UNIVERSITY OF

ILLINOIS LIBRARY

AT URBANA-CHAMPAIGN

BIOLOGY

! 2001

Botany

NEW • S, NO. 37

Austral Hepaticae. 22. The Genus Balantiopsis

in New Zealand, with Observations on

Extraterritorial Taxa and a Phylogeny of

Balantiopsis and the Family

Balantiopsaceae (Jungermanniales)

John J. Engel

G. L. Smith Merrill

January 31, 1997

Publication 1481

PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY

ion for Contributors to Fieldiana

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Part II. Po

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ISI/NISO Z39.48-1992 (Permanence of Paper).

FTELDIANA

Botany

NEW SERIES, NO. 37

Austral Hepaticae. 22. The Genus Balantiopsis

in New Zealand, with Observations on

Extraterritorial Taxa and a Phylogeny of

Balantiopsis and the Family

Balantiopsaceae (Jungermanniales)

John J. Engel

G. L. Smith Merrill

Department of Botany

Field Museum of Natural History

Roosevelt Road at Lake Shore Drive

Chicago, Illinois 60605-2496

U.S.A.

THE LIBRARY OF THE

^ FEB 2 8 1997

Accepted December 6, 1995 Un8f,f £SITY OF ILLINOIS

Published January 31, 1997 URBANA-CHAMPAIGN

Publication 1481

PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY

101 BUWKLL HALL

MAR 0 4 1997

© 1996 Field Museum of Natural History

ISSN 0015-0746

PRINTED IN THE UNITED STATES OF AMERICA

Table of Contents

ABSTRACT 1

INTRODUCTION 1

THE GENUS BALANTIOPSIS IN NEW ZEALAND .... 2

Key to Balantiopsis in New Zealand 5

B. diplophylla 5

B. montana 12

B. lingulata 14

B. verrucosa 16

B. rosea 17

B. convexiuscula 21

B. tumida 24

S YSTEMATICS 26

Choice of Taxa 26

Character Descriptions and Coding 27

Results 41

Character State Reconstructions 44

Discussion of Phylogenetic Relationships .... 47

Intergeneric Relationships 47

Balantiopsis 49

The Systematic Position of Anisotachis

Schust 54

The Systematic Position of Isotachis

subg. Hypoisotachis Schust 55

PHYTOGEOGRAPH Y AND ANCESTRAL AREAS 56

ACKNOWLEDGMENTS 59

LITERATURE CITED 59

INDEX TO TAXA . .62

3. Balantiopsis diplophylla (Hook. f. &

Tayl.) Mitt 8

4. Balantiopsis diplophylla (Hook. f. &

Tayl.) Mitt 10

5. Balantiopsis montana (Col.) Engel &

Merr. 13

6. Balantiopsis verrucosa Engel & Merr. 18

7. Balantiopsis rosea Berggr. 19

8. Balantiopsis convexiuscula Berggr. 22

9. Balantiopsis tumida Berggr. 25

10. Austroscyphus phoenicorhizus (Grolle)

Schust 30

11. Isotachis minima Pears 31

12. Isotachis montana Col 32

13. Isotachis montana Col 36

14. Isotachis lyallii Mitt 38

15. Four most parsimonious cladograms,

showing relative branch lengths 40

16. Consensus of 13 cladograms with in-

clusion of perianth character 42

17. Consensus cladogram showing charac-

ter state transformations 43

18. Area cladogram 57

List of Tables

List of Figures

1 . Total range of Balantiopsaceae Buch

2. Balantiopsis Mitt. General morphology

1. Taxa included in systematic analysis 27

2. Data matrix used in cladistic analysis of

Balantiopsis and related genera 28

3. Proposed classification of higher taxa of

Balantiopsaceae 48

4. Conspectus of Genus Balantiopsis Mitt. .. 49

in

Austral Hepaticae. 22. The Genus Balantiopsis in

New Zealand, with Observations on

Extraterritorial Taxa and a Phylogeny of

Balantiopsis and the Family Balantiopsaceae

(Jungermanniales)

John J. Engel

G. L. Smith Merrill

Abstract

Seven species of Balantiopsis are recognized in

New Zealand, including B. diplophylla (Hook. f.

& Tayl.) Mitt., B. lingulata Schust., B. rosea

Berggr., B. convexiuscula Berggr., and B. tumida

Berggr. The following new taxa are proposed:

Ruizanthoideae Schust., subfam. nov.; Neesios-

cypheae Engel & Merr., trib. nov.; and Balantiop-

sis verrucosa Engel & Merr., the last a new spe-

cies from New Zealand. The following new com-

binations are proposed: Balantiopsaceae subtribe

Isotachidinae (Hatch.) Engel & Merr., Hypoiso-

tachis (Schust.) Engel & Merr., Balantiopsis sect.

Pteridophylla (Hatch.) Engel & Merr., Balantiop-

sis montana (Col.) Engel & Merr., Balantiopsis

splendens (Steph.) Engel & Merr. (Anisotachis

splendens (Steph.) Schust.), Balantiopsis diplo-

phylla var. hockenii (Berggr.) Engel & Merr., and

Hypoisotachis multiceps (Lindenb. & Gott.) Engel

& Merr. (Isotachis multiceps (Lindenb. & Gott.)

Gott.).

Results of a phylogenetic study of the genus

Balantiopsis are presented, together with obser-

vations on intergeneric relationships in the family

Balantiopsaceae. A discussion of character polar-

ity and a reconstruction of ancestral character

states are included. A realignment of the major

taxa of Balantiopsaceae is proposed, based on

morphological evidence that the perigynium of 1s-

otachis and marsupium of Balantiopsis are ho-

mologous structures. An ancestral area analysis

was undertaken to determine the geographical his-

tory of members of the family.

The analysis of 26 taxa and 25 characters, with

Triandrophyllum subtrifidum as the outgroup,

yielded four most parsimonious cladograms, each

with a length of 114 steps, using the heuristic

search option of PAUP. Monophyly of Balantiop-

sis is supported by the possession of a pendant,

geotropic marsupium. The position of subg. Steer-

eocolea as the primitive element in the genus is

supported; however, no support was found for

maintaining Anisotachis as a distinct taxon, and

the single species of the genus is transferred to

Balantiopsis. Two subgenera of Balantiopsis are

recognized (subg. Steereocolea, subg. Balantiop-

sis), as are two sections within subg. Balantiopsis

(sect. Balantiopsis, sect. Pteridophylla) corre-

sponding to units that were consistently resolved

in the phylogenetic analysis.

Keys to the subfamilies, tribes, and subtribes of

Balantiopsaceae and to the Balantiopsis species in

New Zealand are included.

Introduction

The genus Balantiopsis contains 16 species and

is basically south temperate in distribution, ex-

tending northward to Brazil (B. brasiliensis

Steph.) and to Borneo, New Guinea, and the Phil-

ippines (B. ciliaris Hatt. and B. philippinensis H.

Inoue, the latter endemic to the Philippines). Its

distribution in south temperate regions is bicen-

tric, with six species in southern South America

and the remainder in temperate Australasia. The

closely related genus Anisotachis, including the

single species A. splendens (Steph.) Schust., oc-

curs in southern South America. Four species of

Balantiopsis occur in the Magellanian and Valdi-

vian zones and (with one exception) the Falkland

Islands: B. asymmetrica (Herz.) Engel, B. bisbi-

fida (Steph.) Steph., B. cancellata (Nees) Steph.,

and B. erinacea (Hook. f. & Tayl.) Mitt. Two are

FIELDIANA: BOTANY, N.S., NO. 37, JANUARY 31, 1997, PP. 1-62

Valdivian + Juan Fernandez: B. purpurata Mitt,

and B. crocea Herz. For a discussion of the limits

of these zones, see Engel (1978, 1990). For a map

of the geographical range of Balantiopsis and the

other genera of Balantiopsaceae, see Figure 1.

Following Grolle (1983), we use the spellings

Balantiopsaceae (rather than Balantiopsidaceae)

for the family and Balantiopsoideae (rather than

Balantiopsidoideae) for the subfamily. Herbarium

citations follow the abbreviations in Index Her-

bariorum (Holmgren et al., Regnum Vegetabile

120, 1990)

The Genus Balantiopsis in

New Zealand

The genus Balantiopsis was established by Mit-

ten (1867), based on Jungermannia diplophylla

Hook. f. & Tayl., a New Zealand species. Later,

Mitten (1885) described an additional species, B.

knightii, from New Zealand. Colenso (1888) de-

scribed B. glandulifera, as well as two Chiloscy-

phus species (C. heterodontus and C. montanus)

that have since been proven to belong to Balan-

tiopsis. Berggren (1898) added five additional

New Zealand species to the genus: B. aequiloba,

B. convexiuscula, B. hockenii, B. rosea, and B.

tumida. Engel's (1968) revision of Balantiopsis,

the first major contribution to the knowledge of

the genus since Berggren's, recognized 1 1 species

in the genus worldwide. Three species were rec-

ognized from New Zealand: B. convexiuscula (in-

cluding B. aequiloba), B. diplophylla (Hook. f. &

Tayl.) Mitt. (including B. glandulifera, B. rosea, B.

knightii, and B. hockenii), and B. tumida. Schuster

(1968) added an additional species, B. lingulata.

The results of our study of the New Zealand

representatives of the genus are presented in this

monograph. Our conclusions are based on the ex-

amination of specimens collected by Engel, Child,

Hatcher, Hodgson, and others, plus all relevant

type specimens. The result is a considerable re-

finement of species concepts and the acceptance

of several additional taxa not previously recog-

nized. This is followed by a phylogenetic analysis

of members of the family Balantiopsaceae.

Balantiopsis Mitt, in Hook, f., Handb. New Zea-

land Fl. 751, 753. 1867. Anisotachis Schust.,

Nova Hedwigia 8: 282. 1964, syn. nov. Steer-

eocolea Schust., Bull. Natl. Sci. Mus. Tokyo

11(1): 23. 1968.

Plants strongly anisophyllous, usually creeping

or procumbent (unless growing submerged), less

commonly ascending to stiffly erect, dull to quite

nitid, translucent and rather delicate, with rose-red

to deep red pigments or brownish pigmentation

when developed. Branches ventral-intercalary and

terminal, Frullania-type, sparing. Stems slender,

in surface view with narrowly rectangular, firm-

walled, cortical cells, in cross section the cortex

in 1-2 rows of cells typically smaller and thicker-

walled than those of the medulla; medullary cells

± firm-walled. Rhizoids colorless or deep rose,

smooth-walled, from stem near underleaf bases

and (in B. bisbifidd) from stem near base of ven-

tral lobe of leaf. Leaves (in New Zealand taxa)

distinctly complicate-bilobed, with a ± sharp,

well-developed carina (keel) that is curved and

oriented at right angles to stem, the leaves (at least

the ventral half) with an oblique, succubous in-

sertion. Ventral lobe typically widely spreading to

subsquarrose, at least slightly convex, short bifid,

the segments hardly distinguishable in copiously

ciliate plants, the lobe margins entire to conspic-

uously toothed or ciliate, the tips of the teeth, at

least, formed of elongate cells, the ventral margin

straight or recurved at ventral end and comma-

like to J-shaped or distinctly inverted U-shaped.

Dorsal lobe erect or suberect, sometimes dorsally

assurgent, variable in size, much smaller than to

subequal to the ventral lobe, short bifid, the mar-

ginal armature usually similar to that of ventral

lobe. Underleaves smaller than leaves, shallowly

to deeply bifid, commonly with marginal armature

like that of leaves. Cells thin-walled or, at most,

moderately and evenly thick-walled, or the lon-

gitudinal walls sinuous-thickened, trigones lack-

ing or minute, the cells short-rectangular or hex-

agonal, in median to basal parts of leaves elon-

gate-rectangular, typically very narrow, often 3-

5:1, characteristically in transverse, ± regular

tiers; margins of segments and distal sector of

lobes at times with a border of tangentially elon-

gate cells; cuticle finely to strongly striate-papil-

lose or coarsely guttulate. Oil-bodies in all cells,

hyaline or smoky gray to brownish, 2-13 per cell,

granular to botryoidal.

Plants dioecious. Androecia on main shoot, or

leading Frullania-type or ventral-intercalary

branches, initially terminal in position but becom-

ing intercalary; bracts ± similar to leaves; an-

theridia (1)2-4 in axil of dorsal and ventral lobes,

the stalk biseriate. Gynoecia on leading stems or

branches, at time of fertilization with 1 or more

rings of scalelike to densely ciliiform perigynial

FIELDIANA: BOTANY

cc

I I I I I I I

i- N n * in to r-

ENGEL & MERRILL: AUSTRAL HEPATICAE. BALANTIOPSIS

bractlets, no tubular perianth forming but instead

a pendant, cylindrical, fleshy, conspicuously rhi-

zoidous marsupium develops; mature gynoecium

with innermost subgynoecial bracts with dorsal

and ventral lobes nearly equal in size and form,

the apex of each lobe distinctly divided. Calyptra

distinct, carried down within the marsupium, the

sterile archegonia lifted partway up the calyptra

(i.e., a shoot calyptra).

Seta with 17-29 rows of epidermal cells sur-

rounding an inner core of ca. 27-60 cells of ±

similar size. Capsule long-cylindrical, with spiral

dehiscence, the wall uniformly of 3 layers, the

outer layer of cells ca. equal to or thicker than

both interior strata; outer layer (in surface view)

with semiannular bands; innermost layer of cells

with semiannular bands common, the bands

sometimes forked and anastomosing to delimit fe-

nestrae, or the cells with small nodular thicken-

ings largely confined to the longitudinal radial

walls, the thickenings at times extending onto

both inner and outer tangential walls as faint spurs

or semiannular bands.

Spores with exine brownish yellow, with a net-

work of irregular, coalescing vermiform ridges

that anastomose to form numerous irregular are-

olae; sporeielater diameter ratio 0.7-1.4:1. Elaters

tortuous, bispiral, the spirals loosely wound.

The genus Balantiopsis is characterized by 1)

complicate-bilobed leaves, the dorsal lobe smaller

and folded over the ventral, the margins of the

lobes often armed with teeth or cilia (Fig. 2:10);

2) elongate-rectangular leaf cells, thin-walled and

delicate (Fig. 2:13,15) or with sinuous-thickened

walls, lacking trigones, and oriented in transverse,

± regular tiers (Fig. 9:6); 3) secondary pigments,

often well-developed, reddish, purplish, or brown-

ish; 4) the presence of a marsupium (Fig. 4:1,2),

the sporophyte developing at its base; and 5) cy-

lindrical capsules with spirally twisted valves

(Fig. 2:17).

Most Balantiopsis species show considerable

morphological variation. The development of

teeth and/or cilia on the leaf margins is particu-

larly variable, as are the relative size and shape

of the dorsal leaf lobes. These often are highly

variable from one population to the next, and

sometimes even within the same population. For-

tunately, other characters not emphasized in ear-

lier treatments of the genus have proved helpful

not only in delimiting species but also in distin-

guishing major subdivisions within the genus.

These include form and insertion of the dorsal and

ventral lobes, leaf cell size and number of oil-

bodies per cell, cell-wall thickenings, form of the

cuticle, and shape and papillosity of the marginal

cells of the leaf.

The number of oil-bodies per cell has been

found to be an important difference between spe-

cies in the genus. Balantiopsis rosea and B. ver-

rucosa have 2(3) oil-bodies per cell (typically one

at each end of the cell); B. convexiuscula and B.

tumida have 2-7 oil-bodies per cell; and B. diplo-

phylla, B. lingulata, and B. montana, (5)6-11(13)

per cell.

Cell size is also variable in the New Zealand

species of the genus. For example, the leaf cells

of B. diplophylla are up to 3X as large as those

of B. convexiuscula. There are also differences in

areolation between species. In B. convexiuscula

the leaves are densely areolate (apparent even

with a hand lens), the cells short-rectangular (2:

1), and only 12-18 jim wide, with sinuous-thick-

ened walls; the cells are arranged in precise, reg-

ular tiers so that the lobes have a characteristic

ribbed appearance when dry. In B. diplophylla the

cells of the distal third of the lobes are scattered,

firm-walled, and short-rectangular to polygonal

(ca. 2:1); only in the median sector and below do

the cells become somewhat elongated and ar-

ranged in somewhat irregular tiers. The leaves of

B. montana have a delicate, onion-skin-like tex-

ture related to the relatively large, thin-walled

cells of the leaf, which are 23-34 jjim wide and

up to 110(122) jxm long.

The shape and cuticular ornamentation of the

marginal cells of the leaf lobes are also useful in

species delimitation. In Balantiopsis rosea and B.

verrucosa the marginal cells of both the ventral

and dorsal lobes are tangentially elongated, form-

ing a rather distinct border. The border cells have

a finely striate-papillose cuticle, whereas the me-

dian lobe cells are coarsely papillose, with high,

rounded (guttulate) papillae.

Color is also helpful in species differentiation

and is usually retained in dried material: B. rosea

and B. diplophylla are often rose-pink in color,

whereas B. convexiuscula and B. tumida, when

pigmented, are dull purplish; B. montana is typi-

cally pale translucent green to warm brown, with

delicate, often highly nitid leaves.

FIELDIANA: BOTANY

Key to Balantiopsis in New Zealand

1. Leaf cells small (areolation dense), typically 12-16 |xm wide, short-rectangular (2:1), at most 42 jim

long, the cells in the distal sector with sinuous thickened walls; lobe margins minutely denticulate

by projecting septa between the marginal cells, not bordered; dorsal lobe typically as broad as long,

broadly orbicular to suboblate 2

2. Leaf margins (especially lobe apices) coarsely serrate, the teeth often terminating in elongate cilia;

sinuses between apical segments V-shaped; dorsal leaf lobe somewhat smaller than the ventral;

underleaves (often squarrose, spreading when plant suberect) smaller and deeply dissected (bifid to

0.5-0.6), with long, often curved, tapering cilia; plants prostrate to suberect, rather rigid, olive-green

(or sometimes reddish-pigmented), terrestrial B. convexiuscula (p. 21)

2. Leaf margins (especially lobe apices) broadly dentate, with a short apiculus, lacking cilia; sinuses

between apical segments shallow and lunate; dorsal and ventral leaf lobes subequal in size; un-

derleaves rather large, broad, and shallowly bilobed (bifid to 0.4), often longitudinally plicate;

plants typically erect, ± soft-textured, brownish to often vinaceous, often submerged

B. tumida (p. 24)

1. Leaf cells larger (areolation lax), at least 16 |xm to as much as 35(40) |xm wide (exc. where obviously

secondarily divided), short-polygonal or elongate-rectangular, 65 u,m or more long, with evenly thick-

ened walls (not sinuous), often lax and thin-walled; lobe margins without projecting septa, sometimes

with a differentiated border; dorsal lobe variable in shape; broadly to narrowly ovate to ligulate,

typically much smaller than the ventral lobe, sometimes greatly reduced in size and deeply bifur-

cate 3

3. Leaf cells coarsely papillose (often grossly so, the papillae high and spherical to slightly elon-

gated); marginal row of cells elongated and finely striate-papillose, forming a ± distinct border;

ventral lobe often canaliculate to conduplicately folded, the insertion strongly arched (U-shaped);

dorsal lobe transversely inserted, the free margin not decurrent; oil-bodies 2(3) per cell, at opposite

ends of cell 4

4. Leaves copiously armed with cilia, the ventral lobe with up to 20 cilia, the dorsal with up to

16; underleaves narrowly ovate, divided to 0.4, copiously ciliate; plants often clear wine-

red B. rosea (p. 17)

4. Leaves with reduced armature, the ventral lobe 2-9 dentate-ciliate, the dorsal lobe with up to

5 laciniate-lobulate; underleaves broadly ovate to suborbicular, divided to 0.2 or less, sparingly

armed; plants green to brownish or tinged with red B. verrucosa (p. 16)

3. Leaf cells finely striate-papillose (rarely almost smooth) throughout; marginal row of cells not

elongated, not forming a distinct border; ventral lobe flat or at most moderately convex, longi-

tudinally inserted, not or only moderately arched (at most J-shaped); dorsal lobe insertion oblique,

the free margin decurrent; oil-bodies numerous (up to 13), scattered within cell 5

5. Leaf cells typically scattered in distal 0.3-0.5, not in evident transverse rows, the median cells

polygonal to rather short-rectangular (ca. 2.5:1); ventral lobe rather broadly elliptic to broadly

ovate B. diplophylla (p. 5)

5. Leaf cells in ± evident transverse rows, the median cells elongate-rectangular (to 4:1); ventral

lobe narrowly elliptic to Ungulate 6

6. Dorsal lobe dorsally assurgent (at times distinctly so); ventral lobe plane or at most slightly

convex, the insertion ± straight or only slightly recurved (comma-like) at ventral end;

ventral lobe margins with cilia mostly confined to the distal 0.3 of lobe

B. montana (p. 12)

6. Dorsal lobe appressed, the keel sharply folded; ventral lobe convex, the insertion strongly

recurved at ventral end (J-shaped); ventral lobe margins ciliate to the base

B. lingulata (p. 14)

Subgenus Balantiopsis Section Balantiopsis

Balantiopsis subg. Schizophyllon Hatt., J. Jap. Hot. 41: Balantiopsis diplophylla (Hook. f. & Tayl.) Mitt.

129. 1966, syn. nov. Figures 3 and 4.

ENGEL & MERRILL: AUSTRAL HEPATICAE. BALANTIOPSIS 5

6 7 if 48

FIG. 2. Balantiopsis Mitt. General morphology. 1-8, Sectors of stem showing insertion lines, for each species the

dorsal view at left and the lateral view at right (all at scale indicated for Fig. 8). 1, 2, B. bisbifida. 3, 4, B. convexiuscula

(ul = underleaf). 5, 6, B. verrucosa (ul = underleaf). 7, 8, B. erinacea (ul = underleaf; vl = ventral lobe; dl = dorsal

lobe). 9, Sector of main shoot of B. verrucosa, ventral view. 10, Sector of main shoot of B. rosea, ventral view (a

pair of leaves between underleaf and uppermost scar removed for clarity). 11, Cross section of ventral lobe of leaf

FIELDIANA: BOTANY

Plants procumbent to loosely caespitose, whit-

ish pale green to light green to warm brown, oc-

casionally rose-tinted; shoots complanate, to 6

mm wide. Branching rather common, of Frullania

and ventral-intercalary types, the frequency of

each varying with growth habit of the plants.

Leaves loosely to densely imbricate, rather sharp-

ly folded, the ventral lobe obliquely to widely

spreading, to 3.4 mm long, the free margin of dor-

sal lobe ± straight, the opposing dorsal lobes

touching to moderately overlapping. Ventral lobe

moderately convex (dorsally assurgent and con-

cave in var. hockenii), the insertion moderately to

distinctly J- shaped, the margin decurrent on the

stem, the lobes broadly elliptic to broadly ovate,

obscurely to distinctly bifid; segments acuminate,

at times medium to broadly acute (then apiculate),

(7)10-16 cells wide at base, terminating in a uni-

seriate row of 2-3 (var. hockenii) to 3-4 cells, the

walls often thicker than those of the lamina cells;

lamina margins gently and broadly curved, (1)2-

8 dentate-ciliate to dentate-laciniate, occasionally

entire (except for 2 apical teeth), the ventral mar-

gin 1-7 dentate-ciliate, each with a uniseriate row

of 3-4 cells, the dorsal margin more sparingly

armed; no border. Dorsal lobe variable, smaller

than the ventral lobe, to ca. 3X the stem in width

(often small and with a reduced lamina in var.

hockenii), plane, the insertion transverse to some-

what oblique, then often abruptly curved and de-

current on the stem, the lobes narrowly to broadly

ovate, shallowly to distinctly bifid, the segments

often longer and more attenuate than in ventral

lobe; margins 1-6-dentate-ciliate, occasionally

entire, the dorsal (free) margin ± straight, 0-3-

dentate-ciliate. Underleaves large, about equal to

or slightly larger than the dorsal lobes in size,

weakly imbricate, appressed to moderately

spreading, ± plane, the insertion broadly U-sha-

ped, narrowly to broadly ovate (exclusive of cil-

ia), the underleaves bifid to 0.5 (the underleaves

small and more deeply bifid in var. hockenii);

lobes linear to attenuate, ± plane, with several

cilia, the inner margins often entire; lamina mar-

gins ciliate to the base, at times with a lateral lob-

ule approaching the lobes in size and shape, the

ultimate lobe divisions and cilia terminating in a

uniseriate row of 4-6 elongate cells, the margins

decurrent on the stem. Ventral lobe cells in distal

0.35-0.5 of lobe typically scattered, short-rectan-

gular to polygonal (ca. 2:1), and thin- walled but

firm, the median cells in somewhat irregular trans-

verse tiers, somewhat more elongate-rectangular

and narrower, 27-38 u,m wide X 59-80(85) u,m

long; median cells of dorsal lobe similar to those

of ventral lobe or slightly smaller; cuticle smooth

to finely striate-papillose. Oil-bodies hyaline or

gray, (5)6-11(13) per cell, coarsely granular to

distinctly botryoidal (see under varieties).

Androecia terminal on main shoot or ventral-

intercalary branches; bracts similar to leaves or,

occasionally, more imbricate and more pigment-

ed, with 1-2 abaxial bulges; antheridia 3-4 per

bract, the stalk biseriate; paraphyllia present in

bract axils, squamiform. Gynoecia on main shoot

or ventral-intercalary or Frullania-type branches

of varying lengths; bract margins undulate to cris-

pate and sparingly dentate-laciniate. Marsupia

green to deep rose in color, covered with rose-

colored rhizoids.

Sporophyte not seen.

TAXONOMY — The most distinctive feature of

this species is the field of scattered (irregularly

arranged), ± isodiametric cells of the distal third

of the ventral lobe, evident under the dissecting

microscope, as is the complete absence of a dif-

ferentiated border (Fig. 3 : 4). All other New Zea-

land representatives of the genus have the cells

arranged in regular, well-defined, transverse rows.

The cuticle is finely striate-papillose and differs

markedly from the often grossly papillose (gut-

of B. rosea (vm = ventral margin; ds = dorsal surface). 12, 13, Oil-bodies of median ventral lobe cells of B. lingulata

(X575). 14, Oil-bodies of median ventral lobe cells of B. convexiuscula (X1225). 15, Oil-bodies of median ventral

lobe cells of B. rosea (X890). 16, Longitudinal section through distal portion of young marsupium of B. convexiuscula

showing perigynial bractlets at mouth. 17, Capsule of B. bisbifida showing spiral lines of dehiscence. 18, Capsule

wall of B. bisbifida, cross section. (Figs. 1, 2 from Engel 11144, Chile, Prov. Valdivia, Volcdn Quetrupillan; 3, 4, 16

from Engel 21686, New Zealand, South Is., Westland Prov., Westland Natl. Park, track to Lake Gault; 5, 6, 9 from

type of B. verrucosa; 7, 8 from Engel 11978, Chile, Prov. Chiloe, Isla Chiloe\ Cocauque area; 10, 11 from Child

5612, New Zealand, South Is., Southland Prov., Wilmot Pass Road to Deep Cove; 12 from Engel 20954, New Zealand,

North Is., North Auckland Prov., Maungataniwha Range, Mangamuka Walkway; 13 from Engel 21139, New Zealand,

North Is., North Auckland Prov., southern edge of Waipoua Forest; 14 from Engel 20986, New Zealand, North Is.,

North Auckland Prov., Omahuta Forest Kauri Sanctuary; 15 from Engel 21900, New Zealand, South Is., Otago Prov.:

Mt. Aspiring Natl. Park, Blue Valley Track; 17, 18 from Engel 6389, Chile, Prov. Magallanes, Bahia San Nicolas.)

ENGEL & MERRILL: AUSTRAL HEPATICAE. BALANTIOPSIS

FIG. 3. Balantiopsis diplophylla (Hook. f. & Tayl.) Mitt. 1, Sector of main shoot, dorsal view. 2, 3, Leaves. 4,

Distal portion of ventral lobe of leaf. 5, Median cells of ventral lobe. 6, Sector of main shoot, ventral view. 7,

Underleaf. (All from type of B. diplophylla (FH).)

FIELDIANA: BOTANY

tulate) cuticle of B. rosea and B. verrucosa. The

insertion of the ventral lobe is not or only mod-

erately recurved (J-shaped, Fig. 3 : 6). The ventral

lobes are broadly elliptic to broadly ovate.

Balantiopsis diplophylla was given a broad cir-

cumscription in Engel (1968) and would have in-

cluded, in addition to the typical expression, the

taxa we now recognize as B. rosea, B. lingulata,

B. montana, and B. diplophylla var. Hockenii.

This species, B. lingulata, and B. montana to-

gether form a complex of closely related plants

all sharing a similar leaf form, including a ten-

dency for armature development, and leaf cells

with many gray, opaque oil-bodies per cell. Care

should be taken in differentiating these species,

including the selection of optimally developed

shoots within a population for study.

The aspect of this species at times approaches

that of B. convexiuscula, with the ventral lobes

similar in shape, armature, and orientation. The

leaf cells of B. diplophylla, however, are up to 3 X

as large as those of B. convexiuscula, which

makes them easily distinguishable when the two

species occur together. A distinctive growth form

of B. diplophylla that occurs at higher elevations

is often deeply rose-pigmented with strongly dor-

sally assurgent leaves, lending the plant a chan-

neled appearance like that of a species of Junger-

mannia.

DISTRIBUTION — This species is reportedly dis-

tributed from Auckland Island in the south to New

Zealand, Tasmania, Australia, New Caledonia,

and the Philippines, but we have not reevaluated

its variation in areas outside New Zealand. How-

ever, the typical expression of the species (as in

the type, Fig. 3) is found predominantly on the

subantarctic islands and the southern sector of

South Island.

Two varieties may be recognized in New Zea-

land.

Key to Varieties of B. diplophylla

1. Ventral lobe broadly elliptic to broadly ovate, the sinus often conspicuous, often broadly rounded or

angular, the apical segments terminating in a uniseriate row of 3-4 cells, the lobe margins up to

8-ciliate-dentate, the insertion typically J-shaped; dorsal lobe rather large, narrowly to broadly ovate,

usually armed with at least a few divergent cilia; underleaves conspicuous, the sinus reflexed; oil-

bodies smoky grayish brown var. diplophylla

1 . Ventral lobe scarcely longer than wide, subelliptic, the sinus typically a mere narrow, rounded apical

notch, the apical segments terminating in a uniseriate row of 2-3 cells, the lobe margins almost

entire, the insertion nearly longitudinal, scarcely recurved; dorsal lobe often greatly reduced, often

ligulate to vestigial and reduced to a bifurcate appendage: underleaves small and inconspicuous, the

sinus plane, not reflexed; oil-bodies hyaline var. hockenii

Balantiopsis diplophylla (Hook. f. & Tayl.) Mitt,

var. diplophylla

Jungermannia diplophylla Hook. f. & Tayl., Lond. J.

Bot. 3: 377. 1844. Gottschea diplophylla (Hook,

f. &. Tayl.) Nees in G. L. & N., Syn. Hep. 624.

1846. Gymnanthe diplophylla (Hook. f. & Tayl.)

Mitt, in Hook, f., Bot. Antarc. Voy. 2: 230. 1859.

Balantiopsis diplophylla (Hook. f. & Tayl.) Mitt.

in Thorns. & Mum, Rep. Sci. Res. Voy. Chal-

lenger Bot. 1(3): 87. 1885. Lectotype (nov.):

Auckland Is., Hooker (FH!); isolectotype: (NY!).

Chiloscyphus heterodontus Col., Trans. & Proc. New

Zealand Inst. 21: 62. 1888 [1889], ("heterodon-

ta"). Original material: New Zealand, North Is.,

sides of Mt. Tongariro, "County of East Taupo,"

with type of C. montanus, 1887, Hill.

Plants often robust, procumbent to loosely

caespitose. Leaves loosely to densely imbricate;

ventral lobes obliquely spreading, broadly elliptic

to broadly ovate, often distinctly bifid, the apical

segments acuminate to medium or broadly acute,

the margins ciliate-dentate with up to 8 cilia; dor-

sal lobe rather large, narrowly to broadly ovate,

usually armed with up to 7 divergent cilia. Oil-

bodies scattered throughout cell, but with tenden-

cy for several to cluster at each end, pale smoky

gray (chocolate-brown under lower magnifica-

tion), (5)6-1 1 per cell, coarsely granular, the sur-

face appearing papillose, 1 1.8-14.7 X 4.9-8.8 u,m

to 15.7-17.6 X 6.9-7.8 jjim.

DISTRIBUTION AND ECOLOGY — The variety oc-

curs in Auckland Island and New Zealand. The

distribution of the species should be reevaluated

according to the narrower circumscription docu-

mented here.

In New Zealand the variety is primarily a low-

ENGEL & MERRILL: AUSTRAL HEPATICAE. BALANTIOPSIS

FIG. 4. Balantiopsis diplophylla (Hook. f. & Tayl.) Mitt. 1-7, var. diplophylla; 8-13, var. hockenii. 1, Female

inflorescence (X9). 2, Median-longitudinal section of marsupium (a = shoot calyptra; b = unfertilized archegonium;

c = seta) (X 10). 3, Female bract (X9). 4, Antheridial stalk. 5, Segment of ventral lobe of leaf, cuticle shown in part.

6, Sector of ventral margin of ventral lobe. 7, Segments of dorsal lobe (dl) of leaf. 8, 9, Leaves. 10, 11, Distal portion

of ventral lobes (vl). 12, 13, Distal portion of dorsal lobes (dl). (Fig. 1 from Kerr, New Zealand; 2 from

10

FTELDIANA: BOTANY

er- to middle-elevation forest plant of the southern

half of South Island. It typically occurs on damp,

shaded, clayey soil of steep slopes or banks (often

those associated with watercourses) and boggy

niches under shelter of, for example, manuka. It

also occurs under Gleichenia and rushes in paki-

his. The variety is occasionally present in the al-

pine zone, where it may be found under cover of

tussocks.

SELECTED SPECIMENS SEEN— NEW ZEALAND, SOUTH

IS., SOUTHLAND PROV.: Lake Hauroko, ca. 300 m,

Child HI 566 (F); Fiordland Natl. Park, below Mt. Burns,

E of Borland Saddle, S of South Branch of Borland

Burn, 1225-1320 m, Engel 18570 (F); ibid., Burns

Creek, Mt. Burns, ca. 915 m, Child H5192 (F). OTAGO

PROV.: Mt. Cargill, N of Dunedin, ca. 455 m, Child

HI 329 (F); Swampy Hill, Dunedin, ca. 610 m, Child

H1929 (F); Hindon, ca. 300 m, Child H2298—C. mar.

(F). WESTLAND PROV.: Westland Natl. Park, Gilles-

pies Cook River Road, between Tornado Creek and

Whelan Creek, Engel 6590B (F); ibid., S margin of Lake

Wahapo, Engel 6736 (F); 1 km N of White Horse Creek,

ca. 150-300 m, Child H5414, H5385 (F); 10 km S of

Greymouth, ca. 150 m, Child H4946 (F); Rapahoe Range

(or Twelve Apostles Range), N of Greymouth, ca. 150

m, Child H5399 (F); Paparoa Range, Croesus Track,

ridge between Granite Creek and Pagan Creek water-

sheds, 420 m, Engel 19329 (F). NELSON PROV: Pa-

paroa Natl. Park, Inland Pack Track, SW of terminus of

Bullock Creek Road, NE of the Punakaiki, ca. 35 m,

Engel 21659 (F); Dolomite Point Scenic Reserve, 30 m,

Fife 4862 (F); Big Totara River, ca. 60 m, Child H5445

(F); 10 km S of Westport, ca. 30 m, Child H4936 (F).

NORTH IS., WELLINGTON PROV: Mt. Ruapehu,

Blythe Track off Ohakune Mt. Road, ca. 1220 m, Brag-

gins 92/79 as B. rosea (F). TARANAKI PROV: Pukeiti

Bush, near New Plymouth, Hatcher 273 (F). NORTH

AUCKLAND PROV: NE Waitakere Range, Swanson

University Reserve, Tram Valley Road, 95 m, Engel

20451 (F); Rangitoto Is., immediately E of Auckland,

50-160 m, Engel 20793 (F); McElroys Bush Reserve,

western side of Mahurangi Harbour on Cowan Bay

Road, 95 m, Engel 20379 (F); Pohuehue Scenic Reserve,

N of Auckland on State Highway 1, ca. 25 m, Engel

21144 — c. mar. (F); Waipoua Forest, Lookout Track be-

tween Lookout Road and Waipoua River Road, above

forest headquarters, ca. 240 m, Engel 21071 (F); Man-

gamuka Walkway, Maungataniwha Range, ESE of Kai-

taia, saddle on State Highway 1, 390 m, Engel 20963

(F); Radar Bush, WSW of Cape Reinga, S of Mt. Te

Paki, ca. 100 m, Engel 20853.

Balantiopsis diplophylla var. hockenii (Bergrr.)

Engel & Mem, comb. & stat. nov. Figure 4 : 8-

13.

Balantiopsis hockenii Berggr., On New Zealand Hep.

1: 46. / 31. 1898, ("Hockem"). Lectotype (fide

Engel, 1968): New Zealand, South Is., Dunedin,

Berggren 3175 (LD!); isolectotypes: (FH!, GB!,

UPS!).

Balantiopsis knightii Mitt, in Thorns. & Mum, Rep.

Sci. Res. Voy. Challenger Bot. 1(3): 87. 1885.

Original material: New Zealand, without specific

location, Knight (NY!).

Plants typically prostrate. Leaves often dorsally

assurgent, the ventral lobe subelliptic, typically

hardly longer than wide, typically with a narrow,

rounded apical notch, the margins few-toothed to

almost entire; dorsal lobe often greatly reduced,

narrowly ovate to ligulate, sharply bidentate at the

apex, the margins entire, the dorsal lobe often ves-

tigial and reduced to a bifurcate appendage; un-

derleaves small, deeply bifid, sparingly ciliate, of-

ten concealed by dense rhizoids arising from the

stem at their base. Oil-bodies scattered throughout

cell, hyaline, (5)7-11(13) per cell, mostly ovoid

to elliptic, some spherical, coarsely granular (the

surface appearing coarsely papillose) to distinctly,

botryoidal, 11.8-12.7 X 4.9-5.9 (Jim to 11.8-15.7

X 6.9-10.8 u.m, a few 14.7 X 5.9-6.9 |xm.

TAXONOMY — This rather common expression is

distinctive, even though it grades imperceptibly

into more typical forms of the species. The ventral

lobes are broadly rounded at the apex and char-

acteristically ± dorsally assurgent and then re-

flexed, lending the plant its distinctive aspect. The

dorsal lobes are typically reduced in size (Fig. 4:

12, 13) and in extreme cases may be reduced to

a minute, forked appendage at the base of the ven-

tral lobe. A range of variation in development of

the dorsal lobe often can be seen within plants of

a single population or even on the same shoot.

Despite the above characterization of var. hock-

enii as a somewhat suboptimal expression of the

species, it frequently bears marsupia.

NOMENCLATURE — The varietal epithet "hock-

enii" (based on B. hockenii Bergrr.) is adopted

here, since it has been commonly applied in New

Zealand to this plant (cf. discussion in Hodgson,

1958), in preference to one based on B. knightii

Mitt., which is older as a species.

DISTRIBUTION AND ECOLOGY — Occurring in New

Hatcher 303, New Zealand, North Is., North Auckland Prov., Waipoua Kauri Forest; 3 from Weymouth, Tasmania;

4 from Engel 19329, New Zealand, South Is., Westland Prov., Paparoa Range, Croesus Track; 5-7 from type of B.

diplophylla (FH); 8-13 from type of B. hockenii.)

ENGEL & MERRILL: AUSTRAL HEPATICAE. BALANTIOPSIS

11

Zealand, with a range and ecology similar to those

of the typical variety.

SELECTED SPECIMENS SEEN— NEW ZEALAND, SOUTH

IS. SOUTHLAND PROV.:Fiordland Natl. Park, Borland

Road, 14 km by road WNW from Borland Lodge, near

South Branch of Borland Burn, W of Monowai, 855-

870 m, Engel 18688 (F). OTAGO PROV.: Mt. Maun-

gatua, W of Mosgiel, ca. 500 m, Engel 17784 (F); Mor-

risons Creek, N of Dunedin, ca. 300 m, Child H1474

(F); W slope of Flagstaff, NW of Dunedin, 490-520 m,

Engel 17622 (F); S side of Mt. Cargill, just below sum-

mit, N of Dunedin, ca. 670 m, Engel 17572 (F); Trotters

Gorge Scenic Reserve, 1 km N of hut, ca. 1 20 m, Child

s.n. 136 (F); Fiordland, N of McKerrow River, Martin's

Bay, Hatcher 759 (F). WESTLAND PROV.: Turiwhate

Falls on Route 73, Engel 6542 (F); Westland Natl. Park,

S margin of Lake Wahapo, Engel 6741 (F); Arthur's Pass

Natl. Park, Lower Otira Bridge, Engel 6822 (F). CAN-

TERBURY PROV.: Mt. Cook Natl. Park, E facing slope

of Mt. Wakefield, just below Wakefield Falls, 870 m,

Engel 18156 (F). NELSON PROV: Paparoa Range, N

side of Tiropahi or Four Mile River between sea and

Route 6, S of Charleston, 130-170 m, Engel 19252 (F);

Nelson Lakes Natl. Park, NE margin of Lake Rotoroa,

W of St. Arnaud, 520 m, Engel 21521 (F); Stony Creek,

E of Westport, ca. 150 m, Child H3543 (F). MARL-

BOROUGH PROV: Pine Valley, Wairau River, ca. 150

m. Child H4364A (F). NORTH IS., HAWKE'S BAY

PROV: "Kiwi," Wairoa, Hodgson as B. hockenii (F).

NORTH AUCKLAND PROV: Waitakere, Child 2261

(F); Waipoua Kauri Forest, Hatcher 438 (F); McElroys

Bush Reserve, western side of Mahurangi Harbour on

Cowan Bay Road, 95 m, Engel 20377— c. mar. (F); SE

of Goat Is. and between Goat Is. and Cape Rodney, N

of Leigh, 30 m, Engel 20255 (F); junction of Okahu

Stream and unnamed stream, Kiwanis Reserve, ca. 5 km

S of Kaitaia, N edge of Herekino Forest area, ca. 60-

80 m, Engel 20903 (F); Radar Bush, WSW of Cape

Reinga, S of Mt. Te Paki, ca. 100 m, Engel 20844 (F).

Balantiopsis montana (Col.) Engel & Merr.,

comb. nov. Figure 5.

Chiloscyphus montanus Col., Trans. & Proc. New

Zealand Inst. 21: 62. 1888 [1889] ("montana"),

non C. montanus Steph., J. Proc. Roy. Soc. N.

S. W. 48: 103. 1914. Lectotype (nov.): New Zea-

land, North Is., sides of Mt. Tongariro, "County

of East Taupo," 1887, Hill (Colenso a. 1310)

(WELT!).

Balantiopsis glandulifera Col., Trans. & Proc. New

Zealand Inst. 21: 64. 1888 [1889], syn. nov.

Original material: New Zealand, North Is., Mt.

Tongariro, "County of East Taupo," Hill (Col-

enso a. 1428) (BM!, WELT! — c. mar.).

Plants loosely prostrate to ascending, typically

pale translucent green, at times yellowish green to

warm brown, sometimes tinged with red, often

highly nitid; shoots markedly complanate, to 6

mm wide. Branching sparing, of Frullania type;

ventral-intercalary branches sporadic. Leaves

soft-textured, loosely imbricate, not sharply fold-

ed, the ventral lobes widely spreading, to 3.2 mm

long, the dorsal lobes dorsally assurgent, at times

distinctly so, the opposing dorsal lobes slightly

overlapping (excluding cilia) or not. Ventral lobe

plane to feebly convex, longitudinally inserted

and not or typically slightly recurved at ventral

end, not decurrent or weakly so, ± narrowly el-

liptic to lingulate, the apex rather blunt to trun-

cate, the lobes very shallowly bifid; segments at

times barely differentiated from marginal arma-

ture, acuminate, entire or sparsely ciliate, 4-7

cells wide at base, terminating in a uniseriate row

of 3-5 elongate cells (to 3.5:1) with ± thickened

walls, the septa thickened in the corners and

weakly projecting, the sinus hardly differentiated;

lamina margins ± straight, 5-13-ciliate, the ar-

mature often crowded in distal sector, the ventral

margin with 3-8 cilia, weakly contracted at the

base, the dorsal margin typically with fewer cilia;

cilia slender, similar to those of apical segments;

no border. Dorsal lobe much smaller than the ven-

tral lobe, typically longer than wide and to 1.5X

the width of stem (excluding armature), plane, the

insertion ± oblique, weakly and somewhat

abruptly decurrent on the stem, the lobes narrowly

ovate, very shallowly bifid or at times impercep-

tibly so, the segments as in ventral lobe; lobe mar-

gins curved, armed with up to 10 widely diver-

gent cilia. Underleaves about equal to the dorsal

leaf lobes in size, ± distant, appressed, plane, the

insertion broadly inverted U-shaped, the under-

leaves narrowly ovate, bifid to 0.6; lobes attenu-

ate, the margins with several cilia, the sinus

V-shaped to narrowly rounded, not reflexed; lam-

ina margins ciliate to the base, with a lobuliform

process on each side approaching the lobes in size

and shape, the margins decurrent on the stem; cil-

ia with a uniseriate row of 4-5(6) elongate cells.

Ventral lobe cells in evident transverse rows,

short-rectangular in the apical portion to elongate-

rectangular in median and basal portions (ca. 4:1

in median sector), evenly thin-walled, the median

cells 23-34 |im wide X 74-110(122) u,m long;

median cells of dorsal lobe similar to those of

ventral lobe; cuticle finely striate-papillose. Oil-

bodies smoky gray, 8-11(12) per cell, coarsely

granular.

Androecia terminal on leading ventral-interca-

lary branches; bracts similar to leaves but some-

what smaller (especially the dorsal lobe), with 1-

2 abaxial bulges; antheridia 3 per bract, the stalk

biseriate; paraphyllia present in bract axils,

squamiform.

12

FIELDIANA: BOTANY

FIG. 5. Balantiopsis montana (Col.) Engel & Merr. 1, Sector of main shoot, dorsal view. 2, 3, Three leaves (top)

and 4 below. 4, Distal ca. 0.75 of ventral lobe of leaf (vm = ventral margin). 5, Median cells of ventral lobe (cuticular

papillae shown in part). 6, Segment of ventral lobe of leaf. 7, Distal portion of dorsal lobe of leaf. 8, Sector of main

shoot, ventral view. 9, Underleaves. (Figs. 1, 2, 4-9 from Engel 18525, New Zealand, South Is., Westland/Canterbury

Prov. Boundary, Arthur's Pass Natl. Park, Arthur's Pass area, Upper Twin Creek Valley; 3 from Engel 21905, New

Zealand, South Is., Otago Prov., Mt. Aspiring Natl. Park, Blue Valley Track.)

ENGEL & MERRILL: AUSTRAL HEPATICAE. BALANTIOPSIS

13

TAXONOMY — Balantiopsis montana is most

similar to B. lingulata but is notable for the wide-

ly divergent, even more distinctly lingulate (par-

allel-sided, with a broadly rounded apex) but

more sparingly ciliate ventral leaf lobes and the

rather small, dorsally assurgent, spinulose-ciliate

dorsal lobes. The number of cilia on the ventral

lobe margins varies from 5 to 10, whereas B. lin-

gulata may have up to 26 cilia. The ventral lobe

armature tends to be concentrated in the distal

sector of the lobe, accentuating the lingulate ap-

pearance of the lobes, whereas in B. lingulata the

lobe is typically ciliate to the base. The plants are

typically pale translucent green to warm brown,

with delicate, membranous-textured, often highly

nitid leaves. Both species have ciliate lobes that

lack a differentiated border, segments and cilia

with a uniseriate row of 4-5 elongate cells, and

lingulate ventral lobes (at times narrowly elliptic

in B. montana).

Balantiopsis montana is a prostrate to ascend-

ing plant that may develop reddish pigments and

has leaves that are not sharply folded, the dorsal

lobe being dorsally assurgent (Fig. 5:1). The ven-

tral lobes are of a different form than in B. lin-

gulata, being essentially plane or at most slightly

convex, with the insertion (at most) only slightly

recurved at the ventral end (comma-like, Fig. 5:

7), and the median cells of the ventral lobe longer.

NOMENCLATURE — The original material of Chi-

loscyphus montana is a mixture of two elements,

which Colenso himself thought might represent

two species. One element, which Colenso regard-

ed as the "young state" of the species, is "whitish

pellucid, [the] leaves with fewer and more distant

teeth," and is referable to B. diplophylla var.

hockenii. The other element, consisting of "aged"

plants, "longer, dark-brown, more flexuous and

sub-rigid, with its leaves more distinct," is the

major element in the enclosed micropacket la-

beled Chiloscyphus montana in Colenso's own

hand. This latter element is selected as the lecto-

type of the species.

The type of Balantiopsis glandulifera bears

marsupia as well as young gynoecia in the earliest

stages of development and consists of ± flaccid,

plagiotropic shoots with abundant rhizoids. The

underleaves are small. The shape of the ventral

lobes is suggestive of B. lingulata, but the ventral

lobe insertion is essentially straight at the ventral

end, and the dorsal lobes are somewhat dorsally

assurgent and quite narrow. The ventral lobe cells

are evidently tiered and elongate-rectangular (2.5-

3.5:1). These features seem to ally this plant with

B. montana.

Chiloscyphus montana and B. glandulifera ap-

peared in the same publication as did Chiloscy-

phus heterodonta, the type of which we have not

seen. The type material of all three species came

from Mount Tongariro, collected by Hill; C. mon-

tana and C. heterodonta were "picked out both

from a quantity of broken vegetable rejectamen-

ta."

DISTRIBUTION AND ECOLOGY — Endemic to New

Zealand, occurring sporadically on both South

and North Islands. It occurs at a variety of ele-

vations, from lower- to upper-elevation forests to

the subalpine zone, and may be found, for ex-

ample, in deep, well-protected pockets of stream

banks, slopes, vertical banks, etc. In forests the

species also occurs under wet, deeply shaded

overhangs and on heavily shaded, dripping, ver-

tical cliff faces, where it may form large, pure

sheets or masses that are loosely attached and may

even festoon the substrate. The species evidently

requires sheltered, well-shaded, hyperhumid nich-

es (see comments under B. rosed).

SELECTED SPECIMENS SEEN— NEW ZEALAND, SOUTH

IS., SOUTHLAND PROV.: Fiordland Natl. Park, Bor-

land Road, 14 km by road WNW from Borland Lodge,

near South Branch of Borland Burn, W of Monowai,

855-870 m, Engel 18713 (F). OTAGO PROV.: Trotters

Gorge Scenic Reserve, 1 km N of hut, ca. 60 m, Child

H4413 as B. diplophylla (F); tributary of Siberia Stream,

opposite Siberia Hut, WNW of Makarora, ca. 760 m,

Child H2989 as B. diplophylla (F); Mt. Aspiring Natl.

Park, Blue Valley Track, above Blue River just N of

confluence with Makaroa River, 430-480 m, Engel

21905. WESTLAND PROV: Near Fox Glacier, 1940,

Knight as B. rosea (F); Camp Creek, W of Alexander

Range, 320-825 m, Reif C29E, C289D (F); Paparoa

Range, ridge N of Sewell Peak, 890 m, Fife 5163 as B.

convexiuscula (F). WESTLAND/CANTERBURY

PROV. BOUNDARY: Arthur's Pass Natl. Park, Arthur's

Pass area, Upper Twin Creek Valley, 930 m, Engel

18525 (F). CANTERBURY PROV.: Arthur's Pass Natl.

Park, Bealey River, off Bealey Valley Track, 830-850

m, Engel 18460 (F). NELSON PROV: Upper reaches of

Porarari River, ca. 2 km W of the "Lone Hand," 135-

185 m, Fife 6508 (F); lower basin of the Porarari River,

30-45 m, Fife 6673 — c. 6 (F); Paparoa Range, N side

of Tiropahi or Four Mile River, W side of Route 6, ca.

500 m N of bridge over river, S of Charleston, 145 m,

Engel 19274 (F). NORTH IS., SOUTH AUCKLAND

PROV: Mt. Te Aroha, ca. 3 km E of Te Aroha, 900-

940 m, Engel 22125— c. mar.

Balantiopsis lingulata Schust. Figure 2:12, 13.

Balantiopsis lingulata Schust., Bull. Natl. Sci. Mus.

Tokyo 11: 26. 1968. Holotype: New Zealand,

14

FIELDIANA: BOTANY

North Is., Waipoua Kauri Reserve, Waikohatu

Stream, near NS road, Schuster 67-1764 (F!).

Plants caespitose, ascending to stiffly erect,

opaque, dull green, occasionally tinged with red;

shoots to 4.8 mm wide. Branching variable, at

times predominantly terminal, sometimes pre-

dominantly ventral-intercalary. Leaves soft-tex-

tured, loosely imbricate, rather sharply folded, the

ventral lobes widely spreading, to 2.2 mm long,

the opposing dorsal lobes slightly to moderately

overlapping (excluding cilia) or not. Ventral lobe

weakly convex (more strongly so toward base),

the insertion distinctly J-shaped, the margin de-

current on the stem, the lobes distinctly Ungulate,

very shallowly to rather distinctly bifid; segments

at times barely differentiated from marginal ar-

mature, acuminate to ± ciliiform, entire or with a

cilium on lateral margin, 5-6 cells wide at base,

terminating in a uniseriate row of up to 4-5 rather

elongate cells (to 2.5:1) with somewhat thickened

walls, the septa slightly thickened in the corners,

not projecting, the sinus at times hardly differen-

tiated; lamina margins slightly curved, armature

variable: 14-20(26) cilia in optimally developed

plants, often fewer, the lobes typically ciliate to

the base, the ventral margin often sharply deflexed

toward base and with a sector of the dorsal sur-

face of lobe visible in ventral view, the margin

with up to 6-14 cilia, the dorsal margin typically

with fewer cilia; marginal cilia slender, similar to

those of apical segments but the cells of uniseriate

row somewhat longer (2.3-4.4:1); no border. Dor-

sal lobe much smaller than ventral lobe, typically

longer than wide and to 2X the width of stem

(excluding armature), plane, the insertion trans-

verse to ± oblique, weakly decurrent on the stem,

the lobes ovate to oblong-ovate, typically gradu-

ally narrowing toward the apex, shallowly bifid or

at times imperceptibly so, the segments as in ven-

tral lobe except for somewhat longer cells of uni-

seriate row; lobe margins curved, armed with up

to 10-16 widely divergent cilia. Underleaves sub-

equal to the dorsal leaf lobes or somewhat smaller,

contiguous to weakly imbricate, appressed to fee-

bly spreading, plane, the insertion broadly invert-

ed U-shaped, the underleaves narrowly ovate, bi-

fid to 0.5; lobes long attenuate, the margins with

several cilia, the sinus V-shaped to narrowly

rounded, not reflexed; lamina margins ciliate to

the base, often also with a lobuliform process on

each side, the margins decurrent on the stem; cilia

often bifurcate, with a uniseriate row of 4-5(6)

elongate, thin-walled cells. Ventral lobe cells in

evident transverse rows, subisodiametric and ±

hexagonal in the extreme apical portion to elon-

gate-rectangular in median and basal portions (to

ca. 3.8:1 in median sector), evenly thin-walled,

the median cells 20-28 u-m wide X 52-70 u,m

long; median cells of dorsal lobe similar to those

of ventral lobe; cuticle finely striolate-papillose.

Oil-bodies scattered throughout cell, pale, opaque,

smoky gray, (4)5-1 1(14) per cell, mostly ovoid to

elliptic to fusiform, some spherical, finely to

coarsely granular (the surface appearing papil-

lose), most 11.8-14.7 X 4.9-6.9 u,m, some 9.8-

10.8 X 6.9 (Jim, spherical ones 5.9 jim in diameter.

Androecia not seen. Marsupia rose to magenta,

covered with rhizoids.

TAXONOMY — The concept of B. lingulata adopt-

ed here considerably broadens the circumscription

of the species, previously known only from the

type collection. Ciliate forms of this species su-

perficially resemble B. rosea; see comments under

that species. When optimally developed, B. lin-

gulata has up to 26 cilia on the ventral lobe mar-

gins, and this will usually distinguish it from re-

lated species, B. montana and B. diplophylla. The

type collection of B. lingulata consists of gynoe-

cia-bearing shoots that are consequently more co-

piously ciliate than in many collections of this

species, particularly when sterile. Leafy hepatics

with armed leaves are frequently more strongly

armed on 9 shoots or in the vicinity of gynoecia.

Schuster (1968) distinguished this species from

B. rosea, but these species are quite unrelated; B.

lingulata can be readily distinguished by the stri-

ate-papillose leaf cuticle and the absence of a dif-

ferentiated border. Moreover, B. lingulata (in

common with B. diplophylla and B. montana) has

5-13 oil-bodies per cell (Fig. 2: 12, 13), whereas

B. rosea has only 2 (rarely three oil-bodies at op-

posite ends of the cell (Fig. 2:15).

Suboptimal, less ciliate populations are likely

to be confused with B. diplophylla, but the tiered,

elongate-rectangular ventral lobe cells and the

narrower, more Ungulate outline of the ventral

lobe of B. lingulata will aid in distinguishing the

two species.

DISTRIBUTION AND ECOLOGY — Known from a

few sporadic, lower-elevation sites (sea level to

90 m) on South Island and the northern sector of

North Island, where it is rather frequent at ele-

vations mostly under ca. 500 m. The species oc-

curs over rock in streambeds and, like so many

other members of the genus, is also found on the

banks of streams as well as more mesic niches

ENGEL & MERRILL: AUSTRAL HEPATICAE. BALANTIOPSIS

15

such as protected, bryophyte-covered banks and

shaded boulder faces. A population strikingly

similar to the type occurred loosely attached to

the lip of a springy area at the juncture of a road-

side bank and the forest edge in a deeply shaded

niche under Blechnum (Engel 21834). The type is

from a moist Dicksonia-kauri forest (in shade un-

der Cyathea), on smooth boulders or rocks along

the edge of a shallow, clear stream, associated

with Schistochila nitidissima. In shaded niches,

such as those of the Engel collection and the type,

the species does not develop pigmentation.

SELECTED SPECIMENS SEEN— NEW ZEALAND, SOUTH

IS., OTAGO PROV.: Fiordland, near McKerrow Hut,

head of Lake McKerrow, Hatcher 1468 (F). WEST-

LAND PROV.: Cascade Road, just NE of Martyr Saddle

and immediately N of Jackson River, S of Jackson Bay,

ca. 20 m, Engel 21834 (F). NELSON PROV: Track to

German Terrace, 6 km SSE of Westport on Nine Mile

Road, 90 m, Engel 21549 (F). NORTH IS., NORTH

AUCKLAND PROV: Mangatangi Stream, Huana,

Auckland, Child H808 as B. convexiuscula (F); NE Wai-

takere Range, Swanson University Reserve, Tram Valley

Road, 95 m, Engel 20444 (F); Waitakere Hills, Hatcher

153 (F); Trounson Kauri Forest, ca. 120 m, Child H21 11

(F); Waipoua State Forest, ca. 300 m, Child H2124 as

B. rosea (F); ibid., Hatcher 334 — c. mar. (F); ibid., Al-

lison s.n. as B. rosea (F); southern edge of Waipoua For-

est, along Marl borough Road, Engel 21139 (F); SE cor-

ner of Waipoua Forest, just N of Tutamoe, 540 m, Engel

21114 (F); Omahuta Forest Kauri Sanctuary, E of Man-

gamuka Bridge, 260 m, Engel 21014 (F); Mt. Te Aroha,

ca. 3 km E of Te Aroha, 900-940 m, Engel 22137 (F).

Balantiopsls sect. Pteridophylla (Hatch.) Engel &

Mem, comb. nov.

Isotachis sect. Pteridophylla Hatch., Nova Hedwigia

3: 8. 1961; p. 44 of separate. Type: Isotachis

splendens Steph.

Balantiopsis sect. Erinacea Engel, Fieldiana (Bot.)

25: 156. 1990, syn. nov. Type: Jungermannia er-

inacea Hook. f. & Tayl.

For a discussion of the section and its distin-

guishing characteristics, see p. 53.

Balantiopsis verrucosa Engel & Merr., sp. nov.

Figures 2:5,6, 9; 6.

Plantae virides vel brunneae vel rubescentes;

folia lobis dorsalibus transversim insertis, ca. 5

laciniato-lobulatis, ventralibus saepe canaliculatis

vel conduplicatis, ca. 2-9 dentato-ciliatis, inser-

tione valde arcuate (littera 'IT simili); laminae fo-

liorum marginatae, cellulis marginalibus striato-

papillatis; amphigastria vix divisa atque sparsim

armata; cuticula guttulata; corpora oleosa 2(3), in

extremis oppositis cellularum.

Holotype: New Zealand, South Is., Southland

Prov., Fiordland Natl. Park, just SW of Mt.

Burns, S of South Branch of Borland Burn, W

of Monowai, 1010-1170 m, Engel 18678 (F);

isotype: (CHR).

Plants resembling B. tumida, typically procum-

bent to rather stiffly erect, loosely caespitose, light

green to brownish or tinged with red; shoots com-

planate to weakly convex dorsally, the underleav-

es forming a broad ventral keel, the shoots to 3.5

mm wide. Branching sparse, of Frullania type;

ventral-intercalary branches sporadic. Leaves

soft-textured, densely imbricate, not sharply fold-

ed, the ventral lobe obliquely spreading, to 2 mm

long, the opposing dorsal lobes broadly overlap-

ping and extending across the stem. Ventral lobe

convex to ± canaliculate, the insertion strongly

inverted U-shaped, the ventral margin narrowly

decurrent on the stem, the lobes narrowly to

broadly ovate, very shallowly bilobed to biden-

tate, the segments broadly acute, often apiculate,

entire, 11-23 cells wide at base (when bilobed),

terminating in a short apiculus of a single cell or

a uniseriate row of 2(3) at most slightly elongate

cells; lamina margins broadly curved, 2-9-ciliate-

dentate, the ventral margin 4-5-ciliate-dentate, the

armature terminating in a uniseriate row of up to

3 elongate cells, the dorsal margin often few-

toothed to almost entire; lamina between the cilia

± distinctly bordered by elongate cells. Dorsal

lobe at times subequal to the ventral lobe in size

and to 4.5X the width of stem (excluding arma-

ture), plane, the insertion oblique, strongly

curved, becoming longitudinal, the free margin ±

subcordate at base and rather broadly decurrent

on the stem, the lobes broadly ovate to suborbi-

cular, the segments as in ventral lobe; lobe mar-

gins broadly curved, sparingly laciniate-lobulate,

sometimes with lateral teeth approaching the seg-

ments in size and shape, the border cells as in

ventral lobe. Underleaves large, about equal to the

dorsal leaf lobe in size, densely imbricate, ap-

pressed, plane to weakly longitudinally plicate,

the insertion quite narrowly inverted U-shaped,

the underleaves broadly ovate to orbicular, bifid

to 0.2 (rarely more); lobes narrowly acute, the

margins with 1-2 cilia; lamina margins on each

side rather sparingly laciniate-lobulate and ciliate

to the base, the margins long-decurrent on the

stem; ultimate divisions of lobe and marginal ar-

16

FIELDIANA: BOTANY

mature sometimes reflexed and directed toward

the underleaf base, terminating in short, curved

cilia composed of a uniseriate row of 2-3(5) rath-

er short-elongate cells; underleaves bordered as in

leaves. Ventral lobe cells in somewhat irregular

transverse rows, elongate-rectangular (to 3.7:1 in

median sector), thin-walled or the longitudinal

walls moderately thickened but thinning at the an-

gles, the median cells 16-24 |xm wide X (30)40-

60 (Jim long to 20-24 |xm wide X 58-84 jxm long;

median cells of dorsal lobe similar to those of

ventral lobe; cuticle of all but the marginal row

of cells coarsely (often grossly) papillose, the pa-

pillae large, high and rounded, the marginal row

of cells finely striate-papillose, sometimes lighter

in color, forming a distinct border. Oil-bodies at

opposing ends of cell, pale brown, 2 (rarely 3) per

cell, fusiform to crescentic, coarsely granular, 1 1-

13.4 X 5.3-5.8 jjim.

Androecia intercalary on main shoots; bracts

similar to leaves except more imbricate, slightly

pigmented and with 1-2 abaxial bulges; antheridia

2 per bract, the stalk biseriate; paraphyllia lacking.

Gynoecia not seen.

TAXONOMY — This species superficially resem-

bles B. tumida in size and habit and in the broadly

ovate to suborbicular dorsal lobes, the often lon-

gitudinally folded ventral lobes, and reduced leaf

armature. It can be readily distinguished, however,

by the larger, coarsely papillose cells, the presence

of a border, and the oil-body condition, which

mark it as a sparsely ciliate relative of B. rosea.

The specific epithet refers to the strikingly ver-

rucose (guttulate) cuticle of the leaves (Fig. 6 : 5).

When seen in profile (e.g., along a fold in the

leaf), the papillae are high and rounded at the

summit; in surface view they are rounded, con-

trasting with the finely striate-papillose cuticle of

the marginal row of lobe cells (Fig. 6 : 6).

DISTRIBUTION AND ECOLOGY — So far known

only from South Island, New Zealand, in upper-

elevation forests as well as subalpine and alpine

zones. For example, it occurs under tussock cover

in subalpine areas of Chionochloa, Hebe, Dra-

cophyllum, etc.

SELECTED SPECIMENS SEEN— NEW ZEALAND, SOUTH

IS., SOUTHLAND PROV.: Fiordland Natl. Park, Burns

Creek, below Mt. Burns, ca. 915 m, Child 5179 (F);

ibid., head of Gertrude Valley, track to Gertrude Saddle,

NE of Homer Tunnel, 1940-1970 m, Engel 21980 (F).

OTAGO PROV.: Mt. Cargill, N of Dunedin, 700 m,

Child H2674 as B. tumida (F); Swampy Hill, Dunedin,

610m, Child H2645 (F); Fiordland, near McKerrow Hut,

head of Lake McKerrow, Hatcher 1462 (F). WEST-

LAND PROV: Paparoa Range, ridge immediately N of

Sewell Peak, 890 m, Engel 19023 (F). WEST-

LAND/CANTERBURY PROV. BOUNDARY: Arthur's

Pass, 915 m, Child 2099 as B. tumida (F). NELSON

PROV: Paparoa Range, N flank of Mt. Euclid, ca. 0.5

km SE of Morgan Tarn, 1000-1065 m, Fife 6449 as B.

aequiloba (F). MARLBOROUGH PROV: Mt. Rich-

mond, Richmond Range, ca. 1370 m, Child H4304 (F).

Balantiopsis rosea Berggr. Figures 2:10, 11,

15; 7.

Balantiopsis rosea Berggr., On New Zealand Hep. 1:

43, / 28. 1898. Lectotype (nov.)\ New Zealand,

South Is., Westland Prov., Teremakau, Mar.

1874, Berggren 3191 (LD!).

Plants with a hairy, feltlike appearance, pro-

cumbent, loosely caespitose, light green, but typ-

ically rose-tinted to clear wine-red; shoots com-

planate to weakly convex dorsally, the ventral

lobes stiffly but moderately deflexed, the under-

leaves at times forming a ventral keel; shoots of-

ten robust, to 3 mm wide. Branching sparse, of

Frullania type; ventral-intercalary branches not

seen. Leaves soft-textured and delicate, densely

imbricate, not sharply folded, the ventral lobes

obliquely spreading, typically longitudinally fold-

ed and "nested" in the ventral lobe of the leaf

immediately above, to 2.5 mm long, the free mar-

gin of dorsal lobe (excluding cilia) ± aligned

along stem midline, the opposing dorsal lobes

touching to somewhat overlapping. Ventral lobe

typically longitudinally folded and distinctly con-

duplicate, the ventral (free) margin narrowly and

rather sharply reflexed-erect (the lobe thus pli-

cate), the insertion strongly inverted U-shaped,

the ventral margin narrowly long-decurrent on the

stem, the lobes narrowly elliptical, bifid to 0.2;

segments laterally armed with 2-3 cilia (often in

pairs), 8-15 cells wide at base, the segments ter-

minating in a uniseriate row of 2-3(4) elongate

cells; lamina margins rather straight, copiously

ciliate to the base; cilia up to 20, setaceous, each

terminating in a uniseriate row of 3-4 elongate

cells, the septa thickened in the corners and weak-

ly projecting; lamina between the cilia ± distinct-

ly bordered by elongate cells. Dorsal lobe ca.

0.5 X the ventral lobe in length and 2-2.5 X the

width of stem (exclusive of cilia), plane to weakly

longitudinally plicate, the insertion oblique,

strongly curved, becoming longitudinal, the lobe

± subcordate at base and rather broadly decurrent

on the stem, the lobes narrowly ovate to triangu-

lar, the segments as in ventral lobe; margins of

ENGEL & MERRILL: AUSTRAL HEPATICAE. BALANTIOPSIS

17

FIG. 6. Balantiopsis verrucosa Engel & Merr. 1, Sector of main shoot, dorsal view. 2, Leaves. 3, Leaf with lobes

spread, dorsal lobe at left. 4, Distal portion of ventral lobe of leaf. 5, Median cells of ventral lobe showing cuticular

papillae. 6, Sector of ventral margin of ventral lobe showing border. 7, Underleaves. 8, Antheridium. (Figs. 1-7 from

holotype; 8 from Child 2674, New Zealand, South Is., Otago Prov., Mt. Cargill.)

18

FIELDIANA: BOTANY

FIG. 7. Balantiopsis rosea Berggr. 1, Sector of main shoot, dorsal view. 2, Leaves. 3, Dorsal lobe. 4, Distal portion

of dorsal lobes. 5, Median cells of ventral lobe showing (in part) cuticular papillae. 6, Distal portion of segment of

ventral lobe of leaf. 7, Cells of ventral lobe segment base showing cuticular papillae. 8, Sector of ventral margin of

ventral lobe showing border. 9, Sector of ventral margin of ventral lobe. 10, Sector of main shoot, ventral view (note

details of ventral lobe insertion indicated for basalmost leaf pair). 11, Underleaves. (All from lectotype.)

ENGEL & MERRILL: AUSTRAL HEPATICAE. BALANTIOPSIS

19

lobe ± curved, ciliate to base with up to 10 cilia,

the cilia and border cells as in ventral lobe. Un-

derleaves about 0.75 the leaves in length, densely

imbricate, appressed to spreading, strongly con-

cave-channeled at base (ventral view), plane to

longitudinally plicate above, the insertion distinct-

ly narrowly inverted U-shaped, the underleaves

narrowly ovate, bifid to 0.4; lobes narrowly atten-

uate, at times ventrally sulcate, copiously armed

with cilia that are often in pairs; lamina margins

on each side sometimes with a lobuliform process

approaching the main lobes in size, the margins

otherwise copiously ciliate to the base, the mar-

gins dilated near base and long-decurrent on the

stem; ultimate divisions of lobe and marginal ar-

mature terminating in subcapillary cilia composed

of a uniseriate row of 4-5 elongate (5-10:1) cells;

underleaves bordered as in leaves. Ventral lobe

cells in somewhat irregular transverse rows, elon-

gate-rectangular (to 4.4: 1 in median sector), thin-

walled or the longitudinal walls moderately thick-

walled, but thinning at the angles, the median

cells 18-29 |j.m wide X 65-86 jim long; median

cells of dorsal lobe similar to those of ventral

lobe; cuticle of all but the marginal row of cells

coarsely (often grossly) papillose, the papillae

large, high, and somewhat elongate, the marginal

row of cells finely striate-papillose, sometimes

lighter in color, forming a distinct border. Oil-bod-

ies at opposing ends of cell, smoky gray or pale

yellowish brown, 2(3) per cell, irregular in shape:

subcrescentic to bottle-shaped to broad elliptic to

fusiform, finely granular, the surface appearing

finely papillose, 10.8-15.7 x 4.9-5.9 |xm, a few

8.8 X 3.9 |xm.

Androecia not seen. Marsupia long-cylindrical,

with abundant rhizoids. Sporophyte not seen.

TAXONOMY — This handsome species is remark-

able for the densely imbricate, profusely ciliate

leaves and underleaves, which lend the plant a

hairy, feltlike appearance. According to Berggren

(1898, p. 43), "the ventral surface of the plant is

quite like a felt from the close-set cilia." The ven-

tral leaf lobes are typically plicate and strongly

conduplicate-folded (Fig. 2: 10) and nested into the

ventral lobe next above on each side (Fig. 7:10);

the ventral lobe insertion is distinctly U-shaped

(Fig. 2:10). As in the related B. verrucosa, the

cuticle is coarsely papillose with high, rounded

papillae (Fig. 7:7), and the lobes have a differ-

entiated border of elongate, striate-papillose cells

(Fig. 7:6).

Suboptimal expressions of B. rosea (e.g., Engel

21712) often have much less copiously ciliate

lobe margins, with the ventral lobe insertion not

or only weakly recurved, and lack the character-

istic rose-red pigmentation. Plants of this facies

superficially resemble B. montana but can be dis-

tinguished by the guttulate papillae, the differen-

tiated border, and in fresh material, by the 2(3)

oil-bodies per cell, one in each end of the cell.

Balantiopsis rosea is most apt to be confused

with B. lingulata, but the latter differs from B.

rosea in the striate-papillose cuticle, the absence

of a border and the at best moderately convex

ventral lobe, which is J-shaped at the ventral end;

and in the greater number of oil-bodies per cell.

NOMENCLATURE — The lectotype has been cho-

sen on the basis of the original description and

illustration of the species (Berggren, 1898, pp.

43-44), despite the data on the labels of the Berg-

gren specimens at LD, which are contradictory. A

confusion of labels is the most likely explanation.

The leaves and underleaves of B. rosea are de-

scribed as abundantly ciliate, and the plants as

rose-purple in color, growing together with Iso-

tachis lyallii. However, the two specimens labeled

as being from the type locality (Bealey River,

Berggren 3190 and 3683, February 1874) lack

any trace of secondary pigmentation, the leaves

are only moderately ciliate, and no Isotachis is

present. Both specimens are B. lingulata. A third

specimen, labeled as from Teremakau (Berggren

3191, March 1874), contains the Isotachis, is

rose-pigmented and copiously ciliate, and has

been selected as the lectotype of the species. Un-

fortunately, none of the above specimens bear

marsupia, which are described and illustrated by

Berggren.

In his discussion of B. rosea, Berggren (1898,

p. 44) made a puzzling observation: "A peculi-

arity which I did not notice in the other species

... in so abundant a degree is the minute, round-

ed, hyaline granules in the cellules." At first, we

took this to be a reference to oil-bodies, but since

B. rosea typically has only 2 (rarely 3) oil-bodies

per cell, this cannot be the case. Instead, we think

that what Berggren saw were the abundant high,

rounded papillae characteristic of this species (and

B. verrucosa) and that are strikingly evident, even

at moderate magnification.

DISTRIBUTION AND ECOLOGY — Endemic to New

Zealand, where it occurs in middle- to upper-ele-

vation forests as well as the subalpine zone. In

forests the species may occur on well-shaded,

moist cliff faces, where it forms large, pure mass-

es that are loosely attached and may become pen-

20

FIELDIANA: BOTANY

dant from the substrate. It also is present in pro-

tected hollows on slopes or at the bases of banks,

where it is loosely, barely attached to the sub-

strate. Such niches are quite similar to those oc-

cupied by B. montana, and both species seem to

require sheltered, hyperhumid, well-shaded sites;

under such conditions neither species develops

red pigmentation. Along the Mangawhero River,

(Tongariro National Park), B. rosea occurs in hy-

perhumid sites along with Herzogianthus on

steep, moist, peaty, mossy banks in a low Noth-

ofagus solandri var. cliffortioides forest (Engel

21290). On wet roadside banks within forests B.

rosea is frequently associated with Isotachis lyal-

lii. In subalpine areas it occurs on the sides of

rills, etc.

SELECTED SPECIMENS SEEN— NEW ZEALAND, SOUTH

IS., SOUTHLAND PROV.: Fiordland, Wilmot Pass

Road to Deep Cove, ca. 300 m, Child H5612 (F). OTA-

GO PROV.: Mt. Aspiring Natl. Park, Blue Valley Track,

above Blue River just N of confluence with Makaroa

River, 430-480 m, Engel 21900 (F). WESTLAND

PROV: Haast Pass Road, near Gates of Haast, ca. 300

m, Allison s.n. (Child H850) (F); Westland Natl. Park,

track to Lake Gault, NE of Lake Matheson, NW of town

of Fox Glacier, ca. 100-200 m, Engel 21710 (F); ibid.,

track to Alex Knob, off track to Louisa Peak, 1 170 m,

Engel 18975 (F). CANTERBURY PROV: Mt. Cook

Natl. Park, SW facing cliffs off Wakefield Track, over-

looking Hooker River, E of town of Mt. Cook, 720 m,

Engel 18189. NORTH IS., WELLINGTON PROV:

Tongariro Natl. Park, west-facing steep bank of Man-

gawhero River, off Ohakune Mt. Road, ca. 1200 m, En-

gel 21290 (F). SOUTH AUCKLAND PROV: Omatika,

Kaimai Range, 855 m, Bartlett (F).

Balantiopsis convexiuscula Berggr. Figures 2:

3, 4, 14, 16; 8.

Balantiopsis convexiuscula Berggr., On New Zealand

Hep. 1: 44, / 29. 1898. Lectotype (fide Engel,

1968): New Zealand, North Is., Auckland Prov.,

Coromandel, Berggren 3188 (LD!); isolectotypes:

(FH!, GB!, UPS!).

Balantiopsis aequiloba Berggr., On New Zealand

Hep. 1: 46, / 30 H-O. 1898. Lectotype (fide En-

gel, 1968): New Zealand, North Is., Auckland

Prov., Coromandel, Berggren 3184 (LD!); isolec-

totype: (UPS!).

Plants rigid, procumbent, light green to olive-

green to yellowish brown, occasionally rose-tint-

ed, dull when dry; shoots dorsally convex, ± ven-

trally channeled, the ventral lobe apices strongly

decurved; shoots complanate, to 3(4) mm wide.

Branching sparse to common, pseudodichoto-

mous, of Frullania type; ventral-intercalary

branches common. Leaves appearing transversely

ribbed when dry, imbricate, rather sharply folded,

the ventral lobe widely spreading, to 1 .9 mm long,

the free margin of dorsal lobe ± straight, aligned

along the stem midline, the opposing dorsal lobes

touching to somewhat overlapping. Ventral lobe

moderately to strongly convex, the insertion mod-

erately to distinctly inverted U-shaped, the margin

decurrent on the stem, the lobes narrowly elliptic,

bifid to 0.2; segments acuminate, normally entire,

rarely with a tooth, (8)10-15 cells wide at base,

terminating in a uniseriate row of 3-4 cells, the

septa thickened in the corners and feebly project-

ing; lamina margins rather straight to gently

curved, sharply serrate to serrate-laciniate, the

ventral margin with 2-5 (8) stout teeth, each with

a uniseriate row of 3-4 cells, the dorsal margin

± straight in proximal sector, then arched distally,

sparingly armed with 1-4 slender teeth; no bor-

der; margins of ventral lobe and principal teeth

minutely denticulate by projecting septa of mar-

ginal cells. Dorsal lobe somewhat smaller than the

ventral lobe but more than half its length and typ-

ically broader, ca. 2X the stem in width, weakly

convex, the insertion transverse, not decurrent on

the stem, the lobes subquadrate to ± orbicular,

bifid to 0.25, the segments similar to those of ven-

tral lobe; dorsal (free) margin broadly curved, 4-

8(1 l)-toothed, at least some of the opposing teeth

approaching size of the segments; margins of dor-

sal lobe and marginal teeth denticulate as in ven-

tral lobe. Underleaves considerably smaller than

the leaves, contiguous to imbricate, slightly to

widely spreading, ± plane to broadly recurved,

the insertion broadly U-shaped, with the margins

decurrent on the stem, the underleaves narrowly

ovate to subrectangular in outline, appearing la-

ciniate-divided in situ but basically bifid to 0.5-

0.6; lobes parallel-sided to weakly attenuate, ±

plane, the ultimate lobe divisions and cilia curved,

biseriate for much of their length, terminating in

a uniseriate row of 4-6(7) elongate cells, the me-

dian sinus reflexed; lamina margins ciliate, some-

times on each side with a lobuliform process ap-

proaching the main lobes in size and shape. Ven-

tral lobe cells in ± regular transverse tiers, in dis-

tal sector small, short-rectangular (2: 1 ) or almost

isodiametric, the cell walls typically sinuous-

thickened, gradually becoming longer toward

base; median cells 12-15(17) jim wide X 26-40

|im long; median cells of dorsal lobe more uni-

formly short-rectangular; cuticle smooth to mi-

nutely striolate-papillose. Oil-bodies tending to

cluster at opposite ends of cell but sporadically to

commonly with 1 or more in central portion of

ENGEL & MERRILL: AUSTRAL HEPATICAE. BALANTIOPSIS

21

FIG. 8. Balantiopsis convexiuscula Berggr. 1, Portion of plant, dorsal view (Xll). 2, 3, Leaves. 4, Median cells

of ventral lobe. 5, Subapical cells of ventral lobe of leaf; note intermediate thickenings. 6, Segment of ventral lobe

of leaf. 7, Armature of ventral margin of ventral lobe of leaf. 8, Underleaves. 9, 9 Bract (X10). 10, 9 Bracteole

(XlO). (Fig. 1 from material admixed with type of B. diplophylla; 2-4, 6-8 from lectotype of B. convexiuscula; 5

from Hatcher 618, New Zealand, South Is., Otago Prov., trail between Gunn's Hut and Hidden Falls; 9, 10 from

Hatcher 1007, New Zealand, North Is., Pukeiti Bush near New Plymouth.)

22

FIELDIANA: BOTANY

cell, hyaline and glistening, (2)5-7 per cell, short-

to often long-elliptical to fusiform, some spheri-

cal, finely to coarsely granular, 8.8-12.7 X 3.9-

5.9 (xrn, those spherical or nearly so 4.9-5.9 X

3.9 (Jim.

Androecia not seen. Gynoecia on main shoot or

on ventral-intercalary branches of varying

lengths; bract margins very irregular, undulate to

distinctly crispate, occasionally reflexed, dentate-

laciniate. Marsupia green to rose, rhizoids color-

less or magenta.

Capsule long-cylindrical, with spiral dehis-

cence, the wall 31-34 u,m thick, of 3 layers, the

outer layer of cells ca. equal to thickness of both

interior strata; outer layer of cells short-rectan-

gular, large, the longitudinal and transverse walls

with thin, localized, sheetlike thickenings, without

nodular or bandlike thickenings; innermost layer

of cells with longitudinal radial walls with some-

what thick, continuous sheets of wall material,

with semiannular bands narrow.

Spores not seen. Elaters 11.5-13.4 u,m wide,

tapered to tips, bispiral, the spirals 3.8 (Jim wide,

loosely wound.

TAXONOMY — This species is a close ally of B.

tumida, the two sharing a suite of features, in-

cluding 1) dense areolation, the ventral lobe cells

only 12-16 (Jim wide; 2) swollen and projecting

septa, the lobe and segment margins consequently

minutely denticulate (Fig. 8 : 7, 8); 3) sinuous wall

thickenings in the cells of the distal sector of the

leaf lobes (Fig. 8 : 5); and 4) broadly orbicular

dorsal lobes, with at least one lateral tooth on each

side of the margin similar in size and shape to the

apical segments (Fig. 8 : 1-3). Balantiopsis con-

vexiuscula is distinguishable from B. tumida by

its coarsely serrate, often almost laciniate leaves,

the teeth often terminating in an elongate cilium.

The ventral lobes typically are convex and

obliquely spreading, whereas in B. tumida they

are often longitudinally folded and strongly ven-

trally secund. The dorsal lobes are narrower than

in B. tumida and with free margins only narrowly

overlapping versus broadly overlapping and ex-

tending beyond the opposite side of the stem in

B. tumida.

Balantiopsis convexiuscula has a characteristic

appearance: when dry, the plants are a dull, dark

olive-green, the dorsal lobes appearing densely

areolate and ribbed from arrangement of the cells

in regular, undulate transverse rows. It frequently

has been encountered in admixture with B. diplo-

phylla but is immediately distinguishable by its

much smaller cells, dense and regular areolation,

and color. Berggren (1898) describes the areola-

tion of B. aequiloba (a synonym of B. convexius-

cula) as the densest in the genus.

DISTRIBUTION AND ECOLOGY — Similar in distri-

bution to B. tumida, ranging from New Zealand

to Tasmania and southeast Australia, and com-

pared to that species, occurring at lower eleva-

tions and for the most part tolerating more mesic

sites. The species is rather common in lower- (sea

level) to middle-elevation forests, occurring on

moist, shaded (often clayey) soil, particularly of

mossy banks, etc., or over rock of dripping or

seepage areas of cliff faces. It frequents well-pro-

tected sites such as cavelike overhangs of cliffs,

under cover of vegetation (Blechnum,elc.\ and is

found less often on stream banks, boggy ground,

or rotted logs. At stream sites it occurs ascending

from often clayey soil in the splash zone or near

water level, where the plants are subject to inun-

dation. The species also occurs on rock of inter-

mittently dry creek beds. It occurs only sporadi-

cally in subalpine sites.

SELECTED SPECIMENS SEEN— AUCKLAND IS.: Dis-

appointment Is., Macmillan 83/159 (F). NEW ZEA-

LAND, SOUTH IS., SOUTHLAND PROV.: Fiordland

natl. Park, just SW of Mt. Burns, S of South Branch of

Borland Burn, Engel 77597; ibid., Stuart Mts., W shore

of Lake Thomson N of stream draining from Lake Wade,

300 m, Fife 7739 as B. diplophylla (F); W of Monowai,

1010-1 170 m, Engel 17597 (F). OTAGO PROV.: Saddle

W of road to Mt. Cargill, N of Dunedin, 500 m, Engel

17547 (F); Mt. Cargill, just below summit, N of Dune-

din, ca. 670 m, Engel 17568 (F); Flagstaff, NW of Dun-

edin, ca. 610 m, Child HI 663 (F); Fiordland, trail be-

tween Gunn's Hut and Hidden Falls, ca. 30 mi SE of

Lake McKerrow, Hatcher 618, 541 (F); ibid., near

McKerrow Hut, head of Lake McKerrow, Hatcher 1461

(F). WESTLAND PROV.: Haast, 6 mi N of bridge, sea

level, Child H871 (F); near Fox Glacier, 1940, Knight

(F); Westland Natl. Park, track to Lake Gault, NE of

Lake Matheson, NW of town of Fox Glacier, ca. 100-

200 m, Engel 21686 — c. mar. (F); near Gillespies Beach,

W of Fox Glacier, ca. 30 m, Child H4862 (F); Westland

Natl. Park, near Alex Knob track, Engel 6702 (F; ibid.,

Lake Wombat, Engel 6695 (F); Mt. Hercules Scenic Re-

serve, near Hercules Creek on Route 6, Engel 6563 (F);

Arthur's Pass Natl. Park, N of Kellys Creek near High-

way 73, above campground, N of Otira, 420-475 m,

Engel 18346 (F); Camp Creek, W of Alexander Range,

630-880 m, Re if C206E, C278b (F); Paparoa Range,

Croesus Track, ridge between Granite Creek and Pagan

Creek watersheds, 450 m, Engel 19294 (F). CANTER-

BURY PROV.: Arthur's Pass Natl. Park, Bealey River,

off Bealey Valley Track, 830-850 m, Engel 18464 (F).

NELSON PROV: Madman's Creek, ca. 2 mi S of Route

6 bridge over Little Totara River, Fife 4647 (F). NORTH

IS., TARANAKI PROV: Pukeiti Bush, near New Plym-

outh, Hatcher 283 (F). SOUTH AUCKLAND PROV:

ENGEL & MERRILL: AUSTRAL HEPATICAE. BALANTIOPSIS

23

Mt. Te Aroha, ca. 3 km E of Te Aroha, 900-940 m,

Engel 22115 (F). NORTH AUCKLAND PROV.: Huia,

SW of Auckland, ca. 30 m, Linzey (Child 3031) as B.

diplophvlla (F); Omahuta Forest Kauri Sanctuary, E of

Mangamuka Bridge, 260 m, Engel 20986, 21030 (F).

Balantiopsis tumida Berggr. Figure 9.

Balantiopsis tumida Berggr., On New Zealand Hep.

1: 45, / 30A-G. 1898. Holotype: New Zealand,

South Is., "ad flumen Teremakau in montibus

prov. Canterbury," Berggren 3186 (LD!); iso-

types: (FH!, OB!).

Plants ascending to suberect, caespitose, often

submerged, pure green to brownish green, often

rose-tinted to deep wine-red in exposed sectors of

leaves; shoots moderately dorsally convex, to bi-

laterally compressed, with the ventral lobes ven-

trally secund; shoots often robust, to 4.5 mm

wide. Branching sporadic, of Frullania type; ven-

tral-intercalary branches occasional. Leaves soft-

textured, closely imbricate, sharply folded, the

ventral lobes obliquely spreading to rather stiffly

ventrally secund, to 2.8 mm long, the opposing

dorsal lobes broadly overlapping and extending a

little beyond the opposite side of stem. Ventral

lobe strongly convex, the insertion strongly in-

verted U-shaped, the free margin decurrent on the

stem, the lobes typically longitudinally folded and

distinctly conduplicate (at times merely canalicu-

late), elliptical, bifid to 0.2, the segments bluntly

obtuse grading to medium to broad acute, often

terminating in a short apiculus of 1-2 cells, entire,

18-28 cells wide at base; lamina margins ±

straight to gently curved, sparingly armed, the

ventral margin narrowly reflexed-erect, with 1-5

rather slender ciliiform teeth ending in a uniseriate

row of 3-5 cells, the dorsal margin with fewer

teeth or none; no border; margins of ventral lobe

and principal teeth indistinctly denticulate by pro-

jecting septa of marginal cells. Dorsal lobe almost

equal to the ventral lobe in length but much

broader, to 3.5X the width of stem, weakly con-

vex to longitudinally 2-3 plicate, the insertion

somewhat curved, not or slightly decurrent on the

stem, the lobes subquadrate to ± orbicular, broad-

ly dentate, bifid to 0.2, the segments often stron-

ger than those of ventral lobe, entire, the inter-

vening sinus typically lunate; margins of the lam-

ina broadly and ± symmetrically curved to nearly

hemispherical, few-toothed, with at least 1 lateral

tooth on each side similar to the segments; mar-

gins of dorsal lobe and principal teeth denticulate

as in ventral lobe. Underleaves large, equal to the

dorsal leaf lobes in size but often inconspicuous,

hidden among the deflexed ventral leaf lobes, im-

bricate, appressed to spreading, the insertion

broadly U-shaped, with margins decurrent on the

stem, the underleaves broadly ovate to suborbi-

cular in outline, often longitudinally plicate, bifid

to 0.4; lobes attenuate, ± plane, the ultimate lobe

divisions and cilia terminating in a uniseriate row

of 4-5 elongate cells; margins of the lamina rather

sparingly ciliate, rarely with lateral processes ap-

proaching the lobes in size. Ventral lobe cells in

± regular transverse tiers, isodiametric to short-

rectangular (2:1), thin-walled but firm, indistinctly

sinuous-thickened, gradually becoming longer to-

ward base; median cells 12-16(18) |xm wide X

30-42 u,m long; median cells of dorsal lobe

somewhat smaller, more uniformly short-rectan-

gular; cuticle smooth to minutely striolate-papil-

lose. Oil-bodies confined to opposite ends of cell

or at times scattered within cell, hyaline and often

glistening, 2-5(7) per cell, ovoid to elliptical to

fusiform to ± linear, occasionally subcrescentic to

spherical, granular, the spherules somewhat pro-

truding beyond membrane, 5.9-10.8 X 2.9-3.9

u,m, ± linear ones 9.8 X 2.9 u,m, spherical ones

2.9-3.9 UJTI.

Androecia terminal on main shoot or relatively

long ventral-intercalary branches; bracts similar to

leaves or, occasionally, slightly smaller, occasion-

ally with a slight abaxial bulge; antheridia 1-2 per

bract. Gynoecia not seen.

TAXONOMY — When well-developed, B. tumida

is a striking plant, growing ± erect in thick, often

deep wine-red cushions on stream banks or sub-

merged to emergent in flowing water. The apical

segments of the dorsal and ventral lobes are

broadly acute to bluntly obtuse, terminating in a

short apiculus of 1-2 cells (Fig. 9:1, 2, 5). The

ventral leaf lobes are longitudinally folded to dis-

tinctly conduplicate and rather stiffly ventrally se-

cund; the underleaves are large, often longitudi-

nally plicate, and hidden among the deflexed ven-

tral lobes.

Berggren (1898) describes B. tumida as fastig-

iately branched, but most of the robust, caespitose

forms we have seen are only sparsely and sporad-

ically branched. This species is a close relative of

B. convexiuscula (for further discussion see under

that species).

DISTRIBUTION AND ECOLOGY — Known from

New Zealand (sporadic in South and North Is-

lands), Tasmania, and southeast Australia. The

species typically occurs in especially wet sites. In

the forest zone it may be found on silty soil and

24

FIELDIANA: BOTANY

FIG. 9. Balantiopsis tumida Berggr. 1, Portion of plant, dorsal view. 2, Leaves (3). 3, Leaf with lobes spread,

dorsal lobe at left. 4, Distal portion of dorsal lobe of leaf. 5, Distal portion of ventral lobe of leaf. 6, Median cells

of ventral lobe. 7, Underleaves (3). (All from type.)

ENGEL & MERRILL: AUSTRAL HEPATICAE. BALANTIOPSIS

25

rocks associated with rivulet banks, creek beds,

etc., where it may be subject to periodic inunda-

tion. At Waipoua Forest, for example, it occurs

with Schistochila nitidissima on the tops of soft,

conglomerate rocks and boulders in the bed of

Waikohatu Stream, which flows through a kauri

forest with Weinmannia silvicola (Engel 21039).

Also at Waipoua Forest it occurs in mucky areas

with scattered pools in a rich, damp forest domi-

nated by Weinmannia silvicola (Engel 21116). In

the subalpine zone the species typically occurs

over silty soil of rills or over soil between rocks

at rivulet margins, etc. In alpine areas it occurs

over soil at cliff bases, particularly under grass

cover.

SELECTED SPECIMENS SEEN— NEW ZEALAND, SOUTH

IS., SOUTHLAND PROV.: Fiordland Natl. Park, Burns

Creek headwaters, Mt. Burns, ca. 1370 m, Child H5145

(F); ibid., Stuart Mts., W shore of Lake Thomson N of

stream draining from Lake Wade, 300 m, Fife 7687 as

B. convexiuscula (F); ibid., head of Gertrude Valley, on

track to Gertrude Saddle, NE of Homer Tunnel, 1940-

1970 m, Engel 21988 (F). OTAGO PROV.: Summit of

Mt. Maungatua, W of Mosgiel, 825 m, Engel 17722 (F);

Mt. Cargill, N of Dunedin, ca. 455 m, Child H665 as B.

convexiuscula (F); Fiordland, trail between Gunn's Hut

and Hidden Falls, ca. 30 mi SE of Lake McKerrow,

Hatcher 663 (F). WESTLAND PROV: Near Clark's

Bluff, Haast Road, ca. 90-150 m, Child H4487, H4486

(F); Mt. Aspiring Natl. Park, Cross Creek, 1.1 km N of

Haast Pass, 540 m, Engel 21867 (F); Westland Natl.

Park, track to Lake Gault, NE of Lake Matheson, NW

of town of Fox Glacier, ca. 100-200 m, Engel 21681

(F); 8 mi W of Turiwhate, Route 73, Engel 6763 (F);

Camp Creek, W of Alexander Range, 770 m, Reif

C249A (F); Paparoa Range, ridge immediately N of Sew-

ell Peak, 890 m, Engel 19033 (F). WESTLAND/ CAN-

TERBURY PROV. BOUNDARY: Arthur's Pass Natl.

Park, Temple Basin ski area, Arthur's Pass, Engel 6503C

(F). NELSON PROV: Paparoa Range, N flank of Mt.

Euclid, below Morgan Tarn, 1065-1160 m, Fife 6319

(F); Abel Tasman Natl. Park, Porters Rock Track, ca. 915

m, Child H4657 (F). MARLBOROUGH PROV: Wai-

kakaho Valley, W of Tuamarina, ca. 300 m, Child H3273

(F). NORTH IS., NORTH AUCKLAND PROV: Wai-

poua Forest, Waikohatu Stream, on State Highway 2 just

upstream from culvert, 290 m, Engel 21039 (F); SE cor-

ner of Waipoua Forest, just N of Tutamoe, 540 m, Engel

21116 (F).

Systematics

Since its beginnings in the 1950s, phylogenetic

systematics, or cladistics, has become a widely

used procedure for reconstructing phylogenies of

groups of plants and animals and for constructing

classifications. Botanical cladistics was inaugurat-

ed by the work of Koponen (1968), a bryologist,

but until recently few bryologists have made use

of this approach to understanding phylogenetic re-

lationships. Initially, manual Hennigian methods

were employed (e.g., Mishler & Churchill, 1984;

Hyvonen, 1989), but more recently computer pro-

grams, such as PAUP and Hennig86, have been

used. Most of these recent studies have been con-

cerned with mosses (among them Mishler, 1985;

Bruggeman-Nannenga & Roos, 1990; Zander,

1993; Zomlefer, 1993; Hedenas, 1994; and Hy-

vonen & Enroth, 1994), and relatively few have

dealt with hepatics (Thiers, 1993; Hyvonen &

Piippo, 1994).

In the course of our study of the genus Bal-

antiopsis in New Zealand, we developed certain

hypotheses concerning the relationships between

species and the primitive and advanced states (po-

larity) of characters, including some apparent in-

stances of convergence and parallelism. A cladis-

tic analysis was undertaken in order to test these

hypotheses and to reconstruct the evolutionary

history of the genus Balantiopsis in the context

of the Balantiopsaceae as a whole.

The methods used in this analysis are not ref-

erenced extensively here. Manuals to computer

programs for phylogenetic analysis such as PAUP

(Swofford, 1991), MacClade (Maddison & Mad-

dison, 1992), and Hennig86 (Farris, 1988) provide

a good introduction to theoretical concepts and

contain thorough discussions of cladistic meth-

odology with extensive references. The basic

works on the subject are Hennig (1966) and Wiley

(1981); an excellent up-to-date introduction to

cladistic theory and practice is Forey et al. (1992).

Choice of Taxa

For this portion of our study, in addition to the

New Zealand species of Balantiopsis, we sur-

veyed and reevaluated all species of the genus.

(For detailed descriptions of the extraterritorial

species of Balantiopsis, see Engel, 1968.) Repre-

sentatives of all genera of Balantiopsaceae were

included in the analysis as outgroups (Watrous &

Wheeler, 1981; Maddison et al., 1984) in order to

resolve relationships within the genus Balantiop-

sis as well as to assess relationships between the

major subgroupings within the family (Table 1).

With the exception of Isotachis, these genera are

either monotypic (Anisotachis) or have relatively

few species (Ruizanthus with two, Eoisotachis

with two, Austroscyphus with four, Neesioscyphus

26

FIELDIANA: BOTANY

TABLE 1. Taxa included in systematic analysis.

Anisotachis splendens (Steph.) Schust.

Austroscyphus phoenicorhizus (Grolle) Schust.

Balantiopsis asymmetrica (Herz.) Engel

B. bisbifida (Steph.) Steph.

B. brasiliensis Steph.

B. cancellata (Nees) Steph.

B. ciliaris Halt.

B. convexiuscula Berggr.

B. crocea Herz.

B. diplophylla (Hook. f. & Tayl.) Mitt.

B. erinacea (Hook. f. & Tayl.) Mitt.

B. lingulata Schust.

B. montana (Col.) Engel & Merr.

B. philippinensis H. Inoue

B. purpurata Mitt.

B. rosea Berggr.

B. tumida Berggr.

B. verrucosa Engel & Merr.

Eoisotachis stephanii (Salm.) Schust.

Isotachis intortifolia (Hook. f. & Tayl.) Gott.

/. lyallii (Hook. f. & Tayl.) Mitt.

/. montana Col.

Neesioscyphus argillaceus (Nees) Grolle

N. carneus (Nees) Grolle

Ruizanthus venezuelanus Schust.

Triandrophyllum subtrifidum (Hook. f. & Tayl.) Fulf. &

Hatch.

with five). To represent Isotachis, we selected /.

intortifolia, I. lyallii, and /. montana, all New Zea-

land species (see Phytogeography, p. 56) but rep-

resentative of the range of variation in the genus

(for a discussion of monophyly of Isotachis, see

Intergeneric Relationships, p. 47). Ruizanthus ve-

nezuelanus was chosen to represent Ruizanthus

since sporophyte data were available for this spe-

cies (Schuster, 1985a). Similarly, of the four

known species of Austroscyphus, A. phoenicor-

hizus was selected as the placeholder for this ge-

nus since the most complete data (e.g., on oil-

bodies) were available for this species. Neesios-

cyphus is represented by N. argillaceus and N.

carneus (Grolle, 1964), species for which both ga-

metophytic and reproductive structures are

known.

We also chose the primitive, generalized liver-

wort Triandrophyllum subtrifidum as an outgroup

because it is clearly not a member of the Balan-

tiopsaceae but is sufficiently similar in gross mor-

phology to allow polarization of most of the char-

acters used in the analysis. Data were available

for the outgroup on all characters used in the anal-

ysis. To test the effect of additional outgroups on

the analysis, we experimented with inclusion of

two additional taxa, Lepicolea attenuata and Di-

plophyllum taxifolium (for a discussion of the ef-

fect of these taxa on the results of the analysis,

see Results, p. 43). Triandrophyllum and Lepico-

lea occupy a near-basal position relative to other

leafy hepatics in the classification system of

Schuster (1966, 1979); Diplophyllum has compli-

cately bilobed leaves similar to those of Balan-

tiopsis but has a simple perianth with no acces-

sory gynoecial structures and straight valvular

capsule dehiscence. The 26 taxa included in the

analysis are listed in Table 1.

With the exception of R. venezuelanus (Schus-

ter, 1985a), data used in the analysis were verified

by examination of herbarium specimens. Oil-bod-

ies are ephemeral cell inclusions and are soon lost

in drying; data for oil-bodies of many taxa were

necessarily obtained from literature reports. All

characters are treated as unordered to avoid mak-

ing a priori assumptions concerning character po-

larity, which may or may not have a sound phy-

logenetic basis. Over reliance on generalizations

concerning the relative primitiveness or ad-

vancedness of characters has been cited as an ob-

stacle to progress in understanding the evolution

of bryophytes (Mishler & Churchill, 1984, p.

421). Taxa with more than one state of a given

character were treated as polymorphic with re-

spect to that character and assigned both values;

in the analysis, PAUP (p. 41) was instructed to

treat these as instances of polymorphism rather

than uncertainty as to character state. The data

matrix showing the assignment of character states

for each taxon is shown in Table 2.

Character Descriptions and Coding

GAMETOPHYTE

General

1. Symmetry. Anisophylly is typical of most

leafy hepatics, but the Balantiopsaceae include

both anisophyllous and subisophyllous forms.

Pronounced anisophylly is commonly associated

with a plagiotropic habit (e.g., Isotachis minima,

with small, deeply bifid underleaves and abundant

rhizoids; Fig. 11). Unfortunately, isophylly is a

somewhat ambiguous term, since it may refer to

leaves and underleaves that are similar in size

(surface area) only. Strict isophylly, in which un-

derleaves are undistinguishable from leaves, is

rather uncommon in liverworts. The term sub-

isophylly is applied here to taxa in which the un-

ENGEL & MERRILL: AUSTRAL HEPATICAE. BALANTIOPSIS

27

TABLE 2. Data matrix used in cladistic analysis of Balantiopsis and related genera.

derleaves resemble leaves in both form and size

(e.g., Eoisotachis stephanii, hotachis intortifolia).

For example, Anisotachis splendens is scored as

anisophyllous because its deeply conduplicately

bilobed leaves differ strongly from underleaves in

form. In the similar size of the leaf lobes and un-

derleaves (as well as in its turgid, julaceous habit),

this species mimics Eoisotachis. This facies is

characteristic of species that grow erect and sub-

mersed or emergent in small streams or pools, etc.

All Balantiopsis species are considered aniso-

phyllous.

Symmetry is treated as a binary character: sub-

isophyllous (0), moderately to strongly aniso-

phyllous (1). This character could be scored in 25

of 25 (100%) of the ingroup taxa.

2. Pigmentation. Strong pigmentation is char-

acteristic of the family Balantiopsaceae. Balan-

tiopsis rosea and Isotachis lyallii are both notable

for the intense wine-red color that develops in ex-

posed sites. Anisotachis, Eoisotachis, Ruizanthus,

and /. intortifolia are also deeply red-pigmented.

Other species are characteristically brownish in

color and only sporadically reddish-tinged (e.g.,

Balantiopsis montana, B. verrucosa, Isotachis mon-

tana, Austroscyphus phoenicorhizus). The only

taxon not scored with respect to pigmentation was

B. philippinensis, known only from the type, in

which this character is impossible to assess with

certainty.

Pigmentation is treated as a binary character:

red to wine-red (0), brownish or tinged with red

(1). This character could be scored in 24 of 25

(96%) of the ingroup taxa.

3. Branching type. Types of branching modes

and their relative frequency are considered highly

significant in assessing phylogenetic relationships

among hepatics. Although of demonstrated taxo-

nomic utility, for purposes of phylogenetic anal-

ysis, we consider frequency of occurrence to be

less informative than evidence of the ability to

produce a given type of branching. Terminal

branching is lacking in Neesioscyphus (Grolle,

1964), Ruizanthus (Schuster, 1985a), and the out-

group taxon, T. subtrifidum. All other taxa (in-

cluding all Balantiopsis species) produce termi-

nal, Frullania-type, as well as ventral-intercalary,

28

FIELDIANA: BOTANY

branches. The only species not scored was Bal-

antiopsis philippinensis.

Branching type is treated as a binary character:

only intercalary branching present (0), branching

terminal and intercalary ( 1 ). This character could

be scored in 24 of 25 (96%) of the ingroup taxa.

4. Rhizoid origin. In Balantiopsis bisbifida

(Engel, 1968, p. 95, & fig. 65), Neesioscyphus

carneus (Grolle, 1964), and Ruizanthus venezue-

lanus (Schuster, 1985a, p. 73, & fig. 6:6), rhizoids

originate at the bases of the leaves as well as un-

derleaves. In all other taxa included in the anal-

ysis, rhizoids occur only at the bases of under-

leaves.

Rhizoid origin is treated as a binary character:

at bases of leaves and underleaves (0), only at

bases of underleaves (1). This character could be

scored in 25 of 25 (100%) of the ingroup taxa.

Leaves

5. Depth of division between lobes. The "typ-

ical" Balantiopsis leaf is strongly bilobed to at

least 0.6; this character corresponds to the width

of the "keel" in a conduplicate-bilobed leaf. In

B. ciliaris, the keel is only 1-2 cells wide; in B.

purpurata, the dorsal lobe sometimes appears dis-

placed and inserted on the lamina of the ventral

lobe. In B. erinacea, the dorsal lobe is detached

and either immediately juxtaposed to the ventral

or completely free. In Eoisotachis, Isotachis, Rui-

zanthus, and two Balantiopsis species (B. asym-

metrica and B. bisbifida), the leaf is ± regularly

and shallowly bilobed or bisbifid to no more than

0.5. In Neesioscyphus argillaceus and N. carneus

the apex is shallowly and unequally bilobed

(Grolle, 1964, figs. 2:1, 2; 3:3). In Austroscy-

phus phoenicorhizus the leaf apex is unlobed to

retuse (Fig. 10:3); however, in Austroscyphus

iwatsukii (Kitag.) Schust. (not included in the

analysis), the leaves are distinctly bilobed.

Division between lobes is treated as an unor-

dered multistate character with three states: bi-

lobed or bisbifid to 0.5 or less (0), deeply bilobed,

to 0.6 or more (1), dorsal lobe displaced or de-

tached (2). This character could be scored in 25

of 25 (100%) of the ingroup taxa.

6. Relative lobe size. The leaves of leafy he-

patics are fundamentally bilobed from the earliest

stages of development at the shoot apex (Evans,

1939, p. 61.) In nearly all of the taxa treated here,

the leaf is bilobed or bisbifid in varying degrees,

with distinct dorsal and ventral lobes, which differ

in relative size. In the outgroup taxon, Triandro-

phyllum subtrifidum, and in Isotachis, the dorsal-

most lobe commonly exceeds the ventral in size;

in Balantiopsis, as a rule, the dorsal is much

smaller than the ventral lobe. The extreme con-

dition is reached in B. purpurata (Engel, 1968,

fig. 80) and in B. diplophylla var. hockenii; in the

latter, the dorsal lobe may be a slender, deeply

bifurcate appendage of the ventral lobe. In Nee-

sioscyphus (N. argillaceus and N. carneus), the

dorsal lobe is much smaller than the ventral (Grol-

le, 1964, figs. 2:1, 2; 3:3). In Austroscyphus

phoenicorhizus, the leaves are unlobed to retuse;

A. iwatsukii, however, has deeply bilobed leaves.

Relative lobe size is treated as an unordered

multistate character with four states: dorsal lobe

larger than ventral (0), subequal to ventral (1), ca.

0.25-0.5 the ventral (2), 0.2 the ventral or less

(3). Austroscyphus phoenicorhizus, with unlobed

leaves, was scored inapplicable. This character

could be scored in 24 of 25 (96%) of the ingroup

taxa.

7. Leaf insertion. Leaf insertion is essentially

transverse in Isotachis intortifolia and /. montana

(Fig. 12: 1, 7) and weakly to distinctly incubous

in /. lyallii (Fig. 14: 1) as well as in the outgroup

taxon, T. subtrifidum. In all the remaining taxa,

the leaves are weakly to distinctly succubous in

insertion. Assessment of the leaf insertion in

many Balantiopsis species and in Anisotachis is

complicated by the recurvature of the insertion of

the dorsal and ventral lobes (Fig. 2: 1-8), each of

which is treated as a separate character (8 and 9).

Leaf insertion is treated as an unordered mul-

tistate character with three states: transverse (0),

succubous (1), incubous (2). Taxa with variable

insertion (e.g., with transverse to weakly succu-

bous insertion, as in Eoisotachis) were treated as

polymorphic with respect to this character and as-

signed both values. This character could be scored

in 25 of 25 (100%) of the ingroup taxa.

8. Ventral lobe insertion. Seen in ventral as-

pect, the ventral lobe insertion in some Balantiop-

sis species is strongly arched and abruptly re-

curved toward the base of the shoot so that the

line of insertion resembles an inverted U (B. con-

vexiuscula, Fig. 2 : 4; B. verrucosa, Fig. 2 : 6). The

ventral lobe insertion in Anisotachis splendens is

also strongly U-shaped (Schuster, 1972, fig. 5:6).

In other species of Balantiopsis the insertion is

moderately recurved (J-shaped, e.g., B. diplo-

ENGEL & MERRILL: AUSTRAL HEPATICAE. BALANTIOPSIS

29

FIG. 10. Austroscyphus phoenicorhizus (Grolle) Schust. 1, Ventral-lateral view of main shoot. 2, Lateral view of

main shoot with ventral-intercalary branch (vib). 3, Leaves and, to right, an underleaf. 4, Median leaf cells, showing

tiering of cells and intermediate thickenings. 5, Oil-bodies (X615). 6, Teeth of ventral margin of leaf. 7, Distal sector

of underleaf. 8, Antheridium. 9, 10, Portion of lobule of $ bract. 11, Apex of d bract. (Figs. 1, 2 from Engel 15916,

Tasmania, Great Western Tiers, vicinity of Devils Gullet Lookout; 3-7 from Engel 19903, Tasmania, Hartz Mts. Natl.

Park, summit area of Hartz Peak; 8-1 1 from Ratkowsky 78/73, Tasmania, Mt. Wedge.)

30

FIELDIANA: BOTANY

FIG. 1 1 . Isotachis minima Pears. 1, Leading shoot, ventral-lateral view. 2-4, Leaves. 5, Portion of leaf lobe,

showing cells in ill-defined tiers. 6, Leaf lobe apices. 7, Sector of main shoot showing poorly differentiated androe-

cium, dorsal view. 8, Antheridial stalk. 9, Largest inner 9 bracts and, in middle, bracteole; to right are two perigynial

bractlets at same scale. 10, Seta, cross section. (Figs. 1, 4 from Sept. 1939, Hodgson, New Zealand, North Is., South

Auckland Prov., Whakarewarewa Thermal Reserve; 2, 5, 6 from type of/, minima; 3 from Engel 17630, New Zealand,

South Is., Otago Prov, W slope of Flagstaff, NW of Dunedin; 7-9 from Child 47, New Zealand, North Is., South

Auckland Prov., Waiotapu, Rotorua; 10 from Child 1382, New Zealand, South Is., Otago Prov., Flagstaff.)

ENGEL & MERRILL: AUSTRAL HEPATICAE. BALANTIOPSIS

181 WWWLL HALL

MAR 0 4 1997

31

FIG. 12. Isotachis montana Col. 1, Sector of main shoot, the distal sector shown in dorsal aspect, the basal sector

in lateral aspect (1 = leaf; ul = underleaf)- 2-4, Leaves and underleaves (ul). 5, Distal sector of leaf lobe (cuticle not

shown). 6, Median-basal leaf cells. 7, Sector of main shoot with Frullania-type branch (FB; HL = half-leap, dorsal

view. (Figs. 1, 4-6, from type of /. montana; 2, from Hatcher 1429, new Zealand, South Is., Otago Prov., Fiordland,

near McKerrow Hut, head of Lake McKerrow; 3, 7 from Child 4859, New Zealand, South Is., Westland Prov., near

Gillespies Beach, W of Fox Glacier.)

32

FIELDIANA: BOTANY

phylla, Fig. 3 : 6), only weakly recurved (comma-

like, e.g., B. montana, Fig. 5 : 8), or straight.

Ventral lobe insertion is treated as an unordered

multistate character with three states: straight or

comma-like (0), J-shaped (1), U-shaped (2). This

character could be scored in 25 of 25 (100%) of

the ingroup taxa.

9. Dorsal lobe insertion. In some Balantiopsis

species the line of insertion of the dorsal half of

the leaf is an almost uninterrupted extension of

the insertion of the ventral half and is only slightly

bent at the junction between the two. Seen in dor-

sal aspect, the base of the dorsal lobe is strongly

oblique to longitudinal in orientation, with the

free margin long-decurrent on the stem (e.g., B.

rosea, Fig. 7:1; B. verrucosa, Fig. 2:5). In B. bis-

bifida and B. asymmetrica the line of leaf insertion

is essentially straight (Fig. 2:1, 2; Hassel de Me-

nendez & Solari, 1975, fig. 35:11, 12).

In other Balantiopsis species the insertion of

the dorsal lobe is transverse, at right angles to the

axis of the stem, with the free margin scarcely

decurrent (e.g., B. convexiuscula, Figs. 2:3; 8:1,

2). In these species, the insertion line appears

sharply bent at the junction between the dorsal

and ventral lobes (Fig. 2:4). In other Balantiopsis

species and in Anisotachis splendens, the dorsal

lobe insertion is strongly arched, and inverted

U-shaped, like the ventral (e.g., B. erinacea, Fig.

2:7, 8).

Dorsal lobe insertion is treated as an unordered

multistate character with three states: ± transverse

(0), oblique to longitudinal (1), U-shaped (2). This

character could be scored in 25 of 25 (100%) of

the ingroup taxa.

10. Ventral lobe/leaf planation. In B. rosea, B.

tumida, B. verrucosa, and Anisotachis splendens,

the ventral lobe of the leaf is often sharply cana-

liculate or conduplicately folded to strongly pli-

cate (Fig. 2:10, shown in section in Fig. 2:11).

However, in most Balantiopsis species, the ventral

lobe is merely convex, without any sharp folding

or plication (e.g., B. diplophylla, Fig. 3:1, 6). In

B. montana the ventral lobes are widely spread-

ing, horizontally oriented, and essentially flat (Fig.

5:1). In B. asymmetrica, B. bisbifida, Austroscy-

phus, Eoisotachis, Isotachis montana, I. lyallii,

and Ruizanthus, the leaf is moderately to strongly

concave; in Isotachis intortifolia the entire leaf is

often strongly plicate.

Ventral lobe/leaf planation is treated as an unor-

dered multistate character with three states: essen-

tially flat (0), convex or concave (1), canaliculate

to conduplicately folded or plicate (2). Taxa ex-

pressing varying degrees of plication or folding

were treated as polymorphic with respect to this

character and assigned both values. This character

could be scored in 25 of 25 (100%) of the ingroup

taxa.

11. Dorsal lobe orientation. The "typical"

Balantiopsis leaf is deeply conduplicate-bilobed,

with a ± distinct keel (e.g., B. rosea, Fig. 7:1). In

some species, however, the dorsal lobe is dorsally

assurgent (B. montana, Fig. 5:1). In B. asymme-

trica the dorsal lobe margin, seen in dorsal aspect,

is moderately inflexed to narrowly incurved and

erect (Engel, 1968, fig. 1; Hassel de Menendez &

Solari, 1975, fig. 35:1). In B. bisbifida the leaf is

bisbifid, and the leaves are ± symmetrical and

merely concave, as are the leaves of Ruizanthus,

Neesioscyphus, Eoisotachis, and Isotachis spp.

The complicately bilobed leaves of other hepatic

genera (e.g., Scapania, Diplophyllum) are regard-

ed as nonhomologous and independently derived.

Dorsal lobe orientation is treated as an unor-

dered multistate character with three states: leaves

concave, not conduplicately folded (0), keeled but

dorsal lobe assurgent (1), conduplicate-folded, the

dorsal lobe appressed to the ventral (2). This char-

acter could be scored in 25 of 25 (100%) of the

ingroup taxa.

12. Leaf armature. Development of teeth or

cilia on the margins of the leaves is highly vari-

able in Balantiopsis, although individual species

typically have a characteristic range of variation.

The extreme condition occurs in B. ciliaris, in

which the ventral lobe has up to 32 cilia (to 95

on both lobes, Hattori, 1966). Balantiopsis rosea

and B. lingulata have ventral lobe margins with

up to 14 to as many as 20 cilia. Other species,

such as B. convexiuscula, have the lobes moder-

ately armed, the ventral lobe with 2-9, or at most

13, teeth or cilia. Balantiopsis tumida and B. ver-

rucosa at times have almost unarmed margins

(Fig. 9:2, 3).

Leaf armature is treated as an unordered mul-

tistate character with three states: margins copi-

ously armed (0), moderately armed (1), almost

unarmed (2). Taxa with variable armature (i.e.,

moderately armed in some populations to almost

unarmed in others) were treated as polymorphic

with respect to this character and assigned both

values. This character could be scored in 25 of 25

(100%) of the ingroup taxa.

13. Projecting septa. In some Balantiopsis spe-

ENGEL & MERRILL: AUSTRAL HEPATICAE. BALANTIOPSIS

33

cies, the margins of the leaves, particularly in the

distal sector, are minutely denticulate by project-

ing septa of marginal cells. This character is pres-

ent in B. convexiuscula (Fig. 8:6, 7), B. tumida,

B. cancellata, and B. ciliaris (Piippo, 1984, fig.

16c). Neesioscyphus carneus was not scored.

Projecting septa are treated as a binary char-

acter: lobe margins without projecting septa (0),

with projecting septa (1). This character could be

scored in 24 of 25 (96%) of the ingroup taxa.

Underleaves

14. Underleaf lobing. The underleaf apex var-

ies from truncate-entire or more often retuse to

short-bilobed in Anisotachis splendens (Hassel de

Menendez & Solari, 1975, fig. 37:1, 4) to deeply

bifid and dissected to 0.5 or more (e.g., B. con-

vexiuscula, Fig. 8:8). Other taxa (e.g., B. verru-

cosa, Figs. 2:9; 6:7) have underleaves only shal-

lowly bifid, to 0.2-0.4 or less.

Underleaf lobing is treated as an unordered

multistate character with three states: truncate-en-

tire (0), shallowly bifid, 0.2-0.4 or less (1), deeply

bifid to 0.5 or more (2). This character could be

scored in 25 of 25 (100%) of the ingroup taxa.

Cells

15. Leaf areolation. Regular transverse tiering

of leaf and underleaf cells is characteristic of Bal-

antiopsis (Berggren, 1898, p. 44) and, indeed, is

"almost a trademark of the entire suborder"

(Schuster, 1985a, p. 72). This is often accentuated

by the elongate-rectangular to linear shape of the

cells. In many taxa with conspicuous tiering of

cells (e.g., B. rosea, Fig. 7:5), the cells are

grouped in conspicuous "panels" of 6-10 long,

narrow cells that often appear secondarily divid-

ed. In B. diplophylla (Fig. 3:4), B. asymmetrica,

B. bisbifida, and B. purpurata the leaf cells are ±

isodiametric and scattered, without any suggestion

of tiering. The leaf cells of Austroscyphus phoen-

icorhizus are also tiered (Fig. 10:4). In Ruizanthus

the underleaves display noticeable tiering, but the

leaves do not (Schuster, 1985a, fig. 8:15). Nee-

sioscyphus carneus was not scored.

Areolation is treated as an unordered multistate

character with three states: cells typically scat-

tered (0), cells indistinctly tiered (1), leaf cells in

evident transverse rows (2). Taxa with tiering only

sporadically expressed (i.e., present in some

leaves but not others, or only in certain sectors of

leaves) were treated as polymorphic with respect

to this character and assigned both vales. This

character could be scored in 24 of 25 (96%) of

the ingroup taxa.

16. Leaf border. Many Balantiopsaceae have

leaves with a ± distinct border formed by differ-

entiation of the marginal row of cells. In some

species (e.g., B. verrucosd), bordering also takes

the form of a striking difference in the cuticle: the

finely striate-papillose marginal cells are in sharp

contrast to the coarsely guttulate papillae of the

cells immediately adjacent to the marginal row

(Fig. 6:6). It should be noted that Ruizanthus lo-

pezii, which was not included in the analysis, has

a distinct border (Schuster, 1985a, fig. 8:11-13),

but of a different type.

Leaf border is treated as a binary character: no

border (0), marginal cells forming a ± distinct

border (1). Taxa with an indistinct border were

assigned both values. This character could be

scored in 25 of 25 (100%) of the ingroup taxa.

17. Leaf cell size. Some species of Balantiopsis

have relatively small cells, lending the leaves a

dense, subopaque appearance (e.g., B. cancellata,

B. convexiuscula}. Others have much larger cells,

to 40-50 (Jim wide, and lax areolation, with a ±

translucent quality. Comparison of cell width

ranges in different species indicated a disconti-

nuity at ca. 1 8 |xm, which was used as a basis for

scoring this character.

Leaf cell width is treated as a binary character:

12-18 |xm wide (0), 19-53 |xm wide (1). Cell

width measurements were taken from cells that

were not secondarily divided. For consistency,

measurements were made in the median sector of

the leaf, and in the case of deeply bilobed leaves,

from the median sector of the ventral lobe. This

character could be scored in 25 of 25 (100%) of

the ingroup taxa.

18. Wall thickenings. Cell walls in the taxa

studied may be uniformly thin- or thick-walled or

may exhibit characteristic thickenings. In many

species, the median portion of the longitudinal

walls is noticeably thickened, the walls thinning

abruptly to the corners of the cells (e.g., B. rosea,

Fig. 7:7). In species with short-rectangular cells,

the walls of the longitudinally aligned rows of

cells often appear sinuate, due to these unequal

wall thickenings (e.g., B. convexiuscula, Fig. 8:5;

Austroscyphus phoenicorhizus, Fig. 10:4). In Nee-

sioscyphus carneus, intermediate thickenings are

34

FIELDIANA: BOTANY

present, as well as knotlike trigones (Grolle, 1964,

fig. 2:3); in Ruizanthm venezuelanus (Schuster,

1985a), the median leaf cells are thin- walled but

with moderate trigones. Conspicuous, bulging tri-

gones are present in Isotachis intortifolia.

Wall thickening is treated as a binary character:

uniformly thin- or thick-walled (0), intermediate

thickenings (1). Taxa with only sporadic inter-

mediate wall thickenings were assigned both val-

ues. Presence or absence of trigones was not con-

sidered in determining the type of wall thickening.

This character could be scored in 25 of 25 (100%)

of the ingroup taxa.

19. Cuticle. Cuticular patterns in the Balan-

tiopsaceae vary from smooth to striate-papillose

to coarsely verrucose. In leaves of Balantiopsis

cancellata, B. rosea, and B. vermcosa (Fig. 6:5),

as well as Isotachis intortifolia, the papillae are

coarse, high, and spherical, with only a few large

papillae per cell, a condition we term "guttulate":

the papillae resemble spherical droplets of oil or

resin (Stearn, 1966). In other species (e.g., B.

montana, Fig. 5:4), the cuticle is finely and rather

distantly striate-papillose with many papillae per

cell. The leaves of Anisotachis, Austroscyphus

phoenicorhizus, and Eoisotachis are essentially

smooth, without any perceptible cuticle.

Leaf cuticle is treated as an unordered multi-

state character with three states: striate-papillose

(0), guttulate (1), smooth (2). This character could

be scored in 25 of 25 (100%) of the ingroup taxa.

20. Oil-bodies. In the taxa included in the anal-

ysis, the number of oil-bodies ranges from 5-1 1

per cell (e.g., B. montana, Fig. 2:12, 13) to com-

paratively few (2-7 per cell in B. convexiuscula,

Fig. 2:14) to 2 (rarely 3) large oil-bodies per cell,

situated at opposite ends of the cell (e.g., B. rosea,

Fig. 2:15). The state of 2 oil-bodies per cell is

also found in the three Isotachis species included

in the analysis, in Anisotachis splendens (Hassel

de Menendez & Solari, 1975), and in Austroscy-

phus phoenicorhizus (Fig. 10:5). Ruizanthus ve-

nezuelanus has 2-3(4) oil-bodies per cell (Schus-

ter, 1985a). Oil-bodies of Neesioscyphus carneus

and N. argillaceus are not known, but Gradstein

et al. (1977) report 2-3(4) per cell in an undeter-

mined Neesioscyphus species from Ecuador. The

outgroup taxon, T. subtrifidum, is reported to have

4-10 oil-bodies per cell (Hassel de Menendez &

Solari, 1975) and 3-5(6) per cell (Stewart, 1978).

Oil-body data were not available for Eoisotachis,

B. brasiliensis, or B. philippinensis.

Oil-bodies are treated as an unordered multi-

state character with three states: numerous, 8-13

(0), few, 2-7 per cell (1), 2 (rarely 3) per cell (2).

This character could be scored in 20 of 25 (80%)

of the ingroup taxa.

Reproductive

21. Accessory gynoecial structures. Gameto-

phyte-sporophyte interactions associated with the

protection of the developing sporophyte are con-

sidered to be of major significance in the classi-

fication of hepatics. In the outgroup taxon, Trian-

drophyllum, the sporophyte is described as sur-

rounded only by a simple perianth, i.e., a structure

of appendicular (foliar) origin, not involving stem

tissue (Hassel de Menendez & Solari, 1976, p.

102). Austroscyphus, Neesioscyphus, and Ruizan-

thus display stages in the elaboration of a stem

perigynium. Neesioscyphus argillaceus (Grolle,

1964, fig. 3:2) has a well-developed, exerted peri-

anth and a feebly developed basal perigynium.

Austroscyphus phoenicorhizus, A. iwatsukii (Ki-

tagawa, 1984, fig. 1:6, 7). Neesioscyphus carneus

(Grolle, 1964, fig. 1:2), and Ruizanthus venezue-

lanus (Schuster, 1985a, fig. 7:3) have an im-

mersed perianth and short, hypanthium-like peri-

gynium. Gynoecia of Anisotachis or Eoisotachis

are unknown.

Schuster (1984, fig. 30, p. 869) traces the stages

in evolution of a stem perigynium from a simple

perianth condition (fig. 30:3), as the result of "a

ring-like upgrowth of axial tissue below the re-

ceptacle," giving rise initially to what we refer to

as a basal, hypanthium-like perigynium. The ex-

treme development in this sequence is the elabo-

ration of a tubular, fleshy, erect stem perigynium

(fig. 30:5, Isotachis} with what are interpreted as

remnants of the perianth and "perichaetial bracts"

at its mouth.

A well-developed geotropic marsupium is pres-

ent in all species of Balantiopsis for which gy-

noecia are known. Schuster (1984, fig. 30, p. 869)

traces the evolution of the marsupium along four

possible lines, two of which are relevant here. The

first of these derives a marsupium (7) from a stage

with a simple perianth (3); the other derives a

marsupium (6) from a stem perigynium stage (5)

"by initiation of prostrate growth." Of the other

two possibilities, one derives a marsupium (7) di-

rectly from a hypothetical stage (1), "not known

in any extant genus," in which no protective

structures surround the archegonia, proceeding di-

rectly to a marsupium without an intermediate

ENGEL & MERRILL: AUSTRAL HEPATICAE. BALANTIOPSIS

35

200 n

10

FIG. 13. Isotachis montana Col. 1, Leaves and underleaves distant from young gynoecium. 2, Enlarged subgyn-

oecial leaves (basal to young gynoecium). 3, Teeth of leaf margin. 4, Upper-median leaf cells (just below sinus base;

cuticle shown in part). 5, Largest inner 9 bracts and, in middle, bracteole; two perigynial bractlets shown to right.

6, Largest inner 9 bracts and, in middle, bracteole; two perigynial bractlets shown to right. 7, Bractlet, a portion of

lobe enlarged at left, a portion of median-basal area shown at lower left. 8, Inner face of perigynium, distal sector

(up = unistratose portion; cpm = cilium of perigynium mouth; pr = polysratose rib, on which thin-walled cilia (c)

36

FIELDIANA: BOTANY

perianth stage. This is a theoretical possibility,

since no demonstrable perianth is present in Bal-

antiopsis (except perhaps in B. bisbifida), but an

unlikely one, since a well-developed perianth is

present in related, and presumably more primitive,

genera in the family (e.g., Neesioscyphus). The

other option leads to a marsupium (8) accompa-

nied by a "calyptral perigynium" and a perianth

(e.g., Arnellia). No member of the Balantiopsa-

ceae develops such a structure.

In Isotachis, a conical-cylindrical, fleshy, erect

stem perigynium is present (Fig. 14:8), and in

Balantiopsis, a conspicuous, geotropic marsupium

develops (Fig. 4:1, 2). At maturity, both structures

bear numerous, minute scalelike or ciliiform pro-

cesses around the mouth and just within the

mouth (Figs. 14:9, /. lyallii; 13:8, I. montana; 4:

2, B. diplophylla', 2:16, B. convexiusculd). There

is no evidence of a tubular perianth at any stage

of development.

We have studied gynoecia of B. bisbifida and

B. diplophylla from the earliest stages of devel-

opment, when the marsupium is present only as a

small bulbous distention of the apex. We also ex-

amined fully developed marsupia of these species

as well as B. asymmetrica, B. cancellata, B. con-

vexiuscula, and B. lingulata. In B. convexiuscula,

for example, 3-4 cycles of perigynial bractlets

(called "perichaetial bracts" by Schuster [1984,

p. 868]) are present (Fig. 2:16), surrounding the

mouth. The margins of these bractlets are typi-

cally ciliiform, and in B. diplophylla they are

densely so, the cilia consisting of uniseriate rows

of cells up to 13 cells in length.

Examination of developing perigynia of Iso-

tachis spp. reveals that the early stages of devel-

opment are virtually identical to the young mar-

supia of Balantiopsis spp. Several indistinct cy-

cles of perigynial bractlets, 8-10 in number, are

present surrounding the archegonia. As in Bal-

antiopsis, these bractlets are free from one anoth-

er, and there is no trace of a tubular perianth. The

outermost perigynial bractlets are ± regularly bi-

fid (or sometimes bisbifid), with a broadly ovate

disc, surmounted by 2(4) slender lobes (/. mon-

tana, Fig. 13:5, 6; /. minima, Fig. 11:9). The mar-

ginal and submarginal cells of the disc are thin-

walled and irregularly short-rectangular; the cells

of the lobes are more regular in shape, thick-

walled, and elongate (Fig. 13:7). Toward the cen-

ter, the bractlets become progressively smaller and

narrowly ovate-lanceolate, terminating in 4 cilia

consisting of a uniseriate row of 4-6 cells. An

interesting feature observed in /. montana is the

several projecting ribs on the inner perigynium

wall (Fig. 13:8). When fully developed, the outer

surface of the perigynium sometimes bears one or

more bractlets inserted near the base, identical in

shape and areolation to those found at the shoot

apex in young gynoecia; the innermost bractlets

remain at the apex, surrounding the mouth, or in-

serted just within it; mature marsupia of Balan-

tiopsis spp. also occasionally have bractlets in-

serted on the outer wall.

The resemblance between the longitudinal sec-

tions of a stem perigynium of Isotachis (Fig. 14:

8) and a marsupium of Balantiopsis (Fig. 4:2) is

particularly striking. Remarkably similar are the

thickness and general form of the stem calyptra,

the thickness of perigynium and marsupium walls,

and the fleshy, basal portion in which the sporo-

phyte foot is embedded, the position of bractlets

at or near the summit, etc.

Accessory gynoecial structures are treated as an

unordered multistate character with three states:

accessory structures none or with hypanthium-like

basal perigynium (0), erect tubular stem perigyn-

ium (1), marsupium (2). This character could be

scored in 15 of 25 (60%) of the ingroup taxa.

22. Number ofantheridia per bract. Antheridia

vary from 4-5 per bract in Ruizanthus (Schuster,

1985a, p. 71) to solitary in Austroscyphus phoen-

icorhizus. The remaining taxa included in the

analysis had 2-3 antheridia per bract (Fig. 13:10).

Number of antheridia per bract is treated as an

unordered multistate character with three states:

2-3 (0), 4-5 (1), antheridia solitary (2). This char-

acter could be scored in 18 of 25 (72%) of the

ingroup taxa.

are inserted, the cilia crowded, only a few shown for clarity). 9, Sector of main shoot showing differentiation of

androecium, dorsal view. 10, Opposing $ bracts, in situ, dorsal view. (Figs. 1, 2 from type of /. elegans (WELT); 3,

4 from type of /. montana (WELT); 5 from Hatcher 1429, New Zealand, South Is., Otago Prov., Fiordland, head of

Lake McKerrow; 6, 7, 9, 10, from Sorensen s.n., New Zealand, South Is., Southland Prov., Orepuke; 8 from Hatcher

569, New Zealand, South Is., Otago Prov., Fiordland, ca. 30 miles SE of Lake McKerrow.)

ENGEL & MERRILL: AUSTRAL HEPATICAE. BALANTIOPSIS

37

FIG. 14. Isotachis lyailii Mitt. 1, Portion of main shoot, lateral view. 2-4, Leaves. 5, Underleaf. 6, Leaf lobe,

cuticular detail shown in part. 7, 8, Upper-median leaf cells (near lobe bases). 9, Isotachis-type, perigynium and

subtending bracts, median-longitudinal section (A = perianth; B = perigynium; C = calyptra; D = sterile archegonia;

E = seta; F = foot) (X40). 10, Mature capsule showing spiral lines of dehiscence. 11, Capsule after dehiscence

showing spiral valves (X20). 12, Capsule wall, cross section. 13, Spore (X1895). 14, Elater (X340). (Figs.

38

FTELDIANA: BOTANY

SPOROPHYTE

23. Capsule shape. In Balantiopsis, marsupia

are not uncommon, but sporophytes are rarely

seen. Sporophytes are known in five species: B.

asymmetrica, B. bisbifida, B. cancellata, B. con-

vexiuscula, and B. purpurata, four of which are

described here for the first time (see below). Cap-

sules are spherical in Triandrophyllum and Rui-

zanthus and ± cylindrical in Neesioscyphus, Iso-

tachis, and Balantiopsis. Only immature sporo-

phyte are known in Austroscyphus (A. iwatsukii,

Kitagawa, 1984). Sporophytes of Anisotachis and

Eoisotachis are unknown.

Capsule shape is treated as a binary character:

spherical (0), ovate-cylindrical to cylindrical (1).

This character could be scored in 9 of 25 (36%)

of the ingroup taxa.

24. Capsule dehiscence. A remarkable charac-

teristic of the sporophytes of most members of the

suborder Balantiopsineae (sensu Schuster, 1972)

is the spiral dehiscence of the capsule wall (Bal-

antiopsis bisbifida, Fig. 2:17; Isotachis lyallii, Fig.

14:10, 11; AT. bicuspidatus, Grolle 1966, fig. 1).

Capsules of Rhizanthus are unique in the family

for the straight lines of dehiscence (Schuster,

1985a). Spiral capsule dehiscence also occurs in

some Calypogeiaceae, in some Geocalycaceae

(Saccogynidium), and in Plagiochila (e.g., P. gi-

gantea, Schuster, 1966, p. 591), where it is re-

garded as an independent development.

Capsule dehiscence is treated as a binary char-

acter: valves straight (0), valves spiral (1). This

character could be scored in 9 of 25 (36%) of the

ingroup taxa.

25. Capsule wall thickness. In the taxa includ-

ed in the analysis for which sporophytes are

known, capsule wall thickness varies from 2-5

cell layers. In general, the outermost cell layer of

the capsule wall is composed of larger cells that

are about equal in width (in surface view) to 2-3

of the cells of the inner layer(s), but the relative

thickness of the different layers (in cross section)

is variable. In T. subtrifidum the capsule wall is

4-5 layers thick, with the cells of the outer layer

roughly equal in thickness to two of the inner lay-

ers (Hassel de Menendez & Solari, 1975, fig. 21:

10-12). In N. carneus (Grolle, 1964, fig. 1:9), the

wall is 3-4-stratose (3-stratose above, 4-stratose

below), and in N. argillaceus 2-stratose, with the

outer wall only slightly thicker than the inner lay-

er(s). In Ruizanthus venezuelanus (Schuster,

1985a, fig. 6:14), the outer of the 2 (to 3) wall

layers is represented as only slightly thicker than

the inner. The capsule is uniformly 3-stratose in

Balantiopsis (Fig. 2:18) and Isotachis (Inoue,

1974, fig. 5:1), with the outermost layer equal to,

or often exceeding, the combined thickness of the

two inner layers. We elected to score wall thick-

ness merely as number of strata, without respect

to their relative thickness.

Another capsule wall characteristic worthy of

mention is the absence or only weak development

of secondary thickenings on the outer tangential

walls in Balantiopsaceae, seen in Ruizanthus

(Schuster, 1985a, fig. 6:13) and Neesioscyphus

(Grolle, 1964, figs. 1:11; 3:13), as well as in Bal-

antiopsis and Isotachis, where, for the most part,

thickenings are confined to the inner tangential

wall, with thickenings extending, buttress-like,

onto the radial walls (cf. Fig. 2:18 and Inoue,

1974, fig. 5:1). All observed sporophytes of Bal-

antiopsis are of this type. The extreme condition

is seen in Gyrothyra, of the related Gyrothyra-

ceae, where the capsule valves lack secondary

thickenings (Schuster, 1972, p. 361).

Capsule wall thickness is treated as an unor-

dered multistate character with four states: 4-5-

stratose (0), uniformly 3-stratose (1), 2-stratose

(or 2-3-stratose) (2), 3-4-stratose (3). This char-

acter could be scored in 9 of 25 (36%) of the

ingroup taxa.

Other potential characters

26. Perianth. In the outgroup taxon, T. subtri-

fidum, the developing sporophyte is surrounded

only by an involucre consisting of the 9 bracts

and bracteole, plus a simple perianth, presumably

formed by connation of the innermost cycle of

"leaves." Austroscyphus, Neesioscyphus, and

Ruizanthus have a tubular perianth, plus a short,

hypanthium-like stem perigynium. In Isotachis

and Balantiopsis no discernable perianth is pres-

1-2, 5, 6-8 from type of Jungermannia lyallii;3-4 from type of /. rosacea; 9, 11, 14 from Hatcher 1038, New

Zealand; 10, 12-13 from Engel 17916, New Zealand, South Is., Westland Prov., between the confluence of Jackson

River and Arawata River and Lake Ellery; Figs. 9, 11, 14, after Hatcher 1960-61.)

ENGEL & MERRILL: AUSTRAL HEPATICAE. BALANTIOPSIS

39

-^— Trlandrophyllum subtrifidum

— Ruizanthus uenezuelanus

Neesloscyphus argillaceus

— Neesioscyphus carneus

flustroscyphus phoenlcortiizus

Eoisotachis stephanli

Isotachis intortlfolia

Isotachis montana

Isotachis lyallii

flnlsotachls splendens

uerrucosa

rosea

tumida

ehnacea

cancellata

conueniuscula

>— ciliaris

brasillensls

philippinensis

crocea

dlplophylla

lingulata *.

montana

purpurata

bisbiHda

asymmetrica

— — — Triandrophy Mum subtrifidum

— Ruizanthus uenezuelanus

Neesioscyphus argillaceus

— Neesioscyphus carneus

Rustroscyphus phoenicorhizus

Eoisotachis stephanii

Isotachis intortifolia

Isotachis montana

Isotachis lyallii

Rnisotachls splendens

uerrucosa

rosea

tumida

ehnacea

cancellata

conueKiuscula

i— ciliaris

brasiliensis

philippinensis

crocea

diplophylla

montana

lingulata

purpurata

bisbiflda

asymmetrica

FIG. 15. Four most parsimonious cladograms. The length of the branch segments is proportional to the number

of apomorphies supporting that branch. Branches of zero length have no supporting apomorphies. Cladograms 1-3

are identical except for the resolution of the B. diplophylla group (indicated by *). Note also the difference in

placement of B. ciliaris: in 1-3 this species is resolved as a member of the clade representing sect. Pteridophylla; in

4, as a member of sect. Balantiopsis, which alters the topology of the two sections. (A consensus cladogram of these

four trees is shown in Fig. 17.)

ent, and there is no hint of fusion of the minute

bractlets, which surround the opening, into a tu-

bular perianth (Figs. 2:16; 4:2).

Normal 9 bracts and bracteole, in the usual

sense, are also absent in Isotachis and Balantiop-

sis. In leafy hepatics generally, these are formed

from the cycle of leaves immediately adjacent to

the perianth, but here there is no differentiated

inner cycle of bracts and bracteole. Instead, the

innermost cycle of larger subgynoecial leaves

have assumed the appearance and the function of

$ bracts and bracteole and for convenience are

usually described as such.

This character is closely associated with acces-

40

FIELDIANA: BOTANY

I Triandrophyllum subtnfidum

I i— Rulzanthus uenezuelanus

i— Neesioscyphus argillaceus

' Neesioscyphus carneus

r— Rustroscyphus phoenicorhizus

Eoisotachis stephanii

Isotachis intortlfolla

Isotachis montana

Isotachis lyallii

Rnisotachis splendent

uerrucosa

rosea

i— bra

n philipi

tumida

erinacea

cancellata

conuexiuscula

ciliahs

brasiliensls

philippinensls

crocea

Ediplophylla

— montana *

..r

purpurata

blsblHda

asymmetrica

llngulata

i Triandrophyllum subtrifidum

I r— Ruizanthus uenezuelanus

i— Neesioscyphus argillaceus

^* Neesioscyphus carneus

r— Rustroscyphus phoenicorhizus

Eoisotachis stephanii

Isotachis intortifolia

Isotachis montana

Isotachis lyallii

Rnisotachis splendens

uerrucosa

rosea

cancellata

conueKluscula

erinacea

tumida

J brasiliensls

hllipplnensls

iplophylla

....gUlata

_| Th

LJ— mi

• linnn

— — crocea

montana

•— ciliaris

purpurata

bisbifida

asymmetrica

FIG. 15. Continued.

sory gynoecial structures (character 21), since the

absence of a perianth appears to be a result of the

elaboration of these structures which have taken

over its function. We experimented with adding

to the data set a perianth character (perianth pres-

ent vs. perianth none), the results of which are

discussed below (p. 47). In the end, we decided

not to score the presence or absence of a perianth

separately to avoid attaching too much weight to

this character complex.

Results

We used MacClade 3.04 (Maddison & Maddi-

son, 1992) for entering data and editing the data

matrix, PAUP 3.1.1 (Swofford, 1991) for search-

ing for most parsimonious trees, and MacClade

for exploring the resulting cladograms, randomly

resolving polytomies, etc. Both programs were

run on a Macintosh Quadra 840AV.

The analysis of 26 taxa (Table 1) and 25 char-

ENGEL & MERRILL: AUSTRAL HEPATICAE. BALANTIOPSIS

41

- Triandrophyllum subtrifidum

Rulzanthus uenezuelanus

Neesioscyphus argillaceus

— Neesioscyphus carneus

flustroscyphus phoenicorhizus

Eoisotachis stephanii

Rnisotachis splendens

uemicosa

rosea

cancellata

conuenluscula

erinacea

tumida

brasillensis

philippinensis

25.1

— +—

26.1

B

L- d

I lini

— crocea

montana

- ciliaris

purpurata

bisbifida

asymmetrica

Ilsotachis intortifolia

sotachis montana

Isotachis lyallii

diplophylla

lingulata

26.1

21,1

• Triandrophyllum subtrifidum

I •— Ruizanthus uenezuelanus

i— Neesioscyphus argillaceus

"i Neesioscyphus carneus

r- Rustroscyphus phoenicorhizus

Eoisotachis stephanii

Rnisotachis splendens

uerrucosa

rosea

tumida

erinacea

— ^ cancellata

conueKiuscula

25.1

— ciliaris

i— brasiliensis

' philippinensis

'— | |— crocea

diplophylla

lingulata

montana

purpurata

bisbifida

"-TrT

I p III

26.1

asymmetrica

Ilsotachis intortifolia

sotachis montana

Isotachis lyallii

21,1

FIG. 16. Consensus cladograms of 13 trees resulting from the inclusion of character 26 (perianth present vs.

absent): 1, Adams consensus; 2, majority consensus. For discussion see text.

acters, with T. subtrifidum designated as the out-

group, yielded four most parsimonious clado-

grams (Fig. 15), each with a length of 114 steps,

with a consistency index of 0.560, a retention in-

dex of 0.660, and a rescaled consistency index of

0.370, using the heuristic search option of PAUP,

with tree-bisection-reconnection (TBR) branch-

swapping performed and steepest descent option

in effect. Use of a random addition sequence with

100 replicates yielded no shorter trees. In the

cladograms presented here, the ingroup is repre-

sented as monophyletic, with the outgroup as

paraphyletic to the ingroup taxa. Cladograms 1-3

(Fig. 1 5) are identical except for the resolution of

the species complex involving B. diplophylla, B.

lingulata, and B. montana; cladogram 4 differs in

the placement of B. ciliaris, with some differences

in the topology of the clades corresponding to

42

FIELDIANA: BOTANY

22,1

Triandrophyllum subtrifidum

Ruizanthus uenezuelanus

6,2 j— Neesioscyphus argillaceus

* Neesioscyphus carneus

23.1 24.1

I I

3,1 25.1

— I I

rstroscyphus phoenicorhlzus

Eoisotachis stephanii

Isotachis intortifolia

Isotachis montana

Isotachis lyallii

21.2

Rnisotachis splendens

crocea

montana

diplophylla

lingulata

ciliaris

purpurata

— bisbifida

asymmetrica

-6,2

FIG. 17. Adams consensus of the four most parsimonious cladograms presented in Figure 15, showing selected

character state transformations.

sect. Balantiopsis and sect. Pteridophylla (see be-

low for discussion of these taxa). In Figure 17 two

consistently resolved clades are shown, corre-

sponding to subg. Steereocolea and subg. Balan-

tiopsis. Nested within the latter is a polytomy with

three branches: the first of these contains only B.

ciliaris: the others correspond to sect. Balantiop-

sis and sect. Pteridophylla, respectively.

It should be noted that these cladograms are not

robust. A decay index (Bremer, 1988) was cal-

culated by instructing PAUP to save all trees one

step longer than the length of the most parsimo-

nious trees. The number of additional steps re-

quired to collapse a given clade is its decay index.

In the four most parsimonious trees obtained in

this analysis, all but one clade collapsed with the

addition of one step, with only the Austroscy-

phus-Eoisotachis clade retained. Bootstrapping

(Felsenstein, 1985) was attempted but was trun-

cated due to insufficient memory (32 MB).

With the inclusion of a perianth character (26,

above), the analysis yielded 13 most parsimonious

cladograms, each with a length of 127 steps, with

a consistency index of 0.520, a retention index of

0.688, and a rescaled consistency index of 0.314.

The Adams consensus (Adams, 1972) of these 13

cladograms is shown in Figure 16:1. A majority

consensus of these 1 3 cladograms is shown in Fig-

ure 16:2.

Addition of Lepicolea attenuata as an outgroup

yielded 40 most parsimonious cladograms, each

with a length of 1 33 steps. The strict consensus of

these 40 trees showed poor resolution, but succes-

sive reweighting of characters (Farris, 1988) by

maximum value of their rescaled consistency in-

dices yielded three cladograms that were similar in

most respects to those presented here, and conse-

quently Lepicolea was not included as an outgroup.

With the inclusion of both Lepicolea and Diplo-

phyllum taxifolium, monophyly of the family Bal-

ENGEL & MERRILL: AUSTRAL HEPATICAE. BALANTIOPSIS

43

antiopsaceae was not supported. Eleven most par-

simonious cladograms resulted, each with a length

of 136 steps. In the strict consensus of these 11

cladograms, Ruizanthus occupies a near-basal po-

sition, as a sister taxon to a clade containing the

remaining genera of the family and Diplophyllum.

With the exception of the placement of Diplophyl-

lum, the sister group relationships shown in the

cladogram (most importantly, those within Balan-

tiopsis) are unaffected, and consequently we chose

not to include this genus as an outgroup.

A thorough test of the monophyly of the family

is beyond the scope of this monograph. However,

monophyly of the family was assumed as an ini-

tial hypothesis of relationship and used as a basis

for selection of genera to include in the analysis.

Representatives of all genera considered to belong

to the family Balantiopsaceae were included. The

primary goal of the analysis was to resolve in-

group relationships within Balantiopsis. These

genera, plus Triandrophyllum, constitute function-

al outgroups (Watrous & Wheeler, 1981) used for

resolution of relationships within the functional

ingroup, Balantiopsis. Addition of more distant

outgroups (Lepicolea, Diplophyllum) would not

be expected to affect relationships within Balan-

tiopsis if these are already fully resolved.

Character State Reconstructions

In addition to determining the polarity of char-

acters used in the analysis, we used PAUP to re-

construct the ancestral character states at internal

nodes within the cladogram. Two options are pro-

vided in PAUP for this purpose (Swofford, 1991).

The first, accelerated transformation (ACCT-

RAN), tends to favor a single origin for a given

character state, followed by reversal(s). The sec-

ond, delayed transformation (DELTRAN), favors

independent origins of a character state (parallel-

ism), thus requiring fewer reversals. Both result

in the same number of steps on the tree, but the

locations at which changes occur may differ. We

experimented with both methods. The comments

that follow are based on DELTRAN reconstruc-

tions. Where the reconstructions of a character

differ appreciably or one seems preferable to the

other, we present both. The optimal reconstruc-

tions of selected characters are shown in Figure

17, an Adams consensus of the four most parsi-

monious cladograms.

1. Symmetry, CI = 0.333. Isophylly (or sub-

isophylly) is generally regarded as the primitive

condition in leafy hepatics (cf. Schuster, 1972, p.

330). However, the primitive Balantiopsaceae

(Ruizanthus, Neesioscyphus, Austroscyphus) are

strongly anisophyllous, and the julaceous, sub-

isophyllous taxa (Eoisotachis, Isotachis intortifol-

ia) are resolved in the consensus cladogram as

derived. Subisophylly is indicated in the clado-

gram as a homoplasy (probably habitat-related)

and within the context of this family is a derived

state, not a retention of a primitive state.

2. Pigmentation, CI = 0.273. Red to wine-red

pigmentation (0) is plesiomorphic in the ingroup

taxa. Sporadic changes from red to "brownish or

tinged with red" (1) mostly involve terminal taxa.

Pigmentation is variable, and scoring of state (1)

was somewhat subjective. Moreover, the absence

of reddish pigmentation is no indication that the

species is incapable of producing pigment under

certain conditions, and the intensity of pigmenta-

tion, when present, is no doubt related to environ-

mental factors.

3. Branching, CI = 1.000. The plesiomorphic

state is presence of only intercalary branching (0).

The absence of terminal branching distinguishes

Neesioscyphus and Ruizanthus from the other

members of the family. We think it is likely that

terminal branching has been lost in Neesioscyphus

and Ruizanthus and that the polarity 0 > 1 may

be an artifact of the selection of Triandrophyllum

(which lacks terminal branching) as an outgroup.

Schuster (1972, p. 330) indicates "branching both

terminal and intercalary" as generalized and

"branches purely terminal or purely intercalary"

as derived.

4. Rhizoid origin, CI = 0.333. The plesiom-

orphic state in the study taxa is rhizoids present

only at the bases of the underleaves (1). Rhizoids

originating from the bases of lateral leaves, as

well as from underleaves, evolved independently

in three terminal taxa — Ruizanthus, Neesioscy-

phus carneus, and B. bisbifida.

5. Division between lobes, CI = 1.000. The

plesiomorphic state is shallowly bilobed or bis-

bifid (0), evolving to deeply bilobed (1) only

once, as a synapomorphy of subg. Balantiopsis.

Evolution to "dorsal lobe displaced or detached"

(1 > 2) occurs twice, in B. erinacea and within

B. purpurata, suggesting that these may be dif-

ferent states (displaced vs. detached) and thus aut-

apomorphies of their respective species.

6. Relative lobe size, CI = 0.667. Dorsal lobe

44

FIELDIANA: BOTANY

subequal to the ventral (1) is the plesiomorphic

state. Dorsal lobe larger than the ventral (0 or 0

& 1) is a condition occurring only in Isotachis

and in the outgroup taxon, T. subtrifidum, asso-

ciated with incubous leaf insertion. According to

the DELTRAN reconstruction, reduction of the

dorsal lobe (1 > 2), occurred independently three

times — in Neesioscyphus, in subg. Steereocolea

(B. asymmetricd), and in subg. Balantiopsis,

where it supports sect. Balantiopsis. Extreme re-

duction in the dorsal lobe (1 > 3) occurs in B.

purpurata and within B. diplophylla (var. hock-

enii). A reversal to dorsal lobe subequal to the

ventral (2 > 1) occurs in sect. Pteridophylla in

such species as B. tumida and Anisotachis splen-

dens, while other species have relatively reduced

dorsal lobes. The ACCTRAN reconstruction has

the change from 1 > 2 occurring early at the base

of the clade uniting all the ingroup taxa save Rui-

zanthus, necessitating multiple reversals, a much

less satisfactory reconstruction.

7. Leaf insertion, CI = 0.800. Succubous in-

sertion (1) is plesiomorphic in Balantiopsaceae.

Succubous insertion evolves to incubous (2) only

once, in Isotachis. Transverse insertion (0) oc-

curs only as a polymorphism within terminal

taxa (Eoisotachis, Isotachis}. Schuster (1972, p.

353) says of the suborder Balantiopsineae,

"primitively subtransverse (incubous to trans-

verse in Isotachis; weakly succubous in Eoiso-

tachis}, derivatively strongly succubous (Gyroth-

yra, Neesioscyphus).'" Our analysis shows that

the strongly succubous (and strongly anisophyl-

lous) taxa (e.g., Ruizanthus, Neesioscyphus} are

primitive in the family.

8. Ventral lobe insertion, CI = 0.857. The ple-

siomorphic state is straight or comma-like (0),

evolving to U-shaped (2) only once, supporting

sect. Pteridophylla (in trees 1-3 with B. ciliaris

preserving the ancestral state). The J-shaped in-

sertion (1) is characteristic of sect. Balantiopsis,

with most species being either J-shaped or poly-

morphic (0 & 1) with respect to this character. The

most significant indication is that J-shaped and

U-shaped are alternative states, not a continuum

(i.e., the polarity of this character is 1 < 0 > 2,

not 0 > 1 > 2.)

9. Dorsal lobe insertion, CI = 0.571. The ple-

siomorphic state is ± transverse (0). Dorsal lobe

insertion oblique to longitudinal ( 1 ) is a homopla-

sy occurring several times in the cladogram.

U-shaped dorsal lobe insertion (2) is also a ho-

moplasy but confined to sect. Pteridophylla (An-

isotachis, B. erinacea). The polarity of this char-

acter is indicated as 1 < 0 > 2, not a continuum

1 > 0 > 2 as had been expected.

10. Ventral lobelleaf planation, CI = 0.560.

The plesiomorphic state is convex or concave (1).

In tree 4, evolution from convex to flat (1 > 0)

supports subg. Balantiopsis. Flat to plicate, etc.,

(1 > 2) occurs three times: once in /. intortifolia

and twice in sect. Pteridophylla. Canaliculate to

conduplicately folded or plicate leaves/ventral

lobes (2) appears at first glance to be a homopla-

sy. It could be argued that the plicate ventral lobes

of sect. Pteridophylla should not be scored the

same as the plicate leaves of Isotachis intortifolia.

Plicate leaves may simply be another aspect of the

same kind of habitat modification as subisophylly

in this species (see character 1).

11. Dorsal lobe orientation, CI = 1.000.

Leaves concave, not conduplicate (0) is the ple-

siomorphic state, evolving to complicate leaves

with appressed dorsal lobe (2) only once, sup-

porting subg. Balantiopsis. Assurgent dorsal lobes

(1) is a synapomorphy of the B. crocea-B. mon-

tana species pair.

12. Leaf armature, CI = 0.636. Leaves almost

unarmed (2) is the plesiomorphic state of this

character. It evolves from unarmed to moderately

armed (2 > 1 ) near the base of subg. Balantiopsis,

with B. purpurata retaining the primitive condi-

tion, and evolves independently to copiously

armed (0) at several points within the cladogram

in both sect. Balantiopsis and sect. Pteridophylla.

13. Projecting septa, CI = 0.333. Leaf margins

denticulate by projecting septa of the marginal

cells originated independently three times, in B.

ciliaris, in B. convexiuscula-B. cancellata, and in

B. tumida.

14. Underleaf lobing, CI = 0.250. Shallowly

bifid underleaves (1) is the plesiomorphic state in

Balantiopsaceae. This character serves as a good

illustration of the differences between ACCTRAN

and DELTRAN character reconstructions. The

DELTRAN reconstruction has deeply bifid under-

leaves (2) arising independently seven times;

ACCTRAN shows deeply bifid underleaves as the

plesiomorphic state in the family, retained in Rui-

zanthus and Neesioscyphus, but evolving to shal-

lowly bifid once and then reversing five times.

This character is highly homoplasous and phylo-

genetically uninformative.

ENGEL & MERRILL: AUSTRAL HEPATICAE. BALANTIOPSIS

45

15. Areolation, CI = 0.500. Scattered arrange-

ment of cells (0) is plesiomorphic in the family,

evolving to distinctly tiered (2) near the base of

subg. Balantiopsis, with B. purpurata retaining

the primitive condition. Tiered arrangement of

cells apparently also evolved independently in the

common ancestor of the Eoisotachis-Austroscy-

phus clade. A reversal from distinctly tiered to

scattered cells (2 > 0) occurs in B. diplophylla.

16. Border, CI = 0.250. Absence of a differ-

entiated border is the plesiomorphic state in the

family, although a border is present in Ruizanthus

lopezii, which was not included in the data set. A

border is shown evolving four times, at the base

of the B. verrucosa—B. rosea-Anisotachis splen-

dens clade, in B. cancellata, in B. crocea, and in

subg. Steereocolea. However, within sect. Pteri-

dophylla the border is of a special type, always

occurring in association with guttulate papillae

(19,1), and perhaps should not be considered the

same character state as the border in other species

(e.g., B. asymmetrica, B. crocea).

17. Leaf cell size, CI = 0.250. Larger cells (1),

defined as more than 20 |xm wide, is the plesio-

morphic state. Larger cells evolve to small cells

(0) at four points on the cladogram but in Bal-

antiopsis, only in sect. Pteridophylla. In tree 4, it

supports a "small-celled group," consisting of B.

cancellata, B. convexiuscula, B. erinacea, and B.

tumida. Elsewhere, small cells evolve supporting

the I. montana-I. lyallii clade and in Eoisotachis.

18. Wall thickenings, CI = 0.250. Evenly

thickened walls (0) is the plesiomorphic state. In-

termediate thickenings (1) evolved in four places

on the cladogram but only once in Balantiopsis.

In Figure 15:4, intermediate thickenings support

sect. Pteridophylla; in Figure 15:1-3, with B. cil-

iaris included in sect. Pteridophylla, this species

alone lacks intermediate thickenings. Elsewhere,

intermediate thickenings support the Austroscy-

phus-Eoisotachis clade and are present in N. car-

neus and /. lyallii.

19. Cuticle, CI = 0.400. Finely striate-papil-

lose cuticle (0) is the plesiomorphic state. Guttu-

late papillae (0 > 1) evolved three times, in /.

intortifolia, the B. verrucosa-B. rosea clade, and

B. cancellata. Within Balantiopsis guttulate pa-

pillae are confined to sect. Pteridophylla. Smooth

cuticle (2) occurs in the Austroscyphus-Eoiso-

tachis clade and again in Anisotachis.

20. Oil-bodies, CI = 0.333. The plesiomorphic

state in the family is relatively few (2-7) oil-bod-

ies per cell (1). Evolution to 2(3) oil-bodies per

cell ( 1 > 2) occurs twice, supporting the Austros-

cyphus-Eoisotachis-Isotachis clade, and once in

subg. Balantiopsis. Numerous oil-bodies (0) is a

condition occurring only in sect. Balantiopsis, in

three species, B. diplophylla, B. montana, and B.

lingulata. In Figure 15:1-3 these three taxa are

resolved as a single clade.

21. Accessory gynoecial structures, CI = 1.000.

The plesiomorphic condition is "accessory struc-

tures none or with hypanthium-like basal perigyn-

ium" (0). Evolution of an erect stem perigynium

(1) occurs only once, in Isotachis. Balantiopsis is

resolved as monophyletic, based on the presence

of a marsupium (2), the synapomorphy of the ge-

nus. The cladogram supports an independent ori-

gin of an Isotachis-type perigynium and a mar-

supium. However, the early stages of development

suggest a greater degree of homology between

these structures than has been previously sup-

posed; see also the notes on the effect of including

a perianth character in the data set, below.

22. Antheridia, CI = 1 .000. The plesiomorphic

state is 2-3 antheridia per bract (0), evolving to

4-5 per bract (0 > 1) in Ruizanthus, and from 2-

3 to solitary (0 > 2) in Austroscyphus. Solitary

antheridia may be a characteristic of the latter ge-

nus, since A. iwatsukii also has this condition (Ki-

tagawa, 1984).

23. Capsule shape, CI = 1.000. Spherical cap-

sules (0) is the plesiomorphic state and is retained

in Rhizanthus, whereas cylindrical capsules is an

apomorphy of the clade containing all the other

genera of Balantiopsaceae.

24. Capsule dehiscence, CI = 1.000. Straight

capsule valves (0) is the plesiomorphic state, with

spiral valves (1) supporting the clade uniting all

of the ingroup taxa except Rhizanthus.

25. Capsule wall thickness, CI = 1.000. The

outgroup condition, 4-5-stratose (0), does not oc-

cur in the ingroup. The plesiomorphic state in the

Balantiopsaceae is "2-stratose (or 2-3-stratose)"

(2), shown evolving to uniformly 3-stratose (1) at

the base of an Austroscyphus-Eoisotachis-Iso-

tachis— Balantiopsis clade in both Figures 17 and

16:1. This is deceptive, however, since capsule

wall thickness is not reported for Austroscyphus,

and sporophytes of Eoisotachis are unknown. In

Figure 16:2, however, character state (1) is an

apomorphy of an Isotachis— Balantiopsis clade, as

46

FIELDIANA: BOTANY

is perianth absent (26,1). Thus, relatively thin

walls appear to be the plesiomorphic condition in

this group of leafy liverworts, and the regularly

3-stratose condition is characteristic of all but the

most primitive members of the family.

26. Perianth, CI = 0.500. When a perianth

character is included, 13 most parsimonious trees

are obtained. The Adams consensus of these trees

(Fig. 16:1) is almost identical to that shown in

Figure 17 (the result obtained when this character

is not included), except for the presence of a po-

lytomy involving the Austroscyphus-Eoisotachis,

Balantiopsis, and Isotachis clades, respectively.

Note that the majority consensus of these 1 3 trees

(Fig. 16:2) is identical to Figure 15:1, except for

the resolution of Isotachis as a sister taxon to Bal-

antiopsis.

The optimal reconstructions of the accessory

gynoecial structures (21) and perianth (26) char-

acters are indicated on both cladograms. Perianth

present (0) is the plesiomorphic condition, re-

tained in Ruizanthus, Neesioscyphus, and Austros-

cyphus. The Adams consensus shows the loss of

a perianth (1) occurring independently in Balan-

tiopsis and Isotachis; in the majority consensus,

loss of a perianth is an apomorphy of a Balan-

tiopsis-Isotachis clade.

Discussion of Phylogenetic Relationships

Intergeneric Relationships

The genus Balantiopsis was the primary focus

of this study, but the inclusion of representatives

of other genera allows some observations con-

cerning relationships of genera and the phylogeny

of the Balantiopsaceae. Schuster (1972) estab-

lished the suborder Balantiopsineae ("Balantiop-

sidinae") with four "unifying characters": 1) a

tendency to form accessory structures for protec-

tion of the developing sporophyte (a marsupium

or an erect stem perigynium); 2) retention of, in

all cases, at least a vestige of a perianth; 3) bifid

underleaves, with rhizoids for the most part lo-

calized near their base; and 4) spirally dehiscent

capsules. More recently, with the discovery of

Ruizanthus (Schuster, 1985a), the characters of

the suborder have been modified to accommodate

this primitive taxon, which has spherical capsules

with straight valvular dehiscence.

Ruizanthus and Neesioscyphus are resolved as

the earliest divergent lineages on the cladogram

(Fig. 17). They share the lack of terminal branch-

ing (character 3,1) and the development of rhi-

zoids at the bases of the leaves as well as the

underleaves (4,0, also present in B. bisbifida). The

position of Ruizanthus in the cladogram is con-

sistent with the view that this genus is the most

primitive one in the family. In addition to spher-

ical capsules and lack of spiral dehiscence, the

subfamily Ruizanthoideae is supported by an aut-

apomorphy, antheridia 4-5 per bract (22, 1 ). Nee-

sioscyphus is resolved as a separate clade, a sister

taxon to the major clade that includes all of the

other genera (except Ruizanthus); the latter is sup-

ported by a uniformly 3-stratose capsule wall

(24,1).

Schuster (1985b) discusses the similarities of

Ruizanthus and Austroscyphus (as Acroscyphus)

and places the latter in the Ruizanthoideae based

on the prediction that Austroscyphus would prove

to have nonspiral capsule valves. Schuster (1990,

p. 250) suggests that these two genera are "the

remnants of the ancestral complex from which

other Balantiopsidineae, all apomorphic in the

spirally coiled capsule valves, evolved." Imma-

ture sporophytes of A. iwatsukii are illustrated by

Kitagawa (1984), but mature capsules have not

yet been observed in this genus. On the basis of

information in the data set and in the absence of

data on mature sporophytes, the cladogram pre-

sented here does not support the inclusion of Aus-

troscyphus in the Rhizanthoideae.

Austroscyphus, Eoisotachis, and Isotachis make

up a clade, roughly corresponding to the subfam-

ily Isotachidoideae, minus Neesioscyphus. Schus-

ter (1972, 1979) and Grolle (1972, 1983) include

Eoisotachis, Isotachis, and Neesioscyphus (the

last including species now placed in Austroscy-

phus) in the Isotachidoideae, characterized by "an

erect apparatus — perianth, perigynium, or combi-

nation thereof," in contrast to Balantiopsoideae,

where "there has been the evolution of a bilateral

rather than radial gynoecial system" (Schuster,

1972, p. 355). The implication is that a change

from an erect, symmetrical perigynium to a geo-

tropic, bilaterally symmetric marsupium is "a re-

sponse to prostrate growth" (I.e., p. 355).

The relationships between the functional out-

groups (i.e., the other genera of Balantiopsaceae

included in the analysis) are not fully resolved,

and the results of the analysis must be considered

tentative as far as these genera are concerned. A

good example of this is the position of Austros-

cyphus in the cladograms presented here. In habit

and gynoecial features Austroscyphus is closest to

ENGEL & MERRILL: AUSTRAL HEPATICAE. BALANTIOPSIS

47

TABLE 3. Proposed classification of higher taxa of

Balantiopsaceae.

Family Balantiopsaceae Buch

Subfamily Ruizanthoideae Engel & Merr.

Ruizanthus

Subfamily Balantiopsoideae

Tribe Neesioscypheae Engel & Merr.

Neesioscyphus, Austroscyphus

Tribe Balantiopseae

Subtribe Isotachidinae (Hatch.) Engel & Merr.

Isotachis, Eoisotachis, Hypoisotachis

Subtribe Balantiopsinae

Balantiopsis (Anisotachis, Steereocolea)

Neesioscyphus, and yet the two are resolved in

separate clades in Figures. 15-17. Shared features

of these two genera include 1) succubously in-

serted leaves; 2) anisophylly; 3) a tubular, exsert-

ed or immersed perianth; 4) a hypanthium-like

stem perigynium; and 5) a distinct involucre of

bracts and bracteole formed by the innermost cy-

cle of leaves and underleaf (adjacent to the peri-

anth). The data are incomplete, since sporophytes

of Austroscyphus are imperfectly known. In the

classification below, Austroscyphus has been

placed with Neesioscyphus. An Austroscyphus-

Eoisotachis clade is a consistent feature of all the

cladograms generated in the course of this study,

and yet these two genera are strikingly different

in almost all respects: the latter is a julaceous,

subisophyllous plant, similar to many Isotachis

species (e.g., /. intortifolia). Gametangia and spo-

rophytes of Eoisotachis are unknown, but we

think it will eventually prove to have an erect

stem perigynium of the Isotachis-type.

Monophyly of Isotachis is supported by the

erect, fleshy stem perigynium (21,2), incubous

leaf insertion (7,2), and dorsal leaf lobe larger

than the ventral (6,0). Without inclusion of a peri-

anth character (26), the resulting cladogram (Fig.

17) suggests that the marsupium of Balantiopsis

and the Isotachis-iype perigynium evolved inde-

pendently from an ancestor with a Neesioscyphus-

like basal stem perigynium (21,0). However, as

indicated previously, there is persuasive morpho-

logical evidence that these are homologous struc-

tures. In Figure 16:2, Isotachis is resolved as a

sister group to Balantiopsis; this clade is support-

ed by loss of perianth (26,1) and a uniformly

3-stratose capsule wall (25,1). A species of Iso-

tachis that combines both primitive and advanced

features is /. multiceps, the type of Isotachis subg.

Hypoisotachis (see p. 55), which is anisophyllous,

has microphyllous stolons and a 2(3)-stratose cap-

sule wall as in Neesioscyphus spp. and solitary

antheridia (as in Austroscyphus) with an erect

stem perigynium of the Isotachis-type.

It is clear from this discussion that the concept

of a subfamily Isotachidoideae should be aban-

doned in favor of a classification that reflects the

close relationship between Isotachis and Balan-

tiopsis. We propose a realignment of the genera

of Balantiopsaceae, summarized in Table 3. The

following new taxa and combinations are neces-

sary.

Subfamily Ruizanthoideae Schust., subfam. nov.

Balantiopsaceae subfam. Ruizanthoideae Schust. in

Clarke & Duckett, Bryophyte Systematics 75.

1979. New Manual Bryol. 2: 998. 1984. nom.

inval.

Ramificatio solum ventro-intercalaris; folia 2-3 cus-

pidata vel 3-4 lobata; antheridia 4-5 per bracteam; cap-

sulae sphaericae, parietibus 2(3) stratosis, valvis strictis

non spiralibus.

Type: Ruizanthus Schust., Phytologia 39: 240. 1978.

Tribe Neesioscypheae Engel & Merr., trib. nov.

Ramificatio solum ventro-intercalaris; folia succuba,

asymmetrice breviter 2-lobata; perigynium breve, hy-

panthio simile; perianthium praebens, exsurgens vel im-

mersum; parietibus capsularum 2-4 stratosis.

Type: Neesioscyphus Grolle, Oesterr. Bot. Z. Ill: 19.

1964.

Subtribe Isotachidinae (Hatch.) Engel & Merr.,

comb. & stat. nov.

Family Isotachidaceae Hatch., Nova Hedwigia 2: 579.

1960 ("Isotachaceae"). Subfamily Isotachidoi-

deae (Hatch.) Grolle, J. Bryol. 7: 210. 1972.

Type: Isotachis Mitt.

Key to Subfamilies, Tribes, and Subtribes of Balantiopsaceae

1. Capsule spherical, dehiscing by straight valves; antheridia 4-5 per bract

Subfamily Ruizanthoideae Schust. [Ruizanthus]

1. Capsule cylindrical, dehiscing by spiral valves; antheridia 2-3 per bract, or solitary. Subfamily Bal-

antiopsoideae 2

48

FIELDIANA: BOTANY

2. Sporophyte enclosed in a tubular, exserted or immersed perianth at the summit of a shallow,

hypanthium-like stem perigynium; innermost cycle of leaves and underleaf (adjacent to the peri-

anth) forming a distinct involucre of bracts and bracteole that are equal to or larger than vegetative

leaves; capsule wall (Neesioscyphus) 2-4-stratose

Tribe Neesioscypheae Engel & Merr. [Neesioscyphus, Austroscyphus]

2. Sporophyte enclosed in a stem perigynium or marsupium, a tubular perianth not formed; innermost

3-4 cycles of leaves and underleaves present as small, scalelike bractlets that fringe the opening

of the stem perigynium or marsupium, the involucre consisting of large subgynoecial leaves;

capsule wall uniformly 3-stratose. Tribe Balantiopseae 3

3. Sporophyte protected by an erect, symmetrical, tubular stem perigynium; leaves not complicate-

bilobed; leaf cells not, or in indistinct tiers

Subtribe Isotachidinae (Hatch.) Engel & Merr. [Hypoisotachis, Isotachis, Eolsotachis]

3. Sporophyte protected by a pendant, geotropic, asymmetrical marsupium; leaves complicate-

bilobed (rarely simple and bisbifid); leaf cells typically distinctly tiered

Subtribe Balantiopsinae [Balantiopsis (incl. Anisotachis, Steereocolea)]

Balantiopsis

Monophyly of Balantiopsis is supported by a

pendant, asymmetrical, geotropic marsupium

(21,2). We recognize two subgenera (subg. Steer-

eocolea and subg. Balantiopsis) within the genus,

and within the subg. Balantiopsis, two sections

(sect. Balantiopsis and sect. Pteridophylld), cor-

responding to units that were consistently re-

solved in the cladograms presented here. Aniso-

tachis splendens is resolved as a terminal branch

within Balantiopsis sect. Pteridophylla. The sup-

port for each of these infrageneric taxa is dis-

cussed below. The disposition of species is shown

in Table 4.

Subgenus Steereocolea (Schust.) Schust.

Steereocolea Schust., Bull. Natl. Sci. Mus. Tokyo

11(1): 23. 1968. Balantiopsis subg. Steereocolea

(Schust.) Schust., J. Hattori Bot. Lab. 36: 357.

1972 [Jan. 1973].

Leaves shallowly bilobed to bisbifid (to 0.5 or

less), not conduplicately folded; leaf insertion

simple, oblique, the line of insertion ± straight;

dorsal lobe erect to spreading-reflexed toward the

stem base; leaf cells scattered.

Steereocolea was proposed as a monotypic ge-

nus by Schuster (1968), based on the presence of

an "inflated, quite well-developed" perianth in

the early stages of gynoecial development in the

type species, B. bisbifida. We have not seen the

specimen on which his description was based.

Later, Schuster (1972, p. 357) reduced Steereo-

colea to a subgenus of Balantiopsis, citing an En-

gel collection of B. bisbifida from the Hero ex-

pedition to southern Chile with pendant marsupia

and a "vestigial perianth" (cf. Engel, 1978). Schus-

ter suggests that perianth formation may be vari-

able in this species.

At maturity, B. bisbifida has a well-developed

marsupium, as does B. asymmetrica. In neither spe-

cies can we confirm the presence of a tubular peri-

anth at any stage of development. Notwithstanding,

Schuster's characterization of subg. Steereocolea as

"the starting point in evolution within the Balan-

tiopsidoideae" (Schuster, 1972, p. 357) is support-

ed by the position of these species in the clado-

gram. Balantiopsis bisbifida and B. asymmetrica

are resolved as sister taxa, and these two species

TABLE 4. Conspectus of Genus Balantiopsis Mitt.

Subg. Steereocolea (Schust.) Schust.

B. bisbifida (Steph.) Steph.

B. asymmetrica (Herz.) Engel

Subg. Balantiopsis

Sect. Balantiopsis

B. purpurata Mitt.

B. diplophylla (Hook. f. & Tayl.) Mitt,

var. diplophylla

van hockenii (Berggr.) Engel & Mem

B. lingulata Schust.

B. philippinensis H. Inoue

B. brasiliensis Steph.

B. montana (Col.) Engel & Merr.

B. crocea Herz.

B. ciliaris Halt.

Sect. Pteridophylla (Hatch.) Engel & Merr.

B. tumida Berggr.

B. convexiuscula Berggr.

B. cancellata (Nees) Steph.

B. erinacea (Hook. f. & Tayl.) Mitt.

B. splendens (Steph.) Engel & Merr.

B. verrucosa Engel & Merr.

B. rosea Berggr.

ENGEL & MERRILL: AUSTRAL HEPATICAE. BALANTIOPSIS

49

were placed first in the genus by Engel (1968).

Both species are included in subg. Steereocolea by

Hassel de Menendez & Solari (1975).

Balantiopsis asymmetrica and B. bisbifida are

unique in the genus in having an essentially

straight succubous leaf insertion and noncompli-

cately bilobed leaves with erect to spreading-re-

flexed dorsal lobes. The presence of rhizoids at

the bases of leaves as well as underleaves (17,0)

is a homoplasy found in B. bisbifida, Ruizanthus,

and N. carneus.

The two species of the subgenus may be distin-

guished as follows.

Key to Species of Balantiopsis subg. Steereocolea

1. Dorsal lobe equal to or slightly larger than the ventral; median ventral lobe cells 24-36 u,m wide;

underleaves bilobed to 0.5 or more; rhizoids originating at bases of both leaves and underleaves

B. bisbifida

1. Dorsal lobe at most 0.5 the ventral; median ventral lobe cells 31-53 u>m wide; underleaves bilobed

to 0.4 or less; rhizoids only at bases of underleaves B. asymmetrica

Balantiopsis bisbifida (Steph.) Steph.

Isotachis bisbifida Steph., Bih. Kongl. Svenska Veten-

skapsakad. Handl. 26 (in, 17): 24. 1901. Bal-

antiopsis bisbifida (Steph.) Steph., Spec. Hep. 4:

101. 1910. Steereocolea bisbifida (Steph.)

Schust., Bull. Natl. Sci. Mus. Tokyo 11(1):

25.1968. Original material: Chile, Prov. Magal-

lanes, Isla Desolation, Dusen 228 (G!).

DESCRIPTION— Engel (1968, 1978); Schuster

(1968, 1972); Hassel de Menendez & Solari

(1975).

ADDITIONAL DATA — Oil-bodies in all leaf cells,

scattered throughout cell, occupying small frac-

tion of cell volume, light brown, 4-8(1 1) per cell,

ovoid to elliptic to fusiform to crescentic, some

spherical, coarsely papillose, 12-16 X 5-8 u,m, a

few 6-7 X 6 jim.

Seta with 25 rows of epidermal cells surround-

ing an inner core of ca. 40 cells of about equal

size to the outer layer, the epidermal and inner

cells uniformly thin-walled. Capsule long-cylin-

drical, with spiral dehiscence, the wall 31-38 u,m

thick, of 3 layers, the outer layer of cells ca. equal

to thickness of both interior strata; outer layer of

cells short-rectangular, large, the inner tangential

wall with irregular semiannular bands extending

onto the radial longitudinal walls but gradually

thinning and ending short of the outer tangential

wall, the radial walls appearing slightly sinuous

in surface view, the outer tangential wall devoid

of thickenings; innermost layer of cells with semi-

annular bands pale, common, the bands not

forked.

Spores 10.6-11.5 u,m, the exine brownish yel-

low, with a network of very wide, irregular, coa-

lescing vermiform ridges that anastomose to form

irregular but rather deeply set areolae; spore:elater

diameter ratio 0.9-1.2:1. Elaters tortuous, (8.6)10.1-

12 u,m wide, bispiral, the spirals 2.9-4.3(4.8) u-m

wide, loosely wound.

DISTRIBUTION AND ECOLOGY — Engel (1978, 1990).

SPECIMEN SEEN— CHILE, PROV. VALDIVIA: SW

slope of Volcan Quetrupillan, immediately SW of La-

guna Los Patos, Forestal Trafun, 1450-1600 m, Engel

11144 (F).

Balantiopsis asymmetrica (Herz.) Engel

Balantiopsis latifolia var. asymmetrica Herz., Rev.

Bryol. Lichenol. 23: 53. / 11. 1954. Balantiopsis

asymmetrica (Herz.) Engel, Nova Hedwigia 16:

93. 1968. Lectotype (fide Engel, 1968): Chile,

Prov. Aisen, Puerto Magdalena, Schwabe 20a

(JE!).

DESCRIPTION — Engel (1968); Hassel de Menen-

dez & Solari (1975).

ADDITIONAL DATA — Seta with 29 rows of epi-

dermal cells surrounding an inner core of ca. 60

cells of similar size, the outer layer and peripheral

cells of the inner core thicker-walled and with cor-

ner thickenings, the innermost cells delicate and

thin-walled. Capsule long-cylindrical, with spiral

dehiscence, the wall 42-48 u,m thick, of 3 layers,

the outer layer of cells ca. equal to thickness of

both interior strata; outer layer of cells short-rect-

angular, large, the radial walls with sporadic,

small, nodule-like thickenings mostly on longitu-

dinal walls but sometimes on transverse walls, the

longitudinal and transverse walls with thin, slight-

ly sinuous, continuous sheets of pigmented ma-

terial; innermost layer of cells with longitudinal

50

FTELDIANA: BOTANY

radial walls with thick, continuous sheets of wall

material, with semiannular bands pale, common,

rather wide, sometimes forked and anastomosing

to delimit fenestrae.

Spores 12-13 jim, the exine pale yellowish

brown, with a network of irregular, coalescing

vermiform ridges that freely anastomose to form

numerous, shallow areolae; spore:elater diameter

ratio 1.2-1.4:1. Elaters tortuous, 9.1-10.1 |xm

wide, bispiral, the spirals 2.9-3.8 jim wide, loose-

ly wound.

DISTRIBUTION AND ECOLOGY — Occurring in Val-

divian and Magellanian zones of southern South

America and in Juan Fernandez. A plant of mossy

forests (e.g., of Nothofagus betuloides, N. pumilio,

Drimys, and Berberis ilicifolia at Puerto Churru-

ca, Isla Desolacion, or of N. dombeyi and Saxe-

gothaea in the Refugio Antillanca area) at sea lev-

el for most of its range, but to the north extending

to ca. 730 m. It typically occurs on rock walls or

on soil in protected hollows of cliff bases or moist

cliff ledges, at times in densely shaded niches.

SELECTED SPECIMENS SEEN— CHILE, PROV. MA-

GALLANES: Isla Desolacion, Puerto Churruca, head of

Brazo Lobo, Engel 5838 (F, MSC); Fiordo Peel, N shore

of Caleta Amalia, Engel 5435 (F, MSC); S side of Isla

Madre de Dios, head of fiord E of Mt. Roberto, Engel

5141 (F, MSC). PROV. AISEN: At landing cove near gla-

cier on N side of Fiordo Tempano, Engel 4376C (F,

MSC). PROV. OSORNO: Near road to Refugio Antillan-

ca, 12.1 km by road below Refugio, 550 m, Engel 11516

(F); near road (10-11 km) to Refugio Antillanca, be-

tween Laguna El Encanto and Lago Toro, 730 m, Engel

4082 (F, MSC). PROV, CAUTIN: Parque Nacional Vil-

larrica, N slope of Volcan Villarrica, 6.1 km by road S

of park entrance, 1150 m, Engel 11192 (F).

Subgenus Balantiopsis

Leaves deeply bilobed (to 0.6 or more), sharply

keeled and conduplicate-folded; leaf insertion

complex, the line of insertion flexed or sharply

bent; dorsal lobe much smaller than the ventral

and appressed to the ventral (rarely assurgent);

leaf cells in tiers (rarely scattered).

The subgenus is supported by two synapomor-

phies, deeply bilobed leaves (5,1) that are strongly

conduplicately folded ( 1 1 ,2). In this subgenus the

line of leaf insertion is fundamentally succubous

(as in the more primitive subg. Steereocolea), but

it becomes more complex by elaboration of the

insertion of the dorsal and ventral halves of the

leaf. The insertion of the ventral lobe progresses

from essentially straight at the ventral end to

J-shaped (in sect. Balantiopsis) or from straight to

strongly recurved and U-shaped (an apomorphy

of sect. Pteridophylla). The insertion of the dorsal

lobe may be oblique and decurrent on the stem,

± transversely inserted, or strongly arched and

U-shaped (9,2) in Anisotachis and B. erinacea.

The line of leaf insertion is typically bent at the

insertion of the keel between the ventral and dor-

sal lobes. The displacement or complete detach-

ment of the dorsal lobe (5,2) is a homoplasy, oc-

curring in B. purpurata, the basal species in sect.

Balantiopsis, and in B. erinacea, arguably the

most specialized species in sect. Pteridophylla.

The presence of a differentiated leaf border is a

homoplasy found also in subg. Steereocolea and

in Ruizanthus lopezii; unique to several species of

sect. Pteridophylla, however, is the cuticular dif-

ferentiation on the cells of the leaf border.

Variation in leaf armature is taxonomically

bothersome, in some cases varying considerably

within species. Extreme development of leaf ar-

mature (12,0) is a homoplasy originating indepen-

dently several times, in B. lingulata and B. ciliaris

(sect. Balantiopsis) and in B. rosea (sect. Pteri-

dophylla). Sparingly armed leaves (12,1) are ap-

parently the plesiomorphic condition in the family

(Ruizanthoideae, Neesioscypheae, subg. Steereo-

colea), and both sections of subg. Balantiopsis

contain one or more species with almost unarmed

leaves (e.g., B. tumida, B. diplophylla).

In our study of the New Zealand species of Bal-

antiopsis (all members of subg. Balantiopsis), we

initially recognized three groups of species, each

containing two to three species. The first group

centered on B. diplophylla, B. lingulata, and B.

montana, with relatively large, scattered (irregu-

larly arranged) leaf cells, relatively large numbers

of oil-bodies per cell, a striate-papillose cuticle,

and a straight to at most J-shaped insertion of the

ventral leaf lobe. A second group, consisting of

B. rosea and B. verrucosa, was characterized by

guttulate papillae, a differentiated leaf border of

striate-papillose cells, and two oil-bodies per cell,

situated in opposite ends of the cells. A third

group, consisting of B. convexiuscula and B. tum-

ida, had small, short-quadrate leaf cells and

lacked a differentiated border but had distinctly

denticulate margins formed by the projecting sep-

ta of the marginal cells. Inclusion of the extrater-

ritorial species of the genus, however, makes it

clear that this was a geographically biased view

of relationships. For example, B. cancellata, an

American species, combines the small cells, dense

ENGEL & MERRILL: AUSTRAL HEPATICAE. BALANTIOPSIS

51

areolation, and projecting septa of B. convexius-

cula and B. tumida with the coarsely guttulate pa-

pillae and differentiated border of B. rosea and B.

verrucosa.

In the consensus cladogram presented here

(Fig. 1 7), two major groupings are apparent with-

in subg. Balantiopsis, one comprised of general-

ized species (sect. Balantiopsis) and the other of

relatively specialized species (for which the oldest

available name is sect. Pteridophylla). Uncertain-

ty with respect to the placement of B. ciliaris is

reflected in the consensus cladogram by making

this species a member of an unresolved polytomy;

B. purpurata, on the other hand, is resolved as the

basal (earliest divergent) species in the subgenus

(for a discussion of our placement of these species

in sect. Balantiopsis, see below).

Section Balantiopsis

Ventral lobe flat or concave, the insertion ±

straight, comma-like, to J-shaped; dorsal lobe in-

sertion oblique to longitudinal; leaf cells scattered

to indistinctly tiered; leaf cells evenly thin- walled;

cuticle striate-papillose (never guttulate).

The section is characterized by several features:

1) straight to at most J-shaped insertion of the

ventral lobe; 2) generally larger leaf cell size; 3)

evenly thin- or thick-walled leaf cells, lacking in-

termediate thickenings; and 4) relatively large

numbers of oil-bodies per cell. The tendency to-

ward extreme reduction of the dorsal lobe, seen

in B. diplophylla var. hockenii and B. purpurata,

is found only in sect. Balantiopsis.

Balantiopsis purpurata Mitt.

Balantiopsis purpurata Mitt, in Thomson & Murray,

Rep. Scient. Results Challenger. Bot. 1(3): 86.

1885. Original material: Juan Fernandez, Saun-

ders (FH!, NY!).

DESCRIPTION — Engel (1968).

ADDITIONAL DATA — Seta with 17 rows of epi-

dermal cells surrounding an inner core of ca. 27

cells of about the same size, the epidermal cells

firm-walled, the inner cells thin-walled and deli-

cate. Capsule long-cylindrical, with spiral dehis-

cence, the wall delicate, 22-28 (Jim thick, of 3

layers (in median and basal sectors, 2-stratose

above), the outer layer of cells ca. equal to thick-

ness of both interior strata; outer layer of cells

short-rectangular, large, the radial walls exceed-

ingly thin-walled and delicate, with small nodular

thickenings (as seen in surface view) but frequent-

ly extending as spurs onto the inner tangential

wall; innermost layer of cells with small nodular

thickenings largely confined to the longitudinal

radial walls, the thickenings smaller and less

prominent than those of outer layer, the thicken-

ings at times extending onto both inner and outer

tangential walls as faint spurs or semiannular

bands.

Spores 8.6-9.6 jim, the exine pale brownish

yellow with a network of irregular, coalescing

vermiform ridges that freely anastomose to form

numerous, shallow areolae; spore:elater diameter

ratio 0.75-0.9:1. Elaters tortuous, 10.1-12 |xm

wide, bispiral, the spirals 3.8-4.3 (Jim wide, loose-

ly wound.

This species occupies a basal position in subg.

Balantiopsis and is resolved as the sister taxon to

the clade containing all of the other species of the

subgenus. This species seems more appropriately

placed in sect. Balantiopsis than as a monotypic

taxon, coordinate with the other species of the

subgenus. It is also similar in many respects to

subg. Steereocolea but has distinctly (and deeply)

complicately bilobed leaves. The cells are scat-

tered in arrangement as in subg. Steereocolea as

well as in B. diplophylla, which appears to be its

counterpart in temperate Australasia. The leaf

cells in B. purpurata are among the largest in the

genus: 34-45(51) |im wide, similar in size to

those of B. asymmetrica. The marked reduction of

the dorsal lobe in B. purpurata (6,3) is the ex-

treme expression of a tendency that is general in

subg. Balantiopsis but also apparent in B. asym-

metrica. A parallel reduction is seen in B. diplo-

phylla var. hockenii.

The leaf margins in B. purpurata are often un-

armed, or nearly so, and the ventral lobe insertion

is essentially straight. The displacement of the

dorsal lobe to the ventral lobe lamina a short dis-

tance above the base is a unique characteristic of

this species. At times the keel is almost lacking

and represented by a portion of a single cell. The

cuticle of this species is unusual in being very

densely papillose, the papillae being densely

crowded and at times appearing as a more or less

continuous, roughened surface.

DISTRIBUTION — Confined to the Valdivian zone

of southern South America and Juan Fernandez.

SELECTED SPECIMENS SEEN— CHILE, PROV. CHIL-

OE: Isla Chiloe, Cordillera San Pedro, near aserradero

at San Pedro, 320 m, Engel 11848 (F); ibid., Rio Pudeto,

52

FIELDIANA: BOTANY

abandoned rail cut at Pupelde, near sea level, Engel

11782 (F). PROV. VALDIVIDA: W slope of Cordillera

Pelada, 8.8 km by road W of El Mirador on road be-

tween La Union and Punta Hueicolla, 580 m, Engel

12296 (F); near Rio Futa in vicinity of Futa, 5.9 km by

road S of junction of Highways T-60 and T-65, near sea

level, Engel 10852 (F); Rio El Lingue, between Mehufn

and Lleco, Engel 3863 (F, MSC) ; 2.1 km by road N of

Mehum, S of Queule, ca. 15 m, Engel 11398 (F). PROV.

MALLECO: Cordillera Nahuelbuta, Parque Nacional

Contulmo, along trail from western entrance, 7 km by

road E of Contulmo, 330-360 m, Engel 12490 (F).

Balantiopsis ciliaris Hatt.

Balantiopsis ciliaris Hatt., J. Jap. Bot. 41: 129. / 1-

2. 1966. Holotype: North Borneo (Sabah), Mt.

Kinabalu, between Kambaranga Radio Station

and Waterfalls, 2000-2150 m, Mizutani 2347

(MICH!).

DESCRIPTION — Hattori (1966, figs. 1, 2); Piippo

(1984, fig. 16a-d).

This paleotropical species exhibits the extreme

development of leaf armature in the genus, with

up to 30 cilia on the ventral margin in the speci-

men seen (to 95 on both lobes in subsp. novogui-

neensis, fide Hattori, 1966). In general aspect and

leaf armature, B. ciliaris resembles the New Zea-

land B. lingulata but differs in the extremely nar-

row keel, only 1-2 cells wide.

This species shares with the species of sect.

Pteridophylla the condition of 2 oil-bodies per

cell but has numerous features characteristic of

sect. Balantiopsis: straight or comma-like ventral

lobe insertion, larger leaf cells, and no interme-

diate thickenings. The species seems best placed

in sect. Balantiopsis. The species is divided into

two subspecies by Hattori (1966), subsp. ciliaris

of North Borneo and the Philippines and subsp.

novoguineensis, endemic to New Guinea (Piippo,

1984).

DISTRIBUTION. — New Guinea (1500 — 2500 m),

Borneo, and the Philippines.

SPECIMEN SEEN— NEW GUINEA, MOROBE PROV.:

Cromwell Mts., 7 km SE of Indagen airstrip, 2450-2555

m, Koponen 30985 (F).

Section Pteridophylla (Hatch.) Engel & Merr.

Ventral lobe insertion U-shaped, the lobe often

canaliculate to conduplicately folded or plicate;

leaf cells with intermediate thickenings; cuticle

distinctly papillose (often guttulate, rarely

smooth), often with a contrasting border of striate-

papillose cells; oil-bodies 2 per cell, at opposite

ends of the cell.

This section includes the more specialized spe-

cies of the genus characterized by a greater elab-

oration and complexity of leaf form and insertion.

In the cladogram, the section is supported by in-

termediate wall thickenings (18,1) and the strong-

ly recurved, U-shaped insertion of the ventral lobe

(8,2). Other features found only in sect. Pterido-

phylla (but not in all species) include ventral lobe

longitudinally folded and plicate (10,2), insertion

of dorsal lobe transverse to U-shaped (9,2), small

leaf cells (17,0), guttulate papillae (19,1), and a

leaf border (16,1) accentuated by differentiation

of the cuticle. With the exception of B. convex-

iuscula and B. tumida, oil-bodies are consistently

two per cell, situated in opposite ends of the cell.

Intermediate wall thickenings (18,1) are an apo-

morphy of this section but also occur in other gen-

era of the family.

The origin of the sectional epithet Pteridophyl-

la ("fern leaf") is obscure and was not explained

by Hatcher (1960-61), who proposed it as a

monotypic section of Isotachis, including only /.

splendens (Anisotachis splendens).

Balantiopsis cancellata (Nees) Steph.

Ptilidium cancellatum Nees in G. L. & N., Syn. Hep.

251. 1845. Balantiopsis cancellata (Nees)

Steph., Hedwigia 32: 145. 1893. Original mate-

rial: "In Peruvia ad Plagiochilam hookerianam

repens . . . ," sin. coll. (STR!).

DESCRIPTION— Engel (1968, 1978).

ADDITIONAL DATA — Oil-bodies in all leaf cells,

occupying small fraction of cell volume, at op-

posing ends of cell, tan or buff to pale yellowish,

brown, glistening, 2 per cell (even near leaf base),

subspherical to ovoid to elliptical to crescentic,

coarsely papillose, the globules slightly protrud-

ing, 5-7 X 4-5 u>m to 8-1 1 X (4)5, spherical ones

5 |jun.

Seta with 25 rows of epidermal cells surround-

ing an inner core of ca. 40 cells of about equal

size, the epidermal layer and the outermost layer

of the core moderately thick-walled and with cor-

ner thickenings, the inner core cells thin-walled

and delicate. Capsule long-cylindrical, with spiral

dehiscence, the wall 40-46 |im thick, of 3 layers,

the outer layer of cells exceeding thickness of

both interior strata combined; outer layer of cells

short-rectangular, large, in surface view the lon-

gitudinal and transverse walls with localized

sheetlike thickenings without nodular or bandlike

thickenings, in cross section the cells with short

spurlike thickenings extending buttress-like from

ENGEL & MERRILL: AUSTRAL HEPATICAE. BALANTIOPSIS

53

radial walls a short distance onto the inner tan-

gential wall; innermost layer of cells with longi-

tudinal and transverse radial walls with strongly

thickened, continuous sheets of wall material,

with semiannular bands pale, common, very wide,

closely spaced and numerous, often forked and

anastomosing, the close and forked bands delim-

iting pitlike fenestrae (resembling scalariform pit-

ted tracheids).

Spores 9.1-10.1 u.m, the exine pale yellowish

brown, with a network of irregular, coalescing

vermiform ridges which freely anastomose to

form numerous, shallow, small areolae; spore:el-

ater diameter ratio 0.7-0.9:1. Elaters tortuous,

10.1-12.5 u.m wide, bispiral, the spirals 3.8-4.8

|xm wide, loosely wound.

DISTRIBUTION — Occurring on the Falkland Is-

lands, the Valdivian and Magellanian zones of

southern South America, and on Juan Fernandez

(see Engel, 1978, 1990).

SELECTED SPECIMENS SEEN— CHILE, PROV. MA-

GALLANES: Isla Riesco, E side of Bahia Borja, Paso

Tortuoso, Engel 6157B (F, MSC); Isla Desolacion, head

of Brazo Lobo, Engel 5837 (F, MSC); head of Fiordo

Peel, Engel 5494 (F, MSC); Fiordo Peel, N shore of Caleta

Amalia, Engel 5398 (F, MSC); Isla Chatham, N shore of

Bahia Wide, Engel 5349 (F, MSC); S side of Isla Madre

de Dios, head of fiord E of Mt. Roberto, Engel 5143 (F,

MSC); near E shore of Isla Pilot (Puerto del Morro, Canal

Trinidad), Engel 4760B (F, MSC); Puerto Charnia (S side

of Isla Wellington), head of inlet, Engel 4816 (F, MSC).

PROV. AISEN: Near glacier on N side of Fiordo Tem-

pano, Engel 4376D (F, MSC). PROV. CHILOE: Isla Chi-

loe, Cocauque area, across Estero Yaldad from village

of Yaldad, sea level, Engel 11986 (F); ibid., Cordillera

San Pedro, near aserradero at San Pedro, 320 m, Engel

11841 (F); ibid., Cordillera San Pedro, Butalcura, near

Rio Butalcura, 100 m, Engel 11801 (F); ibid., Aguas

Buenas area, 4.7 km E along Aguas Buenas road from

Ancud-Quemchi road, 100 m, Engel 12185 (F). PROV.

OSORNO: Road below Refugio Antillanca, 3000 ft, En-

gel 3942 (F, MSC). PROV. VALDIVIDA: Cordillera Pe-

lada, summit of El Mirador, ca. 100 m SW of refugio,

near road between La Union and Punta Hueicolla, 1000

m, Engel 12362 (F); Corral, Quebrada El Boldo, 55 m,

Mahu 11939 (F); near Rio Futa in vicinity of Futa, 10.5

km by road S of junction of Highways T-60 and T-65,

10 m, Engel 11009 (F); Rio El Lingue, between Mehufn

and Lleco, Engel 3884 (F, MSC). PROV. MALLECO:

Cordillera Nahuelbuta, Parque Nacional Contulmo,

along trail from western entrance, 7 km by road E of

Contulmo, 330-360 m, Engel 12528 (F). PROV. MAL-

LECO/PROV ARAUCO: Parque Nacional Nahuelbuta,

Aguas Calientes, 10.2 km W of park entrance, 1240 m,

Engel 12622 (F).

Balantiopsis erinacea (Hook. f. & Tayl.) Mitt.

Jungermannia erinacea Hook. f. & Tayl., London J.

Bot. 3: 462. 1844. Gottschea erinacea (Hook. f.

& Tayl.) Nees in G. L. & N., Syn. Hep. 624.

1846. Gymnanthe erinacea (Hook. f. & Tayl.)

Mitt, in Hook, f., Bot. Antarc. Voyage 3(2): 230.

1859. Balantiopsis erinacea (Hook. f. & Tayl.)

Mitt, in Hook, f., Handb. New Zealand Fl. 753.

1867. Lectotype (Engel, 1968): Falkland Is.,

Hooker s.n. (NY!).

DESCRIPTION— Engel (1968, 1990).

TAXONOMY — This species is remarkable for the

detachment of the dorsal lobe of the leaf (5,2) and

the dorsal lobe insertion, which appears almost

circular, due to the dilated, sinuate-lobulate mar-

gins at the base of the lobe. The insertion of both

the dorsal and ventral lobes is strongly arched and

U-shaped, with at least one stem cortical cell in-

tervening between the long-decurrent insertion

lines of the lobes (Fig. 2:7,8).

This species was considered the type of a

monotypic sect. Erinacea by Engel (1990), but its

position in the cladogram (Fig. 17) indicates that

it properly belongs in sect. Pteridophylla.

DISTRIBUTION AND ECOLOGY — Engel (1990).

SELECTED SPECIMENS SEEN — CHILE, PROV. CHIL-

OE: Isla Chiloe, Cocauque area, across Estero Yaldad

from village of Yaldad, sea level, Engel 11978 (F).

PROV. VALDIVIA: Valdivia, Isla Teja, Parque de Ex-

posiciones Saval, sea level, Engel 10924 (F).

The Systematic Position of Anisotachis

Schust.

The close relationship between Anisotachis

splendens and Balantiopsis was emphasized by

Stephani (1895) when the species was first pro-

posed. For a full discussion of this problematical

taxon and the history of its interpretation, see

Schuster (1964). Hatcher (1960-61) misinterpret-

ed this plant, treating it as an isolated element

within the genus Isotachis (cf. his phylogenetic

diagram, I.e., fig. 764) and the type and only spe-

cies of his Isotachis sect. Pteridophylla. Engel

(1968, p. 91) emphasized tiering of cells, complex

leaf insertion, and conduplicately bilobed succu-

bous leaves in this species as strongly indicative

of a close relationship to Balantiopsis but stopped

short of making the formal transfer of the species

because of the absence of reproductive structures.

No support was found in the cladogram for

maintaining Anisotachis as a distinct genus. Rath-

er, A. splendens is consistently resolved as a mem-

ber of Balantiopsis sect. Pteridophylla, under-

scored by the strongly U-shaped insertion line of

both the ventral and dorsal lobes (Schuster, 1972,

54

FIELDIANA: BOTANY

fig. 5:4, 6), plication of the ventral lobe (I.e., fig.

5:1, 11, 12), and pronounced tiering of cells (I.e.,

fig. 5:7), all features shared with its sister taxa B.

rosea and B. verrucosa. This species most closely

resembles B. verrucosa in its sparsely armed

leaves and underleaves, dorsal and ventral lobes

subequal in size, rounded to truncate-retuse lobe

apices (cf. A. splendens, Hassel de Menendez &

Solari, 1975, p. 163; B. verrucosa, Figs. 2:9; 6:3,

4), and retuse-bidentate to shallowly bifid under-

leaves (cf. Hassel de Menendez & Solari, fig. 37:

1, 4; Figs. 2:9; 6:7). Smooth leaf cuticle (19,2) is

a homoplasy, occurring here and in the clade con-

taining Austroscyphus and Eoisotachis.

The following new combination is required:

Balantiopsis splendens (Steph.) Engel & Mem,

comb. nov.

Isotachis splendens Steph., Hedwigia 34: 49. 1895.

Anisotachis splendens (Steph.) Schust., Nova

Hedwigia 8: 282. 1964. Original material: Chile,

Isla Desolacion, Bahia Tuesday, Cunningham

(non vidi).

DESCRIPTION — Hassel de Menendez & Solari

(1975), Hatcher (1960-61), Schuster (1964,

1972).

The aspect of this species is notable for the

striking resemblance in size and shape of the leaf

lobes to the underleaves. The deeply bilobed na-

ture of the leaves is often not evident without

careful dissection, and in situ the "leaves" often

appear 5-ranked.

DISTRIBUTION AND ECOLOGY — A species of the

Magellanian moorland, a region occurring south

of 48°S to Cabo de Hornos, and limited by the

Magellanian rain forest to the east and Pacific

Ocean to the west. The moorland is characterized

by being extremely wet, with a permanently sat-

urated peaty soil, and with only scattered and dis-

continuous forests chiefly on coastal fringes or in

sheltered gullies (see Godley, 1960; Engel, 1978).

The species occurs in wet depressions and pools,

where it is often submerged, and ranges from Ti-

erra del Fuego northward in the wet Patagonian

Channel region to 50°39'S (Isla Juan).

SELECTED SPECIMENS SEEN— CHILE, PROV. MA-

GALLANES: Isla Desolacion, Puerto Churruca, head of

Brazo Lobo, Engel 5845B (F, MSC); Isla Desolacidn,

Puerto Churruca, peninsula on N side of Fondeadero

Nassau, Engel 5909 (F, MSC); Isla Desolacion, Bahia

Tuesday, head of inner harbor, Engel 5779 (F, MSC); E

side of Puerto Bueno, Engel 5612 (F, MSC); E side of

Isla Juan, Bahia Wide, Engel 5262 (F, MSC).

The Systematic Position of

Isotachis subg. Hypoisotachis Schust.

The sole species of this monotypic subgenus,

Isotachis multiceps (Lindenb. & Gott.) Gott., is

discussed and illustrated by Schuster (1978;

1985a, figs. 4,5). We have studied a specimen of

this plant from Brazil (Parque National do Itatiaia,

2140 m, Vitt 21587, F), with both young and ma-

ture perigynia. Some additional comments are in

order.

This species is remarkable for the production

of 9 and often 8 gametangia on short ventral-

intercalary branches, a feature found in no other

member of suborder Balantiopsinae. Terminal,

Frullania-type, vegetative branches are produced,

but sparingly; geotropic, microphyllous stolons

are also present.

Other features are also noteworthy. The leaf in-

sertion is weakly succubous, compared to weakly

to distinctly incubous (or transverse) in Isotachis.

The leaf cells have well-developed intermediate

wall thickenings and are noticeably tiered, al-

though not as regularly as in some Balantiopsis

species. The d bracts are monandrous, as in Aus-

troscyphus, compared to 2-4 per bract in Isotach-

is.

Hypoisotachis has a slender Isotachis-\ike per-

igynium that bears Balantiopsis-like rhizoidal out-

growths from its surface, as well as a few peri-

gynial bractlets inserted on the distal third to half

of the outer perigynium wall. The young perigyn-

ium is broadly ovoid, shallowly plicate, and

strongly tapered toward the ± contracted mouth;

the whole is strikingly perianth-like in appear-

ance. However, a cycle of unfused bractlets is in-

serted just within the mouth of the perigynium

proper, which is pluristratose in the basal half.

Additional unfused bractlets, external to this, are

distinguishable as bractlets by their different are-

olation, being made up of ± regularly quadrate

cells. The capsule is cylindrical and spirally de-

hiscent, but the capsule wall is bistratose (or lo-

cally 3-stratose) as in Ruizanthus and M carneus,

rather than uniformly 3-stratose; the latter is the

apomorphy supporting the Isotachis-Balantiopsis

clade in Figure 16:2, the majority consensus of

the "perianth" data set.

We experimented with including this species in

the data set and scoring the same characters and

character states as in the other taxa (p. 27), in-

cluding the perianth. Thirty-seven most parsimo-

nious trees were obtained, each with a length of

120 steps. In the strict consensus of these trees,

ENGEL & MERRILL: AUSTRAL HEPATICAE. BALANTIOPSIS

55

Hypoisotachis is resolved as the sister taxon to the

Austroscyphus-Eoisotachis clade. Unfortunately,

the most striking feature of this curious plant, the

production of gametangia on ventral-intercalary

branches, is an autapomorphy and thus uninform-

ative in a cladistic analysis. Schuster (1978, p.

240) calls attention to the numerous distinctive

features of Hypoisotachis, remarking that "per-

haps the plant should be placed into an autono-

mous genus." The following nomenclatural

changes are required.

Hypoisotachis (Schust.) Engel & Mem, comb. &

stat. nov.

Isotachis subg. Hypoisotachis Schust., Phytologia 39:

240. 1978.

Type: Isotachis coilophylla Herz.

Hypoisotachis multiceps (Lindenb. & Gott.) En-

gel & Merr., comb. nov.

Jungermannia multiceps Lindenb. & Gott. in G. L. &

N., Syn. Hep. 687. 1847. Isotachis multiceps

(Lindenb. & Gott.) Gott., Mexikanske Lever-

moser 105. 1863. Original material: Mexico,

"apud Sempoaltepec, Talea et San Jago Ama-

tlan" (not seen).

Isotachis coilophylla Herz., Feddes Repert. 21: 25.

1925. Original material: Brazil, Serra dos Or-

gaos, Morro Assu, Luetzelburg 6035b (not seen).

DISTRIBUTION — Mexico, Costa Rica, Antilles

(Guadeloupe, Jamaica, Puerto Rico), northern

South America (Colombia, Venezuela), and south-

east Brazil.

Hypoisotachis is strongly anisophyllous but has

evolved an erect stem perigynium of the Isotachis

type. The relationship of Hypoisotachis to Iso-

tachis is unclear: the latter includes both radial,

subisophyllous species (e.g., I. intortifolia} and

prostrate, strongly anisophyllous forms (e.g., /.

minima, Fig. 11:1). Schuster (1972, p. 355) sug-

gests that the erect, fleshy stem perigynium

evolved in plants with an erect radial habit of

growth. The question of which habit is plesio-

morphic in the genus is largely unsettled, except

for the indication that subisophylly in /. intorti-

folia is derived (p. 44).

Phytogeography and Ancestral Areas

An ancestral area analysis was undertaken as

part of this study in an attempt to explain the pres-

ent geographical distribution of Balantiopsis and

the Balantiopsaceae (Fig. 1). An area cladogram

is shown in Figure 18, based on the consensus

cladogram (Fig. 17) of the four most parsimoni-

ous trees obtained in the core phylogenetic anal-

ysis. The area cladogram suggests that the South

American continent was the ancestral area for the

Balantiopsaceae, since the oldest, most plesio-

morphic lineages (Ruizanthus, Neesioscyphus) are

South American. Within Balantiopsis, the earliest

divergent lineages (subg. Steereocolea, B. pur-

puratd) are also South American. However, both

sections of subg. Balantiopsis contain Austral-

asian as well as South American species. Several

examples of amphi-Pacific species pairs in Balan-

tiopsis are indicated in the area cladogram; of

these, Balantiopsis convexiuscula (Australia, Tas-

mania, New Zealand) and B. cancellata (southern

South America) are resolved as sister taxa in all

four most parsimonious cladograms (Fig. 15), as

are B. splendens (southern South America) and B.

rosea and B. verrucosa, a New Zealand species

pair. Two of the three tropical species of the genus

(B. brasiliensis and B. philippinensis) are also

consistently resolved as sister taxa.

Reconstructions of past distributions of anti-

podal taxa rely heavily on assumptions drawn

from vicariance biogeography. Groups showing

amphi-Pacific distribution patterns of this type are

usually regarded as "Gondwanalandic" — the

most recent common ancestor is assumed to have

been widely distributed in Gondwanaland, with

fragmentation of the ancestral range as a result of

the breakup of the supercontinent, and subsequent

speciation. Dispersal is invoked to explain pre-

sumably more recent migration of descendants to

lower latitudes (e.g., to New Guinea, Borneo, and

the Philippines, and to the northern Andes). In

other words, the ancestor was more widely dis-

tributed than its descendants, and the direction of

dispersal was away from the ancestral area in the

south to (often geologically younger) areas to the

north. In the case of Balantiopsaceae, however,

the area cladogram seems to point to a more re-

stricted ancestral area, and the phylogenetically

deepest branches on the cladogram are presently

neotropical, implying migration in the opposite

direction.

Ruizanthus, the most primitive genus in the

family, occurs in high-altitude neotropical para-

mos from northern Peru to Colombia and Vene-

zuela and extending into Central America (Grad-

stein et al., 1994). The distribution of Neesioscy-

phus extends from the northern Andes and the

56

FIELDIANA: BOTANY

BALANTIOPSACEAE

Area

Temp. So. America

Australasia

Neotropics

Paleotropics

Steps RP

11 0. 90

11 0. 90

10 1 . 00

12 0.83

BALANTIOPSIS

Area

Temp. So. America

Australasia

Neotropics

Paleotropics

Steps RP

8 1. 00

9 0. 89

10 0.80

12 0.66

FIG. 18. Area cladogram showing the distribution of each terminal taxon in the data set (based on Fig. 17) with

the geographical distribution substituted for the name of the taxon (A = Australasia; N = Neotropics; P = Paleo-

tropics; S = Temperate South America). The tables give the number of steps on the cladogram required to account

for the distribution of the terminal taxa and the reversible parsimony (RP) indices (Ronquist, 1994) for each of the

putative ancestral areas. Values are given for Balantiopsaceae and for the Balantiopsis clade. (The RP index is the

reciprocal of the number of steps for a given ancestral area, multiplied by the smallest number of steps for any area

on the cladogram.) Bars show the transformation from the "plesiomorphic" South American "state" to Australasia

(1>0), according to a DELTRAN reconstruction of this "character" (see text).

Antilles to Brazil and northern Argentina (Reiner-

Drehwald & Drehwald, 1995). The possibility that

these taxa (and hence the family as a whole) had

an indigenous neotropical origin, however, seems

inconsistent with the classic Gondwanalandic pat-

tern shown by the two largest genera in the family

(Balantiopsis, Isotachis), which have their great-

est diversity in both southern South America and

Australasia (cf. Schuster, 1990, p. 250). Further,

at least two elements in the family (Austroscy-

phus, Eoisotachis) are restricted to the Austral-

asian sector of former Gondwanaland.

We approached this problem in two ways, the

first using the ancestral character state function of

PAUP (see p. 44), and the second employing the

"reversible parsimony" method proposed by

Ronquist (1994). In addition to morphological

characters, the geographical distribution of each

taxon was entered as part of the data set, although

this "character" was excluded when the PAUP

search was run. Geographical distribution was

treated as a binary character: Australasia (0) or

South America (1). A DELTRAN reconstruction

of this "character" was then obtained, showing

the ancestral "state" at each of the internal nodes

within the tree. The "plesiomorphic" state is

South America, changing to Australasia (1 > 0)

at two points (indicated in Fig. 18), thus support-

ing a South American ancestral area for the Bal-

antiopsaceae as well as for Balantiopsis.

Ronquist (1994) proposed the method of re-

versible parsimony for determining ancestral ar-

eas, based solely on the number of steps on an

area cladogram necessary to account for the pres-

ent distribution of the terminal taxa. It should be

emphasized that the method does not examine

"the possibility that distribution areas may be his-

torically related to each other" (I.e., p. 273) and

is explicitly performed outside the context of vi-

cariance biogeography (see critique in Lavin &

Sousa, 1995, pp. 30-32). Ancestors are assumed

to have been no more widespread than their de-

scendants, and the direction of dispersal events is

assumed to be reversible. This is analogous to

treating morphological characters as unordered

(see p. 27) to avoid making a priori assumptions

as to polarity. The area that requires the smallest

ENGEL & MERRILL: AUSTRAL HEPATICAE. BALANTIOPSIS

57

number of steps is the one most likely to have

been the ancestral area of the taxon.

For the reversible parsimony analysis we dis-

tinguished four putative ancestral areas — temper-

ate South America, Australasia, the Neotropics,

and the Paleotropics. The results for each of these

areas (Fig. 1 8) support a neotropical origin for the

Balantiopsaceae (i.e., in the present neotropical

sector of former Gondwanaland), but also show

only a slightly lower probability (RP = 0.90) of

a southern South American or Australasian origin.

All three of these areas are historically related, so

the most likely explanation remains that the over-

all distribution of the family is due largely to the

breakup of formerly continuous areas. Even if

parsimony does not provide conclusive evidence

as to the area in which the Balantiopsaceae ulti-

mately originated, it is at least strongly suggested

that much of the earliest evolution and diversifi-

cation in the family took place in the South Amer-

ican sector of Gondwanaland, simply because so

many basal splits occurred there.

The picture is complicated by the fact that the

modern representatives of these earliest divergent

lineages occur in high-altitude habitats that only

recently came into existence as a result of the up-

lift of the Andean cordillera in the late Tertiary.

Schuster (1985a, p. 52), in discussing the distinc-

tive high-elevation flora of the northern Andes,

argues that "few taxa — indeed a very small mi-

nority— are clearly derived from subantarctic or

Laurasian-temperate antecedents; the majority,

clearly, are derived from forest-zone, neotropical

ancestors." Nevertheless, we think it is likely that

these taxa (Ruizanthus, Neesioscyphus) belong to

the group of high-altitude neotropical bryophytes

of austral origin, which have extended their range

northward along the cordillera in relatively recent

times, many of which also have outliers in south-

eastern Brazil.1 Ruizanthus is said to be "highly

1 For example, Pseudocephalozia quadriloba (Steph.)

Schust. belongs to a nearly exclusively cool- to cold-

temperate Gondwanalandic Lepidoziaceae subfam.

Zoopsidoideae but extends northward at high elevations

to Costa Rica (Schuster & Engel, 1974). Clasmatocolea

is exclusively south temperate-subantarctic in range

with the exception of C. vermicularis (Lehm.) Grolle,

which extends north in the neotropics via the high Andes

to Costa Rica and north at higher elevations in Africa

to Burundi. The species is frequently fertile. The moss

genus Polytrichadelphus (C. Mull.) Mitt, has its major

center of species diversity in the paramos of the northern

Andes, with one species (P. costaricensis Bartr.) in Cen-

tral America and another (P. pseudopolytrichum (Raddi)

G. Sm.) in southeast Brazil (Smith, 1971, fig. 59). How-

fertile" (Gradstein et al., 1994, p. 75) and hence

may be readily dispersable.

The most probable ancestral area for Balan-

tiopsis is temperate South America (Fig. 18), a

result that is consistent with the present distribu-

tion of representatives of the earliest divergent lin-

eages (subg. Steereocolea, B. purpuratd) in the

genus. We are postulating an ancestral area for

Balantiopsis in the sector of Gondwanaland that

included present south temperate South America

and possibly adjacent Antarctica. The genus has

only a limited extension northward to Brazil (B.

brasiliensis) and is lacking in the Andes. Both

sections of subg. Balantiopsis, however, contain

Australasian as well as American species, indi-

cating that the most recent ancestors of these taxa

were probably more widely distributed and that

the evolution of sectional diversity in subg. Ba-

lantiopsis took place before the breakup of Gond-

wanaland.

Austroscyphus is exclusively Australasian, ex-

tending from New Zealand and Tasmania north to

New Caledonia (A. i\vatsukii) and Java (A. tjiwi-

deiensis (Sande-Lac.) Schust.; for treatment see

Engel, 1980). Eoisotachis is also limited in dis-

tribution to cold-temperate Australasia (E. ste-

phanii of extreme southern South Is., and Stewart

Is., New Zealand, and E. nigella (Herz.) Schust.

of Tasmania). The area cladogram (Fig. 18) shows

the Austroscyphus-Eoisotachis-Isotachis clade as

an Australasian sister group to Balantiopsis. This

is obviously an oversimplification, however, since

the only Isotachis species included in the analysis

are Australasian. Schuster (1985b, 1990) places

Austroscyphus in the Ruizanthoideae. In our clas-

sification (see discussion on pp. 47-49), Austro-

scyphus is placed with Neeioscyphus in the Neeio-

scypheae, which (if borne out by future study)

gives this tribe an amphi-Pacific distribution. It

should be emphasized, however, that the cladistic

analysis presented in this paper does not support

this placement of Austroscyphus. The neotropical

Hypoisotachis, with a distribution similar to that

of Neeioscyphus, may be the sister taxon to the

ever, the genus is also represented in southern South

America (P. magellanicus (Hedw.) Mitt.), in Australia

and New Zealand (P. innovans (C. Mull.) Jaeg.), and

in New Guinea (P. archboldii Bartr.). Notoligotrichum

G. Sm. has a similar distribution (I.e., fig. 77), extend-

ing northward to Mexico in the New World and to New

Guinea (Hyvonen, 1986). These taxa represent exten-

sions of the "Antarctic radiant" pattern proposed by

Smith (1971, 1972).

58

FIELDIANA: BOTANY

Australasian Austroscyphus-Eoisoitachis clade

(see above).

Isotachis, the most widely distributed genus of

Balantiopsaceae, occurs in the neotropics, south-

ern South America, Africa and Madagascar, tem-

perate Australasia, and north in the Old World to

Japan (Fig. 1). Unlike Balantiopsis, Isotachis has

a major center of diversity in tropical America

(see key in Schuster, 1985a); Balantiopsis, how-

ever, has only one representative in the neotropics

(B. brasiliensis, in southeastern Brazil). There are

ten species of Isotachis in the neotropics, five in

southern South America, seven in temperate Aus-

tralasia; three are Indo-Malayan (/. japonica

Steph. extends north to southernmost Japan) and,

according to Vana (1982), only one occurs in Af-

rica (and that species, /. aubertii (Schwaegr.)

Mitt., is a tropical alpine species also occurring in

tropical America; Gradstein, Pocs & Vana, 1983).

The poor representation of Isotachis in Africa

suggests that the occurrence of the genus there is

relatively recent. The total distribution of Isota-

chis (as opposed to that shown in the area clado-

gram) suggests a history similar to that of Ba-

lantiopsis, with major diversification occurring

prior to the breakup of Gondwanaland.

Acknowledgments

The senior author gratefully acknowledges the

support of the National Science Foundation

(grants BMS76-03616 and DEB-8 109680), which

funded field work in New Zealand, Tasmania, and

Australia. The senior author also gratefully ac-

knowledges support of the National Geographic

Society, which supported field studies in southern

Chile (grant 1527) and New Zealand (grant 5375-

94). The authors thank Marlene Werner and Zor-

ica Dabich for preparing the habitus figures for

the illustrations, and Drs. Peter Crane and Greg-

ory Mueller for advice and for reading the manu-

script. We also thank the following individuals

and institutions for the loan of specimens: Dr. P.

J. Brownsey and Ms. Fiona D. H. Pitt (WELT), Dr.

D. H. Pfister (FH), Dr. Per Lassen (LD), and Dr.

Barbara Thiers (NY).

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ENGEL & MERRILL: AUSTRAL HEPATICAE. BALANTIOPSIS

61

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.

Anisotachis Schust. 2, 54

Anisotachis splendens (Steph.) Schust. 55

Austroscyphus iwatsukii (Kitag.) Schust. 58

Austroscyphus phoenicorhizus (Grolle) Schust. 27, 30

Austroscyphus tjiwideiensis (Sande-Lac.) Schust. 58

Balantiopsaceae

Subfam. Ruizanthoideae Schust. ex Engel &

Mem 48

Subfam. Isotachidoideae (Hatch.) Grolle 48

Tribe Neesioscypheae Engel & Merr. 48

Subtribe Isotachinae (Hatch.) Engel & Merr. 48

Balantiopsis Mitt. 2, 49

Subg. Balantiopsis 5, 51

Subg. Schizophyllon Hatt. 5

Subg. Steereocolea (Schust.) Schust. 49

Sect. Balantiopsis 5, 52

Sect. Erinacea Engel 16

Sect. Pteridophylla (Hatch.) Engel & Merr. 16,

53

Balantiopsis aequiloba Berggr. 21

Balantiopsis asymmetrica (Herz.) Engel 50

Balantiopsis bisbifida (Steph.) Steph. 50

Balantiopsis cancellata (Nees) Steph. 53

Balantiopsis ciliaris Hatt. 53

Balantiopsis convexiuscula Berggr. 21

Balantiopsis diplophylla (Hook. f. & Tayl.) Mitt. 5

Balantiopsis diplophylla var. diplophylla 9

Balantiopsis diplophylla var. hockenii (Bergrr.) En-

gel & Merr. 1 1

Balantiopsis erinacea (Hook. f. & Tayl.) Mitt. 54

Balantiopsis glandulifera Col. 12

Balantiopsis hockenii Berggr. 1 1

Balantiopsis knightii Mitt. 1 1

Balantiopsis latifolia var. asymmetrica Herz. 50

Balantiopsis lingulata Schust. 14

Balantiopsis montana (Col.) Engel & Merr. 12

Balantiopsis purpurata Mitt. 52

Balantiopsis rosea Berggr. 17

Balantiopsis splendens (Steph.) Engel & Merr. 55

Balantiopsis tumida Berggr. 24

Balantiopsis verrucosa Engel & Merr. 16

Chiloscyphus heterodontus Col. 9

Chiloscyphus montanus Col. 12

Eoisotachis nigella (Herz.) Schust. 58

Eoisotachis stephanii (Salm.) Schust. 27, 58

Gottschea diplophylla (Hook f. & Tayl.) Nees 9

Gottschea erinacea (Hook. f. & Tayl.) Nees 54

Gymnanthe diplophylla (Hook f. & Tayl.) Mitt. 9

Gymnanthe erinacea (Hook. f. & Tayl.) Mitt. 54

Hypoisotachis (Schust.) Engel & Merr. 56

Hypoisotachis multiceps (Lindenb. & Gott.) Engel &

Merr. 56

Isotachidaceae Hatch. 48

Isotachis

Subg. Hypoisotachis Schust. 56

Sect. Pteridophylla Hatch. 16

Isotachis aubertii (Schwaegr.) Mitt. 59

Isotachis bisbifida Steph. 50

Isotachis coilophylla Herz. 56

Isotachis intortifolia (Hook. f. & Tayl.) Gott. 27

Isotachis japonica Steph. 59

Isotachis lyallii (Hook. f. & Tayl.) Mitt. 27, 38

Isotachis minima Pears. 31

Isotachis montana Col. 27, 32, 36

Isotachis multiceps (Lindenb. & Gott.) Gott. 56

Isotachis splendens Steph. 55

Jungermannia diplophylla Hook. f. & Tayl. 9

Jungermannia erinacea Hook. f. & Tayl. 54

Jungermannia multiceps Lindenb. & Gott. 56

Neesioscyphus argillaceus (Nees) Grolle 27

Neesioscyphus carneus (Nees) Grolle 27

Ptilidium cancellatum 53

Ruizanthus venezuelanus Schust. 27

Steereocolea Schust. 2, 49

Steereocolea bisbifida (Steph.) Schust. 50

62

FffiLDIANA: BOTANY

A Selected Listing of Other Fieldiana: Bota?

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Publication 14!

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