu,m, 4-6 per cell (X600). 2.

Laminar cells with oil-bodies and, below, chloroplasts; oil-bodies 4-5 to 4.5-5 X 6 to 5 X 7.5-9 up to 5 X 15 to 6

X 11 /nm, (1-2) 3-4 (5-6) per cell (X675). 3. Laminar cells with oil-bodies; oil-bodies 4.5 X 8 to 5 X 1 1-14 to 5.5

X 9-10 /xm, (2)3-5(6) per cell (X725). 4. Stem underleaf and ventral cortical cells; note size difference and 4 cell-

broad merophyte (X235; 100 /xm scale). 5. Dorsal half of leaf (X235; 100 yum scale). 6. Shoot sector, dorsal aspect

(X57.5; 1 mm scale). 7. Same, ventral aspect; note bifid first branch underleaf, BUL1, at base of Frullania-type

branch, FB; at HL, part of dorsal half-leaf visible (X57.5; 1 mm scale). 8. Same, but with basal branch underleaf

(bul1) unlobed (X57.5; 1 mm scale). 9. Shoot sector, dorsal aspect, the Frullania-type branch (FB) with stem half-

leaf (HL) at its base (X57.5; 1 mm scale). 10, 11. Stem leaves (X 1 10; 150 /im scale). 12. Dorsal fourth of leaf, distal

cells with cuticular papillae drawn in (X235; 100 /tm scale). 13. Stem cross section; at arrows, merophyte margins

(X337; 80 /im scale). (Fig. 1 from Schuster 74-128, New Zealand, Fiordland, Mt. Burns; 2 from Schuster 84-146,

New Zealand, Fiordland, Mt. Burns; 3 from Schuster 84-1027, New Zealand, Westland, Pakahi; 4-13 from Schuster

48802b, New Zealand, Fiordland, Falls Cr.)

72

FIELDIANA: BOTANY

FIG. 27. Lepidozia acanlha Engel. 1. Sector of main shoot, dorsal view. 2. Stem, cross section. 3. Sector of main

shoot with Frullania-type branch, ventral view (BUL = first branch underleaf). 4. Underleaves. 5. Half-leaves (HL),

3- and 4-lobed primary branch leaves (BL), and, at left, first branch underleaf (FBU) (cellular detail of first branch

underleaf at far left). 6. Leaves. 7. Leaf lobes (cuticular detail indicated). 8. Median leaf cells (cuticular detail

indicated). (All from type.)

Plants procumbent, in thin mats, slender, flex-

uous, with spreading branches, pale green, nitid,

delicately spinescent when dry, the shoots small

to medium, to 1 cm wide (stem to branch extrem-

ities). Branching nearly exclusively of Frullania

type, somewhat irregularly and loosely pinnate,

the branches slender, short to elongated and dis-

tinctly whiplike, long flagelliform and microphyl-

lous, the nonmicrophyllous leaves deeply subsym-

metrically 3-4-lobed; secondary branches not

seen; branch half-leaf symmetric, linear, bilobed

to 0.5; first branch underleaf undivided and su-

bulate, rarely bilobed, inserted on ventral or ven-

tral-lateral face of main axis and aligned with

branch underleaves. Ventral-intercalary branches

sporadic, leafy. Stems slender, flexuous. Leaves

when dry strongly ventrally secund, with the

lobes straight to slightly incurved; leaves when

moist strongly spreading, plane or (when lobes

slightly incurved) slightly concave, contiguous to

distant, with much of stem exposed in dorsal as-

pect, 0.45-0.65 mm long at longest point X 0.5-

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

73

0.65 mm wide at widest point, the insertion dis-

tinctly incubous; leaves ± symmetrical, deeply

and subequally 4-lobed, divided to ca. 0.5-0.6

(median sinus), the distance from dorsal sinus

base to insertion slightly greater than that from

ventral sinus to insertion. Lobes long and narrow-

ly attenuate, subequally divergent, the dorsal lobe

5-7 cells wide at the base, the ventral lobes sim-

ilar to the dorsal in size, the lobes entire, termi-

nating in a single cell or more commonly in a

uniseriate row of 2-4 cells; cells of uniseriate row

somewhat elongated (up to 2.5 : 1), evenly thick-

walled; cuticle of lobes closely and distinctly

short striate-papillose. Disc subsymmetrical, sub-

deltoid, 8-10 cells high at dorsal sinus, 5-8 cells

high at ventral sinus, the margins entire, the dorsal

margin straight (at most feebly ampliate), the ven-

tral straight, not much shorter than the dorsal.

Cells of disc middle moderately and evenly thick-

walled, subquadrate to somewhat longitudinally

elongated (particularly those aligned with 2nd

lobe), 18-23 /xm wide X 27-41 /^m long; median

basal cells larger, in one to several rows; cuticle

of disc closely striate papillose, becoming long

striate at disc base. Underleaves inserted on 5-7

rows of stem cells, rather large for plant size, ca.

1-1.5 X stem width, plane, spreading, symmetri-

cally quadrifid to ca. 0.5-0.6, the lobes long and

slenderly attenuate, ending in a single cell or more

often a uniseriate row of 2-3 cells, the cells of

uniseriate row quadrate to elongated; disc 4-6

cells high at median sinus; disc margins entire.

Androecia and gynoecia unknown.

DIFFERENTIATION — Resembling L. bidens in the

narrowly attenuate lobes, the elongation of unis-

eriate row cells, and the feebly ampliate dorsal

margin. It differs in the distinctly incubous leaf

insertion (Fig. 27 : 1), the deeply and subequally

lobed leaves (Fig. 27 : 6), the closely papillose cu-

ticle (Fig. 27 : 7, 8), the larger underleaves with

slenderly attenuate lobes, and the smaller size.

The species has the leaf shape and symmetry

of L. spinosissima but is a much smaller, procum-

bent plant, lacking the bipinnate branching, the

strongly ventrally secund primary branches, and

the transverse leaf insertion of that species.

The disc cuticle is closely striate-papillose (Fig.

27 : 8), as in L. laevifolia and its var. acutiloba.

Lepidozia acantha differs from the latter in the

narrowly attenuate leaf lobes and from the former

in the ± symmetrical leaves, the lobes subequally

spreading, and the ventral lobes not widely diver-

gent from the paired dorsal lobes. Suboptimal

populations of L. laevifolia may key here; see

comments under that species.

The species differs from all other New Zealand

members of Lepidozia except L. elobata by the

consistently undivided first branch underleaves

(Fig. 27 : 3, 5). The name is an allusion to these

structures, which resemble miniature "thorns" at

the base of each branch. The first branch under-

leaf is subulate in L. acantha, versus rather broad-

based and strongly tapered in L. elobata. The dis-

tinction of these two species is dealt with under

L. elobata. In L. elobata, the half-leaf is only 0.2-

0.25 bifid versus 0.4-0.45 in L. acantha; leaf

lobes are much less tapered, and their end cells

are scarcely elongated. It is notable that undivided

first branch underleaves are also found in many

Telaranea species, as is the presence of 3-lobed

branch leaves (Fig. 27 : 5); L. acantha can be dis-

tinguished from superficially similar diminutive

Telaranea species by the absence of a hyaloderm

(Fig. 27 : 2) and the papillose cuticle of the

leaves.

DISTRIBUTION-ECOLOGY — Known only from a

limited number of New Zealand collections.

Found at a single station on South Island at 990

m in Westland Province. North Island populations

are from the Rotorua area, at 1 85 m under manuka

and from edges of shaded paths in thermal spring

areas, and at ca. 30 m on a very rotted, decorti-

cated, bryophyte-covered log in a mixed broad-

leaf— podocarp lowland forest (Pakiri Scenic Re-

serve).

SPECIMENS SEEN— NEW ZEALAND. SOUTH IS-

LAND. WESTLAND PROY: Camp Creek, W of Al-

exander Range, 990 m, Reif C260A (F). NORTH IS-

LAND. SOUTH AUCKLAND PROV.: Rotorua, hot

springs area, Hatcher 34 (F); Waimangu, Rotorua, Child

412 (F). NORTH AUCKLAND PROV.: Pakiri Scenic

Reserve, SSE of Mangawhai Heads, ca. 30 m, Engel

20300 (F).

Lepidozia elobata Schust., sp. nov. Figure 28

Amphigastrium primarium ramulinum regulariter non

lobatum, ovato-lanceolatum; folia dimidia caulina bifida

sub 0.25 longitudinis folii attingens; folia ramulina 4-

loba. Folia quadrifida, sub 0.5 longitudinis folii attin-

gentia; lobi triangulares, sub lente 3 X longitudine lati-

tudinem aequante; sinus dorsalis plerumque usque 0.2-

0.3 longitudinis folii; lobi in cellulas non elongatas de-

sinentes; cellulae loborum maximam partem

subquadratae.

HOLOTYPE — New Zealand, North Is., Auckland

Harbor, Rangitoto Is., summit area, at tree base,

Schuster 95-401 (F).

74

FIELDIANA: BOTANY

FIG. 28. Lepidozia elobata Schust. 1. Apex of plant, ventral aspect, with three Frullania-type branches; at arrows,

unlobed basal branch underleaves (HUL1) (X36; 1 mm scale). 2. Stem half-leaf, at branch base (X54; 400 /im scale).

3, 4. Leaves (X54; 400 //.m scale). 5. Dorsal half of leaf (leaf at arrow, Fig. 8) (X 170; 150 /j.m scale). 6. Three leaves

(X68; 250 ^m scale). 7. Leaf + underleaf (X90; 200 ^m scale). 8. Three leaves (X54; 400 /urn scale). 9. Part of

main shoot, dorsal aspect, with Frullania-type branch; at HL, stem half-leaf (X68; 250 pm scale). (All from type.)

Plants creeping, light green, whitish when dry,

only feebly nitid when dry, small, the main shoots

with leaves ca. 430-640 to 700 /tm broad; shoot

apices and branch apices decurved, especially in

drying. Branches almost exclusively of Frullania

type, 1 -pinnate, rather irregularly and remotely so,

the branches usually of limited length, but spo-

radic branches elongating and becoming flagelli-

form; branch leaves normally similar to stem

leaves, quadrilobed like them; first branch under-

leaf inserted in-line with succeeding branch un-

derleaves, consistently unlobed, ovate lanceolate

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

15

to narrowly ovate, ending in a single cell, 80-85

X 130-140 /xm (flattened), usually standing stiffly

away from stem. Ventral-intercalary branches

sporadic, flagelliform. Stems with hyaloderm

well-developed, the cortical cells rather thin-

walled, oblong (on ventral merophytes ca. 20-22

/xm broad X 25-56 /xm long). Leaves adaxially

concave, the dorsal lobes decurved or deflexed,

the ventralmost lobe usually strongly incurved;

leaves of leading stems variable, typically quad-

rilobed for ca. 0.2-0.25(0.3) dorsally to 0.4-0.45

ventrally, the dorsal lobes usually somewhat

smaller than the ventral, the lobes acute but never

longly acute, ending in 1 or 2 superposed cells

that are not or weakly elongated; some to many

leaves almost symetrically quadrifid, the lobes

then subequal but the two outer lobes usually

somewhat smaller; leaf at dorsal base at most

weakly cordate. Cells in lobes subquadrate, av-

eraging ca. 11-14 X 13-16 /im, in leaf middle

scarcely larger, grading toward base to oblong and

15-18 X 22-27 [j,m', cells of leaf base much larg-

er, rather inflated, 21-30 X 27-50 /mi, forming a

small basal field; cuticle weakly papillose above,

weakly striolate basally. Underleaves inserted on

ca. 5 rows of stem cells, standing stiffly away

from stem or subsquarrose, under 0.45 quadrifid,

the lobes usually 2 cells broad basally.

No asexual reproduction (leaf lobe apices not

caducous). Otherwise unknown.

DIFFERENTIATION AND VARIATION — Similar to L.

acantha in (1) the basal branch underleaf is always

unlobed; (2) leaves do not consistently show a clear

division between a pair of dorsal lobes, set off from

the ventral lobes. At times the leaf tends to be al-

most equally quadrifid, although as in all Lepidozia

species, the disk is lower ventrally than dorsally.

Lepidozia elobata shows very considerable

malleability, often on one shoot, in form and lob-

ing of the leaves. The dorsal pair of lobes is typ-

ically separated by a shallower sinus than the si-

nus separating it from the ventral pair (Fig. 28 :

8), but in L. acantha, these two sinuses are usu-

ally subequal (Fig. 27 : 6). Lepidozia elobata

sporadically develops 5- or even 6-lobed leaves

(Fig. 28 : 6). On vigorous main stems the three

dorsal lobes are often subequal in size and length

and only the ventral lobe, which is typically

somewhat incurved, is appreciably smaller. How-

ever, even the ventral lobe may be rather large,

and the leaf may then be subequally quadrifid.

Figures 28 : 3, 4, 6-8 shows only part of the var-

iation seen.

In leaf form, especially form of the lobes and

depth of sinuses, L. elobata approaches L. pumila.

That species, however, has rather consistently

more asymmetrical leaves; the dorsal pair of lobes

is consistently much longer than the ventral pair

(compare Fig. 29 : 5). By contrast, Lepidozia

pumila has first branch underleaves consistently

2-4-fid.

Distinct from L. acantha in the relatively

broad-based, ovate-lanceolate first branch under-

leaf; the shorter and broader leaf bases, not long-

tapered as in L. acantha; the subisodiametric cells

of leaf-lobe apices; the obscurely papillose cuti-

cle; and the relatively high disk, about as high as

or higher than the length of the leaf lobes. In L.

acantha, branch leaves are typically trifid; in L.

elobata, they are almost uniformly quadrifid, like

the main axis leaves. We have checked branching

on several scores of plants: the Frullania-type

branches uniformly have an unlobed first branch

underleaf, and this is ovate or ovate-lanceolate,

relatively short and usually rather broad-based

compared with that of L. acantha.

The first branch underleaf exhibits limited vari-

ation. Usually it is broadly lanceolate (Fig. 28 :1,

2), but at times it is unusually small and only 2-3

cells broad at the base; the apex is often uniseriate

for 2-3 cells. On weaker shoots, the underleaf is

only 2 cells broad for 2-3(4) cell tiers and ends in

2 or 3 single cells. The largest underleaves are 4(5)

cells broad basally and contracted to an apex

formed of 2(3) superposed cells.

Underleaves in L. elobata are uniformly strong-

ly patent, often at right angles to the ventral stem

surface; they typically are seen almost edge-on

when unflattened plants are examined ventrally.

The flattened underleaf is ca. 0.4 quadrifid vs. ca.

0.5-0.6 quadrifid in L. acantha. Similarly, the si-

nuses on stem leaves (and stem half-leaves) are

uniformly much shallower in L. elobata (cf. Fig.

28 : 2-4, 7, 8, L. elobata; compare Fig. 27 : 5, 6,

L. acantha).

Lepidozia elobata is part of a complex of small

species that includes L. pumila, L. acantha, L. no-

vae-zelandiae, and L. bidens. It can be separated

from all taxa of that complex except L. acantha

by the consistently unlobed first branch under-

leaves.

Lepidozia pumila Engel, sp. nov. Figure 29

Plantae minutae vel breves, 365-490 /urn latae; discus

folii ad sinum dorsalem altitudine per 10-12 cellulas

extensus e cellulis quadratis vel parum longioribus quam

76

FIELDIANA: BOTANY

FIG. 29. Lepidozia pumila Engel. 1. Portion of main shoot with Frullania-lype branch (= FB; HL = half-leaf;

FBU = first branch underleaf), dorsal view. 2. Portion of main shoot with 2 Frullania-type branches, ventral view.

3. Portion of main shoot with Acromastigum- (= AB; note half underleaf) and Frullania-lype ( = FB) branches,

ventral view. 4. Three leaves. 5. Leaf, cellular detail. 6. Dorsal pair of leaf lobes (ms = median sinus). 7. Median

disc cells. 8. Marginal cells of dorsal margin of disc showing thickened free wall. 9. Underleaf (at same scale as Fig.

4). 10. Portion of stem with underleaf; note ventral merophyte width of 6 cells. 11. First branch underleaves. 12.

Half-leaf. (All from type.)

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

77

latioribus compositus; lobi dorsales cellulis series unis-

eriatae subisodiametris vel parum longioribus quam la-

tioribus instructi; cutis cellularis disci foliaris laeva; cel-

lulae lobi laeves vel non distincte papillosae; amphigas-

tria in 5-6 seriebus cellularum caulinarum inserta.

HOLOTYPE — New Zealand, South Is., Fiordland

Natl. Park, Moraine Creek, Hollyford Valley, ca.

400-650 ft., Schuster 4811 la (F); isotype: (CHR).

Plants closely prostrate, flexuous, with spread-

ing branches, green, nitid when dry, the shoots

minute to small, 365-490 //,m wide (leaf tip to

leaf tip), plant to 5 mm wide, including branches.

Branching nearly exclusively of Frullania type,

rather short, usually irregularly and distantly pin-

nate to ± regularly pinnate, the primary (and sec-

ondary) branches often becoming whiplike, fla-

gelliform, and microphyllous, the nonmicrophyl-

lous leaves ± symmetrically 4-lobed; primary

branches often developing into new leading

shoots; secondary branches occasional, 1—4 per

primary branch; branch half-leaf subsymmetrical,

cordate, 2-lobed to ca. 0.3; first branch underleaf

(l)2-4-lobed, inserted on ventral side of branch

base to the ventral-lateral side of junction of main

axis and branch, in both cases the first branch un-

derleaf aligned with underleaves of branch. Ac-

romastigum-type. branches (?) occasional. Ventral-

intercalary branching occasional, leafy. Stems

flexuous. Leaves cup-shaped (with the tips of the

lobes incurved and not visible in dorsal view) to

concave, contiguous, with much of stem visible

in dorsal view, somewhat longer than wide,

(0.25)0.4-0.5 mm long and wide, spreading, the

insertion weakly to distinctly incubous; leaves

usually strongly asymmetrical, 4(5)-lobed, the

leaves divided to ca. 0.45-0.55 (median sinus),

the distance from dorsal sinus base to insertion

much greater than that from ventral sinus to in-

sertion. Lobes attenuate, somewhat apiculate, the

dorsal lobes sometimes paired, the two ventral

shorter and somewhat divergent from the dorsal

lobes, the lobes entire, terminating in a uniseriate

row of 2-4 cells, the cells of uniseriate row ±

isodiametric or slightly longer than wide, thick-

walled; dorsal lobe (2)3-5 cells wide at base; cu-

ticle of lobes smooth or faintly and indistinctly

papillose. Disc asymmetrical, obliquely truncate

and deltoid, 10-12 cells high at dorsal sinus, 4-

6 cells high at ventral sinus, the margins entire,

the dorsal margin ± straight, abruptly cordate

near the base, the ventral margin shorter than the

dorsal, at times distinctly so. Cells of disc middle

evenly and distinctly thick-walled, occasionally

with small trigones, 20-33 ^tm wide and long;

median basal cells in 1 (locally 2) rows of en-

larged cells; marginal cells of disc and lobes with

a thickened outer wall; cuticle of disc smooth.

Underleaves inserted on 5-6 rows of stem cells,

spreading, short, broader than high, symmetrically

4-fid to ca. 0.4-0.55 (median sinus), the lobes

plane, slenderly acuminate, often only 2-3 cells

wide at base, entire, terminating in a uniseriate

row of 3—5 cells; disc 4-5(6) cells high at median

sinus, the margins plane, entire.

Plants dioecious. Androecia not seen. Gynoecia

on abbreviated ventral-intercalary branches issu-

ing from main stem; gynoecium base bulbous,

fleshy, polystratose and rhizoidous; bracts of in-

nermost series much larger than leaves, erect and

sheathing the perianth, the bracts deeply concave,

broadly ovate; apices 2-4-dentate-lobulate, each

lobule terminating in a single cell or a uniseriate

row of 2 cells, the cells of lobule base may have

a cell on each side that projects at its distal end

to form a crenulation, the lobules otherwise com-

posed of subrectangular to subsinuate cells; lam-

ina composed of ± regularly subrectangular cells,

the margin bordered by thin- to slightly thick-

walled cells of variable shape and orientation,

some hardly longer than wide, others ± sinuate-

rhomboidal, the apical or free end of marginal

cells slightly divergent and forming a short pro-

jection, the margin irregularly and often sparingly

crenulate, the margin otherwise entire except for

a sporadic single-celled tooth; bracteole similar in

size and form. Perianth rather long and prominent,

slenderly cylindrical-fusiform, straight, terete be-

low, obscurely trigonous above, distinctly and

deeply 3-plicate toward mouth, the perianth grad-

ually narrowing toward the strongly contracted,

shallowly 3-lobed mouth, the lobes composed of

± tiered, narrowly subrectangular cells, the dis-

talmost tier with cells that at the apical end are

laterally free for varying lengths, the lobes cre-

nate-subdenticulate; perianth cells ± regularly

subrectangular below mouth, and here the cuticle

is finely striate; perianth 2-4 stratose near base.

Seta with 8 rows of outer cells surrounding an

inner core of numerous much smaller cells. Cap-

sule oblong, the wall 38-41 /mi thick, of 3 layers;

outer layer of cells with two-phase development,

the longitudinal walls with sinuous, sheetlike

thickenings and several large nodulelike to spine-

like thickenings alternating with those that are de-

void of thickenings (or, rarely, are locally thick-

ened), the transverse walls usually devoid of

thickenings or sporadically with an isolated nod-

78

FIELDIANA: BOTANY

ule; semiannular bands sporadic; innermost layer

of cells ± tiered, narrowly rectangular, semian-

nular bands common, close and numerous, some-

times incomplete, not forked.

Spores 13.9-14.9 ^tm, exine brown, with dense,

rather coarse but low, sharply defined papillae and

simple or furcate vermiculate markings. Elaters

rigid, nontortuous, 7.7-9.6 yum wide, only slightly

tapering toward tips, bispiral to tips, the spirals

3.4-3.8 /Am wide.

DIFFERENTIATION — This smallest New Zealand

member of the laevifolia group is evidently rather

rare. Suboptimal expressions of L. laevifolia with

poorly developed cuticular papillae may well be

mistaken for this species. For further details, see

the discussion under L. laevifolia.

A minute phenotype of the species occurs in

median and upper elevations on cliff faces and

boulders and forms pure, tight, compact mats of

usually wirelike, upright shoots. Examples are En-

gel 1 7564 from a cliff face at 670 m on Mt. Car-

gill (Otago Prov.), and Engel 18668 from a sub-

alpine boulder at 1010 m in the Mt. Burns area

(Southland Prov.). Shoots are ca. 350-400 /mi

(not including branches), and leaves are minute

(400-500 /Am wide and long) and in dry plants

quite inconspicuous. Despite their seemingly im-

poverished condition, these specimens bear ma-

ture perianths.

Lepidozia pumila and L. bidens are closely re-

lated in many respects: they share similar leaf

shapes, with the dorsal pair of lobes much longer

than the ventral (compare Figs. 29 : 4, 5 and 30 :

2, 3); both have median-basal disc cells relatively

large versus distal and peripheral cells (Figs. 29 :

5; 30 : 3); both have leaf lobes acute, ending in

2-3 superposed cells. The former, however, is a

minute to small species (the smallest member of

sect. Notholepidozia), yet has a leaf disc 10-13

cells high at the dorsal sinus, while the latter is a

medium-sized plant but has a leaf disc 8-10 cells

high at the dorsal sinus. Also, L. pumila has un-

derleaves inserted on 5-6 rows of stem cells (Fig.

29 : 10), whereas L. bidens has underleaves in-

serted on 8 rows of stem cells (Fig. 30 : 10). The

species are for the most part separated ecologi-

cally: L. pumila typically occurs from 570 m into

the subalpine zone, whereas L. bidens is strictly a

forest species (on South Island only at low ele-

vations).

This species and L. spinosissima are the only

members of the genus Lepidozia known to have

ventral-terminal branching (Fig. 29 : 3). Branches

of this type are diagnostic for the genus Acro-

mastigum but otherwise occur only sporadically

in the family — in, for example, Kurzia mollis,

Neogrollea, and Paracromastigum. Taxa with this

branch type typically are erect (K. mollis) or, oc-

casionally, loosely prostrate (Acromastigum cavi-

folium Schust.), but are not closely prostrate as in

L. pumila.

DISTRIBUTION-ECOLOGY — Auckland Is., New

Zealand (North Is., South Is.). A species occurring

over a broad elevational gradient. It occurs in for-

ests ranging from 570 m to the upper reaches of

the forest and found corticolous or on shaded,

moist, cliff faces and boulders. It ranges up to

1 370 m in the subalpine zone on rock (particularly

with some soil accumulation) or in rock crevices.

Occasionally (type) in lower elevation forests

( 1 20-200 m) over wet humus of the shaded forest

floor, and then occurring with L. pendulina.

SELECTED SPECIMENS SEEN— AUCKLAND IS.: 1 mile

W of Trinity Cove, Carnley Harbour, Johnson 19/8

(WELT). NEW ZEALAND. SOUTH ISLAND.

SOUTHLAND PROV.: Fiordland Natl. Park, valley just

SW of Mt. Burns, S of South Branch of Borland Burn,

W of Monowai, 1010-1 170 m, Engel 18668 (F); Garvie

Range, East Dome, ca. 1370 m, Child H 3892 (F). OTA-

GO PROV.: S side of Mt. Cargill, just below summit, N

of Dunedin, ca. 670 m, Engel 17564 (F); Kakanui Range,

ca. 1525 m, Child H 2827 (F); Mt. Aspiring Natl. Park,

ridge below and W of Mt. Shrimpton, 1250 m, Engel

17857 (F). OTAGOAVESTLAND PROV. BOUNDARY:

Summit area of Haast Pass, 570 m, Engel 18003 (F).

WESTLAND PROV: Arthur's Pass Natl. Park, Lower

Otira Bridge, Engel 6799 (F). WESTLAND/CANTER-

BURY PROV. BOUNDARY: Arthur's Pass Natl. Park,

Arthur's Pass, near Temple Basin Ski area, Engel 6449A

(F). CANTERBURY PROV: Mt. Cook Natl. Park, Gov-

ernors Bush, SW of town of Mt. Cook, 760-800 m,

Engel 18193 (F); Arthur's Pass Natl. Park, immediately

below Punchbowl Falls, Engel 6900 (F); summit of Mt.

Tourlesse, Hatcher 1214 (F). MARLBOROUGH PROV:

Branch River, ca. 305 m, Child H 4367 (F). NORTH

ISLAND. WELLINGTON PROV: Tongariro Natl. Park,

Taranaki Falls Track, E of Whakapapa Village, 1080 m,

Engel 22441 (F). TARANAKI PROV: Pukeiti Bush,

near New Plymouth, Hatcher 261 (F). GISBORNE

PROV: Lake Waikaremoana, Hatcher 1308 (F).

Lepidozia bidens Engel, sp. nov. Figure 30

Plantae regulariter pinnatae, amplitudine medianae,

700-900 /ur» latae; folia lobis dorsalibus distincte binatis

instructa, anguste attenuata, in seriem uniseriatam e cel-

lulis elongatis (usque 2: 1 ) compositam terminantia; dis-

cus foliaris ad sinum dorsalem altitudine per 8-10 cel-

lulas extensus e cellulis plus minusve longitudinaliter

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

79

FIG. 30. Lepidozia bidens Engel. 1. Sector of main shoot with Frullania-lype branch, ventral view. 2. Leaves. 3.

Leaf. 4, 5. Dorsal pairs of leaf lobes. 6, 7. Dorsal lobes. 8. Underleaves. 9. Underleaf. 10. Portion of stem with

underleaf; note ventral merophyte width of 8 cells. 11. Two-, 3- and 4-lobed first branch underleaves (drawn to same

scale as Fig. 8). 12, 13. Half-leaves (3). 14, 15. Outlines of two shoots at low magnification. (Figs. 1 from Engel

19280, New Zealand, Nelson Prov., Paparoa Range, N side of Tiropahi or Four Mile River; 2, 3, 5-1 1,15 from type;

4, 12-14 from Child 358, New Zealand, South Is., Westland Prov., Fox Glacier.)

80

FIELDIANA: BOTANY

elongatis compositus; amphigastria in seriebus octo cel-

lularum caulinarum inserta.

HOLOTYPE — New Zealand, South Is., Nelson/

Westland Prov. boundary, Paparoa Range, S side

of Porarari River, up river from gorge and ca. 500

m WSW of ford on inland track to Bullock Creek,

10-20 m, Engel 19204 — c. per. (F); isotype:

(CHR).

Plants procumbent, in thin mats, slender, flex-

uous, with spreading branches, pale green, nitid,

delicately spinescent when dry, the shoots small

to medium, to 1.5 cm wide (stem to branch ex-

tremities). Branching exclusively of Frullania

type, regularly pinnate, the branches slender, short

to greatly elongated and distinctly whiplike, long-

flagelliform, microphyllous, the nonmicrophyl-

lous leaves subsymmetrically 4-lobed, the pri-

mary branches occasionally developing into new

leading shoots; secondary branches sporadic to

occasional, 1(2) per primary branch; branch half-

leaf symmetrical, linear (at times slightly broad-

ening toward base), 2 (rarely 3)-lobed to 0.3-0.45;

first branch underleaf (2)3-4-lobed, inserted on

ventral side of branch base or at juncture of

branch and main axis, aligned with branch under-

leaves. Ventral-intercalary branches not seen.

Stems slender, flexuous. Leaves when dry slightly

concave, with tips of lobes straight to slightly and

broadly incurved; leaves when moist spreading,

slightly concave, contiguous to distant, with much

of stem exposed in dorsal aspect, 0.45-0.5 mm

long X 0.35-0.5 mm wide, the insertion subtrans-

verse to weakly incubous; leaves distinctly asym-

metrical, unequally 4(5)-lobed, divided to ca. 0.5-

0.65 (median sinus), the distance from dorsal si-

nus base to insertion much greater than that from

ventral sinus to insertion. Lobes long and narrow-

ly attenuate (the dorsal lobes especially so), the

dorsal lobes parallel to somewhat divergent (rare-

ly approaching 90°), ± distinctly paired, 3-5 cells

wide at base, the ventral lobes often shorter,

curved and spreading (often at right angles to the

dorsal margin), the lobes entire, terminating in a

uniseriate row of 2-3 cells; cells of lobes and un-

iseriate row somewhat elongated (up to 2:1, but

the terminal 1-2 cells sometimes subquadrate),

evenly thick-walled; cuticle of lobes smooth to

faintly striate-papillose. Disc strongly asymmet-

rical, subdeltoid, 8-10 cells high at dorsal sinus,

4-5 cells high at ventral sinus, the margins entire,

the dorsal margin straight (at most feebly amplia-

te), the ventral straight, much shorter than the dor-

sal. Cells of disc middle moderately and evenly

thick-walled, ± longitudinally elongated, partic-

ularly those in the dorsal half of leaf, 1 8-3 1 fim

wide X 30-50 /tm long; median basal cells dis-

tinctly larger, in 1 to several rows; marginal cells

of disc and lobes with a thickened outer wall; cu-

ticle of disc smooth to faintly striate-papillose.

Underleaves inserted on 8 rows of stem cells,

small, ca. 1-1.1 X stem width, plane, strongly

spreading to squarrose, symmetrically quadrifid to

ca. 0.4-0.5, the lobes attenuate, ending in a single

cell or a uniseriate row of 2-3 cells, the cells of

uniseriate row often elongated; disc 4-6 cells high

at median sinus; disc margins entire.

Plants dioecious. Androecia not seen. Gynoecia

on abbreviated ventral-intercalary branches issu-

ing from main stem; bracts of innermost series

much larger than leaves, erect and sheathing the

perianth, the bracts deeply concave, narrowly

ovate to suborbicular; apices with 2-4 short, ir-

regular lobes composed of ± regularly rectangular

+ irregularly and feebly sinuate-rhomboidal cells,

the apical end of the marginal cells often feebly

to distinctly diverging, the lobes crenate-denticu-

late to spinose dentate; lamina composed of ±

regularly short-rectangular cells, the margin bor-

dered by cells of variable shape, some long and

narrow, others hardly longer than wide, the apical

or free end of marginal cells variously divergent

and forming a short projection or a tooth, the mar-

gin irregularly crenate-denticulate to the base, the

teeth at times composed of a few to several cells;

bracteole similar in size and form. Perianth long

and prominent, slenderly cylindrical-fusiform,

slightly curved, terete below, obscurely trigonous

above, distinctly and deeply 3-plicate toward

mouth, the perianth gradually narrowing toward

the strongly contracted, shallowly 3-lobed mouth,

the lobes composed of feebly tiered, long, narrow,

subrectangular to sinuate-rhomboidal, thick-

walled cells; mouth cells thick-walled, especially

at the summit, at the apical end laterally free for

varying lengths, the mouth thus crenate-denticu-

late; perianth cells ± regularly rather short sub-

rectangular below mouth, and here the cuticle is

long striate; perianth 2-stratose near base.

Sporophyte not seen.

DIFFERENTIATION — The species may be imme-

diately distinguished by the distinctly paired,

long, narrowly attenuate, dorsal lobes, which lend

a delicately spinescent appearance to dried plants

(Fig. 30 : 3-5). Related to this feature is the dis-

tinctive appearance of the half-leaves, which are

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

81

linear and deeply divided (up to 0.5; Fig. 30 : 12,

13). The ventral lobes are typically shorter than

the dorsal, curved and spreading, often at nearly

right angles to the dorsal margin (Fig. 30 : 2, 3).

The species also has distinctive areolation; medi-

an cells of the disc are often elongated, particu-

larly in the dorsal half of the leaf, and the median

basal cells are larger in contrast to those along the

dorsal and ventral margins (Fig. 30 : 3).

Lepidozia bidens belongs to a complex of small

to minute species that includes L. novae-zelan-

diae, L. acantha, and L. laevifolia; however, L.

bidens differs from the other three in its regularly

pinnate branching (Fig. 30 : 14, 15) and generally

larger size.

DISTRIBUTION-ECOLOGY — A forest plant of both

South and North Islands. On South Island it oc-

curs in wet, rich, lower-elevation forests (sea level

to 330 m) in banks of bryophytes, or it may be

corticolous (on, for example, Fuchsia). It also

may occur in open forests (again, below 330 m)

of, for example, Nothofagus fusca and Dacrydium

cupressinum, and here it occurs in large masses

at tree bases. On North Island it occurs, for ex-

ample, on rotted, decorticated, bryophyte-covered

logs in the mature podocarp forest at Whirinaki

Forest Park. The species reaches higher elevations

on North Island, occurring at 920 m on bryo-

phyte-covered wood in the beach-remu-totara

forests at Waikareiti Track (Urewera Natl. Park),

and (also Urewera Natl. Park), at 720 m, on the

sides of old Sphagnum mounds in a low, open,

boggy area with Sphagnum hummocks, Lepto-

spermum scoparium, and Dracophyllum subula-

tum at Waipai Swamp.

SELECTED SPECIMENS SEEN— NEW ZEALAND.

SOUTH ISLAND. OTAGO PROV.: Morrisons Creek, N

of Dunedin, 330 m, Engel 17709 (F). WESTLAND

PROV.: Cascade Road, Cascade ultramafic moraine, W of

Martyr Saddle, SSW of Jackson Bay, ca. 35 m, Engel

21775 (F); Fox Glacier, sea level, Child 358 (F); Westland

Natl. Park, Lake Wombat, 250 m, Engel 6671 (F). NEL-

SON PROV.: Paparoa Range, N side of Tiropahi or Four

Mile River, W side of Route 6 ca. 500 m N of bridge

over river, 145 m, Engel 19280 (F). NORTH ISLAND:

WELLINGTON PROV.: Tree Trunk Gorge, W bank of

Tongariro River, E border of Tongariro Natl. Park, boun-

dry of Kaimanawa State Forest Park, ca. 700 m, Engel

21191 (F). GISBORNE PROV: Urewera Natl. Park, Wai-

kareiti Track between track entrance and Lake Ruapani,

N of eastern extremity of Lake Waikaremoana, 650-920

m, Engel 20586 (F); ibid., Waipai Swamp, Waikareiti

Track between track entrance and Lake Ruapani, N of

eastern extremity of Lake Waikaremoana, 720 m, Engel

20601 (F). TARANAKI PROV: Pukeiti Bush near New

Plymouth, Hatcher 410 (F). SOUTH AUCKLAND

PROV: Whirinaki Forest Park, Waterfall Loop track, near

Whirinaki River, SSW of Minginui, Engel 20689 (F).

Sect. Kirkii Schust.

Lepidozia sect. Kirkii Schust., Beih. Nova Hedwigia

118: 197. 2000.

Plants primarily 1 -pinnate, but longer branches

sometimes with 1-2(3) secondary branches; Frul-

lania-type branches with basal branch underleaf

normally trifid, often with a few marginal teeth,

oriented "in line" with succeeding underleaves

and clearly on ventral side of plant. Leaves im-

bricate, clearly incubous, asymmetrical, with am-

pliate dorsal margin, relatively shallowly and sub-

equally quadrilobed (ventral sinus clearly deeper,

descending 0.35-0.5 leaf length), the dorsal mar-

gin of disc (usually) and dorsal 1-2 lobes (spar-

ingly), rather remotely, coarsely and variably

toothed.

TYPE — Lepidozia kirkii Steph.

Including L. kirkii and L. hirta. The two species

are clearly similar in most respects, including the

coarse, often oblique dentition of the dorsal leaf

margins and the large lacinium on each side of

the underleaf, so that underleaves appear 6-lobed.

Distinctive for the group is the position of the

initial branch underleaf; this is aligned with suc-

ceeding branch underleaves (Fig. 31 : 3, 4), and

the underleaf is very similar in form to succeeding

branch underleaves, differing chiefly in being tri-

lobed.

Lepidozia kirkii Steph. Figure 31

Lepidozia kirkii Steph., Spec. Hep. 3: 598. 1909.

Original material: New Zealand, South and North

Is., Krone, Beckett, Kirk (non vidi).

Lepidozia dentifolia Steph., Spec. Hep. 3: 599. 1909,

syn. nov. Original material: New Zealand, Jacksons

Track, Goebel (o!).

Plants ascending, rather flexuous, with ventrally

secund branches, pale green, the shoots medium,

to 2 cm wide, including branches. Branching

mostly of Frullania type, regularly pinnate, the

primary branches often becoming whiplike, fla-

gelliform, microphyllous, and rooting in the sub-

strate, the primary branches often developing into

new leading shoots; secondary branches occasion-

ally present, 1-3 per primary branch, often arising

in distal half of primary branch, the secondary

branches normally not longer than the primary;

branch half-leaf ± symmetrical to slightly asym-

metrical, broadly ovate, shallowly 2(3)-lobed to

0.3, irregularly toothed nearly to the base; first

82

FIELDIANA: BOTANY

FIG. 31. Lepidozia kirkii Steph. 1, 2. Three stem leaves and two underleaves (X32; 400 /im scale). 3. Shoot

sector, ventral aspect, with two Frullania-type branches; at HL, the dorsal half stem leaf (X32; 1 mm scale). 4. Shoot

sector, ventral aspect; the dorsal half stem leaf (HL) visible; at BUL, the first branch underleaf, which is already in

the "normal" position vis a vis the branch (X30; 700 /xm scale). 5. Shoot sector, dorsal aspect (X30; 700 /j.m scale).

6, 8. Cells of lobe apex, middle, and base (X153; 150 /im scale). 9. Stem leaf and adjacent first branch underleaf

(X30; 700 /im scale). 10. Stem cross section (X140; 200 ^m scale). 11. Cells of lobe middle (X ca. 780). (All from

Schuster 67-254, New Zealand, South Is., Westland, Omeroa River.)

branch underleaf 4(5)-lobed, inserted on ventral

to ventral-lateral (side facing shoot base) side of

branch and aligned with underleaves of branch.

Ventral-intercalary branching rare and sporadic,

leafy. Stems soft, flexuous, 10-12 cells in diam-

eter, the cortical cells in 1 layer of thick-walled

cells somewhat larger than the medullary cells;

medullary cells thick-walled, rather uniform in

size. Leaves rigid, concave, with incurved lobes,

imbricate and completely hiding stem in dorsal

view, 0.6-0.95 mm long at longest point, 0.8-1.4

mm wide at widest point, patent, the insertion dis-

tinctly incubous and slightly recurved at dorsal

end; leaves distinctly asymmetrical, unequally 4-

lobed, the leaves divided to ca. 0.4-0.55 (median

sinus), the distance from dorsal sinus base to in-

sertion greater than that from ventral sinus to in-

sertion. Lobes of differing shape, the dorsal pair

of lobes acute, the ventral lobe narrowly acumi-

nate to sublinear, the lobes ± divergent, the dorsal

pair of lobes partially united, the lobes entire or

sometimes with 1 to several short teeth, the dorsal

lobe 9-14 cells wide at base, the lobes terminat-

ing in a single cell or a uniseriate row of 2-3 cells

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

83

(the ventral often somewhat longer); cells of un-

iseriate row ± isodiametric to slightly longer than

wide, thick-walled. Disc distinctly asymmetrical,

22-27(30) cells high at dorsal sinus, 10-14 cells

high at ventral sinus; dorsal margin broadly am-

pliate, the dorsal margin and the confluent margin

of dorsal lobe irregularly (often doubly) dentate

or serrate (erose) by teeth of 1 to several cells (the

margin at times with a few weak teeth); ventral

margin much shorter than the dorsal, entire or

sometimes with a process. Cells of disc middle

evenly thick-walled, ± isodiametric to longitudi-

nally elongate, 14-22 X 18-30(36) /urn; median

basal cells enlarged, in 1-2 rows, with irregularly

thickened walls; cuticle smooth to indistinctly

striate-papillose. Underleaves spreading, symmet-

rically 4-fid to ca. 0.45-0.6 (median sinus), the

sinus bases somewhat reflexed, the lobes in-

curved, ± parallel, gradually attenuate, entire or

with a tooth or spine, usually blunt at the tip or

terminating in a single cell or short uniseriate row

of 2-3(5) cells; disc 10-15 cells high at median

sinus, disc margins somewhat reflexed, on each

side with a lobelike process distally, the under-

leaves then appearing 6-lobed.

Plants dioecious. Androecia on inconspicuous,

short, determinate, tightly spicate, cernuous ven-

tral-intercalary branches mostly from main shoot

and occasionally from primary branches; bracts

ventricose-cucullate, 2-lobed to ca. 0.3-0.4, the

lobes acute, the lateral margins entire or with a

tooth and a few slime papillae; antheridia 1-2 per

bract, the stalk biseriate. Gynoecia on abbreviated

ventral-intercalary branches issuing from main

stem; bracts of innermost series much larger than

leaves, erect and sheathing the perianth, the bracts

deeply concave, broadly ovate; apices with 4

short, ± regular lobes composed of ± regularly

rectangular and (especially at the lobe extremities)

irregularly sinuate-rhomboidal cells, the apical

end of the marginal cells often diverging and

forming a tooth, the lobes thus crenate-denticu-

late; lamina composed of ± regularly subrectan-

gular cells, the margin bordered by thin-walled

cells of variable shape and orientation, some hard-

ly longer than wide, others ± sinuate-rhomboidal,

the apical or free end of marginal cells variously

divergent and forming a short projection or a

tooth, the margin irregularly crenate-denticulate

(+ slime papillae) to the base, the teeth at most

of 3 superposed cells; bracteole similar in size and

form. Perianth long and prominent, slenderly cy-

lindrical-fusiform, slightly curved, terete below,

obscurely trigonous above, distinctly and deeply

3-plicate toward mouth, the perianth gradually

narrowing toward the strongly contracted, shal-

lowly 3-lobed, somewhat contorted mouth, the

lobes composed of irregularly sinuate-rhomboidal

cells that at the apical end are laterally free for

varying lengths, the lobes crenate-denticulate;

perianth cells ± regularly subrectangular below

mouth area and here the cuticle is conspicuously

striate; perianth 2-3 stratose near base.

Seta with 10-14 rows of outer cells surround-

ing an inner core of numerous much smaller cells.

Capsule elliptical to oblong, the wall 52-54 yarn

thick, of (4)5 layers; outer layer of cells with two-

phase development, the longitudinal walls with

sinuous, sheetlike thickenings and several large

nodules alternating with walls that are devoid of

thickenings (or are sporadically locally thick-

ened), the transverse walls usually devoid of

thickenings or sporadically have an isolated nod-

ule; innermost layer of cells with ± tiered, nar-

rowly rectangular, semiannular bands common,

close and numerous, sometimes incomplete, rare-

ly forked.

Spores 14.4-15.8 /urn, exine brown, with

coarse, high, sharply defined, widely spaced pa-

pillae and simple or furcate vermiculate markings

that sometimes coalesce (but do not delimit are-

olae). Elaters rigid, nontortuous, 9.6-11.5 /urn

wide, only slightly tapering toward tips, bispiral,

the spirals 3.4-3.8 /urn wide.

DIFFERENTIATION — The dorsal margin of the

leaf disc and the confluent margin of the dorsal

lobe are typically coarsely and irregularly (often

doubly) dentate or serrate (Fig. 31 : 1), rather than

armed with discrete spines (terminating in a uni-

seriate row of 2 or more cells), as in L. hirta (Fig.

32 : 2, 4, 10). Presence of teeth, even when few

in number, will distinguish the species. Lepidozia

hirta also differs in the more deeply lobed under-

leaves with the lobes often variously and irregu-

larly dissected.

DISTRIBUTION-ECOLOGY — Auckland Is., New

Zealand (South Is., North Is.).

The species occurs in forests over a variety of

substrates (rock, tree bark, rotten logs) over a

broad altitudinal range (ca. 30-1230 m). It is pre-

sent in the upper limits of Nothofagus menziesii

forests as well as in forests dominated by Noth-

ofagus solandri var. cliffortioides. It also extends

to the upper reaches of the subalpine zone (1040-

1110 m), where it forms tufts on rock. In forested

sites it occurs with Goebeliella, Lepicolea scolo-

pendra, Lepidolaena taylori, Bazzania adnexa,

84

FIELDIANA: BOTANY

and others. In the southern sectors of South Is-

land, it occurs at 100 m or lower; for example,

the species occurs over soil of vertical, bryophyte-

covered banks in Nothofagus truncata forests (90

m in Nelson Prov., Engel 21545) to the matai-

totara flood plain forests (110 m in Westland

Prov., Engel 18923), where it is corticolous. On

North Island it is may be found in open forests

dominated by Weinmannia racemosa, Kunzea er-

icoides, and Agathis australis, and it occurs over

bare rock of cliff faces (820 m in South Auckland

Prov., Engel 22409).

SELECTED SPECIMENS SEEN— AUCKLAND IS.: Head

of North Arm, Carnley Harbour, 15 m, Johnson 22/8

(welt). NEW ZEALAND. SOUTH ISLAND. SOUTH-

LAND PROV.: Fiordland Natl. Park, off track to Island

Lake, just W of Borland Saddle, S of South Branch of

Borland Burn, W of Monowai, 800 m, Engel 18642 (F);

ibid., near Lake Thompson Hut, on track to George

Sound, W of Lake Te Anau, ca. 290 m, Schuster 48297c

as L. ulothrix (F). OTAGO PROV.: Fiordland Natl. Park,

head of Lake McKerrow, near McKerrow Hut, Hatcher

1446 (F). Mt. Aspiring Natl. Park, Blue Valley Track,

above Blue River just N of confluence with Makaroa

River, 430-480 m, Engel 21920 (F). WESTLAND

PROV: Jackson Bay, between confluence of Jackson

River and Arawata River and Lake Ellery, off Jackson

River Road, ca. 30 m, Child H4267 (F); Mt. Aspiring

Natl. Park, Cross Creek, 1.1 km N of Haast Pass, 540

m, Engel 21885 (F); Westland Natl. Park, Franz Josef

Glacier Valley, off Roberts Point Track, SW of Mt.

Gunn, ca. 510-570 m, Engel 18136 (F); ibid., along Gil-

lespies Cook River Road, between Tornado Creek and

Whelan Creek, Engel 6604 (F); near Waiho River be-

tween Lake Wombat Terrace and Canavans Knob, NW

of town of Franz Josef Glacier, off Hwy 6, 110m, Engel

18923 (F); Camp Creek, W of Alexander Range, 500-

905 m, ReifC62D, C229F (F); Arthur's Pass Natl. Park,

Kelly Range, off track to Carroll Hut, above Kellys

Creek, N of Otira, 1,040-1,1 10 m, Engel 18431 (F); Pa-

paroa Range, road to Sewell Peak, 710 m, Engel 19067

(F). WESTLAND/CANTERBURY PROV. BOUND-

ARY: Arthur's Pass Natl. Park, Arthur's Pass, near Tem-

ple Basin Ski area, Engel 6495— c. 6 (F). NELSON

PROV: Track to German Terrace, 6 km SSE of Westport

on Nine Mile Road, 90 m, Engel 21545 (F). NORTH

ISLAND. WELLINGTON PROV: Tongariro Natl. Park,

Blyth Track, along small stream ca. 0.5 km from Ohak-

une Mt. Road, ca. 1230 m, Engel 21306 (F). TARAN-

AKI PROV: Pukeiti Bush, near New Plymouth, Hatcher

249 (F). SOUTH AUCKLAND PROV: Coromandel

State Forest Park, ridge just W of summit of Table Mt.,

820 m, Engel 22409 (F).

Lepidozia hirta Steph. Figure 32

Lepidozia hirta Steph., Spec. Hep. 3: 599. 1909. Orig-

inal material: New Zealand, without specific loc.,

Kirk (o!).

Lepidozia angelii Gott. ex Steph., Spec. Hep. 3: 596.

1909. Original material: New Zealand, without spe-

cific loc., (Angel) (o!).

Plants delicate, loosely procumbent, rather flex-

uous, soft and plumose, with weakly ventrally se-

cund branches, pale green, the shoots medium, to

1 .7 cm wide, including branches. Branching most-

ly of Frullania type, closely and regularly pin-

nate, primary branches becoming whiplike, fla-

gelliform, microphyllous, and rooting in the sub-

strate; secondary branches rarely present; branch

half-leaf ± symmetrical to slightly asymmetrical,

broadly ovate, shallowly 2-lobed, ± regularly spi-

nose-dentate nearly to the base; first branch un-

derleaf 4(5) lobed, inserted on ventral to ventral-

lateral (side facing shoot base) side of branch and

± aligned with underleaves of branch; second

branch underleaf in a strong morphologically ven-

tral position. Ventral-intercalary branching occa-

sional, stoloniform. Stems rather soft, flexuous.

Leaves rigid, concave, with incurved lobes, im-

bricate and nearly or completely hiding stem in

dorsal view, 0.4-0.5 mm long at longest point,

0.6-0.8 mm wide at widest point, patent, the in-

sertion distinctly incubous and feebly recurved at

dorsal end; leaves distinctly asymmetrical, un-

equally 4-lobed, the leaves divided to ca. 0.4-0.65

(median sinus), the lobes ± divergent, the dis-

tance from dorsal sinus base to insertion greater

than that from ventral sinus to insertion, the si-

nuses gradually becoming deeper ventrally. Lobes

of differing shape, the dorsal pair of lobes cau-

date, the ventral lobe attenuate, often spreading as

a claw, the dorsal lobes not noticeably paired, the

lobes entire or with a single prominent spine or

with a few small, often opposing teeth, the dorsal

lobes (5)9-1 1 cells wide at base, terminating in a

uniseriate row of 2-3(6) cells (the ventral often

somewhat longer); cells of uniseriate row ± iso-

diametric to somewhat longer than wide (2:1 or

less), not noticeably longer than basal cells in

lobe, thick-walled. Disc weakly asymmetrical,

10-16 high at dorsal sinus, 5-8 cells high at ven-

tral sinus; dorsal margin broadly curved to subam-

pliate, the dorsal margin and the confluent margin

of the dorsal lobe ± regularly spinose-dentate

with 3-6 discrete, short, acuminate, often curved,

multicellular spines (the margin often also with

several few-celled teeth); ventral margin much

shorter than the dorsal, entire. Cells of disc middle

thick-walled, ± isodiametric to elongate, 12-18 X

18-25 /Ltm; median basal cells enlarged, in 1-2

rows, with trigones and irregularly thickened

walls; cuticle striate-papillose. Underleaves

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

85

FIG. 32. Lepidozia hirta Steph. 1. Sector of main shoot, dorsal view (hi = half-leaf). 2. Leaves. 3. Dorsalmost

lobe of leaf (s — sinus). 4. Dorsal margin of leaf (B = leaf base). 5. Half-leaf. 6, 7. Underleaves. 8. Outline of plant

at low magnification. 9. Seta, cross section. 10. Two leaves and to right, two underleaves, all at same scale. (Figs.

1-8 from Engel 19205, New Zealand, South Is., Nelson/Westland Prov. boundary, Paparoa Range, S side of Porarari

River; 9 from Hatcher 136, New Zealand, South Is., Canterbury Prov., Mt. Tourlesse; 10 from Braggins 94/241, New

Zealand, North Auckland Prov., SE corner of Waipoua Forest.)

spreading, symmetrically 4-fid to ca. 0.55-0.8

(median sinus), the sinus bases somewhat re-

flexed, the lobes broadly incurved, somewhat di-

vergent, narrowly attenuate, variable: entire to

variously and irregularly dissected (with one to

several lobuliform spines), the lobes terminating

in a uniseriate row of 3-6 cells; disc 6-9 cells

high at median sinus, disc margins on each side

with a lobelike spine, the underleaves then ap-

pearing 6-lobed.

Plants dioecious. Androecia not seen. Gynoecia

on abbreviated ventral-intercalary branches issu-

ing from main stem; bracts of innermost series

much larger than leaves, erect and sheathing the

86

FIELDIANA: BOTANY

perianth, the bracts deeply concave to canalicu-

late, ovate to suboblate; apices with 2-4 teeth or

short, irregular lobes or unlobed, the distal cells

of bracts as well as cells of teeth and lobes com-

posed of irregularly sinuate-rhomboidal cells that

at their apical end are laterally free for varying

lengths, the lobes crenate-denticulate; lamina

composed of ± regularly subrectangular cells, the

margin bordered by thin-walled cells of variable

shape and orientation, some hardly longer than

wide, others ± sinuate-rhomboidal, the apical or

free end of marginal cells variously divergent and

forming a short projection or tooth, the margin

irregularly and at times sparingly crenate-dentic-

ulate to the base, the teeth at most of 2 superposed

cells; bracteole similar in size and form. Perianth

long and prominent, slenderly cylindrical-fusi-

form, slightly curved, terete below, obscurely tri-

gonous above, distinctly 3-plicate toward mouth,

the perianth gradually narrowing toward the dis-

tinctly contracted, obscurely ca. 6-lobulate mouth,

the lobules composed of irregularly sinuate-rhom-

boidal, thick-walled cells that at the apical end are

laterally free for varying lengths, the lobules thus

irregularly crenate-denticulate; perianth cells ±

regularly subrectangular below mouth area; peri-

anth 2 stratose near base.

Seta with 9 rows of outer cells surrounding an

inner core of numerous much smaller cells. Cap-

sule oblong, the wall 50-53 /im thick, of 4 layers;

outer layer of cells with two-phase development,

the longitudinal walls with sinuous, sheetlike

thickenings and several low, broad nodules alter-

nating with walls that are devoid of thickenings,

the transverse walls usually devoid of thickenings

or sporadically have 1-2 isolated nodules; inner-

most layer of cells ± tiered, narrowly to rather

broadly rectangular, the longitudinal walls with

nodular to spinelike thickenings and, often, some-

what sinuous sheetlike thickenings, semiannular

bands sporadic, never forked.

Spores 38.4-42 yum, exine yellow-brown, thin,

with low, dense, close papillae and short, some-

times branched, vermiculate markings that some-

times coalesce but do not delimit areolae. Elaters

rigid, nontortuous, 24-30 /urn wide, only slightly

tapering toward tips, bispiral, the spirals 9.6-10.8

/am wide.

DIFFERENTIATION — Lepidozia hirta differs from

L. ulothrix in its smaller leaves, its closely and

regularly pinnate branching (Fig. 32 : 8), the

branches often becoming whiplike and rooting in

the substrate. Typically, L. ulothrix is a more ro-

bust plant, with distant, abbreviated, somewhat

curved branches.

For comments distinguishing this species from

L. ulothrix and L. kirkii, see under those species.

Among the traditionally cited synonyms of L.

ulothrix are two Stephani names that apply to this

taxon, L. hirta and L. angelii, published simulta-

neously in 1909. The types of both names have

been examined and are quite similar; L. hirta is

considered the more appropriate name.

DISTRIBUTION-ECOLOGY — Endemic to New

Zealand (South Is., North Is.). It occurs, for ex-

ample, at the edges of rills in the subalpine zone

in areas with mostly snow tussock, but with some

Dracophyllum longifolium, Celmisia, Empodisma

minor, and others (Engel 19022). At the Manga-

whero River site (Engel 21288), it occurred in low

Nothofagus solandri var. cliffortioides forest with

steep, moist, peaty, mossy banks; in the Paeroa

Range (Allison H2995), it occurred on very old

logs in an upper-elevation forest. At the Waipoua

Forest site (Braggins 94/241), the species oc-

curred over rotted wood in a very wet Weinman-

nia silvicola-Syzygium maire forest.

SELECTED SPECIMENS SEEN— NEW ZEALAND.

SOUTH ISLAND. WESTLAND PROV.: Paparoa

Range, ridge immediately N of Sewell Peak, 890 m,

Engel 19022 (F). NELSON/WESTLAND PROV.

BOUNDARY: Paparoa Range, S side of Porarari River,

up river from gorge and ca. 500 m WSW of ford on

inland track to Bullock Creek, 10-20 m, Engel 19205—

c. sporo. (F). CANTERBURY PROV.: Mt. Tourlesse,

Hatcher 136— c. sporo. (F). NORTH ISLAND. WEL-

LINGTON PROV: Tongariro Natl. Park, west-facing

steep bank of Mangawhero River, off Ohakune Mt.

Road, ca. 1200 m, Engel 21288 (F). SOUTH AUCK-

LAND PROV: Paeroa Range, S of Rotorua, ca. 915 m,

Allison H2995 (F). NORTH AUCKLAND PROV: SE

corner of Waipoua Forest, Mataraurau Plateau, 540 m,

Braggins 94/241 (AKU).

Subg. Austrolepidozia Schust.

Lepidozia subg. Austrolepidozia Schust., Beih. Nova

Hedwigia 1 18: 197. 2000. Type: Lepidozia ulothrix

(Schwagr.) Lindenb.

Lepidozia HI. Incisae Lindenb. in G. L. & N., Syn.

Hep. 210. 1845, syn. nov. Lectotype (nov.): Lepi-

dozia ulothrix (Schwagr.) Lindenb.

Plants normally 1 -pinnate, the branches of lim-

ited length, sporadically to frequently the lower

becoming flagelliform; stem half-leaf ovate, shal-

lowly (to 0.35-0.4) bilobed, lobes with 1-3 cilia,

disc copiously spinose-dentate; basal branch un-

derleaf variably (2)3-4-lobed, bearing or ending

in 1-2(3) cilia. Stem with hyaloderm relatively

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

87

distinct, weakly firm-walled, the medulla relative-

ly uniform, smaller-celled, prominently thick-

walled. Leaves asymmetrically ovate, the dorsal

margin strongly convex and ampliate, the ventral

margin short, straight to concave-sided; dorsal si-

nus descending ca. 0.4-0.45 leaf length, the disc

high dorsally; ventral sinus descending less than

0.65 the leaf length; dorsal margin copiously cil-

iate-dentate to ciliate, the cilia mostly shorter than

those of lobes; ventral margin usually eciliate;

dorsal 1-2 lobes mostly with (1)2-3(4) cilia or

teeth, mostly alternate, sporadically in 1(2) op-

posed pairs; ventral 1-2 lobes with mostly only

1-2(3) long cilia, alternate. Cells in disc and cilia

thick-walled, rigid, in cilia in large part strongly

elongated (2-5: 1); cuticle smooth to obscurely

striolate. Underleaves obdeltoid to obtrapezoidal,

0.6-0.7 quadrifid; disc 1 -ciliate to edentate; lobes

with 1-3 cilia, if 2 often opposed.

Apparently monotypic.

We have debated placing Austrolepidozia as a

mere section of subg. Chaetolepidozia Schust.12

Both groups bear strongly spinose-ciliate leaves,

the cilia formed in large part of prominently elon-

gated, rigid cells. Chaetolepidozia, s. str., is quite

different in (1) the very deeply quadrifid leaves

in which the dorsal lobe is not strongly ampliate

basally; (2) the lobes all freely ciliate, even the

ventral with up to 10-12 cilia, often secondarily

bifid; (3) cilia, except for the distalmost, usually

clearly oriented in opposed pairs; (4) underleaves

freely ciliate, each usually with (1)2 pairs of op-

posed cilia; (5) stem half-leaf at branch bases nor-

mally trifid, asymmetrical, the sinuses descending

to 0.75-0.8, the lobes with (1)2-3 pairs of op-

posed cilia; and (6) basal branch underleaves

mostly deeply bifid, sporadically one lobe again

bifid, with sinuses descending to within 0.2-0.25

of the base, freely ciliate with often opposed pairs

of long cilia.

Lepidozia ulothrix (Schwagr.) Lindenb. Figures

33 and 34

Jungermannia ulothrix Schwagr., Hist. Muse. Hep.

Prodr. 21. 1814. Lepidozia ulothrix (Schwagr.) Lin-

denb. in G. L. & N., Syn. Hep. 210. 1845. Masti-

gophora ulothrix (Schwagr.) Trev., Mem. 1st.

Lomb. Sci. Lett. III. 4: 416. 1877. Original mate-

rial: "N. Hollandia," without specific locality,

"Herb. Ldbg," sin. coll. (G!).

12 For subg. Chaetolepidozia, see Schuster (2000).

Piippo (1984) recognized the group as Lepidozia at the

section level.

Jungermannia albula Hook. f. & Tayl., London J.

Bot. 3: 387. 1844, syn. nov. Lepidozia albula

(Hook. f. & Tayl.) G. L. & N., Syn Hep. 211. 1845.

Mastigophora albula (Hook. f. & Tayl.) Trev.,

Mem. 1st. Lomb. Sci. Lett. III. 4: 416. 1877. Orig-

inal material: Auckland Is., "on Gottschea lehman-

niana," Nov., 1840, Hooker (FH!).

Plants rather delicate, flexuous, with weakly

ventrally secund branches, pale olive green to

whitish green, the shoots medium-sized, to 2 cm

wide, including branches. Branching exclusively

of Frullania type, distantly markedly and regular-

ly pinnate, the primary branches short, somewhat

curved or hooked at the tip, occasionally becom-

ing flagelliform but not extensively so; secondary

branches only sporadically present, then 1 per pri-

mary branch, the secondary branches not longer

than the primary; branch half-leaf asymmetrical

to ± symmetrical, ovate, 2-lobed, regularly ciliate

nearly to the base; first branch underleaf (2)3-4

lobed, inserted on ventral-lateral (side facing

shoot base) side of branch (often at extreme base

of branch) and aligned with underleaves of

branch; second branch underleaves in a ± strong

morphologically ventral position or the 2nd-4th

cycles of branch underleaves gradually becoming

truly ventral. Ventral-intercalary branching lack-

ing. Stems rather soft and flexuous, 10-13 cells

in diameter, the cortical cells in 1 layer of weakly

to clearly thick-walled cells (the exposed wall

sometimes particularly thickened) that are slightly

to conspicuously larger than medullary cells;

medullary cells distinctly thick-walled in cross

section, often with marked trigonelike thicken-

ings. Leaves rigid, strongly concave, with in-

curved lobes lending the leaves a ± cuplike as-

pect, imbricate and nearly or completely hiding

stem in dorsal view, 0.4-1.2 mm long at longest

point, 0.7-2.2 mm wide at widest point, patent,

the insertion distinctly incubous and slightly re-

curved at dorsal end; leaves distinctly asymmet-

rical, unequally 4-lobed, the leaves divided to ca.

0.4-0.65 (median sinus), the lobes suberect or

only slightly divergent, the distance from dorsal

sinus base to insertion much greater than that

from ventral sinus to insertion, the dorsal lobes

not noticeably paired, the sinuses gradually be-

coming deeper ventrally. Lobes of differing

shape, the dorsal pair of lobes caudate, the ventral

lobe linear to attenuate, the lobes with 1 to (less

often) several pairs of opposing spines (1[2], non-

opposing spines per lobe in suboptimal plants) the

dorsal lobes 7-10 to 13-15 cells wide at base, the

ventral lobe 6-9 cells wide at base, the lobes ter-

88

FIELDIANA: BOTANY

Xo^o^sAcAJV

^>OC?OC

^OP

FIG. 33. Lepidozia ulothrix (Schwaegr.) Lindenb. 1. Part of main axis, ventral aspect, with bases of two Frullania-

type branches; at BUL1, first branch underleaf; at BUL2, second branch underleaf (X34; 1 mm scale). 2. Part of main

stem with underleaf; at right base of Frullania-type branch (FB) with (BUL1) basal branch underleaf; at UL, stem

underleaf (X34; 1 mm scale). 3, 5. Main stem leaves (X45; 700 /j,m scale). 6. Stem half-leaf associated with branch

(X45; 700 /Am scale). 7. Optimal-sized basal branch underleaf (X45; 700 /tin scale). 8, 9. Stem underleaves (X45;

700 /urn scale). 10. Apex of dorsal leaf lobe (X195; 150 /Am scale). 11. Tip of ventral leaf lobe (X195; 150 urn

scale). 12. Part of dorsal leaf margin (X 195; 150 /urn scale). 13. Stem cross section and surface view of cortical cells

(X195; 150 /Am scale). 14. Cladograph; double lines = leafy shoot sectors; single lines = flagella; at fb = initiation

of flagellar branch (X 1.75). (All from Schuster 95-1924, New Zealand.)

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

89

FIG. 34. Lepidozia ulothrix (Schwaegr.) Lindenb. 1. Sector of main shoot, dorsal view. 2. Leaves. 3. Underleaf;

at right, leaf. 4. Underleaves. 5. Sector of dorsal margin of leaf. 6. Portion of dorsal lobe. 7. Distal sector of dorsalmost

leaf lobe. 8. Median disc cells of leaf. 9. Sector of main shoot with Frullania-type branch, ventral view (BUL = first

branch underleaf). 10. Half-leaf. 11. First branch underleaves. 12. 8 Bract (DM = dilated dorsal margin, slime

papillae at arrows). (Figs. 1, 2, 4-6, 8-11 from Fife 8987, New Zealand, South Is., Arthur's Pass Natl. Park, Bealey

Glacier Track; 3, 7 from type of L. albula.)

90

FIELDIANA: BOTANY

minating in a uniseriate row of 5-7 to 8-10(15)

cells (the ventral typically the longest); cells of

uniseriate row capillary, elongated (3-4: 1 ), thick-

walled and with the septa thickened and swollen.

Disc distinctly asymmetrical, primarily due to

strong dilation of dorsal sector, the dilated portion

up to a third of the disc area, the disc 18-26 cells

high at dorsal sinus, 6-13 cells high at ventral

sinus; dorsal margin and the confluent margin of

the dorsal lobe with 7-8 to 12-19 spinose teeth

or cilia that terminate in a uniseriate row of 2-

5(6) cells; ventral margin much shorter than the

dorsal, entire or sporadically with 1-2 well-de-

veloped spines or laciniae. Cells of disc middle

evenly thick-walled, elongate, 12-20 /im X 24-

42 ^im; cells in the ampliate sector and median

lobe cells shorter, 14-18 /im X (17)20-26 /on;

median basal cells enlarged and in a rather distinct

field; cuticle smooth to finely striate-papillose.

Oil-bodies 2-4 per cell, ovoid or globose, botry-

oidal, mostly 4-6 yum in greater diam. Underleav-

es 1.5-2 X stem width, spreading, symmetrically

basically 4-fid to ca. 0.45-0.65 (median sinus),

the sinus bases plane, the lobes distinctly broadly

incurved, the lobe base ± parallel-sided, 8-9 cells

wide, the summit of median lobes truncate and ±

regularly ciliiform 2-3-fid, the distal sector seta-

ceous, terminating in a uniseriate row of 5-8 to

8-13 cells; disc 10-13 cells high at median sinus,

disc margins on each side with 1-2 cilia.

Plants dioecious. Androecia on inconspicuous,

short, determinate, tightly spicate, cernuous ven-

tral-intercalary branches from main shoot and pri-

mary and secondary branches (the androecial

branches often not exceeding tips of underleaf

lobes); bracts ventricose-cucullate, 2-lobed to ca.

0.4, the lobes acuminate, terminating in a uniser-

iate row of 2—4 elongated cells, the lobe margins

with several spinose teeth; dorsal margin of disc

slightly dilated in basal sector to form a feebly to

moderately rounded projection, the dilated portion

with several slime papillae; ventral margin of disc

with several spinose teeth or laciniae; antheridia

1 per bract, the stalk biseriate. Gynoecia not seen.

DIFFERENTIATION AND VARIATION — New Zea-

land representatives of the genus Lepidozia with

margins of the leaf disc and lobes armed with dis-

crete spines and/or cilia heretofore have been

treated as a single species, L. ulothrix. In fact, in

New Zealand, three distinct elements are repre-

sented: L. ulothrix s. str. (including L. albula), L.

kirkii, and L. hirta. The first is broadly distributed

in Australasia, the last two are endemic to New

Zealand.

Lepidozia ulothrix is a robust plant, with dis-

tant, abbreviated, somewhat curved branches. The

leaf armature of L. ulothrix is more copious than

in L. hirta, with longer and more numerous

spines, and/or cilia on the dorsal margin and con-

fluent margin of the dorsal lobe, the lobe margins

are armed with several, often opposing, spines,

and the cells of the uniseriate row are more elon-

gated.

The types of Jungermannia ulothrix and J. al-

bula are somewhat less well developed than in

some populations of the species, such as Fife

8987, which display a more pronounced devel-

opment of armature on the dorsal leaf margin and

lobes. These often have larger leaves with longer,

more numerous (up to 19) cilia on the dorsal mar-

gin and confluent margin of the dorsal lobe, and

leaf lobes with one to several pairs of opposing

cilia, and terminating in a uniseriate row of as

many as 14 cells.

More commonly the dorsal margin and the con-

tiguous margin of the dorsal lobe bear only 1 1-

1 3 cilia or teeth, those of the disc margin shorter

than those of the lobe. Such plants may bear only

1-3 cilia of the dorsal lobe (Fig. 33 : 3-5), and

these are only 6-7 cells long (Fig. 33 : 10, 11).

DISTRIBUTION-ECOLOGY — Auckland Is., New

Zealand (South Is.), Tasmania, southeast Austra-

lia.

Known from scattered sites in the southern half

of the South Island. In the Haast River area (sea

level) the species occurs in mucky niches at the

sides of mounds in open swampy areas with

Sphagnum and scattered Leptospermum, etc. It is

also present in the Cascade ultramafic moraine (S

of Haast along Cascade Road), an area at low el-

evation (ca. 35-135 m) characterized by ultra-

mafic rocks and outcrops with rather open vege-

tation consisting mainly of Gleichenia, Lycopo-

dium, Juncus, the lichen Cladina, and scattered

Leptospermum; at this site, the species is terrico-

lous among Restionaceae. The plant from Bealey

Glacier Track occurred at a considerably higher

elevation, 915 m, at the bottom of an earth hum-

mock, forming loose wefts over duff in a rain for-

est on a flat river terrace dominated by Nothofa-

gus solandri, Coprosma pseudocuneata, Draco-

phyllum longifolium, and Phyllocladus alpinus.

SELECTED SPECIMENS SEEN— NEW ZEALAND.

SOUTH ISLAND. WESTLAND PROV.: Cascade Road,

Cascade ultramafic moraine, W of Martyr Saddle, SSW

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

91

of Jackson Bay, ca. 35 m, Engel 21799 (F); ca. 4 km N

of Haast River, sea level, Engel 21769 (F). CANTER-

BURY PROV.: Arthur's Pass Natl. Park, Bealey Glacier

Track, 915 m, Fife 8987— c. S (F).

Subg. Glaucolepidozia Schust.

Lepidozia subg. Glaucolepidozia Schust., J. Hattori

Bot. Lab. 36: 386. 1973 (1972).

Lepidoziopsis Hodgs., Rec. Domin. Mus. 4: 105.

1962.

Plants procumbent to creeping, forming thin,

often extensive and pure, opaque, whitish patches,

usually in deeply shaded sites. Similar to subg.

Notholepidozia in the strongly incubously inserted

and oriented leaves, but the leaves are usually

only weakly convex and only slightly asymmet-

rically 4-lobed. Cells covered with an opaque

whitish waxy substance, the leaf cells obscured,

the cell walls difficult to resolve, even under the

microscope. Plants not rigid, the stem not strongly

corticated, usually regularly 1- or 1-2-pinnately

branched; lateral branches all terminal, Frullania

type.

TYPE — Lepidozia glaucophylla Steph.

The subg. Glaucolepidozia consists of four spe-

cies that collectively are very distinctive in aspect.

In the field, the plants look almost white and are

quite opaque, owing partly to the peculiar cutini-

zation, which is not seen in any other New Zea-

land hepatic. The strongly incubously inserted

leaves are almost horizontally oriented, quadrate-

oblong, and subsymmetrically 4-lobed, with eden-

tate margins; they are much larger than the remote

underleaves.

The subgenus is restricted to Australasia and

occurs from Stewart Island north to North Island,

at least to the Coromandel Peninsula. Plants are

often found in areas shaded by shrubs or dwarfed

trees, or in sheltered loci under ledges, at bases

of shaded cliffs, or in pockets at tree bases. They

are usually in diffuse but often extensive and very

thin patches.

Both ecology and reproductive biology of

Glaucolepidozia are distinctive. Three of the four

taxa are known only sterile or from plants of one

sex; only L. glaucescens is known with perianths.

No taxon is known with sporophytes. In view of

the rather wide distribution of several species and

their rather regular occurrence in extensive mats

in deeply shaded sites, their means of dissemina-

tion is of considerable interest. The species tend

to occur on humus-rich soil or decayed wood in

deeply shaded sites, such as hollows of decayed

stumps, or on the ground on the shaded side of

fallen and decayed trees. In at least L. digitata,

weak forms may regularly have fragmenting or

irregularly caducous leaf lobes (Fig. 38 : 1,3, ar-

rows), and even underleaf lobes may be caducous.

Presumably dissemination is by such leaf frag-

ments.

In Glaucolepidozia, leaf cells tend to show dis-

tinct middle lamellae (Fig. 38 : 10), and "Tangl's

canals" may be evident (Fig. 36 : 9). In all taxa

known from living plants, the chloroplasts are un-

usually dense and large (the whitish aspect is de-

rived at least in part from reflection from chlo-

roplast surfaces), and the oil-bodies are unusually

small and often obscure (Figs. 36 : 9, 10; 38 : 9),

always much smaller than the chloroplasts. They

may be, in part, vestigial (Fig. 38 : 10) and

formed of only 3-4 obscure segments. Stems, al-

ways creeping or procumbent, are relatively soft-

textured; both cortical and medullary cells are vir-

tually thin-walled (Figs. 36 : 1 1 ; 37 : 7).

Lepidozia glaucophylla (Hook. f. & Tayl.) G. L.

& N. Figure 35

Jungermannia glaucophylla Hook. f. & Tayl., London

J. Bot. 3: 580. 1844. Lepidozia glaucophylla

(Hook. f. & Tayl.) G. L. & N., Syn. Hep. 207.

1845. Mastigophora glaucophylla (Hook. f. &

Tayl.) Trev., Mem. 1st. Lomb. Sci. Lett. III. 4: 416.

1877. Lepidoziopsis glaucophylla (Hook. f. &

Tayl.) Hodgs., Rec. Domin. Mus. 4: 106. 1962.

Lectotype (nov.): Tasmania, 1840, Hooker (FH!).

Plants prostrate, in loosely interwoven pure

mats or as conspicuous, isolated strands growing

among other matted bryophytes, slender, with

spreading branches, glaucous, whitish to cera-

ceous, the older sectors subnitid and brownish (as

if scorched), the surface dull, waxy, and water re-

pellent, the shoots medium, to 1.5 cm wide, in-

cluding branches. Branching mostly of Frullania

type, ± distantly l(2)-pinnate, the branches grad-

ually tapering, occasionally whiplike, flagelliform,

and microphyllous, the nonmicrophyllous leaves

subsymmetrically 4-lobed, the dorsal margin cor-

date at the base, the primary branches rarely de-

veloping into new leading shoots; secondary

branches occasional in well-developed shoots;

branch half-leaf broadly ovate, covering branch

axil (as in Bazzania), slightly asymmetrical, cor-

date at base, 2-lobed to ca. 0.2-0.3, the margin

sinuate or bluntly toothed; first branch underleaf

large, mostly undivided, occasionally 2- or 3-

92

FIELDIANA: BOTANY

n m^P^

u

FIG. 35. Lepidozia glaucophylla (Hook. f. & Tayl.) G. L. & N. 1. Sector of main shoot, dorsal view (HL = half-

leaf). 2-4. Leaves (all at same scale). 5. Dorsalmost lobe (above) and dorsal pair of lobes, the cuticle shown for lobe

at right. 6. Median lobes of leaves (all at same scale). 7-9. Marginal cells of ampliate sector (dorsal, basal portion)

of leaf disc. 10. Median disc cells. 11. Portion of ventral margin of leaf disc. 12. Sector of main shoot with Frullania-

type branch, ventral view (note first branch underleaf, BUL). 13. Underleaves. 14. Underleaf, cellular detail. (Figs.

1, 2, 5, 8, 10, 12-14 from type; 3, 7 from Fife 6442, New Zealand, South Is., Paparoa Mts. Ml. Euclid; 4, 6, 9 from

Engel 23170, New Zealand, South Is., Southland Prov., Fiordland Natl. Park, S end of Lake Marion.)

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

93

lobed, inserted on ventral-lateral side of branch at

or near junction of branch and main shoot, typi-

cally obliquely to horizontally inserted and

aligned with underleaves of branch. Ventral-inter-

calary branches sporadically produced. Stems

flexuous. Stem epidermal cells in surface view ±

thick-walled, with a waxy coating like that of the

leaves, the stems 11-14 cells in diameter, the cor-

tex in 1 layer of thin to slightly thick-walled cells

± distinctly larger than the medullary cells (the

cortical cells well-defined in longisection, much

shorter than the medullary); medullary cells

slightly thickened. Leaves involute and subam-

plexicaul when dry, spreading and divergent when

moist, rather strongly concave, contiguous to im-

bricate, at times nearly or completely hiding stem

in dorsal view, uniform in size (not in sequential

sectors of varying size), 0.6-0.7 mm long and

wide, the insertion broad, moderately to strongly

incubous, recurved at dorsal end; leaves asym-

metrical, ± equally 4-lobed, the leaves divided to

ca. 0.4-0.5 (median sinus), the distance from dor-

sal sinus base to insertion not much greater than

that from ventral sinus to insertion, the sinuses

gradually becoming deeper ventrally. Lobes sub-

caudate, not noticeably in pairs, entire, abruptly

narrowing to a uniseriate row of 3-5(6) cells; cells

of uniseriate row and often the subtending lobe

cells elongated (to 2:1), thick-walled, the terminal

cell tapering to a point; dorsal lobe 5-7 cells wide

at base. Disc distinctly asymmetrical because of

ampliation of dorsal margin, the dorsal sector to

one-third the disc area, 14-18 cells high at dorsal

sinus, 10-14 cells high at ventral sinus, the dorsal

margin subentire to sinuate to distantly and irreg-

ularly armed with 1-2-celled teeth, the margin au-

riculate at the base, the ventral much shorter than

the dorsal, subcordate at the base, occasionally

with a small tooth or short spine. Cells of disc

and lobes rounded quadrate, evenly thick-walled,

trigones absent, the median disc cells 20-26 ym

X 22-30 /Jim; marginal cells of disc and lobes

typically with a thickened outer wall; cuticle a

dense and coarsely granular coating. Underleaves

inserted on 8-9 rows of stem cells, distinctly

spreading, ca. 1-1. 3 X stem width, symmetrically

4-fid to ca. 0.4 (median sinus), the lobes plane,

entire, narrowly attenuate and consisting of sev-

eral tiers of 2 laterally juxtaposed cells and then

rounded at the tip or, more commonly, terminating

in a uniseriate row of 2-4 cells; disc 9-12 cells

high at median sinus, the margins plane, occa-

sionally with a 1-2-celled tooth that terminates in

a slime papilla.

Androecia and gynoecia not seen.

DIFFERENTIATION — Lepidozia species with a

waxy, water-repellent "bloom" on the surface of

the stems and leaves have been understood as

constituting a single species, L. glaucophylla. Pri-

marily because of this striking feature, the species

has been treated as the type of a monotypic genus,

Lepidoziopsis Hodgs. Schuster (1973) listed a

number of additional distinctive features of this

taxon, which he treated as Lepidozia subg. Glau-

colepidozia Schust. In fact, in New Zealand, four

distinct elements are represented: L. glaucophylla

s. str., L. bisbifida Steph., L. digitata Herz., and

L. glaucescens, described here.

Examination of the Tasmanian type of Lepido-

zia glaucophylla (FH!) shows it to be distinct

from the familiar New Zealand plant that has long

been known by this name. True L. glaucophylla

appears to be the only species of this complex

present in Tasmania and Australia, but it is quite

rare in New Zealand; L. bisbifida Steph. (appar-

ently the oldest name for the New Zealand plant)

is "widely distributed in New Zealand, but no-

where common" (Allison & Child, 1975, p. 65)

and is probably endemic.

Lepidozia glaucophylla is similar to L. bisbifida

in the conspicuous water-repellent "bloom" on

the surface of the stem and leaves and the strongly

incubous, subsymmetrically lobed leaves (Fig.

35 : 1-3). It is immediately distinguishable, how-

ever, by the broadly ampliate and strongly auric-

ulate dorsal margin of the leaf (Fig. 35 : 1-4, 8);

the irregularly sinuate to toothed leaf margins

(Fig. 35 : 7-9); the evenly thick-walled disc cells

(Fig. 35 : 7, 9, 10); the subcaudate, abruptly ta-

pering leaf lobes, which terminate in a uniseriate

row of 3-5 elongated cells (Fig. 25 : 4-6); and

the oblique to longitudinal insertion of the first

branch underleaf at the base of the branch, rather

than transversely inserted on the ventral-lateral

face of the stem, as in L. bisbifida.

DISTRIBUTION-ECOLOGY — New Zealand, Tas-

mania, Australia (Victoria, N. S. W.). Apparently

very rare in New Zealand, and known only from

a few sites on South Island. The Engel plant oc-

curred at 695 m in a rather open forest dominated

by Nothofagus menziesii, with frequent large

boulders that are often largely bryophyte covered.

Plants of L. glaucophylla, together with hymen-

ophylls, formed a thick mass that was loosely at-

tached to a huge boulder; the boulder otherwise

was covered with bryophytes and hymenophylls.

Interesting ecological partitioning occurred here:

94

FIELDIANA: BOTANY

Lepidozia bisbifida occurred on soil of the forest

floor deep in a recess formed by a ledgelike over-

hang of the same boulder. The Fife plant occurred

on humus over rock in a north-facing, moist Noth-

ofagus menziesii forest with abundant Dracophyl-

lum traversii at 1000-1065 m.

SPECIMENS SEEN— NEW ZEALAND. SOUTH IS-

LAND. SOUTHLAND PROV.: Fiordland Natl. Park, S

end of Lake Marion, W of Hollyford River, 695 m, En-

gel 23170 (F). NELSON PROV: Paparoa Mts., N flank

of Mt. Euclid, ca. 0.5 km SE of Morgan Tarn, 1000-

1065 m, Fife 6442 (F).

Lepidozia bisbifida Steph. Figure 36

Lepidozia bisbifida Steph.. Spec. Hep. 3: 593. 1909.

Original material: New Zealand, without specific

locality. Kirk s.n. (c!).

Lepidozia subquadrata Steph., Spec. Hep. 6: 341.

1922. Original material: New Zealand, "alpine in-

terior." without specific locality, Colenso "inter

2064" (c!).

Lepidozia brevipinna Pears., Univ. Calif. Publ. Bot.

10: 321. pi 94. 1923. Original material: New Zea-

land, North Is., Wairakei, Waiora Valley, 4 May

1904, Setchell 9 (uc!).

Plants prostrate to loosely procumbent, the

stems loosely interwoven, flexuous, with spread-

ing branches, glaucous and ash-gray to ivory, the

older shoot sectors burnt orange or lightly tinged

with brown as if scorched, the surface dull and

water repellent; shoots medium-sized, to 1.5 cm

wide, including branches (sporadically large and

to 2 cm wide [including branches] X 7.5 cm

long). Branching nearly exclusively of Frullania

type, rather short, closely and regularly pinnate to

locally bipinnate, the branches abruptly tapering,

occasionally becoming ± whiplike, flagelliform,

and microphyllous, the nonmicrophyllous leaves

shallowly and ± symmetrically 4(5)-lobed, the

primary branches sometimes developing into new

leading shoots; secondary branches occasional, 1-

3 per primary branch; branch half-leaf broadly

ovate, covering branch axil (as in Bazzania), sub-

symmetrical, subcordate at the base, 2-lobed to ca.

0.2-0.3, the dorsal lobe often smaller; first branch

underleaf large, often squarrose reflexed, 2-4-

lobed (rarely undivided), usually transversely in-

serted on lateral or ventral-lateral side of stem,

often somewhat below the branch, aligned with

leaves of main shoot or underleaves of branch.

Ventral-intercalary branching occasional, leafy.

Stems flexuous. Stem epidermal cells thin-walled,

with a waxy coating like the cells of the leaves.

Leaves involute-triangular and distinctly amplex-

icaul when dry, when moist rigid, distinctly con-

cave to nearly cuplike, approximate to contiguous,

intermittently large and small along length of

stem, those in well-developed sectors 0.8-1.3 mm

long and wide, the leaves of smaller sectors as

small as 0.5 mm long and wide, the leaves spread-

ing, nearly horizontal, the insertion broad, strong-

ly incubous, recurved at dorsal end; leaves sub-

symmetrical to moderately asymmetrical, typical-

ly ± equally 4-lobed (rarely bisbifid), the lobes

sometimes in turn divided, the leaves then ap-

pearing 5-8-lobed, the leaves divided to ca. 0.2-

0.5 (median sinus), the distance from dorsal sinus

base to insertion subequal to or not much greater

than that from ventral sinus to insertion, the si-

nuses of subequal depth or gradually becoming

deeper ventrally. Lobes acute to apiculate, entire,

terminating in a single cell or more commonly a

uniseriate row of 2-3 cells; cells of uniseriate row

± isodiametric or slightly longer than wide, thin-

walled, the terminal cell strongly tapering; median

pair of lobes somewhat larger, 6-8(1 1) cells wide

at base, the dorsal lobes (2)3-5 cells wide at base.

Disc subsymmetrical, (13)15-22(28) cells high at

dorsal sinus, 9-14 cells high at ventral sinus, the

margins usually entire, the dorsal margin moder-

ately ampliate, sporadically with 1 or more blunt

teeth or sinuate, cordate at the base, the ventral

margin not much shorter than the dorsal, occa-

sionally with a slender lobuliform process. Cells

of disc and lobes uniformly thin-walled, with con-

spicuous intercellular pits, trigones minute to me-

dium, the median disc cells (18)24-40 X 28-40

yum; median basal cells not differentiated; margin-

al cells of disc smaller, those of disc and lobes

sporadically with thickened outer walls, but rarely

consistently so; cuticle a dense granular coating

that ultimately develops cracks or fine fissures (es-

pecially over the vertical cell walls), the cell out-

lines obscured by the scurfy, water-repellent coat-

ing. Oil-bodies small and inconspicuous, 2-5 to

(0)4-6(9) per cell, few (often only 3-6[9]) seg-

mented, coarsely botryoidal, much smaller than

chloroplasts, often 6 //.m long X 3 /im wide and

consisting of a large spherule at each end and 4-

6 smaller ones centrally. Underleaves inserted on

9-10 rows of stem cells, widely spreading, ca.

0.7- IX stem width, asymmetrically or symmet-

rically 4-fid to ca. 0.3-0.45 (median sinus), the

underleaves often with 1, 2, or all 3 sinuses nar-

row, slitlike and the lobes adnate by the surface

covering, the underleaves then in situ appearing 2

or 3 lobed, the lobes plane, entire, attenuate to

acuminate, consisting of serveral tiers of 2 later-

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

95

FIG. 36. Lepidozia bisbifida Steph. 1. Part of plant, dorsal aspect (X 10.2; 2.5 mm scale). 2. Sector of main stem,

dorsal view; half-leaf (HL) and first branch underleaf (UL1) labeled (X21; 1.0 mm scale). 3. Same, ventral aspect;

lateral leaves in part 5-lobed (L5), in part 6-lobed (L6); Frullania-lype branches (FB) with basal branch underleaf at

right (UL1) inserted below seeming branch base; at left, stem underleaf (SUL) and branch underleaf (BUL1) faintly

connate at juxtaposed bases (X27; lower 1.0 mm scale). 4, 5. Stem leaf and underleaf (X32; 700 /xm scale). 6. Leaf

lobe cells (X160; 150 ^m scale). 7. Cortical cells (X160; 150 ^m scale). 8. Raphides (X650; 20 pm scale). 9. Cells

with oil-bodies; right, chloroplasts (X190). 10. Oil-bodies (X190). 11. Stem cross section (X190).

ally juxtaposed cells and then rounded at the tip

or, more commonly, terminating in a single cell

or a uniseriate row of 2-4 cells that often termi-

nate in a slime papillae; disc 5-9 cells high at

median sinus, the margins plane, entire.

Plants dioecious. Androecia on inconspicuous,

short, determinate, tightly spicate, often cernuous

ventral-intercalary branches from main shoot;

bracts ventricose-cucullate, 2-lobed to ca. 0.3, the

lobes acute to short acuminate; dorsal margin of

disc dilated in basal sector to form a feebly to

moderately rounded projection, the dilated portion

96

FIELDIANA: BOTANY

crenulate and with several slime papillae, the disc

otherwise entire; antheridia 1 per bract, the stalk

biseriate. Gynoecia not seen.

DIFFERENTIATION — This is the species long

known in the New Zealand literature as L glau-

cophylla; for the correct application of this name,

see the discussion under that species. Lepidozia

bisbifida is readily distinguished from L. glauco-

phylla by the broadly inserted, deeply concave, in-

volute leaves (Fig. 36 : 1 ); by the leptodermous

leaf cells with small to minute trigones (Fig. 36 :

9); and by the ± transverse insertion of the first

branch underleaf on the stem, often some distance

below the associated branch (Fig. 36 : 3). The thin

walls of the leaf cells are traversed by delicate in-

tercellular connections (plasmodesmata, Fig. 36 :

9). Another feature of this species is the variability

in leaf size on a single shoot and the characteristic

appearance of the branches (Fig. 36 : 1 ), which are

abruptly tapering and have much smaller leaves

than those of the main shoot. The first branch un-

derleaves in this species are inserted well below

the apparent branch base (Fig. 36 : 2, 3).

The marginal cells of disc and lobes are typi-

cally thin-walled like those of the leaf interior.

Marginal cells, however, sporadically have thick-

ened outer walls and at times these may be locally

feebly crescentic. In Engel 22930 the marginal

cells of both disc and lobes have strongly thick-

ened, concave outer walls, with the wall thick-

ening crescentic and bulging into the cell lumen.

In water mounts, the waxy, granular coating on

the leaf surface renders the leaves almost opaque

and obscures the outlines of the cells. An inter-

esting feature of the species is the fine network of

hairline cracks in the cuticle, which is visible over

the cell walls of dried specimens. When the leaves

are mounted in Hoyer's solution, the waxy coating

is apparently dissolved and forms delicate, needle-

like crystals (Fig. 36 : 8).

DISTRIBUTION-ECOLOGY — Confined to New

Zealand (South Is., North Is., Antipodes Is.).

A species occurring in forests ranging from as

low as 450 m, but more often from over 750 m

to the upper reaches of the forest. It is found on

rotted wood (particularly where there is cover of

other bryophytes, along with, at times, hymeno-

phylls) or deep in shaded, moist, sheltered pockets

of tree bases, boulders (see also note under L.

glaucophylla), or rotted logs. Also on shaded cliff

bases, particularly where soil has accumulated,

and plants are notable as scattered glaucous patch-

es among other bryophytes. The species at times

covers large areas but typically does not become

matted or pure. However, pure mats are at times

formed deep in sheltered recesses. In the subal-

pine zone over soil deep in protected recesses be-

tween boulders or under boulder overhangs, as

well as on the sides of rills. In the alpine zone on

shaded, damp slopes under cover of snow tussock.

This is the only species of subg. Glaucolepi-

dozia known from the geothermal areas of North

Island, New Zealand. The presence of L. bisbifida

at such sites is somewhat of an anomaly, because

higher plants from geothermal areas are for the

most part cosmopolitan plants of the tropics;

moreover, few or no higher plants are endemic to

geothermal areas. Rather, they are weedy in na-

ture, high light loving, and, if they occur else-

where in New Zealand, they typically are present

only in warm sites, such as in the vicinity of

Auckland or Northland (J. Braggins, pers. com.).

All of the specimens of L. bisbifida cited here

from geothermal areas are from, at least to some

degree, shaded sites; the Braggins plant, for ex-

ample, occurred in light shade under Kunzea.

The species is quite distinct in the field due to

the large plant size coupled with the glaucous

condition, with older portions of the plants often

burnt orange or lightly tinged with brown and

then appear as if scorched. It often grows loosely

attached, and may be very common locally, such

as at the Moraine Creek Track site.

SELECTED SPECIMENS SEEN— NEW ZEALAND.

SOUTH ISLAND. SOUTHLAND PROV: Fiordland

Natl. Park, off track to Island Lake, just W of Borland

Saddle, S of South Branch of Borland Burn, W of Mon-

owai, 800 m, Engel 18621 (F); ibid.. Central Earl Mts.,

Mistake Creek, between Triangle Peak and Melita Peak,

NE of northern end of Lake Te Anau, 740-800 m, Engel

18784 (F); ibid., Gertrude Valley, near track entrance to

Gertrude Saddle, E of Homer Tunnel, 1740 m, Engel

21947 (F); ibid., S end of Lake Marion, W of Hollyford

River, 695 m, Engel 23168 (F); Lake Marian Valley, ca.

305 m. Child H2950 (F); Fiordland Natl. Park. Moraine

Creek Track, area N of Moraine Creek, W of Hollyford

River, 610m, Engel 23227 (F). SOUTHLAND/OTAGO

PROV. BOUNDARY: Lake Howden. ca. 760 m, Child

2557 (F). OTAGO PROV.: Ajax Swamp, ca. 1 km N of

Ajax Hill, Catlins River area, ca. 610 m. Child H5494

(F); ML Cargill, N of Dunedin, ca. 455 m. Child H1706A

(F); Rees Valley, Invincible Creek, ca. 455 m. Child

H375 (F); tributary of Siberia Stream, opposite and

downstream from Siberia Hut, WNW of Makarora, ca.

760 m. Child H2873 (F); Fiordland Natl. Park, near

McKerrow Hut, head of Lake McKerrow. Hatcher 1446

(F); Ml. Aspiring Natl. Park, below and W of ML

Shrimpton, 1370-1470 m, Engel 17880 (F); ibid.. Blue

Valley Track, above Blue River just N of confluence

with Makaroa River, 430-480 m, Engel 21908 (F).

OTAGO/WESTLAND PROV. BOUNDARY: Mt. As-

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

97

piring Natl. Park, Cross Creek, 1 km N of Haast Pass,

510 m, Engel 23105 (F); ibid., summit area of Haast

Pass, 570 m, Engel 17986 (F); Mt. Brewster, ca. 1370

m, Child 4188 (F). WESTLAND PROV.: 2 km N of

White Horse Creek, ca. 305 m, Child H5425 (F). WEST-

LAND/CANTERBURY PROV. BOUNDARY: Arthur's

Pass Natl. Park, Arthur's Pass, near Temple Basin Ski

area, Engel 6499A (F); ibid., Bridal Veil Track, E side

of Bealey River and just N of town of Arthur's Pass,

760-825 m, Engel 22930 (F). CANTERBURY PROV:

Cass, Woolshed Hill, Visch s. n. (F); Arthur's Pass Natl.

Park. Bealey River, off Bealey Valley Track, 830-850

m, Engel 18516— c. <S (F). NELSON PROV: Paparoa

Mts., N flank of Mt. Euclid, ca. 0.5 km SE of Morgan

Tarn, Fife 6412, 6442 (F). NORTH ISLAND: WEL-

LINGTON PROV: Tongariro Natl. Park, Blyth Track,

along small stream ca. 0.5 km from Ohakune Mt. Road,

ca. 1230 m, Engel 21310 (F); Tokaanu Hot Springs, S

end of Lake Taupo, Braggins—c. 6 (F). SOUTH AUCK-

LAND PROV: Rotorua thermal area, Hatcher 36 (F);

ibid., Barnard NZ-24 (F). ANTIPODES ISLAND: ca.

150 m, Godley (F).

Lepidozia digitata Herz. Figures 37 and 38

Lepidozia digitata Herz., Trans. & Proc. Roy. Soc.

New Zealand 68: 45. pi. 5, m-n. 1938. Holotype:

New Zealand, near Atiamuri, Allison 72 (JE!).

Plants with the superficial facies of Telaranea

centipes, prostrate, the stems loosely interwoven,

flexuous, with widely spreading branches, glau-

cous to ivory, the older shoot sectors tinged with

brown, the surface dull and water repellent; shoots

small, to 0.6 cm wide, including branches.

Branching exclusively of Frullania type, the

branches rather short, rather distant and irregular-

ly pinnate, the branches short and not tapering or

only sporadically becoming flagelliform and mi-

crophyllous, the nonmicrophyllous leaves ± sym-

metrically (3)4-lobed, to 3-lobed in distal sectors;

branch half-leaf narrowly ovate, subsymmetrical,

subcordate at the base, 2-lobed to ca. 0.2-0.3; first

branch underleaf 2-lobed (occasionally undivid-

ed), inserted on ventral-lateral side of main shoot

at branch base and obliquely to subvertically in-

serted. Ventral-intercalary branching not seen.

Stems flexuous. Stem epidermal cells thin-walled,

with a waxy coating like the cells of the leaves,

the cortex in a single row of cells that are slightly

smaller to or at most feebly larger than those of

the medulla. Leaves with margins somewhat in-

volute when dry, plane to slightly concave when

moist, distant to contiguous, ± uniform in size,

0.35-0.55 mm wide X 0.4-0.55 mm long, spread-

ing, nearly horizontal, the insertion broad, strong-

ly incubous, at most feebly recurved at dorsal end;

leaves at most moderately asymmetrical, (3)4-

lobed to ca. 0.35 (median sinus), the distance

from dorsal sinus base to insertion not much

greater than that from ventral sinus to insertion,

the sinuses gradually becoming deeper ventrally.

Lobes subacuminate, entire, terminating in a un-

iseriate row of 2-3(4-5) cells; cells of uniseriate

row ± isodiametric or somewhat longer than

wide, thin-walled, the terminal cell tapering to a

rounded apex; median pair of lobes somewhat

larger, 3-5(6) cells wide at base, the dorsal lobe

2-3(4) cells wide at base. Disc subsymmetrical,

(6)7-13(16) cells high at dorsal sinus, 6-13 cells

high at ventral sinus; dorsal margin scarcely am-

pliate, entire or somewhat sinuate, sporadically

with a 1 -celled tooth, feebly cordate at the base;

ventral margin somewhat shorter than the dorsal,

entire. Cells of disc and lobes uniformly thin-

walled, trigones none or minute, the median disc

cells 24-35 /urn wide X 29-42 /i,m long; median

basal cells somewhat elongated but otherwise not

differentiated; marginal cells of disc (especially

the dorsal) often with a somewhat thickened, con-

cave outer wall (the wall thickening somewhat

crescentic and bulging into the cell lumen); cuticle

finely granular, the cell outlines obscured by the

scurfy, water-repellent coating. Oil-bodies small

or tiny, in some cells rudimentary (and obscurely

few-septate), much smaller than chloroplasts, very

variably developed, ranging from tiny and few-

septate to ellipsoidal or fusiform and clearly bot-

ryoidal. Underleaves inserted on 3-4 rows of

stem cells, small, slightly spreading, ca. 1-1. 3X

stem width, ± symmetrically 3-4-fid to ca. 0.5-

65 (median sinus), the sinuses broad to narrow,

the lobes entire, slender, 2-3(4) cells wide at the

base and often uniseriate for most of their length,

the uniseriate row of 2-3 cells and often termi-

nating in a slime papilla; disc (1.5)2-4 cells high

at median sinus, the margins plane, entire. Asex-

ual reproduction sporadically present, via frag-

menting or caducous leaf lobes, in which bases

persist as truncate stubs.

Androecia and gynoecia not seen.

DIFFERENTIATION — Lepidozia digitata is the

smallest of the four New Zealand species of subg.

Glaucolepidozia, with leaves at most 0.5 mm in

longest dimension. Hodgson (1956) treated L. dig-

itata as a small-leaved expression of "L. glauco-

phylla" (i.e., L. bisbifida), but it differs in being

only one-third the size. The leaves are uniformly

small, not in alternate sectors of large and small

leaves along the length of the stem as in L. bis-

bifida. Curiously, Herzog (1938), who described

98

FIELDIANA: BOTANY

FIG. 37. Lepidozia digitata Herz. 1. Sector of main shoot, dorsal view. 2. Leaves and, above, underleaves. 3.

Leaf, cellular detail. 4. Leaf lobes, cuticle indicated in part. 5. Median disc cells, cuticular detail indicated in part. 6.

Underleaf, cellular detail. 7. Stem, cross-section. 8. Sector of main shoot with 2 Frullania-type branches, ventral

view (note undivided and, lower, bilobed first branch underleaves; the latter is the typical condition). (All from type.)

the leaf cells as "subopaco-brunneolae" and the

cuticle as "minutissime et densissime punctula-

ta," did not associate this species with L. glau-

cophylla; the type is distinctly glaucous and water

repellent.

Lepidozia digitata is not only the smallest spe-

cies of Glaucolepidozia, it has the most remote

leaves (Figs. 37 : 1, 8; 38 : 3). It apparently repro-

duces only via fragmenting leaf lobes, in which

bases persist as truncated stubs. Such asexual re-

production may be widespread (Fig. 38 : 1, 3) or

sporadically evident (Fig. 37 : 8). The leaf lobes

mostly end in 3 superposed cells (Fig. 37 : 3, 4)

but on weak phases sometimes in 4-5 cells (Fig.

38 : 1, 4, 6). Underleaves on branches or (weak

phases) even on main stems may be bifid or trifid

in part, or predominantly so (Figs. 37 : 8; 38 : 3).

Although the disc may be 3-4(5) cells high on

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

99

FIG. 38. Lepidozia digitata Herz. 1. Shoot sector, ventral aspect; at right, a Frullania-lype branch, with bifid first

branch underleaf (BUL1) (X93; 400 ,u,m scale). 2. Underleaf from stem (X 170; 150 /im scale). 3. Part of plant, ventral

aspect; of the four Frullania-type branches, the upper three bear bifid first branch underleaves, the lower (arrow) a

monocrural one; at arrows, leaf lobes that have fragmented (X50; 600 yum scale). 4. Median lobe of leaf (X 170; 150

/u,m scale). 5. Underleaf (X170; 150 yum scale). 6. Ventral lobe of leaf (X170; 150 /am scale). 7. Stem cross section

(X300; 80 /tm scale). 8. Individual oil-bodies (X1770). 9. Two cells with oil-bodies (X925). 10. Three laminar cells

with oil-bodies; at, upper left, chloroplasts (X925). (All from Schuster 95-314, New Zealand, North Is., Urewera

Natl. Park; FAA preserved plants.)

vigorous phases, weak phases may have it only

2-3 cells high (Fig. 38 : 2). Stems are weaker in

L. digitata (compare Figs. 37 : 7 and 38 : 7 with

Fig. 36 : 1 1, L. bisbifida) than in other Glaucolep-

dozia species; on weak phases they may be only

6 cells high, ranging to 8-9 cells high in vigorous

phases. Oil-bodies in this species are often vesti-

gial (stippled ones in Fig. 38 : 8, 10 are almost

100

FIELDIANA: BOTANY

unresolvable, even under oil immersion) and typ-

ically occur only 2-4 per cell; they are often ob-

scured by the large, often dense chloroplasts.

DISTRIBUTION-ECOLOGY — New Zealand (South

and North Is.).

The few, scattered stations for the species are

for the most part within Nothofagus solandri van

cliffortioides and Nothofagus menziesii forests

ranging from 630 to 1390 m. It occurs in shaded,

protected, often deep pockets (such as the hol-

lowed recesses of tree bases); in crevices and re-

cesses of outcrops; or over soil of steep, mossy

banks, where it may occur, for example, in a pro-

tected niche under the lip of the forest edge at the

upper extremity of a stream bank (Engel 22637).

Also known from 455 m in Otago Prov. (Invin-

cible Creek, leg. Child); the vegetation type is not

provided.

SPECIMENS SEEN— NEW ZEALAND. SOUTH IS-

LAND. OTAGO PROV: Rees Valley, Invincible Creek,

ca. 455 m. Child H375 (F). CANTERBURY PROV: Ar-

thur's Pass Natl. Park, Scotts Track to Avalanche Peak,

W of town of Arthur's Pass. 950 m, Engel 22081 (F).

NELSON PROV: Nelson Lakes Natl. Park, Pinchgut

Track, W of southern sector of Lake Rotoiti, SSW of St.

Arnaud, ca. 1280-1390 m, Engel 21424 (F); ibid., off

Lakehead Track, near juncture with southern end of

Loop Track, NE end of Lake Rotoiti, 630 m, Engel

22637 (F). NORTH ISLAND, WELLINGTON PROV:

Tongariro Natl. Park, west-facing steep bank of Man-

gawhero River, off Ohakune Mt. Road, ca. 1200 m, En-

gel 21287 (F). GISBORNE PROV: Urewera Natl. Park,

Huiarau Range, summit area of Te Rangaakapua, 1230-

1320 m, Engel 23470 (F).

Lepidozia glaucescens Engel, sp. nov. Figure 39

Folia infirme vel distincte incuba insertione angusta

exorientia, distincte asymmetrica, margine dorsali sub-

recta et disco piano et lobis patentibus praedita; cellulae

et disci et loborum foliarium parietibus distincte crassis,

papillis striatis, velamineque continue ceraceo granuloso

instructae.

HOLOTYPE — New Zealand, South Island, Nel-

son/Westland Prov. boundary, Porarari River

Track, 5 km from road, ca. 500 ft., Child H4966

(F); isotype: (CHR).

Plants prostrate, in loosely interwoven mats,

the shoots slender, fragile, with widely spreading

branches, fragile, brownish and subnitid to glau-

cous and ivory-white (at least on new growth),

the surface then dull and water repellent, the

shoots medium, to 2.5 cm wide, including branch-

es. Branching nearly exclusively of Frullania

type, ± regularly 1 -pinnate varying to remotely

2-pinnate, the branches gradually tapering, be-

coming whiplike, flagelliform and microphyllous,

the nonmicrophyllous branch leaves ± symmet-

rically 4-lobed, the 2 median lobes somewhat

larger, the primary branches occasionally devel-

oping into new leading shoots; secondary branch-

es sporadic, 1-2 per primary branch; branch half-

leaf narrow, linear, subsymmetrical, subcordate at

the base, 2-lobed to ca. 0.4; first branch underleaf

mostly undivided, less commonly 2(4)-lobed, in-

serted on ventral-lateral side of branch base and

aligned with underleaves of branch. Ventral-inter-

calary branching rare, leafy. Stems flexuous. Stem

epidermal cells in surface view rather thick-

walled, sometimes with a waxy coating like that

of the leaves. Leaves fragile, when dry moderate-

ly concave, with the lobes curved ventrally;

leaves when moist explanate, with spreading

lobes, ± distant to contiguous, with much of stem

visible in dorsal view, uniform in size (not in se-

quential sectors of varying size), narrowly insert-

ed and somewhat longer than wide, 0.3-0.5(0.6)

mm long and wide, spreading, the insertion vary-

ing from weakly to distinctly incubous; leaves

moderately to distinctly asymmetric, unequally 4-

lobed, the leaves divided to ca. 0.4-0.55 (median

sinus). Lobes narrowly attenuate to acuminate, the

dorsal lobes ± paired, the ventral shorter than the

dorsal lobes and somewhat divergent, the lobes

entire, terminating in a uniseriate row of 3-4(5)

cells; cells of uniseriate row isodiametric to ±

elongated (to 2:1), thick- walled and often with

somewhat swollen septa, the terminal cell often

moderately to distinctly elongated, tapering to a

point; the dorsal lobe 3-4(5) cells wide at base;

cuticle of lobes striate papillose, the papillae

mostly obscured by the thin and continuous gran-

ular coating. Disc moderately to distinctly asym-

metrical, obliquely truncate and deltoid, 8-12

cells high at dorsal sinus, 5-9 cells high at ventral

sinus; dorsal margin ± straight, abruptly cordate

at the base, entire; ventral margin shorter than the

dorsal, at times distinctly so, entire. Cells of disc

middle thick-walled, with trigones small, the cells

somewhat longitudinally elongated, 18-24(28)

H.m wide X 21-30 /im long, the cells of the nar-

row dorsal sector smaller and quadrate; median

basal cells in 1 (locally 2) rows of enlarged cells;

marginal cells of disc and lobes typically with a

thickened outer wall (the wall thickening often

crescentic and bulging into the cell lumen); cuticle

of disc as in lobes. Underleaves inserted on 4-6

rows of stem cells, spreading, small, ca. 0.9-1 X

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

101

FIG. 39. Lepidozia glaucescens Engel. 1. Sector of main shoot, dorsal view. 2. Sector of main shoot, ventral view.

3, 4. Leaves. 5. Underleaves. 6. Half-leaves. 7. Dorsal half of leaf (ms = median sinus). 8. Outer (dorsalmost) lobe

of dorsal half of leaf (cuticular detail not shown). 9. Median lobe of dorsal half of leaf (cuticular detail not shown).

10. Portion of dorsal lobe showing cuticular papillae, granular coating, and lamellated cell walls (cross-hatched

shading). 11. Median cells of dorsal half of disc showing striate cuticle. (Figs. 1-3, 5-7, 10 from type; 4, 8, 9, 11

from Engel 21087, New Zealand, North Is., North Auckland Prov., SE corner of Waipoua Forest.)

stem width, symmetrically 4-fid to 0.5 (median

sinus), the lobes plane, slenderly acuminate, en-

tire, terminating in a uniseriate row of 2-3(5)

cells; disc 3-6 cells high at median sinus, the mar-

gins plane, entire.

Plants dioecious. Androecia not seen. Gynoecia

on abbreviated ventral-intercalary branches issu-

ing from main stem, the gynoecium base bulbous,

fleshy, polystratose, and rhizoidous; bracts of in-

nermost series much larger than leaves, erect and

sheathing the perianth, the bracts deeply concave,

broadly ovate to ± orbicular; apices with 4 short,

lobes composed of ± regularly rectangular and

(especially at the lobe extremities) irregularly sin-

102

FIELDIANA: BOTANY

uate-rhomboidal cells, the apical end of the mar-

ginal cells often diverging and forming a slight

projection, the lobes thus crenulate; lamina com-

posed of ± regularly subrectangular cells, those

in distal sector rather thick-walled, those of re-

mainder of bract (particularly the cells in a large

median-basal field) thin-walled, the margin bor-

dered by cells of variable shape and orientation,

some hardly longer than wide, others ± sinuate-

rhomboidal, the apical or free end of marginal

cells variously divergent and forming a short pro-

jection or a tooth, the margin irregularly crenate-

denticulate to the base, the teeth at most of 3 su-

perposed cells; bracteole similar in size and form.

Perianth long and prominent, slenderly cylindri-

cal-fusiform, slightly curved, terete below, ob-

scurely trigonous above, distinctly and deeply 3-

plicate toward mouth, the perianth gradually nar-

rowing toward the strongly contracted, shallowly

3-lobed mouth, the lobes composed of rather

thick-walled, irregular, long, narrow, at times ±

sinuate, papillose cells that at the apical end are

laterally free for varying lengths, the lobes cre-

nate-denticulate; perianth cells subrectangular be-

low mouth, and here the cuticle is striate; perianth

2-stratose near base.

Sporophyte not seen.

DIFFERENTIATION — Lepidozia glaucescens dif-

fers from the other glaucous-leaved species of the

genus by having distinctly asymmetrical leaves,

with a ± straight dorsal margin (Fig. 39 : 1, 3, 4,

7). The leaf cells and the cortical cells of the stem

(in surface view) are distinctly thick-walled (Fig.

39 : 11), rather than leptodermous as in L. bisbi-

fida and L. digitata, and the wall thickenings have

a distinctive lamellated (layered) appearance (Fig.

39 : 10). The first branch underleaf is typically un-

divided in both L. glaucescens and L. glauco-

phylla. The leaf insertion is narrow and varies

from weakly to distinctly incubous, often on the

same shoot.

The leaves of L. glaucescens are remarkable

because of the unique combination of cells with

discrete elliptical papillae and a ± uniform waxy,

granular coating (Fig. 39 : 10). The papillae are

hemispherical to short on the lobes (Fig. 39 : 10)

but distinctly elongate in the median and basal

sectors of the disc (Fig. 39 : 11). The waxy coat-

ing is poorly developed in many collections, how-

ever, and is often evident only on the new growth

at the tips of the shoots. Consequently, Lepidozia

glaucescens might be mistaken for L. novae-ze-

landiae, which also has a striate-papillose cuticle,

similar leaf shape, and lobes terminating in a dif-

ferentiated uniseriate row of elongated, thick-

walled cells. Under the compound microscope,

however, the leaf surface of L. glaucescens has a

semiopaque, granular appearance, even if the

waxy coating is not apparent under the dissecting

microscope.

Plants with an apparent smooth cuticle but pos-

sessing ± flat discs with spreading lobes, ±

straight dorsal margin of the leaf, and thick-

walled lobe and disc cells, should be treated with

great care. Some populations of the species have

cuticlar papillae suboptimally developed; prefer-

ably, several leaves from various areas of several

shoots should be examined (including leaves from

near the shoot apex). Verification under oil im-

mersion can be helpful at times.

DISTRIBUTION-ECOLOGY: Known only from

Chatham Island and a few stations on South and

North Islands, New Zealand.

Apparently a lower-elevation species; the South

Island stations are all from between 120 and 300

m, while those from the North Island are from ca.

340-540 m. The type is from a damp forest, par-

tially under log cover; the White Horse Creek

specimens include a site beside a tarn in a pakihi

area under manuka (Leptospermum scoparium;

Child H5436) as well as boggy areas. In the

southern portion of Westland (Monkey Puzzle

Gorge), the species occurred in a very protected

niche: over soil deep in a pocket under a mass of

fallen trees in a forest dominated by Nothofagus

menziesii and Dacrydium. Known from North Is-

land only in the Waipoua Reserve area; the Engel

collection from just north of Tutamoe is from a

wet forest dominated by Weinmannia silvicola,

occurring on the top of a bryophyte-covered log,

while that from Te Matua Ngahere is from an

Agathis forest with Dacrydium and other podo-

carps, with a strong understory of Gahnia xantho-

carpa, where the species occurred in a shaded,

protected niche on the ventral-lateral side of a log.

The Schuster specimen is from a wet kauri forest.

The Chatham Island plant is from damp rocks and

banks in a dense broadleaf forest.

SELECTED SPECIMENS SEEN:— NEW ZEALAND.

SOUTH ISLAND. WESTLAND PROV.: Monkey Puz-

zle Gorge, Cascade Road, Martyr River near Martyr

Saddle, 120 m, Engel 23042 (F); 1 km N of White Horse

Creek, ca. 150 m, Child H5416 as L. glaucophylla (F);

2 km N of White Horse Creek, ca. 150-300 m, Child

H5425, H5436, both as L. glaucophylla (F). NORTH IS-

LAND. NORTH AUCKLAND PROV.: NE Waitakere

Ranges, Spraggs Bush, NE end of Turanga Rd. ca. 340

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

103

m, Braggins 99/298 (AKU); SE corner of Waipoua Forest,

just N of Tutamoe, 540 m, Engel 21087 (F); same loc.,

Braggins 98/318 (AKU, f); Waipoua Forest, track to Te

Matua Ngahere, ca. 340 m, Engel 22572 (F); ibid., Big

Tree Kauri area, near n-s road, Schuster 67-3798 (F).

CHATHAM IS.: Hattwoods Hole, Tuku a tamatea River,

Given (F).

Acknowledgments

Field work by the senior author was supported

by the National Science Foundation (grant Nos.

BMS76-03616 and DEB-8 109680) and National

Geographic Society (grant Nos. 5375-94 and

5795-96). We jointly thank the institutions and

foundations involved in supporting this research.

The senior author also extends thanks to Zorica

Dabich and Marlene Werner for preparing some

of the habitus figures, and to Gary Merrill for ad-

vice and assistance. Figures 1-5, 7, 13-18, 20-

21, 23-24, 27, 29-30, 32, 34-35, 37, and 39 were

prepared by the senior author; the remainder were

prepared by R. M. Schuster. We thank Patricia

Eckel for preparing the Latin diagnoses, Olga M.

Schuster for typing and revising portions of the

manuscript, and Elizabeth S. Engel for data pro-

cessing.

The junior author (in 1961-1962) conducted

field work while he was Fulbright Exchange Pro-

fessor at the University of Otago and (in 1 967) as

a recipient of a second Guggenheim Fellowship;

field work (and subsequent research) in 1983-

1984 was supported by a National Science Foun-

dation grant (No. DEB-8 1035 16).

We wish to thank the following individuals and

institutions for the loan of specimens to the Field

Museum: Mr. S. Blackmore and Dr. L. T Ellis

(BM), Dr. John Braggins, Matt von Konrat and Ka-

ren Sheath (AKU), Dr. P. J. Brownsey and Ms.

Fiona D. H. Pitt (WELT), Dr. Barbara Ertter (uc),

Dr. Patricia Geissler (G), Dr. F. K. Meyer (JE), and

Dr. D. H. Pfister (FH).

Literature Cited

ALLISON, K. W., AND J. CHILD. 1975. The Liverworts of

New Zealand, pp. 1-300. Univ. of Otago Press, Dun-

edin.

BOESEN, D. F. 1982. The taxonomy of Drucella Hodgs.

and Drucelleae, trib. nov. and their position within

Lepidozioideae Limpr. Lindbergia, 8: 77-88, figs.

1-8.

DEGENKOLBE, W. 1938 [1937]. Brutorgane bei beblatter-

ten Lebermoosen. Ann. Bryol., 10: 43-96, f. 1-112.

DUMORTIER, B. C. 1831. Sylloge Jungermannidearum

Europae Indigenarum, earum genera et species syste-

matice complectens, pp. 1-100, pis. 1-2. Tournay.

DUMORTIER, B. C. 1835. Recueil d'observations sur les

Jungermanniacees. Fasc. I, pp. 3-27. Tournay.

ENGEL, J. J., AND G. MERRILL. 1996a [1995]. Austral

Hepaticae 23. New taxa and new combinations in Te-

laranea Spruce ex Schiffn. (Lepidoziaceae). Phytolo-

gia, 79: 250-253.

. 1996b. Studies of New Zealand Hepaticae. 14-

19. Kurzia and Lepicolea. J. Hattori Bot. Lab., 80:

217-231, f. 1-3.

ENGEL, J. J., AND R. M. SCHUSTER. 1988. Studies of New

Zealand Hepaticae. 1-6. Brittonia, 40: 200-207, figs.

1-18.

FRYE, T. C., AND L. CLARK. 1937-1947. Hepaticae of

North America. Univ. Wash. Publ. Biol., 6: 1-1018.

FULFORD, M. 1966. Manual of the leafy Hepaticae of

Latin America. Part II. Mem. New York Bot. Card.,

11: 173-276, pis. 38-65.

FULFORD, M., AND J. TAYLOR. 1959. The segregate gen-

era of the Lepidozia complex (Hepaticae) Part 1 . Spru-

cella Steph. and Neolepidozia gen. nov. Brittonia, 11:

77-85, figs. 1-41.

FURUKI, T, AND M. HIGUCHI. 1997. Oil bodies and oil

droplets of some hepatics from Sichuan, China. Bull.

Natl. Sci. Mus. Tokyo, 23: 81-102, pi. 1-8.

GOTTSCHE, C. M., J. B. G. LlNDENBERG, AND C. G. NEES

AB ESENBECK 1844-1847. Synopsis Hepaticarum.

Meissner, Hamburg, pp. 1-834. (Fasc. I, pp. 1-144,

1844; Fasc. II-III, pp. 145-464, 1845; Fasc. IV, pp.

465-624, 1846; Fasc. V, pp. 625-834, 1847).

GROLLE, R. 1964 [1963]. Uber Kurzia v. Martens. Rev.

Bryol. Lichenol., 32: 166-180, pi. 1-2.

GROLLE, R. 1967. Lebermoose aus Neuguinea. 6. Dritte

Fundliste. J. Hattori Bot. Lab., 30: 113-118, pi. 1-2.

GROLLE, R. 1976. Verzeichnis der Lebermoose Europas

und benachbarter Gebiete. Feddes Repert., 87: 171-

279.

HATTORI, S. 1953. Oil-bodies of Japanese Hepaticae (2).

J. Hattori Bot. Lab., 10: 63-78, pi. 1-3.

HERZOG, TH. 1938. Descriptions of new species of New

Zealand hepatics, II. Trans. Roy. Soc. New Zealand,

68: 40-46, pis. 3-7.

HODGSON, E. A. 1956. New Zealand Hepaticae (Liver-

worts)— IX. A review of the New Zealand species of

the genus Lepidozia. Trans. Roy. Soc. New Zealand,

83: 589-620, figs. 1-2.

HODGSON, E. A. 1962a. Hepatics from the subantarctic

islands of New Zealand including "Cape Expedition"

collections from the Auckland and Campbell Islands.

Rec. Domin. Mus., 4: 101-132.

. 1962b. New Zealand Hepaticae (Liverworts) —

XIV. Drucella, a new genus of the Lepidoziaceae.

Trans. Roy. Soc. New Zealand, Bot., 2: 45-47, figs.

1-9.

HODGSON, E. A. 1964. New Zealand Hepaticae (Liver-

worts)— XV. A new monotypic family of the thalloid

Hepaticae Phyllothalliaceae Hodgson fam. nov. Trans.

Roy. Soc. New Zealand, Bot., 2: 247-250, fig. 1.

104

FIELDIANA: BOTANY

HODGSON, E. A. 1965. New Zealand Hepaticae (Liver-

worts)— XVI. A miscellany of new genera, new spe-

cies and notes, part I. Trans. Roy. Soc. New Zealand,

Hot., 3: 67-97, figs. 1-2.

MIZUTANI, M. 1968. Studies of little known Asiatic spe-

cies of Hepaticae in the Stephani Herbarium. 6. On

some interesting tropical species of Lepidozia. J. Hat-

tori Bot. Lab., 31: 176-188, f. I-VII.

PIIPPO, S. 1984. Bryophyte flora of the Huon Peninsula,

Papua New Guinea. VI. Lepidoziaceae subfam. Lep-

idozioideae, Calypogeiaceae, Adelanthaceae, Cephal-

oziaceae subfam. Cephalozioideae and subfam. Odon-

toschismatoideae and Jubulaceae. Ann. Bot. Fennici,

21: 309-335, f. 1-12.

SCHUSTER, R. M. 1963. Studies on Antipodal Hepaticae.

I. Annotated keys to the genera of Antipodal Hepati-

cae with special reference to New Zealand and Tas-

mania. J. Hattori Bot. Lab., 26: 185-309.

SCHUSTER, R. M. 1966. The Hepaticae and Anthocerotae

of North America east of the hundredth meridian. Vol.

I, pp. i-xvii, 1-802, figs. 1-84. Columbia University

Press, New York.

SCHUSTER, R. M. 1969. The Hepaticae and Anthocerotae

of North America east of the hundredth meridian. Vol.

II, pp. i-xii, 1-1062, figs. 85-301. Columbia Univer-

sity Press, New York.

SCHUSTER, R. M. 1973 [1972]. Phylogenetic and taxo-

nomic studies on Jungermanniidae. J. Hattori Bot.

Lab., 36: 321-405, figs. 1-11.

SCHUSTER, R. M. 1980. Studies on Hepaticae, LIV-LVIII.

Kurzia v. Mart. \Microlepidozia (Spr.) Joerg.], Me-

galembidium Schust., Psiloclada Mitt., Drucella

Hodgs., and Isolembidium Schust., J. Hattori Bot.

Lab., 48: 337-421, figs. 1-19.

SCHUSTER, R. M. 1982. Exogenous branching and its

phylogenetic significance in Calobryales and Junger-

manniales. J. Hattori Bot. Lab., 51: 1-50, figs. 1-6.

SCHUSTER, R. M. 1984. Evolution, phylogeny and clas-

sification of the Hepaticae, pp. 892-1070, figs. 36-

100. In Schuster, R. M. ed., New Manual of Bryology.

Vol. 2. Hattori Botanical Laboratory, Nichinan.

SCHUSTER, R. M. 2000. Austral Hepaticae. Beih. Nova

Hedwigia, 118: 1-524, figs. 1-211.

SCHUSTER, R. M., AND J. J. ENGEL. 1974. A monograph

of the genus Pseudocephalozia (Hepaticae). J. Hattori

Bot. Lab., 38: 665-701, figs. 1-17, tabs. 1-2.

SCHUSTER, R. M., AND J. J. ENGEL. 1996. Austral He-

paticae. XXI. Paracromastigum fiordlandiae, sp. nov.

and the delimitation of Paracromastigum and Hyalole-

pidozia (Lepidoziaceae). Brittonia, 48: 165-173, f. 1.

SPRUCE, R. 1884-1885. Hepaticae Amazonicae et An-

dinae. Trans. & Proc. Bot. Soc. Edinburgh, 15: 1-308,

pis. 1-4, 1884 (part I); i-xi, 309-589, pis. 5-22, 1885

(part II).

STEPHANI, F. 1892. A revision of Colenso's New Zealand

Hepaticae, with descriptions of new species collected

by him. J. Linn. Soc., Bot., 29: 263-280, pis. 26-28.

STEPHANI, F. 1898-1924. Species Hepaticarum. Geneve

& Bale, 6 vols.

ENGEL & SCHUSTER: AUSTRAL HEPATICAE. LEPIDOZIA

105

Index to Taxa

Recognized taxa are in roman type and illegitimate or synonymous taxa are in italics. New taxa and

new combinations are in boldface.

Drucella integristipula (Steph.) Hodgs. 2

Herbertia spinosissima (Hook. f. & Tayl.) Trev. 26

Herpetium sect. Lepidozia Nees 4

Jungermannia albula Hook. f. & Tayl. 88

Jungermannia glaucophylla Hook. f. & Tayl. 92

Jungermannia laevifolia Hook. f. & Tayl. 56

Jungermannia microphylla Hook. 28

Jungermannia pendulina Hook. 32

Jungermannia ulothrix Schwagr. 88

Kurzia calcarata (Steph.) Grolle 2

Kurzia compacta (Steph.) Grolle 2

Kurzia hippuroides (Hook. f. & Tayl.) Grolle 2, 57

Kurzia mollis (Steph.) Engel & Schust. 13

Kurzia setiformis (De Not.) Engel & Schust. 13

Lepidozia (Dum.) Dum. 4

III. Incisae Lindenb. 87

I. Microphyllae Lindenb. 28

Subg. Austrolepidozia Schust. 87

Subg. Chaetolepidozia Schust. 13, 19, 88

Subg. Cladolepidozia Schust. 13, 19

Subg. Dendrolepidozia Schust. 23

Subg. Glaucolepidozia Schust. 92

Subg. Mastigolepidozia Schust. 27

Subg. Notholepidozia Schust. 3 1

Sect. Kirkii Schust. 82

Sect. Microphyllae (Lindenb.) Grolle 28

Sect. Notholepidozia 40

Sect. Pendulinae Schust. 32

Sect. Setigerae Schust. 36

Lepidozia acantha Engel 71

Lepidozia albula (Hook. f. & Tayl.) G. L. & N. 88

Lepidozia allisonii Herz. 2

Lepidozia angelii Gott. ex Steph. 85

Lepidozia asperifolia Steph. 56

Lepidozia bidens Engel 79

Lepidozia bisbifida Steph. 95

Lepidozia breviloba Steph. 41

Lepidozia brevipinna Pears. 95

Lepidozia bursifera Hatt. & Grolle 12, 14

Lepidozia calcarata Steph. 2

Lepidozia centipes Tayl. 2

Lepidozia cladorhiza (Reinw. et al.) G. L. & N. 13

Lepidozia colensoana Steph. 52

Lepidozia compacta Steph. 2

Lepidozia concinna Col. 52

Lepidozia corticola Steph. 2

Lepidozia cupressina (Sw.) Lindenb. 5

Lepidozia cupressina var. T pendulina (Hook.) Lehm. &

Lindenb. 32

Lepidozia dentifolia Steph. 82

Lepidozia digitata Herz. 98

Lepidozia dispar (Mont, ex Tayl. & Hook, f.) Mont. 2,

57

Lepidozia elobata Schust. 74

Lepidozia ferdinandi-muelleri Steph. 13

Lepidozia fugax Engel 63

Lepidozia gibbsiana Steph. 2

Lepidozia gigantea Steph. 32

Lepidozia glaucescens Engel 101

Lepidozia glaucophylla (Hook. f. & Tayl.) G. L. & N.

92

Lepidozia gottscheana Lindenb. 2

Lepidozia herzogii Hodgs. 2

Lepidozia hippuroides (Hook. f. & Tayl.) Steph. 2

Lepidozia hirta Steph. 85

Lepidozia integristipula Steph. 2

Lepidozia kirkii Steph. 82

Lepidozia lacerifolia Steph. 14

Lepidozia laevifolia (Hook. f. & Tayl.) G. L. & N. 56

Lepidozia laevifolia var. acutiloba Engel 61

Lepidozia laevifolia var. alpina Schust. & Engel 62

Lepidozia laevifolia var. laevifolia 59

Lepidozia latiloba Col. 52

Lepidozia leptodictyon Herz. 2, 3

Lepidozia longifissa Steph. 26

Lepidozia longiscypha (Tayl.) Carring & Peras. 2

Lepidozia martinii Hodgs. 2

Lepidozia meridiana Hodgs. 2

Lepidozia microphylla (Hook.) Lindenb. 28

Lepidozia minuta Col. 56

Lepidozia multipinna Steph. 28

Lepidozia novae-zelandiae Steph. 31, 66

Lepidozia novae-zelandiae var. heterostipa Schust. 70

Lepidozia novae-zelandiae var. minima Schust. 71

Lepidozia novae-zelandiae var. novae-zelandiae 70

Lepidozia obtusiloba Steph. 43

Lepidozia obtusiloba var. obtusiloba 48

Lepidozia obtusiloba var. parvula Engel 48

Lepidozia ornata Engel 49

Lepidozia papillata Steph. 56

Lepidozia parvitexta Steph. 43

Lepidozia patentissima (Hook. f. & Tayl.) G. L. & N. 2

Lepidozia pearsonii Spruce 5

Lepidozia pendulina (Hook.) Lindenb. 32

Lepidozia pinnaticruris Spruce ex Steph. 12

Lepidozia portoricensis Fulf. 27

Lepidozia praenitens (Lehm. & Lindenb.) G. L. & N. 2

Lepidozia procera Mitt. 41

Lepidozia pulcherrima Steph. 2

Lepidozia pumila Engel 76

Lepidozia radiata Steph. 2

Lepidozia remotifolia Hodgs. 2

Lepidozia reptans (L.) Dum. 4, 5, 8, 13, 14, 15, 16, 17,

18, 31, 32

Lepidozia retrusa Col. 56

Lepidozia robusta Steph. 16

Lepidozia roseana Steph. 2

Lepidozia setigera Steph. 36

Lepidozia spinosissima (Hook. f. & Tayl.) Mitt. 23

Lepidozia subdichotoma Spruce 28

Lepidozia subquadrata Steph. 95

Lepidozia subtransversa Steph. 16

Lepidozia tetradactyla (Hook. f. & Tayl.) G. L. & N. 2

Lepidozia ulothrix (Schwagr.) Lindenb. 88

106

FIELDIANA: BOTANY

Lepidoziopsis Hodgs. 4, 92

Lepidoziopsis glaucophylla (Hook. f. & Tayl.) Hodgs.

92

Mastigophora albula (Hook. f. & Tayl.) Trev. 88

Mastigophora glaucophylla (Hook. f. & Tayl.) Trev. 92

Mastigophora laevifolia (Hook. f. & Tayl.) Trev. 56

Mastigophora microphylla (Hook.) Mont. 28

Mastigophora pendulina (Hook.) Trev. 32

Mastigophora ulothrix (Schwagr.) Trev. 88

Paracromastigum longiscyphum (Tayl.) Schust. & Engel

2

Pleuroschisma sect. Lepidozia Dum. 4

Pseudocephalozia lepidozioides Schust. 2, 3

Sendtnera spinosissima Hook. f. & Tayl. 26

Telaranea centipes (Tayl.) Schust. 2, 8

Telaranea corticola (Steph.) Hodgs. 2

Telaranea gibbsiana (Steph.) Hodgs. 2

Telaranea gottscheana (Lindenb.) Hodgs. 2

Telaranea herzogii (Hodgs.) Hodgs. 2

Telaranea inaequalis Schust. 5

Telaranea martinii (Hodgs.) Schust. 2

Telaranea meridiana (Hodgs.) Hodgs. 2

Telaranea mooreana (Steph.) Schust. 5, 8

Telaranea nematodes (Gott. ex Aust.) Howe 8

Telaranea patentissima (Hook. f. & Tayl.) Hodgs. 2

Telaranea praenitens (Lehm. & Lindenb.) Hodgs. 2

Telaranea remotifolia Hodgs. 2, 5

Telaranea tetradactyla (Hook. f. & Tayl.) Hodgs. 2

Telaranea tetrapila var. roseana (Steph.) Engel & Merr.

2

Telaranea tuberifera Engel & Schust. 8

INDEX

107

A Si ' :ted ;. ,i ' i:ijj

I iti te 22 Hie Gc •

Tax;i ;-;ony of /GV

-el and G. L. Smith Merrill

A Re : i] i South inierican Ipecie; of i

: • ••• : »usly by S. Kriapp and ' R Press

Piease order Fieldiana by publication no. Inquiries and

Peter roM. ;• ;

Fortsas Books, Ltd.

5435 Mo. Lovejoy

Chicago, Illinois 6063'

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