FIELDIANA

Botany

NEW SERIES, NO. 44

Austral Hepaticae. 35. A Taxonomic and Phylogenetic

Study of Telaranea (Lepidoziaceae), with a

Monograph of the Genus in Temperate Australasia

and Commentary on Extra-Australasian Taxa

John J. Engel

G.L. Smith Merrill

November 30, 2004

Publication 1531

PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY

DEC 0 8 2004

UBRAIU

FIELDIANA

Botany

NEW SERIES, NO. 44

Austral Hepaticae. 35. A Taxonomic and

Phylogenetic Study of Telaranea (Lepidoziaceae),

with a Monograph of the Genus in

Temperate Australasia and Commentary

on Extra-Australasian Taxa

John J. Engel

G. L. Smith Merrill

Department of Botany

The Field Museum

1400 South Lake Shore Drive

Chicago, Illinois 60605-2496

Accepted June 27, 2003

Published November 30, 2004

Publication 1531

PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY

© 2004 Field Museum of Natural History

ISSN 0015-0746

PRINTED IN THE UNITED STATES OF AMERICA

Tabl

able of Contents

ABSTRACT ............................................... 1

INTRODUCTION .......................................... 1

Evolution of a Generic Concept ............... 2

Telaranea Spruce ex Schiffn ................... 5

THE GENUS TELARANEA IN AUSTRALASIA ......... 6

Key to Australasian Taxa of Telaranea ..... 6

Taxonomic Treatments .......................... 11

DISCUSSION OF EXTRA-AUSTRALASIAN

SPECIES .......................................... 120

Key to Telaranea Species of Temperate

South America ............................. 120

SYSTEMATICS ......................................... 201

Choice of Taxa .................................. 202

Character Descriptions and Coding ........ 202

Results ............................................. 220

Character-State Reconstructions ............ 224

Discussion of Phylogenetic Relationships . . 232

Synoptic key to the subgenera and sec-

tions of Telaranea ..................... 236

Key to species of sect. Tenuifoliae ..... 24 1

The status of Arachniopsis subg. Mono-

dactylopsis Schust ..................... 241

The systematic position of Arachniop-

sis "sect. Dissotrichae" ............. 242

PHYTOGEOGRAPHY AND ANCESTRAL AREAS ... 249

TELARANEA EXCLUDENDA .......................... 254

ACKNOWLEDGMENTS ................................ 255

LITERATURE CITED .................................. 256

INDEX TO TAXA ..................................... 262

List of Figures

1. Telaranea clatritexta (Steph.) Engel &

Mem ............................................... 14

2. Telaranea meridiana (Hodgs.) Hodgs ..... 16

3. Telaranea capilligera (Schwaegr.)

Schust ............................................. 19

4. Telaranea pennata Engel & Merr. ........ 21

5. Telaranea quadristipula (Steph.) Engel

& Merr. ........................................... 23

6. Telaranea tridactylis (Lehm. & Lin-

denb.) Engel & Merr. ......................... 26

7. Telaranea tridactylis (Lehm. & Lin-

denb.) Engel & Merr. ......................... 27

8. Telaranea consobrina Engel & Merr. .... 30

9. Telaranea palmata Engel & Merr. ........ 33

10. Telaranea praenitens (Lehm. & Lin-

denb.) Fulf. var. praenitens + var. den-

tifolia Engel & Merr. ......................... 36

11. Telaranea glbbsiana (Steph.) Hodgs ..... 40

12. Telaranea gibbsiana (Steph.) Hodgs ..... 41

13. Telaranea grossiseta (Steph.) Engel &

Schust 42

14. Telaranea grossiseta (Steph.) Engel &

Schust. 43

15. Telaranea hodgsoniae Engel & Merr. .... 44

16. Telaranea patentissima (Hook. f. &

Tayl.) Hodgs. var. patentissima + var.

ampliata Engel & Merr. 49

17. Telaranea patentissima var. zebrina

Engel & Merr. + var. patentissima 51

18. Telaranea tetrapila (Hook. f. & Tayl.)

Engel & Merr. var. tetrapila 55

19. Telaranea tetrapila var. roseana

(Steph.) Engel & Merr. + var. cancel-

lata (Col.) Engel & Merr. -I- Telaranea

paludicola (Hodgs.) Hodgs 56

20. Telaranea elegans (Col.) Engel &

Merr. (1-6) + T. perfragilis Engel &

Merr. 63

21. Telaranea centipes (Tayl.) Schust 66

22. Telaranea centipes (Tayl.) Schust 68

23. Telaranea tuberifera Engel & Schust 69

24. Telaranea remotifolia Hodgs : 75

25. Telaranea pulcherrima (Steph.) Schust.

var. pulcherrima + var. mooreana

(Steph.) Engel & Merr. 78

26. Telaranea lindenbergii (Gott.) Engel &

Merr. + Telaranea tetradactyla (Hook.

f. & Tayl.) Hodgs 82

26A. Telaranea lindenbergii var. papillata

Engel & Merr. 84

27. Telaranea tetradactyla (Hook. f. &

Tayl.) Hodgs 93

28. Telaranea quadriseta (Steph.) Engel &

Merr. 97

29. Telaranea martinii (Hodgs.) Schust 100

30. Telaranea martinii (Hodgs.) Schust 101

31. Telaranea granulata Engel & Merr. .... 104

32. Telaranea tasmanica (Steph.) Engel &

Merr. 108

33. Telaranea tasmanica (Steph.) Engel &

Merr. 109

34. Telaranea quinquespina (Engel &

Merr.) Engel & Merr. 110

35. Telaranea herzogii (Hodgs.) Hodgs 114

36. Telaranea herzogii (Hodgs.) Hodgs 116

37. Telaranea inaequalis Schust. tex Engel

& Merr. 119

38. Telaranea bicruris (Steph.) Howe +

Telaranea nematodes (Gott. ex Aust.)

Howe 126

39. Telaranea blepharostoma (Steph.) Fulf. 130

40. Telaranea chaetocarpa (Pears.) Grolle .. 133

41. Telaranea chaetocarpa (Pears.) Grolle .. 134

in

42. Telaranea chaetophylla (Spruce)

Schiffn 137

43. Telaranea chaetophylla (Spruce)

Schiffn 138

44. Telaranea coactilis (Spruce) Engel &

Mem 141

45. Telaranea coactilis (Spruce) Engel &

Mem 142

46. Telaranea diacantha (Mont.) Engel &

Mem 144

47. Telaranea disparata Engel & Mem .... 148

48. Telaranea europaea Engel & Mem. .... 152

49. Telaranea europaea Engel & Mem .... 154

50. Telaranea longifolia (Howe) Engel &

Mem 156

51. Telaranea longifolia (Howe) Engel &

Mem 157

52. Telaranea neesii (Lindenb.) Fulf. 170

53. Telaranea nematodes (Gott. ex Aust.)

Howe 172

54. Telaranea oligophylla (Lehm. & Lin-

denb.) Engel 175

55. Telaranea plumulosa (Lehm. & Lin-

denb.) Fulf 182

56. Telaranea plumulosa (Lehm. & Lin-

denb.) Fulf 183

57. Telaranea sejuncta (Angstr.) S. Arnell .. 188

58. Telaranea trichocoleoid.es (Herz.)

Schust 195

59. Telaranea trisetosa (Steph.) Grolle 196

60. Telaranea verruculosa Engel & Mem .. 198

61. Kurzia hippuroides (Hook. f. & Tayl.)

Grolle 203

62. Kurzia hippuroides (Hook. f. & Tayl.)

Grolle 204

63. Lepidozia obtusiloba Steph 205

64. Lepidozia obtusiloba Steph 206

65. Lepidozia obtusiloba Steph 207

66. Lepidozia spinosissima (Hook. f. &

Tayl.) Mitt 208

67. Triandrophyllum symmetricum Engel .. 210

68. Triandrophyllum symmetricum Engel . . 211

69. SEM micrographs of spores (Telaranea

chaetophylla, Telaranea diacantha, Te-

laranea lindenbergii, Telaranea longi-

folia) 217

70. SEM micrographs of spores (Telaranea

nematodes, Telaranea europaea, Telar-

anea tuberifera) 218

7 1 . SEM micrographs of spores (Telaranea

tridactylis, Telaranea patentissima, Te-

laranea tetrapila) 219

72. SEM micrographs of spores (Telaranea

plumulosa, Telaranea coactilis, Telara-

nea pallescens) 220

72A. Oil-bodies 221

73. Strict consensus of 485,487 most parsi-

monious trees 222

74. Majority Rule consensus of 485,487

most parsimonious trees 223

75. Best tree from 10,000 replicates 225

76. Cladograms obtained from subsets of

data 226

77. Strict consensus of 306 most parsimo-

nious trees 227

78. Unrooted array based on strict consen-

sus of 306 trees 228

79. Distance tree based on total character

distance 233

80. Comparison of classifications, I 234

81. Comparison of classifications, II 235

82. Amazoopsis diplopoda (Pocs) Engel &

Mem 243

83. Amazoopsis diplopoda (Pocs) Engel &

Mem 244

84. Amazoopsis gracilis Engel & Mem 248

85. Amazoopsis dissotricha (Spruce) Engel

&Mem 250

86. Area cladogram 252

List of Tables

1. Taxa included in systematic analysis 212

2. Data matrix used in cladistic analysis of

Telaranea and outgroup taxa 213

3. Conspectus of Telaranea Spruce ex

Schiffn 253

4. Geographical distribution of major taxa

of Telaranea and numbers of species in

each region 254

IV

Austral Hepaticae. 35. A Taxonomic and Phylogenetic

Study of Telaranea (Lepidoziaceae), with a

Monograph of the Genus in Temperate Australasia

and Commentary on Extra-Australasian Taxa

John J. Engel and G. L. Smith Merrill

Abstract

Telaranea is the third largest genus of Lepi-

doziaceae, after Bazzania (over 450 species) and

Lepidozia (over 300 species). In this monograph,

98 species of Telaranea are recognized, 62 of

which are extra-Australasian. Included are 1 1 new

species, 2 new varieties, 27 new combinations (25

species and one variety), and one new name. New

taxa are fully described and illustrated. Two sub-

genera of Telaranea are recognized (subg. Acro-

lepidozia, subg. Telaranea), and 7 sections within

subg. Telaranea (sect. Neolepidozia (Fulf. & J.

Tayl.) Engel & Merr. comb, nov., sect. Cancella-

tae Engel & Merr., sect. Ceraceae Engel & Merr.

sect, nov., sect. Tricholepidozia (Schust.) Engel &

Merr. comb, nov., sect. Transversae Engel &

Merr., sect. Telaranea, sect. Tenuifoliae (Schust.)

Engel & Merr. comb. nov.).

A major part of this work is devoted to a phy-

logenetic study of the genus Telaranea, with a

discussion of character evolution in the genus and

a reconstruction of ancestral character states. An

ancestral area analysis was also undertaken, using

phylogeny to trace the geographical history of

members of the genus. Differing concepts of phy-

logeny as applied to leafy hepatics are also briefly

considered. The phylogenetic analysis included 56

taxa and 32 characters, with representative species

of Lepidozia, Kurzia, and Arachniopsis as out-

groups. Cladograms obtained from parsimony

analyses of the full data set and subsets of taxa,

as well as distance (NJ) trees, were strikingly sim-

ilar in topology. Monophyly of Telaranea is sup-

ported by the possession of a hyaloderm (en-

larged, usually hyaline stem cortical cells) and an

undivided first branch underleaf. Progressive re-

duction in morphological complexity as a central

evolutionary tendency in the genus is strongly

supported. The phylogeny indicates that Kurzia,

as defined by the presence of Microlepidozia-iype

branching, is polyphyletic, and 6 species previ-

ously assigned to Kurzia are transferred to Telar-

anea. No support was found for maintaining Ar-

achniopsis as a distinct genus, and 6 species of

the genus are transferred to Telaranea. The new

genus Amazoopsis Engel & Merr. is described,

with 3 species: A. diplopoda (Pocs) Engel & Merr.

comb, nov., A. dissotricha (Spruce) Engel & Merr.

comb, nov., and A. gracilis Engel & Merr. sp. nov.

The status of Monodactylopsis (Schust.) Schust.

is discussed; the genus includes M. monodactyla

(Spruce) Schust. and M. minima Schust. ex Engel

& Merr., sp. nov. Paracromastigum vastilobum

(Steph.) Engel & Merr. is a new combination.

Keys to the subgenera and sections of Telaranea,

the Australasian species, as well as the southern

South American species of Telaranea, and the

species of Telaranea sect. Tenuifoliae are includ-

ed.

Introduction

Until the mid-1950s, the genus Telaranea was

frequently understood narrowly to include only

the type, T. chaetophylla (Spruce) Schiffn., usu-

ally treated as a synonym of either T. nematodes

(Gott. ex Aust.) Howe or T. sejuncta (Angstr.) S.

Arnell. With the inclusion of other elements, how-

ever, the genus now ranks third among the genera

of the family Lepidoziaceae, after Bazzania, with

over 450 described species and Lepidozia, with

well over 300. Kurzia, with 32-36 taxa (Schuster,

1980) is the next largest genus.

FIELDIANA: BOTANY, N.S., NO. 44, November 30, 2004, PP. 1-265

As treated here, Telaranea (incl. Arachniopsis)

comprises 98 species in both the Northern and

Southern Hemispheres. The greatest diversity in

numbers of species is in western Melanesia and

Australasia. Temperate Australasia, and in partic-

ular New Zealand, has the largest number of spe-

cies and the greatest diversity of distinct morpho-

logical types. Thirty-six species and 8 varieties of

Telaranea are treated here from temperate Aus-

tralasia.

The bulk of the Australasian species now as-

signed to Telaranea were formerly included in the

genus Lepidozia. Hodgson's (1956) treatment of

the New Zealand species of Lepidozia is the only

revision of the Australasian Telaranea taxa to

date. Fourteen of the 33 Lepidozia species treated

by Hodgson are now placed in Telaranea. Fulford

and Taylor (1959) removed 16 species from Lep-

idozia and placed them in a new genus Neolepi-

dozia, with Jungermannia capilligera Schwaegr.

as its type. Hodgson (1962) treated Neolepidozia

as a synonym of Telaranea and transferred 10

Australasian species of the Neolepidozia group to

Telaranea; Schuster (1963) transferred seven

more regional species to Telaranea. Engel and

Merrill (1996a), added four sections (sect. Can-

cellatae, sect. Capillares, sect. Latifoliae, and

sect. Transversae), two new species (T. hodgson-

iae, T. pennata), three new varieties (T. linden-

bergii var. complanata, T. lindenbergii var. mel-

lea, and T. praenitens var. dentifolia) and nine

new combinations (T. complanata, T. elegans, T.

lindenbergii, T. paludicola, T. tasmanica, T. te-

trapila, T. tetrapila var. roseana, T. tetrapila var.

cancellata, and T. tridactylis), all from Australa-

sia. Engel and Merrill (1999) described three ad-

ditional species from the region, T. consobrina, T.

fragilis, and T. palmata, and transferred one Aus-

tralasian species, T. clatritexta, from Lepidozia to

Telaranea.

The results of our study of the Australasian rep-

resentatives of the genus Telaranea are presented

in this monograph. Our conclusions are based on

the examination of numerous specimens collected

by Engel, Child, Hatcher, Hodgson, Allison, and

others, plus all of the relevant type specimens.

The result has been a considerable refinement of

species concepts, and the acceptance of several

additional taxa not previously recognized as be-

longing to Telaranea. The first section of the work

deals with taxonomic treatments of the Austral-

asian species. This is followed by a review of the

extra-Australasian representatives of the genus.

The third portion of the work presents the results

of a phylogenetic analysis of the genus Telaranea.

The monograph concludes with a discussion of

geographical relationships, and a list of excluded

taxa.

Evolution of a Generic Concept

I. Telaranea in the restricted sense

Telaranea first appears as a synonym (Spruce,

1885, p. 360, as "Telaranea nobis nov. gen?")

under subhead 2. of Lepidozia subg. Micro-Lepi-

dozia, based on a single species, Lepidozia chae-

tophylla, with "Telaranea chaetophylla Spruce

Mst nov. gen." listed as a synonym (p. 365). Te-

laranea as a genus name dates from 1893, when

validly published by Schiffner (1893). It remained

essentially monotypic in works published as late

as the 1950's (e.g., Schiffner, 1893; Howe, 1902;

Stephani, 1909). The detailed study of T. longi-

folia (Howe) Engel & Merr. by Schuster and

Blomquist (1955, as T. nematodes) included a dis-

cussion of the limits of the genus and concluded

that "... our present concept of the genus is

based largely upon the genotype [i.e., generitype]

T. nematodes" (p. 592).

The characteristics of Telaranea s. str. are those

of T. chaetophylla, as stated by Spruce: stem cor-

tical cells in 8-10 rows and medulla 4X7 cells

wide; 3-lobed leaves, the lobes 4-7 cells long,

uniseriate (or biseriate) at the base, with their bas-

al cells connate to about Vs their height, half-

leaves almost always undivided and underleaves

2- (rarely 3-) lobed, the lobes erect-incurved, from

a rhizoid-bearing base.

II. Lepidozia sect. Capillares and

Microlepidozia

Karl Miiller (1914) was apparently the first to

expand Telaranea (as Lepidozia subg. Telaranea)

beyond Spruce's original concept to include other

Lepidozia species with large-celled, uniseriate leaf

lobes and transversely inserted leaves. Among the

species included by Miiller were L. blepharosto-

ma, L. lawesii, L. neesiana (T. neesii), L. nema-

todes, L. sejuncta, and L. trisetosa (as well as L.

chaetophylla), all now placed in Telaranea. The

subgenus was not treated by Miiller since it in-

cluded only extra-European taxa. Telaranea nem-

atodes was not reported from Europe until 1936

(Miiller, 1956).

FIELDIANA: BOTANY

Mullet's treatment was foreshadowed by the

Synopsis Hepaticarum (Gottsche, Lindenberg and

Nees, 1845), whose Lepidozia sect. Capillares in-

cluded species with deeply divided leaves, capil-

lary (uniseriate) lobes, and a lacinate-ciliate peri-

anth mouth. Four of the 6 species included in the

section are now placed in Telaranea, namely T.

plumulosa, T. neesii, T. lindenbergii, and T. tetra-

dactyla; the others are Kurzia species (for typifi-

cation of sect. Capillares, see p. 79).

Sect. Capillares and two others were adopted

by Spruce ( 1 876) as subgenera of Lepidozia, the

others (sect. Communes, Microphyllae) having in-

cubous leaves, divided to the middle. Hodgson

(1956, p. 589) decried the fact that "with 30 or

more years of priority, these names to all intents

and purposes have been dropped because Spruce,

in addition to ignoring them, replaced them with

his own 3 sections . . . and placed these on a per-

manently higher level by calling them subgen-

era." Spruce's Lepidozia subg. Microlepidozia

"corresponds nearly to the § Capillares of 'Syn.

Hep.'" (quoting Spruce, 1876, p. 165), and is

"almost distinct enough to form a genus apart."

In addition to deeply-divided leaves and ciliate-

laciniate perianth mouth (characters of sect. Cap-

illares), Spruce adds "leaves transverse, neither

succubous nor incubous" and unistratose peri-

anths.

Telaranea has had a long history of association

with Microlepidozia (Spruce) Joerg. (= Kurzia v.

Mart.), although the two are now generally con-

sidered to be distinct. Miiller (1956) united the

two elements (Telaranea and Microlepidozia), us-

ing Spruce's generic name for the combined ge-

nus, distinguished from Lepidozia by the nearly

transversely inserted leaves, divided up to the

base, with setaceous lobes; unistratose perianth;

seta cross-section with 8 large, outer cells sur-

rounding 4-16 small, inner cells; and 2-, occa-

sionally 3-stratose capsule valves. Oil-bodies, list-

ed as absent by Miiller, are now known to be pre-

sent in both Kurzia and Telaranea species. Schus-

ter and Blomquist (1955) distinguished the genus

Telaranea (sensu Spruce) from Microlepidozia

primarily by the absence of Microlepidozia-iypc

branching in Telaranea, and the presence of a dis-

tinct hyaloderm. Fulford (1963a) also emphasized

Microlepidozia-lype branching as a character dis-

tinguishing Microlepidozia from Telaranea.

Schuster (1969, p. 31) reviewed the distinctions

between Telaranea and Microlepidozia ( = Kur-

zia).

III. Lepidozia sect. Communes and Neolepidozia

Lepidozia sect. Communes G. L. & N. (Gott-

sche, Lindenberg and Nees, 1845, = sect. Lepi-

dozia, since it included the type of the genus, L.

reptans) contained species with leaves having a

higher disc and non-capillary lobes. Six of the 21

included species are now placed in Telaranea,

namely T. capilligera, T. centipes, T. gottscheana,

T. patentissima, T. praenitens, and T. wallichiana.

A seventh species, T. oligophylla, was placed in

sect. Microphyllae by Gottsche, Lindenberg and

Nees.

These and similar, Lepidozia-\'\ke Telaranea

species remained in Lepidozia until Fulford and

Taylor (1959) removed 16 species from the genus

and placed them in a new genus Neolepidozia,

with Jungermannia capilligera as its type. Neo-

lepidozia was distinguished from Lepidozia by

fewer, larger stem cortical cells forming a distinct

hyalodermis, leaves with entire margins, the mar-

gins straight or convex and without auricles at the

base, a much greater uniformity of leaf cell size

than in Lepidozia, and a first branch underleaf

usually undivided and ventral in position. The re-

lationship between Neolepidozia and Telaranea

was not discussed by Fulford and Taylor.

Fulford (1963b, in key, pp. 17-19) further dis-

tinguished Neolepidozia from Lepidozia by leaf

cells tending to be in longitudinal rows and stems

with 12 rows of cortical cells, vs. leaf cells not

necessarily in rows and cortical cells more nu-

merous in Lepidozia. Fulford (1966) added an

oblique, nearly longitudinal line of leaf insertion

and triangular leaf and underleaf lobes as char-

acters of Neolepidozia.

In a review of the Telaranea species in South

America, Fulford (1963a) distinguished the genus

from Lepidozia by a stem with 18 to as few as 6

large cortical cells, leaves transversely inserted,

2-6 uniseriate leaf lobes, and a disc "4, 3, 2, 1

or even Vi (or less) rows of cells high." Included

were T. apiahyna, T. blepharostoma, T. neesii, T.

plumulosa, T. pseudozoopsis, T. sejuncta, and T.

tetradactyla (the last a Neolepidozia in Fulford

and Taylor, 1959), all species "which are closely

related to one another and which show many char-

acters in common with T. sejuncta" (understood

by Fulford as synonymous with T. nematodes).

Fulford's key (1963b, pp. 17-19) separates Telar-

anea from Lepidozia by leaves transversely in-

serted or nearly divided to the base or approxi-

mately one-half their length, uniseriate, "capilla-

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

ceous" leaf lobes, and the margins of the leaves

and underleaves entire.

In summary, Ful ford's concept of Telaranea is

ultimately traceable to Lepidozia sect. Capillares

of the Synopsis Hepaticarum, and subg. Telara-

nea of Miiller (1914). Subsequently, Piippo

(1984) has returned the Lepidozia-\ike species to

the parent genus as Lepidozia sect. Neolepidozia,

consisting of species with almost symmetrical

leaves, leptodermous cells, and a conspicuous hy-

aloderm, while maintaining Telaranea in the more

limited sense of Miiller and Fulford.

IV. Merging of concepts: Telaranea in the

modern sense

Hodgson ( 1 962) treated Neolepidozia as a syn-

onym of Telaranea and transferred 10 Australa-

sian species of the Neolepidozia group to Telar-

anea, without according it subgeneric rank.

Schuster (1963) combined all three elements dis-

cussed above under Telaranea, listing Frullania-

type branching, a stem with a hyaloderm, leaf

cells large and pellucid, leaf lobes narrow (1-2,

or rarely 3-4 cells broad at base), disc 1-3(4)

cells high, capsule-wall 3-stratose (as in Telara-

nea s. str.), and finely areolate spores as charac-

teristics of the combined genus. Schuster recog-

nized 3 subgenera, of which subg. Telaranea is

"exactly equivalent in concept to [the] genus Te-

laranea of Schuster and Blomquist (1955)." Sub-

genus Neolepidozia comprises the other low-disc,

capillary-lobed species of Telaranea in the sense

of Miiller and Fulford, as well as those of Neo-

lepidozia. A third element, subg. Tricholepidozia,

is discussed below.

V. Other new elements: Acrolepidozia,

Tricholepidozia, Chaetozia

Schuster (1963) proposed the genus Acrolepi-

dozia, based on Lepidozia longitudinalis, empha-

sizing a dimorphic mode of growth "quite unlike

any of the New Zealand taxa [of Lepidozia]

known to me." The genus was reduced to a sub-

genus of Telaranea by Schuster (1969, see foot-

note, p. 30), and considered derived from subg.

Neolepidozia. Subgenus Acrolepidozia is de-

scribed and discussed in detail in Schuster (1973,

p. 389), who called attention to the diminuitive

half-leaf and first branch underleaves, and deem-

phasized the mode of growth in this species.

Telaranea subg. Tricholepidozia was proposed

by Schuster (1963) for 4 species with a "Tricho-

colea-\ike fades" and 8-13-lobed leaves, which

had not previously been associated with Telara-

nea. Besides the type, T. mooreana, T. pulcher-

rima, T. radiata, and T. trichocoleoides comprise

the subgenus. Schuster (1969, 1973) added a

weakly differentiated hyaloderm of ca. 24-28

rows of cortical cells, and a 5-stratose capsule

wall as characteristics of this subgenus. Schuster

(2000, p. 240) cites relative subisophylly as the

chief reason for retaining Tricholepidozia as a

subgenus.

Grolle (1966) proposed the new subgenus

Chaetozia, with T. chaetocarpa of New Caledonia

as its type. This species has the distal portion of

the perianth covered with ciliform outgrowths. A

second species, T. trisetosa, was originally in-

cluded in the subgenus by Grolle, but it was later

found not to have this feature (Grolle, 1968) and

was excluded. The subgenus was synonymized

with subg. Telaranea by Schuster (1969).

VI. Telaranea and Arachniopsis

The striking similarity between Telaranea s. str.

and Arachniopsis Spruce (Spruce, 1882) has been

noted since the former was first proposed. Spruce

(1885, p. 366) distinguished Arachniopsis from

Lepidozia chaetophylla, citing exclusively posti-

cal (ventral-intercalary) branching, leaf lobes

quite free from one another (or solitary), and the

lack of underleaves in Arachniopsis. In fact, un-

derleaves are present in all species, but can con-

sist of as few as 2 adjacent cells, each capped by

a slime papilla. In others (A. diacantha) they are

bilobed and caliper-like, resembling those of e.g,

Telaranea herzogii. Likewise, terminal, Frullan-

/a-type branching has since been reported from

many Arachniopsis species. In more recent times,

a 2-stratose capsule wall (already noted by

Spruce, 1885, p. 354), 8+4 seta (Schuster, 1965),

and the proliferation of rhizoids from near the tips

of some leaf lobes (Schuster, 1973, p. 388; illus-

trated by Fulford, 1968, pi. 96, fig. 2h in A. coac-

tilis) have been cited as additional characters dis-

tinguishing Arachniopsis from Telaranea.

Only one species of Telaranea (T. herzogii) has

been transferred formally to Arachniopsis (Hodg-

son, 1964). The supposed lack of terminal branch-

ing in Arachniopsis and the fact that the genus is

rare and poorly represented in herbaria have so

far precluded discussion of combining the two

FIELDIANA: BOTANY

genera, although according to Grolle (1975), the

type of Blepharostoma sejuncta Angstr. (Telara-

nea sejuncta (Angstr.) S. Arnell, a name exten-

sively used in the literature as a synonym of T.

nematodes) is Arachniopsis diacantha (Mont.)

Howe.

VII. Evolutionary speculation

Comments regarding evolutionary development

in Telaranea and related genera are scattered in

the literature and relatively few. Telaranea sens,

str. has been widely regarded as "reduced" from

more structurally complex members of the family,

with Arachniopsis as a "still further departure

along the same lines from the Lepidozia type"

(Howe, 1902, p. 284). Fulford (1963a) likewise

referred to a "drastic reduction which has oc-

curred in the T. sejuncta evolutionary line," evi-

denced by a decrease in the number of rows of

cortical cells in the stem, reduction in leaf size

and complexity (both in number of lobes and in

the height and width of the disc), reduction in the

size of underleaves, weaker development of thick-

ening bands on the inner wall of the capsule, and

a seta with only 8-10 large outer cells.

Fulford (1963a, p. 80) characterized the genus

Telaranea as "probably the most clearly defined

reduction series among the leafy Hepaticae," a

"retrogressive evolutionary series of considerable

scope." The South American T. plumulosa was

identified as the "most primitive condition" by

virtue of its radial symmetry, with similar leaves

and underleaves, and T. sejuncta as "the most re-

duced and most simple species of the series."

Schuster (1965, p. 46) considered Arachniopsis as

derived by loss of terminal branching from Telar-

anea-\ike ancestors.

In some cases, an evolutionary progression is

implied, as, for example, in the diagrams of axial

anatomy in Schuster (1984, p. 813, fig. 15 leg-

end). The leaf of subg. Tricholepidozia (T. pul-

cherrima, fig. 7) is shown arising from a quadrifid

leaf of Telaranea (fig. 6), so that "by an accen-

tuation of this process an 8- or even 1 2-lobed leaf

may ensue." For further discussion see section on

phylogenetic relationships (p. 232).

Familial placement is no longer in dispute, al-

though some authors (Evans, 1939; Fulford, 1968)

formerly placed Arachniopsis in the Cephalozia-

ceae. Verdoorn (1932) listed Arachniopsis as a ge-

nus of Trigonanthaceae, together with Lepidozia,

Telaranea, Zoopsis, and other genera of Lepido-

ziaceae in the modern sense. Schuster (1965) re-

garded the presence of a "cephalozioid" (8+4)

seta in Arachniopsis as an instance of "secondary

and independent reduction."

Telaranea Spruce ex Schiffn.

Telaranea Spruce ex Schiffn. in Engler & Prantl, Die

Natiirl. Pflanzenfam. 1(3,1): 103. 1893, nom. cons.

Telaranea Spruce, Trans. & Proc. Bot. Sex:. Edin-

burgh 15: 358, 360, 365. 1885, nom. inval. in syn.

Lepidozia subg. Telaranea (Spruce ex Schiffn.) K.

Mull, in Rabenh.. Krypt.-Fl. 6(2): 276. 1914.

(Type: Lepidozia chaetophylla Spruce).

Arachniopsis Spruce, On Cephalozia 84. 1882, nom.

rejic. Lectotype (fide Schuster, 1965): Arachniopsis

coactilis Spruce.

Neolepidozia Fulf. & J. Tayl., Brittonia 1 1: 81. 1959.

Type: Jungermannia capilligera Schwaegr.

Acrolepidozia Schust., J. Hattori Bot. Lab. 26: 254.

1963. Type: Lepidozia longitudinalis Herz.

Plants soft-textured and often lax, to ± stiff and

wiry, to thread-like and minutely prickly, mostly

prostrate or creeping, occasionally ascending to

suberect, pale, whitish or yellowish to pure green,

rarely brownish yellow to rust brown, often nitid,

at times glaucous and water repellent. Branching

l(2)-pinnate, usually rather regularly so, the

branches predominantly of Frullania type, rarely

to frequently becoming flagelliform, rarely absent;

Microlepidozia-lype and Acromastigum-type

branches in some species; ventral-intercalary

branches present, leafy or stoloniform, or absent;

lateral-intercalary branches rarely present. Stems

with 9-18 rows of cortical cells (as few as 4 in

some species of sect. Tenuifoliae), mostly thin-

walled (the exposed wall rarely thickened), form-

ing a conspicuous hyaloderm (up to 30 rows of

weakly differentiated cells in sect. Tricholepido-

zia); medullary cells numerous (as few as 1 in

some species of sect. Tenuifoliae). Rhizoids from

basal and/or distal cells of underleaf. Leaves in-

cubously inserted and oriented, varying to trans-

versely inserted or almost longitudinal in some

species, rarely weakly succubous, highly variable

in form, ± symmetrically lobed, 4-lobed (or 4-6-

lobed) in most species, 2-3, as few as 1, or as

many as 1 2-lobed (sect. Tricholepidozia), the

leaves shallowly to deeply divided; branch leaves

similar to those of main axis, typically with 1 lobe

fewer than stem leaves, in some species the

branches differentiated, the branch leaves (and

underleaves) differing from those of the stem in

insertion and orientation, and form. Lobes ranging

from acute, caudate, acuminate to ciliiform, typi-

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

cally uniseriate for much of their length, to unis-

eriate almost or quite to the base. Disc (when pre-

sent) variable, from 2 cells high (consisting of a

single tier of cells along the insertion, plus the

basal cells of the lobes) to as many as 6(14) cells

high, in some species the "disc" consisting of the

connate basal cells of the lobes; margins ±

straight to slightly curved, normally entire (very

rarely denticulate by projecting septae). Cells of

disc typically in regular rows and tiers, the num-

ber of cell rows equal to twice the number of

lobes, often large, usually leptodermous, occa-

sionally firm- to evenly thick-walled, often rather

turgid, in a few species collapsing on drying; cu-

ticle mostly smooth, in some species variously pa-

pillose, finely scabrous (with minute, sharp prick-

les), or with a uniform glaucous coating. Oil-bod-

ies hyaline or grayish, the surface variable, ± ho-

mogeneous in some species, coarsely granular in

other species, some species coarsely papillose,

(4)5-9(12) per cell. Underleaves small and incon-

spicuous (rarely rudimentary) to large and similar

to leaves in form but with fewer, and often shorter

lobes; margins entire. Asexual reproduction rare,

by tubers or caducous and fragmenting leaf lobes

in a few species.

Dioecious, less commonly autoecious. Androe-

cia either terminal on short to moderately long

Frullania-type branches or, more often, on short,

spicate ventral-intercalary branches from leading

shoots; bracts typically monandrous, the anther-

idial stalk 1- or 2-seriate; bracteolar antheridia

lacking. Gynoecia on short ventral-intercalary

branches lacking normal vegetative leaves. Bracts

and bracteoles similar, crenulate to dentate, lobu-

late to shallowly to deeply lobed. Perianth large

for plant size, typically assurgent, ovoid to fusi-

form to cylindrical, terete below, usually bluntly

trigonous to shallowly to deeply plicate above

(rarely eplicate), gradually or abruptly narrowed

to the mouth; perianth surface smooth (very rarely

with prorate cells or ciliate); mouth variable, den-

ticulate to shallowly lobulate to lobulate-ciliate to

subfimbriate; perianth 2-4(5 )-stratose in basal

portion, the median portion (l)2(3)-stratose.

Seta with outer layer of 8 cells or exceptionally

as few as 4, the inner cells 15-24(34) to 12 or

fewer, rarely as few as 6 or 4. Capsule short- or

exceptionally long-elliptic, the wall typically 3-

stratose, less often 2-, 4-, or 5-stratose; outer layer

of cells in tiers, rectangular, with two-phase de-

velopment, the longitudinal walls with nodule-like

thickenings alternating with primary walls devoid

of thickenings; innermost cells usually with semi-

annular bands, at times forked and anastomosing

and delimiting local fenestrae, the bands com-

plete, less often incomplete, or bands weakly de-

veloped or lacking, the radial walls with nodular

thickenings and short spinelike extensions onto

the exposed tangential wall.

Spores (SEM) reticulate-areolate, with enclosed

polygonal areolae bounded by a network of low

ridges, or papillose-vermiculate, with short dis-

continuous ridges formed by coalesced papillae,

or the surface irregularly roughened. Elaters little

tapered, typically bispiral to tips.

The Genus Telaranea in Australasia

Key to Australasian Taxa of Telaranea1

1. Leaves and underleaves with 2-6 lobes; stem with a distinct hyaloderm formed of 6-18 cell rows;

capsule wall normally 2-4-stratose 2

2. Leaves (on main shoots) with 4 or more lobes (or if 3-lobed, then with disc present); leaf disc

present and variously developed, 2-6 (to 9) cells high, or in deeply divided leaves, with at least

one tier of disc cells along the insertion, plus the basal cells of the lobes; leaf lobes 2-4(8)

cells wide at base; half-leaf associated with terminal branches with 2 or more lobes (undivided

in T. fragilifolia and T. martinii) 3

3. Lobes caudate, with a broad triangular base abruptly contracted to a uniseriate row, the

cells of the uniseriate row distinctly elongated (ca. 4-5:1) and capillary; lobe cells thick-

walled in comparison to the cells of the disc, the septa thickened in the corners and swollen

. 4

1 An earlier version of this key was sent to R. M. Schuster ca. 1995 as part of a projected "Flora of New Zealand."

That key was appropriated, with minor modifications, by Schuster (2000, p. 212-226). For example, T. obscura Engel

& Mem, a manuscript name, appears in the key.

FIELDIANA: BOTANY

4. Leaf disc 7-9 cells high; plants glaucous and water repellent; first branch underleaf

undivided, ciliiform, at times inserted on main axis. Tasmania, Australia (NSW) ....

T. grossiseta (p. 39)

4. Leaf disc 4-6 cells high (median sinus); plants not glaucous or water-repellent; first

branch underleaf of terminal branches mostly bifid (occasionally 3-fid or undivided),

uniformly inserted on base of branch. New Zealand (primarily North Island)

T. gibbsiana (p. 37)

3. Lobes acute, gradually attenuate to acuminate, the lobe cells shorter (to 3:1, except T.

elegans); lobe cell walls ± equal in thickness to those of disc cells, the septa often thickened

in the corners but not swollen (except T. praenitens) 5

5. Plants glaucous and water repellent; lobe tips (and sometimes disc) fragmenting . . 6

6. Disc cells thin- to firm-walled, larger, to 54 jxm wide; disc and lobe margins not

denticulate; plants whitish, glaucous, larger (to 1 cm wide with branches) .... 7

7. Underleaf disc 3-4 cells high; leaf disc margins entire (cells not bulging).

New Zealand (primarily South Island) T. tuberifera (p. 67)

7. Underleaf disc 1-2(3) cells high; leaf disc margins crenulate by bulging cells

8

8. Lobes submoniliform throughout, the lobe cells short, barrel-shaped;

leaves 4-lobed (or at times 5-6[7]-lobed), not commonly fragmenting

(lobes sometimes missing but disc usually intact). Tasmania, Australia

(Victoria, NSW) T. centipes (p. 64)

8. Lobes straight-sided in distal portion; leaves (3)4-lobed, fragmenting, of-

ten only portions of disc present. New Zealand (North Island)

T. perfragilis (p. 72)

6. Disc cells uniformly thick-walled, smaller, to 45 u>m wide; disc and lobe margins

finely denticulate by projecting upper ends of cells; plants green to olive, ± nitid,

often not glaucous (except in youngest parts of plant), small (to 6 mm wide with

branches). New Zealand (North Island); Australia (NSW, Queensland)

T. elegans (p. 61)

5. Plants not glaucous; lobe tips persistent 9

9. Disc 14-16 cells wide at the insertion (or if ca. 8, then the disc more than 6 cells

high), the leaves often ± horizontally inserted and oriented; leaf lobes (4)5-6(8)

cells wide at base; cells of disc and lobes ± undifferentiated 10

10. Disc narrowing to 8(9) cells wide at the insertion; stem cortical cells in 12-

1 4 rows 11

1 1. Median leaf cells 16-26 fim wide, evenly thick- walled and firm; branch

leaves subfalcate, appearing as if brushed toward tip of branch; leaves on

main axis incubously inserted and obliquely oriented. New Zealand (West-

land Prov. only) T. pennata (p. 18)

1 1 . Median leaf cells 24-38 u,m wide, thin-walled, lacking trigones; branch

leaves not subfalcate, not brushed toward branch tip; leaves on main axis

distinctly incubous and almost longitudinally inserted and oriented. Tas-

mania, Australia (Victoria) T. consobrina (p. 29)

10. Disc (1 1)14-16(18) cells wide at insertion; stem cortical cells in ca. 24 rows

12

12. Flagelliform terminal branches with at most a few normal leaves at base;

leaf disc 8-14 cells high (median sinus), often longer than'broad; rhizoids

originating from basal cells of underleaf disc; leaf lobes often unequal in

size: the dorsal smallest, the two ventral lobes larger. Forest species, on

soil. Western Australia T. clatritexta (p. 11)

12. Flagelliform terminal branches leafy for much of their length; leaf disc

6-8 cells high, broader than high; rhizoids originating from underleaves

at bases of lobes and distal cells of disc; leaves ± symmetrically lobed.

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA .1

Hygric, often in bogs. Auckland Is., New Zealand (South Island, unknown

N of Westland Prov.) T. meridiana (p. 13)

9. Disc typically 8 cells wide at the insertion (6 cells wide in 3-lobed leaves), the

leaves transversely to incubously inserted (if horizontally oriented, then leaves

glaucous, cf. couplet 5); leaf lobes 4 cells wide at base or fewer; cells of disc and

lobes variously differentiated (in size, shape, wall thickening, and cuticle) . . 13

13. Stem leaves all or predominately 3-lobed (or if 4-lobed then plants distinctly

glasslike when dry, and appearing etiolated, the leaves shrunken) 14

14. Microlepidozia-type branches present; underleaves asymmetrically 3—4-

lobed, with one or more lobes abbreviated 15

15. Leaves with lobes persistent, the leaves of main stems 3(4)-lobed, the

disc 2-3 cells high; branch leaves mostly 3-lobed; cuticle smooth or

finely striate-papillose. Lateral-intercalary branches occasional. New

Zealand T. trilobata (p. 89)

15. Leaves with lobes freely fragmenting, the leaves on main stems all

3-lobed, the disc 1-1.5 cells high; branch leaves asymmetrically bifid

(dorsal lobe shorter); cuticle striolate. New Zealand (North Island,

Little Barrier Is.), Tasmania T. fragilifolia (p. 90)

14. Microlepidozia-type branches absent; underleaves symmetrically lobed (or

if asymmetric, 2-lobed, the longer lobe uniseriate, resembling the lobes

of the leaves 16

16. Leaf disc 1.5 cells high, with a single tier of cells along the insertion,

plus the partially connate basal cells of the lobes; cuticle of leaf lobes

minutely scabrous; first branch underleaf undivided. New Zealand

(North Island) T. granulata (p. 103)

16. Leaf disc 2-4(5) cells high; cuticle of leaf lobes finely striate papillose

(T. remotifolia) or smooth; first branch underleaf bilobed, often asym-

metrically 17

17. Leaves 3-4-lobed, shrunken and glass-like when dry, the lobes

often sharply decurved and clawlike; branch half-leaf bilobed;

stems fleshy, with cortical cells in 10-13 rows, the medullary

cells in 17-25 rows; dioecious. New Zealand (sporadic on South

and North Islands) T. remotifolia (p. 73)

17. Leaves on main shoot consistently 3-lobed, the lobes ± straight,

not clawlike; branch half-leaf undivided; stems slender, with 9

cortical cells, the medullary cells in 9-10 rows; monoecious. New

Zealand T. martinii (p. 98)

13. Stem leaves 4-6-lobed, never 3-lobed on main shoots; plants not glasslike,

not etiolated in aspect, the leaves not appearing shrunken. Lobes not claw-

like, about equal in size 18

18. Leaf disc 1.5-3(4) cells high 19

19. Cuticle of leaf lobes smooth 20

20. Basal tier of disc cells small, short-rectangular to quadrate; Mi-

crolepidozia-type branches present; underleaves asymmetrically

lobed, with 1 or more lobes abbreviated. New Zealand, Tasmania

T. pallescens (p. 87)

20. Basal tier of disc cells narrowly elongate; Microlepidozia-type

branches not produced; underleaves symmetrically lobed ... 21

21. Disc of stem leaves (incl. basal tier of lobe cells) 2 (rarely

3) cells high; cells of uniseriate row of leaf lobes straight-

sided and without or with weakly protruding septa, the cells

thicker walled than those of disc; leaves (except van com-

planata) of both stems and branches widely spreading to

squarrose, typically transversely inserted and oriented

FIELDIANA: BOTANY

(branch leaves at times weakly ihcubous); stem medullary

cells thick-walled, in 40-67 rows; branches (except var. com-

planata) terete; sporophytes common. New Zealand (com-

mon throughout) T. lindenbergii (p. 80)

21. Disc of stem leaves 3(4) cells high (rarely 2); cells of unis-

eriate row of leaf lobes typically constricted at the septa, the

cells not noticeably thicker walled than those of disc; leaves

of both stems and branches obliquely spreading, incubously

inserted and oriented; stem medullary cells thin-walled, in

18-36 rows; branches complanate; sporophytes unknown.

New Zealand (primarily South Island)

T. tetradactyla (p. 91 )

19. Cuticle of leaf lobes scabrous, roughened by minute, ± evenly spaced,

sharp prickles. Australia (NSW, Queensland)

T. quadriseta (p. 95)

18. Leaf disc 4 or more cells high from medium sinus base to leaf base . . 22

22. Leaves densely areolate, the cells small (18-40 u.m wide in median

disc), never turgid and pillow-like, typically firm-walled, trigones of-

ten present, small to medium 23

23. Cells of uniseriate row of leaf lobes with septa thickened and

projecting; disc margins (especially the dorsal) minutely serrulate

by projecting distal ends of cells. Cuticle of lobe cells finely to

distinctly striate-papillose. New Zealand (widespread)

T. praenitens (p. 32)

23. Cells of uniseriate row of leaf lobes with septa thickened but not

distinctly swollen and projecting; disc margins entire 24

24. Leaves transversely inserted, palmately divided to 0.6; cells

of uniseriate row of lobes elongate (to 2.3: 1 ). Tasmania . . .

T. palmata (p. 31)

24. Leaves incubously (to almost longitudinally) inserted and ori-

ented, not palmately lobed; cells of uniseriate row of lobes

isodiametric to short rectangular (to 1.3:1) 25

25. Branch leaves asymmetrically lobed, subfalcate and

swept toward the tip of branch, the dorsal lobe smallest

26

26. Leaf lobes 4(5) cells wide at base, often with up to

3 additional 4-celled tiers, the lobe tips not cadu-

cous; disc 15-16 cells wide in distal portion, the disc

cells 16-24 u,m wide. Australia (New South Wales)

T. quadristipula (p. 20)

26. Leaf lobes biseriate at the base, at most with an ad-

ditional biseriate tier, the lobe tips often caducous;

disc 8 cells wide throughout, the disc cells 26-32

(Jim wide. Australia (Queensland)

[T. disparata p. 147]

25. Branch leaves not or only weakly asymmetrically lobed,

not subfalcate ; 27

27. Cuticle of leaf disc distinctly papillose, the papillae

hemispherical near lobe bases and distinctly elon-

gated and striolate below; branch leaves strongly in-

cubous to almost laterally inserted, the median dor-

sal cortical cells ± exposed. Australia (Queensland)

[T. verruculosa p. 197]

27. Cuticle of leaf disc smooth (except T. patentissima

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

van zebrina); branch leaves moderately to distinctly

incubous, but the insertion reaching the branch mid-

line 28

28. Disc cells small, the largest to 25 |xm wide, the

areolation dense, the cells somewhat irregularly

arranged; branches complanate, wider than the

main shoot, the branch leaves typically closely

and regularly shingled. Tasmania, Australia

(Victoria, Queensland), Campbell and Auckland

Is. (rare) T. tridactylis (p. 24)

28. Disc cells larger, the largest to 40 jxm wide, the

areolation more ordered and regular (cells in ±

regular rows and tiers); branches not complanate,

the branch leaves not tightly shingled 29

29. Lobe cells thin- to moderately thick-walled,

with septa thickened in the corners; disc

cells with small but distinct trigones; lobes

about equal to or shorter than the disc, the

uniseriate row 2-4(5) cells long (typically

less than half the length of lobe). Cuticle of

leaf lobes often striate-papillose. New Zea-

land, Tasmania, Australia (NSW, rare) ....

T. patentissima (p. 46)

29. Lobe cells distinctly and evenly thick-

walled; disc cells with walls evenly thick-

ened, lacking trigones; lobes longer than the

disc, the uniseriate row 4-5(6) cells long

(more than half the length of lobe). Cuticle

of leaf lobes smooth. Tasmania

T. capilligera (p. 17)

22. Leaves laxly areolate, the cells larger (40 (Jim or more wide in median

disc), often appearing turgid and distinctly pillow-like (even under

low magnification), typically ± thin-walled and with trigones lacking

30

30. Leaves (disc) ± symmetrical; leaf lobes acute to acuminate, with

1-2 tiers of paired cells at base; disc cells isodiametric .... 31

31. Cells in median sector of disc thin- walled, to (45)50-70 |xm

wide; uniseriate portion of leaf lobe 5-6 cells long; plants

varying from medium to robust (to 1 .5 cm wide), highly nit-

id; disc weakly to distinctly cuneate and (9)14-21 cells broad

distally (approaching 8 cells wide throughout in van ro-

seand). New Zealand (common) T. tetrapila (p. 52)

3 1 . Cells in median sector of disc evenly thick- walled and firm,

rather dense, to 38 jxm wide; uniseriate portion of leaf lobe

short, 2—3 cells long; plants small and stenotypic in stature,

ca. 0.6 cm wide (including branches), rather dull; disc sub-

quadrate, 8(9) cells wide throughout. Auckland Is., New Zea-

land (South Island and extreme southern sector of North Is-

land) T. hodgsoniae (p. 45)

30. Leaves (disc) asymmetrical; leaf lobes long attenuate, often fal-

cate and hooked at the tips, with 3-5 tiers of paired cells at base;

disc cells ± elongate. New Zealand T. paludicola (p. 60)

10 FIELDIANA: BOTANY

2. Leaves with 2-3 lobes, deeply divided; leaf "disc" formed of the partly connate basal lobe

cells and lacking a tier of disc cells along the insertion; leaf lobes from a 1 -seriate or biseriate

base; half-leaf undivided 32

32. Stems appearing fleshy, the cortex in 9-10 cell rows and the medulla in 18-24 rows; leaf

lobe bases biseriate, the paired cells elongate, connate for more than 0.5 their length, the

cells of the uniseriate row barrel-shaped, constricted at septa; underleaves 3-4 lobed. Alpine

tussock grassland plant. New Zealand T. nivicola (p. 102)

32. Stems slender and wiry, the cortex in 6-9 cell rows and the medulla in 6-12(18) rows;

leaf lobes uniseriate or biseriate at base, the basal cells connate for 0.1-0.4 their length,

the cells of the uniseriate row with septa thickened at the corners, feebly swollen; under-

leaves bilobed. Lowland plants 33

33. Underleaves typically asymmetrically bilobed, the longer lobe resembling a leaf lobe,

with a uniseriate row 4(5) cells long, the other abbreviated, with a pair of short basal

cells, a ± elongated cylindrical cell, capped by a slime papilla; Microlepidozia-type

and lateral-intercalary branches present. New Zealand .... T. quinquespina (p. 109)

33. Underleaves symmetrically bilobed, small and caliper-like; Microlepidozia-lype and

lateral-intercalary branches absent 34

34. Lobes not tapering, flexuous, 6-9 cells long; first branch underleaf equally and

symmetrically bilobed; leaves consistently bilobed, the lobes unequal in length;

lobe cells about equal in length, the terminal cell small and button-like; cortical

cells moderately and evenly thick walled. New Zealand, Tasmania

T. inaequalis (p. 1 1 7)

34. Lobes tapering, stiff, (4)5-6 cells long; first branch underleaf asymmetrically bi-

lobed, one lobed long and divergent, the other short and appressed, resembling an

underleaf lobe; leaves 2-3-lobed, when bilobed the lobes ± equal in length; lobe

cells gradually shorter toward tip, the terminal cell not small and button-like; cor-

tical cells thin walled 35

35. Leaf lobes uniseriate to base (only sporadically with 1 or more lobes biseriate

at base), the basal cell 95-120 jxm long. New Zealand, Tasmania, Australia

(Victoria) T. herzogii (p. 1 1 2)

35. Leaf lobes nearly always biseriate at base, the paired cells of the basal tier

54-74 |xm long. Tasmania T. tasmanica (p. 106)

1. Leaves with 8-12(13) lobes; stem with a weakly differentiated hyaloderm of ca. 24-28 cell rows;

capsule wall 5-stratose. New Zealand (Stewart Is., Southland and Westland Prov. of South Is.),

Tasmania, Australia (rare in Victoria) T. pulcherrima (p. 76)

Taxonomic Treatments

Telaranea subg. Acrolepidozia (Schust.) Schust.

Acrolepidozia Schust., J. Hattori Bot. Lab. 26: 254.

1963. Telaranea subg. Acrolepidozia (Schust.)

Schust., J. Hattori Bot. Lab. 36: 389. 1973 (1972).

Telaranea subg. Acrolepidozia (Schust.) Schust.,

Hep. Anthoc. N. Amer. 2: 30. 1969, nom. inval.

basionym non cit. Type: Lepidozia longitudinalis

Herz.

Telaranea sect. Latifoliae Engel & Merr, Phytologia

79: 251. 1996 (1995). Type: Lepidozia meridiana

Hodgs.

Note that "sect. Meridianae" (Schuster, 2000,

p. 212), was a provisional name for the section

published by Engel and Merrill (1996) as Sect.

Latifoliae.

For a discussion of subgenera and sections and

their distinguishing characteristics, see the phy-

logenetic section which follows (p. 232).

Telaranea clatritexta (Steph.) Engel & Merr.

Lepidozia clatritexta Steph., Spec. Hep. 3: 583. 1909.

Neolepidozia clatritexta (Lindenb.) Fulf. & J. Tayl.,

Brittonia 11: 85. 1959. Telaranea clatritexta

(Steph.) Engel & Merr., Novon 9: 339. 1999. Type:

Australia, Western Australia, Swan River, Drum-

mond, ex Herb. Kew (G!).

Lepidozia whiteleggeana Steph., Spec. Hep. 3: 584.

1909, syn. nov. Type: Australia, Cook River, Bot-

any Bay, Whitelegge. comm. F. v. Miiller No. 34

(G!).

Lepidozia complanata Herz., Memoranda Soc. Fauna

Fl. Fenn. 27(1950-1951): 92. / 39. 1952, syn. fide

Engel & Merrill, 1999. Telaranea complanata

(Herz.) Engel & Merr., Phytologia 79: 251. June,

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

11

1996 (1995). Type: Australia, Western Australia,

without specific loc., Goebel (M — non vidi); iso-

type: (JE!).

Plants soft and flexuous, complanate and re-

sembling T. centipes, loosely creeping, in rather

dense mats, pale green, highly nitid when dry,

with age becoming dull and opaque brown; plants

medium, to 1.0 cm wide, including branches.

Branching rather regularly and laxly 1 -pinnate,

often with numerous long, geotropic flagelliform

branches (to 2 cm long), which have few or no

normal leaves at the branch base, the branches

nearly exclusively of the Frullania-type; branch

half-leaf shallowly 2(3)-fid, obliquely inserted and

in axil of main axis and branch, narrowly rect-

angular to elliptic; first branch underleaf undivid-

ed and subulate (or less often to 3-lobed), longi-

tudinally inserted on ventral-lateral side of main

axis at or somewhat below branch base. Acro-

mastigum-type branches sporadic, stoloniform,

the branch half-underleaf shallowly bifid. Ventral-

intercalary branches sporadic, leafy. Stems with

cortical cells rather weakly differentiated, thin

walled, in 15-26 rows; cortical cells in section

moderately larger than the numerous (to ca. 95)

medullary cells, the medullary cells moderately

thin-walled, the walls finely pitted. Leaves on

main shoot rigid, distant to loosely imbricate,

when dry with margins revolute (the leaves ap-

pearing channelled), when moist the leaves wide-

ly spreading, the disc plane or nearly so but the

lobes somewhat ventrally decurved, the leaves

strongly longitudinally inserted and oriented in

well-developed shoot sectors and nearly in same

plane as dorsal surface of stem; leaves 455-645

u,m wide X 610-750 u,m long, rather asymmet-

rically 4(6) lobed to (0.25)03-0.45, the lobes

tending to subdivide, straight to feebly conver-

gent, often distinctly shorter than the disc. Lobes

narrowly attenuate, sporadically weakly acumi-

nate, the dorsal lobe at times smallest and the two

ventral lobes larger, the largest lobes 5-6(7) (rare-

ly 4) cells wide at extreme base, tapering to a

short uniseriate row of 2-3(4) cells; lobe cells ±

isodiametric to somewhat elongate, thin- to ±

thick-walled, the cell walls of uniseriate row not

much thickened in the corners, the basal cell of

the uniseriate portion 20-36 u,m wide X 24-38

u,m long (0.9-1 .5: 1 ), the next cell slightly narrow-

er and shorter, 1 8-30 u,m wide X 1 8-3 1 u,m long

(0.8-1.6:1), the terminal cell shorter than the pen-

ultimate cell, short conical and rounded at the tip;

cuticle smooth. Disc variable: ± symmetrically

broad- to narrow-rectangular, (6)8-14 cells high

(from median sinus base to leaf base), wider than

high to higher than broad, 14-22 cells wide in

distal portion narrowing to (10)14-16(18) cells

wide in basal portion; dorsal margin gently curved

and subcordate at the base, the ventral ± straight,

entire or rarely with a small accessory lobe or

tooth, often strongly decurrent. Cells of disc thin-

to slightly and evenly thick-walled or the free wall

at margins ± thickened, trigones lacking to mi-

nute, the median cells elongate, 28-39(43) jxm

wide X 53-65 u,m long, the cells in ± regular

longitudinal rows, not noticeably tiered; basal row

of disc cells somewhat larger; cuticle smooth. Un-

derleaves variable in size, ca. 0.8-2X the stem in

width, strongly spreading, distant, plane or slight-

ly convex, 4(6) lobed to 0.3-0.45, the lobes

straight to curved at the tips, weakly divergent,

highly variable, the lobes consisting of a base of

2(3) cells, sporadically with a biseriate tier above

the base, and a uniseriate row of 2-3(4) not or

somewhat elongate cells, or the lobes narrowly

acute, 4-5 cells wide at base and resembling the

leaf lobes, terminating in a slime papilla; disc

symmetrically subrectangular (wider than high),

variable in height and width, 3-6(7) cells high

(median sinus), 12-15(19) cells wide at widest

point; margins entire. Rhizoid initial cells small,

subquadrate, forming a bistratose pad or band at

the base of the disc. Asexual reproduction by cla-

vate tubers at the tips of stonoliferous branches.

Dioecious. Androecia not seen. Gynoecia

strongly dorsally assurgent, slightly swollen and

sparsely rhizoidous at base; bracts of innermost

series erect to appressed to the perianth, ovoid to

elliptic to subrectangular, ± regularly but un-

equally 4(6)-lobed to 0.3-0.4, the lobes narrowly

attenuate, 4-6 cells wide at base, ending in a sin-

gle cell or a short uniseriate row of 2-4 cells, the

terminal cell with a rounded tip or surmounted by

a slime papilla, the lobe margins sporadically with

a 1 -several-celled tooth; lamina composed of lax,

± regularly elongate-rectangular cells, the mar-

ginal cells similar, the margin with several sessile

slime papillae, otherwise entire; bracteole of in-

nermost series smaller but similar in form to

bracts. Perianth ca. 0.75 emergent, stout cylindric,

terete in basal sector, the distal sector trigonous,

with 3 distinct plicae, narrowing toward the con-

tracted mouth; mouth shallowly ca. 10-12 lobu-

late, the cells thin-walled but firm, the lobules

ending in a single elongate cell or in a uniseriate

row of 2 cells, then with the transverse septa

thickened in the corners and weakly swollen and

12

FIELDIANA: BOTANY

projecting, the terminal cell broadly rounded, not

ending in a slime papilla; perianth 4-5-stratose in

basal portion, the median and upper portions 1-

stratose.

Sporophyte not seen.

DIFFERENTIATION AND VARIATION — The original

description and figure of T. complanata (Herzog,

1952) gives only a hint of the extreme variability

of this species. The impression given is of a plant

"aus der Verwandschaft von L. centipes," as not-

ed by Herzog (ibid., p. 93): subsymmetrical, elon-

gate rectangular leaves, divided to the middle,

slender leaf lobes that are 3-4 cells wide at the

base, small underleaves with a low disc (2 cells

high) and short uniseriate lobes. All of the above

features vary considerably. Our study of a number

of specimens from several localities in Western

Australia has resulted in a considerable broaden-

ing of our concept of the species. In particular, the

leaves are often distinctly broader than long, shal-

lowly divided, with the lobes 5-6(7) cells wide at

the base. The underleaves are often twice the

width of the stem, up to 6 cells high (at median

sinus), with lobes 4-5 cells wide at the base, and

resembling those of the leaves.

Despite a superficial resemblance to T. centi-

pes, on closer examination the leaves are striking-

ly like those of T. meridiana in form, areolation,

and mode of insertion. In particular, the leaves are

typically 14-16 cells wide at the insertion (Fig. 1:

2, 3), with broad lobes, leptodermous cells, and a

weakly differentiated hyaloderm (Fig. 1 : 11). Un-

like T. meridiana, however, most plants show at

least a tendency toward the curious leaf symmetry

seen in T. pennata, with the dorsal lobe smallest

and the ventral lobes largest and ± paired. The

similarities between the species of subg. Acrolep-

idozia, with their highly fragmented distribution,

argues a considerable antiquity for these very un-

Telaranea-\ike species.

A highly unusual feature of this species, apart

from its variability, is the production of abruptly

long-flagelliform, strongly positively geotropic,

terminal branches, which resemble the ventral-in-

tercalary stoloniform branches of many Bazzania

species (Fig. 1:1).

This remarkable species has a disjunct distri-

bution in Western Australia and in New South

Wales, where it is represented by the type of L.

whiteleggeana. The latter is a luxuriant plant with

stems (as noted by Stephani) to 5 cm long. The

leaves are 12-14 cells wide along the insertion,

the lobes are 4-5 cells wide at the base, and the

disc is up to 11 cells high. Regrettably, the iden-

tity of L. whiteleggeana was not known to us at

the time the combination based on L. clatritexta

was made (Engel and Merrill, 1999). The type of

Lepidozia clatritexta consists of only three stems,

and it is a weakly developed expression of the

species. The leaves are described as 8 cells wide,

but in fact the disc is 10 cells wide at the insertion

and 14-17 cells wide in the distal portion. The

leaf lobes are 4(5) cells wide at the base. Never-

theless, the distinctive features of this taxon are

expressed, namely, the abruptly flagelliform ter-

minal branches, the longitudinally inserted leaves,

and underleaves with rhizoids originating from

the basal cells of the disc.

DISTRIBUTION AND ECOLOGY — Disjunct in New

South Wales and in Western Australia. In the

west, the species is found from near sea level to

650 m, in forests of Eucalyptus calophylla, Aca-

cia alata, Jacksonia, or of kauri. The species oc-

curs over soil (typically loamy) in humid situa-

tions, e.g., shaded banks, stream banks, protected

sites near cascades or in boulder fields.

SELECTED SPECIMENS SEEN — AUSTRALIA. WEST-

ERN AUSTRALIA: Churchman Brook, off Soldier's

Road, Roleystone, just below Churchman Brook Res-

ervoir, SE of Perth, 90 m, Engel 13319 (F); Darling

Range, Little Dandalup Creek, near intersection of Tor-

rens Road with Delpark Road, Engel 13356 (F); The

Cascades, Lefroy Brook, S of Pemberton, Engel 13418

(F); Walpole-Nornalup Natl. Park, near intersection of

Hilltop and Gully Roads, Engel 13425 (F); ibid.. Brainy

cutoff near intersection with Tingle Drive East, just be-

yond bridge over Frankland River, Engel 18436 (F); Po-

rongurup Natl. Park, Porongurup Range, Devils Slide,

440-650 m, Engel 13476 (F); Waterfall Beach, Two

Peoples Bay Nature Reserve. 30 km E of Albany, Sto-

neburner & R. Wyatt 3680 as T. centipes (MELU).

Telaranea meridiana (Hodgs.) Hodgs.

a meridiana Hodgs.. Trans. Roy. Soc. New

Zealand 83: 61 1. pi. 2, f. 24. 1956. Telaranea mer-

idiana (Hodgs.) Hodgs., Rec. Domin. Mus. 4: 107.

1962. Type: Auckland Is., "Cape Expedition," No.

2 Camp. 31 Oct. 1944, Turbott x.n. (CHR!).

Plants soft and flexuous, brittle, ascending to

suberect, in dense cushions, pale green, nitid

when dry; shoots medium, to 1.5 cm wide, in-

cluding branches. Branching somewhat regularly

but loosely 1(2) pinnate, often becoming flagelli-

form, the branches typically flexuous and some-

what contorted, of the Frullania-type; branch

half-leaf 2-lobed, distinctly obliquely inserted,

narrowly rectangular, the lobes ± parallel; first

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

13

FIG. 1. Telaranea clatritexta (Steph.) Engel & Merr. 1. Sector of main shoot with Frullania-type branches, one,

at upper-right, becoming long flagelliform. dorsal view. 2, 3. Leaves, cellular detail. 4-6. Leaves (dm = dorsal

margin; fig. 5 with an underleaf at left at same scale). 7. Main shoot with Acromastigum-type stoloniform branch (at

left; note half-underleaf) and Frullania-lype branch (at right). 8-10. Underleaves, cellular detail, showing variation

in size and lobe development; stippled cells are those that each give rise to a rhizoid. 1 1. Stem, cross section. 12-

13. Primary branch leaves (= bl). 14. Basal portion of shoot showing stoniferous branch and a tuber. 15. Portion of

perianth mouth. 16. Gynoecial bracts. (Figs. 1-2, 5. 7, 9, 1 1, 14-16 from Engel 13425, Australia, Western Australia.

Walpole-Nornalup Natl. Park, near intersection of Hilltop and Gully Roads; 3-4, 10, 13 from Engel 13446, same

loc.; 6. 8. 12 from Engel 13418. Australia. Western Australia, The Cascades. Lefroy Brook.)

14

FIELDIANA: BOTANY

branch underleaf undivided, broadly acuminate

(rarely bilobed), inserted on ventral-lateral side of

branch at juncture of branch and main axis, grad-

ing to longitudinally inserted on stem just below

the branch. Ventral-intercalary branches not seen.

Stems with cortical cells rather weakly (on stron-

gest main axis) to distinctly differentiated, thin

walled, in 17-24 rows; cortical cells in section

slightly to distinctly larger than the numerous (ca.

120) medullary cells, the medullary cells moder-

ately thin walled, the walls finely pitted. Leaves

on main shoot rigid, widely spreading, distant to

contiguous (on compact shoots), the disc plane or

nearly so, rarely convex, the lobes ventrally de-

curved and claw-like (not visible in dorsal view),

the insertion incubous (on elongated main shoots

the leaves often strongly longitudinally inserted

and oriented, with the disc broader than high and

nearly in same plane as dorsal surface of stem);

leaves 520-700 jim wide X 545-770 u,m long,

the leaves 4(6) lobed to 0.4-0.6, the lobes straight

to feebly divergent, slightly shorter than the disc.

Lobes weakly acuminate, the largest leaf lobes

(4)5-6(8) cells wide at extreme base, 4-5 cells

wide for 4-5 tiers, biseriate for 1-2 tiers, termi-

nating in a short uniseriate row of 2-3(4) cells;

lobe cells ± isodiametric to short rectangular,

thin-walled, the cell walls of the uniseriate row

not or very weakly thickened in the corners, the

basal cell of the uniseriate portion 1 8-29 jim wide

X 23-36 (Jim long (1-1.5:1), the next cell 14-24

u,m wide X 17-26 jim long (1-1.5:1), the termi-

nal cell normally about equal to the penultimate

cell in length or a little shorter, tapering to the

apex; cuticle smooth, rarely feebly striate papil-

lose. Disc moderately asymmetrically rectangular,

the dorsal margin longer than the ventral, 6-8

cells high (from median sinus base to leaf base),

21-24 cells wide in distal portion narrowing to

(11)14-16 cells wide in basal portion; margins

entire, the dorsal margin straight or weakly sub-

cordate at the base, the ventral straight. Cells of

disc thin-walled, trigones lacking, the median

cells elongate, 30-42 u,m wide X 38-5 1 \im long,

the cells in ± irregular rows; basal row of disc

cells somewhat larger; cuticle smooth. Underleav-

es much smaller than leaves, ca. 1-1.2X stem

width, firmly inserted, strongly spreading, distant,

plane, 4(6) lobed to 0.35-0.65, the lobes straight

to curved at the tips, acuminate, 3-4 cells wide

for 1 -several tiers at base and biseriate for 2-3

tiers, ending in two laterally juxtaposed cells, or

more commonly in a uniseriate row formed of

2(3) short cells, terminating in a slime papillae;

disc symmetrically subrectangular (wider than

high), 5-6 cells high (median sinus), 15-18 cells

wide at widest point, the cells ± regularly ar-

ranged; margins entire, moderately curved. Rhi-

zoid initial cells small, subquadrate, forming a

continuous bistratose pad or band often including

the basal portion of the lobes as well as the apex

of the disc. Asexual reproduction lacking.

Plants dioecious. Androecia either on short

Frullania-lypc branches with a few cycles of re-

duced leaves prior to androecial formation, or on

short, abbreviated, ventral-intercalary branches

from leading shoots; bracts closely imbricate, dor-

sally assurgent, deeply concave, 2-lobed, the

lobes acuminate, terminating in a uniseriate row

of 2 not or hardly elongated cells; lamina cells

irregular in arrangement, the dorsal margin of

lamina feebly dilated and slightly incurved, entire

or crenulate, devoid of slime papillae; bracts mon-

androus; antheridial stalk short, ca. 6 cells high,

biseriate; bracteolar antheridia absent. Gynoecia

not seen.

DIFFERENTIATION AND VARIATION — In its most

characteristic expression (well represented by the

type), T. meridiana is a distinctive plant, very un-

Telaranea like in aspect. Particularly diagnostic

are the flexuous, rather loosely pinnate shoots,

and the broad-based, longitudinally inserted and

oriented leaves (particularly on elongated main

shoots, Fig. 2: 1), which may be up to 16 cells

wide in the basal sector. Also distinctive are the

broad lobes, (4)5-6(8) cells wide at base, which

are ventrally decurved and claw-like, and the uni-

form, dense areolation of the disc and lobes. In

more compact expressions (e.g., Child H3669),

however, the insertion is variable, from distinctly

to rather weakly incubous, and the leaves are rath-

er markedly convex (Fig. 2: 6).

DISTRIBUTION AND ECOLOGY — Known only from

Auckland Is. and South Island, New Zealand (un-

known north of Westland Prov.). The species oc-

curs at lower to median elevations, and is pri-

marily terricolous in damp or boggy sites on the

forest floor. It less commonly is found in rather

open sites, such as in low, mucky niches in

swampy areas with Sphagnum and scattered Lep-

tospermum, etc., at sea level just north of the

Haast River. At Ship Creek (north of the Haast

River) it grows in low, wet areas just above the

water level in a mature kahikatea (Dacrycarpus

dacrydioides) swamp forest. In the Lake Kaniere

area (125 m, Westland) the species is very com-

mon on saturated, rich, peaty soil in and espe-

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

15

FIG. 2. Telaranea meridiana (Hodgs.) Hodgs. 1. Sector of main shoot, dorsal view. 2. Leaves. 3. Leaf, cellular

detail. 4. Leaf lobe. 5. Leaf base (dm = dorsal margin). 6. Shoot sector of vigorous plant, dorsal view. 7. Sector of

main shoot, ventral view. 8. Underleaves. 9. Underleaf, cellular detail, stippled cell walls indicate rhizoid initials. 10.

Stem, cross section. 1 1. Antheridial stalk. (Figs. 1-4, 7-10 from type; 5 from Johnson 175. New Zealand, South Is.,

Southland Prov., Lake Manapouri: 6, from Child H3669, New Zealand. South Is.. Westland Prov., Mahinapua; 1 1

from Child H3101, New Zealand, South Is., Otago Prov., Ml. Cargill.)

16

FIELDIANA: BOTANY

cially at the edges of narrow, shallow, ± stagnant

pools in the shade of a mixed podocarp-broadleaf

forest. Vegetation at this site consists of Dacry-

dium cupressinum, Weinmannia racemosa, Po-

docarpus totara, Metrosideros umbellata and an

open understory dominated by Pseudowintera. It

also occurs in pakihis under rushes, etc., associ-

ated with Sphagnum. The species is associated

with a variety of other hepatics that occur in forest

niches, e.g., Telaranea tetradactyla, Lepidozia

procera, L. spinosissima, L. laevifolia, etc.

SPECIMENS SEEN— NEW ZEALAND. SOUTH IS-

LAND. SOUTHLAND PROV.: Hope Arm, Lake Man-

apouri, ca. 185 m, Johnson 175 (CHR); Fiordland Natl.

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

stream draining from Lake Wade, 300 m, Fife 7782 (F).

OTAGO PROV: Blue Mts., S end, E of Garden Gully

Road, ca. 610 m, Child s.n.l 53 (F); W slope of Flagstaff,

NW of Dunedin, 490-520 m, Engel 17624 (F); Mt. Car-

gill, N of Dunedin, ca. 455 m, Child H3101—C. 6 as

Lepidozia laevifolia (F); Lee Stream, ca. 305 m. Child

H2504 as T. gottscheana (F); Lammermoor Range, ca.

610 m, Child H3855 (F); N of McKerrow River, Mar-

tin's Bay, Hatcher 760, 853 (F). WESTLAND PROV.:

Ca. 4 km N of Haast River, sea level, Engel 21770 (F);

Ship Creek, 14.5 km N of Haast River, sea level, Engel,

21726, (F); near Lake Paringa, near west coast, ca. 15

m, Child H1796, 1801 as Lepidozia microphylla (F); Mt.

Aspiring Natl. Park, Cross Creek, 1.1 km N of Haast

Pass, 540 m, Engel 21876 (F); Mahinapua, S of Hoki-

tika, ca. 60 m. Child H3669 as L. microphylla (F); Lake

Kaniere Scenic Reserve, Lake Kaniere Rd, 1 25 m, Engel

24881 (F); 10 km S of Greymouth, ca. 150 m, Child

H4947 (F).

Telaranea subg. Telaranea

Telaranea sect. Neolepidozia (Fulf. & J. Tayl.)

Engel & Mem, comb. nov.

Neolepidozia Fulf. & J. Tayl., Brittonia 11:81. 1959.

Telaranea subg. Neolepidozia (Fulf. & J. Tayl.)

Schust., J. Hattori Bot. Lab. 26: 255. 1963. Lepi-

dozia sect. Neolepidozia (Fulf. & J. Tayl.) Piippo,

Ann. Bot. Fennici 21: 314. 1984. Type: Junger-

mannia capilligera Schwaegr.

Telaranea capilligera (Schwaegr.) Schust.

Jungermannia capilligera Schwaegr, Hist. Muse.

Hep. Prodr. 21. 1814. Lepidozia capilligera

(Schwaegr.) Lindenb. in G. L. & N., Syn. Hep. 204.

1845. Mastigophora capilligera (Schwaegr.) Trev.,

Mem. 1st. Lomb. Sci. Lett. III. 4: 416. 1877. Neo-

lepidozia capilligera (Schwaegr.) Fulf. & J. Tayl.,

Brittonia 11: 84. 1959. Telaranea capilligera

(Schwaegr.) Schust., J. Hattori Bot. Lab. 26: 256.

1963. Lectotype (nov.): Tasmania, Labillardiere,

hb. Montagne, Lindenberg Hep. no. 4639 (W!);

isolectotypes: (G!, STR!).

Plants rather stiff and wiry, in herbarium clear

bronze, shoots small for subgenus, the main

shoots to 775 u.m wide, the plants to 6 mm wide

with branches. Branching ± regularly pinnate,

the branches of the Frullania type, rather short,

at times flagelliform; branch half-leaf bifid, sub-

transversely to obliquely inserted, the lobes

straight to ± divergent; first branch underleaf un-

divided, biseriate in basal portion, with a uni-

seriate row of 2-3 submoniliform cells, inserted

on ventral side of branch near juncture of branch

and main axis. Ventral-intercalary branches pre-

sent, leafy. Stems with cortical cells in 9-12

rows, rather thick-walled and firm in surface

view, in section larger than the medullary cells,

rather thick-walled; medullary cells in 40-42

rows, the cells much smaller, with moderately

thickened walls. Rhizoids sparse, from distal

cells of underleaf disc. Leaves on main axis rig-

id, obliquely spreading, loosely imbricate, the

disc gently incurved, the lobes moderately to dis-

tinctly incurved, the insertion moderately to

strongly incubous; leaves 330-440 u,m wide

(measured between tips of lobes) x 290-390 u,m

long, subsymmetric, 4-lobed to 0.5-0.6, the

lobes moderately spreading, with some leaves on

main shoot with lateral lobes widely and asym-

metrically divergent (at times forming an angle

of up to 1 80° with each other), the lobes longer

than the disc. Lobes slender, 2-4 cells wide at

extreme base followed by 1(2) biseriate tiers, ter-

minating in a uniseriate row of 4-5(6) cells; cells

of the uniseriate portion short, submoniliform

(lobes constricted at the septa), ± isodiametric,

evenly and distinctly thick-walled (even the ter-

minal cell decidedly thick walled), the basal cell

of the uniseriate portion 31-40 u,m wide and

long (at most 1.3:1), the next cell smaller, 25-31

u-m wide and 30-34 u,m long, the terminal cell

a little smaller than the penultimate cell, not sec-

ondarily divided, broadly rounded at the tip. Disc

± symmetrically cuneate, 4(5) cells high (from

median sinus to leaf base), 12-15 cells wide in

distal portion narrowing to 8 cells wide in basal

tier; margins entire, ± straight. Cells of disc

moderately and evenly thick-walled, lacking tri-

gones, indistinctly tiered; largest cells in median

portion of disc 26-36 u,m wide, (28)35-48 n-m

long, the cells in distal tiers narrower; cuticle

smooth. Underleaves much smaller than leaves,

erect-spreading to nearly at 90° to stem, distant,

plane, 3-4 lobed to 0.45, the lobes rather short,

2 cells wide at the base, terminating in a uni-

seriate row of 2-3 ± isodiametric, submonili-

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

17

form, somewhat thick-walled cells, sometimes

terminating in a slime papilla; disc short cuneate,

3 cells high (median sinus), the cells in ± regular

tiers, the disc 8-10 cells wide in distal portion

(6-8 cells wide in 3-lobed underleaves), narrow-

ing to 8 cells wide at base (6 cells wide in 3-

lobed underleaves); margins entire, straight.

Asexual reproduction lacking.

Androecia and gynoecia not seen.

DIFFERENTIATION AND VARIATION — The type of

Jungermannia capilligera is a striking plant. We

have examined large numbers of New Zealand

and Tasmanian specimens, previously named as

well as unidentified, and have not found any

which match the type of T. capilligera in char-

acteristic disc areolation, cell size and wall thick-

ness, and form of the lobes. Accordingly, we have

chosen to define T. capilligera rather narrowly, so

far represented only by the type. More study is

needed to understand the variability of this spe-

cies and its distribution.

The leaves of the main shoot are deeply dis-

sected (to 0.6), with a disc 4 cells high, and at

least some leaves have the lobes widely spreading

(splayed) as in T. tridactylis (Fig. 3: 1, 2). The

disc cells, however, are larger, 26-36 |xm wide vs.

1 8-25 u,m wide in T. tridactylis. The cells are

firm, uniformly thick-walled and turgid, and are

somewhat more regularly arranged than in T. tri-

dactylis. The cells are comparable to those of T.

patentissima in size, but the leaf lobes are more

slender, terminating in a uniseriate row of 4-5(6)

cells (Fig. 3: 1-4), vs. 2-4(5) cells in T. patentis-

sima. In addition, the disc cells are quite evenly

thick-walled (Fig. 3: 3), vs. thinner-walled and

with small but distinct trigones in T. patentissima.

In T. capilligera, the lobe cells in particular are

evenly and distinctly thick walled (Fig. 3: 4). The

terminal cell is also markedly thick walled, more

so than in any other Australasian member of sect.

Neolepidozia.

Telaranea palmata (p. 31), another Tasmanian

species, is very similar to T. capilligera, both spe-

cies having notably thick-walled leaf cells, partic-

ularly in the uniseriate row of the lobes. The most

obvious difference is in the leaf insertion, which

in T. capilligera is distinctly incubous (Fig. 3: 1,

2), vs. subtransverse in T. palmata. The cells of

the leaf lobes are also more elongate in the latter

species, and the septa often weakly projecting.

Oil-bodies of a specimen from Australia (Syd-

ney, leg. Heim, not seen) were described by Jovet-

Ast (1949) as botryoidal, 3-4 per cell, 8-10(1 1)

u>m long, composed of 5-8 granules. The speci-

men in question was determined by Hodgson as

this species, but the identity of the specimen

should be checked, in light of the more restricted

sense of the species used here.

NOMENCLATURE — Telaranea capilligera is the

earliest described species in the genus Telaranea

(Schwaegrichen, 1814), and is the nomenclatural

type of the genus Neolepidozia (Fulford and Tay-

lor, 1 959), which is treated here as Telaranea sect.

Neolepidozia (see p. 237). The original material

of Jungermannia capilligera represents a quite

different plant from the species as it has previ-

ously been understood. We have examined several

duplicates of the type (G, STR, W); the W spec-

imen is here designated as the lectotype of the

species. Piippo (1984, p. 314) stated that a lec-

totype of Jungermannia capilligera was "chosen

by J. Taylor 1958." The Geneva specimen bears

an annotation by Taylor in 1958 as "original," but

to our knowledge this lectotypification was not

published.

Schwaegrichen (1814) gives the type locality

simply as "Australasia." Synopsis Hepaticarum

(G. L. & N., 1845, p. 204) cites "Insulis austral-

ibus (Labillardiere); in Terra van Diemen (Hb.

M.); in Nova Hollandia (Hb. Berolin.)." The lat-

ter, however, refers to the type of Jungermannia

tridactylis, considered a synonym of J. capilligera

by G. L. & N. The lectotype designated here is

labelled "Hb. Mont.," and is therefore presum-

ably the Tasmanian plant cited.

The G, STR, and W plants are identical, and

almost certainly represent the same collection. All

are a clear, bronze color, something like that of T.

lindenbergii van mellea (p. 86). It is uncertain,

however, if this is the natural color of the plant.

In the van mellea the color is consistently present

in all collections seen.

DISTRIBUTION — Stephani (1909, p. 583) listed

the distribution as Australia, Tasmania, New Zea-

land, and Auckland Is., and "ubique communis."

As treated here, the species is apparently endemic

to Tasmania and known only from the type.

Schuster (2000), however, keyed the species un-

der "Neotropical taxa," but stated that the species

was "doubtfully in Patagonia/Magallanes." We

are discounting records from southern South

America (see Engel, 1978).

Telaranea pennata Engel & Merr.

Telaranea pennata Engel & Merr., Phytologia 79:

252. June, 1996 [1995]. Holotype: New Zealand,

IS

FIELDIANA: BOTANY

FIG. 3. Telaranea capilligera (Schwaegr.) Schust. 1, 2. Leaves, in xiiu. 3. t-eaf. 4. Dorsalmost lobes of leaf. 5.

Underleaf, in situ. 6. Underleaf. 7. First branch underleaves. 8. Stem, cross section. (All from type, hb. W.)

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

19

South Island, Westland Prov., Route 73, 8 miles W

of Turiwhate, Engel 6754 (F); isotype: (CHR).

Plants rather stiff, loosely prostrate, loosely

matted, pale green to olive-green, nitid when dry;

plants medium, to 1 cm wide, including branches.

Branching very regularly 1 -pinnate, the branches

of the Frullania-type, rather long for plant size,

to 775 [Jim wide, not becoming flagelliform;

branch half-leaf bifid, usually obliquely inserted,

narrowly rectangular, the lobes parallel to weakly

diverging; first branch underleaf undivided, sub-

ulate, (rarely bilobed) inserted on ventral side of

branch at juncture of branch and main axis. Ven-

tral-intercalary branches occasional, becoming

leading leafy shoots. Stems with cortical cells dis-

tinctly differentiated, rather thick walled, in 12-

13 rows; cortical cells in section much larger than

the numerous (ca. 80) medullary cells, the med-

ullary cell walls moderately and somewhat un-

evenly thick-walled. Leaves on main shoot rigid,

obliquely spreading, contiguous to imbricate, the

disc ± plane, the lobes somewhat ventrally de-

flexed, the insertion strongly incubous; leaves

330-390 (Jim wide X 315-435 jxm long, moder-

ately asymmetric (the dorsal margin shorter), 4(5)

lobed to 0.4-0.45, the lobes feebly divergent,

shorter than the disc. Leaves on branches closely

and regularly imbricate, the ventral lobes subfal-

cate, appearing as if brushed toward shoot apex.

Lobes acute to weakly acuminate, the dorsal lobe

shortest, the 2 ventral lobes largest, somewhat

paired and often subfalcate (especially on branch-

es), the largest leaf lobes (4)5-8 cells wide at ex-

treme base, terminating in a short uniseriate row

of 2-4 cells (ca. half or less the length of lobe);

lobe cells short, ± isodiametric to short rectan-

gular, thick-walled, the cell walls of uniseriate

row somewhat thickened in the corners but the

septae not swollen and projecting at the lobe mar-

gin, the basal cell of the uniseriate portion 15-20

u>m wide X 17-23 |xm long (1.1-1.4:1), the next

cell 12-17 |xm wide X 13-21 u,m long (1.1-1.4:

1), the terminal cell a little shorter than the pen-

ultimate cell, the apex ± rounded; cuticle smooth

to finely striate-papillose at lobe tips. Disc sub-

symmetrically quadrate-rectangular to obliquely

trapezoidal, 6-9 cells high (8-10 cells high in

branch leaves), 16-19 cells wide in distal portion

narrowing to 8-9 cells wide in basal portion; mar-

gins entire, the dorsal straight to weakly incurved,

the ventral margin rounded. Cells of disc small,

evenly thick-walled and firm, trigones lacking, the

median cells elongate, 16-26 u.m wide X 30-40

u,m long, often elongated longitudinally, follow-

ing the contour of the leaf; basal row of disc cells

larger (especially in ventral-basal sector); cuticle

smooth. Underleaves much smaller than leaves,

0.9-1 X stem width, strongly spreading, distant,

plane, 4-lobed to 0.5-0.6, the lobes somewhat di-

vergent, narrowly acuminate, the uniseriate por-

tion formed of 3-4 short cells, terminating in a

slime papilla; disc symmetrically subrectangular

(wider than high), 3(4) cells high (median sinus),

the cells ± regularly arranged, the disc 8 cells

wide; margins entire, usually straight. Asexual re-

production lacking.

Androecia and gynoecia not seen.

DIFFERENTIATION AND VARIATION — The highly

regular, neatly combed appearance of the branches

(Fig. 4: 1), and the characteristic asymmetry of

the leaves — the dorsal lobe shortest, and the ±

paired ventral lobes appearing brushed toward the

branch tips (Fig. 4: 3, 4) — will immediately dis-

tinguish this apparently rare species. The leaves

of T. clatritexta of Western Australia are similarly

asymmetric, but not to such a marked degree (cf.

Fig. 1: 1, 4, 5). Many of our Telaranea taxa have

asymmetrical Lepidozia-\ike leaves (e.g., T. pal-

udicola), with the dorsal lobes paired and often

distinctly larger than the ventral. The leaves in T.

pennata, however, are almost a mirror-image of

the typical Lepidozia-type leaf.

The leaf lobes in T. pennata are broader than

in any other New Zealand Telaranea (up to 8 cells

wide at the base). The disc, however, is shorter

than in T. clatritexta, 6-9 cells high (Fig. 4: 3)

vs. 8-10(14) cells high in in T. clatritexta.

DISTRIBUTION AND ECOLOGY — Known only from

the type locality, a dryish forest in Westland

Prov., where the species occurs both over rock

and on fallen logs in a stream bed.

SPECIMEN SEEN— NEW ZEALAND. SOUTH IS-

LAND. WESTLAND PROV.: Route 73, 8 miles W of

Turiwhate, Engel 6765 (F).

Telaranea quadristipula (Steph.) Engel & Merr.,

comb. nov.

Lepidozia quadristipula Steph. in Stephani & Watts,

J. & Proc. Roy. Soc. New South Wales 48: 116.

1914. Type: Australia, New South Wales, Rotunda,

Neate's Glen, Blackheath, 4 Jan. 1911, Watts 1009

(G!).

Plants flexuous, in loose mats, pale green, dis-

tinctly nitid when dry; plants medium, to 8 mm

20

FIELDIANA: BOTANY

FIG. 4. Telaranea pennata Engel & Merr. 1. Sector of main shoot with f-'rullania-lype branches, dorsal view. 2.

Leaves (dorsal lobe [= dl] at right) and to right an underleaf (= ul). 3. Leaf, cellular detail (dl - dorsal lobe). 4.

Sector of branch, ventral view. 5. Underleaf, cellular detail. 6. Branch leaves (= bl; dorsal lobe at right). 7. Stem,

cross section. (All from holotype.)

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

21

wide, including branches. Branching ± regularly

1 -pinnate, the branches of the Frullania-type, at

times abruptly flagelliform; branch half-leaf bifid

to 0.4, inserted a little above branch axil; first

branch underleaf undivided, subulate, with up to

4 biseriate tiers and a uniseriate row of up to 3

cells, ventral-lateral in position on branch base.

Ventral-intercalary branches occasional, leafy.

Stem cortical cells thick walled in surface view,

in 14 rows, in cross section the outer wall thick-

ened, a distinct cortical layer confined to the dor-

sal side of stem (ventral cells smaller and thicker

walled); medullary cells ca. 37, smaller, firm

walled, the outermost medullary cells on ventral

side of stem somewhat thicker walled. Leaves on

main shoot obliquely spreading, contiguous, the

insertion strongly incubous, the leaves ± sym-

metrically 4-lobed to 0.4, the disc cuneate, con-

spicuously narrowed to the base, complanate,

plane to weakly convex, the lobes broadly acu-

minate, shorter than the disc, the lobe tips ven-

trally decurved; branch leaves imbricate, subfal-

cate, appearing "combed" toward the apex, on

stronger branches 4-lobed, the dorsal lobe(s)

smaller and more slender, the ventral lobe + ven-

tral disc margin broadly arched. Lobes narrowly

acute, often ± abruptly narrowed in the upper

half, 4(5) cells wide at extreme base, with up to

3 additional 4-celled tiers, then contracted to bi-

seriate at the base of the uniseriate row of 3-4

cells; branch leaves with dorsal 1-2 lobes more

slender, 2-3 cells wide at base, with 1-2 biseriate

tiers or 2 cells wide at the base and otherwise

uniseriate; lobe cells isodiametric to short rect-

angular, firm-walled, about equal to the disc cells

or a little smaller, the cells of uniseriate row some-

what thickened in the corners, straight or weakly

constricted at the septa, the terminal cell normally

about equal to the penultimate cell in length or a

little shorter, the apex rounded; cuticle of lobe tips

indistinctly papillose. Disc rather broadly cuneate,

(4)5-6(7) cells high (from median sinus base to

leaf base), 15-16 cells wide in distal portion nar-

rowing to 8 cells wide at the insertion, the mar-

gins entire; disc of branch leaves asymmetrically

cuneate, (5)6-7 cells high, the dorsal margin lon-

ger than the ventral and ± straight, the ventral

margin broadly arched, not decurrent on the stem.

Cells of disc moderately thick-walled, in ± irreg-

ular rows, 16-24 u,m wide X 23-31 jxm long, the

basal row of disc cells often conspicuously larger,

about twice the width of the cells of the disc prop-

er, which are evidently smaller by vertical subdi-

visions; cuticle smooth. Underleaves about the

same width as the stem or a little less, obliquely

spreading, plane, 4-lobed to 0.5 or a little less;

lobes 2-3 cells wide at base, at times with an

additional biseriate tier, ending in a uniseriate row

of 2-3 short cells, terminating in a slime papilla;

disc parallel-sided, 3 cells high, 8 cells wide. Rhi-

zoids originating from distal portion of disc.

Androecia and gynoecia not seen.

DIFFERENTIATION AND VARIATION — This appar-

ently rare Australian species has features sugges-

tive of both T. pennata and T. tridactylis, among

them the small cell size (16-26 u,m wide) and the

flattened, ribbon-like branches, with a character-

istic reversed asymmetry of the branch leaves: the

dorsal lobe(s) smaller, the ventral larger and

broadly arched (Fig. 5: 3, 9), the leaves appearing

brushed toward the branch tips. They differ chief-

ly in the differentiation of the branch leaves from

those of the main axis and the height of the leaf

disc. In T. pennata both the stem and branch

leaves show the characteristic asymmetry, but in

T. quadristipula the leaves on the main shoot are

± symmetrically 4-lobed (Fig. 5: 1, 4). The stem

leaves of T. tridactylis are highly variable, but

characteristically include those which are ± lon-

gitudinally inserted and deeply divided (disc as

little as 3 cells high), with distinctive, widely

splayed lobes. The lobes of T. tridactylis are also

more slender: 2-4 cells wide at the base, with 1-

2 biseriate tiers. In T. quadristipula the leaf disc

is (4)5-6(7) cells high (Fig. 5: 1,4) vs. 6-9 cells

high (8-10 cells high in branch leaves) in T. pen-

nata. An interesting feature of the leaf lobes of T.

quadristipula is the often abrupt narrowing from

several 4-seriate tiers to biseriate at the base of

the uniseriate row (Fig. 5: 1,5).

Telaranea disparata of Queensland has branch

leaves with reversed symmetry, but the disc cells

are larger (26-32 (im wide), the disc is 8 cells

wide throughout, and the lobes are biseriate at the

base.

NOTE — The specific epithet, "quadristipula," is

probably not what Stephani originally intended.

The type (Watts 1009, cited in the protologue) is

labeled "Lepidozia quadratistipa" ; the Icones

(Lepidozia 108, Australia, leg. Watts) is labeled

"Lepidozia quadristipa." None of these names is

particularly informative.

DISTRIBUTION AND ECOLOGY — Known only from

the type. New South Wales; admixed are Zoopsis

sp., Balantiopsis sp., and the moss Distichophyl-

lum sp.

22

FIELDIANA: BOTANY

10

FIG. 5. Telaranea quadristipula (Steph.) Engel & Merr. 1. Portion of main shoot with a Frullania-type branch

(= FB; hi = half-leaf); dorsal aspect. 2. Portion of shoot showing base of terminal branch (fbu = first branch underleaf;

bl = branch leaf base); base of main shoot leaf included at right. 3. Portion of primary, Frullania-lype branch, dorsal

aspect; note the dorsal pair of lobes are narrower and often shorter. 4, 5. Leaves of main shoot (both drawn to same

scale). 6, 7. Median lobes of leaf, a complete lobe at left, only the distal portion shown at right. 8, 9. Branch leaves;

note the two dorsalmost lobes (= dl) are smaller and narrower. 10. Stem, cross section. (All from type.)

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

23

Telaranea tridactylis (Lehm. & Lindenb.) Engel

& Merr.

Jiingermannia tridactylis Lehm. & Lindenb. in Leh-

mann, Nov. Min. Cogn. Stirp. Pug. 4: 41. 1832.

Lepidozia tridactylis (Lehm. & Lindenb.) Mont, in

Dumont d'Urville, Voy. Pole Sud, Bot. 1: 245.

1845. Telaranea tridactylis (Lehm. & Lindenb.)

Engel & Merr., Phytologia 79: 253. June, 1996

[1995]. Type: "Nova Hollandia," without specific

loc., sin. coll., "Hb. Berol[inensis] 46" (W!, Lin-

denberg Hepat. no. 4636).

Lepidozia oldfieldiana Steph., Spec. Hep. 3: 581.

1909, syn. nov. Lectotype (nov.): Tasmania, without

specific loc., Oldfield, ex hb. Kew (G!— c. 6*).

Plants rather stiff and wiry, prostrate in com-

pact mats, light olive green, highly nitid when

dry, not water repellent; shoots very small for

subgenus, the main shoots 350-560 |xm wide (0.5

cm wide with branches). Branching irregularly to

regularly pinnate, occasionally locally 2-pinnate,

the branches of the Frullania type, rather short (to

moderately long but normally determinate), at

times flagelliform; branch half-leaf bifid, trans-

versely to obliquely inserted, the lobes sometimes

widely divergent; first branch underleaf undivid-

ed, subulate, inserted on ventral or ventral-lateral

side of branch near juncture of branch and main

axis. Ventral-intercalary branches common, leafy

or sporadically flagelliform. Stems with cortical

cells in 12-13 rows, often rather thick-walled and

firm in surface view, in section larger than the

medullary cells, the radial walls somewhat thick-

ened, the exposed wall at times distinctly so; med-

ullary cells in ca. 38 rows, the cells much smaller,

with thin to very thick walls. Rhizoids sparse,

from distal cells of underleaf disc. Leaves on main

shoot rigid, fragile, widely spreading, but not at

right angles to stem, loosely imbricate to conspic-

uously distant, plane or slightly concave resulting

from occasional gentle incurving of lobes, the in-

sertion variable: subtransverse to strongly incu-

bous to almost longitudinal; leaves (145)265-385

u,m wide X (120)235-335 |xm long, subsymme-

tric, 4- (rarely 5-6-) lobed to 0.45-0.6, the lobes

moderately to widely (and at times asymmetrical-

ly) divergent (on more distant leaves on the main

axis the lateral lobes often widely spreading,

forming an angle of 180° with each other); branch

leaves typically closely and regularly imbricate,

weakly asymmetrical. Lobes narrowly attenuate,

tapering from 2-4 cells wide at extreme base fol-

lowed by 1-2(4) biseriate tiers, terminating in a

uniseriate row of (2)3-5 cells; cells of the uni-

seriate portion short to weakly elongate, moder-

ately thick-walled, the basal cell of the uniseriate

portion 19-26 jxm wide, (24)28-34 |xm long, (1-

1 .6: 1 ), the next cell shorter and on the whole nar-

rower, being 14-19(24) (xm wide and 19-25 |xm

long (1-1.3:1), the terminal cell about the size of

the penultimate cell or a little shorter, tapering to

the tip. Disc ± symmetrically weakly cuneate, 3-

5 cells high (disc of branch leaves to 6(7) cells

high), 11-16 cells wide in distal portion narrow-

ing to (7)8(9) cells wide in basal tier; margins

entire, straight to broadly curved. Cells of disc

typically evenly and moderately thick-walled, the

areolation rather compact and lacking the regular

arrangement of other Telaranea species (second-

ary divisions of disc cells random in both planes);

median cells small, 18-23 |xm wide, 26-38(42)

(xm long; basal cells considerably larger and

forming an obvious tier; cuticle with a network of

fine irregular striae. Underleaves much smaller

than leaves, spreading nearly or at 90° to stem,

distant, slightly convex (ventral view), 3-4 lobed

to 0.45-0.55, the lobes often gently curved dor-

sally, narrowly attenuate to subciliiform, 2 cells

wide at the base, terminating in a uniseriate row

of 2-3 slightly elongated, somewhat thick-walled

cells, or consisting entirely of the uniseriate row,

terminating in a slime papilla; disc weakly cune-

ate, 2-3(4) cells high (median sinus), the cells in

± regular tiers; disc 9-10 cells wide in distal por-

tion (6-8 cells wide in 3-lobed underleaves), nar-

rowing to 8 cells wide at base (6 cells wide in 3-

lobed underleaves), the basal row of cells on the

whole longer and forming an obvious tier; mar-

gins entire, straight. Asexual reproduction lack-

ing.

Dioecious. Androecia either on short Frullania-

type branches with a few to several (to 5) cycles

of normal vegetative leaves prior to androecial

formation, or on short, abbreviated, ventral-inter-

calary branches lacking normal vegetative leaves;

bracts closely imbricate, strongly dorsally assur-

gent, the entire bract deeply concave, 2(3) lobed,

each lobe terminating in a uniseriate row of 2-3

not to hardly elongated cells; lamina cells non-

tiered, irregular in shape and arrangement, the

lamina margins sometimes denticulate, often with

stalked or sessile slime papillae; antheridia 1 per

bract, large for bract size, the stalk short, 5-6 cells

high, uniseriate; bracteolar antheridia absent. Gy-

noecia strongly dorsally assurgent, with a vestigial

stem perigynium barely present, not swollen, rhi-

zoidous; bracts and bracteoles in 3 series, inserted

on the vestigial perigynium; bracts becoming pro-

gressively larger and less deeply lobate towards

24

FIELDIANA: BOTANY

the perianth, those of innermost series deeply con-

cave, with apices rounded, irregularly denticulate

with sporadic 1-2-celled teeth; bract margins

crenulate to irregularly repand, with a few sessile

slime papillae, the bracts with a border formed of

1 row of cells longer, narrower and more irregular

than those within; bracteole nearly identical in

size and form. Perianth not extending above veg-

etative axes and entangled by them. 0.65-0.8

emergent, narrowly fusiform, 5-7. 2X longer than

wide, terete in basal half, the distal half obscurely

trigonous and with 5-6 deep plicae, gradually and

distinctly narrowing toward the contracted mouth;

mouth with 6 narrowly triangular lobes, each lobe

fringed with thick- walled, contorted, coarsely pa-

pillose cilia, the cilia consisting of 2-4 elongated

cells with thickened and swollen septa, often with

a knob-like swelling at the basal end of the cells

just above the septa; perianth 2-4 stratose in basal

portion, the median portion 2 (locally 1 ) stratose.

Seta with 8 rows of outer cells (each with their

free face bulging), surrounding an inner core of

17-18 much smaller thin-walled cells, with corner

thickenings resembling bulging and knotlike tri-

gones. Capsule rather short elliptic, the valves

980-1050 u,m long, the wall 24-30 u,m thick, of

3 layers, the outer layer subequal to the combined

2 inner layers, or slightly less thick; outer layer

of cells in tiers, rather regularly short-rectangular,

with 2-phase development, the longitudinal walls

with well-defined sheetlike thickenings and nod-

ule-like thickenings (4-6 per cell) alternating with

walls that are devoid of thickenings (or with very

few sporadic, local, nonpigmented, nodular swell-

ings), the transverse walls devoid of thickenings;

intermediate layer thinner than outer or inner lay-

ers; innermost layer of cells ± tiered, irregularly

narrowly to broadly rectangular, with semiannular

bands rather narrow, ± irregular, short or infre-

quently complete, rarely forked and anastomosing

to delimit ill-defined, local fenestrae.

Spores 14.9-16.8 u,m, exine yellow brown,

with low, delicate papillae, coalescing to form

short ridges that rarely delimit areolae. Elaters rig-

id, somewhat sinuous, (7.7)8.6-10.1 u,m wide,

only slightly tapering toward tips, bispiral to tips,

the spirals 2.9-3.4 u,m wide.

DIFFERENTIATION AND VARIATION — This species

is the smallest member of sect. Neolepidozia. It is

distinguished from T. patentissima by its smaller

size, smaller leaf cells (at most 25 u,m wide), and

less regular arrangement of disc cells, vs. 25-40

|xm wide in T. patentissima, with a more T. tet-

rapila-like areolation of rather regular rows and

tiers.

The species shows considerable variation in

leaf form and insertion. When optimally devel-

oped, the branches are complanate and ribbon-

like, and broader than the main shoot, with closely

shingled leaves (Fig. 6: 3, 4). The branch leaves

have a rather high disc (to 7 cells high) and are

often 4-lobed. By contrast, the leaves of the main

shoot are often distant, longitudinally-inserted,

and deeply divided, with widely splayed lobes,

often oriented as much as 180° to each other

(Figs. 6: 7; 7: 2). Weaker shoots may have both

stem and branch leaves splayed-lobed, and lack

the flattened, ribbon-like branches (Fig. 6: 1, 2).

The leaves of the flattened, ribbon-like branch-

es are suggestive of those of T. pennata (p. 18)

in that the ventral lobes are weakly arched toward

the apex of the branch, and the dorsal lobes more

slender than the ventral. Telaranea pennata, how-

ever, is a more robust plant, with the comb-like

branches to 775 u,m broad, with leaf lobes to 8

cells wide at the base. Both stem and branch

leaves tend to show the characteristic asymmetry,

and the stems do not have the small, longitudi-

nally inserted, splayed-lobed leaves as in T. tri-

dactylis. The disc of T. pennata is higher (6-9

cells high), and the disc cells are longitudinally

elongate. For further comparisons of T. tridactylis

with T. pennata and T. quadristipula, all species

with small disc cells in the range of 16-24 |xm

wide, see comments under T. quadristipula.

DISTRIBUTION AND ECOLOGY — The type of T. tri-

dactylis is from Australia (Nova Hollandia). It is

a common species in Tasmania, but in New Zea-

land is known only from Auckland and Campbell

Is. and one station at Milford South on South Is-

land. Reports of this species as T. capilligera from

mainland New Zealand are based on misdeter-

minations (confirming Hodgson, 1956, p. 610, sub

Lepidozia gottscheana). Conversely, T. patentis-

sima (p. 46), also a small plant, is common in

New Zealand, but rather rare in Tasmania. The

species is also known from Victoria. Australia and

one station in Queensland. The species was re-

ported for the Strait of Magellan by Montagne

(1845) and Kiihnemann (1937, 1949). We have

not seen the collections on which these reports are

based.

Telaranea tridactylis is primarily a forest plant.

In Tasmania the species occurs, for example, in

lower elevation forests such as of Acacia melan-

oxylon and Leptospermum lanigerum or of Lep-

tospermum scoparium-Eucalyptus in the north-

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

25

FIG. 6. Telaranea tridactylis (Lehm. & Lindenb.) Engel & Merr. 1-5. Portion of main shoot with one or more

Frullania-lypc branches, dorsal view (note especially the variation in orientation and planation of branch leaves and

the imbrication of main shoot leaves). 6-8. Leaves. 9. Leaf lobe. (Figs. 1 , 6, 7, 9 from type of T. tridactylis; 2 from

Norris 29804, Tasmania, Arve Road; 3, from Norris 27281, Tasmania, above Lake Harrington near Forth Falls; 4, 8

from Engel 19725, Tasmania, Newhaven Road, just S of Doughboy Hill; 5 from Norris 29470, Tasmania, Kermandie

River at North Creek.)

26

FIELDIANA: BOTANY

FIG. 7. Telaranea tridactylis (Lehm. & Lindenb.) Engel & Merr. I . Main shoot, with four Frullania-typc branches,

dorsal view. 2, 5. Leaves. 3. Rhizoid with ramified tip. 4. Leaf lamina cross section. 6. First branch underleaf. 7. 9

Bracts (top to bottom), from innermost, second, and lowermost series. 8. Antheridial stalk. 9. Underleaf. 10. Half-

leaf (b = adjoining branch cell; s = adjoining stem cell). 1 1. Distal portion of innermost female bract. 12, 13. Leaf

outlines. 14. Portion of main shoot, dorsal view with only leaf (L) and half-leaf (HL) bases indicated (FB = Frullania-

type branch). 15. Stem, cross section + underleaf. 16. Portion of main shoot, dorsal view, showing leaf arrangement

(note six male branches toward base). (AH from Norris 29470, Tasmania, Kermandie River.)

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

27

western sector of the state. On the east coast it

may occur in forests of Eucalyptus obliqua to-

gether with varying combinations of other vas-

cular plant forest species. On Mt. Raoul (380 m),

for example, it is found in a forest consisting of

E. obliqua over Pomaderris apetala, Olearia ar-

gophylla, Bedfordia salicina, Hakea lissosperma,

Pimelea nivea, P. drupacea and Coprosma quad-

rifida. The species also occurs in middle-elevation

Nothofagus-Eucalyptus-Atherosperma or Notho-

fagus cunninghamii-Eucryphia forests, as well as

upper-elevation forests of Nothofagus-Athrotaxis.

In forests the species often occurs on bryophyte-

covered logs (especially where some soil has ac-

cumulated), or less frequently, is terricolous or

saxicolous on outcrops in niches where soil has

acumulated. The species is rare and sporadic in

subalpine and alpine zones. For example, it occurs

on soil deep under thicket cover in subalpine

scrub of Richea scoparia, Nothofagus gunnii, Di-

selma and Athrotaxis selaginoides, etc. In alpine

areas it occurs over rock at the margins of creeks

or with Astelia at margins of seepage areas.

The plant from Youngs Creek, Victoria, oc-

curred in a cool temperate rainforest with Noth-

ofagus cunninghamii and Acacia melanoxylon co-

dominant, Eucalyptus regnans emergent, and

Blechnum wattsii and Dicksonia antarctica the

main understorey species. At this site the species

occurred on the base of Dicksonia antarctica. The

Scott specimen from Gippsland was found on the

trunk of Nothofagus cunninghamii in a cool rain

forest.

The plant from Milford Sound, New Zealand

occurred on exposed, moist, vertical cliffs with a

dense bryophyte cover at the forest margin.

SELECTED SPECIMENS SEEN— CAMPBELL IS.: Bee-

man Hill, "Meterological Party" as T. dispar (CHR).

AUCKLAND IS.: Hooker sub Jungermannia dispar

(BM, FH); Ewing Is., Fineran 1358 as T. patentissima

(CHR). NEW ZEALAND. SOUTH IS., SOUTHLAND

PROV.: Fiordland Natl. Park, Milford Sound, track to

Bowen Falls, sea level, Engel 22007 (F). TASMANIA:

Waterfall Creek State Reserve, South Bruny Range, 100

m, Moscal 25217 as T. praenitens (HO); E side of Tas-

man Peninsula, upper slope of Tatnells Hill, S of Wa-

terfall Bay, 500-530 m, Engel 13164 (F); Cape Raoul

State Reserve, Mt. Rauol, 380 m, Moscal 24890 as T.

tetradactyla (HO); Sandspit River, 8.5 km WNW of

Cape Bernier, 180 m, Moscal 16838 as T. tetradactyla

(HO); Humbug Point State Recreation Area, Higgin-

bothams Creek, sea level, Moscal 25655 as T. tetradac-

tyla— c. per. (HO); south coast, Deadmans Bay, 10 m,

Moscal 14180C (2), both as T. tetradactyla (HO);

Adamsons Track, 150 m, Ratkowsky HI 540 as T. gotts-

cheana (HO); near Manuka Flat on trail to Adamsons

Peak, 300-400 m, Norris 26878 (F); Adamsons Peak,

below hut, Ratkowsky 80/152 (HO); near Jacques Creek

near end of Styx River Road, Norris 29037 (F); Ker-

mandie River at North Creek, ca. 50 m, Norris 29470

(F); Arve Road ca. 1 mile W of Willies Saddle, ca. 300

m, Norris 29804 (F); Tahune Forest Reserve, near Huon

River, 70 m, Engel 19860 (F); E facing slope of Mt.

Wellington, O'Gradys Falls, 525 m, Engel I2752B (F);

Mt. Wellington, 305-365 m, Rodway s.n. as both as L

praenitens (HO); ibid., 610 m, Ratkowsky B79e (F);

ibid., "Pipeline," above Silver Falls, Ratkowsky H1676

as T. gottscheana (HO); Glenorchy Water Reserve, 400

m, Moscal 19900 as T. praenitens (HO); Mt. Drome-

dary, Dean Brook, Weymouth 851 as L. praenitens — c.

S (HO); Myrtle Forest, SW of Collinsvale and ca. 2.3

km N of Collins Bonnet, 700 m, Engel 12773 (F); Mt.

Field Natl. Park, Lake Belcher, ca. 900 m, Norris 28690

(F); Mt. Wedge, Ratkowsky H1539 as T. gottscheana

(HO); ibid., Rodway Range, between Rodway Ski Tow

and K Col area, 1240-1310 m, Engel 14415 (F); Main-

waring River, 1 km from mouth, sea level, 1 m, Moscal

9768 A as T. praenitens (HO); near Scotts Peak Road,

2.2 km S of junction with Gordon River Road, E of S

end of Lake Gordon, 580 m, Engel 13800 (F); Serpen-

tine River, below Lake Pedder dam, Scott & P. Dalton

s.n. as T. tetradactyla (MELU); Cradle Mtn.-Lake St.

Clair Natl. Park, Overland Track between Watersmeet

and Echo Point, W side of Lake St. Clair, 760 m, Engel

14207 (F); Mt. King William I, 1200 m, Ratkowsky

H1538 as T. gottscheana (HO); Surprise Valley, above

both Surprise River and Lyell Highway, 2.1 km W of

King William Saddle, 750 m, Engel 19446 — c. sporo.

(F); Frenchman's Cap, 1893, Moore 40, syntype of Lep-

idozia oldfieldiana (G); Franklin River at Frenchman's

Cap Trail crossing, ca. 400 m, Norris 31157 (F); west

coast, Sophia Point, Moore s.n. as L. praenitens (HO);

Zeehan-Renison Bell State Reserve, on Murchison

Highway 6.7 km N of intersection with road to Zeehan.

250-320 m, Engel 20079 (F); Mt. Read, S of Rosebery,

990-1010 m, Engel 20040 (F); Wilson River, 26 km N

of Zeehan, 140 m, Moscal 21606 as T. tetradactyla

(HO); Alfred River, 200 m, Moscal 21869 as T. tetra-

dactyla (HO); Mersey River, 2.5 km S of Lewis Falls,

750 m, Moscal 75267 as T. praenitens (HO); Blue Tier,

Rodway s.n. as T. praenitens (HO); Cradle Mt., Rodway

s.n. (2), both as L. praenitens (HO); ibid., near Lake

Dove, Rodway s.n. as L. praenitens (HO); Cradle Mtn.-

Lake St. Clair Natl. Park, Weindorfers Forest, along

track from Waldheim Chalet to Hounslow Heath Track,

975 m, Engel 14058 (F); ibid.. Cradle Mt. area, Plateau

Creek, between Overland Track and Kitchen Hut Track,

W of N end of Lake Dove, 1250 m, Engel 19659 (F);

ibid., above Pencil Pine Creek and off track between

Quailes Hill and Pencil Pine Lodge, E of Pencil Pine

Lodge, 770-800 m, Engel 19620 (F); above Lake Bar-

rington near Forth Falls, ca. 150 m, Norris 27281 (F);

Foot Mt. Roland, Rodway s.n. as L. praenitens (HO);

Newhaven Road, just S of Doughboy Hill, 0.4 km E of

junction with Mawbanna Road, S of Port Latta, 45 m,

Engel 19725 (F); Detention Falls, SSW of Sisters Creek,

W of Wynyard, 190 m, Engel 19673 (F); Ferndene, Nor-

ris 33970 (F); Deadwood, Grafs Tree Hill, Oldfield

29(H) (G); Flinders Is., Mileara Valley, Bob Smith's

Gully, Scott s.n. as T. dispar (MELU). AUSTRALIA.

VICTORIA: Otway Ranges, Youngs Creek, Phillips

Track, Turner s.n. as T. dispar (F); Gippsland, West

28

FIELDIANA: BOTANY

Branch Creek, Gunyah, Scon s.n. as T. patentissima (F,

MELU); Errinundra Plateau, track down to First Creek

Falls, Scott & Chesterfield s.n. as T. dispar — c. sporo.

(MELU); Welshpool, near Gunyah Junction, Scott s.n.

as T. Idispar — c. cJ (MELU); Marysville, Talbot Drive,

Scott s.n. as T. tdispar (MELU). QUEENSLAND: Mi-

mosa Creek, Blackdown Tableland, Stone s.n. (MELU).

Telaranea consobrina Engel & Merr.

Telaranea consobrina Engel & Merr., Novon 9: 339.

/ /. 1999. Holotype: Tasmania, eastern slope of

Black Bluff just below summit, S of Burnie, 1250

m, Engel 15799 (F); isotype: (HO).

Plants soft and flexuous, ascending to suberect,

in soft cushions, pale green, distinctly nitid when

dry; plants medium, to 6 mm wide, including

branches. Branching rather regularly 1 -pinnate,

the branches of the Frullania-iype, at times be-

coming flagelliform; branch half-leaf 2-lobed,

obliquely inserted, narrowly rectangular, the lobes

± parallel to slightly diverging; first branch un-

derleaf undivided (very rarely bilobed), broadly

acuminate to lanceolate, inserted on ventral side

of branch at base of branch. Ventral-intercalary

branches not seen. Stems with cortical cells dis-

tinctly differentiated, thin-walled, in 13-14 rows;

cortical cells in section slightly to distinctly larger

than the numerous (49-53) medullary cells, the

medullary cells slightly thick walled. Leaves on

main shoot obliquely spreading, contiguous, the

disc plane or weakly convex, the lobes ventrally

decurved, the insertion distinctly incubous, the

disc broader than high, nearly in same plane as

dorsal surface of stem; leaves 475-600 u.m wide

X 440-525 \im long, the leaves 4(5)-lobed to

0.4-0.5, the lobes straight to moderately diver-

gent, slightly shorter than the disc. Lobes narrow-

ly acute to acuminate, 4-5 cells wide at base, of-

ten 4 cells wide in basal sector, then biseriate for

2-3 tiers, terminating in a short uniseriate row of

2-3 cells (or sporadically a single cell or 2 later-

ally juxtaposed cells), the tip sometimes with a

slime papilla; lobe cells ± isodiametric to short

rectangular, thin-walled, the cell walls of uniser-

iate row not or weakly thickened in the corners,

the basal cell of the uniseriate portion 20-25 u,m

wide X 24-31 jim long (1-1.3:1), the next cell

17-22 u,m wide X 18-26 u.m long (1-1.2:1), the

terminal cell normally about equal to the penul-

timate cell in length or a little shorter, the apex

rounded; cuticle smooth. Disc somewhat asym-

metrically cuneate, the dorsal margin somewhat

longer than the ventral, the disc (5)6-7(8) cells

high (from median sinus base to leaf base), 13-

16 cells wide in distal portion narrowing to 8(9)

cells wide in basal portion; margins entire, the

dorsal margin weakly to broadly arched, the ven-

tral margin ± straight to moderately arched, dis-

tinctly decurrent on the stem. Cells of disc thin-

walled, trigones lacking, the cells of the distal half

to one-third smaller by vertical subdivisions or all

but the basal row of disc cells smaller, the smaller

cells 16-22 (Jim wide, the largest 24-38 u,m wide

X 42-49 |xm long, the cells in ± irregular rows;

cuticle smooth. Underleaves somewhat smaller

than leaves, 1.7-2.3X stem width, obliquely

spreading, distant, plane, 4(5)-lobed to 0.4-0.45,

the lobes straight to moderately divergent, nar-

rowly acute, 3-4 cells wide at base and biseriate

for 1-2 tiers, ending in a uniseriate row of 2-3

short cells, terminating in a slime papilla; disc

symmetrically broadly cuneate (wider than high),

4-5 cells high (median sinus), 14-18 cells wide

at widest point, the cells in ± irregularly rows;

margins entire, moderately arched. Rhizoid initial

cells small, subquadrate, in distal portion of disc

and basal cells of lobe. Asexual reproduction

lacking.

Androecia and gynoecia not seen.

DIFFERENTIATION AND VARIATION — This rare

species of Tasmania and Victoria resembles T.

meridiana of New Zealand in being rather soft in

texture, with broad leaves and ventrally decurved

leaf lobes in situ, and disc cells arranged in some-

what irregular rows (Fig. 8: 1-3). The epithet con-

sobrina ("cousin") was chosen to evoke this re-

semblance. It differs most notably in the shape of

the leaf disc, which narrows to 8 cells wide at the

insertion (Fig. 8: 1) vs. 14-16 cells wide in T.

meridiana (Fig. 2: 3, 5), and in the narrower

lobes, which are only 4-5 cells wide at base (Fig.

8: 1-3), vs. as many as 6(8) cells wide in T. mer-

idiana (Fig. 2: 3, 4). The underleaves of T. con-

sobrina are larger, roughly twice the width of the

stem (Fig. 8: 6), and somewhat shorter (vs. 1.2X

the stem diameter in T. meridiana).

DISTRIBUTION AND ECOLOGY — Known only from

Tasmania and Victoria, Australia. The species is

known from a few stations in Tasmania, two in

the northwest, both at 1250 m, occurring in the

crevice of a dripping cliff face in an area with

alpine vegetation (type) and in the bed toward the

side of Plateau Creek, the area within a mosaic of

cushion plants, Diselma, etc., scattered pools and

small streams. The Moore plant is from a consid-

erably lower elevation on the west coast. On Flin-

ders Island the species occurs on creek banks in

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

29

200 n

FIG. 8. Telaranea consobrina Engel & Merr. 1. Leaf, in situ, dorsal view (sp = slime papilla). 2-4. Leaves (sp

= slime papilla). 5. Leaf lobe. 6. Sector of main shoot with Frullania-type branch (= FB; FBU = first branch

underleaf), ventral view. 7. Underleaf; note small-celled rhizoid initials in distal part of disc and in basal portion of

a lobe. 8. First branch underleaves. 9. Stem, cross section. (All from holotype.)

30

FIELDIANA: BOTANY

a blue gum/tea tree forest. Also known from a

single site in Victoria (Grampians Natl. Park),

where it is found on wet rocks adjacent to water-

falls.

SPECIMENS SEEN— TASMANIA: Cradle Mtn.-Lake St.

Clair Natl. Park, Plateau Creek area, between Cradle

Plateau and Marions Lookout, NNW of Cradle Mtn.,

1250 m, Engel 13960 (F); Macquarie Harbour, Sophia

Point, Moore 134 (HO); Flinders Is., Bass Strait, Cron-

ley's Creek, foot of Ml. Strzelecki, Scott s.n. as T. pa-

tentissima (F, MELU). AUSTRALIA. VICTORIA:

Grampians Natl. Park, Roses Gap, Beehive Falls, Scott

s.n. as T. Ipatentissima (F, MELU).

Telaranea palmata Engel & Merr.

Telaranea palmata Engel & Merr., Novon 9: 344. /

3. 1999. Holotype: Tasmania, ridge SE of Black

Bluff near junction of access road to plateau area

and road to Devonport gold mines, S facing slope,

S of Burnie, 1000 m, Engel 16251 (F); isotype:

(HO).

Plants with a rather stiff and wiry appearance,

densely to loosely interwoven in compact mats,

yellowish green, nitid when dry; shoots small, to

0.6 cm wide, including branches. Branching

somewhat irregularly and loosely l(2)-pinnate,

the branches remaining rather short, normally de-

terminate, not much differing in length (rather

than plumose), the branches of the Frullania-lype,

occasionally to frequently flagelliform; branch

half-leaf 2-lobed, usually obliquely inserted, nar-

rowly rectangular to cuneate, the lobes diverging;

first branch underleaf undivided and subulate, in-

serted on ventral side of branch near base. Ven-

tral-intercalary branches common, leafy, often be-

coming leading shoots. Stems with cortical cells

markedly differentiated, the radial walls thin, the

outer wall somewhat thickened, in 12 rows, those

on ventral side of stem a little smaller; cortical

cells in section larger than the numerous (ca. 50)

medullary cells, the medullary cell walls slightly

thickened and finely pitted. Rhizoids sparse, from

distal cells of underleaf disc. Leaves on main

shoot rigid, the disc widely spreading to squar-

rose, distant to loosely imbricate, moderately con-

cave to hand-like, the lobes erect and incurved, at

times subfalcate, the insertion transverse to weak-

ly incubous; leaves 480-665(700) u,m wide (mea-

sured between tips of lobes) X 400-510 u,m long,

moderately asymmetric, ± equally palmately 4-

lobed to ca. 0.6, the lobes often widely divergent

(the lateral lobe then forming an angle of up to

180° with adjacent median lobe), subequal to

somewhat longer than the disc in length. Lobes

attenuate to subcaudate, 2-4 cells wide at base

(when 4 cells wide sometimes with an additional

3-4 seriate tier), then biseriate for 1(2) tiers, ter-

minating in a uniseriate row of 4-6 cells (typi-

cally more than half the length of lobe); lobe cells

± firm, often distinctly thick-walled (to 7 ftm

thick), the septa thickened in the corners and at

times weakly projecting- from the lobe margins,

the basal cell of the uniseriate row 30-37 u,m

wide X 38-54 u,m long, the next cell 24-31 u,m

wide X 36-48 u,m long, the terminal cell shorter

than the penultimate cell, tapering to a rounded

point; cuticle smooth, the lobe tips rarely weakly

striate-papillose. Disc symmetrically to somewhat

asymmetrically short cuneate, 4 (rarely 5) cells

high (from median sinus base to leaf base), 13-

16 cells wide in distal portion, narrowing to 8

cells wide at the insertion; disc margins entire or

with feebly projecting septa, ± straight. Cells of

disc moderately to distinctly thick-walled, tri-

gones minute or none, the cells in ± regular tiers,

the median and basal disc cells 31-36 u,m wide

X 39-48 jxm long, the distal tiers often longitu-

dinally divided and 21-25 u,m wide (2-2.3:1); cu-

ticle smooth. Underleaves much smaller than

leaves, obliquely to widely spreading, distant,

plane, 4-lobed to ca. 0.5 or a little more, the lobes

divergent, ciliiform, straight, basically 2 cells

wide at the base, the uniseriate portion formed of

3(4) slightly elongated, ± thick-walled cells with

septa thickened in the corners, often terminating

in a slime papilla; disc symmetrically subquadrate

to weakly cuneate, 3(4) cells high (median sinus),

the cells in ± regular tiers; disc 8-1 1 cells wide

in distal portion, 8 cells wide at base; margins

entire, usually straight. Asexual reproduction

lacking.

Plants apparently dioecious. Androecia either

terminal on short, primary, FrulUinia-lype branch-

es with a few cycles of normal vegetative leaves

prior to androecial formation, or on short, excep-

tionally abbreviated, ventral-intercalary, spicate

branches; bracts closely imbricate, strongly dor-

sally assurgent, deeply concave, bilobed to ca.

0.5, the lobes acute, terminating in a uniseriate

row of 2-3 cells, the basal cell not or barely lon-

ger than wide, the terminal cell rather sharp and

variable in length, to 3:1; dorsal margin of lamina

somewhat dilated, crenulate, with slime papillae

single and sessile or lacking; bracts monandrous;

antheridial stalk short, 6 cells high, uniseriate;

bracteolar antheridia absent. Gynoecia not seen.

DIFFERENTIATION AND VARIATION — This species

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

31

is similar to T. patentissima and T. capilligera,

but differs from both in the transverse insertion

of the leaves (Fig. 9: 1 ). It differs from T. paten-

tissima in several respects: the uniseriate row of

the leaf lobes is 4-6 cells long, typically more

than half the length of lobe, vs. 2-3(4) cells long

in T. patentissima; the disc is 4(5) cells high vs.

4-6(7) cells high; the leaves are palmately 4-

lobed to 0.6 vs. more shallowly 4(6)-lobed to 0.4-

0.5 in T. patentissima; and the cells of both disc

and lobes are distinctly thick-walled. It resembles

T. capilligera in the shape of the leaves and in

the thick-walled leaf cells, but differs in the elon-

gate cells of the uniseriate row (to 2.3:1; Fig. 9:

4, 5), vs. barrel-shaped (at most 1.3:1) in T. cap-

illigera (Fig. 3: 3, 4), and in the weakly projecting

septa of lobes (and disc margins) vs. lobe cells

barrel-shaped in T. capilligera, the lobes contract-

ed at the septa.

Telaranea palmata, T. capilligera and T. tri-

dactylis all have at least some leaves with widely

spreading lateral lobes (often at an angle of 180°

or more), but in the latter two species the leaves

of this type are typically almost longitudinally in-

serted and oriented, whereas all the leaves of T.

palmata are transversely inserted or only feebly

incubous.

The leaves of this species are palmately lobed

(to 0.6) like those of T. praenitens (p. 32). Both

species have protruding septa of the lobe and disc

margins, although this character is only weakly

developed in T. palmata. In addition, the cuticle

is smooth vs. distinctly striate papillose in T.

praenitens.

DISTRIBUTION AND ECOLOGY — Endemic to Tas-

mania, and for the most part a subalpine-alpine

species, occurring above 1000 m in protected,

moist niches. It is found, for example, on soil un-

der shrubby cover, as well as on the sides of rills,

or with Lophozia sp., Pachyschistochila parvistip-

ula, Adelanthus sp. and Haplomitrium gibbsiae in

seepage channels. The type is from 1000 m in a

mosaic of Gymnoschoemus (button grass), subal-

pine shrubs, Nothofagus cunninghamii, and rocky

outcrops. The Surprise Valley station, however, is

at 610 m and heavily forested, with Nothofagus

cunninghamii, Eucryphia and Anodopetalum and

an understory of Richea; T. palmata occurred

over humus on the forest floor. In the southwest

it was collected near sea level (Mainwaring River)

in a riverine forest, occurring in carpets of moss,

hepatics, and lichens on a downed Nothofagus

cunninghamii.

SELECTED SPECIMENS SEEN — TASMANIA: Mt. Field

Natl. Park, Tarn Shelf, below and E of Rodway Range,

1270 m, Engel 14358 (F); Mt. Field, Clemes Tarn, 1220

m, Moscal 23024 as T. tetradactyla (HO); Lake Gordon,

Walters (Ratkowsky 79/182) (F, HO); South West Dis-

trict, Mainwaring River, 2 km E of river mouth, 2 m,

Moscal 9671B as T. tetradactyla (HO); Cradle Mtn.-

Lake St. Clair Natl. Park, Lake St. Clair area, between

Mt. Rufus and Mt. Hugel, 1 120-1 130 m, Engel 19363-

c. 6* (F); Surprise Valley, above both Surprise River and

Lyell Highway, 2.6 km W of King William Saddle, 610

m, Engel 19427 (F); Cradle Mtn.-Lake St. Clair National

Park, Weindorfers Forest, along track from Waldheim

Chalet to Hounslow Heath Track, 975 m, Engel 14046

(F).

Telaranea sect. Cancellatae Engel & Merr.

Telaranea sect. Cancellatae Engel & Merr., Phytolo-

gia 79: 250. 1996 (1995). Type: Lepidozia tetrapila

Hook. f. & Tayl.

Telaranea sect. Capillares Engel & Merr., Phytologia

79: 251. 1996 (1995) non Lepidozia sect. Capilla-

res G. L. & N., Syn. Hep. 211. 1845. Type: Lepi-

dozia grossiseta Steph.

Telaranea praenitens (Lehm. & Lindenb.) Hodgs.

Jungermannia praenitens Lehm. & Lindenb. in Leh-

mann, Nov. Min. Cogn. Stirp. Pug. 6: 27. 1834.

Lepidozia praenitens (Lehm. & Lindenb.) G. L. &

N., Syn Hep. 206. 1845. Mastigophora praenitens

(Lehm. & Lindenb.) Trev., Mem. 1st. Lomb. Sci.

Lett. III. 4: 416. 1877. Telaranea praenitens

(Lehm. & Lindenb.) Hodgs., Rec. Domin. Mus. 4:

107. 1962. Type: New Zealand, South Is., Dusky

Bay, Menzies (G!, S! [3 seen]).

Plants with a rather spiny appearance, soft,

flexuous yet firm, loosely prostrate, often in

dense, compact mats, dirty yellowish green, nitid

when dry; shoots small to medium, to 0.9 cm

wide, including branches. Branching mostly reg-

ular and loosely to less often quite densely 1 -pin-

nate, the branches remaining rather short, nor-i

mally determinate, not much differing in length

(rather than plumose), the main axis often bearing

only 2-4 normal leaves (on each side) between

branches, the branches of the Frullania-type, oc-

casionally flagelliform; branch half-leaf bilobed,

usually obliquely inserted, narrowly rectangular to

cuneate (then with lobes diverging), entire (or

with one or both margins toothed in var. dentifol-

ia); first branch underleaf undivided and ciliiform,

inserted on ventral side of branch near juncture of

branch and main axis. Ventral-intercalary branch-

es occasional, often becoming leading shoots and

leafy throughout. Stems with cortical cells mark-

edly differentiated, the radial walls thin, the outer

wall somewhat thickened, in 12-13 rows, those

32

FIELDIANA: BOTANY

FIG. 9. Telaranea palmata Engel & Merr. 1 . Portion of main shoot (the terminal branches not shown, but note

half-leaves), dorsal view. 2, 3. Leaves. 4, 5. Leaf lobes. 6. Underleaf. 7. Antheridial stalk. (Figs. 1-6 from type; 7

from Engel 19363, Tasmania. Cradle Mtn.-Lake St. Clair Natl. Park, between Mt. Rufus and Mt. Hugel.)

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

33

on ventral side of stem a little smaller; cortical

cells in section larger than the numerous (ca. 50-

65) medullary cells, the medullary cell walls

slightly thickened and finely pitted. Rhizoids not

seen. Leaves on main shoot rigid, at least the disc

widely spreading, distant to loosely imbricate,

plane to moderately concave, the lobes somewhat

incurved, the insertion moderately to distinctly in-

cubous; leaves 510-870 (Jim wide X 455-595 u,m

long, moderately to distinctly asymmetric (the

dorsal lobes paired), ± equally and rather deeply

palmately 4-lobed to ca. 0.5-0.65 (very rarely [1

population] 5-6-lobed), the lobes often widely di-

vergent, subequal to somewhat longer than the

disc in length. Lobes attenuate, 4 cells wide for

1-3 tiers at extreme base (5-6 cells wide at base

in var. dentifolia), biseriate for much of their

length, terminating in a short uniseriate row of 2-

4(5) cells (typically less than half the length of

lobe), the lobe margins contracted at the transition

from 4 cells wide to 2 and from 2 cells wide to

1 ; lobe cells ± firm, moderately thick- walled, the

septae thickened in the corners and swollen and

projecting from the lobe margins, the lobe cells

rectangular and not narrowing distally, the cells

in basal portion of lobe 14-20 (Jim wide X 20-30

u,m long, the cells of the median (biseriate) por-

tion 16-23 |xm wide X 25-40 u,m long, the ter-

minal cell of uniseriate row 9-13 |xm wide X 19-

23 (jtm long, tapering to a rounded point; cuticle

finely to distinctly striate-papillose. Disc some-

what asymmetrically short cuneate, 4-5 cells high

(from median sinus base to leaf base), 15-16(19)

cells wide in distal portion narrowing to 8-10

cells wide in basal portion; margins entire (spo-

radically toothed in var. dentifolia), the dorsal

margin often bluntly denticulate by projecting

septa at the ends of cells, ± straight. Cells of disc

moderately thick-walled, trigones minute to small,

the median disc cells 22-38 u.m wide X (38)43-

60 (Jim long, in ± regular tiers or somewhat ir-

regularly arranged; basal 1-2 rows of disc cells

larger; cuticle smooth or faintly striate-papillose

distally. Underleaves much smaller than leaves,

strongly spreading to subsquarrose, distant, plane,

4(rarely 5)-lobed to ca. 0.5, the lobes divergent,

ciliiform, straight, 2-3 cells wide at the base, the

uniseriate portion formed of 3-4(5) elongated, ±

thick-walled cells with thickened septae, often ter-

minating in a slime papilla; disc symmetrically

cuneate, 3-4(5) cells high (median sinus), the

cells in irregular tiers, the disc 11-17 cells wide

in distal portion narrowing to 8-9(12) cells wide

at base; margins entire, usually straight.

Plants dioecious. Androecia either terminal on

short to moderately long primary or secondary

Frullania-type branches or on short, abbreviated,

ventral-intercalary, spicate branches; bracts close-

ly imbricate, strongly dorsally assurgent, deeply

concave, bilobed to ca. 0.4, the lobes acuminate,

terminating in a uniseriate row of 2-4 not to

slightly elongated cells; dorsal margin of lamina

irregularly crenulate, with a few stalked or sessile

slime papillae; bracts monandrous; antheridia

large for bract size, the stalk short, 6 cells high,

uniseriate; bracteolar antheridia absent. Gynoecia

feebly to strongly dorsally assurgent, slightly

swollen and densely rhizoidous at base; bracts

small for perianth size, those of innermost series

closely ensheathing the perianth, concave, broad

ovate to suborbicular, ± regularly 3-4-lobulate,

the lobules terminating in a pair of laterally jux-

taposed cells or a single cell; lamina composed of

± regularly subrectangular cells, the margin bor-

dered by cells of variable shape, often long and

narrow, the apical or free end of marginal cells

often divergent and forming a crenulation, the

margins thus irregularly crenulate, otherwise with

a sporadic unicellular tooth and a few slime pa-

pillae; bracteoles of innermost series similar in

form and size to bracts. Perianth long emergent,

fusiform, terete in basal and median sectors, the

distal sector obscurely trigonous and with 4-5 pli-

cae, the perianth narrowing toward the strongly

contracted mouth; mouth cells thick walled, often

sinuate, partially or wholly laterally free, occa-

sionally with a laterally free uniseriate row of 2

cells, the mouth thus denticulate-subciliate, the

surface cells immediately below mouth often with

their apical end projecting as a blunt, thick-walled

tooth; perianth 2-3-stratose in basal portion, the

median portion 2-stratose.

Seta with 8 rows of outer cells (each with their

free face bulging), surrounding an inner core of

19-20 much smaller cells. Capsule rather long el-

liptic, 980-1260 X 420-560 u, the wall 26-34

u,m thick, of 3 layers, the outer layer subequal to

the combined 2 inner layers, or slightly less thick;

outer layer of cells in tiers, rather regularly short-

rectangular, with 2-phase development, the lon-

gitudinal walls with well-defined sheetlike thick-

enings and nodule-like thickenings (4-6 per cell)

alternating with walls that are devoid of thicken-

ings (or with sporadic, local, nonpigmented, nod-

ular swellings), the transverse walls devoid of

thickenings; intermediate layer thinner than both

outer and inner layers; innermost layer of cells ±

tiered, irregularly narrowly to broadly rectangular,

34

FIELDIANA: BOTANY

with semiannular bands common, rather narrow,

close, usually complete, at times forked and anas-

tomosing to delimit ill-defined, local fenestrae.

Spores 12-13 u,m, exine yellow brown, areolate

(with a low, delicate, close network of furcate

ridges that coalesce and delimit areolae). Elaters

rigid, nontortuous, 9.6-12 |j.m wide, only slightly

tapering toward tips, bispiral to tips, the spirals

2.4-3.8 u,m wide.

DIFFERENTIATION AND VARIATION — Judging

from identifications of herbarium specimens, T.

praenitens has been widely misunderstood. It is,

however, one of the easiest of our species to rec-

ognize, marked by the rather deeply palmately-

lobed leaves (to 0.6), and the minute denticula-

tions formed by the swollen projecting septa on

the lobes and disc margins (Fig. 10: 5, 6). In ad-

dition, the cuticle of the lobes is almost always

distinctly papillose (Fig. 10: 5, 6). Telaranea ver-

ruculosa (Fig. 60, p. 197), a recently discovered

species from Queensland, also has a distinctly pa-

pillose cuticle, although in that species the disc is

also papillose.

A noteworthy feature of this species is the sca-

brous condition of the perianth surface. The cells

immediately below the mouth are prorate, their

apical ends projecting as blunt, thick-walled teeth

(Fig. 10: 9). Other species with similarly rough-

ened perianths are T. mamillosa (p. 166), and T.

jowettiana (p. 161). Telaranea chaetocarpa

(Pears.) Grolle of New Caledonia may represent

an extreme of this condition. There, the surface

of the perianth bears numerous simple or bifurcate

capillary bristles which are similar in appearance

to the leaf lobes (Pearson, 1922, p. 27; pi. 2: 45-

47; Fig. 41: 1, 2).

Among the specimens assigned to this species

are some which have broader-based leaf lobes and

discrete teeth on the disc margins of both leaves

and half-leaves, a rather startling anomaly in Tel-

aranea, a genus thought to have perfectly entire

leaves. These are recognized here as a variety, var.

dentifolia.

Key to Varieties of T. praenitens

1. Margins of leaves and half-leaves devoid of

teeth; leaf lobes 4 cells wide at base, the cells

in basal portion of lobe elongate (ca. 2:1).

Throughout New Zealand . . . var. praenitens

1 . Margins of leaves and half-leaves (and rarely

the lobes) sporadically with 1(2) multicellular

teeth; leaf lobes (at least the median) often 5-

6 cells wide at base, the cells of lobe bases

somewhat shorter, at times ± isodiametric. Lo-

cal, southern half of South Island

var. dentifolia

Telaranea praenitens (Lehm. & Lindenb.) Hodgs.

var. praenitens

Lepidozia beckettiana Steph., Spec. Hep. 3: 593.

1909. Lectotype (nov.): New Zealand, South Is.,

Westland, Kellys Range, 30 Jan. 1903, Beckett

4310— c. per. (G!).

Leaf and half-leaf margins devoid of teeth; leaf

lobes 4 cells wide at base, the cells in basal por-

tion of lobe elongate (ca. 2:1); margins of disc

and lobe typically minutely denticulate by pro-

truding end walls of the marginal cells.

DISTRIBUTION AND ECOLOGY — Endemic to New

Zealand, found on Stewart Island and rather wide-

spread on South and North Islands. The species

was reported in the earlier literature from Tas-

mania (Bastow, 1888; Hooker, 1867; Mitten,

1859; Rodway, 1916; Stephani, 1909) and West-

ern Australia (Lehmann, 1844-47); Ratkowsky

(1987) also listed the species for Tasmania. We

have seen no collections of the species from Tas-

mania or Australia.

A species of wet, rich lower- to upper-elevation

forests, where often on rotted logs, particularly

when largely bryophyte-covered. It occurs, for ex-

ample, in rich, mixed Nothofagus forests with a

tree fern understory (Tutoko River, Fiordland); in

matai/totara flood-plain forests (Waiho River area

in Westland); in Nothofagus menziesii-Dacrydium

cupressinum-Podocarpus forests along the Four

Mile River (Nelson Prov.); and in N. menziesii-

Pseudopanax simplex-Podocarpus totora-Draco-

phyllum forests (Panekiri Range, Urewera Natl.

Park). It only sporadically inhabits soil or humus

of the forest floor, or moist banks, etc.

The species rarely is subalpine. It was recently

found at 1520 m on Mt. Arthur in communities

of subalpine cushion vegetation, rocky herb fields,

and Dracophyllum, where it occurred over decay-

ing blades at the base of a grass immediately ad-

jacent to a small, stagnant pool.

SELECTED SPECIMENS SEEN— NEW ZEALAND.

STEWART ISLAND: Port Adventure, Schuster, Scott &

Taylor (CHR). SOUTH ISLAND. SOUTHLAND

PROV: Dusky Sound, Supper Cover, Zotov as T. paten-

tissima (CHR); Doubtful Sound, Simpson (CHR); Fiord-

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

35

FIG. 10. Telaranea praenitens (Lehm. & Lindenb.) Fulf. (Figs. 1-10, var. praenitens; 11-12, var. dentifolia.) 1.

Sector of main shoot showing androecia on primary and secondary terminal branches, dorsal view. 2, 3. Leaves (note

symmetry). 4. Leaf, cellular detail. 5, 6. Dorsal lobes of leaves. 7. Underleaves. 8. Underleaf, cellular detail. 9. Sector

of perianth surface immediately below mouth. 10. Antheridial stalk. 11. Leaf. 12. Half-leaf. (Figs. 1-2, 4, 6-8, 10

from Engel 18935, New Zealand, South Is.. Westland Prov., near Waiho River between Wombat Terrace and Canavans

Knob; 3, 5 from Engel 6667B, New Zealand, South Is., Westland Prov., Lake Wombat; 9 from Reif C63B, New

Zealand, South Is., Westland Prov., Camp Creek; 11,12 from type of van dentifolia.)

36

FIELDIANA: BOTANY

land Natl. Park, Stillwater River, Zotov (CHR); ibid.,

Cleddau Valley, The Chasm, Burrell, Scott & Tavlor

(CHR); ibid., Tutoko River. W of Milford Sound, 50 m,

Engel 18841 (F). OTAGO PROV.: Fiordland Natl. Park,

near head of Lake McKerrow, Hatcher 727 (F); N of

McKerrow River, Martin's Bay, Hatcher 735b (F).

WESTLAND PROV.: Cascade Road, just W of Jackson

River, ca. 8-12 km SW of confluence of Jackson and

Arawata Rivers, 25-90 m, Engel 22992— c. 6 (F); Lake

Ellery, ca. 30 m. Child H4569 (F); Haast Pass Road,

Robinson Creek, between Haast Pass and gates of Haast

bridge, ca. 305 m. Child H1852 (F); Haast, 5 miles N

of bridge, ca. 30 m. Child H521 (F); Westland Natl.

Park, Fox Glacier, S side of Fox River, Engel 6618A (F);

Weheka, near Fox Glacier Hotel, Whitehouse 29704 (F);

Westland Natl. Park, Franz Josef Glacier Valley, Roberts

Point, SW of Mt. Gunn, ca. 620-670 m, Engel 18087

(F); ibid.. Lake Wombat, 250 m, Engel 6667B (F); near

Waiho River between Lake Wombat Terrace and Cana-

vans Knob, NW of town of Franz Josef Glacier, off Hwy

6, 110 m, Engel 18935—c. S (F); Arthur's Pass Natl.

Park, N of Kellys Creek near Hwy 73, N of Otira, 420-

475 m, Engel 18340 — c. sporo. (F); Otira Gorge, Berg-

gren 3012 (S); Lake Kaniere Scenic Reserve, Lake Kan-

iere Rd, 125 m, Engel 24857— -c. per. (F); along Route

73, 8 miles W of Turiwhate, Engel 6746 (F); Camp

Creek, W of Alexander Range, 190-840 m, Reif C9C,

CllG—c. sporo., C170C (F); 2 km N of White Horse

Creek, ca. 305 m, Child H5429 (F). CANTERBURY

PROV: Arthur's Pass Natl. Park, near Bealey Glacier

Vista, Engel 6843 (F). NELSON/WESTLAND PROV.

BOUNDARY: Paparoa Range, S side of Porarari River,

up river from gorge and ca. 500 m WSW of ford on

inland track to Bullock Creek, 10-20 m, Engel 19172

(F). NELSON PROV: Paparoa Natl. Park. Bullock

Creek Road, along Bullock Creek, NE of Punakaiki. ca.

25 m, Engel 21616 (F); Four Mile River (Tiropahi)

Track, 215 m, Fife 4674 (F); track to German Terrace,

6 km SSE of Westport on Nine Mile Road, 90 m, Engel

21554 (F); Nelson Lakes Natl. Park, off Lakehead Track,

near juncture with southern end of Loop Track, NE end

of Lake Rotoiti, 630 m, Engel 22729 (F); Kahurangi

Natl. Park, Mt Arthur, track to summit of Mt. Arthur,

1520 m, Engel 24943 (F). NORTH ISLAND. WEL-

LINGTON PROV: Oroua Valley, Western Ruahine

State Forest, ca. 670 m, Hodgson s.n. (CHR); Mt. Ara-

waru, Zotov (CHR); Tararua Mts., Orongorongo River,

ca. 610 m, Zotov (CHR); ibid., Mangahao Downs,

Hodgson 10645 (CHR); ibid., N of Field Hut, ca. 795

m, Zotov (CHR); Ruahine Mts., above Upper Pohangina

Valley, Allison H7590 (CHR); near E border of Tonga-

riro Natl. Park along road to Tree Trunk Gorge, ca. 0.5

km W from gorge, 750 m, Engel 21213 (F). GISBORNE

PROV: Urewera Natl. Park, Panekiri Range, summit

area of Pukenui in vicinity of Punekiri Bluff, S of Lake

Waikaremoana, 1 180 m, Engel 23341 (F); Urewera Natl.

Park, Huiarau Range, summit area of Te Rangaakapua,

1230-1320 m, Engel 23476 (F). TARANAKI PROV:

Below North Egmont Mt. Hut. Hodgson J0243(CHR):

Mt. Egmont, 915 m, Hodgson (S); Pukeiti Bush, near

New Plymouth, Hatcher 336 (F). SOUTH AUCKLAND

PROV: Pukerimu Bush, E of Taupo, ca. 760 m, Allison

H5908 (CHR); Paeroa Range, S of Rotorua, ca. 915 m,

Allison H3035 (CHR); Ohau-iti River, Zotov (CHR).

Telaranea praenitens var. dentifolia Engel &

Merr.

Telaranea praenitens var. dentifolia Engel & Merr..

Phytologia 79: 253. June, 1996 [1995]. Holotype:

New Zealand, South Island, Fiordland, Dusky

Sound, Supper Cove, 1 1 Feb. 1946. Allan, as Lep-

idozia gottscheana (CHR); isotype: (F).

Leaves, half-leaves, and rarely lobes with disc

margins sporadically armed with 1(2) multicellu-

lar teeth; leaf lobes (at least the median) often 5-

6 cells wide at base; cells of lobe bases at times

± isodiametric; end walls of marginal cells of disc

and lobes often indistinctly or not swollen and

protruding.

DIFFERENTIATION AND VARIATION — This variety

and the typical expression of the species both

have rather deeply divided, palmately lobed

leaves, which distinguish T. praenitens from other

members of the sect. Neolepidozia, although the

septa between cells are not so distinctly swollen

and protruding as in var. praenitens. The combi-

nation of rather broad leaf lobes, marginal teeth

(Fig. 10: 11), and papillose cuticle is reminiscent

of Temnoma spp.; the well-developed stem hy-

aloderm, however, marks this plant as a Telaran-

ea.

DISTRIBUTION AND ECOLOGY — Known only from

a limited number of collections, all from the wet,

southwestern sector of South Island. The type oc-

curred intimately intermixed with Riccardia sp.,

and the Simpson collection is mixed with Sema-

tophyllum amoenum. The variety appears to be a

lower-elevation forest plant.

SELECTED SPECIMENS SEEN— NEW ZEALAND.

SOUTH ISLAND. SOUTHLAND PROV: Fiordland.

Doubtful Sound, Hall Arm, Simpson 1328 as T. gottsch-

eana (CHR). WESTLAND PROV: Westland Natl. Park,

250 m, Engel 6662 (F); near Lake Parina. W coast, cf.

15 m. Child HI 798 (F).

Telaranea gibbsiana (Steph.) Hodgs.

Lepidozia gibbsiana Steph., Spec. Hep. 6: 328. 1922.

Telaranea gihhsiana (Steph.) Hodgs., Trans. Roy.

Soc. New Zealand, Bot. 3: 70. 1965. Type: New

Zealand, North Is., without specific loc., Gibbs

1041 (G!).

Plants subisophyllous, soft, with a hairy ap-

pearance, flexuous yet firm, prostrate to ascend-

ing, in dense, compact mats, pale green to (deep

olive green in herb.), highly nitid when dry; plants

medium, to 0.8 cm wide, including branches.

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

37

Branching regularly rather densely and closely

pinnate, occasionally locally 2- or 3-pinnate, near-

ly exclusively of the Frullania type, the branches

short to moderately long but normally determi-

nate, sometimes flagelliform; branch half-leaf 2-

4-lobed, usually obliquely inserted, cuneate (the

lobes diverging, even when bifid); first branch un-

derleaf undivided and ciliiform or 2(3)-fid, insert-

ed on ventral or less often ventral-lateral side of

branch near or at juncture of branch and main

axis. Ventral-intercalary branches occasional, of-

ten becoming leading shoots and leafy through-

out. Stems with cortical cells distinctly differen-

tiated, thin-walled, in 13-15 rows, those on ven-

tral side of stem a little smaller; cortical cells in

section much larger than the numerous (34-47)

medullary cells, the medullary cell walls uniform-

ly slightly thickened. Rhizoids observed only on

stoloniform axes. Leaves on main shoot rigid,

suberect to widely spreading, loosely to closely

imbricate, concave reflecting the incurving of

lobes, the insertion moderately to distinctly incu-

bous; leaves 720-1275 |im wide X 630-890 u,m

long, subsymmetric to less often asymmetric, 4-

6(7) lobed to (0.6)0.65-0.75, the lobes widely di-

vergent, longer than the disc. Lobes ciliform, rig-

id, the base subtriangular, (3)4(5) cells wide at

extreme base, terminating in a uniseriate row of

5-7(8) cells; cells of the uniseriate portion ±

thick-walled, the septa thickened in the corners

and swollen, the basal cell of the uniseriate por-

tion 36-46 u,m wide X 84-108 jim long

([1.6]2.4-4.1:1), the next cell narrower but of

about the same length, 29-36 (im wide X 84-118

u,m long ([2.3]3-4.4:l), the terminal cell normally

about the size of the penultimate cell, or a little

smaller. Disc ± symmetrically cuneate, 4-6 cells

high (from median sinus base to leaf base), 15-

22 cells wide in distal portion narrowing to 8-15

cells wide in basal portion; margins entire, ±

straight to less often slightly curved. Cells of disc

thin-walled but not delicate, trigones minute to

small, the median cells large, 36-65 jxm wide,

50-72(78) u,m long; basal 1-2 rows of disc cells

considerably larger (wider and on the whole a lit-

tle longer), often not in regular tiers; cuticle with

network of fine irregular, elongate striae. Under-

leaves somewhat smaller than leaves, strongly

spreading to subsquarrose, contiguous to loosely

imbricate, plane, 4-6(8)-lobed to 0.6-0.8, the

lobes ± symmetrically divergent, ciliiform,

straight to arched, the uniseriate portion formed

of 5-7(8) elongated, ± thick-walled cells with

swollen septa, not terminating in a slime papilla;

disc symmetrically cuneate, 3-4(5) cells high

(median sinus), the cells often not in regular tiers,

the disc 9-17 cells wide in distal portion narrow-

ing to (8)10-12 cells wide at base; margins entire,

usually straight. Asexual reproduction lacking.

Plants dioecious. Androecia on short, abbrevi-

ated, ventral-intercalary, spicate branches from

main shoot or primary, Frullania-lype branches;

bracts closely imbricate, strongly dorsally assur-

gent, ± cucullate, bilobed to ca. 0.3, the lobes

acuminate, terminating in a uniseriate row of 2-

3(4) cells, the basal cell isodiametric to rather

elongated, the terminal cell ± thick-walled, often

curved, rather elongated, to 3.5:1; dorsal margin

of lamina somewhat dilated and incurved, entire,

bordered by 1-2 rows of elongated, very thin-

walled cells, no slime papillae; bracts monan-

drous; antheridial stalk rather long, ca. 9 cells

high, uniseriate; bracteolar antheridia absent.

Gynoecia oriented laterally and at best only weak-

ly dorsally assurgent, with a vestigial stem peri-

gynium present, swollen, rhizoidous; bracts be-

coming progressively larger and less deeply lo-

bate towards the perianth, those of innermost se-

ries ensheathing the perianth, deeply concave, the

apical portion canaliculate, the bracts ± suborbic-

ular, with apices irregularly 4-lobulate, the lobules

terminating in a uniseriate row of 1-2 cells; bract

margins crenate to dentate, the armature frequent-

ly sharply inflexed, often terminating in a slime

papilla, the bracts with an obscure border formed

of 1 to several rows of cells longer, narrower and

more irregular than those within; bracteoles of in-

nermost series nearly identical in form to bracts

although a little smaller. Perianth not extending

above vegetative axes, 0.5-0.55 emergent, ovoid-

cylindrical, terete in basal half, the distal half ob-

scurely trigonous and with 6-9 plicae, the sulci

shallow to deep; perianth narrowed toward a de-

cidedly contracted mouth, the mouth with 6 nar-

rowly triangular lobes, each lobe fringed with

slightly thick-walled, contorted, crowded, spar-

ingly papillose cilia, the terminal cell of each cil-

ium coarsely papillose.

Seta seen only in collapsed state. Capsule wall

32-40 u,m thick, of 3-4 layers, the outer layer

subequal to 2 of inner layers; outer layer of cells

weakly tiered, rather regularly short-rectangular,

with 2-phase development, the longitudinal walls

with well-defined sheetlike thickenings and nod-

ule-like thickenings (4-6 per cell) alternating with

walls that are devoid of thickenings (or with oc-

casional local, not or weakly pigmented, nodular

swellings), the transverse walls devoid of thick-

38

FIELDIANA: BOTANY

enings or with 1-2 nodular swellings; innermost

layer of cells in weakly defined tiers, irregularly

narrowly rectangular; semiannular bands com-

mon, rather wide, close, usually complete, at

times forked and anastomosing to delimit ill-de-

fined, local fenestrae.

Spores and elaters not seen.

DIFFERENTIATION AND VARIATION — Telaranea

gibbsiana is one of our largest Telaranea species,

and is most closely allied to T. grossiseta of Tas-

mania. Both are relatively robust plants (for the

genus), with a high disc, ciliiform leaf lobes, and

the cells of the uniseriate row distinctly elongated

and capillary. The septa are thickened in the cor-

ners and swollen and projecting (Fig. 11: 7). Sim-

ilar projecting septa occur in T. praenitens. The

most obvious differences between T. gibbsiana

and T. grossiseta are the lower disc (4-6 cells

high vs. 7-9 cells in T. grossiseta), and the char-

acteristic asymmetry of the disc and decidedly

water-repellent cuticle in T. grossiseta.

DISTRIBUTION AND ECOLOGY — The protologue

and the type at G give no specific locality within

New Zealand, other than North Island. Gibbs

(1911), however, stated that the type was collected

at Te Aroha, Thames District, south of Auckland,

2500 ft. on stones and rotten wood, in forest in

November, 1907.

The species appears to be locally and sporadi-

cally distributed on North Island, New Zealand.

Known from several sites in the Lake Waikare-

moana-Urewera area of North Island; there it oc-

curs on ground, exposed roots, rotted stumps, and

bryophyte-covered logs, etc., in forests between

540 and 1320 m (where recorded). It is also lo-

cally abundant in the Waipoua Kauri forests,

where it occurs associated with Hymenophyllum,

Dicranoloma and Trichocolea over rotten logs.

The plant is known from a single station on South

Island, occurring abundantly on humus in a Po-

docarpus-Nothofagus forest (280-350 m, Nelson

Prov.).

SELECTED SPECIMENS SEEN— NEW ZEALAND.

SOUTH ISLAND. NELSON PROV.: Maimai State For-

est, 3 m NW of Reefton, 280-350 m, Macmillan 76/12,

76/154 both as T. gottscheana (CHR). NORTH IS-

LAND. GISBORNE PROV.: Lake Waikaremoana, 610

m, Hodgson s.n. (F); ibid., Ngamoko Track, Hodgson

402 (CHR); ibid.. Hatcher 1301— c. <J, 1310 (F); near

Lake Waikaremoana, trail to Lake Waikare-iti. White-

house 29196 (F); Urewera Natl. Park, Waikareiti Track

between track entrance and Lake Ruapani, N of eastern

extremity of Lake Waikaremoana, 650-920 m, Engel

20513 — c. sporo. (F); ibid., Maungapohatu, ca. 1 100 m.

Moore HI 3 1 (CHR); ibid., Huiarau Range, summit area

of Te Rangaakapua, 1230-1320 m, Engel 23503 (F).

SOUTH AUCKLAND PROV.: Colomandel Peninsula,

Waihi, "ex herb. D. Petrie No. 73"— c. <J (CHR).

NORTH AUCKLAND PROV.: Track to Mahuta, Wai-

poua State Forest, ca. 305 m. Child H 2137 — c. <J,

H2153 (F); SE corner of Waipoua Forest, just N of Tu-

tamoe, 540 m, Engel 21 136 (F); Waima Forest, Hauturu

Highpoint Track in vicinity .of summit area of Hauturu

Trig, off Waiotemarama Track, 650-680 m. Engel 22609

(F).

Telaranea grossiseta (Steph.) Engel & Schust.

Lepidozia grossiseta Steph., Spec. Hep. 3: 584. 1909.

Neolepidozia grossiseta (Steph.) Fulf. & J. Tayl.,

Brittonia 11: 85. 1959. Telaranea grossiseta

(Steph.) Engel & Schust., Fieldiana, Bot. n.s. 14:

3. 1983. Type: Tasmania, West Coast, near Moore's

Lookout, Mar. 1900, T. B. Moore 5794 (G!).

Plants subisophyllous, with a rather hairy ap-

pearance, flexuous yet firm, prostrate in loose

mats, glaucous, whitish green to ceraceous, the

surface dull, waxy and water repellent, the old

shoots brownish and subnitid; plants large, the

main shoot to 2.2 mm wide, to 1.1 cm wide, in-

cluding branches. Branching loosely to somewhat

irregularly 1 -(sporadically 2) pinnate, mostly of

the Frullania type, frequently becoming flagelli-

form; branch half-leaf bifid, transversely to incu-

bously inserted, the lobes often divergent; first

branch underleaf undivided, ciliate to subulate,

(rarely bilobed), inserted on ventral-lateral side of

stem at or somewhat below the juncture of branch

and stem. Acromastigum-[ype branches present

but rare. Ventral-intercalary branches common,

either leafy throughout or stoloniform, the stolons

normally present toward shoot base. Stems with

cortical cells distinctly differentiated, in 12-17

rows, much larger than the numerous (ca. 40-90

rows) medullary cells, both the cortical and med-

ullary cells thin or evenly and slightly thick

walled. Rhizoids from distal cells of underleaf

disc. Leaves on main shoot rigid, widely spread-

ing, nearly but normally not at right angles to the

stem, contiguous to loosely imbricate, plane or es-

sentially so, distinctly incubously inserted and ori-

ented, 1135-1650 |im wide X 1160-1650 u.m

long, moderately asymmetric, 4(6}-lobed to 0.5-

0.65, the lobes somewhat divergent, often asym-

metrically so, the dorsal lobe typically stiffly dor-

sally assurgent and out of plane of leaf, often ori-

ented toward shoot apex, the other lobes ± par-

allel with the disc margins, the lobes about as long

as the disc. Lobes ciliiform, rigid, consisting of a

short subtriangular base 4-6 cells wide at extreme

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

39

FIG. 11. Telaranea gibbsiana (Steph.) Hodgs. 1. Shoot with 6 Frullania-type branches, dorsal view (note 2-, 3-

and 4-lobed half-leaves). 2. Shoot with 3 young gynoecia toward base, dorsal view. 3-6. Leaves. 7. Distal portion

of leaf lobe. 8. First branch underleaves. 9. Stem, cross-section. 10, 11. Capsule wall cross sections. (Figs. 1-9 from

Hodgson s.n.. New Zealand, North Is., Lake Waikaremoana [F]; 10, 1 1 from Engel 20513, New Zealand, North Is.,

Gisborne Prov., Urewera Natl. Park, Waikareiti Track.)

40

FIELDIANA: BOTANY

FIG. 12. Telaranea gibbsiana (Steph.) Hodgs. 1. Gynoecium. 2. Cross section through distal portion of perianth.

3. Portion of perianth mouth. 4. Distal portion of innermost bract. 5. Underleaves of main shoot. 6. Underleaf. 7.

Half-leaves. 8. Four-lobed half-leaf. 9. Antheridium. (Figs. 1-8 from Hodgson s.n.. New Zealand, North Is., Lake

Waikaremoana [F]; 9 from Hatcher 1301, same loc.)

base and abruptly contracted to a bristle-like uni-

seriate portion 6-7(8) cells long; dorsal lobe often

widely divergent, 1-2-seriate at base (if uniseriate

to base, then "embedded" in disc), often appear-

ing displaced midway onto the dorsal margin of

disc; cells of the uniseriate row thick-walled (in

comparison with disc cells), elongated, subcapil-

lary, the septa thickened at the corners and typi-

cally swollen, the basal cell of the uniseriate por-

tion 36-48 |xm wide X 137-168 u,m long, the

next cell narrower and longer, 26-34 u,m wide X

149-180 jjim long, the longest cells of the uni-

seriate row up to 7.5:1, the terminal cell very

short, tapering to a sharp point. Disc asymmetri-

cally cuneate to short rectangular, the dorsal mar-

gin (insertion to dorsal lobe base) distinctly short-

er than the ventral, the disc 7-9HO) cells high

(from median sinus base to leaf base), 14-19 cells

wide in distal portion narrowing to 8-12 cells

wide in basal portion; margins entire, ± straight

to slightly curved, sometimes asymmetrically so.

Cells of disc thin-walled but not delicate, trigones

minute or lacking, median disc cells large, 46-60

Urn wide, 77-94 u,m long; cells of disc margins

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

41

FIG. 13. Telaranea grossiseta (Steph.) Engel & Schust. 1. Plant, dorsal view. 2. Portion of main shoot with

Acromastigum-lype branch (= AB). ventral view; note 6-fid underleaf (rare), (HUL = half-underleaf; BL = first

branch leaf; BUL = branch underleaf"). 3. Stem, cross section. 4. Uniseriate portion of leaf lobe. (All from Engel

14728, Tasmania. Gordon River.)

42

FIELDIANA: BOTANY

FIG. 14. Telaranea grossiseta (Steph.) Engel & Schust. 1. Portion of main shoot with Frullania-iype branch ( =

FB), dorsal view (HL = half-leaf; BUL = first branch underleaf). 2. First branch underleaves. 3. Underleaf. 4-5.

Leaves. 6. Dorsal lobe of leaf. (All from Engel 14728, Tasmania, Gordon River.)

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

43

200 n

FIG. 15. Telaranea hodgsoniae Engel & Merr. 1. Leaves and to left, an underleaf. 2. Three leaves, cellular detail.

3. Median leaf lobes. 4. Median disc cells. 5. Underleaves, cellular detail, stippled walls indicate formation of rhizoid

initials. (All from holotype.)

often narrowly elongate, forming an indistinct

border; disc cells in ± regular tiers to irregularly

arranged; cuticle with a hazy to faintly granular

appearance caused by the continuous waxy, water-

repellent coating, eventually developing a net-

work of fine, irregular lines. Oil-bodies (fide Rat-

kowsky 81/14) ca. 4-6 per cell, botryoidal, small,

located around cell periphery, the chloroplasts

likewise located. Underleaves somewhat smaller

than the leaves, strongly spreading, distant to

weakly imbricate, plane, 4-6(8)-lobed to

(0.6)0.65-0.7; lobes symmetrically divergent, cil-

iiform, rigid to tortuous, the uniseriate portion

formed of 5-9 elongate, ± thick-walled cells, the

distal cells often secondarily divided, the lobes

commonly terminating in a slime papilla; disc

symmetrically cuneate, 3-4 cells high (from me-

dian sinus base to underleaf base), the cells in

regular or irregular tiers; disc 13-23 cells wide in

distal portion narrowing to 8-13 cells wide at the

base; margins entire, usually straight, sometimes

gently curved. Asexual reproduction lacking.

Androecia and gynoecia not seen.

DIFFERENTIATION AND VARIATION — Telaranea

grossiseta is a handsome plant, immediately dis-

tinguishable by the regularly incubously shingled,

dull, glaucous leaves, with divergent, bristle-like

lobes. This species has no close relatives in Tas-

mania, but is closely allied to T. gibbsiana of New

Zealand. They both have a distinctive "hairy" as-

pect, and leaf lobes of similar form and orienta-

tion. The New Zealand plant, however, is not

glaucous, but highly nitid, at least when dry. For

44

FIELDIANA: BOTANY

additional comparisons between these species, see

under T. gibbsiana.

As in other glaucous members of the Lepido-

ziaceae (e.g., Bazzania tayloriana, Lepidozia

glaucophylla, Telaranea tuberifera) the leaves

have a waxy, water-repellent coating, although the

surface is not as evidently coarsely granular as in

some of the foregoing species.

DISTRIBUTION AND ECOLOGY — Limited to Tas-

mania and New South Wales, Australia; in the for-

mer confined to the wet, western sector of the

state. The species typically occurs over soil in

cool, shaded niches of lower to middle-elevation

forests. The species is rare and sporadic in distri-

bution, and nowhere, in our experience, common,

with one exception. It is locally common on the

sides of yabby (crayfish) holes in the Trowutta

Caves area (Engel 19760). At Kelly Basin Road

it forms large, cushion-like, pure patches on the

shaded, vertical rock walls of an abandoned rail-

road line (Engel 14969).

SELECTED SPECIMENS SEEN— TASMANIA: Cradle

Mtn.-Lake St. Clair Natl. Park, W shores of Lake St.

Clair about 1 mile N of Echo Hut, ca. 750 m, Norris

28050 (F); Franklin River at Frenchman's Cap Trail

crossing, ca. 400 m, Norris 31194 (F); Gordon River,

vicinity of Sir John Falls, just up river from Butler Is.,

ca. 50 m, Engel 14728 (F); Kelly Basin Road, between

junction with Franklin River Road and Kelly Basin,

above Nora River, 90 m, Engel 14969 (F); King River.

11.3 km by road from Regatta Point and 13.3 km by

road from Strahan Harbour, sea level, Engel 14927 (F):

Mt. Lyell, Ratkowsky 81/14 (HO); near Zeehan, up

Dunkley's tram line, 185 m, Weymouth 1445 (F); road

from Melba Flats to Confidence Saddle, 200-400 m.

Norris 31643 (F); Murchison River at Murchison Hwy,

Norris 33735 (F); vicinity of Trowutta Caves (Arch), 4.5

km by road from intersection of Reynolds Road and

Reids Road and 6.9 km from Trowutta Post Office, ca.

~ 180 m, Engel 19760 (F); Dismal Swamp, between Brit-

tons Swamp and W coast along Bass Hwy. (2), 34.4 km

by road SW of Smithton, 30 m, Engel 16176 (F). AUS-

TRALIA. NEW SOUTH WALES: Macquarie River,

Jan. 1836, Ball (mixed with Jungermannia procumhens,

nom. hb. [Balantiopsis]) (FH).

Telaranea hodgsoniae Engel & Merr.

Telaranea hodgsoniae Engel & Merr.. Phytologia 79:

251. June, 1996 [1995]. Holotype: New Zealand,

South Is., Canterbury Prov.. Peel Forest, ca. 1500

ft., Child H2084—C. cJ (F); isotype: (CHR).

Plants rather stiff and firm, prostrate, in com-

pact mats, glaucescent, whitish green, the surface

rather dull and water repellent, subnitid when

dry; plants ± stenotypic in stature, small, to 0.6

cm wide, including branches. Branching irregu-

larly to rather regularly once-pinnate, the branch-

es of the Frullania type, short, hardly tapering,

sporadically to rather commonly flagelliform for

much of their length; branch half-leaf 2-lobed,

strongly obliquely inserted, linear-rectangular,

the lobes parallel to weakly diverging; first

branch underleaf undivided, subulate, inserted on

ventral side of branch at juncture of branch and

main axis. Ventral-intercalary branches sporadic,

leafy, becoming leading shoots. Stems with cor-

tical cells distinctly differentiated, thin-walled, in

13 rows; cortical cells in section much larger

than the numerous (ca. 52) medullary cells, the

medullary cell walls slightly thickened and mi-

nutely pitted. Leaves on main shoot rigid,

obliquely spreading, rather closely imbricate,

moderately convex (especially when dry), the in-

sertion distinctly incubous; leaves 315-350(420)

u,m wide X 350-525 u,m long, subsymmetric,

(3)4-lobed to 0.4-0.5. the lobes straight, much

shorter than the disc. Lobes narrowly acute, 2-

3(4) cells wide at extreme base, terminating in a

short uniseriate row of 2-3 cells; lobe cells short.

± isodiametric to short rectangular, bulging,

moderately thick-walled, the cell walls weakly

thickened in the corners but the septae not swol-

len and projecting, the basal cell of the uniseriate

portion 32-42 u,m wide X 34-48 |xm long

(1.1 [1.3]:!), the next cell 22-32 jxm wide X 22-

40 u,m long (0.9-1.3:1), the terminal cell nor-

mally about equal to the penultimate cell in

length or a little shorter, secondary divisions oc-

casional, the apex narrowly rounded; cuticle as

in disc. Disc symmetric, subquadrate, 4-5 cells

high (from median sinus base to leaf base), 8(9)

cells wide in both distal and basal portion; mar-

gins entire, ± parallel to less weakly curved;

cells of margins often bulging. Cells of disc

evenly and moderately thick-walled and firm, tri-

gones minute, the cells ± isodiametric, those in

median sector 38-50(55) p-m wide X 43-62(70)

u,m long, the cells in ± regular longitudinal

rows, but somewhat offset transversely; cuticle

with a hazy, granular appearance. Underleaves

much smaller than leaves, ca. 1.3X stem width,

patent to strongly spreading, distant, plane, 4-

lobed to 0.5-0.6, the lobes straight, consisting of

a uniseriate row of 3(4) short cells (the lobes 2

cells wide at extreme base) terminating in a

slime papilla; disc subquadrate to weakly cune-

ate, 3 cells high (median sinus), the cells ± reg-

ularly arranged, the disc 8 cells wide, the cells

in distal tier sporadically secondarily divided to

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

45

form rhizoid initials; margins entire, usually

straight.

Asexual reproduction lacking.

Plants apparently dioecious. Androecia on

short, abbreviated, ventral-intercalary branches

from leading shoots or from flagelliform branch-

es; bracts closely imbricate, dorsally assurgent,

deeply concave, 2-lobed, each lobe terminating in

a single cell or a uniseriate row of 2-3 not to

hardly elongated cells; lamina cells irregular in

shape and arrangement, the dorsal lamina margin

feebly dilated, feebly and irregularly crenulate, no

slime papillae seen; bracts monandrous; anther-

idial stalk uniseriate; bracteolar antheridia absent.

Gynoecia not seen.

DIFFERENTIATION AND VARIATION — Plants of T.

hodgsoniae often appear somewhat dull under the

dissecting microscope but are never glaucous as

in T. tuberifera and T. centipes. Under the com-

pound microscope the cells have a hazy, granular

appearance, rather like frosted glass.

The plant resembles a diminutive T. tetrapila.

The parallel-sided leaf disc (Fig. 15: 1, 2) is rem-

iniscent of the var. roseana of that species, being

only 8 cells wide throughout, although only 4-5

cells high (Fig. 15: 2). However, the cells appear

small and firm-walled, and the lobes are rather

stout and stubby, and not attenuate. The plants are

rather uniform in size, small, and not more than

0.6 cm wide, including the branches.

Our concept of this species apparently coin-

cides with T. roseana sensu Hodgson (1956, p.

606), since all of the specimens identified as T.

roseana by Hodgson in the Christchurch Herbar-

ium (CHR) are this taxon. The true T. roseana,

however, is a different plant, treated here as a va-

riety of T. tetrapila (p. 57).

"Telaranea obscura" (Schuster, 2000, p. 213)

was our provisional name for this species, sub-

sequently published as T. hodgsoniae (Engel and

Merrill, 1996).

DISTRIBUTION AND ECOLOGY — Known only from

Auckland Is. and scattered sites on South Island,

and the extreme southern sector of North Island

(Tararua Range). It apparently is a forest species,

where it may cover many square yards on steep,

shaded, silty, damp banks in, for example, ma-

nuka or mahoe bush (Otago Prov.) or Nothofagus

fusca-N. solandri forest (Nelson Prov.).

SELECTED SPECIMENS SEEN — AUCKLAND IS.: Mus-

grave Peninsula. Dawbin as T. roseana (CHR). NEW

ZEALAND. Without specific loc., ex hb. Petrie (Allison

no. H3063 as T. roseana) (CHR). SOUTH ISLAND.

OTAGO PROV.: Near Outram, Taieri Plains, S of Dun-

edin. Allison H3208 as T. roseana (CHR); Lake Waiho-

la, S of Dunedin, ca. 30 m, Allison H3207 as T. roseana

(CHR); Town Belt, Mornington, Dunedin, Allison

H3206 as T. roseana (CHR). NELSON PROV.: Nelson

Lakes Natl. Park, Pinchgut Track, W of southern sector

of Lake Rotoiti, SSW of St. Arnaud, ca. 1280-1390 m,

Engel 21390 (F). NORTH ISLAND. WELLINGTON

PROV: Upper Hut, N of Wellington, Druce 914 as T.

roseana (CHR); Tararua Range, Mt. Holdworth, above

Mountain House, ± 850 m, Butler 2584 as T. roseana

(CHR).

Telaranea patentissima (Hook. f. & Tayl.) Hodgs.

Jungermannia patentissima Hook. f. & Tayl., London

J. Bot. 3: 386. 1844, (3: 286 [sic] in errore pro

386). Lepidozia patentissima (Hook. f. & Tayl.) G.

L. & N., Syn. Hep. 204. 1845. Mastigophora pa-

tentissima (Hook. f. & Tayl.) Trev., Mem. 1st.

Lomb. Sci. Lett. III. 4: 416. 1877. Telaranea pa-

tentissima (Hook. f. & Tayl.) Hodgs., Rec. Domin.

Mus. 4: 107. 1962. Type: Auckland Is., Hooker

(BM! [seen 4], FH!).

Plants soft, often spinescent, flexuous yet firm,

prostrate, in loose to compact mats, pale green to

yellowish green, nitid when dry; plants rather

stenotypic in stature, small, to 0.8 cm wide, in-

cluding branches. Branching irregularly to some-

what regularly 1 -pinnate, never 2-pinnate, the

branches of the Frullania type, short and hardly

tapering to long and flagelliform; branch half-leaf

bifid, usually obliquely inserted, narrowly rect-

angular, the lobes parallel to weakly diverging;

first branch underleaf undivided and ciliiform to

subulate, inserted on ventral side of branch at

juncture of branch and main axis. Ventral-inter-

calary branches occasional to common, leafy, of-

ten becoming leading shoots. Stems with cortical

cells distinctly differentiated, thin-walled, in 1 1-

12 rows, the outer walls somewhat thicker; cor-

tical cells in section larger than the numerous (ca.

25-45) medullary cells, the medullary cell walls

uniformly slightly thickened. Rhizoids from distal

cells of underleaf disc. Leaves on main shoot rig-

id, obliquely spreading, distant (esp. in small

phases) to imbricate, plane to moderately convex,

noticeably narrowing to the base, the insertion

typically distinctly incubous; leaves 335-545 u,m

wide X 345-580 u,m long, subsymmetric to mod-

erately asymmetric, 4(6)-lobed to 0.4-0.5, the

lobes usually somewhat divergent, about equal to

or shorter than the disc. Lobes narrowly acute to

attenuate, 2-4(5) cells wide at extreme base, the

two median lobes typically somewhat larger (and

at times 3-4 cells wide in basal sector), often bi-

46

FIELDIANA: BOTANY

seriate for 1-2(3) tiers, terminating in a uniseriate

row of 2-4(5) cells, typically less than half the

length of lobe (exc. var. zebrina); lobe cells short,

± isodiametric to short rectangular, thin- to mod-

erately thick- walled, the septa somewhat thick-

ened in the corners but not swollen and project-

ing, the basal cell of the uniseriate portion 22-36

u,m wide X 26-46 u,m long (1-1.8:1), the next

cell 17-30 jim wide X 18-34 jxm long, the ter-

minal cell at times secondarily divided, tapering

to a rounded summit; cuticle smooth to finely stri-

ate-papillose (at least the distal half of lobes).

Disc symmetrically to somewhat irregularly cu-

neate, 4-5 cells high from median sinus base to

leaf base (up to 6[7] cells high in var. ampliata),

10-15 cells wide in distal portion (8 cells wide in

smaller leaves) narrowing to (5)8(12) cells wide

in basal portion; margins entire, ± straight to

broadly curved, cells of disc and lobe margins

often bulging. Cells of disc thin- to moderately

and evenly thick-walled and firm, with small but

distinct trigones, the cells in median sector of disc

subquadrate to somewhat elongate, 25-42 u,m

wide X 38-55 u,m long; cuticle smooth (coarsely

striolate-papillose in var. zebrina). Underleaves

somewhat to much smaller than leaves, variable

in size, ca. 1-1.5X stem width, strongly spread-

ing, distant to less often contiguous, plane, (3)4-

lobed to 0.35-0.45, the lobes weakly divergent,

biseriate at base (exc. var. zebrina), with a uni-

seriate row of 2-4 rather short cells, terminating

in a slime papilla; disc symmetrically subquadrate

to weakly cuneate, 3-4 cells high (median sinus),

(8)10-14 cells wide in distal portion narrowing to

8 cells wide at base (8 cells wide throughout in

var. zebrina); margins entire, usually straight to

somewhat curved. Asexual reproduction lacking.

Plants dioecious. Androecia mostly on short,

abbreviated, ventral-intercalary, spicate branches

from main shoot or, occasionally, from primary,

flagelliform Frullania-type branches; bracts close-

ly imbricate, dorsally assurgent, deeply concave,

bilobed to ca. 0.35, the lobes short acuminate, ter-

minating in a uniseriate row of 2(3) cells, the bas-

al cell slightly longer than wide, the terminal cell

to ca. 2:1; dorsal margin of lamina somewhat di-

lated, crenulate, with a few slime papillae; bracts

monandrous; antheridial stalk 8 cells high, uni-

seriate; bracteolar antheridia absent. Gynoecia

feebly to strongly dorsally assurgent, swollen and

densely rhizoidous at base; bracts small for peri-

anth size, those of innermost series closely en-

sheathing the perianth, concave, oblong to sub-

orbicular, very shallowly and irregularly 3-4-lob-

ulate, the lobules terminating in single curved cell

or several sinuous, laterally juxtaposed cells,

sometimes irregularly denticulate; lamina com-

posed of ± regularly subrectangular cells, the

margin bordered by narrower elongated cells, the

apical or free end of marginal cells often diver-

gent and forming a blunt tooth, especially distally,

the margins otherwise with a sporadic unicellular

tooth and a few slime papillae; bracteoles of in-

nermost series smaller but similar in form to

bracts. Perianth long emergent, fusiform, terete in

basal and median sectors, the distal sector ob-

scurely trigonous and with 3-plicae, the perianth

narrowing toward the contracted mouth; mouth

cells thick-walled, often sinuate, partially or whol-

ly laterally free, occasionally with a laterally free

uniseriate row of 2 cells, the mouth thus indis-

tinctly denticulate-subfimbriate; perianth 2- to lo-

cally 3-stratose in basal portion.

Seta with 8 rows of outer cells surrounding an

inner core of 21-24 much smaller cells. Capsule

rather short elliptic, 1070 X 525 u,m, the wall 41-

46 u,m thick, of 4 layers, the outer layer equiva-

lent to 1.5-1.8X thickness of any one of the in-

terior layers; outer layer of cells rather regularly

short-rectangular, with 2-phase development, the

longitudinal walls with well-defined sheetlike

thickenings and nodule-like thickenings (4-6 per

cell) alternating with walls that are devoid of

thickenings (or with sporadic, pigmented or non-

pigmented, nodular swellings), the transverse

walls often with thickenings; intermediate layer

thinner than outer or inner layers; innermost layer

of cells ± tiered, irregularly narrowly to broadly

rectangular, with semiannular bands common,

rather wide, close, usually complete, rarely forked

and anastomosing to delimit ill-defined, local fe-

nestrae.

Spores 12-13.9(15.4) p.m, exine pale brown,

with low but sharply defined, close papillae and

short-vermiculate markings which at times coa-

lesce. Elaters rigid, nontortuous. 8.6-10.6 jim

wide, only slightly tapering toward tips, bispiral

to tips, the spirals 2.4-3.8 fim wide.

NOMENCLATURE — The type of Jungermannia

patentissima, as stated in the prptologue, was

"creeping on other Hepaticae or on Musci,"

namely Bazzania adnexa, Heteroscyphus sinuo-

sus, Lepidozia sp., a hypnoid moss, and a hy-

menophylloid fern. A portion of the type is pre-

sent in Thomas Taylor's herbarium (FH) under the

manuscript name Jungermannia patentispina,

with the same admixture of cryptogamic species.

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

47

Examination of the type of Lepidozia gottsch-

eana discloses that this widely used name is a

synonym of T. patentissima and has been misap-

plied to the plant that is properly called T. tetra-

pila (p. 52). The protologue of Lepidozia gottsch-

eana states only that the plant came from New

Zealand. The type (W, "no. 28") is labelled

"Dusky Bay," and therefore was presumably col-

lected by Menzies. Both perianths and mature

sporophytes are present, as per the original de-

scription. The specimen at Geneva ("Original,"

ex herb. Bescherelle) also bears the notation

"Dusky Bay," and is a probable isotype, although

gynoecia are not present.

DIFFERENTIATION AND VARIATION — Telaranea

patentissima is a relatively common plant in New

Zealand, but its variability may cause difficulties

in determination. At times the species can resem-

ble a smaller T. tetrapila. The leaf disc is ± reg-

ularly cuneate, with the disc cells arranged in reg-

ular rows and tiers (Fig. 16: 4). Another feature

shared with T. tetrapila and its varieties is the

occasional occurrence of secondary cell divisions

at the tips of the lobes (Fig. 16: 3). However, the

smaller plant size, the relatively small cells (40

u,m in width or less), and the firm, rather evenly

thickened cell walls with small but distinct tri-

gones ally this species with sect. Neolepidozia.

Hodgson (1956, p. 609) says, "cells 30-40 u,,

showing as large in such a small plant." Hodgson

provided a description based on the type, although

the plant is considerably more common than her

discussion would indicate. The concept of T. pa-

tentissima in Allison & Child (1975, p. 79) also

appears to be correct.

Lepidozia gracillima Carr. & Pears. (1888) of

New South Wales is a synonym of T. patentissi-

ma, as shown by the type (G!). The disc cells are

23-28 u,m wide, somewhat smaller than in typical

T. patentissima. Telaranea quadristipula, also of

New South Wales, has disc cells 16-24 |x, and has

lobes 4(5) cells wide at the base, often with up to

3 additional 4-celled tiers, often somewhat abrupt-

ly contracted to the uniseriate row (Fig. 5: 1, 5).

Stephani (1909, p. 581) remarks that the illustra-

tion of the leaves of L. gracillima in the original

publication is "missgliickt und irrefiirend." In

particular, the lobes are shown as terminating in

a single cell, rather than uniformly ending in a

uniseriate row of 2-3 cells as in the type, and the

leaf outlines filled in with oddly shaped cells.

Three expressions of T. patentissima are rec-

ognized here at the varietal level:

48

Key to the Varieties of T. patentissima

1. Leaf disc typically 4 cells high, ± regularly

cuneate, the areolation regular, the cell divi-

sions in distal sector mostly longitudinal, pre-

serving a regular tiered arrangement; disc and

lobe margins straight; leaves 4-lobed 2

2. Cells of disc coarsely striolate-papillose;

uniseriate portion of leaf lobes (4)5 cells

long; underleaf disc 8 cells wide through-

out, the lobes uniseriate, 2-3 cells long . .

var. zebrina

2. Cells of disc smooth; uniseriate portion of

leaf lobes 2-4(5) cells long; underleaf disc

(8)10-14 cells wide in distal portion nar-

rowing to 8 cells wide at base

var. patentissima

1 . Leaf disc typically 5-6(7) cells high, becoming

irregularly and often asymmetrically cuneate,

the areolation (esp. in distal sector) ± irregular

due to divisions in various planes; disc and

lobe margins irregular, with bulging cells;

leaves often 5-6-lobed var. ampliata

Telaranea patentissima (Hook. f. & Tayl.) Hodgs.

var. patentissima

Lepidozia gottscheana Lindenb. in G. L. & N., Syn.

Hep. 206. 1845, syn. nov. Mastigophora gottsch-

eana (Lindenb.) Trev., Mem. 1st. Lomb. Sci. Lett.

III. 4: 416. 1877. Neolepidozia gottscheana (Lin-

denb.) Fulf. & J. Tayl., Brittonia 1 1: 85. 1959. Te-

laranea gottscheana (Lindenb.) Hodgs., Rec. Dom-

in. Mus. 4: 106. 1962. Type: New Zealand, Dusky

Bay, "(Hb. Hk., L. et Lg. n. 28)," sin. coll.—c.

per. + sporo. (G!, W!, Lindenberg Hepat. no.

4748).

Lepidozia gracillima Carr. & Pears., Proc. Linn. Soc.

New South Wales sen 2. 2: 1045. pi. XXVII. 1888,

syn. nov. Type: Australia, New South Wales, "New

Sydney," Botany Bay, Whitelegge — c. c? (G!).

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

syn. nov. Neolepidozia corticola (Steph.) Fulf. & J.

Tayl., Brittonia 11: 85. 1959. Telaranea corticola

(Steph.) Hodgs., Rec. Domin. Mus. 4: 107. 1962.

Type: New Zealand, South Is., Mt. Winterslow,

(Canterbury), Mar. 1906, Beckett 457 (G!).

Leaves 4-lobed, the disc ± regularly cuneate,

the disc and lobe margins straight; disc typically

4 cells high, with areolation regular, the cell di-

visions in distal sector mostly longitudinal, pre-

serving a regular tiered arrangement.

Telaranea patentissima var. ampliata Engel &

Merr., var. nov.

Folia saepe 5-6 lobata, lobis anguste acutis (ad basin

FIELDIANA: BOTANY

imm

FIG. 16. Telaranea patentissima (Hook. f. & Tayl.) Hodgs. (Figs. 3-5, 9, var. patentissima; 1, 2, 6-8, var. am-

pliata.) 1. Sector of main shoot, dorsal view. 2-4. Leaves, cellular detail. 5. Underleaf, cellular detail. 6. Stem, cross

section. 7, 8. Dorsal lobe and dorsal margin of disc. 9. Antheridal stalk. (Figs. 1-2, 7 from holotype of var. ampliata;

3 from Kirk 183, New Zealand, Great Barrier Is.; 4-5, 9 from Zotov New Zealand, North Is., Wellington Prov.,

Tararua Mts., Field Hut; 6, 8 from Hatcher 29, New Zealand, North Is., South Auckland Prov., Rotorua.)

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

49

3-4 cellulas latis), disco plerumque 5-6(7) cellulas alto,

usymmetricae irregulariterque cuneato, disco lobisque

margine cellulis tumidis.

HOLOTYPE — New Zealand, Otago Prov., Mt.

Maungatua, W of Mosgiel, 500 m, Engel 17777—

c. sporo. (F); isotype: (CHR).

Leaves often 5-6-lobed, the disc becoming ir-

regularly and often asymmetrically cuneate, the

disc and lobe margins irregular, with bulging

cells; disc typically 5-6(7) cells high, with areo-

lation (esp. in distal sector) ± irregular due to

divisions in various planes.

This distinctive expression of the species has

broad, somewhat irregularly cuneate leaves, with

the cells of the lobe and disc margins often bulg-

ing (Fig. 16: 7, 8). The leaf disc is rather high (to

6 or 7 cells, Fig. 16: 2), the cells are somewhat

irregularly arranged, and the lobes are narrowly

acute (often 3-4 cells wide for several tiers above

the extreme base).

The distal broadening of the cuneate leaf disc

results from additional cell divisions, which are

mostly confined to the distal portion of the disc;

the remainder of the disc has ± regular rows and

tiers of comparatively large, ± isodiametric to

slightly elongated cells.

Specimens referrable to the var. ampliata in-

clude Engel 22818 from Nelson Lakes Natl. Park

and Hatcher 29 from Rotorua, both New Zealand,

and Engel 13783 from near Lake Gordon, and En-

gel 14016 from Lake St. Clair Natl. Park, Tas-

mania.

Telaranea patentissima var. zebrina Engel &

Mem, var. nov.

Var. patentissima simile, differt cellulis disci striatulo-

papillosis, disco ubique 8 cellulas lato, lobis foliorum

parte uniseriata (4)5 cellulas longa, lobis amphigastrior-

um 2-3 cellulas longis uniseriatis.

HOLOTYPE — New Zealand, North Island, Puketi

Forest, 6 Dec. 1993, Braggins 93/172 (AKU); iso-

type: (F).

Leaves distinctly papillose, the dorsal half of

disc coarsely striolate-papillose, the lobes with a

uniseriate row (4)5 cells long (more than half the

length of lobe); underleaf disc 8 cells wide

throughout, the lobes uniseriate, 2-3 cells long.

This variety is similar to var. patentissima in

almost all respects but has a more pronounced de-

velopment of cuticle on the leaves. Papillae are

not confined to the tips of the lobes but occur also

on the disc (Fig. 17: 1), which is conspicuously

striolate (Fig. 17: 9). Among the other Austral-

asian species of the genus, only T. verruculosa of

Queensland (Fig. 60: 6, p. 197) has a striolate

disc. The variety also differs by the underleaves

with uniseriate lobes 2-3 cells long and disc 8

cells wide throughout (Fig. 17: 11), and the some-

what longer leaf lobes, with a uniseriate portion

(4)5 cells long (Fig. 17: 1, 2).

The varieties of T. patentissima are not distin-

guished in the discussions of ecology and distri-

bution and specimens cited.

DISTRIBUTION AND ECOLOGY — Telaranea paten-

tissima occurs on Auckland, Campbell, and Stew-

art Islands, and is rather common throughout the

South Island and North Island, New Zealand. Also

present (but much less common) in Tasmania, as

well as in New South Wales, at present repre-

sented by a single collection, the type of L. gra-

cillima. In New Zealand the species occurs in

middle- to upper-elevation forests (typically

above 400 m) to the subalpine-alpine zone. In the

forest zone (often Nothofagus) it often occurs on

very rotted logs, typically on the side or lower

half of prostrate, bryophyte-covered logs, or in

moist, sheltered niches such as cliff faces and soil

of boulders of stream edges, etc. In the subalpine

and alpine zones it occurs under snow tussock

cover or over soil in protected niches of cliffs, etc.

In the southern extremity of South Island (Milford

Sound) it may be found at sea level with Treubia

lacunosa, Pellipsis, Verdoornia succulenta, and

Geocalyx novaezealandiae, etc. It also, though

rarely, occurs at lower elevations in pakihis (un-

der Gleichenia).

In Tasmania the species has a similar altitudinal

range to that of New Zealand; for example, it oc-

curs in Nothofagus-Eucalyptus-Atherosperma- or

Nothofagus-Athrotaxis forests. In forests the plant

occurs on bryophyte-covered logs and stumps,

and in rock crevices of dryish stream beds; in sub-

alpine scrub it occurs over soil at edges of tarns.

SELECTED SPECIMENS SEEN— AUCKLAND IS.: Horn-

ing SA444 (CHR); Kirk 559 (BM). CAMPBELL IS.:

Poppleton M14 (CHR). NEW ZEALAND. SOLANDER

IS.: East Bay, 10 m, Johnson as T. gottscheana (CHR);

SE Peninsula, 100 m, Johnson as T. gottscheana (CHR);

SE Ridge, 160 m, Johnson as T. gottscheana (CHR).

STEWART ISLAND: Rakiura Natl. Park, directly be-

hind Belltopper Falls, ca. 30 m, Engel 24164; ibid., Port

Pegasus, ca. 200 m, Engel 24178 (F). SOUTH ISLAND.

50

FIELDIANA: BOTANY

FIG. 17. Telaranea patentissima (Hook. f. & Tayl.) Hodgs. (Figs. 1, 2, 9-11, var. zebrina; 3-8, 12, 13, van

patentissima). 1-6. Leaves, the leaf of fig. 1 with cuticle shown in part. 7, 8. Leaf lobes. 9. Marginal cells of leaf

base showing papillae. 10-13. Underleaves (all drawn to scale of fig. 10). (Figs. 1, 2, 9-1 1 from type of var. zebrina;

3, 4, 7, 8, 12 from type of T. patentissima (BM); 5, 6, 13 from type of T. gottscheana [W].)

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

51

SOUTHLAND PROV.: Waikaia State Forest, junction of

VVaikaia River Track and Tison Peak Track, upstream

from Piano Flat Recreational Area, ca. 260 m, Engel

24705 (F); Dusky Bay, 1791 \Menzies] (BM); Lake

Hauroko, ca. 215 m, Child HI 559 (F); Mt. Burns, ca.

1280 m. Child H505I (F); near road between Homer

Tunnel and Te Anau, Hatcher 1528 (F); Cascade Creek,

near Lake Gunn, tributary of Eglinton River, ca. 365 m,

Child H2597 (F); Fiordland Natl. Park, Lake Gunn, Zo-

tov as T. gottscheana (CHR); ibid., Central Earl Mts.,

Mistake Creek, between Triangle Peak and Melita Peak,

740-800 m, Engel 18785 (F); ibid., Gertrude Valley, off

track to Gertrude Saddle, NE of Homer Tunnel, 1 880-

1900 m, Engel 18899 (F); ibid., Milford Sound, along

path to lower end of Bowen Falls, sea level, Schuster

55548b as Lepidozia concinna (F); ibid.. Moraine Creek

Track, area N of Moraine Creek, W of Hollyford River,

610 m, Engel 23210 (F). OTAGO PROV.: Mt. Cargill,

just below summit, N of Dunedin, ca. 670 m, Engel

17576 (F); ibid., W of summit road, ca. 455 m, Child

H2044 as Lepidozia tetradactyla (F); Silver Peaks

Range, N of Dunedin, Allison H5911 (CHR); Lee

Stream, Hindon Road, ca. 305 m, Child H929 (F); near

Herbert, Oamaru District, Allison H5907 as T. praeni-

tens (CHR); Rock and Pillar, ca. 915 m, Child H2751

as Temnoma palmatum (F); Paradise, Dart Valley, N of

N end of Lake Wakatipu, ca. 365 m, Child HI 193 as T.

meridiana (F); trail between Gunn's Hut and Hidden

Falls, ca. 30 m SE of Lake McKerrow, Hatcher 580 as

Lepidozia meridiana (F); Mt. Aspiring Natl. Park, below

and W of Mt. Shrimpton, 1370-1470 m, Engel 17874

(F); Makarora River, near base of Haast Pass, Allison

H5968—C. sporo. as T. gottscheana (CHR). OTAGO/

WESTLAND PROV. BOUNDARY: Mt. Aspiring Natl.

Park, summit area of Haast Pass, 570 m, Engel 17984 —

c. sporo. (F). WESTLAND PROV: Kellys Range, Beck-

ett 4304, 4599, syntypes of Lepidozia beckettiana-(both)

c. per. (G); Turiwhate Falls on Route 73, Engel 6531 A

(F); Mt. Aspiring Natl. Park, Blue Valley Track, above

Blue River just N of confluence with Makaroa River,

430-480 m, Engel 21919 (F); ibid., Cross Creek, 1.1

km N of Haast Pass, 540 m, Engel 21871 (F); Camp

Creek, W of Alexander Range, 630 m, Reif C270B (F);

Paparoa Range, ridge immediately N of Sewell Peak,

910m, Engel 19042 (F). WESTLAND/CANTERBURY

PROV: Arthur's Pass Natl. Park, Arthur's Pass area, Up-

per Twin Creek Valley, 930 m, Engel 18528 (F); ibid.,

Halpin Creek, at junction with main road, Hair & Beu-

zenberg (CHR). CANTERBURY PROV: Arthur's Pass,

Moore H77 (CHR); ibid., Hair 3&4pp (CHR); Arthur's

Pass Natl. Park, Bealey River, off Bealey Valley Track,

830-850 m, Engel 18509 (F). NELSON PROV: Big

Totara River, ca. 60 m, Child H5448 (F); Nelson Lakes

Natl. Park, NE margin of Lake Rotoroa, W of St. Ar-

naud, 520 m, Engel 21500 (F); ibid., E slope of Robert

Ridge in vicinity of Mt. Robert Skifield, W end of Lake

Rotoiti, 1400-1480 m, Engel 22828 (F); ibid., Upper

Travers River, Simpson 3332 (CHR); Kahurangi Natl.

Park, Mt. Arthur, Mt. Arthur Hut Track, 1050 m, Engel

24927 (F). NORTH ISLAND. WELLINGTON PROV:

Tararua Mts., Field Hut, ca. 825 m, Zotov as L. praen-

itens — c. d (CHR); Tararua Range, Ohau Stream, Moore

H171 as T. gottscheana (CHR); ibid., Mt. Holdsworth,

Wormald 14pp as T. gottscheana (CHR); Te Matawai,

Zotov as L. capilligera; Tongariro Natl. Park, Soda

Springs, Mangatepopo Stream, NE of Whakapapa Vil-

lage, 1350 m, Engel 22491 (F). SOUTH AUCKLAND

PROV: Ruahakune Bush, E of Taupo, ca. 760 m, Allison

H5893 as T. gottscheana (CHR); Rotorua, hot springs

area. Hatcher 29 (F); Whareorino Forest, start of track

to Leitch's Hut, 278 m, Engel 23766 (F); Mt. Te Aroha,

ca. 3 km E of Te Aroha, 900-940 m, Engel 22146 (F).

NORTH AUCKLAND PROV: NE Waitakere Ranges,

Swanson University Reserve, Tram Valley Road, 95 m,

Engel 20470 (F); Waitakere, ca. 120 m, Child H2234

(F); SE corner of Waipoua Forest, just N of Tutamoe,

540 m, Engel 21119 (F); ibid., Mataraurau Plateau,

Waoku Road, Braggins 94/240 (AKU). LITTLE BAR-

RIER IS.: Hynes as T. gottscheana (CHR). GREAT

BARRIER IS.: Kirk 176, 183, both as L. gottscheana

(CHR). TASMANIA: Trail to Adamsons Peak between

Manuka Flat and the hut, 500-800 m, Norris 27048 (F);

along Old Hartz Trail about 1 mile beyond Kermandie

Plains, ca. 650 m, Norris 29674 (F); Geeveston Forestry

District, just S of Spur 2 on Hermans Road, 410 m,

Blanks (HO); Mt. Wellington, Deep Creek, Rodway

n.s. — c. sporo. as L. praenitiens (HO); same loc., 490

m, Weymouth 1516 — c. sporo. as L. praenitiens (HO);

Manuka Road, W of Tahune Bridge, 1 30 m, Jarman 99/

4—c. S + per. (HO); near Scotts Peak Road, 2.2 km S

of junction with Gordon River Road, E of S end of Lake

Gordon, 580 m, Engel 13783 (F); plateau region E and

SE of Mt. Eliza summit, E of Lake Pedder, ca. 1225 m,

Engel 13668G (F); Cradle Mtn.-Lake St. Clair Natl.

Park, Ballroom Forest, SW side of Lake Dove, 950-

1050 m, Engel 14016 (F); Paddys Lake, below and E of

Black Bluff, SW of Nietta, 1090 m, Engel 19784 (F);

Flinders Is., Mt. Strzelecki, Cronley's Ck, Scott s.n.

(MELU).

Telaranea tetrapila (Hook. f. & Tayl.) Engel &

Merr.

Lepidozia tetrapila Hook. f. & Tayl. in Taylor, Lon-

don J. Bot. 5: 370. 1846. Mastigophora tetrapila

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

III. 4: 416. 1877. Telaranea tetrapila (Hook. f. &

Tayl.) Engel & Merr., Phytologia 79: 253. June,

1996 [1995]. Lectotype (nov.): New Zealand,

Hooker "n. 119. 1844" (FH!); isolectotype: (W!,

Lindenberg Hep. no. 4739).

Plants soft, flexuous yet firm, loosely prostrate

to suberect, in dense, compact mats, pale green to

yellowish green to olive green (to pale amber in

van cancellata), nitid when dry, often somewhat

water-repellent; plants variable in stature, small to

medium, to 1.5 cm wide, including branches.

Branching regular and rather densely pinnate,

sometimes 2-pinnate, the main axis often bearing

only 2-4 normal leaves (on each side) between

branches, the branches of the Frullania type, short

to moderately long but normally determinate, of

about equal length or (especially in robust plants)

gradually becoming longer from shoot apex to

base (e.g., plumose), hardly tapering, sporadically

flagelliform; branch half-leaf 2(4)-lobed, usually

52

FIELDIANA: BOTANY

obliquely inserted, narrowly rectangular, the lobes

parallel to weakly diverging; first branch under-

leaf undivided and ciliiform or, less often, bifid,

sporadically trifid, inserted on ventral side of

branch near or (often) at juncture of branch and

main axis. Ventral-intercalary branches occasion-

al, leafy, often becoming leading shoots. Stems

with cortical cells distinctly differentiated, thin

walled, in 11-18 rows, those on ventral side of

stem a little smaller: cortical cells in section much

larger than the numerous (35-65) medullary cells,

the medullary cell walls uniformly slightly thick-

ened. Rhizoids from distal cells of underleaf disc.

Leaves on main shoot rigid, obliquely spreading,

distant (small phases) to loosely to closely imbri-

cate, plane to moderately convex, the insertion

weakly to distinctly incubous; leaves 630-980 u,m

wide X 630-840 jxm long, subsymmetric, 4-6(8)

lobed to 0.4-0.55, the lobes straight to widely di-

vergent, typically shorter than the disc (longer

than the disc in var. cancellala). Lobes typically

subcaudate, the base subtriangular, 2-4 cells wide

at extreme base, terminating in a uniseriate row

of (4)5-6(8) cells (typically more than half the

length of lobe); cells of the uniseriate portion

short, often ± barrel shaped (longer cells of the

uniseriate portion to 2.5:1 in var. cancellata), rath-

er thin-walled, the septae thickened in the corners

but the septa not swollen and projecting, the basal

cell of the uniseriate portion 34-52 (Jim wide X

48-72 |o,m long (1.1-1.6:1), the next cell narrower

but of about the same length, 28-40 u,m wide X

41-62 (im long (1.2-2.2:1), the terminal cell nor-

mally smaller than the penultimate cell, the distal

lobe cells often short quadrate and more numer-

ous resulting from secondary cell divisions; cuti-

cle of at least the distal half of lobes finely to ±

distinctly striate-papillose. Disc symmetrically to

somewhat asymmetrically broadly subrectangular

to cuneate, 4-6(7) cells high (from median sinus

base to leaf base), (9)14-21 cells wide in distal

portion narrowing to 8-12 cells wide in basal por-

tion (disc typically 8 cells wide throughout in var.

roseana); margins entire, ± straight to less often

curved; cells of disc margins often bulging. Cells

of disc thin-walled but firm, trigones lacking or

minute, large, convex, "pillow-like" (obvious

even under low magnification), the largest cells

(in median sector of disc) large, often ± isodia-

metric, (45)50-70 u.m wide X 55-90 u,m long;

basal 1-2 rows of disc cells considerably larger

(wider), irregularly arranged or in ± regular tiers;

cells of disc margins (especially the dorsal) some-

times forming an indistinct border of elongate.

narrower cells; cuticle smooth. Oil-bodies grayish

(var. tetrapila) or hyaline (var. cancellata),

coarsely papillose, the spherules protruding be-

yond membrane, the median disc cells with 5-1 1

oil-bodies per cell (8-9 per in lobe cells), narrow-

ly to broadly elliptic to subcrescentic and 8.3-

1 1.6 X 5.5-5.8 UJTI, less often ca. 14.7 X 5.3 u.m,

the subspherical ones 5.9-6.6 u,m in diam. Un-

derleaves normally smaller than leaves (subequal

to leaves in var. cancellata). strongly spreading,

contiguous to loosely imbricate, plane, 4-6-lobed

to 0.4-0.6, the lobes somewhat divergent, cili-

iform, the uniseriate portion formed of 3-5(8)

somewhat elongated cells; disc symmetrically cu-

neate, (3)4-5 cells high (median sinus), the cells

in ± regular tiers or irregularly arranged, the disc

(10)12-19 cells wide in distal portion narrowing

to 8-10 cells wide at base; margins entire, usually

straight. Rhizoid initial cells small, subquadrate,

at bases of lobes or forming a continuous bistra-

tose band across the apex of the disc. Asexual

reproduction lacking.

Plants dioecious. Androecia on short, abbrevi-

ated, ventral-intercalary branches from leading

shoots, as well as from flagelliform or stoloniform

branches; bracts closely imbricate, dorsally assur-

gent, deeply concave, 2-3 lobed, each lobe ter-

minating in a single cell or a uniseriate row of 2

not to hardly elongated cells; lamina cells irreg-

ular in shape and arrangement, the dorsal margin

of lamina feebly dilated and slightly incurved, ir-

regularly crenulate, at times with a few sessile

slime papillae; bracts monandrous; antheridia

large for bract size, the stalk short, ca. 7 cells

high, uniseriate; bracteolar antheridia absent. Gy-

noecia oriented laterally and at best only weakly

dorsally assurgent, with a vestigial stem perigyn-

ium present, swollen, rhizoidous; bracts and brac-

teoles in 2-3 series, inserted on the vestigial peri-

gynium, becoming progressively larger and less

deeply lobate towards the perianth, those of in-

nermost series ensheathing the perianth, deeply

concave, the apical portion canaliculate; bracts of

innermost series ± suborbicular, irregularly 4-lob-

ulate, the lobules terminating in a uniseriate row

of 1-2 cells; bract margins crenate to dentate, the

armature frequently sharply indexed, often ter-

minating in a slime papilla, the bracts with an ob-

scure border formed of 1 to several rows of cells

longer, narrower and more irregular than those

within; bracteoles of innermost series nearly iden-

tical in form" to bracts although a little smaller,

free from bracts. Perianth not extending above

vegetative axes, 0.5-0.55 emergent, ovoid-cylin-

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

53

drical, terete in basal half, the distal half obscurely

trigonous and with 6-9 plicae, the sulci shallow

to deep; perianth narrowed toward a decidedly

contracted mouth, the mouth with 6 narrowly tri-

angular lobes, each lobe fringed with slightly

thick-walled, contorted, crowded, sparingly papil-

lose cilia, the terminal cell of each cilium coarsely

papillose.

Seta with 8-9 rows of outer cells, surrounding

an inner core of 20-34 much smaller cells. Cap-

sule short-cylindric, 1204-1470 u.m X 560-805

fxm, the wall 41-50 jxm thick, of 4 layers (locally

3), the outer layer subequal to the combined thick-

ness of the inner layers, or slightly less thick; out-

er layer of cells in tiers, ± regularly short-rect-

angular, with 2-phase development rather indis-

tinct, the longitudinal secondary walls with sheet-

like thickenings and nodule- to spine-like

thickenings (4-6 per cell) alternating with pri-

mary walls also with similar thickenings, the

transverse walls almost devoid of thickenings, or

with local, nonpigmented, nodular swellings; in-

termediate layers (when 2 in number) thinner than

the outer or innermost layers; innermost layer of

cells ± tiered, rather regularly narrowly rectan-

gular, with semiannular bands common, rather

narrow, complete, only rarely forked to delimit

local fenestrae.

Spores 12-13.4 jxm in diam., exine rather thick

in optical section, dark yellow brown, areolate

(with a low, close network of well-defined, rather

thick, furcate ridges that anastomose to form are-

olae). Elaters rigid, ± straight, 9.1-12.5 jxm wide,

only slightly tapering toward tips, bispiral to tips,

the spirals 3.8-5.3 jxm wide.

NOMENCLATURE — The original specimen of

Lepidozia tetrapila in Taylor's herbarium (FH) in-

cludes plants of Telaranea tridactylis and a larger

Telaranea with prominent, pillow-like disc cells.

The statements in the protologue, "cells remark-

ably large and strongly defined" and "stems

about 1 one inch long," and the reference to "fla-

gelliform shoots" can only apply to the larger

plant. This element is therefore selected as the lec-

totype. Also admixed with the type are Lepidozia

pendulina, Chiloscyphus spiniferus, Haplomitrium

gibbsiae, and Isotachis lyallii, the last three seg-

regated by Taylor into separate micropackets.

The type locality of T. tetrapila is not given in

the protologue, however the presence of T. tri-

dactylis, a species known to occur only on Camp-

bell Is., Auckland Is. and Tasmania, strongly sug-

gests that the plant came from the subantarctic

islands. Hodgson (1956, p. 597), who had not

seen the type, mentioned this name in passing as

a possible older name for Lepidozia laevifolia.

DIFFERENTIATION AND VARIATION — Telaranea

tetrapila is the oldest name for the species which

has generally been called Telaranea (or Lepido-

zia) gottscheana in the New Zealand literature

(e.g., Hodgson, 1956; Allison & Child, 1975).

The type of the name L. gottscheana, however, is

T. patentissima. At its best, T. tetrapila is one of

the most handsome of our Telaranea species, and

is common throughout New Zealand. The plants

are highly variable in size, color, and general as-

pect, as well as leaf form and number of lobes.

The most useful features for distinguishing the

species are the large, pillow-like, almost geomet-

rically hexagonal disc cells (Figs. 18: 4; 19: 3),

which are conspicuous and easily observed, even

under the dissecting microscope, the bulging cells

of the lobe bases and the short, secondarily divid-

ed cells at the lobe tips (Figs. 18: 5, 6; 19: 9).

One marked expression of T. tetrapila (van ro-

seana) bears a strong resemblance to T. tuberi-

fera, due to its widely spreading, parallel-sided

leaves (Fig. 19: 1). Another (var. cancellata)

somewhat resembles T. gibbsiana in the finely

elongated leaf lobes, and large, ciliiform-lobed

underleaves.

Examination of herbarium specimens indicates

that T. tetrapila has often been confused with T.

praenitens, which differs by its more deeply-

lobed, palmate leaves and the characteristic mi-

nute denticulations formed by the swollen pro-

jecting end-walls on the lobes and disc margins.

Key to the Varieties of T. tetrapila

\. Leaves rather flat and widely spreading, the

disc parallel-sided and somewhat higher than

wide (resembling T. tuberiferd), 8 cells wide

in both distal and basal portions; lobes often

straight and aligned with the disc margins . . .

var. roseana

1. Leaves weakly to distinctly convex, .and

obliquely spreading, the disc weakly to dis-

tinctly cuneate, (9)14-21 cells broad distally,

the margins often curved; lobes divergent . . 2

2. Lobes typically shorter than the disc, the

cells of the uniseriate row short, often ±

barrel-shaped; underleaves smaller than

leaves; oil-bodies greyish, 5-8 per cell.

Common throughout New Zealand ....

var. tetrapila

54

FIELDIANA: BOTANY

11

FIG. 18. Telaranea tetrapila (Hook. f. & Tayl.) Engel & Merr. var. letrapila. I. Sector of leading shoot, dorsal

view. 2, 3. Leaves. 4. Leaf, cellular detail. 5, 6. Median lobes of leaf; note secondary septa. 7. Underleaves. 8.

Underleaf disc, cellular detail, stippled cell walls indicate rhizoid initial cells. 9. Stem, cross section. 10. Plant, dorsal

view. 1 1. Antheridium (as seen in optical section). (Figs. 1, 3, 6-7, 10 from Engel 18029, New Zealand, South Is.,

Otago/Westland Prov. boundary, Mt. Aspiring Natl. Park, summit area of Haast Pass; 2, 5, 8 from Reif CJ5IA; 4,

from Reif C93E; 9, from Reif C80F, all Reif coll. from New Zealand, South Is., Westland Prov., Camp Creek; 1 1

from Engel 19298, New Zealand, South Is., Westland Prov., Paparoa Range, Croesus Track.)

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

55

FIG. 19. Telaranea tetrapila (Hook. f. & Tayl.) Engel & Merr. Var. roseana (Steph.) Engel & Merr (1-3); var.

cancellata (Col.) Engel & Merr. (9-12); Telaranea paludicola (Hodgs.) Hodgs. (4-8). 1. Sector of main shoot, dorsal

view. 2. Leaves. 3. Leaf, cellular detail. 4. Leaves. 5, 6. Leaves, cellular detail. 7. Sector of main shoot with branch

bases and associated half-leaves, dorsal view. 8. Leaves of extreme. 9. Leaf lobe showing secondary cell divisions

in distal portion. 10. Underleaf. 11. Leaves (note, figure drawn at 50% larger than other leaf outlines). 12. Distal

sector of leaf. (Figs. 1-3 from Fife 4964, New Zealand, South Is., Nelson Prov., Fox River; 4, 6-7 from type of

56

FIELDIANA: BOTANY

2. Lobes longer than the disc, the cells of the

uniseriate row ± elongate (to 2.5:1), not

barrel-shaped; underleaves subequal to

leaves in size; oil-bodies hyaline, 8-1 1 per

cell. Scattered sites on North Island

var. cancellata

Telaranea tetrapila var. roseana (Steph.) Engel &

Merr.

Lepidozia roseana Steph., Spec. Hep. 3: 590. 1909.

Neolepidozia roseana (Steph.) Fulf. & J. Tayl.,

Brittonia 11: 85. 1959. Telaranea roseana (Steph.)

Hodgs., Rec. Domin. Mus. 4: 106. 1962. Telaranea

tetrapila var. roseana (Steph.) Engel & Merr., Phy-

tologia 79: 253. June, 1996 [1995]. Type: New

Zealand, without specific loc., 1898, Petrie, "com.

Rose" (G!).

Plants resembling T. tuberifera; leaves rather

flat and widely spreading; leaf lobes typically

shorter than the disc, subcaudate, ± parallel with

the disc margins; leaf disc parallel-sided and

somewhat higher than wide, typically 8 cells wide

in both distal and basal portions, the cells con-

spicuously large, hexagonal (as in var. tetrapila).

DIFFERENTIATION AND VARIATION — Plants as-

signed to this variety often bear a striking resem-

blance to Telaranea tuberifera, but differ by not

having the glaucous cuticle or the strongly hori-

zontal leaf orientation of that species; it also dif-

fers by the nonfragmenting lobe apices, and the

absence of tubers at the tips of flagelliform sto-

lons. The leaves are obliquely oriented, as in var.

tetrapila, but are typically rather flat, and diverg-

ing at a wider angle with the stem. As indicated

in the species description, T. tetrapila is very var-

iable, and even well-developed plants will have

some leaves which are only 8 cells wide through-

out the disc, particularly on the branches. The var.

roseana, when well developed, has a very char-

acteristic appearance, due to the conspicuously

parallel-sided disc (somewhat higher than wide),

which is typically 8 cells wide in both distal and

basal portions (Fig. 19: 3), and the lobes are often

straight and aligned with the disc margins.

The var. roseana differs from Telaranea palu-

dicola by the more slender lobes, which are 2 (and

only sporadically 3-4) cells wide at the base, and

the very prominent, hexagonal disc cells are ar-

ranged in precise geometrical fashion.

Hodgson (1956, p. 608) treated this plant as T.

corticola, describing the leaf cell size as 50-60

u,m, but referring to Stephani's (1909) measure-

ments as "much smaller, being only 27 X 36 u,."

Examination of the type of T. corticola, however,

reveals that this species' is a synonym of T.

patentissima (p. 46).

DISTRIBUTION AND ECOLOGY — The var. roseana

is known primarily from South Island, New Zea-

land in lower- to middle-elevation forests, espe-

cially on damp, clayey banks. Known in the North

Island only from Wellington, South and North

Auckland Provinces.

SELECTED SPECIMENS SEEN — NEW ZEALAND.

SOUTH ISLAND. SOUTHLAND PROV.: Waikaia

State Forest, junction of Waikaia River Track and Tison

Peak Track, upstream from Piano Rat Recreational

Area, ca. 260 m, Engel 24721 (F); Fiordland Natl. Park,

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

draining from Lake Wade, 300 m, Fife 7729 as T. cf.

patentissima (F). OTAGO PROV.: Blue Mts., ca. 455 m,

Child Hs.n.159 (F); Waipori Gorge, S of Dunedin, Al-

lison H7809 (CHR); Paradise, Dart Valley, N of N end

of Lake Wakatipu, ca. 365 m. Child H1248 as T. praen-

itens (F). WESTLAND PROV.: Lake Kaniere Scenic

Reserve, Lake Kaniere Rd, 125 m, Engel 24826 (F);

Rapahoe Range (or Twelve Apostles Range), N of Grey-

mouth, ca. 150 m. Child H5393 as T. centipes (F); Pa-

paroa Range, along Croesus Track, ESE of Barrytown,

between Granite Creek and Pagan Creek watersheds,

250 m, Engel 19287 (F); Hari Hari, Langridge as T.

praenitens (CHR). NELSON PROV.: Track on N bank

of Fox River, ca. 1.5 km E of Route 6 Fife 4964 — c.

per. (F). NORTH ISLAND. WELLINGTON PROV.:

Tongariro Natl. Park, Mangawhero River near juncture

of Rimu Track and Forest Walk, just NE of Ohakune,

625 m, Engel 22711 (F). SOUTH AUCKLAND PROV.:

Whareorino Forest, start of track to Leitch's Hut, 280 m,

Engel 23746 (F); Coromandel State Forest Park, Kau-

raeranga River, just SE of suspended footbridge, 150 m,

Engel 22287 (F); ibid., Mt. Moehau, below summit of

"Little Moehau," ca. 800 m; Engel 23670 (F). NORTH

AUCKLAND PROV: Waipoua Forest, Lookout Track

between Lookout Road and Waipoua River Road, above

forest headquarters, ca. 240 m, Engel 21082 (F); Wai-

poua Kauri Forest, Hatcher 441 (F); Omahuta Forest

Sanctuary, E of Mangamuka Bridge, 320 m, Engel

21035 (F).

Telaranea tetrapila var. cancellata (Col.) Engel

& Merr.

L paludicola; 5, 8 from Child 5446, New Zealand, South Is., Nelson Prov., Big Totara River; 9-12 from Zotov 7083,

New Zealand, North Is., Wellington Prov., Ohau-iti River.)

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

57

ti cancellata Col., Trans. & Proc. New Zea-

land Inst. 18: 244. 1886. Telaranea tetrapila var.

(•(incellata (Col.) Engel & Merr, Phytologia 79:

253. June, 1996 [1995]. Type: New Zealand, Wai-

pawa Co., near Norsewood, edge of Bartramia

Creek, 1885, Colenso a.1418 (BM!, WELT!).

Psiloclada digitata Col., Trans. & Proc. New Zealand

Inst. 18: 243. 1886, syn. nov. Type: New Zealand,

Waipawa Co., near Norsewood, 1885, Colenso

a. 1378 (BM!, WELT = 0).

Plants resembling T. gibbsiana, often pale am-

ber in color; plants medium, to 1.1 cm wide, in-

cluding branches; branching somewhat irregularly

to regularly 1 -pinnate; leaf lobes longer than the

disc, subcaudate, rather abruptly tapering to a un-

iseriate row of cells, ± straight, the cells of the

uniseriate row ± elongate (to 2.5:1), not barrel-

shaped; leaf disc at most moderately asymmetric,

the disc cells mostly isodiametric; oil-bodies gray-

ish, 5-8 per cell; underleaves subequal to the

leaves in size, with long ciliiform lobes, the ter-

minal cells often secondarily divided.

DIFFERENTIATION AND VARIATION — Colenso

(1886, p. 245) aptly described this as "a truly el-

egant plant." It is chiefly distinguished by the

rather regularly 1 -pinnate shoots; the often pale

amber pigmentation; the slender leaf lobes, which

are subcaudate as in the var. tetrapila, but with a

uniseriate row composed of elongated (to 2.5:1)

cells; and the rather large underleaves, with long,

ciliiform lobes, rather like those of T. gibbsiana.

It differs from T. paludicola in the ± isodiametric

cells of the disc and lobe bases, and the abruptly

tapering lobes, with a long, uniserate row, which

are ± straight, versus lobes biseriate, often falcate

and hooked at the tips in T. paludicola.

DISTRIBUTION AND ECOLOGY — Known only from

several scattered sites on North Island, "on trees

and logs, forming large and thick patches" (Col-

enso, I.e., p. 245). A slip of newspaper with the

type (WELT) indicates that the plant was associ-

ated with the moss Cyathophorum. On Mt. Te

Aroha, plants occurred at 880-890 m loosely on

the floor as well as over soil deep in a protected

niche under the lip of the forest overhang, both

populations in a stunted Nothofagus menziesii for-

est associated with Dracophyllum and Quintinia

serrata. In the Herangi Range the species oc-

curred at ca. 720-750 m on an exposed plateau

with a mosaic of subalpine bog vegetation, stunt-

ed Quintinia serrata, Dracophyllum-heath (to 1 m

tall), rocky outcrops and small water channels. At

this site plants were found in moist niches, e.g.,

on the floor or beneath blades of Poa sp.

SELECTED SPECIMENS SEEN— NEW ZEALAND.

NORTH ISLAND. WELLINGTON PROV.: Ohau-iti R.

(Ohauite Stream), Zotov 7083 (CHR!— c. per.); Tararua

Mts., Akatarawa Saddle, Zotov 9294 (CHR— c. sporo.).

GISBORNE PROV.: Waikaremoana, Wairoa, Hodgson

111 (CHR). NEAR GISBORNE/SOUTH AUCKLAND

PROV. BOUNDARY: Urewera Natl. Park, track to

Whakataka Hut from Lake Waikaremoana, N of western

extremity of Lake Waikaremoana, 1 160 m, Engel 23284

(F). SOUTH AUCKLAND PROV.: Southern extent of

Herangi Range, plateau area S of Te Whakapatiki, W of

Te Kuiti township, ca. 720-750 m, Engel 25127 (F);

Kaimai-Mamaku Forest Park, Mt. Te Aroha, S facing

slope, 880-890 m, Engel 23828— c. 6 (F). GREAT

BARRIER IS.: Coffin's Creek Camp, 245 m, Lloyd

260pp (CHR).

Telaranea tetrapila (Hook. f. & Tayl.) Engel &

Merr. var. tetrapila

Plants with shoots to 1.5 cm wide (including

branches); branching regularly and rather densely

pinnate (often plumose in well-developed plants);

leaf lobes typically shorter than the disc, subcau-

date, ± straight, the cells rather short, barrel-

shaped; leaf disc at most moderately asymmetric,

often distinctly cuneate, (9)14-21 cells broad dis-

tally, composed mostly of large, hexagonal cells,

except for distal tier at base of lobes; oil-bodies

hyaline, 8-1 1 per cell.

DISTRIBUTION AND ECOLOGY — The typical vari-

ety is common throughout New Zealand in low-

to middle-elevation, forests (typically below 700

m), where it can be particularly luxuriant. It can

occur in the upper limits of Nothofagus menziesii

forests or in subalpine scrub, but at these upper

elevations the plants often represent a small or

weak phase of the species (it nevertheless may

produce sporophytes and androecia and may be

well developed in such niches). It grows on an

array of substrates, but typically can be found on

the forest floor, on soil of mossy banks, on rotted

logs, or, at times, on upright tree trunks. It is able

to tolerate some degree of exposure and can occur,

for example, in open boggy areas with Juncus

squarrosus, Coprosma, Leptospermum, Cassinia,

etc. (Mt. Cargill area), or in pakihis, with Glei-

chenia, Lycopodium, Juncus, Drosera spp., Schiz-

aea and Leptospermum, etc. (Tiropahi River area).

SELECTED SPECIMENS SEEN— NEW ZEALAND. SO-

LANDER IS.: Summit, 330 m, Johnson (CHR); ibid.,

plateau, 300 m, Johnson (CHR); ibid., SW Bay, 100 m,

Johnson (CHR). STEWART ISLAND: Cockayne 8383

as L. remotifolia (CHR); Rakiura Natl. Park, Port Peg-

asus, immediately adjacent Belltopper Falls, ca. 10-80

m, Engel 24137 (F); Pegasus, Schuster, Scott & Taylor

58

FIELDIANA: BOTANY

(CHR); Mt. Rocky summit area, 530 m, Engel 24292

(F); Ulva Is., Paterson's Inlet, Hatcher 1589 (F); near

Oban, Half Moon Bay, Hatcher 1635 (F); Christmas Vil-

lage, Mt. Anglem, 170 m, Scott as T. roseana (CHR).

SOUTH ISLAND. SOUTHLAND: Bluff Hill, 90-120

m, Allison H5163 (CHR); ibid.. Kirk 527 (CHR); Fiord-

land Natl. Park, Mt. Burns, Burns Creek, ca. 1280 m,

Child H5117 (F); ibid., Stuart Mts., W shore of Lake

Thomson N of stream draining from Lake Wade, 300 m,

Fife 7607, 7726 both as T. cf. patentissima (F); ibid.,

off track along East Branch of Eglinton River, SE of Mt.

Eglinton, 440 m, Engel 18748 (F). OTAGO PROV.:

Lake Wilkie, Catlins District, Burrell (CHR); Catlins

River area, near Purakanui, NE of Maclennan, ca. 150

m. Child 794— c. <J (F); Blue Mts., S end, E of Garden

Gully Road, Child Hs.n.158 (F); near Akatore Stream,

coastal, S of Dunedin, Allison H5898 (CHR); S of Taieri

River, S of Dunedin, ca. 180 m, Allison H5897 (CHR);

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

Engel 17529 (F); immediately N of Mt. Cargill, N of

Dunedin, 525 m, Engel 17593— c. sporo. (F); Mt. Car-

gill, Mt. Holmes, N of Dunedin, ca. 550 m, Child H1075

as T. corticola (F); W slope of Flagstaff, NW of Dun-

edin, 560 m, Engel 17634 (F); Silver Peak, Simpson as

L praenitens (CHR); Leith Valley, near Dunedin, Child

H102 (F); Route Burn, ca. 520 m, Child Hs.n.67 (F);

Mt. Aspiring Natl. Park, Blue River near confluence

with Makarora River, NNE of Makarora, 310 m, Engel

18914 (F). OTAGOAVESTLAND PROV. BOUNDARY:

Mt. Aspiring Natl. Park, summit area of Haast Pass, 570

m, Engel 17987 (F). WESTLAND PROV.: Between

confluence of Jackson River and Arawata River and

Lake Ellery, off Jackson River Road, sea level, Engel

17944— c. per. (F); Lake Paringa, ca. 15 m, Child HI 886

(F); Ship Creek, 14.5 km N of Haast River, sea level,

Engel 21760 (F); Westland Natl. Park, terminal moraine

of Fox Glacier, S side of Fox River, Engel 6617 A (F);

ibid., Franz Josef Glacier Valley, Roberts Point, SW of

Mt. Gunn, ca. 620-670 m, Engel 18089— c. per. (F);

ibid., track to Alex Knob just below juncture with track

to Louisa Peak, 1150-1170 m, Engel 19002 (F); ibid.,

trail to Lake Wombat, Engel 6694 (F); Arthur's Pass

Natl. Park, Bealey River, off Bealey Valley Track, 830-

850 m, Engel 18469 — c. sporo. (F); ibid., Upper Twin

Creek Valley, 930 m, Engel 18529A—C. sporo. + <J (F);

ibid., N of Kellys Creek near Hwy 73, N of Otira, 420-

475 m, Engel 18329 (F); Camp Creek, W of Alexander

Range, 190-990 m, Reif C3F— c. <J, C260E (F); White

Horse Creek, ca. 150 m. Child H54I1— c. 6 (F); Pa-

paroa Range, road to Sewell Peak, 710 m, Engel 19075

(F); ibid., along Croesus Track, ESE of Barrytown, be-

tween Granite Creek and Pagan Creek watersheds, 450

m, Engel 19298— c. d (F); Mt. Aspiring Natl. Park,

Cross Creek, 1 km N of Haast Pass, 510m, Engel 231 14

(F). WESTLAND/CANTERBURY PROV. BOUND-

ARY: Arthur's Pass Natl. Park, Bridal Veil Track, E side

of Bealey River and just N of town of Arthur's Pass,

760-825 m, Engel 22939 (F); ibid., Bealey Valley Track,

ca. 875-900 m, Engel 22875 (F). NELSON/WEST-

LAND PROV. BOUNDARY: Paparoa Range, S side of

Porarari River, up river from gorge and ca. 500 m WSW

of ford on inland track to Bullock Creek, 10-20 m. En-

gel 19209 (F); Lower Basin (Syncline area) of Porarari

River, 30-45 m, Fife 6667 (F); track between Punakaiki

and Porarari Rivers, 150 m, Fife 6078 (F). NELSON

PROV.: Paparoa Natl. Park, Bullock Creek Road, along

Bullock Creek, NE of Punakaiki, ca. 25 m. Engel 21578

(F); Paparoa Range, upper slopes of N side of Tiropahi

or Four Mile River, between sea and Route 6, S facing

side of gorge, 130-170 m, Engel 19246 (F); Maimai

State Forest, 3 m NW of Reefton, 280-350 m, Macmil-

lan 76/169 as T. praenitens (CHR); upper Buller River

gorge, near Lyell, Allison 6464 (CHR); track to German

Terrace, 6 km SSE of Westport on Nine Mile Road, 90

m, Engel 21551 (F); Nelson Lakes Natl. Park, NE mar-

gin of Lake Rotoroa, 520 m, Engel 21496 (F); ibid., off

Lakehead Track, near juncture with southern end of

Loop Track, NE end of Lake Rotoiti, 630 m, Engel

22736 (F); Big Bush State Forest 48, Donald Creek,

480-600 m, Macmillan 77/40 (CHR); Able Tasman

Natl. Park, Mt. Evans, 1067 m. Child 4703B (F); ibid..

Porters Rock Track, ca. 915 m. Child H4680 as L. cf.

concinna (F). NORTH ISLAND. WELLINGTON

PROV.: Orongorongo River, ca. 610 m, Zotov 7148 as

L. praenitens (CHR); Tararua Mts., Field Hut, 825 m,

Zotov 7633, 7374 as L praenitens (CHR); ibid.. Table

Top, Moore HI 75 (CHR); ibid., Akatarawa Saddle, Zo-

tov 9295 as L. praenitens (CHR); ibid., Ruamahanga

Valley, Zotov 9267 as L praenitens (CHR); Ohau-iti

River, Zotov 7090 as L. praenitens (CHR); Ruahine

Range, Pohangina Valley, ca. 855 m, Child HI94—C.

sporo. (F); Tongariro Natl. Park, ca. 8 km from Ohakune

on Ohakune Mt. Road, ca. 950 m, Engel 21326 (F);

ibid., near Ohakune Mt. Hut, Mt. Ruapehu, 1370 m,

Sainsbury (CHR); ibid., Mt. Hauhungatahi, Mt. Ruape-

hu, Moore (CHR); ibid., Mahuia Track, Barr (CHR).

HAWKES BAY PROV.: "Kiwi," Wairoa, Hodgson 203

(CHR). TARANAKI PROV.: Mt. Egmont, Ngaroto

Track, Barr 385 (CHR); Pukeiti Bush, near New Plym-

outh, Hatcher 293 — c. per. (F); Mt. Messenger, ca. 185

m, Allison H5905 (CHR). NEAR GISBORNE/SOUTH

AUCKLAND PROV. BOUNDARY: Urewera Natl.

Park, crest trail from highway 38 towards Whakataka

summit, N of northern extremity of Lake Waikaremoana,

930-1030 m, Engel 20665 (F). GISBORNE PROV:

Lake Waikaremoana, Hodgson as L. praenitens (F);

Urewera Natl. Park, Huiarau Range, summit area of Te

Rangaakapua, 1230-1320 m, Engel 23461 (F). SOUTH

AUCKLAND PROV.: W of Lake Taupo, Matthews

(CHR); Mamaku Plateau, W of Lake Rotorua on Rt. 5,

400 m, Engel 20507 (F); southern extent of Herangi

Range, vicinity of plateau area S of Te Whakapatiki, W

of Te Kuiti township, ca. 720-750 m, Engel 25158 (F);

Herangi Range, Wharcorino Forest, near tributary of

Awakino River, W of Leitchs Road, and SW of Te Kuiti

township, 285 m, Engel 23934 (F); Kaimai Range, net-

work of tracks at the end of Wrights Rd. off the Katikati-

Tauranga highway. Swimming Holes Track, 60 m, Engel

23537 (F); Mt. Te Aroha, ca. 3 km E of Te Aroha, 900-

940 m. Engel 22150 (F); Coromandel State Forest Park,

summit of Table Mt., 835 m, Engel 22378 (F). NORTH

AUCKLAND PROV.: Waitakere Hills, W of Auckland,

Hatcher 207 — c. per. (F); NE Waitakere Ranges, Swan-

son University Reserve, Tram Valley Road, 95 m, Engel

20447 (F); Auckland, N shore. Kauri Point Centennial

Park, up from W end of Kendalls Bay, Cameron 5562

as T. praenitens (F); Auckland, Kirk 559 (CHR); Kauri

Grove Track, upper Piha Valley, near Auckland, Barr

223 (CHR); Waipoua Forest, Lookout Track between

Lookout Road and Waipoua River Road, above forest

headquarters, ca. 240 m, Engel 21067; ibid., track from

Lookout to forest headquarters, Braggins 94/278B

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

59

(AKU); ibid., track to Te Matua Ngahere, ca. 340 m,

i.n^el 22560 (F); Waipoua State Forest, Allison H763

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

Bridge, 260 m, Engel 21012 (F); Tributary of Manga-

muka River. Maungataniwha Range, SE of Kaitaia on

State Highway 1, 200 m, Engel 20812 (F).

Telaranea paludicola (Hodgs.) Hodgs.

Lepidozia meridiana var. paludicola Hodgs., Trans.

Roy. Soc. New Zealand 83: 61 1. pi. 2, f. 21. 1956.

Telaranea paludicola (Hodgs.) Hodgs., Trans. Roy.

Soc. New Zealand, Bot. 3: 69. 1965; Engel &

Merr., Phytologia 79: 252. 1996 (1995). Type: New

Zealand, North Is., Tararuas, Oriwa Lake Hollow,

bog, 3300 ft., 15 April 1933, Zotov (CHR!; "No.

6619 in Bot. Divn. Herb.").

Plants with a rather spinose appearance, resem-

bling T. praenitens, soft, flexuous yet firm, loose-

ly prostrate, often in dense, compact mats, pale

green to olive-green or at times brownish yellow

to rust-brown, nitid when dry; shoots medium, to

0.9 cm wide, including branches. Branching

somewhat irregularly to regularly 1 -pinnate, the

branches rather short, normally determinate, of

the Frullania type, often flagelliform; branch half-

leaf 2(3-4)-lobed, usually obliquely inserted, nar-

rowly rectangular to linear, the lobes parallel to

slightly diverging; first branch underleaf undivid-

ed and ciliiform or less often 2-3-lobed, inserted

on ventral or ventral-lateral side of branch near

base. Ventral-intercalary branches occasional, of-

ten becoming leading shoots and leafy through-

out. Stems with cortical cells markedly differen-

tiated, the radial walls thin, the outer wall slightly

thickened, in 12-14 rows, those on ventral side of

stem a little smaller; cortical cells in section larger

than the numerous (ca. 65) medullary cells, the

medullary cell walls in ventral half of stem slight-

ly thickened, those in dorsal half of stem distinctly

thicker. Leaves on main shoot rigid, distant to

loosely imbricate, widely spreading, moderately

concave, the lobes incurved, the insertion mod-

erately incubous; leaves 770-980(1260) jxm wide

(from lobe tip to lobe tip) X 700-1050 |xm long,

usually distinctly asymmetric, or less often sub-

symmetric, ± equally 4(5-6)-lobed to ca. 0.4-0.6,

the lobes typically somewhat longer than the disc,

often distinctly falcate and hooked at the tip.

Lobes slender, long attenuate, divergent, the dor-

sal lobes often ± paired, the lobes 3-4 cells wide

for 1-2 tiers at extreme base, biseriate for 2-4(5)

tiers, terminating in a long uniseriate row of (4)5-

7(8) cells (at times more than half the length of

lobe); lobe cells ± firm, thin to moderately thick-

ened, the septa thickened in the corners but not

swollen and projecting from the lobe margins (the

lobes at times somewhat constricted at the septa),

the lobe cells all elongated (to 2.5:1), the basal

cell of uniseriate row (28)34-50 u,m wide X 78-

88 u,m long, the next cell 36-42 u,m wide X 76-

78 \im long, the terminal cell normally about

equal to the penultimate cell in length but some-

times secondarily divided, the terminal cell of un-

iseriate row tapering to a blunt tip; cuticle smooth

or finely striate-papillose at lobe tips. Disc often

distinctly asymmetric, typically longer than broad

and narrowing to the base, 5-7 cells high (from

median sinus base to leaf base), 15-16 cells wide

in distal portion narrowing to 8-1 1 cells wide in

basal portion; disc margins ± straight, entire, or

the marginal cells somewhat bulging. Cells of disc

moderately thin-walled, trigones small or lacking,

median disc cells longitudinally elongate, (38)45-

52 (xm wide X 65-1 10 jxm long, in ± regular,

somewhat obliquely arranged tiers; cuticle

smooth. Underleaves much smaller than leaves,

strongly spreading to subsquarrose, distant, plane,

4(5)-lobed to ca. 0.6, the lobes divergent, cili-

iform, straight to subfalcate and hooked at tip, 2

cells wide at the base, often with an additional

biseriate tier at base of lobe, the uniseriate portion

formed of (3)5-8 short to elongated cells with

septae thickened at the corners; disc symmetri-

cally cuneate, 4-5(6) cells high (median sinus),

the cells in ± regular tiers; disc 9-12 cells wide

in distal portion narrowing to 8 cells wide at base;

margins entire, usually straight. Rhizoid initial

cells small, subquadrate, at bases of lobes or

forming a continuous bistratose, dense band

across the apex of the disc.

Gametangia and sporophytes not seen.

DIFFERENTIATION AND VARIATION — Telaranea

paludicola is a striking plant when well devel-

oped. Although described as a variety of T. mer-

idiana (Hodgson, 1956), T. paludicola has little

to do with that species. It is more likely to be

confused with T. tetrapila but is distinguished pri-

marily by the pronounced leaf asymmetry (Fig.

19: 5, 6, 8), the longitudinally elongated disc'cells

(Fig. 19: 5, 6), and the slender, caudate leaf lobes,

which are often falcate and hooked at the tips

(Fig. 19: 6-8).

Hodgson (1956) referred to the "3-5 rows of

twin cells [biseriate tiers] below the 5 single

ones" of the lobes. In this, it resembles T. praeni-

tens, but lacks the characteristic swollen, project-

ing septa of that species and has larger leaf cells

60

FIELDIANA: BOTANY

(median disc cells (38)45-52 jxm wide vs. 22-38

|j.m in T. praenitens).

DISTRIBUTION AND ECOLOGY — This plant is spo-

radic in distribution in middle and upper eleva-

tions (usually not below 500 m) of both North and

South Islands, New Zealand. As the specific epi-

thet suggests, the plants typically occur in boggy

sites, at times creeping among Sphagnum (for ex-

ample, on the sides of old Sphagnum mounds, or

mixed with Chiloscyphus spiniferus deep in shad-

ed pockets among bryophyte masses). At Omo-

eroa Saddle it occurred in a bog with dwarf/shrub

heath vegetation including Juncas sp., Carex sp.,

and stunted Libocedrus bidwillii surrounded by

mature Dacrydium cupressinum forest. In subal-

pine areas it occurs in sheltered, protected pock-

ets, particularly in moist sites, such as rills in

scrub consisting of Chionochloa, Dracophyllum

and Hebe or of Dracophyllum and Olearia; also

found under cover of tussock blades in mosaic

areas of stunted Olearia colensoi and Chiono-

chloa. Also at the lips of tarns in low alpine veg-

etation (Mt. Robert Skifield).

The species also is present on Stewart Island,

but as is the case with a number of vascular and

nonvascular plants, it occurs at considerably low-

er elevations on this island as compared to coun-

terparts on the South and North Islands (see War-

die, 1991). Telaranea paludicola was found at 5

m (track to Mason's Bay) in a mosaic of stagnant

ponds, Sphagnum bog, open Leptospermum sco-

parium-Dracophyllum heath (to 1-2 m tall), and

dense communities of Gleichenia dicarpa and

Empodisma. At this site the species occurs in ±

protected, moist niches, e.g., on the floor (partic-

ularly in pockets), on vertical, shaded small banks

under Leptospermum or at the margin of bryo-

phyte-covered mounds beneath Leptospermum.

SELECTED SPECIMENS SEEN— NEW ZEALAND.

STEWART ISLAND: Rakiura Natl. Park, Fresh Water

Swamp, track to Mason's Bay, ca. 1 km W of Freshwater

Hut, 5 m, Engel 24405 (F). SOUTH IS. SOUTHLAND

PROV.: Fiordland Natl. Park, Moraine Creek Track, area

N of Moraine Creek, W of Holly ford River, 610m, En-

gel 23217 (F). OTAGO PROV: Ajax Swamp, ca. 1 km

N of Ajax Hill, Catlins River area, ca. 610 m. Child

H5505 (F); Lee Stream, Hindon Road, ca. 305 m, Child

H931 (F); between Waitati and Dunedin, ca. 365 m, Al-

lison H5907 (CHR); E slopes of Mt. Cargill, Dunedin,

ca. 520 m, Allison H5826 as T. praenitens (CHR); Mt.

Cargill, N of Dunedin, ca. 610 m. Child H677 (F); Be-

thune's Gully, Dunedin, ca. 490 m, Allison 4683 (CHR);

Mt. Maungatua, W of Mosgiel, 760 m, Engel 17762 (F);

Mt. Aspiring Natl. Park, below and W of Mt. Shrimpton,

1370-1470 m, Engel 17906 (F). OTAGO/WESTLAND

PROV. BOUNDARY: Haast Pass, ca. 610 m. Child

H4507 (F). WESTLAND PROV.: Westland Natl. Park,

track to Alex Knob just below juncture with track to

Louisa Peak, 1 150-1 170 m, Engel 18998 (F); ibid., Om-

oeroa Saddle, Highway 6, 330 m, Engel 24812 (F); 2

km N of White Horse Creek, Child H5426 (F). WEST-

LAND/CANTERBURY PROV. BOUNDARY: Arthur's

Pass Natl. Park, Bealey Valley Track, ca. 875-900 m,

Engel 22882 (F). NELSON PROV.: Big Totara R., ca.

60 m. Child 5446 (F); Mt. Richmond Forest Park, Red

Hills, track to Maitland Hut, NE of St. Arnaud, ca. 700-

920 m, Engel 21469 (F); Nelson Lakes Natl. Park, E

slope of Robert Ridge in vicinity of Mt. Robert Skifield,

W end of Lake Rotoiti, 1400-1480 m, Engel 22827 (F).

NORTH IS.: WELLINGTON PROV.: Tararua Mts.,

Field Hut, 825 m, Zotov as L. praenitens (CHR); Ton-

gariro Natl. Park, Mt. Ruapehu, off Ohakune Mountain

Road, ca. 890 m, Braggins 92/88A (F); ibid., ca. 7 km

from Ohakune on Ohakune Mt. Road, 900 m, Engel

22708 (F); ibid., Blyth Track, along small stream ca. 0.5

km from Ohakune Mt. Road, ca. 1230 m, Engel 21308

(F); ibid., Taranaki Falls Track, E of Whakapapa Village,

1080 m, Engel 22429 (F). GISBORNE PROV.: Urewera

Natl. Park, Huiarau Range, summit area of Te Rangaak-

apua, 1265-1320 m, Engel 23452 (F); ibid., Waipai

Swamp, Waikareiti Track between track entrance and

Lake Ruapani, N of eastern extremity of Lake Waika-

remoana, 720 m, Engel 20600 (F). TARANAKI PROV:

Pukeiti Bush, near New Plymouth, Hatcher 274 (F).

SOUTH AUCKLAND PROV: Plateau E of Waiotapu

Valley, Rotorua Region, ± 550 m, Allison H5896

(CHR); near Atiamuri, S of Rotorua, ca. 305 m, Allison

H3026 (CHR); Whakarewarewa, Rotorua, Allison

H59I3 (CHR).

Telaranea sect. Ceraceae Engel & Merr., sect,

nov.

Plantae in omnes paries glaucedlnae insignis, folia ±

longitudine inserta, discus foliorum altus marginibus

parallelibus, lobi foliorum plerumque fragiles atque cad-

uci, amphigastrium primum ramorum plerumque distans,

pedicellus antheridialis 2-seriatus.

Type: Lepidozia centipes Tayl. in G. L. & N.,

Syn. Hep. 204. 1845.

Telaranea sect. Glaucolepidozia Schust., Beih. Nova

Hedwigia 118: 212. 2000, nom. nud.

For a discussion of this section and its distin-

guishing characteristics, see p. 238.

Telaranea elegans (Col.) Engel & Merr.

Lepidozia elegans Col., Trans. & Proc. New Zealand

Inst. 21: 65. 1889 (1888). Telaranea elegans (Col.)

Engel & Merr., Phytologia 79: 251. June, 1996

[1995]. Type: New Zealand, Great Barrier Is., Frith

of Thames, 1888, Winkelmann (Colenso a. 1355)

(BM!, WELT!).

Lepidozia tripilosa Steph. in Stephani & Watts, J. &

Proc. Roy. Soc. N. S. W. 48: 116. 1914, syn. nov.

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

61

Type: Australia, New South Wales, Centennial

Glen, Blackheath, 10 Jan. 1911, Watts 1043 (G!).

Plants delicate, flexuous, prostrate in loosely

creeping, thin mats, faintly glaucous and water re-

pellent (particularly shoot tips), yellowish green

to olive-green and ± nitid, becoming brownish

with age; shoots medium, to 6 mm wide, includ-

ing branches. Branching loosely but ± regularly

and often sparingly pinnate, the branches of the

Frullania type, at right angles to the shoot, often

becoming leading shoots, rarely becoming flagel-

liform; branch half-leaf bifid, linear, obliquely to

almost longitudinally inserted; first branch under-

leaf undivided, subulate, inserted on ventral-lat-

eral side of branch near juncture of branch and

main axis. Ventral intercalary branches occasion-

al, both stoloniform and leafy and becoming lead-

ing shoots. Stems with cortical cells thin-walled

but firm, in 12 rows, in section much larger than

the numerous (26-29) medullary cells. Rhizoids

from distal cells of underleaf disc. Leaves on main

shoot spreading at right angles to stem, approxi-

mate to contiguous but scarcely overlapping,

plane, strongly horizontally oriented, the disc in

the same plane as the dorsal surface of the stem

or nearly so, the insertion almost longitudinal, the

lobe tips often broken; leaves 200-285 (Jim wide

X 385-575 jxm long, longer than broad, (3)4-

lobed to 0.4, the lobes strictly parallel with disc

margins, shorter than the disc. Lobes setaceous,

2-3(4) cells wide at extreme base, uniseriate or

biseriate in basal tier, the uniseriate portion 3-4

cells long, the lobe cells successively narrower in

width (the cells scarcely tapered); cells of uni-

seriate portion elongate-rectangular, the walls

straight-sided and not bulging, the basal cell 22-

34 u,m wide X 36-55 |xm long, the next cell of

similar length but narrower, 14-20 u,m wide X

35-46(60) u,m long, the terminal cell slender, 8-

14 u,m X 24-40 u,, rounded at the tip. Disc ±

symmetrically rectangular, parallel-sided, 5-6

cells high (from median sinus base to leaf base),

8 cells wide throughout; margins entire or finely

serrulate by projecting distal ends of the marginal

cells (esp. the dorsal margin); ventral margin ±

straight, the dorsal weakly arched. Cells of disc

firm, in regular longitudinal rows, uniformly and

often distinctly thick-walled, trigones none; me-

dian disc cells isodiametric to short rectangular,

23-36 (im wide, 33-49 fim long; basal row of

disc cells considerably longer and forming an ob-

vious tier; cuticle with a hazy to faintly granular

appearance, rarely glaucous. Underleaves much

smaller than leaves, widely spreading, 4-lobed to

0.75-0.85, the lobes divergent, ciliiform, consist-

ing of a uniseriate row of 2-3 elongated, thin- to

slightly thick-walled cells, inserted on 2 disc cells,

terminating in a slime papilla; disc abbreviated,

2(3) cells high, 8 cells wide. Asexual reproduction

probably by broken tips of leaf lobes.

Gametangia and sporophytes not seen.

DIFFERENTIATION AND VARIATION — This is an in-

teresting species, both in its morphology and its

disjunct geographical distribution. In overall as-

pect, leaf shape, and slender, bristle-like leaf lobes

it resembles a small T. tuberifera (p. 67), but the

lobes in T. elegans are typically shorter, only 3-

4 cells in length (Fig. 20: 2, 3) vs. (4)5-6(7) cells

in T. tuberifera, and the disk cells are smaller, 36-

45 X 42-55 |xm vs. 41-54 X 60-74 u.m in T.

tuberifera. Its most notable features are the evenly

thick-walled cells of the disc and the finely ser-

rulate disc margins (Fig. 20: 1, 2). Telaranea ele-

gans also typically lacks the conspicuous glau-

cous, water-repellent cuticle characteristic of T.

centipes, T. perfragilis and T. tuberifera. The

Queensland specimen cited below (Windolf 689)

is distinctly glaucous and whitish. Typically, how-

ever, the plants are only faintly glaucous, most

evident in the youngest portions of the plant. In

herbaria the plants are yellowish-green to olive-

green in color and somewhat nitid (on the label

of the Waipoua specimen cited below, K. W. Al-

lison notes "glaucous when fresh"). Telaranea

elegans, as used in the key in Schuster (2000, p.

212), is equivalent to T. tuberifera, and is not T.

elegans sensu the type.

Lepidozia elegans Col. was put in the synony-

my of L. centipes by Stephani (1892). They differ

chiefly in the disc margins, which in T. elegans

are entire or finely serrulate by projecting distal

ends of the marginal cells, vs. distinctly crenutete

by bulging marginal cells in T. centipes. In both

T. centipes and T. elegans the leaf disc is rather

low, 5-6 cells high vs. 6-9(10) cells in T. tuber-

FIG. 20. Telaranea elegans (Col.) Engel & Merr. (1-6) and T. perfragilis Engel & Merr. (7-14). 1, 2. Leaves. 3.

Distal portion of leaf. 4. Leaf lobe. 5. Underleaf. 6. Stem, surface view. 7. Sector of main shoot, dorsal view. 8. Leaf

62

FIELDIANA: BOTANY

lobes (cuticle shown in part). 9. Underleaf. 10. Sector of branch showing (top leaf) fragmenting of leaf tip forming

a gemma-like body (at arrow) and (lower leaf) a disc in process of partially breaking away (fracture at arrow). 11.

Half-leaf. 1 2. Branch base (FB = Frullania-type branch; FBU = first branch underleaf), the underleaf of main shoot

shown in part. 13. First branch underleaf shown in fig. 12. 14. Antheridial stalk. (Figs. 1, 4-6 from type of T. elegans,

WELT; 2-3, from Engel 17216, Australia, New South Wales, Morton Natl. Park, Fitzroy Falls; 7-14 from type of T.

perfragilis.)

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

63

if era. The underleaf disc in T. elegans is only 1-

2 cells high, as in T. centipes.

Lepidozia tripilosa Steph. (Stephani & Watts,

1914) of New South Wales is a synonym of T.

elegans. Stephani's illustration (Icones, Lepidozia,

111) shows a leaf with 3 lobes and a disc 6 cells

wide throughout, and probably represents a

branch leaf. The protologue describes the leaves

as 3-lobed. Our examination of the type (G!) re-

veals that the leaves of the main shoot are 4-

lobed.

DISTRIBUTION AND ECOLOGY — Known from a

few stations in the northern sector of North Island,

New Zealand and from New South Wales and

Queensland, Australia. The New Zealand popu-

lations occur in Nothofagus menziesii-Griselinia

littoralis forests or Leptospermum-Weinmannia

silvicola-Dacrydium cupressinum scrub forests or

(Waipoua area) Agathis forests with Weimannia

silvicola. At the Aongatete River site, the species

occurred in a mixed Dacrydium cupressinum and

Beilschmedia tawa forest with Cyathea understo-

ry. Plants occur in shaded, moist, protected nich-

es, often with only marginal light, as, for example,

within pockets on ± vertical stream banks. Plants

for the most part are very loosely creeping, and

form thin, ± pure, feltlike sheets that hardly touch

the substrate. At Waikohatu Stream in Waipoua

Forest the species occurs not only over soil under

overhanging vertical banks well above the stream,

but also, at this hyperhumid site, growing epi-

phytically on fronds of Trichomanes elongatum.

The Waipoua plant collected by Allison formed a

pure colony on very shady bare ground above a

creek in the forest; the niche was "too shady for

other plants" (label data). In New South Wales

the species occurs in shaded niches in forests be-

tween 475 and 640 m. The plant from Fitzroy

Falls occurred over soil among exposed fibrous

roots of a vertical bank; the plant from Tobys

Glen on soil over a boulder near a waterfall. In

Queensland, the plant was collected at 80 m on

the bank of small stream in a wet sclerophyll for-

est.

SPECIMENS SEEN— NEW ZEALAND. NORTH IS-

LAND: SOUTH AUCKLAND PROV.: Kaimai Range,

network of tracks at the end of Wrights Rd. off the Ka-

tikati-Tauranga highway. Swimming Holes Track, 60 m,

Engel 23539 (F); ibid., Aongatete River, upstream from

intersection of North-South Track and river, 425 m. En-

gel 23638 (F); Mt. Te Aroha, ca. 3 km E of Te Aroha,

900-940 m, Engel 22124 (F); Coromandel State Forest

Park, ridge between Webb Creek Track and Billy Goat

Track, 510-540 m, Engel 22338 (F). NORTH AUCK-

LAND PROV.: Waipoua Forest, Waikohatu Stream at

Waikohatu Kauri Bridge, 290 m, Engel 22657, 22662

(F); Waipoua Forest, Allison H3202 as T. centipes

(CHR). AUSTRALIA. NEW SOUTH WALES: Morton

Natl. Park, Fitzroy Falls, 530-640 m, Engel 17216 (F);

Blue Mts. Natl. Park, Tobys Glen, W of Glenbrook, 475

m, Engel 17226 (F); Sydney, [Gore Cove], Jul. 1885,

Whitelegge, com. Pearson (BM). QUEENSLAND: Coo-

roora, 80 m, Windolf 689 as T. centipes (MELU).

I da nine <i centipes (Tayl.) Schust.

Lepidozia centipes Tayl. in G. L. & N., Syn. Hep. 204.

1845. Mastigophora centipes (Tayl.) Trev., Mem.

1st. Lomb. Sci. Lett. III. 4: 416. 1877. Neolepidozia

centipes (Tayl.) Fulf. & J. Tayl., Brittonia 11: 85.

1959. Telaranea centipes (Tayl.) Schust., J. Hattori

Bot. Lab. 26: 256. 1963. Type: Terra van Diemen

(Tasmania), 1824, Spence (FH!).

Plants soft, flexuous, prostrate and loosely

creeping in noncompact mats, glaucous, common-

ly greenish white, dull and distinctly water repel-

lent; shoots medium, to ca. 1 cm wide, including

branches. Branching normally loosely and some-

what irregularly pinnate, the branches of the Frul-

lania type, frequently elongating and remaining

leafy, rarely stoloniform or becoming flagelli-

form; branch half-leaf bifid, linear, obliquely in-

serted and bisecting the angle between the branch

and main axis; first branch underleaf undivided,

ciliate to subulate (rarely asymmetrically bilobed),

inserted on ventral-lateral side of branch near

juncture of branch and main axis, or displaced

toward lateral or ventral-lateral side of stem, often

somewhat below the branch. Ventral intercalary

branches common, both leafy and stoloniform.

Stems with cortical cells distinctly differentiated,

in 12 rows, those on ventral side of stem some-

what smaller, the dorsal cortical cells much larger

than the ca. 25-27 medullary cells, both the cort-

ical and medullary cells evenly and slightly thick

walled. Rhizoids issuing from distal cells of un-

derleaf disc, or basal cell(s) of first branch under-

leaf. Leaves on main shoot rigid, fragile (the lobes

at times partly broken off), widely spreading, at

times nearly at right angles to stem, distant to

loosely imbricate (more densely shingled in plants

growing in drier conditions), plane, strongly hor-

izontally oriented, the disc in the same plane as

the dorsal surface of the stem or nearly so, the

insertion distinctly incubous; leaves 390-525 u,m

wide X 410-575 u,m long, subsymmetric, mostly

4-lobed to ca. 0.35-0.45 (leaves on robust main

shoots at times 5-6(7) lobed, somewhat distant

and wider than high), the lobes ± parallel with

disc margins or only slightly divergent, shorter

64

FIELDIANA: BOTANY

than the disc. Lobes attenuate to subcaudate, 2-4

cells broad at base, often biseriate for 1(2) tiers,

terminating in a uniseriate row of (2)3-4(5) cells

that are distinctly constricted at the septa; lobe

cells thin- to somewhat thick-walled, short to

somewhat elongated, barrel-shaped and bulging

lending the lobes a submoniliform aspect, the bas-

al cell of the uniseriate portion 25-42 jim wide

X 37-59 n,m long (ca. 1.1-1.8:1), the next cell

narrower and shorter, 18-26 u,m wide X 34-49

u,m long (ca. 1.5-2.2:1), the terminal cell variable,

short to somewhat elongate. Disc ± symmetrical-

ly quadrate to subrectangular to distinctly cuneate,

5-6(7) cells high (from median sinus base to leaf

base), 8-15 cells wide in median portion of disc,

the basal tier consisting of 8-12 cells; margins

about equal in length, entire to crenulate due to

the bulging marginal cells, the ventral ± straight,

the dorsal often distinctly arched and contracted

to the insertion (esp. seen in situ). Cells of disc

in regular longitudinal rows, the walls moderately

thickened, trigones minute or absent; median disc

cells large, subquadrate grading to subrectangular,

36-54 (Jim wide, 60-84 jxm long, the basal row

somewhat larger (much longer but only occasion-

ally a little wider) and forming an obvious tier;

cuticle a dense granular and faintly striate coating.

Underleaves much smaller than leaves, widely

spreading, distant, often gently curved dorsally,

4(6) lobed to 0.75-0.85, the lobes divergent, cili-

iform, consisting of a uniseriate row of 2-4 elon-

gated, thin- to slightly thick-walled cells, inserted

on 2(3) disc cells, terminating in a slime papilla;

disc abbreviated, 1-2 cells high, (7)8(10) cells

wide. Asexual reproduction probably by caducous

tips of leaf lobes.

Plants dioecious. Androecia either on short

Frullania-type branches with a few to several cy-

cles of normal vegetative leaves prior to androe-

cial formation or on short, abbreviated, ventral-

intercalary branches lacking normal vegetative

leaves; bracts closely imbricate, strongly dorsally

assurgent, deeply concave, (2)3-4-lobed, each

lobe terminating in a uniseriate row of 2-3 not to

hardly elongated cells; lamina cells irregular in

shape and arrangement, the lamina margins some-

what incurved, irregularly crenulate to denticu-

late, often with stalked or sessile slime papillae,

occasionally with a cilium near the base; bracts

monandrous; antheridia large for bract size, wide

ovoid, the stalk short, 6-7 cells high, biseriate;

bracteolar antheridia absent. Gynoecia strongly

dorsally assurgent, slightly swollen and densely

rhizoidous at base; bracts small for perianth size,

those of innermost series closely ensheathing the

perianth, the tips somewhat spreading, the bracts

bistratose at extreme base but soon thinning to

unistratose and remaining so, concave, orbicular

to suboblate; apices irregularly 4-6 ciliate-lobu-

late, the lobules consisting of a uniseriate row of

(2)3-4 cells, inserted on. a base of 2-4 cells, ter-

minating in a slime papilla; margins irregularly

repand, with a few short cilia and several sessile

slime papillae; bracteoles of innermost series

nearly identical in form and size to bracts. Peri-

anth ca. 0.75 emergent, narrowly obovoid-cylin-

drical to fusiform, terete in basal half, the distal

half obscurely trigonous and with 4-8 plicae, the

sulci shallow to deep, the perianth narrowing to-

ward the mouth; mouth short ciliate, the cilia

formed of 1-3 rather elongate, coarsely striolate-

papillose cells; cells immediately below the

mouth with well defined long striae; perianth 3-

4 stratose in basal portion, the median portion

2(3) stratose.

Seta not examined. Capsule rather short elliptic,

the valves 735 jim long, the wall 24 u.m thick, of

3 layers, the outer layer ± equal to the thickness

of the combined 2 inner layers, or slightly less

thick; outer layer of cells in tiers, rather regularly

short-rectangular, with 2-phase development, the

longitudinal walls with well-defined sheetlike

thickenings and nodule-like thickenings (4-6 per

cell) alternating with walls that are devoid of

thickenings (or with sporadic, local, nonpigment-

ed, nodular swellings), the transverse walls devoid

of thickenings; intermediate layer thinner than

outer or inner layers; innermost layer of cells ±

tiered, irregularly narrowly to broadly rectangular,

with semiannular bands common, rather narrow,

close, usually complete, at times forked and anas-

tomosing to delimit ill-defined, local fenestrae.

Spores not seen. Elaters rigid, nontortuous, 8.6-

10.6 u,m wide, only slightly tapering toward tips,

bispiral to tips, the spirals 4.3 u,m wide.

DIFFERENTIATION AND VARIATION — Examination

of numerous Tasmanian specimens of Telaranea

centipes has shown them to be distinct from the

New Zealand plants referred here to T. tuberifera.

Three species of sect. Ceraceae with a distinctly

glaucous appearance are recognized here: T. cen-

tipes, T. perfragilis and T. tuberifera. True T. cen-

tipes appears to be confined to Australia (Tasman-

ia, Victoria, New South Wales). Particularly di-

agnostic of T. centipes are the short (to 2.2:1),

bulging, bead-like (submoniliform) cells of the

lobes (Fig. 20: 1 1 ). The dorsal margin of the disc

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

65

FIG. 21. Telaranea centipes (Tayl.) Schust. 1. First branch underleaves. 2. Portion of main shoot with branch,

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

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

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

showing cuticular detail. 9. Perianth, cross section through median third. 10. Perianth bearing shoot. 1 1. Median lobe

of leaf. 12. Female bract and (below) bracteole of innermost series. (Figs. 1-7 from Engel 15593, Tasmania, track

from Ferndene to Mt. Dial; 8-12 from Engel 15191, Tasmania, Lillydale Falls.)

66

FIELDIANA: BOTANY

is often crenulate, due also to the bulging margin-

al cells. Compared to T. tuberifera, the leaf disc

in T. centipes is lower, 5-6(7) cells high vs. 6-

9(10) cells high, and the underleaf disc is also

lower, 1-2 cells high vs. 3-4 in T. tuberifera. The

leaves of well-developed leading shoots differ

from those of the branches in being much broader

than high, up to 12 cells wide at the insertion, and

at times 5-6(7)-lobed (Fig. 22: 2, 3), with the dor-

sal margins broadly arched and contracted to the

insertion. In T. tuberifera, the leaves are (3)4-

lobed and tend to be parallel-sided, with the disc

longer than wide (Fig. 23: 5-7).

The rare T. perfragilis (p. 72) of North Island,

New Zealand also has bulging cells of the leaf

margins, but has parallel-sided leaves which are

highly fragmented and broken — typically not only

the lobes but also much of the disc is missing.

The basal cells of the lobes are barrel-like and

bulging as in T. centipes, but the intact lobes taper

to a slender distal portion, with narrower, elon-

gate, nonbulging cells.

Oil-bodies of T. centipes from Australia were

described by Jovet-Ast (1949) as botryoidal, 3-4

per cell, composed of 6(7-9) very large granules.

The specimen, from Sydney, determined by

Hodgson, was probably this species.

DISTRIBUTION AND ECOLOGY — Occurring in Tas-

mania and in Australia (Victoria, New South

Wales). In Tasmania the species occurs sporadi-

cally in lower- to middle-elevation, dry Eucalyp-

tus forests (usually under 400 m). It occurs nearly

exclusively in the drier, eastern half of the island,

and in the wet, southern and central sectors it ap-

parently is rare or lacking west of ca. 146° 30' E

(we have seen only 1 specimen west of that lon-

gitude— Engel 16509 from the Williamsford-

Rosebery area). It extends west only in the north-

ern sector of the island (e.g., Ferndene-Mt. Dial

area). It occurs on soil, particularly of banks and

shaded, moist, cliff faces, particularly where soil

has accumulated. It rarely occurs in wet Notho-

fagus forests or in the subalpine zone. Scott

(1985) stated that the species is common in wet

sclerophyll forests of Victoria. In Victoria (as in

Tasmania) the species can occur rarely in upper

elevations (1450 m on Mt. Buffalo, Engel 17084).

SELECTED SPECIMENS SEEN — TASMANIA: E coast of

Tasman Peninsula, Waterfall Bay, Camp Falls, 150-240

m, Engel 12720 (F); Freycinet Natl. Park, trail to Mt.

Amos, 100-500 m, Norris 30603 (F); Mt. Cameron,

300-350 m, Norris 28484 (F); Lilydale Falls, 2.6 km by

road N of Lilydale, NNE of Launceston, ca. 200 m. En-

gel 15191 — c. per. + 6 (F); Kermandie River, 140 m,

Ratkowski H1546 (HO); Kermandie River at North

Creek, ca. 50 m, Norris 29380 (F); Huon River near

Codeine Creek, ca. 150 m, Norris 30157 (F); Mt. Wel-

lington, Silver Falls pipeline, 580 m, Ratkowski 1545

(HO); E slope of Mt. Wellington, McRobies Gully, ca.

100-300 m. Norris 29291 (F); ibid., headwaters of New

Town Rivulet, ca. 600-800 m, Norris 29337 (F); Mt.

Field Natl. Park, Tyenna River, ca. 200 m, Norris 28825

(F); Styx River Road at intersection with Gordon River

Road, Norris 28858 (F); North East Dundas Tramway

(abandoned), W of Williamsford and SW of Rosebery,

ca. 400 m, Engel 16509 (F); Great Western Tiers, pla-

teau near Lake Salome. W of Walls of Jerusalem, 1050

m, Engel 15991 (F); Leven Gorge, along River Leven,

10 km by road from Nietta, 400 m, Engel 15741 — c. d

(F); above Cethana Dam, ca. 200-300 m, Norris 27348

(F); above Lake Harrington near Forth Falls, ca. 150 m,

Norris 27242 (F); near Devil's Gate Dam, ca. 150 m,

Norris 27219 (F); track from Ferndene to Mt. Dial, SSW

of Penguin, 30-60 m. Engel 15593— c. cj (F). AUS-

TRALIA. VICTORIA: Mt. Buffalo Natl. Park, Dick-

son's Falls, ENE of The Horn (Mt. Buffalo), 1450 m,

Engel 17084 (F); Mackley Creek Road, which leads N

from road between Powelltown and Noojee, just E of

Powelltown, Engel 16832 (F); Great Dividing Range,

Watts River near Fernshaw, NE of Melbourne, Engel

16775 (F); Marysville, Olsens Track, Meagher.689 (F).

NEW SOUTH WALES: Monga, Scott s.n. (MELU

1300).

Telaranea tuberifera Engel & Schust.

Telaranea tuberifera Engel & Schust., Fieldiana, Bot.

n.s. 14: 2. fig. 1. 1983. Holotype: New Zealand,

South Island, Fjordland Natl. Park, Falls Creek,

Upper Hollyford River Valley, along Milford Road,

Schuster 48775 (F!).

Plants soft, flexuous, prostrate, often in inter-

woven, ± thick mats, glaucous, greenish white to

ceraceous, becoming light brown with age, dis-

tinctly water repellent; shoots medium, to 1 cm

wide, including branches. Branching loosely and

somewhat irregularly pinnate, occasionally 2-3

pinnate, the branches of the Frullania type, fre-

quently elongating and remaining leafy, rarely

stoloniform or becoming flagelliform; branch

half-leaf bifid, linear, obliquely inserted; first

branch underleaf undivided, ciliate to subulate, in-

serted on ventral side of branch base, at times

displaced toward the ventral-lateral side of the

stem-branch juncture. Ventral intercalary branch-

es common, leafy or stoloniform and rooting in

the substrate. Stems with cortical cells thin-

walled, with a granular coating like that of the

leaves, in 12-14 rows, those on ventral side of

stem somewhat smaller, in section much larger

than the numerous (ca. 25), evenly and slightly

thick-walled medullary cells. Rhizoids from distal

cells of underleaf disc. Leaves on main shoot rig-

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

67

fffffy.

pipr ...>..A^M$$im

FIG. 22. Telaranea centipes (Tayl.) Schust. 1. Leaf of main shoot. 2, 3. Leaves of main shoot. 4. Plant with 3

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

dorsal view. 8. Antheridium. 9. Androecial branch. (Figs. 1, 4-9 from Engel 15593, Tasmania, track from Ferndene

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

68

FIELDIANA: BOTANY

FIG. 23. Telaranea tuberifera Engel & Schust. 1. Plant, dorsal view. 2. Germinating tuber; note scattered leaf

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

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

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

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

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

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

69

id, fragile (the lobe tips frequently caducous),

spreading ± at right angles to stem, imbricate,

plane, strongly horizontally oriented, the disc in

the same plane as the dorsal surface of the stem

or nearly so, the insertion strongly incubous;

leaves 595-770 u,m wide X 775-840 jim long,

subsymmetric, longer than broad, (3)4-lobed to

0.4-0.5, the lobes strictly parallel with disc mar-

gins, equal to or shorter than the disc. Lobes se-

taceous, typically 2 cells wide at extreme base (or

rarely 4 cells wide, and then biseriate for 1[2]

tiers), terminating in a uniseriate portion of (4)5-

6(7) cells only weakly constricted at the septa;

cells of uniseriate portion thin- to somewhat thick-

walled, elongated, the walls not bulging, the basal

cell 35-42(-50) UJTI wide X 60-92 u,m long (1.8-

2.4:1), the next cell much narrower and of similar

length, 20-30 |xm wide X 72-94 jxm long (3-4.6:

1), the terminal cell very thin-walled and often

collapsed, variable in size, short to somewhat

elongate. Disc ± symmetrically rectangular, often

rather narrowly so, 6-9(10) cells high (from me-

dian sinus base to leaf base), 8-10(1 1) cells wide

in distal portion, 8(1 1) cells wide at base; margins

entire, the ventral ± straight, the dorsal curved

(esp. as seen in situ). Cells of disc in regular lon-

gitudinal rows, the cell walls thin to slightly thick-

ened, trigones minute or absent; median disc cells

large, short rectangular to somewhat elongated,

41-54 u,m wide, 60-74 u,m long; basal row of

disc cells considerably larger (much longer and

generally a little wider) and forming an obvious

tier; cuticle a dense granular and faintly striate

coating. Underleaves much smaller than leaves,

strongly spreading, distant, often gently curved

dorsally, 3-4-lobed to 0.55-0.7, the lobes diver-

gent, ciliiform, consisting solely of a uniseriate

row of (2)3(4) elongated, thin-walled cells, the

lobes terminating in a slime papilla; disc narrowed

toward base, (2)3-4 rows of cells high and 8(9)

cells wide, the basal tier of cells larger. Rhizoid

initials in a band at base of underleaf lobes. Asex-

ual reproduction by elliptic to ovoid tubers at the

tips of stoloniform branches and probably by ca-

ducous leaf lobes.

Plants apparently dioecious. Androecia not

seen. Gynoecia feebly to strongly dorsally assur-

gent, weakly swollen and rhizoidous at base;

bracts small for perianth size, those of innermost

series closely ensheathing the perianth, the tips

somewhat spreading, the bracts concave, broad

ovate, ± regularly 4-6 ciliate-lobulate, the lobules

2(4) cells wide at base, with a uniseriate row of

2(3) cells, terminating in a slime papilla; lamina

composed of ± regularly subrectangular cells, the

margins each with 1 -several weak teeth formed

by the apical or free divergent end of marginal

cells, terminating in a slime papilla, otherwise en-

tire; bracteoles of innermost series similar in form

and size to bracts. Perianth long emergent, fusi-

form, terete in basal and median sectors, the distal

sector obscurely trigonous and with 3(4) plicae,

the perianth narrowing toward the strongly con-

tracted mouth; mouth cells thin- walled, often sin-

uate, partially or wholly laterally free, occasion-

ally with a laterally free uniseriate row of 2 cells,

the mouth thus shortly denticulate-subciliate; peri-

anth 2-3-stratose in lower portion, the median

portion 2-stratose.

Seta with 5-7 rows of outer cells (each with

their free face bulging), surrounding an inner core

of 9-17 much smaller cells. Capsule short elliptic,

910-980 X 476-497 jim, the wall 24 u,m thick,

of 3 layers, the layers of ± equal thickness; outer

layer of cells in tiers, rather regularly short-rect-

angular, with 2-phase development, the longitu-

dinal walls with well-defined sheetlike thicken-

ings and nodule-like to spinose thickenings (4-6

per cell) alternating with walls that are devoid of

thickenings (or with sporadic, local, nonpigment-

ed to pigmented, nodular swellings), the trans-

verse walls devoid of thickenings, or rarely with

a few nodular swellings; innermost layer of cells

± tiered, irregularly narrowly rectangular, with

semiannular bands common, rather narrow, close,

usually complete, at times short and spinose, or

rarely forked.

Spores 13.9-15.8 u,m, exine yellow brown, are-

olate (with a network of sharply defined furcate

ridges that coalesce and delimit areolae). Elaters

rigid, nontortuous, 8.2-10.6 u>m wide, only slight-

ly tapering toward tips, bispiral to tips, the spirals

2.9-4.8 u,m wide.

DIFFERENTIATION AND VARIATION — This is the

common New Zealand glaucous Telaranea, which

has been known in the New Zealand literature as

T. centipes (Hodgson, 1956; Allison & Child,

1975). As treated here, however, T. centipes sejis.

str. is confined to Tasmania, Victoria and New

South Wales. The chief differences between T.

tuberifera and T. centipes are in the shape of the

disc, the disc margins, and the form of the lobes.

The lobes in T. tuberifera are setaceous and whis-

ker-like (Fig. 23: 6, 13), composed of a uniseriate

row of 5-6 cells, typically inserted on a base of

two cells, which are a continuation of the highly

regular longitudinal rows of cells of the disc (Fig.

70

FIELDIANA: BOTANY

23: 5, 7). The cells of the uniseriate row are elon-

gated (to 4.6:1) and subcapillary (Fig. 23: 14).

They are much longer than those of T. centipes,

and are never bulging, nor are the lobes strongly

constricted at the septa (cf. Fig. 21: 11).

The leaf disc of T. tuberifera is parallel-sided,

and the lobes strictly aligned with the disc mar-

gins as in T. tetrapila van roseana (p. 57), but T.

tuberifera is distinctly glaucous, greenish white to

ceraceous, and almost tediously water repellent,

whereas T. tetrapila and its varieties are never so.

As in T. tetrapila, the disc cells of T. tuberifera

are large and prominent, even under the dissecting

microscope, but unlike T. tetrapila, the leaves are

horizontally oriented, with the disc lying in essen-

tially the same plane as the stem. For additional

differences, see comments under T. tetrapila.

Conspicuously glaucous leaves are also seen in

other regional Lepidoziaceae, for example Baz-

zania tayloriana and L. glaucophylla. Interesting-

ly, these glaucous species also have strongly hor-

izontally-oriented leaves, spreading at nearly right

angles with the stem, suggesting a possible adap-

tive significance to this combination of features.

Telaranea tuberifera was described as a new

species by Engel and Schuster (1983), citing the

presence of caducous leaf lobes, and the produc-

tion of tubers at the tips of prostrate, microphyl-

lous intercalary stolons, issuing from older sectors

of the plant (Fig. 23: 2, 10). Caducous, fragment-

ing leaf lobes are even more strongly developed

in another glaucous species, T. perfragilis (p. 72).

We have observed tubers in several additional col-

lections of T. tuberifera. Two other regional Te-

laranea species are now known to produce tubers:

T. clatritexta (Fig. 1: 14; subg. Acrolepidozia) and

T. tasmanica (Fig. 32: 1, 2; sect. Telaranea). Te-

laranea europaea (p. 150) also produces tubers.

Hassel de Menendez (1984) has described similar

tubers in several different genera of Lepidozia-

ceae, as well as in another Telaranea species, T.

blepharostoma (Steph.) Fulf. of southern South

America.

DISTRIBUTION AND ECOLOGY — Endemic to New

Zealand where mostly on Stewart Island and

South Island and infrequent and sporadic on

North Island. The species is for the most part ter-

ricolous in lower to middle to upper elevation for-

ests of, for example, Podocarpus, or of Nothofa-

gus fusca, Dacrydium cupressinum, or of Notho-

fagus menziesii associated with Dracophyllum and

Quintinia serrata or of dense, broadleaf forests of

Coprosma australis, Rhipogonum scandens, Rho-

palostylis sapida and ferns, etc. It typically oc-

cupies very shaded niches in forests, where it may

form pure, at times extensive colonies, often in

sites too shaded for other plants. It is often found

deep in shaded pockets or holes and on steep-

sided banks, as, for example, under the liplike

overhang of the forest edge at the top of the bank

of Waikohatu Stream (Engel 22653). It is notable,

for example, that Hodgson 632 from Mt. Drury,

was collected on the sides of a cave. It may be

subalpine, where it may line deep pockets or deep

crevices in ledges in scrub of, for example, Olear-

ia, Dracophyllum longifolium, Gaultheria and

Cassinia vauvilliersii, etc.

The Butterfield Beach station on Stewart Island

is of interest. Plants formed large sheets over a

thin layer of sandy, loamy soil on bedrock near

the mouth of a sea cave just a few meters above

sea level on an exposed shoreline. The niche like-

ly would be exposed to periodic salt spray (see

Engel & Schuster, 1973).

SELECTED SPECIMENS SEEN— NEW ZEALAND.

STEWART ISLAND: Rakiura Natl. Park, Port Pegasus,

immediately adjacent Belltopper Falls, ca. 10-80 m. En-

gel 24133 (F); ibid., Fresh Water Swamp, track to Ma-

son's Bay, c. 1 km W of Freashwater Hut, ca. 5 m, Engel

24461 (F); Port Pegasus, Schuster, Scott & Taylor as T.

centipes (CHR); Butterfield Beach, 1-2 m above sea lev-

el, Engel 24109 (F). SOUTH ISLAND: OTAGO PROV.:

Fiordland, head of Lake McKerrow, near McKerrow

Hut, Hatcher 1493 (F). WESTLAND PROV: Monkey

Puzzle Gorge, Cascade Road, Martyr River near Martyr

Saddle, 120 m, Engel 23021 (F); Cascade Readjust NE

of Martyr Saddle, N of Jackson River, S of Jackson Bay,

75-1 10 m, Engel 23054 (F); ca. 10 km along Cascade

Rd, 45 m, Engel 24774 (F); Arthur's Pass Natl. Park, N

of Kellys Creek near Hwy 73, N of Otira, 420-475 m,

Engel 18323 (F); ibid., Otira River Gorge near parking

area for track to Mt. Rolleston, 915 m, Fife 5409 (F);

Camp Creek, W of Alexander Range, 310-800 m, Reif

C86B, C124D as T. centipes (F); Paparoa Range, along

Croesus Track, ESE of Barrytown, ridge between Gran-

ite Creek and Pagan Creek watersheds, 250 m, Engel

19307— c. sporo. (F). WESTLAND/CANTERBURY

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

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

18526 (F). NELSON/WESTLAND PROV. BOUND-

ARY: Paparoa Range, S side of Porarari River, up river

from gorge and ca. 500 m WSW of ford on inland track

to Bullock Creek, 10-20 m, Engel 19193 (F). NELSON

PROV: Big Bush State Forest 48, Donald Creek, 480-

600 m, Macmillan 77/95 as T. centipes (CHR). NORTH

ISLAND. WELLINGTON PROV: Tararua Mts., Rua-

mahanga, Zotov as T. centipes (CHR). SOUTH AUCK-

LAND PROV.: Mt. Drury, Tauranga, sea level, Hodgson

632 as T. centipes (CHR); Kaimai-Mamaku Forest Park,

Mt. Te Aroha, S facing slope, 880-890 m, Engel 23858

(F); Coromandel Peninsula, Matthews 42 as T. centipes

(CHR). NORTH AUCKLAND PROV.: Waipoua Forest,

Waikohatu Stream at Waikohatu Kauri Bridge, 290 m,

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

71

.' //v</ -265.? (F); Waipoua Forest, Waikohatu Stream,

Mlison H708 as T. centipes (CHR).

Telaranea perfragilis Engel & Mem, nom. nov.

Telaranea perfragilis Engel & Merr, nom. nov. pro

Telaranea fragilis Engel & Merr., Novon 9: 34 \.f.

2. 1999 non T. fragilis Miz., J. Hattori Bot. Lab.

40: 449. / /. 1976 (Philippines). Holotype: New

Zealand, North Is., North Auckland Prov., NE Wai-

takere Ranges, Swanson University Reserve, Tram

Valley Road, 95 m, Engel 20465 (F); isotype:

(CHR).

Plants delicate, flexuous, prostrate in thin strag-

gling mats or as isolated strands among other

bryophytes, glaucous, whitish to bluish green, dull

and distinctly water repellent; shoots medium, to

ca. 1 cm wide, including branches. Branching

loosely and irregularly 1 -pinnate, the branches of

the Frullania type, rarely becoming flagelliform;

branch half-leaf bifid, linear, obliquely inserted

and bisecting the angle between the branch and

main axis; first branch underleaf undivided, sub-

ulate, inserted on the ventral side of branch at

branch base. Ventral-intercalary branches present,

both leafy and stoloniform. Stems dorsiventrally

flattened, with cortical cells distinctly differenti-

ated, in 9-12 rows, thin walled, those on ventral

side of stem much smaller and similar in size to

medullary cells, the dorsal cortical cells much

larger than medullary cells; medullary cells ca.

16, moderately thick-walled. Rhizoids issuing

from distal tier of underleaf disc cells. Leaves

fragile, typically erose-truncate (the lobes all or

mostly broken off, and often the distal tiers of disc

cells missing), widely spreading, at times nearly

at right angles to stem, distant to loosely imbri-

cate, plane, strongly horizontally oriented, the

disc in the same plane as the dorsal surface of the

stem or nearly so, the insertion distinctly incu-

bous; leaves 260-290 u,m wide, the fragmented

leaf 280-390 u,m long (including basal cell of

lobe), 465-505 u,m long with lobes, leaves sub-

symmetric, 4-lobed to ca. 0.4, the lobes ± parallel

with disc margins or only slightly divergent,

shorter than the disc. Lobes (when present) sub-

caudate, 2 cells broad at base, terminating in a

uniseriate row of 6-7 cells, distinctly constricted

at the septa (esp. in basal portion); lobe cells thin

walled, the basal pair of lobe cells and the basal

cell of uniseriate row barrel-shaped and bulging,

42-50 |xm wide X 50-61 u,m long (1-1.3:1), the

next cell a little narrower but not much longer,

25-31 JUTI wide X 40-54 u,m long (1.3-2.1:1),

the distal cells of uniseriate row slender, ± elon-

gate, the terminal cell variable: short to somewhat

elongate. Disc ± symmetrically quadrate to sub-

rectangular, 5-6(7) cells long (from median sinus

base to leaf base), mostly 8 cells wide throughout;

margins about equal in length, distinctly crenulate

due to bulging marginal cells, ± straight to weak-

ly arched. Cells of disc in regular longitudinal

rows, thin- walled but firm or moderately thick

walled, trigones minute or absent; median disc

cells large, subquadrate, 33-45 u,m wide, 40-52

|xm long, the basal row somewhat longer, forming

an obvious tier; cuticle a finely granular and faint-

ly striate coating. Underleaves much smaller than

leaves, widely spreading, distant, often gently

curved dorsally, 4-lobed to 0.75-0.85, the lobes

widely divergent, ciliiform, consisting of a uni-

seriate row of 3-4 elongated, thin-walled cells,

inserted on 2 disc cells, terminating in a slime

papilla; disc abbreviated, 2 cells high, 8 cells

wide. Asexual reproduction evidently by fragmen-

tation of leaf lobes and disc.

Plants apparently dioecious. Androecia on

short, abbreviated, ventral-intercalary branches

lacking normal vegetative leaves; bracts rather

closely imbricate, strongly dorsally assurgent,

deeply concave, 2-3-lobed, each lobe terminating

in a uniseriate row of (2)3 moderately elongated,

thick-walled cells; lamina margins irregularly mi-

nutely crenulate by few cells with the free margin

bulging, otherwise with a few slime papillae;

bracts monandrous; antheridia large for bract size,

the stalk 8 cells high, biseriate; bracteolar anther-

idia absent. Gynoecia not seen.

DIFFERENTIATION AND VARIATION — Commenting

on the North Auckland specimens cited below,

Hodgson (1956, p. 606) observed that the leaves

"present a curious appearance, in that the apices

are either crenulate with protruding cells of the

discus, or with 1 or 2 segments showing as re-

duced to one roundly quadrate cell, sitting as it

were on 2 terminal cells of the lengthwise rows,"

which aptly describes the leaves of this species.

In shoots freshly mounted in water the leaves of-

ten can be observed breaking apart (Fig. 20: 10,

arrow), and it is likely that detached lobe and disc

cells function as gemmae. The populations cited

by Hodgson ( 1 956) were included under her treat-

ment of Lepidozia centipes.

Telaranea perfragilis resembles T. centipes in

the conspicuously turgid, bulging cells of the disc

margins and lobes, but the leaves are 4-lobed in

T. perfragilis and the disc is typically parallel-

sided and 8 cells wide throughout (Fig. 20: 7). In

72

FIELDIANA: BOTANY

T. centipes the disc is cuneate and up to 15 cells

wide in the distal portion, and the leaves are

sometimes 5-6(7 )-lobed on robust main shoots. In

addition, the basal cells of the uniseriate row in

T. perfragilis are fat and bulging, but the distal

portion of the uniseriate row, when present, is

slenderly tapering, with narrower, more elongate

cells (Fig. 20: 8).

DISTRIBUTION AND ECOLOGY — Known from a

few scattered sites in North and South Auckland

Provinces, New Zealand. The species occurs on

moist, clayey banks or at times over rock in for-

ests. The type occurred on a vertical clayey bank

above a small stream in an old Kunzea forest with

Agathis and (common) Phyllocladus trichoma-

noides.

SPECIMENS SEEN— NEW ZEALAND. NORTH IS-

LAND: SOUTH AUCKLAND PROV.: Coromandel

State Forest Park, ridge just W of summit of Table Mt.,

820 m, Engel 22410 (F). NORTH AUCKLAND PROV.:

Orakei, Auckland, Langridge (Allison 3197) as T. cen-

tipes (CHR); Waipoua Forest, Allison H711 (CHR).

Telaranea sect. Tricholepidozia (Schust.) Engel

& Mem, comb. & stat. nov.

Telaranea subg. Tricholepidozia Schust., J. Hattori

Bot. Lab. 26: 256. 1963. Type: Lepidozia moo-

reana Steph.

Telaranea remotifolia Hodgs.

Telaranea remotifolia Hodgs., Rec. Domin. Mus. 4:

107. 1962. Lepidozia remotifolia Hodgs., Trans.

Roy. Soc. New Zealand 83: 603. pi. 2, f. 14. 1956,

non L remotifolia Horik., J. Sci., Hiroshima Univ.,

Sen B, Div. 2, 2: 202. 1934 (Taiwan). Type: New

Zealand, North Is., Tararuas, Ruamahanga V., 3

Dec. 1933, Zotov 9275 (MPN!); isotype: (CHR!).

Plants soft, appearing etiolated and often rather

flaccid, loosely prostrate and straggling, in pure,

intermingled loose mats or as isolated wisps

among other bryophytes, pale green to grass-

green, the plants when dry glasslike, the stems

sinuous and the leaves shrunken and inconspicu-

ous; plants stenotypic in stature, small, to 1.2 cm

wide, including branches. Branching very irregu-

larly and distantly pinnate, the branches of the

Frullania type, typically long, weak and filiform;

branch half-leaf undivided and subulate or bifid

and narrowly rectangular, usually obliquely in-

serted and bisecting the angle between branch and

main axis; first branch underleaf asymmetrically

bifid, the dorsal lobe leaf-lobe-like, the ventral

lobe shorter and terminating in a slime papilla, the

first branch underleaf inserted on ventral to ven-

tral-lateral side of branch near juncture of branch

and main axis. Ventral-intercalary branches rare,

leafy. Stems often appearing fleshy and large for

plant size, with cortical cells in 10-13 rows, dis-

tinctly differentiated, thin-walled, in section much

larger than the numerous ( 17-25(34]), thin-walled

medullary cells, the medullary strand distinctly

visible in surface view of stem. Rhizoids not seen.

Leaves on main shoot delicate, brittle, obliquely

to widely spreading, conspicuously remote and al-

ternately arranged, weakly convex, the insertion

distinctly incubous to almost longitudinal; leaves

265-385 u,m wide X 440-565 jxm long, asym-

metric, 3-4-lobed to ca. 0.55, the lobes straight to

weakly divergent, about equal to disc or a little

longer. Lobes narrowly attenuate, 2-4 cells wide

at extreme base, terminating in a uniseriate row

of 2-3(4) cells, the dorsal lobe(s) typically some-

what stronger and with 1(2) biseriate tiers be-

tween the uniseriate portion and the extreme base,

the ventral lobe typically weaker; cells of the uni-

seriate portion somewhat elongate, thin-walled,

the septa neither thickened nor swollen, the basal

cell of uniseriate portion 36-50 u,m wide X 60-

79 u-m long, the next cell 24-38 u,m wide X 43-

67 (Jim long, the terminal cell hardly tapering,

rounded at the tip; cuticle of lobe tips finely stri-

ate-papillose, otherwise smooth. Disc somewhat

asymmetric, parallel-sided to weakly cuneate, 2-

3(4) cells high (from median sinus base to leaf

base), 6-1 1 cells wide in distal portion narrowing

to 6-8 cells wide in basal portion; margins entire,

± straight. Cells of disc thin-walled, trigones

lacking, the basal tier of disc cells at times con-

siderably longer (particularly when disc 2 cells

high), the distal tier(s) shorter, ± elongated,

(35)42-53 u,m wide X 72-90 |xm long; cuticle

smooth. Oil-bodies hyaline, coarsely papillose,

the spherules somewhat protruding beyond mem-

brane, the median disc cells with 6 oil-bodies per

cell, globose to subglobose and 5-5.9 u,m in

ili. mi., the cells at sinus base with 5-6 oil-bodies

per cell, elliptic to subfusiform, 5.3-6.4 X 9.5-

12.2 u,m, a few ca. 6.4 X 14.2 UJTI. Underleaves

much smaller than leaves, roughly equal in width

to the medullary strand of the stem, spreading,

distant, slightly convex (ventral view), 3-4-Iobed

to 0.45-0.55, the lobes straight to weakly diver-

gent, consisting of a uniseriate portion of 1-2(3)

somewhat elongated cells, terminating in an elon-

gate slime-papilla, and a base of 2 laterally jux-

taposed cells; disc rectangular (broader than high)

to weakly cuneate, 2 cells high (median sinus), 6-

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

73

8 cells wide in distal and basal portions; margins

entire, straight. Asexual reproduction lacking.

Plants dioecious. Androecia delicate and easily

detached, either on short primary Frullania-lype

branches with a few cycles of normal vegetative

leaves prior to androecial formation, or on short,

abbreviated, ventral-intercalary branches from

leading shoots, the androecium exceptionally long

for genus, with up to 9 pairs of bracts; bracts

closely to rather loosely imbricate, dorsally assur-

gent, deeply concave, bilobed to ca. 0.5, the lobes

acuminate and terminating in a single cell or a

uniseriate row of 2 cells, the tip cell of the lobe

rather elongated (to 84 u,m long, to 3.2:1); dorsal

margin somewhat dilated and with a large tooth/

lobule with its free margin crenulate, at times with

a slime papilla; ventral margin with a conspicuous

tooth; bracts monandrous; antheridial stalk short,

ca. 5 cells high, biseriate; bracteolar antheridia ab-

sent. Gynoecia not seen.

DIFFERENTIATION AND VARIATION — Telaranea

remotifolia has a lax, etiolated aspect when dry.

The pale green, loosely matted stems appear

fleshy and out of proportion to the leaves, which

are distant, shrunken, distinctly glasslike and

strongly convex. The leaf lobes are often sharply

decurved and clawlike. Another of our species

with a similar lax, etiolated appearance is T. niv-

icola (p. 102), but this has a greatly reduced disc,

consisting only of the partially united paired basal

cells of the leaf lobes.

Optimally developed shoots have the aspect of

an etiolated T. tetrapila. Leaves on such shoots

are frequently 4-lobed, and the disc is 3-4 cells

high (Fig. 24: 1, 4, 5), and the lobes are biseriate,

at least in part (Fig. 24: 4, 5). The leaves of some-

what weaker shoots have a disc 2-3 cells high and

are mostly 3-lobed (Fig. 24: 2). In all cases, how-

ever, the ventral leaf lobe is the smallest. Also

distinctive are the underleaves, which are partic-

ularly small and are about equal in width to the

medullary strand of the stem (Fig. 24: 8), which

is clearly visible through the cells of the hyalo-

derm.

Plants with abbreviated leaf discs (2 cells high)

could be confused with T. tetradactyla, which

also has a rather glasslike appearance, but even

when suboptimal, the transverse leaf insertion of

this species will distinguish it from T. remotifolia.

Hodgson (1956) stated that perianths are pre-

sent in the type material. Careful examination of

the type revealed only a single, old, decayed peri-

anth.

DISTRIBUTION AND ECOLOGY — Sporadically oc-

curring on Stewart Island as well as South and

North Islands, New Zealand. The species seems

to have a rather broad ecological amplitude, at

least with respect to elevation. For example, on

Stewart Island (Fern Gully Track) the species was

found at 20-30 m on very old wood of a long-

abandoned bridge in dense shade of a forest dom-

inated by Fuchsia excorticata and Griselinia lit-

toralis canopy with a Blechnum discolor under-

story. Yet it occurs above tree line on stream

banks and submerged boulders (915m, Lake Shir-

ley) as well as over peaty soil in shallow pools at

bog edges (having abundant Sphagnum) in an area

of tussock grassland with a mosaic of scattered

bogs and patches of Dracophyllum, Dacrydium

bidwillii, Hebe and Donatia (825 m, summit of

Mt. Maungatua, Otago Prov.). The species also

occurs in mixed Nothofagus forests (310 m, Blue

River), as well as in dense Nothofagus forests of

the southern portion of South Island, where it is

terrestrial or on rotted logs, occurring at times

with Acromastigum cunninghamii, Telaranea tub-

erifera, Lepidozia kirkii, etc. At the Kaimai Range

station it is found over thin soil on vertical bed-

rock along the Aongatete River within a mixed

Dacrydium cupressinum and Beilschmedia tawa

forest with Cyathea understory (425 m). In the

Waipoua area (290 m) it occurs in Agathis forests

(with Weinmannia silvicola) over soil of stream-

bed rocks or of stream banks within the steep-

sided valley of Waikohatu Stream (290 m).

SELECTED SPECIMENS SEEN— SOLANDER IS.: SE

Peninsula, W side, 100 m, Johnson (CHR). LONG IS.

(Taukiepa Is.), Murders Cove, Fineran 889 (CHR).

NEW ZEALAND. STEWART ISLAND: Rakiura Natl.

Park, Fern Gully Track, 20-30 m, Engel 24055 (F).

SOUTH ISLAND. SOUTHLAND PROV.: Dusky

Sound, Supper Cove, Zotov (CHR); SE of head of Lake

Shirley, 915m, Given 69459 as T. gottscheana (CHR);

Milford Track, Sutherland Falls, Morice et al. (CHR).

OTAGO PROV.: Leith Valley, Allison, Scott & Taylor

(CHR); summit plateau of Mt. Maungatua, W of Mos-

giel, 825 m, Engel 17736 (F); Maungatua, W of Mos-

giel, ca. 915 m, Child H4099 (F); Fiordland, beyond

Lake Howden, Burrell, Scott & Taylor (CHR); trail be-

tween Gunn's Hutt and Hidden Falls, ca. 30 miles "SE

of Lake McKerrow, Hatcher 539 (F); near McKerrow

Hut, head of Lake McKerrow, Hatcher 1431 (F); Mar-

tins Bay, N of McKerrow River, Hatcher 858 (F); Mt.

Aspiring Natl. Park, Blue River near confluence with

Makarora River, NNE of Makarora, 310 m, Engel

18921— c. d (F). WESTLAND PROV: Cascade Road,

just W of Jackson River, ca. 8-12 km SW of confluence

of Jackson and Arawata Rivers, 25-90 m, Engel 22982

(F); Haast Pass Road, Robinson Creek, between Haast

Pass and gates of Haast bridge, ca. 305 m, Child H1838

74

FIELDIANA: BOTANY

FIG. 24. Telaranea remotifolia Hodgs. 1-6. Leaves, cellular detail. 7. Dorsal lobe of leaf. 8. Main shoot (fbu =

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

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

12, 13 from Engel 18921, New Zealand, South Is., Otago Prov., Mt. Aspiring Natl. Park, Blue River; 2-4, 7-9 from

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

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

75

(F): Copland Valley, ca. 90 m, Child H4731 (F); 5 km

N of Runanga, ca. 60 m, Child H5366—C. $ (F). NEL-

SON: Madman's Creek, ca. 2 miles S of Route 6 bridge

over Little Totara River, ca. 30 m, Fife 4645 (F). NEW

ZEALAND. TARANAKI PROV.: Pukeiti Bush, near

New Plymouth, Hatcher 219A (F). SOUTH AUCK-

LAND PROV.: Kaimai Range, Aongatete River, up-

stream from intersection of North-South Track and river,

425 m, Engel 23628 (F). NORTH AUCKLAND PROV.:

Waipoua River, ca. 70 m upstream from State Highway

12 bridge, ca. 95 m, Engel 21061 (F); Waipoua Forest,

Waikohatu Stream at Waikohatu Kauri Bridge, 290 m,

Engel 22652 (F).

Telaranea pulcherrima (Steph.) Schust.

Lepidozia pulcherrima Steph., Spec. Hep. 3: 600.

1909. Telaranea pulcherrima (Steph.) Schust., J.

Hattori Bot. Lab. 26: 256. 1963. Type: New Zea-

land, Okarito, Kirk 588 (G!, CHR!— Bot. Div.

Herb. no. 4844).

Plants subisophyllous, with a soft and woolly

appearance, flexuous, prostrate, superficially ap-

pearing like Trichocolea mollissima, markedly

pale green, distinctly nitid when dry; shoots me-

dium, to 1.3 cm wide (including branches).

Branching regularly and rather densely pinnate, 2-

pinnate in broadest portion of plant, the main axis

often bearing only 2(-4) normal leaves (on each

side) between branches, the branches of the Frul-

lania type, rather short, determinate, rarely be-

coming flagelliform; branch half-leaf 4-5(6)-

lobed, often obliquely inserted, cuneate, the lobes

diverging; first branch underleaf 3-4-lobed, in-

serted on ventral-lateral side of juncture of branch

and main shoot. Ventral-intercalary branches fre-

quent, leafy, often becoming leading shoots.

Stems with cortical cells poorly differentiated, in

24-30 rows of thin-walled cells, the dorsal and

lateral somewhat larger than the exceedingly thin-

walled medullary cells but hardly forming a dif-

ferentiated hyaloderm, the 6-8 or more ventral

cortical cells not or scarcely larger than medullary

cells; medullary cells numerous (ca. 75), thin-

walled, in 8-9 vertical tiers medially. Rhizoids

originating from basal tier of underleaf cells.

Leaves soft yet firmly attached, the disc spread-

ing, the lobes becoming suberect, the leaves

closely imbricate, feebly convex due to slight de-

curving of lobes, the insertion subtransverse to

weakly incubous; leaves 1015-1825 (Jim wide X

1015-1400 (xm long, very slightly asymmetric, ±

equally (7)9-12(13)-lobed to 0.55-0.65, the lobes

± symmetrically spreading, longer than the disc.

Lobes ciliiform, widely spreading, uniseriate

throughout, inserted on a triangular base com-

posed of 2 disc cells, the uniseriate portion 6-

9(10) cells long, the cells ± thin-walled, the septa

somewhat thickened in the corners, moderately to

distinctly constricted, the basal cell 40-66 (xm

wide X (110)120-140 u,m long (1.7-3.3:1), the

longest cells 86-130(160) u,m long and 3.6-

5.5(6.3):! in van pulcherrima, 125-160 u,m long

and 7.4-9.6:1 in var. mooreana, terminal cell

shorter than penultimate cell, rather sharply ta-

pering to a point. Disc ± symmetrically to slightly

asymmetrically short cuneate, 4-5 cells high in

dorsal sector (including intermediate cells and

paired cells at bases of lobes), 3-4 cells high in

ventral sector, 18-20(24) cells broad at midpoint

of disc; disc margins entire, ± straight. Cells of

disc in ± regular tiers, thin-walled but not deli-

cate, with no trigones, the median disc cells large,

38-48 |xm wide X 62-84 jim long; basal row of

disc cells (intermediate cells) narrower; cuticle

smooth. Oil-bodies (fide Schuster, 2000), 9-12 per

cell, finely granular. Underleaves similar to

leaves, the disc spreading to subsquarrose (the

lobes erect), imbricate, plane, symmetrically 6-

10-lobed to 0.8, the lobes symmetrically spread-

ing; disc short cuneate, 4-5(6) cells high (median

sinus), (15)19-23 cells broad in median portion,

the cells in ± regular tiers; margins entire; cuticle

as in leaves. Asexual reproduction lacking.

Dioecious. Androecia either terminal on short,

secondary Frullania-type branches or on short,

abbreviated, ventral-intercalary subspicate

branches; bracts rather closely imbricate, strongly

dorsally assurgent, 3 lobed, the dorsal somewhat

inflexed (and ± oriented toward shoot apex), the

lobes 3-4 cells wide at base, terminating in a uni-

seriate row of 3-4 elongate cells; lamina with dor-

sal margin somewhat incurved to form a weak

pocket, with a few slime papillae or small teeth

(one at times large, lobuliform and inflexed);

bracts monandrous; antheridial stalk biseriate;

bracteolar antheridia absent. Gynoecia strongly

dorsally assurgent; bracts very small for perianth

size, those of innermost series appressed to peri-

anth, strongly concave and ± abruptly involute-

tubular forming a subcullate apex, the bracts

broad elliptic to broad obovate; apices very shal-

lowly and irregularly 3-4-lobulate, the lobules

composed of cells similar to those of lamina, ter-

minating in a short uniseriate cilium of 2-3 elon-

gated cells, often with a terminal slime papilla;

lamina composed of ± regularly subrectangular

cells, the margins curved, the marginal cells

obliquely oriented and sinuous, the free apical

ends diverging and forming irregular denticula-

76

FIELDIANA: BOTANY

tions, with slime papillae frequent; bracteoles of

innermost series similar in form to bracts but

slightly smaller. Perianth large for plant size, te-

rete for most of its length, trigonous and 3-plicate

only near the summit, the perianth gradually ta-

pering toward the mouth; mouth crenate-denticu-

late by the variably projecting, bluntly rounded,

marginal cells.

Capsule (Schuster, 1969, p. 33; 2000, rig. 81)

wall 5-stratose, the outer layer ± equal in thick-

ness to intermediate layers; outer layer of cells in

tiers, with primary cells subdivided by 3 longi-

tudinal walls with sheetlike sinuous thickenings,

the transverse walls devoid of thickenings; inner-

most layer of cells ± tiered, narrowly rectangular,

with semiannular bands common, mostly com-

plete.

Spores areolate. Elaters rigid, not or weakly tor-

tuous, 8.6-10.6 jim wide, only slightly tapering

toward tips, bispiral to tips, the spirals 4.3 jxm

wide.

NOMENCLATURE — Stephani's L. pulcherrima

and L. mooreana were published in the same

work (Stephani, 1909). Telaranea pulcherrima is

the name which is current in the Australasian lit-

erature (Hodgson, 1956; Scott, 1985; Glenny,

1998), and is the most expressive and appropriate

name for this truly beautiful plant.

DISTRIBUTION AND ECOLOGY — Present in New

Zealand (Stewart Is. and Southland and Westland

Prov. of South Island), Tasmania, and Australia

(rare in Victoria). The species is much more com-

mon in Tasmania than in New Zealand, where it

is apparently somewhat frequent only on Stewart

Is. This species forms pure, translucent light yel-

low-green to whitish green mats in very wet sites.

On Stewart Is. it is associated with quiet pools

and small, quiet rills or narrow creek systems

where it occurs within the pools or loosely over

very deep humus at the creek margins. The hu-

midity is always very high in such sites, which

are in the shade of podocarp-hardwood forests of

Dacrydium cupressinum and Weinmannia race-

mosa, with a subcanopy of Dicksonia squarrosa

and woody liane of Ripogonum scandens (Engel

24201) or of stunted and dense Leptospermum

scoparium-Dracophyllum-Weinmannia forest (to

3-4 m tall) (Engel 24357). In the Haast River area

the plant occurs at sea level in open swampy areas

with Sphagnum and scattered Leptospermum, etc.,

in low areas between hummocks (Engel 21771).

Among the most hygrophytic of the Telaranea

species, usually confined to sites (damp rocks near

water, perhaps subject to inundation, and peaty

ground where submersion could occur) where

never subject to drying out. In Tasmania the spe-

cies is confined to forests in the wet western half

of the island (the sole exception being the Mt.

Arthur station).

DIFFERENTIATION AND VARIATION — The pale yel-

lowish green, soft-textured, rather spongy, regu-

larly 2(3)-pinnate, plumose shoots, and the mul-

tiple leaf lobes, radiating fan-like from the disc

(Fig. 25: 1-3) will immediately distinguish this

species. In all these respects, the plants have the

aspect of a small, delicate Trichocolea.

Based chiefly on Stephani's (1909) descrip-

tions, T. pulcherrima and T. mooreana have pre-

viously been maintained as separate species

(Schuster, 1963). The latter has been regarded as

a somewhat weaker plant with 8-lobed rather than

12-lobed leaves, but this distinction is not borne

out by examination of collections. The New Zea-

land and Tasmaman plants are distinguishable,

however, in the form of the lobes, and are treated

here as varieties of the species (see below).

An extreme form of T. pulcherrima is possibly

represented by an illustration in Stephani's Icones

(Lepidozia 122), Tasmania, leg. Weymouth, as

Lepidozia multipila, an unpublished name. The

stem leaf is strongly asymmetric and 16-lobed,

but the plants we examined, labeled T. multipila

in Stephani's herbarium (G), are fairly typical of

the Tasmanian var. mooreana. In our experience,

the common run of lobe numbers in T. pulcher-

rima is 8-12, and never as many as 16. In the

related T. trichocoleoides of Borneo and New

Guinea (p. 194), the strongly asymmetric leaves

are associated with a distinct ventral decurrency

of the leaf, a condition which we have not ob-

served in T. pulcherrima. Stephani's figure of L.

multipila shows a disc which is 6-7 cells high in

the dorsal sector, diminishing to only 2-3 cells

high in the attenuated ventral portion, whereas the

most we have seen in T. pulcherrima is 4-5 cells

high dorsally and 3-4 cells ventrally; his figure

of T. mooreana (ibid., 102) is of a 13-lobed, sym-

metric leaf with a disc 3 cells high throughout.

Key to Varieties

1. Cells in distal half of leaf lobes shorter and

larger in diam., the longest 86-130(160) (Jim

long and 3.6-5.5(6.3):!. New Zealand ....

var. pulcherrima

1 . Cells in distal half of leaf lobe slender and cap-

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

77

11

FIG. 25. Telaranea pulcherrima (Steph.) Schust. Van pulcherrima (1, 2, 6, 7, 9); var. mooreana (Steph.) Engel

& Merr. (3-5, 8, 10, 11). 1. Sector of main shoot, dorsal view (FB = Frullania-type branch; HL = half-leaf). 2, 3.

Leaves. 4-7. Leaf lobes. 8. Terminal cells of leaf lobes. 9. Sector of main shoot, ventral view (FB = Frullania-type

branch: FBU = first branch underleaf). 10. Apex of innermost 9 bract, with (at arrows) 2 of lobules shown in detail.

11. Stem, cross section. (Figs. 1, 6, 9 from Engel 21771, New Zealand, South Is., Westland Prov., ca. 4 km N of

Haast River; 2, 7 from type of Lepidozia pulcherrima; 3, 4, 8 from type of Lepidozia mooreana; 10, 1 1 from Engel

78

FIELDIANA: BOTANY

illary, the longest 125-160 u,m long and 7.4-

9.6:1. Tasmania, Australia . . . var. mooreana

Telaranea pulcherrima (Steph.) Schust. var. pul-

cherrima

Longest cells in distal portion of lobes shorter

than in var. mooreana, 86-130(160) u,m long,

3.6-5.5(6.3):! (see Fig. 25: 6, 7). Stephani's illus-

tration (Icones Lepidozia, 199) gives the dimen-

sions of the lobe cells as 36 X 108 u,m.

NOTE — The protologue (Stephani, 1909) and

Icones (Lepidozia, 199) cite the collector as Kirk.

A specimen from Okarito at CHR determined as

Lepidozia lindenbergii bears the number 588 (as

does the type at G); the plants are labeled "A.

H." but are strikingly similar in appearance to the

type, and are probably parts of the same collection

(see also Hodgson, 1956, p. 614).

DISTRIBUTION — Confined to New Zealand.

SELECTED SPECIMENS SEEN— NEW ZEALAND.

STEWART ISLAND: Rakiura Natl. Park, Port Pegasus,

ca. 100-200 m, Engel 24201 (F); ibid., Mt Rocky Track,

390-475 m, Engel 24357 (F); Pegasus, track to tram

line, Schuster, Scott & Tavlor (CHR); Rakiahua Flat,

Moore (CHR). SOUTH ISLAND. SOUTHLAND

PROV.: Supper Cove, Zotov s.n. (CHR). WESTLAND

PROV.: Ca. 4 km N of Haast River, sea level, Engel

21771 (F).

Telaranea pulcherrima var. mooreana (Steph.)

Engel & Mem, comb. & stat. nov.

Lepidozia mooreana Steph., Spec. Hep. 3: 585. 1909.

Telaranea mooreana (Steph.) Schust., J. Hattori

Bot. Lab. 26: 256. 1963. Type: Tasmania, West

Coast, King River, T. B. Moore s.n. (G!).

Lobe cells in distal half of lobe slender and

capillary, the longest 125-160 u,m long (7.4-9.6:

1) (see Fig. 25: 4, 5).

DISTRIBUTION — Confined to Tasmania and Aus-

tralia (Victoria).

SELECTED SPECIMENS SEEN — TASMANIA: Without

specific loc., Moore as Lepidozia multipila [nom. herb.]

(G); N slope of Mt. Victoria, ca. 800 m, Norris 28324

(F); along tributary of Styx River about 10 miles W of

Maydena, Norris 28933 (F); Tahune Forest Reserve, off

Pine Track, along Huon River, 70 m, Engel 19927 (F);

Florentine River, near Gordon River Road. 1 km W of

junction with Scotts Peak Road, Engel 15084 (F); Clear

Hill Road, 0.9-9.2 km N of Gordon River Road on E

side of Lake Gordon. 350-425 m, Engel 16718 (F);

Gordon River just E of confluence with Serpentine Riv-

er, WNW of Strathgordon, 350 m. Engel 13885— c. 6

(F); Gordon River. Sir John Falls, just up river from

Butler Is., sea level, Engel 14696 (F); Gordon River,

across river and slightly W of Gorge Creek and Pine

Landing, sea level. Engel 14849 (F); Cradle Mtn.-Lake

St. Clair Natl. Park, Pine Valley, between Pine Valley

Hut and Cephissus Falls, NNW of Lake St. Clair, 850

m. Engel 14233 (F); Surprise Valley, above Surprise

River and below Lyell Highway, 2.6 km W of King

William Saddle, 610 m, Engel 19424 (F); Franklin River

at Frenchman's Cap Trail crossing, ca. 400 m, Norris

31195 (F); Kelly Basin Road, between junction with

Franklin River Road and Kelly Basin, above Nora River,

90 m, Engel 14952 (F); Allans Creek, NE slope of Mt.

Darwin, off Kelly Basin Road (Crotty Track). 410 m,

Engel 16575 (F); King River, 11.3 km by road from

Regatta Point and 13.3 km by road from Strahan Har-

bour, sea level, Engel 14938 (F); road between Zeehan

and Trial Harbour, 250 m, Engel 16407— c. per. (F);

Dubbilbarril, Weymouth s.n., 1098, as Lepidozia multip-

ila (G); North East Dundas Tramway (abandoned), W

of Williamsford and SW of Rosebery, ca. 400 m, Engel

16477 (F); Cradle Mtn.-Lake St. Clair Natl. Park. Wein-

dorfers Forest, along track from Waldheim Chalet to

Hounslow Heath Track, 975 m, Engel 14059 (F); S shore

of Lake Lea, ca. 800 m, Norris 27459 (F); southern spur

of St. Valentines Peak, SSW of Burnie. 700-790 m. En-

gel 15680 (F); Emu (Companion) River near Guildford

Road, SSW of Burnie. 500 m, Engel 15833 (F): between

Arthur River and Julius River along Sumac Road, S of

Roger River West, S of Smithton, 100 m, Engel 16114

(F); vicinity of Trowutta Caves (Arch), 4.5 km by road

from intersection of Reynolds Road and Reids Road and

6.9 km from Trowutta Post Office, ca. 180 m, Engel

19761 (F); Newhaven Road. 9.4 km by road E of junc-

tion with Mawbanna Road, S of Detention River mouth,

70 m. Engel 19708 (F). AUSTRALIA. VICTORIA: Ot-

way Ranges, headwaters of Calder River, Turner s.n. (F).

Telaranea sect. Transversae Engel & Merr.

Telaranea sect. Transversae Engel & Merr.. Phyto-

logia 79: 251. 1996 (1995). Type: Lepidozia lin-

denbergii Gott.

Lepidozia sect. Capillares G. L. & N.. Syn. Hep. 211.

1845. "IV. Capillares." Lectotype (nov.): Lepido-

zia lindenbergii Gott. non Telaranea sect. Capil-

lares Engel & Merr., Phytologia 79: 251. 1996

(1995). Type: Lepidozia grossiseta Steph.

Kurzia subg. Leucolepidozia Schust.. J. Hattori Bot.

Lab. 48: 356. 1980. Type: Kurzia auadriseta Grolle.

Note that "sect. Gibbsianae" (Schuster, 2000,

p. 214) was our provisional manuscript name for

El 6407, Tasmania, road between Zeehan and Trial Harbour: 5 from Engel El 9927: Tasmania. Tahune Forest Reserve,

along Huon River.)

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

79

the section later published by Engel and Merrill

(1996) as sect. Capillares. "Lepidozia IV. Capil-

lart-.s" G. L. & N. was first treated as a section

("§ Capillares") by Spruce (1876).

Telaranea lindenbergii (Gott.) Engel & Merr.

Lepidozia lindenbergii Gott. in G. L. & N., Syn. Hep.

213. 1845. Mastigophora lindenbergii (Gott.)

Trev., Mem. 1st. Lomb. Sci. Lett. III. 4: 416. 1877.

Telaranea lindenbergii (Gott.) Engel & Merr., Phy-

tologia 79: 252. June, 1996 [1995]. Type: "Nova

Zeelandia inter L. praenitentem (Hb. Hk.)" (W!,

Lindenberg Hepat. no. 4849).

For synonymy, see under varieties.

Plants delicate, often with a bristly appearance,

soft, flexuous yet firm (straight and ± wiry in var.

mellea), prostrate, pale green to pale yellow-green

(or brownish yellow to rust brown in var. mellea),

nitid when dry; plants small, to 5 mm wide, in-

cluding branches. Branching rather regularly 1-2

pinnate, the branches rather closely spaced, often

at right angles to main axis, often ± equal in

length or the branching pattern ± plumose, with

branches becoming gradually longer toward shoot

base, terete (complanate-foliate in var. complan-

ata), scarcely tapering (long-flagelliform in var.

mellea); branch leaves transversely inserted (in-

cubous in var. complanata and var. papillata);

branch half-leaf 2-lobed; first branch underleaf

undivided and ciliiform, inserted on ventral side

of branch at juncture of branch and main axis.

Ventral-intercalary branches occasional, leafy.

Stems flexuous, often appearing stout for plant

size; cortical cells distinctly differentiated, some-

what thick-walled, in 12-18 rows; cortical cells in

section much larger than the numerous (ca. 40-

67), moderately to distinctly thick-walled medul-

lary cells. Rhizoids often copiously produced

from cells of underleaf disc. Leaves on main

shoot rigid, widely spreading to squarrose, rather

distant on main stems to closely imbricate on the

branches, plane or somewhat handlike due to in-

curving of lobes, typically transversely inserted

and oriented (at times weakly incubous), trans-

verse or weakly incubous on branches; leaves

245-280 |xm wide X 175-225 (Jim long ranging

to 390-440 u,m wide X 235-320 u.m long (4-

lobed) to 525-735 fim wide X 370-460 u,m long

(6-lobed), symmetric to weakly asymmetric, 4-

6(7)-lobed to 0.65-0.8, the lobes divergent, longer

than disc height. Lobes ciliiform, typically uni-

seriate to the base, inserted on a triangular base

composed of 2 disc cells (or sporadically with an

additional biseriate tier between base and uniser-

iate row), the uniseriate portion 5-6(7) cells long

(4-5 cells long in var. papillata), with cells mod-

erately thick-walled and firm, straight-sided and

without or with weakly bulging septa, the basal

cell of the uniseriate portion 20-32 jim wide X

42-65 |xm long (1.8-2:1), the next cell narrower

and a little shorter, 17-24 (Jim wide X 38-48 |im

long (2.6-2.8:1), the terminal cell a little smaller

than the penultimate cell, evenly tapered to the

tip; cuticle smooth. Disc semicircular, 2 cells high

(locally 3, exceptionally 4 in var. mellea) (from

median sinus base to leaf base), including paired

cells at bases of lobes, the basal tier of cells elon-

gate and 8 cells wide in 4-lobed leaves to 12 cells

wide in 6-lobed leaves; margins entire, straight to

somewhat incurved. Underleaves much smaller

than leaves, inconspicuous, 4-5-lobed nearly to

base, the lobes ciliiform, the uniseriate portion

formed of 3-4 somewhat elongated cells, termi-

nating in a slime papilla; disc abbreviated, 2 cells

high, 8 cells wide (in 4-lobed underleaves); mar-

gins entire. Rhizoid initial cells small, subquad-

rate, formed from some or all cells of underleaf

disc. Cells of disc somewhat thinner-walled than

lobe cells but firm, trigones lacking, the largest

cells (in median sector of disc) 17-24 u,m wide

X 40-60 |xm long (basal tier); basal tier of disc

cells often longitudinally elongated; cuticle

smooth.

Plants dioecious. Androecia either terminal on

short to moderately long primary or secondary

Frullania-type branches or on short, abbreviated,

ventral-intercalary, spicate branches; bracts close-

ly imbricate, strongly dorsally assurgent, deeply

concave-subcucullate, 2(3)-lobed to ca. 0.5, the

lobes subcaudate, terminating in a uniseriate row

of 3-4 rather elongated, moderately thick-walled

cells, the terminal cell tapering to summit, to 3:1;

dorsal margin of lamina somewhat dilated and in-

curved, crenulate and with a few slime papillae

or with a 1- to several-celled, often sharp tooth;

bracts monandrous; antheridial stalk short, 6 cells

high, uniseriate; bracteolar antheridia absent. Gy-

noecia not dorsally assurgent and lying in_ the

same plane as main shoot or feebly to strongly

dorsally assurgent, scarcely swollen and often

densely rhizoidous at base; bracts small for peri-

anth size, those of innermost series closely en-

sheathing the perianth, concave, short oblong to

ovate; apices shallowly (3)4-lobulate, the median

sinus a little deeper, the lobules acuminate, ter-

minating in a uniseriate cilium of 2-4 somewhat

elongated cells, often with a terminal slime papil-

80

FIELDIANA: BOTANY

la; lamina composed of ± regularly subrectangu-

lar cells, the margins curved, with a few blunt

teeth formed by the divergent apical or free end

of marginal cells, or margins subentire or with a

few small 1-2-celled teeth; bracteoles of inner-

most series similar in form to bracts, but 0.5-0.75

their size. Perianth long emergent, terete in basal

sector, trigonous above, basically 3-plicate but

with several accessory smaller plicae, the perianth

narrowing toward the contracted mouth; mouth

with 9-13 caudate lobes, the lobes each with a

base composed of 2-4 laterally juxtaposed, elon-

gate cells, the lateral cells of base often with the

distal end protruding to form a "shoulder," the

lobes each terminating in a flexuous or stiff setose

cilium composed of a uniseriate row of 3-5 elon-

gate, rather thick-walled cells, the basal 1-2 cells

13-20 fjtm wide X 96-130 u,m long, (5.7-8.3: 1)

(van lindenbergii) or 14-22 u,m wide X 60-90

u,m long, (3.1-4.6:1) (var. complanata); perianth

2-stratose in basal portion, the cells in section ra-

dially elongate.

Seta with 8 rows of outer cells, surrounding an

inner core of 15-18 much smaller cells. Capsule

short elliptic, 1120-1295 u,m X 553-616 u,, the

outer wall undulate in transverse section, the pri-

mary walls shorter than the thickened secondary

walls, alternatively 29-31 |xm to 38-41 u,m thick,

of 3 layers, the outer layer at its thickest point

equivalent to the combined thickness of the two

inner layers; outer layer of cells (surface view) in

tiers, thin-walled, rather regularly short-rectangu-

lar, with 2-phase development, the longitudinal

walls with rather weakly developed nodule-like

thickenings (4-6 per cell) alternating with walls

that are devoid of thickenings, the transverse

walls also devoid of thickenings; intermediate lay-

er thinner than outer or inner layers; innermost

layer of cells ± tiered, somewhat irregularly nar-

rowly rectangular, with semiannular bands com-

mon, rather narrow, close, usually complete, spo-

radically forked and anastomosing to delimit fe-

nestrae.

Spores 13.4-14.9(15.4) u,, exine pale brown,

with a network of low but sharply defined papillae

and short-vermiculate markings that coalesce and

delimit areolae. Elaters rigid, ± sinuous, 7.7-9.6

u,m wide, only slightly tapering toward tips, bis-

piral to tips, the spirals 3.8-4.8 |xm wide.

DIFFERENTIATION AND VARIATION — This com-

mon and widespread species was treated by

Hodgson (1956) as a synonym of T. tetradactyla.

The name Lepidozia lindenbergii has been used

extensively for the New Zealand plant. All of the

published species in this complex have proved to

be T. lindenbergii, whereas T. tetradactyla s. str.

(p. 91) has no synonyms. For a discussion of the

differences between these two species, and the

typification of T. tetradactyla, see the nomencla-

tural notes under that species.

Telaranea lindenbergii is readily distinguish-

able from other Telaranea species by the distant,

widely spreading, deeply dissected stem leaves

(Fig. 26: 1), with a low disc, for the most part

only 2 cells high (Fig. 26: 2, 3). The leaves of the

main axis are ± transversely inserted and orient-

ed, although the orientation of the branch leaves

is variable. Three varieties of the species are rec-

ognized, which can be distinguished as follows:

Key to Varieties of Telaranea lindenbergii

1. Stems flexuous, often appearing somewhat

fleshy for plant size, the branches more closely

spaced; branches short, of about equal length,

hardly tapering, only exceptionally flagelli-

form, the secondary branches often frequent;

plants pale green 2

2. Branch leaves ± transversely inserted and

oriented. Common throughout New Zea-

land var. lindenbergii

2. Branch leaves incubously inserted and ori-

ented 3

3. Branches terete; lobe cells papillose . . .

var. papillata

3. Branches complanate-foliate; lobe cells

smooth var. complanata

1. Stems slender, straight, rather wiry, the

branching ± distant; branches long, flagel-

liform, the secondary branches typically

few; plants brownish yellow to rust brown.

Primarily southern South Is. . . var. mellea

Telaranea lindenbergii (Gott.) Engel & Merr. var.

complanata Engel & Merr.

Telaranea lindenbergii var. complanata Engel &

Merr., Phytologia 79: 252. June, 1996 [1995]. Ho-

lotype: New Zealand, South Is., Otago Prov.,

Whare Flat, W of Dunedin, 70 m, Engel 17641—

c. sporo. (F).

Branches distinctly complanate-foliate, the

branch leaves imbricate, obliquely spreading, in-

cubously inserted and oriented; leaves of main

axis variable in insertion, mostly subtransverse to

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

81

15

FIG. 26. Telaranea lindenbergii (Gott.) Engel & Merr. (1-8, 11, 13-16) + Telaranea tetradactyla (Hook. f. &

Tayl.) Hodgs. (9-10, 12). 1. Sector of main shoot and primary branch, dorsal view. 2. Three leaves. 3. Two leaves.

4, 5. Leaf lobes. 6. Stem, cross section. 7. Distal sector of perianth. 8. Portion of perianth mouth. 9. Distal half of

perianth. 10. Portion of perianth mouth, the uniseriate row at right shown at higher magnification. 11.9 Bracts and,

in middle, bracteole. 12. 9 Bracts and bracteole. 13. Capsule profile. 14. Capsule wall, outer layer, showing 2-phase

development. 15. Capsule wall, cross section. 16. Seta, cross section. (Figs. 1-2, 4, 7-8, 1 1 from Engel 19300, New

82

FIELDIANA: BOTANY

moderately incubous, sporadically strongly

oblique to almost longitudinal.

DIFFERENTIATION AND VARIATION — Similar in

aspect and branching to var. lindenbergii but dif-

fering chiefly in the flattened, comblike branches

with obliquely spreading, incubously oriented

leaves. This plant is likely to be confused with T.

tetradactyla, which also has strongly complanate-

foliate branches, but differs in the higher disc of

the leaves of the main axis, which is typically 3

or often 4 cells high, vs. 2 or only rarely 3 cells

high in var. complanata. In addition, the stems of

var. complanata are stouter, with more numerous,

thicker-walled medullary cells. Telaranea tetra-

dactyla is also a much softer, laxer plant with

more irregular branching, and is essentially an ex-

treme southern South Island plant.

The branches of var. complanata are complan-

ate-foliate, and the branch leaf insertions extend

to the middle of the adjacent cortical cells along

the midline, thus lacking a conspicuous median

strip. Broad, dorsiventrally flattened branches are

also a feature of T. chaetophylla (p. 1 36), the type

species of the genus, but there the branch leaves

do not extend to the midline, leaving a prominent

2-cell-wide dorsal median strip of quadrate corti-

cal cells (Fig. 42: 2). Several other extraterritorial

species (T. fissifolia, T. kogiana) show this type

of branches.

DISTRIBUTION AND ECOLOGY — Known from

scattered sites at lower elevations on Stewart Is-

land and South and North Islands. The variety oc-

curs in humid niches within forests of various

types and may be on rotten logs and on rich hu-

mus or clayey soil of banks. It also is present on

trunks of Cyathea dealbata and Dicksonia squar-

rosa and in such niches may form large colonies;

in particularly humid niches it is also found on

old, very rotted, prostrate tree fern trunks.

The Lee Bay site on Stewart Island is of inter-

est. Plants occurred over soil of a drainage area

just above the shoreline at the edge of very ex-

posed coastal scrub that includes Brachyglottis ro-

tundifolia, Carpodentus serrata and Coprosma

robusta; the coastal scrub is at the margin of a

podocarp/hardwood forest. The niche likely

would be exposed to periodic salt spray (see En-

gel & Schuster, 1973).

SELECTED SPECIMENS SEEN — NEW ZEALAND.

STEWART ISLAND: Rakiura Nail. Park, Port Pegasus,

ca. 100-200 m, Engel 24192 (F); Lee Bay, head of Rak-

iura Track to Christmas Hut, 5 m, directly above shore-

line, Engel 24105 (F). SOUTH ISLAND. OTAGO

PROV: Leith Valley Bush. Morrison's Creek Gully,

Dunedin, Allison H7713 as T. tetradactyla — c. sporo.

(CHR); Cosy Dell, Dunedin, ca. 150 m. Child HI 509 as

T. tetradactyla — c. sporo. (F). NORTH ISLAND:

SOUTH AUCKLAND PROV: Puaiti Bush, S of Roto-

rua, Allison H25—c. per, H3236—C. per. (CHR); Her-

angi Range, Whareorino Forest, near tributary of Awak-

ino River, W of Leitchs Road, SW of Te Kuiti township,

285 m. Engel 23950— c. 6 (F); Whareorino Forest, start

of track to Leitch's Hut, 280 m. Engel 23752— c. 6 (F);

Mangaotaki Catchment, Waitanguru Scenic Reserve,

track to Waitanguru Falls, 180 m, Engel 23794 (F); Kai-

mai Range, spur ESE of Ngatamahinerua, headwaters of

Poupou Stream, ca. 2 km W of North-South Track, 400-

425 m, Engel 23567— c. per. (F).

Telaranea lindenbergii (Gott.) Engel & Mem var.

papillata Engel & Merr., var. nov.

Foliis ramulorum incubis sed ramuli vix complanatis,

lobis foliorum 4-5 cellulas longis, cellulis loborum pap-

illosis diversa.

Holotype: New Zealand, North Island, South

Auckland Prov., Whareorino Forest, track to

Leitch's Hut, 280 m, Engel 23773 — c. sporo. (F);

isotype: (AK).

DIFFERENTIATION — Known only from the type,

the plants are similar to var. complanata in the

incubous insertion of the branch leaves, but differ

in the terete, not complanate-foliate branches and

the roughened cuticle of the leaf lobes (Fig. 26A:

6). The lobes are rather short, 4-5 cells long (Fig.

26A: 1-5) vs. 5-6(7) cells long in var. linden-

bergii (Fig. 25: 2-5).

DISTRIBUTION AND ECOLOGY — The type and

only known plant occurred on clay banks under a

Blechnum novae -zealandiae overhang, with Wein-

mannia and Dicksonia squarrosa as the dominant

forest canopy and with occasional Leptospermum

scoparium.

Zealand, South Is., Paparoa Range, along Croesus Track; 3, 5 from Allison H79, New Zealand, North Is., South

Auckland Prov., Puaiti Bush; 6 from Allison 3236, same loc. as Allison H79; 9-10. 12 from Child H2353, New

Zealand, South Is., Otago Prov., Ross Creek; 13-15 from Reif C86E; New Zealand, South Is., Westland Prov., Camp

Creek; 16 from type of var. complanata.)

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

83

FIG. 26A. Telaranea lindenbergii var. papillata Engel & Merr. 1 . Portion of shoot showing a leaf and 2 half-

leaves (= hi) each at the base of a Frullania-type branch (- FB), dorsal aspect. 2-4. Leaves (cuticular detail not

shown); all drawn to same scale. 5. Two leaf lobes, cuticular detail not shown. 6. Distal portion of leaf lobes showing

cuticular papillae (X 1065). 7. Underleaf. 8-11. Underleaf lobes. 12. Innermost 9 bracts and, in middle, bracteole.

13. Distal portion of 9 bract. 14-16. Lobes of perianth mouth, all drawn to same scale. (All from type.)

84

FIELDIANA: BOTANY

Telaranea lindenbergii (Gott.) Engel & Merr. var.

lindenbergii

Lepidozia subverticillata Col., Trans. & Proc. New

Zealand Inst. 18: 245. 1886. Lectotype (ntn:): New

Zealand, Waipawa Co.. near Norsewood, 1885,

Colenso a. 1420 (WELT!— c. per. + sporo. + cJ);

isolectotype: (BM! — c. d).

Lepidozia leucocarpa Col., Trans. & Proc. New Zea-

land Inst. 21: 65. 1888 (1889). Lectotype (nm:):

New Zealand, Waipawa Co., near Norsewood,

"Shingle splitter's swamp," 1885, Colenso a. 1320

(WELT! — c. per. + sporo.); isolectotype: (BM! —

c. per. + sporo.).

Lepidozia minutissima Col., Trans. & Proc. New Zea-

land Inst. 21: 66. 1888 (1889). Type: New Zealand,

Waipawa Co., near Dannevirke, 1888, Colenso a.

1319 (BM!, WELT = 0).

Lepidozia occulta Col., Trans. & Proc. New Zealand

Inst. 22: 456. 1889 (1890). Type: New Zealand,

Waipawa Co., South of Dannevirke, 1889, Colenso

a. 1517 (BM!— c. 6*. WELT = 0).

Lepidozia hepaticola Steph., Spec. Hep. 3: 592. 1909.

Type: New Zealand, without specific loc., Goehel

s.n. (G!).

Lepidozia radiata Steph., Spec. Hep. 6: 339. 1922,

syn. nov. Telaranea radiata (Steph.) Schust., J.

Hattori Bot. Lab. 26: 256. 1963. Type: New Zea-

land, sin. loc., "inter Colenso 1095" (G!,

WELT!— c. <J).

Plants pale green; stems flexuous, often ap-

pearing fleshy for plant size; branches rather

closely spaced, of about equal length, hardly ta-

pering, often at right angles to main axis, with

secondary branches often frequent; branches te-

rete, not complanate, with leaves typically trans-

versely inserted and oriented (at times weakly in-

cubous), plane or somewhat handlike due to in-

curving of lobes; leaf disc 2-3(4) cells high; leaf

lobes typically uniseriate to the base, the cells of

uniseriate row weakly constricted at septa or

sometimes with feebly protruding septa.

DIFFERENTIATION AND VARIATION — Telaranea

lindenbergii var. lindenbergii has a distinctive as-

pect due to the ± regular 1-2-pinnate branching

and the rather short primary branches of roughly

equal length. The leaf insertion is typically trans-

verse and the disc is only 2 (locally 3) cells high,

consisting of a basal tier of narrowly elongate

cells, and a distal tier, formed by the bases of the

lobes. The lobe cells are commonly moderately

thick-walled, and the lobes are either straight-sid-

ed or have weakly protruding septa (Fig. 26: 4, 5)

and a firm, subcapillary appearance, suggestive of

T. praenitens. The var. lindenbergii is noteworthy

for the high incidence of fruiting, with perianths

(and often sporophytes) present in a large number

of the specimens examined, and androecia also

commonly present.

Plants of T. lindenhergii often bear a striking

resemblance to T. pallescens. Both species are

very pale to whitish green in color, with trans-

versely inserted, deeply dissected, handlike

leaves. The branches are . often subopposite, at

right angles to the main shoot, with an abundance

of secondary branches. The presence of a hyalo-

derm and absence of Microlepidozia-[ype branch-

ing will immediately mark T. lindenbergii. In ad-

dition, the disc cells of T. pallescens are quite

small, short-rectangular to quadrate, and the disc

is 2-4(5) cells high.

The protologue of Lepidozia radiata Steph.

(Stephani, 1922) describes a plant with 8-lobed

leaves, and a disc only 2 cells high; Stephanies

Icones 177 also depicts a plant with 8-lobed

leaves. Hodgson (1956, p. 614) compared this

species to T. pulcherrirna, but said that "Colen-

so's specimen No. 1095, ex Herb. Stephani, la-

beled T. radiata is L tetradactyla." Hamlin

(1972, p. 284) listed this species as a synonym of

T. tetradactyla, citing "Colenso a. 1095 in

WELT." The latter specimen is typical T. linden-

bergii var. lindenbergii, with leaves 4-6-lobed

and a disc 2 cells high. It is associated with Sym-

phyogyna sp., Schistochila balfouriana and Het-

eroscyphus coalitux, etc.. but no other Telaranea

species are present. We also examined the speci-

men at G labeled "inter Colenso 1095," which is

also var. lindenbergii, a plant known to occasion-

ally produce 7-lobed leaves but never, in our ex-

perience, 8-lobed leaves. Lepidozia radiata is

treated here as a synonym of T. lindenbergii.

DISTRIBUTION AND ECOLOGY — Auckland Is. and

New Zealand (Stewart Island and South and North

Islands). The species was reported as Lepidozia

lindenbergii in the earlier literature from Tasman-

ia (Bastow, 1888; Hooker, 1867; Mitten, 1859);

we have seen no collections of the species from

Tasmania or Australia.

The plant is common throughout New Zealand

in low- to middle-elevation forests (typically be-

low 700 m, and commonly at sea level). It occurs

on a variety of substrates but commonly may be

found on rotted, especially decorticated wood

(rather frequently on crumbly wood on the forest

floor), bryophyte-covered logs, tree fern bases

(e.g., Cyathea), shaded, clayey, mossy banks,

damp leaf litter, or, less commonly corticolous

(e.g., bark of Kunzea), etc. It often forms yellow-

green, feltlike, at times dense mats.

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

85

Si i.txTKD SPECIMENS SEEN— NEW ZEALAND. With-

out specific loc., Cunningham, ex hb. Pearson as Lepi-

ilozia chaetophylla van tenuis (BM). AUCKLAND IS.:

Ranui Cove, Fineran 1967 (CHR). MUTTONBIRD IS.:

Long Island, Boat Harbour Gully, Fineran 732 (CHR).

STEWART ISLAND: Rakiura Natl. Park, Mt Rocky

summit area, 530 m, Engel 24310 (F); between Half

Moon Bay and Deep Bay, Hatcher 1581 (F). SOUTH

ISLAND. SOUTHLAND: Waikaia, Heriot Rd, ca. 610

m, Child 3924a (F); Preservation Inlet, Revolver Cove,

Allan as L. hepaticola (CHR); Doubtful Sound, Simpson

(F); Bligh Sound, Fiordland, Simpson H3221 (CHR);

Head of Milford Sound, Fiordland, Allison H6594 — c.

per. (CHR). OTAGO PROV.: Conical Hill State Forest,

near Tapanui, Rowlings (CHR); Taieri River, south of

Dunedin, near Henley, Allison H5852 (CHR); Swampy

Hill, Dunedin, Child H2644 (F); Morrisons Creek, N of

Dunedin, ca. 150 m, Child H s.n. 146 as T. gottscheana

(F); trail between Gunn's Hutt and Hidden Falls, ca. 30

miles SE of Lake McKerrow, Hatcher 668 — c. per. (F);

N of McKerrow River, Martin's Bay, Hatcher 793 (F).

WESTLAND PROV.: Ship Creek, 14.5 km N of Haast

River, sea level, Engel 21765 (F); Westland Natl. Park,

Gillespies Cook River Road, between Tornado Creek

and Wheland Creek, Engel 6587 (F); ibid., trail to Lake

Wombat, Engel 6692 (F); ibid., Lake Wahapo, Engel

6734 (F); ibid., Franz Josef Glacier Valley, Roberts

Point, SW of Mt. Gunn, ca. 620-670 m, Engel 18095

(F); Lake Kaniere Scenic Reserve, Lake Kaniere Rd,

125 m, Engel 24846— c. $ (F); Arthur's Pass Natl. Park,

N of Kellys Creek near Hwy 73, N of Otira, 420-475

m, Engel 18359 (F); Camp Creek, W of Alexander

Range, 280-440 m, Re if C88F— c. per., C86E—C. 6 +

sporo., C111C (F); Paparoa Range, along Croesus Track,

ESE of Barrytown, between Granite Creek and Fagan

Creek watersheds, 450 m, Engel 19300 — c. per. (F).

OTAGO/WESTLAND PROV. BOUNDARY: Mt. As-

piring Natl. Park, summit area of Haast Pass, 570 m,

Engel 17966 (F). NELSON/WESTLAND PROV.

BOUNDARY: Paparoa Range, S side of Porarari River,

E of Punakaiki, 10 m, Engel 19146. NELSON PROV:

Paparoa Natl. Park, Bullock Creek Road, along Bullock

Creek, NE of Punakaiki, ca. 25 m, Engel 21618 (F);

Paparoa Range, N side of Fox River in vicinity of cave,

NE of Punakaiki, 15 m, Engel 19133 — c. per. (F); Little

Wanganui, W coast, ca. 15 m, Child H414 (F); Able

Tasman Natl. Park, Porters Rock Track, ca. 915 m, Child

H4693 (F). MARLBOROUGH PROV: Kaiuma, Pelorus

Sound, NEN of Havelock, ca. 30 m, Child H4461 (F).

NORTH ISLAND. Without specific loc., "inter Colenso

1095," as Lepidozia radiata. KAPITI IS.: Wilkinson—

c. cJ (CHR). WELLINGTON PROV: Wellington, East-

bourne, Butler H96 (CHR); Wilton Bush, Mackenzie

Burn to Boundry, Mason 36711 as L. lindenbergii — c.

per. (CHR); Wairongomai River, Zotov 7231 — c. per.

(CHR); Tararua Mts., Ruamahanga Valley, Zotov

9273— c. sporo. (CHR); Ohau-iti River, Zotov 7261

(CHR); upper Tiritea River, Zotov 6873 — c. sporo.

(CHR); Mt. Marima, 185 m, Zotov 6943— c. sporo. +

6 (CHR); Tongariro Natl. Park, N side of Lake Roto-

pounamu, S of western side of Lake Taupo, 750 m, En-

gel 22420 (F). GISBORNE PROV: Urewera Natl. Park,

Waikareiti Track between track entrance and Lake Rua-

pani, N of eastern extremity of Lake Waikaremoana,

650-920 m, Engel 20577 (F). TARANAKI PROV: Mt.

Messenger, S of Ahititi, 185 m, Whitehouse 29875 (F);

Pukeiti Bush, near New Plymouth, Hatcher 351 (F);

Waihau Bay and Raukokore River, Bay of Plenty,

O'Malley & Sainsbury (CHR). SOUTH AUCKLAND

PROV: 'Whirinaki Forest Park, Waterfall Loop track,

near Whirinaki River, SSW of Minginui, Engel 20687

(F); Roto-a-kui Bush, E of Taupo, ca. 670-760 m, Al-

lison H3048—C. per. + 6, H3246-*. sporo. (CHR);

Run 95, Kaingaroa S. F. Plantation, Rotorua Region,

H3050—C. per. + 6" (CHR); Mangaiti Bush, near Atia-

muri, S of Rotorua, ca. 460 m, Allison H3049 (CHR);

Puaiti Bush, S of Rotorua, ca. 490 m, Allison H79 — c.

per. + c?, H3237—C. per. (CHR); opposite 7 mile hill

on Rotorua-Atiamuri Road, Allison H3238 (CHR); Ma-

maku Plateau, W of Lake Rotorua on Rt. 5, 400 m,

Engel 20495 (F); Rotorua Region, Allison H3240—C.

per. (F); Ngongotaha Mt., Rotorua, Allison H3243 — c.

sporo. (CHR); southern extent of Herangi Range, plateau

area S of Te Whakapatiki, W of Te Kuiti township, ca.

720-750 m, Engel 25145 (F); Pongakawa Valley, near

coast, Bay of Plenty, Allison H3231—C. per. (CHR);

Coromandel Peninsula, Mercury Bay, Matthews &

Langridge 160 (CHR); Coromandel, Berggren s.n.

(CHR); Coromandel Forest Park, Mt. Moehau, below

summit of "Little Moehau," ca. 800 m, Engel 23648

(F). NORTH AUCKLAND PROV: Anawhata Stream,

near Auckland, Moore (CHR); Hunua, near Auckland,

Moore (CHR); NE Waitakere Ranges, Swanson Univer-

sity Reserve, Tram Valley Road, 95 m, Engel 20417 (F);

Waitakere Hills, W of Auckland, Hatcher 162 — c. per.

(F); Waitakere, ca. 120 m, Child H2223 (F); SE of Goat

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

30 m, Engel 20266 (F); Omaha, just E of causeway to

Omaha Beach, 2 m, Engel 20352 (F); Pakiri Scenic Re-

serve, SSE of Mangawhai Heads, ca. 30 m, Engel 20298

(F); Puketi Forest, Mokau Ridge, Braggins (F); Waipoua

Forest, Allison H762 — c. sporo. (CHR); Waipoua Kauri

Forest, Hatcher 311 — c. per. (F); Waipoua, track to Ma-

huta, ca. 305 m, Child H2142 (F); Waima Forest, Hau-

turu Highpoint Track in vicinity of summit area of Hau-

turu Trig, off Waiotemarama Track, 650-680 m, Engel

22615 (F); Omahuta Forest Kauri Sanctuary, E of Man-

gamuka Bridge, 260 m, Engel 20994 (F); Kiwanis Re-

serve, junction of Okahu Stream and unnamed stream,

ca. 5 km S of Kaitaia, N edge of Herekino Forest area,

ca. 60-80 m, Engel 20941 (F). GREAT BARRIER IS.:

Kirk 169 (CHR). KERMADEC IS. RAOUL: Denham

Bay, Sykes 1201 K (F).

Telaranea lindenbergii (Gott.) Engel & Merr. var.

mellea Engel & Merr.

Telaranea lindenbergii var. mellea Engel & Merr.,

Phytologia 79: 252. June, 1996 [1995]. Holotype:

New Zealand, South Is., Otago Prov., near Herbert,

Allison H5674 (CHR); isotype: (F).

Plants brownish yellow to rust brown; stems

slender, straight, rather wiry; branches rather dis-

tant, often long-flagelliform, the secondary

branches typically few; branches terete, not com-

planate, the leaves transversely oriented, some-

what handlike (at least on branches) due to in-

curving of lobes; leaf disc 2-3(4) cells high; lobes

86

FIELDIANA: BOTANY

not infrequently with a biseriate tier of cells be-

tween the uniseriate row and extreme base, the

cells of the uniseriate row ± thick-walled, weakly

constricted or sometimes with feebly protruding

septa.

DIFFERENTIATION AND VARIATION — When

viewed by transmitted light under the compound

microscope, plants of var. mellea are often suf-

fused with a brownish yellow to rust-brown pig-

ment, rather like that of honey. The plant also has

a distinctive aspect, differing from the typical va-

riety by its straight, wiry stems and distant, long-

flagelliform branches. The same pigment is de-

veloped in T. ferruginea (p. 159) and T. blephar-

ostoma (p. 1 29) of southern South America.

DISTRIBUTION AND ECOLOGY — Known only from

a limited number of collections, primarily from

the southern portion of South Island, and from

Stewart Is. Known also from a single station in

North Island, near Rotorua.

SPECIMENS SEEN— NEW ZEALAND. STEWART IS-

LAND: Kirk 600 as Lepidozia lindenbergii (CHR).

SOUTH ISLAND. OTAGO PROV.: Conical Hill, Rowl-

ings s.n. as Lepidozia hepaticola (CHR); Akatore, coast-

al S of Dunedin, Allison H578I—C. per. (CHR); same

loc. Child 995— c. per. (F); Swampy Hill, Dunedin, ca.

610 m. Child H5218 (F); saddle W of road to Mt. Car-

gill, N of Dunedin, 500 m, Engel 17545 (F). NELSON:

Stony Creek, E of Westport, ca. 150 m, Child H3556

(F). NORTH ISLAND: SOUTH AUCKLAND PROV:

Edge of Urewera Country: Run 95, beyond Rotorua, Al-

lison H3241—C. per. + 6 (CHR).

Telaranea pallescens (Grolle) Engel & Mem,

comb. nov.

Kurzia pallescens Grolle, Rev. Bryol. Liche'nol. 32:

177. 1963 (1964). Microlepidozia quadriseta fo.

pallescens (Grolle) Schust., Nova Hedwigia 15:

456. 1968. Holotype: New Zealand, North Is., NW

Ruahines, Otupae Station, 1948, Druce 8223 (non

vidi).

Kurzia quadriseta Grolle, Rev. Bryol. et Lich6nol. 32:

175. pi. 2. 1963 (1964) non Lepidozia quadriseta

Steph., Spec. Hep. 3: 582. 1909 (= Telaranea

quadriseta (Steph.) Engel & Mem). Microlepidozia

quadriseta (Grolle) Schust., Nova Hedwigia 15:

456. 1968. Holotype: Tasmania, Williamsford, near

Mt. Read, 1 Jan. 1900, Weymouth 5755 as Lepi-

dozia longiscypha (M, non vidi); isotype: (G!).

Plants with the aspect of T. lindenbergii, pros-

trate and densely interwoven, rarely subcaespi-

tose, pale whitish green or at times pure green,

nitid when dry; shoots to 6 mm wide including

branches. Branching irregularly 1 -pinnate to ±

regularly l-2(3)-pinnate, the branches dorsally as-

cending to suberect, the tips sporadically becom-

ing flagelliform, typically with Frullania-type

branches on one side of the main axis and Micro-

lepidozia-iype on the other; branch half-leaf of

Frullania-type branches (l)2-3-lobed nearly to

the base, the first branch urjderleaf 2-3-lobed, in-

serted on the lateral side of branch base or the

juncture of main shoot and branch, aligned with

underleaves of branch. Ventral-intercalary branch-

es occasional; lateral-intercalary branches not

seen. Stem cortical cells moderately thick-walled,

in 12-16 rows, the medullary cells 15-20, some-

what smaller, thin-walled. Leaves stiffly and wide-

ly spreading, transversely inserted, symmetrically

(3)4-6-lobed, 280-700 u,m wide (between tips of

lobes) X 260-350 u,m long, the lobes widely di-

vergent; branch leaves 4-lobed, subcontiguous,

feebly succubously oriented. Lobes uniseriate to

base, inserted on a triangular base composed of

2-3(4) disc cells, the uniseriate portion 5-6(7)

cells long, the cells firm and rather thick-walled,

gradually smaller distally, the cells 2-4.7:1. grad-

ually shorter distally; cuticle smooth. Disc 10-12

cells broad at base, 2-4(5) cells high (including

base of lobes). Cells of disc differentiated in size

and shape from those of the lobes, compact, short-

rectangular to subquadrate, 16-24 X (16)24-

28(40) |xm; cuticle smooth. Oil-bodies (Schuster,

1980, fig. 2: 2) (1)2-3 in cells of disc, 3-5 per

cell in uniseriate row of lobe, weakly to distinctly

botryoidal. Underleaves 4-6-lobed, often with 1

or more lobes abbreviated, terminating in a slime

papilla, the disc 2-3 cells high.

Dioecious. Androecia on inconspicuous, short,

determinate, tightly spicate. ventral -intercalary

branches from main shoot or terminal on rather

short to long primary or (more often) secondary

terminal branches; bracts concave, the disc espe-

cially so, 2-3-lobed to ca. 0.5, the lobes 2 or more

cells wide at base, apiculate to short acuminate,

terminating in a uniseriate row of several cells;

dorsal margin of disc slightly dilated, with a few

crenulations and slime papillae; antheridia 1 per

bract, the stalk uniseriate. Gynoecia on abbrevi-

ated ventral-intercalary branches from main axis;

bracts of innermost series much larger than

leaves, erect and ensheathing the perianth, the

bracts concave, narrowly ovate; apices with 2-4

narrow lobes that terminate in a single cell or a

uniseriate row of 2-3 cells, the lobes composed

of ± regularly rectangular cells, the apical end of

the marginal cells sometimes feebly diverging, the

margins at times with a partially or wholly later-

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

87

ally free cell, the lobe margins thus finely and

sparingly crenulate-spinose dentate; lamina com-

posed of ± regularly short rectangular cells, the

margin bordered by cells of variable shape, most

long and narrow, a few only slightly longer than

wide, the apical or free end of marginal cells at

times divergent and forming a short projection or

an apically oriented tooth, the margin irregularly

and sparingly crenate-denticulate, the teeth rarely

composed of more than 1 cell; bracteole similar

in size and form. Perianth large, fusiform, hyaline,

± terete below, bluntly trigonous only in the distal

portion, the mouth strongly contracted and pluri-

plicate, dentate-ciliate, the teeth free for varying

lengths (at times ca. 0.5 of cell free), at times

consisting of a uniseriate row of 2 cells; perianth

unistratose to base.

Seta with 8 rows of large epidermal cells sur-

rounding an inner core of numerous, leptoder-

mous, smaller cells. Capsule ellipsoid-cylindric,

985-1200 X 490-515 fim, the wall largely 3-stra-

tose, 32-36 u,m thick (locally 4-stratose and 35-

38 u,m thick); outer layer with somewhat irregular

2-phase development, the primary cells with walls

remaining colorless, the secondary walls with

strong, brown, sinuous-nodular thickenings; in-

nermost layer with cells ± irregular in shape, with

semiannular bands remote, well developed and al-

most always complete.

Spores 12-14 (im, exine rather thin, honey-col-

ored, with close vermiculate to granular-vermic-

ulate ridges, which are often furcate but do not

delimit areolae. Elaters feebly twisted, 7.2-9.5

(xm wide, bispiral, the spirals 2.8-3 |xm wide.

DIFFERENTIATION AND VARIATION — Telaranea

pallescens is a rather uncommon plant, and it is

most likely to be mistaken for T. lindenbergii, a

common species in New Zealand. The two species

are remarkably similar in appearance, due to their

± wiry, bristly appearance, regularly pinnate

branching and pale whitish green color. The

leaves are transversely inserted and deeply 4-6-

lobed, the lobes uniseriate to the base and inserted

on 2 disc cells, with a disc 2-4 cells high. Dif-

ferences include the presence of Microlepidozla-

type branches and asymmetrically lobed under-

leaves in T. pallescens. The most conspicuous dif-

ference, however, may be in the basal tier of cells

of the leaf disc, which in T. lindenbergii are thin-

walled and longitudinally elongated (17-24 jxm

wide X 40-60 jim long, Fig. 26: 2, 3) vs. small,

compact and subquadrate in T. pallescens (ca. 24-

28 Jim wide and long, Schuster, 1980, fig. 2: 9-

1 1 ). Telaranea trilobata differs from T. pallescens

in the mostly 3-lobed leaves, and less strongly

differentiated cells of the leaf disc.

DISTRIBUTION AND ECOLOGY — Known from

New Zealand and Tasmania. In New Zealand it

occurs sporadically, over a broad ecological am-

plitude (often in humid sites), from the southern

part of South Island to the northern extremity of

North Island. It occurs in forests of several dif-

ferent types on both South and North Islands. For

example, on South Island the species grows in

seepage areas (with T. remotifolia) among cliffs

and boulders in a Nothofagus menziesii forest

(Haast Pass area), on the floor of open boggy ar-

eas in a rather open Leptospermum scrub forest in

a serpentine area (Red Hills), as well as in humid

niches with the moss, Distichophyllum kraussei,

in forest dominated by Phyllocladus alpinus,

Nothofagus solandrii, Leptospermum scoparium,

Olearia lacunosa and Weinmannia racemosa (Pa-

paroa Range). It is present in several different

types of lowland bogs, and is found, for example,

in Sphagnum-Hypolaena-Dacrydium bogs with

Kurzia helophila, Megalembidium insulanum and

Riccardia sp. (Omoeroa River, Schuster 67-246),

in pakihis along the west coast of South Island,

as well as on wet soil among sedges and Glei-

chenia in an ecotonal area between a Typha

swamp and a Leptospermum swamp (south of

Cape Reinga). On North Island, it grows, for ex-

ample, over soil on stream banks just above water

level in old Kunzea forest with Agathis and Phyl-

locladus trichomanoides (Waitakere Ranges) and

on well-shaded, bryophyte-covered stream banks,

again not far above water level, in kauri forest

with Weimannia silvicola (Waipoua Forest). The

Aongatete River plant (Kaimai Range) occurred

at 425 m on soil over bedrock deep in a pocket-

like drainage channel at the river margin where it

occasionally may be inundated. The river is at the

bottom of a narrow ravine with a dense overhang

of Blechnum and Freycinetia baueriana in a

mixed Dacrydium cupressinum and Beilschmedia

tawa forest with a Cyathea understory. The Mt.

Moehau plants were found at ca. 800 m growing

intermingled in Sphagnum on the bank of a very

moist seepage area in a mosaic of Sphagnum bog

and small communities of shrub heath, including

Dracophyllum recurvum, Lepidothamnus laxifol-

ius, Coprosma foetidissima, Oreobolus pectinatus,

Corokia, and occasional stunted Weinmannia sil-

vicola and stunted Dacrydium cupressinum.

The species extends into the subalpine and al-

pine zones and is present in a variety of humid

88

FIELDIANA: BOTANY

sites, including the sides of rills and watercourses,

soil on tussock bases and protected shaded niches

under shrub cover. In the Arthur's Pass area it is

found in a mosaic of subalpine scrub and alpine

vegetation at the sides of tussock plants that grow

at the edges of pools and, interestingly, T. palles-

cens may form large masses semi submerged in

the pools. In the alpine vegetation of Gertrude

Valley (South Island) it occurs on protected ledges

in seepage areas of cliffs.

SELECTED SPECIMENS SEEN— NEW ZEALAND.

SOUTH ISLAND. SOUTHLAND PROV.: Fiordland

Natl. Park, plateau and slopes below Mt. Burns, E. of

Borland Saddle, S of South Branch of Borland Burn,

W of Monowai, 1225-1320 m, Engel 18572 (F); ibid.,

head of Gertrude Valley, on track to Gertrude Saddle,

NE of Homer Tunnel, 1940-1970 m, Engel 21984.

OTAGO PROV.: Fiordland, N of McKerrow River,

Martin's Bay, near sea level. Hatcher 805 (F); Maun-

gatua, W of Mosgiel, ca. 760 m. Child 4120 (F).

WESTLAND PROV: Ca. 3 km. along Cascade Rd, just

W of Jackson River, SW of confluence of Jackson and

Arawata Rivers, 75 m, Engel 24761 (F); bog N of Om-

oeroa River, S of town of Franz Josef, toward Weheka

Hills, Schuster 67-246— c. <J (F); 10 km S of Grey-

mouth, ca. 150 m, Child H4945 (F); Paparoa Range,

ridge immediately N of Sewell Peak, 890 m, Engel

19032— c. S + sporo. (F); Mt. Aspiring Natl. Park,

Cross Creek, 1 km N of Haast Pass, 510 m, Engel

21877 (F). WESTLAND/CANTERBURY PROV.

BOUNDARY: Arthur's Pass Natl. Park, Dobson Nature

Walk, summit area of Arthur's Pass, 920 m, Engel

22029 (F); ibid., Arthur's Pass, near Temple Basin Ski

Area, Engel 65008—c. per. (F). NELSON PROV: Pa-

paroa Range, N flank of Mt. Euclid, ca. 1-1.5 km E of

Morgan Tarn, 915-1000 m, Fife 6482 as Kurzia quad-

riloba (F). MARLBOROUGH NEAR BOUNDARY

WITH NELSON PROV: Mt. Richmond Forest Park.

Red Hills, track to Maitland Hut, NE of St. Arnaud,

ca. 700-920 m, Engel 21471— c. 8 (F). NORTH IS-

LAND. TARANAKI PROV: Pukeiti Bush, near New

Plymouth, Hatcher 226 (F). SOUTH AUCKLAND

PROV: Kaimai Range, Aongatete River, upstream

from intersection of North-South Track river, 425 m,

Engel 23623 (F); Coromandel State Forest Park, ridge

between Webb Creek Track and Billy Goat Track, 510-

540 m, Engel 22347 (F); ibid., Mt Moehau, below sum-

mit of "Little Moehau," ca. 800 m, Engel 23657 (F).

NORTH AUCKLAND PROV: NE Waitakere Ranges,

Swanson University Reserve, Tram Valley Road, 95 m,

Engel 20487 (F); McElroys Bush Reserve, western side

of Mahurangi Harbour on Cowan Bay Road, 95 m. En-

gel 20382 (F); Waipoua Forest, Waikohatu Stream at

Waikohatu Kauri Bridge, 290 m, Engel 22640 (F);

Omahuta Forest Kauri Sanctuary, E of Mangamuka

Bridge, 260 m, Engel 21013 (F); track to Te Werahi

Beach, directly S of Cape Reinga, ca. 10 m, Engel

20882B (F).

Telaranea trilobata (Schust.) Engel & Mem,

comb. nov.

Kurzia quadriseta var. trilobata Schust., J. Hattori.

Bot. Lab. 48: 363. / 3. 1980. Holotype: New Zea-

land. North Is.. Tongariro Natl. Park. Soda Springs,

head of Mangatepopo Stream, near saddle between

Mts. Ngauruhoe and Tongariro, ca. 4500 ft., Schus-

ter 511 Oa (F).

Plants light green to grass green, with whitish

stems, in densely interwoven mats or caespitose

and erect, nitid when dry; shoots to 5 mm wide

(including branches). Branching irregularly and

rather sparingly 1 -pinnate, with branches of the

Frullania-type on one side, Microlepidozia-lype

on the other, the branches suberect to ascending;

branch half-leaf of Frullania-lype branch undivid-

ed to 2-lobed nearly to base, the first branch un-

derleaf undivided to 2-3-lobed, inserted on the

ventral-lateral side of branch base or the juncture

of main shoot and branch and aligned with un-

derleaves of branch; first appendage (ventral half-

leaf) of Microlepidozia-lype branch bilobed or un-

divided. Ventral-intercalary branches occasional,

leafy or geotropic and stoloniform; lateral-inter-

calary branches occasional to rather freely pro-

duced. Stem with cortical cells moderately thick-

walled, in 9-12 rows, larger than the medullary

cells, which are thin-walled, in 6-7 to 12-14

rows. Leaves strongly spreading, approximate,

transversely inserted, symmetrically 3(4)-lobed,

350-475 p.m (500-550 u,m wide in 4-lobed

leaves) X 350-450 u,m long, the lobes divergent;

branch leaves subcontiguous, weakly succubously

oriented, mostly 3-lobed. Lobes ciliiform, uni-

seriate to the base, inserted on a triangular base

composed of 2-4 cells, sporadically with an ad-

ditional biseriate tier above the base, the uniser-

iate portion 4-5 cells long, the lobe cells elon-

gated (2-3:1), gradually shorter distally, rather

thick-walled; cuticle smooth or finely striate pa-

pillose. Disc 6-8 cells broad at insertion, 2(3)

cells high, including basal cells of lobes. Cells of

disc not much differentiated from lobe cells, ob-

long to rectangular, 22-30 jxm wide X 48-64(72)

u,m long; cuticle smooth or finely striate papillose.

Underleaves 3(4)-lobed, the lobes mostly 4 cells

long, uniseriate or 2-seriate in basal tier, formed

of elongated cells as on leaves, the middle lobe

often abbreviated, terminating in a slime papilla;

disc 6-8 cells wide, 1-2 cells high, the cells small

and ± isodiametric.

Dioecious. Androecia not seen. Gynoecia (only

juvenile known) on short lateral-intercalary

branches.

DIFFERENTIATION AND VARIATION — Telaranea

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

89

irilohata most closely resembles T. pallescens,

differing chiefly in the predominantly 3-lobed

leaves and less sharply differentiated disc cells.

The plants are regularly and suboppositely

branched, with Microlepidozia-type branches on

one side of the shoot and Frullania-type on the

other. The underleaves in T. trilobata are dissim-

ilarly 3-4-lobed, with 1-2 lobes abbreviated and

ending in a slime papilla, and like the leaves, have

a distinct basal tier of cells at the base of the disc.

The cortical cells of the stem are in 9-12 rows,

with 6-7 to 12-14 rows of medullary cells.

DISTRIBUTION AND ECOLOGY — New Zealand,

where known from only a few collections, all in

humid microniches. On Stewart Island the spe-

cies was found near sea level in a mosaic of stag-

nant ponds, Sphagnum bog, open Leptospermum

scoparium-Dracophyllum heath to 1-2 m tall and

dense communities of Gleichenia dicarpa and

Empodisma. At this site the species occurred on

the floor under very dense Gleichenia cover ad-

jacent to a water channel. In the southern sector

of the South Island the species occurred at 680

m on a protected rock face at the margin of a

waterfall within a Nothofagus forest (Southland

Prov., Fiordland Natl. Park, above South Branch

of Borland Burn, Engel 18735). In Mt. Aspiring

Natl. Park the species occurred at 430-480 m on

vertical, dripping, cliff faces under Blechnum

cover in a rich forest dominated by Nothofagus

menziesii (Otago Prov, Blue Valley Track, above

Blue River just N of confluence with Makaroa

River, Engel 21906). In Tongariro Natl. Park

(North Island) it occurs over soil on the lip of a

stream bank within a forest dominated by Noth-

ofagus solandri van cliffortioides (Blyth Track,

ca. 0.5 km from Ohakune Mt. Road, ca. 1230 m,

Engel 21312). The type was growing at 1370 m

on moist volcanic soil in rocky areas along a

stream, in a subalpine open area, associated with

Temnoma quadrifidum and Triandrophyllum sub-

trifidum.

SPECIMENS SEEN— NEW ZEALAND. STEWART IS-

LAND: Rakiura Natl. Park, Fresh Water Swamp, track

to Mason's Bay, ca. 1 km W of Freashwater Hut, 5 m,

Engel 24394— c. 6 (F). SOUTH ISLAND. SOUTH-

LAND PROV.: Fiordland Natl. Park, 11.3 km. by road

WNW of Borland Lodge, above South Branch of Bor-

land Burn, W of Monowai, 680 m, Engel 18735 (F).

OTAGO PROV: Mt. Aspiring Natl. Park, Blue Valley

Track, above Blue River just N of confluence with Mak-

aroa River, 430-480 m, Engel 21906 (F). NORTH IS-

LAND. WELLINGTON PROV: Tongariro Natl. Park.

Blyth Track, along small stream ca. 0.5 km from Ohak-

une Mt. Road, ca. 1230 m, Engel 21312 (F); ibid., Tar-

anaki Falls Track, E of Whakapapa Village, 1240 m,

Engel 22458 (F).

Telaranea fragilifolia (Schust.) Engel & Mem,

comb. nov.

Kurzia fragilifolia Schust., J. Hattori Bot. Lab. 48:

364. / 4. 1980. Holotype: New Zealand, Little Bar-

rier Is., summit track to Mt. Hauturu, SE of summit

of Mt. Herekohu, 1700-1900 ft., Schuster 57896

(F).

Plants delicate, loosely creeping to suberect,

pale green, nitid; shoots to 560 |xm wide, with

leaves. Branching irregularly and remotely

branched, with Frullania- and Microlepidozia-

type branches and lateral-intercalary branches

produced; shoots occasionally becoming micro-

phyllous, flagelliform; lateral-intercalary branches

at times present nearly to the exclusion of other

branch types; ventral-intercalary stolons or flagel-

la rarely present; half-leaf of Frullania-lype.

branch undivided or bilobed, the first branch un-

derleaf undivided, inserted on ventral side of

branch at base; ventral half-leaf of Microlepido-

z/a-branch undivided. Stems slender and delicate,

the cortical cells in 8-9 rows, faintly striolate,

thin-walled, much larger than the medullary cells,

which are in 6-13 rows. Main shoots and branch-

es with 2-3 cortical cells intervening between

successive leaves on either side, the leaves on

prolonged, slender axes more distant, to 9 cells

apart. Leaves remote, rather widely spreading, the

insertion transverse, asymmetrically to ± sym-

metrically 3-lobed almost to the base, the lobe

bases (basal tier) biseriate, connate for ca. 0.5 or

somewhat less, or the basal cells completely unit-

ed, with the tier above partly united, forming a

disc 1 .5 cells high, occasionally with an additional

biseriate tier at base of lobes; branch leaves bifid.

Lobe tips caducous, at times only the basal cells

of the lobes remaining, the lobes subequal to un-

equal, the dorsalmost lobe then somewhat shorter;

lobes gradually tapering from base to apex, ±

straight, biseriate at the base (at times with an

additional biseriate tier), the intact lobes with a

uniseriate row of 4-6 cells; cells of basal tier 24-

31 |xm wide X 48-60 up to 86-95 u,m long, the

basal cell of uniseriate row 28-34 X 72-84 u,m,

the next cell shorter and narrower; terminal cell

(in intact lobes) not distinctly shorter than the

penultimate cell; lobe cells rather firm-walled, the

transverse septa not projecting, the lobe straight-

sided or at times weakly constricted at the septa;

cuticle faintly to distinctly striolate. Underleaves

90

FIELDIANA: BOTANY

trifid, with 1-2 lobes abbreviated, consisting of a

uniseriate row of 3-4 cells arising from a gemi-

nate base, terminating in a slime papilla.

Dioecious. Androecia not seen. Gynoecial

branches (type) very short, all ventral-intercalary

in origin.

DIFFERENTIATION — This species is notable for

its highly fragile, caducous leaf lobes. Species of

Telaranea with fragmenting, caducous lobes are

widely separated taxonomically and geographi-

cally. They include the regional T. perfragilis and

T. tuberifera (sect. Ceraceae), T. mamillosa (sect.

Cancellatae) of New Guinea and Borneo, T. eu-

ropaea (sect. Telaranea) of western Europe, and

T. diacantha (sect. Tenuifoliae) of tropical Amer-

ica.

Telaranea tasmanica also commonly produces

lateral- as well as ventral-intercalary branches, but

differs from T. fragilifolia in the nonfragmenting

lobes and smooth cuticle. The abbreviated disc (to

1.5 cells high) sometimes present in leaves of T.

fragilifolia is reminiscent of the eastern North

American T. longifolia (p. 163; compare Schuster,

1980, fig. 4: 6, T. fragilifolia, and Schuster &

Blomquist, 1955, figs. 10, 11, as T. nematodes,

see also Fig. 50: 3, 4).

The underleaves of T. fragilifolia, T. pallescens

and T. trilobata, T. blepharostoma of southern

South America (Fig. 38: 1, 3, 4), and T. kogiana

(Grolle, 1966, fig. 3f-g) of New Caledonia and

New Guinea are asymmetrically lobed, with one

or more lobes abbreviated (Schuster, 1980, fig. 2,

7), as in many Kurzia species (see Schuster, 1969,

fig. 92: 11, AT. sylvatica). Telaranea quinquespina

has asymmetrically bilobed underleaves with one

lobe abbreviated, the other resembling the lobes

of the leaves (Fig. 34: 1, 3), as do the extra-ter-

ritorial T. anomala and T. monocera (sect. Ten-

uifoliae). Telaranea herzogii (Fig. 35: 2, 3) and

T. tasmanica (Fig. 32: 10-14) differ in having

symmetrically bilobed, caliper-like underleaves.

Schuster (1980) reported the absence of lateral-

intercalary branching in T. fragilifolia, but in our

material (Clarkson 19) lateral-intercalary branch-

es are abundantly produced, to the almost com-

plete exclusion of other branch types (occasional

ventral-intercalary, but only 2 terminal, Frullania-

type branches seen). In the Tasmanian plants (Jar-

men 99/2), only lateral-intercalary branches were

observed, arising from the axil of nearly every

leaf on the main axis.

DISTRIBUTION AND ECOLOGY — The species is

known only from three stations in New Zealand

and one in Tasmania. The type (Little Barrier Is.)

reportedly occurred with material of Lembidium

longifolium Schust. on shaded, permanently moist

rock walls, in transition from Kauri Forest to

Tawa-Ta where Forest. The plant from Cascade

Road, Westland, occurred over soil in a protected

pocket on a large vertical roadside bank at the

margin of a mature Nothofagus menziesii forest at

45 m. The banks are bryophyte-covered (domi-

nated by Isotachis sp.) with Fuchsia excorticata

at the base of the bank. The plant from Hauraki

Plains, North Island, New Zealand, occurred in

ground cover with Goebelobryum unguiculatum

and Riccardia crassa under Sporadanthus ferru-

gineus (Restionaceae)-dominated canopy. Other

associates were Empodisma minus (Restionaceae)

and Leptospermum scoparium (Myrtaceae) (see

de Lange et al., 1999). The Tasmanian collection

occurred with T. herzogii in a Eucalyptus obliqua

wet sclerophyll forest at 130 m.

SPECIMENS SEEN— NEW ZEALAND. SOUTH IS-

LAND. WESTLAND PROV.: Ca. 10 km. along Cascade

Rd, just W of Jackson River, SW of confluence of Jack-

son and Arawata Rivers, 45 m, Engel 24775 (F).

NORTH ISLAND: Hauraki Plains, Kopuatai Peat Dome,

5 m, Clarkson 19 (F). TASMANIA. Manuka Road. W

of Tahune Bridge, 130 m, Jartnan 99/2 (HO).

Telaranea tetradactyla (Hook. f. & Tayl.) Hodgs.

Jungermannia tetradactyla Hook. f. & Tayl., London

J. Bot. 3: 286. 1844 (3: 286 [sic] in errore pro

386). Lepidozia tetradactyla (Hook. f. & Tayl.) G.

L. & N., Syn Hep. 213. 1845. Mastigophora tetra-

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

Sci. Lett. III. 4: 416. 1877. Neolepidozia tetradac-

tyla (Hook. f. & Tayl.) Fulf. & J. Tayl., Brittonia

11: 84. 1959. Telaranea tetradactyla (Hook. f. &

Tayl.) Hodgs., Rec. Domin. Mus. 4: 106. 1962.

Lectotype (nov.): Auckland Is., Nov. 1840, on Jun-

germannia coalita. Hooker (FH!); isolectotypes:

(BM! [2 seen)).

Telaranea longii J. A. Paton, J. Bryol. 17: 289. fig.

1. 1992. syn. nov. Holotype: Scotland, Argyll, near

Dunoon. foot of Glen Massan, Younger Botanic

Garden. 40 m. Long 14231 (E!); isotypes: (E!,

Herb. Paton; NY!).

Plants delicate, flexuous yet firm, procumbent,

in compact, soft, silky, felt-like mats, normally

silvery green to pure green; plants small, to ca. 5

mm wide, including branches. Branching some-

what irregularly l(2)-pinnate, the branches of the

Frullania type, typically of somewhat unequal

length, complanate, the branch leaves incubously

inserted and shingled; branch half-leaf bifid, usu-

ally obliquely inserted, the lobes diverging; first

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

91

branch underleaf undivided and ciliiform, inserted

on ventral side of branch at juncture of branch

and main axis. Ventral-intercalary branches oc-

casional, leafy, sometimes becoming leading

shoots. Stems with cortical cells distinctly differ-

entiated, thin walled, in 12-13 rows; cortical cells

in section much larger than the numerous (ca. 18-

36) medullary cells, the medullary cell walls thin.

Rhizoids often copiously produced from cells of

underleaf disc. Leaves on main shoot obliquely

spreading, contiguous to loosely imbricate on

main stems and closely imbricate on the branches,

plane to moderately convex (due to broad, slight

incurving of lobes), the leaves (of main axis) with

insertion weakly to distinctly incubous, the branch

leaves moderately to distinctly incubously insert-

ed and oriented; leaves 220-320 (when lobes par-

allel) to 390-540 (when lobes divergent) (Jim wide

X 490-630 u,m long, symmetric to weakly asym-

metric, 4(6)-lobed to 0.6-0.75, the lobes ± par-

allel to, more often, divergent, longer than disc

height. Lobes ciliiform, uniseriate throughout and

inserted on a triangular base composed of 2-3

disc cells or, often, with a base of (2)3-4 laterally

juxtaposed cells and 1(2) additional biseriate tiers

between the base and uniseriate row, the uniser-

iate portion (5)6(7) cells long, with cells ± thin-

walled, often weakly to moderately constricted at

the septa, the basal cell of the uniseriate portion

variable: 29-54 u,m wide X 67-98 jim long (1.7-

2.5:1), the next cell narrower and a little shorter,

23-43 p-m wide X 54-86 u,m long (1.8-2.7:1),

the terminal cell a little smaller than the penulti-

mate cell, rounded at the tip; cuticle smooth. Disc

± symmetrically short cuneate, (2)3-4 cells high

(from median sinus base to leaf base) including

paired cells at bases of lobes, 8 cells wide in 4-

lobed leaves, 12 cells wide in 6-lobed leaves;

margins entire, ± straight. Cells of disc thin-

walled but firm, ± equal in thickness to those of

lobe cells, trigones lacking, the largest cells (in

median sector of disc) 28-41 u,m wide X 48-64

jim long; basal tier of disc cells often longitudi-

nally elongate; cuticle smooth. Underleaves much

smaller than leaves, strongly spreading, distant,

plane to slightly incurved, 4-lobed to ca. 0.8, the

lobes somewhat divergent, ciliiform, the uniser-

iate portion formed of 3-4 somewhat elongated

cells, terminating in a slime papilla; disc abbre-

viated, 2 cells high, 8 cells wide; margins entire.

Rhizoid initial cells small, subquadrate, formed

from some distal cells of underleaf disc.

Plants dioecious. Androecia either terminal or

intercalary on short to moderately long primary

or secondary Frullania-type branches or on short,

abbreviated, ventral-intercalary, spicate branches;

bracts closely imbricate, dorsally assurgent, deep-

ly concave-subcucullate, 2-lobed to ca. 0.5-0.6,

the lobes acuminate to subcaudate, terminating in

a uniseriate row of 4-5 somewhat elongated, thin-

walled cells, the terminal cell rounded at the sum-

mit; dorsal margin of lamina somewhat dilated

and incurved, crenulate and with a few slime pa-

pillae or with a 1- to several-celled, sharp tooth;

bracts monandrous; antheridial stalk short, 5 cells

high, uniseriate; bracteolar antheridia absent. Gy-

noecia strongly dorsally assurgent; bracts small

for perianth size, those of innermost series closely

ensheathing the perianth, concave, short ovoid,

shallowly (3)4-lobed, the 2 median lobes normal-

ly larger, the median lobes subcaudate, terminat-

ing in a uniseriate cilium of 3-4 elongated cells,

with a terminal slime papilla, the lateral processes

shorter, the ciliiform process consisting of a single

cell or a uniseriate row of 2 cells; lamina com-

posed of ± regularly subrectangular cells, the

margins curved, with a few crenations and blunt

teeth each formed by the protruding apical end of

a marginal cell which is free for varying lengths,

a single cell at times entirely laterally free, rarely

with a tooth consisting of a uniseriate row of 2

cells, the margins thus crenate-dentate; bracteoles

of innermost series similar in form and size to

bracts. Perianth long emergent, terete in basal sec-

tor, obscurely trigonous and basically 3-plicate

above, the perianth narrowing toward the con-

tracted mouth; mouth fringed with 12 caudate

lobes, the lobes each with a base composed of 3-

5 laterally juxtaposed, moderately elongate cells

and often a lateral, single-celled spinose tooth, the

lobes each terminating in a cilium composed of a

uniseriate row of (2)3-5 elongate, thin-walled

cells, the basal 1-2 cells 11-14 u,m wide x* 42-

76 u.m long, (4.1-6.3:1).

Sporophyte unknown.

FIG. 27. Telaranea tetradactyla (Hook. f. & Tayl.) Hodgs. 1. Sector of main shoot with several terminal branches,

dorsal view. 2. Four- and 5-lobed leaves. 3, 4. Leaves. 5. Leaf lobe. 6. Stem, cross section. 7. Sector of main shoot

(hi = half-leaf: fbu = first branch underleaf; vib = ventral-intercalary branch), ventral view. 8. Underleaves; stem

92

FIELDIANA: BOTANY

cells (= s) stippled in at right. 9. Sector of main shoot, ventral view, showing base of Frullania-type branch (= FB)

and first branch underleaf (= fbu) (fbl = bases of first branch leaves). 10. Leaf pair of primary branch, dorsal view.

11. Antheridium. (Figs. 1, 2, 5-10 from type of T. tetradactyla [FH]; 3, 4 from Engel 17967, New Zealand, South

Is., OtagoXWestland Prov. Boundary, summit of Haast Pass, 570 m, Engel 17967; 1 1 from Child 991, New Zealand,

South Is., Otago Prov., Akatore, 20 mi. SW of Dunedin.)

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

93

DIFFERENTIATION AND VARIATION — This name

has previously been loosely applied to the com-

mon, widespread, and weedy plant here recog-

nized as T. lindenbergii (p. 80). Telaranea tetra-

dactyla s. str. has a more restricted distribution

and is best characterized by the lax, incubously

shingled stem and branch leaves (Fig. 27: 1, giv-

ing the plants a soft, silky aspect. The leaf disc is

typically 3 (and often 4) cells high, vs. 2 (rarely

3) cells high in T. lindenbergii. Plants of this spe-

cies are rarely encountered with perianths, and in

our experience, sporophytes are unknown. This is

in sharp contrast to T. lindenbergii, which char-

acteristically bears perianths (and often sporo-

phytes), as well as androecia.

Both T. tetradactyla and T. lindenbergii have

the perianth mouth long ciliate (Fig. 26: 7, 9);

however, in T. tetradactyla and T. lindenbergii

var. complanata, the cells of the uniseriate cilia

are thin walled and relatively short, at most 90

u,m long (Fig. 26: 10), vs. thick walled and 96-

130 |xm long in T. 1. var. lindenbergii (Fig. 26: 8).

A distinctive expression of T. tetradactyla oc-

curs in the far south, growing in compact, silky

mats, which when dry have a somewhat glasslike

appearance, and shrunken leaves, reminiscent of

T. remotifolia (p. 73). Hodgson (1956, p. 605) re-

ferred to a form subplumulosa, characterized by

"longer, smoother, silky looking plants, mainly

from the south, with a discus of at least 4 cells

high, and each segment based on 2 cells," which

seems to coincide with this expression of T. tetra-

dactyla. Hodgson does not formally describe this

taxon, and we have not seen any of the relevant

specimens cited, mostly from Stewart Is.

Telaranea longii is known from two localities

in Great Britain, in Scotland (Argyll) and in Sur-

rey, in both cases associated with botanic gardens,

"almost certainly introduced with imported hor-

ticultural plants but country of origin unknown"

(Paton, 2000, p. 59). Only 9 plants are known.

At the type locality, the plant grows on shaded

logs and on moist peaty soil "on a woodland

slope near the old fernery." According to Long

(pers. com., cited by Paton, 1992, p. 294) the

plant was originally found on the wall of the old

fernery but does not grow there now. The type

agrees in almost all respects with T. tetradactyla

(compare Paton, 1992, fig. 1; 2000, fig. 11 with

Fig. 26: 9-10, 12 and Fig. 27). Telaranea tetra-

dactyla is said to differ by having "almost trans-

versely inserted lateral leaves" and "shoots al-

ways terete." This description applies to T. lin-

denbergii, which is common and widely distrib-

uted in New Zealand, but not to T. tetradactyla,

which has incubously inserted leaves and distinct-

ly complanate branches.

Schuster (1963, p. 255) referred to the capsule-

wall cross section in T. tetradactyla as being "in

perfect agreement with Telaranea s. str.," which

we take to be a reference to T. lindenbergii, since

that species is common and weedy, and typically

produces sporophytes in abundance.

"Telaranea trilobata" (Schuster, Beih. Nova

Hedwigia 118: 220. / 70. 2000, nom. inval. sin.

descr. lat.) was based on a specimen from New

Zealand (Doubtful Sound, Schuster 52883). Ac-

cording to Schuster (p. 220), this plant is very

close to T. tetradactyla. We have not seen the

specimen.

NOMENCLATURE — Examination of the relevant

specimens in Thomas Taylor's herbarium (FH)

and comparison with the protologue reveal that

Jungermannia tetradactyla included two disparate

elements: (1) an Auckland Is. plant, collected in

November, 1840; and (2) a specimen from New

Zealand, collected by Hooker in 1843, both

mounted on the same sheet. The details of the

gametophyte are derived from the Auckland Is.

collection, as is clear from the description of the

shoots as "foliis distichis patentibus laxiusculis,"

and the comparison with T. centipes, a species

with strongly incubously, almost horizontally ori-

ented leaves. The protologue includes details of

the perianth and androecia, which "occur on New

Zealand specimens" (Hooker & Taylor, 1844, p.

386). The New Zealand element (2) bears abun-

dant perianths and included sporophytes, whereas

the Auckland Is. plants are barren. The New Zea-

land plants, moreover, are typical Telaranea lin-

denbergii, which has terete shoots and transverse-

ly inserted and oriented leaves. We therefore have

no hesitation in designating element ( 1 ), the plant

from Auckland Is., as lectotype of J. tetradactyla.

DISTRIBUTION AND ECOLOGY — Telaranea tetra-

dactyla, as here delimited, is restricted in New

Zealand to the southern sector of South Island

(with only two stations on North Island, at 930-

1 160 m in Urewera Natl. Park), and to Auckland,

Campbell, Snares, Solander, and Antipodes Is.,

and to Stewart Is. (fide Hodgson, 1956, p. 605).

Introduced in Britain, as T. longii.

In New Zealand the species occurs in shaded,

damp, protected niches in lower- to middle-ele-

vation forests as well as in subalpine sites (for

example, it occurs on rock, deep in protected

niches between boulders at 1010-1170 m in a

subalpine stream valley with Chionochloa, Hebe,

94

FIELDIANA: BOTANY

Dracophyllum, etc., near Mt. Burns). Telaranea

tetradactyla is not an "abundant species" as stat-

ed by Schuster (2000, p. 216).

Telaranea tetradactyla has been previously un-

derstood to be an amphi-Paciric species, with the

broadest reported range of any of our Telaranea

species: Auckland Is., New Zealand, Kermadec

Is., Falkland Is. (see Engel, 1990), southern South

America, Juan Fernandez, Falkland Is. (Fulford,

1963a, 1966; Solari, 1987; Schuster, 2000), Fiji

(Hurlimann, 1985; Miller et al., 1983), and Lord

Howe Is. (Miller et al., 1983). At least some of

the records of the species from the Americas are

based on misidentifications of T. plumulosa (see

Engel, 1978, p. 109). Records from Juan Fernan-

dez are probably misdeterminations of T. fernan-

deziensis (see p. 155).

Fulford (1963a, p. 77) stated that T. tetradac-

tyla is "widespread in southeastern Australia

[and] Tasmania . . . ," and made a similar state-

ment with regard to Australia (Fulford, 1966).

Scott & Bradshaw (1986) reported the species for

Victoria and South Australia, and Ratkowsky

(1987) listed the species for Tasmania. We have

not seen specimens of this species from either

Tasmania or Australia.

SELECTED SPECIMENS SEEN — NEW ZEALAND. With-

out specific loc., Petrie s.n., syntype of Lepidozia he-

paticola (G). CAMPBELL ISLAND: South East Har-

bour, Meurk (F); Dent Is., ca. 100 m, Meurk (WELT).

AUCKLAND IS.: Ranui Cove, Fineran 1942 (CHR);

No. 2 Camp, Turbott s.n. (CHR); NW corner of Water-

fall Inlet, Dawbin (4 coll.) (CHR); Ewing Is., Fineran

1347 as T. corticola (CHR); ibid.. Black (WELT).

SNARES IS.: Homing SA299 (CHR); Mutton Bird

Creek, Horning SA72 (CHR, F); ibid., Fineran I2pp as

Telaranea dispar (CHR). SOLANDER ISLAND: SE

peninsula, W side, 100 m, Johnson as T. gottscheana

(CHR); SW Bay, 80 m, Johnson as T. gottscheana

(CHR). BIG SOLANDER ISLAND: Northeast Bay,

Wilson as T. gottscheana (CHR). MUTTON-BIRD IS-

LANDS. LONG ISLAND: Murder's Cove, Fineran 585

as T. corticola (CHR). SOUTH ISLAND. SOUTH-

LAND PROV.: Bluff Hill, Allison H5162 (CHR); Open

Bay Islands, Burrows as Telaranea dispar (CHR);

Fiordland Natl. Park, SW of Mt. Burns, S of South

Branch of Borland Burn, W of Monowai, 1010-1 170 m,

Engel 18671 (F); ibid., Mackinnon Pass, ca. 915 m,

Morice et al. (CHR); ibid.. Moraine Creek Track, area

N of Moraine Creek, W of Hollyford River, 400-530 m,

Engel 23245A (F). OTAGO PROV.: Akatore, 20 mi. SW

of Dunedin, 15 m, Child 991— c. per. (F); Orakanui

Home, Waitati, N of Dunedin, ca. 90 m. Child H324 (F);

Frasers Gully, Dunedin, Child 282 as T. gottscheana,

H283 (F); Town Belt, Dunedin, Allison H3226 (CHR);

same loc., Child 1408 (F); Ross Creek, Dunedin, ca. 185

m, Child H2353—C. per. (F); Mt. Cargill, N of Dunedin,

Child 781 (F); Orakanui Home, Waitati, N of Dunedin,

ca. 90 m. Child 321 (F); Morrison's Creek, N of Dun-

edin, 330 m, Engel 17690 (F); ibid., Leith Valley, Dun-

edin, Allison H6803 (CHR); Allan's Beach, Otago Pen-

insula, Allison H6846 (CHR); Bethune's Gully, Dune-

din, Allison H6520 (CHR). OTAGOAVESTLAND

PROV. BOUNDARY: Mt. Aspiring Natl. Park, summit

of Haast Pass, 570 m, Engel 17967 (F). WESTLAND

PROV: Mt. Aspiring Natl. Park, Roaring Billy Nature

area, along Haast River on road between Haast and

Haast Pass, 70 m, Engel 21859 (F). CANTERBURY

PROV: Rock wood, 610m, Beckett 449, syntype of Lep-

idozia hepaticola (G). NORTH ISLAND. NEAR GIS-

BORNE/SOUTH AUCKLAND PROV. BOUNDARY:

Urewera Natl. Park, crest trail from highway 38 towards

Whakataka summit, N of northern extremity of Lake

Waikaremoana. 930-1030 m, Engel 20626 (F); ibid.,

track to Whakataka Hut from Lake Waikaremoana, N of

western extremity of Lake Waikaremoana, 1 160 m. En-

gel 23287 (F). ANTIPODES ISLAND: Stack Bay, Tay-

lor 64 as T. plumulosa (AK 227343). SCOTLAND: Ar-

gyll, Benmore Arboretum, foot of Glen Massan, v.-c. 98,

30 m. Long 14231 — c. young per. (F).

Telaranea quadriseta (Steph.) Engel & Merr.,

comb. nov.

Lepidozia quadriseta Steph., Spec. Hep. 3: 582. 1909.

Type: Australia, New South Wales, East Ballina,

Watts 433 (G!).

Lepidozia quadricilia Steph., Spec. Hep. 6: 338. 1922,

syn. nov. Telaranea quadricilia (Steph.) Engel &

Merr., Novon 9: 344. 1999. Type: Australia, New

South Wales, Clarence, Verreaux (com. Becherelle,

Herb. Mus. Paris) — c. per. (G!).

Plants resembling T. lindenbergii, delicate, soft,

flexuous, prostrate, pale; plants •small, to 2.5 mm

wide, including branches. Branching rather regu-

larly 1-2-pinnate, the branches terete, rather

closely spaced, often at right angles to main axis

and subopposite, often ± equal in length or the

branching subplumose, the branches sporadically

becoming flagelliform; branch half-leaf bifid; first

branch underleaf undivided and ciliiform, inserted

on ventral side of branch at juncture of branch

and main axis. Ventral-intercalary branches oc-

casional, forming leading, leafy shoots. Stems

with cortical cells distinctly differentiated, mod-

erately thickened, in 13 rows; cortical cells in sec-

tion much larger than the numerous (ca. 42), dis-

tinctly thick-walled medullary cells. Rhizoids of-

ten copiously produced from cells of underleaf

disc. Leaves rather stiff, obliquely spreading, con-

tiguous on main stems and imbricate on the

branches, ± plane, moderately to distinctly incu-

bous; leaves 540-600 u,m wide (between tips of

lobes) X 350-440 u,m long, ± symmetrically 4-

lobed to 0.8, the lobes widely divergent, much

longer than disc height. Branch leaves 4-lobed at

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

95

L.IM initially, weakly incubous, the insertion ex-

tending dorsally almost to the midline of the

branch. Lobes ± straight, ciliiform, distinctly ta-

pering, uniseriate to the base, inserted on a base

composed of 2 short disc cells (rarely with an ad-

ditional biseriate tier between base and uniseriate

row), the uniseriate portion 6-8(9) cells long, the

cell walls moderately thickened and firm, thick-

ened in the corners and moderately to distinctly

constricted at the septa, the basal cell of the uni-

seriate portion 29-35 u,m wide X 53-62 jxm long

(1.6-2:1), the next cell narrower, 22-30 u,m wide

x 54-70 u,m long (1.8-3.2:1), the terminal cell a

little shorter than the penultimate cell, evenly ta-

pered to the tip; cuticle finely but distinctly sca-

brous, roughened by minute, ± evenly spaced,

sharp prickles. Disc short-cuneate, 2 cells high

(including paired cells at bases of lobes, some-

times with 1 or both basal cells of the dorsal lobe

directly inserted on the stem), 8 cells wide; mar-

gins entire, straight. Underleaves much smaller

than leaves, 4-lobed nearly to base, the lobes cili-

iform, the uniseriate portion formed of 2(3) short

to somewhat elongated, often bulging cells, the

lobes terminating in a slime papilla; disc abbre-

viated, 2 cells high, 8 cells wide; margins entire.

Rhizoid initial cells small, subquadrate, formed

from some or all cells of underleaf disc. Branch

underleaves 2-3-lobed, rather large and at times

approaching the branch leaves in size. Cells of

disc (basal tier) somewhat thinner-walled than

lobe cells, trigones lacking, 20-26 u>m wide X

41-50 Jim long; cuticle smooth.

Plants dioecious. Androecia on short to mod-

erately long primary Frullania-type branches with

up to 4 pairs of normal vegetative leaves, subspi-

cate; bracts loosely imbricate, dorsally assurgent,

moderately concave, 2-lobed to ca. 0.6, the lobes

subcaudate, abruptly narrowing from a base of 3-

4 laterally juxtaposed cells to a uniseriate row of

4-5 moderately elongated cells, the terminal cell

tapering to the summit, the lobe cells papillose

like the leaves; dorsal margin of lamina slightly

dilated, with a small tooth; bracts monandrous;

antheridial stalk short, uniseriate; bracteolar an-

theridia absent. Gynoecia feebly dorsally assur-

gent, scarcely swollen at base; bracts of innermost

series closely ensheathing the perianth, concave,

elliptic, irregularly divided to ca. 0.5; lobes un-

equal in size, caudate, terminating in a uniseriate

cilium of 7-8 elongated, thick-walled cells that

resemble those of the leaf lobes, without a ter-

minal slime papilla; bracteole of innermost series

similar in form to bracts, but slightly smaller. Peri-

anth ± terete in basal sector, obscurely trigonous

above; mouth with ca. 10 caudate lobes, the lobes

each with a base composed of 2-4 laterally jux-

taposed, elongate cells, the lobes each terminating

in a uniseriate row of 7-8 elongate, thick-walled

cells, the basal cell 26-36 u,m wide X 139-173

u,m long (4.7-6:1), the septa weakly projecting;

lateral cells of lobe base often with their distal

end protruding to form a small tooth, or prolonged

into one or more slender accessory cilia that are

much narrower than the lobes and ca. 0.5 their

length.

Sporophyte not seen.

DIFFERENTIATION AND VARIATION — This species

is similar to T. lindenbergii but is distinguished

by the incubously inserted stem leaves (Fig. 28:

1 ), the longer leaf lobes with often distinctly con-

stricted septa, and the scabrous cuticle (Fig. 28:

3). As in T. lindenbergii, the leaf disc is only 2

cells high, consisting of a single tier of cells, plus

the basal tier of the lobes. The uniseriate row of

the lobes of the 9 bracts is much longer, 7-8 cells

long and without a terminal slime papilla in T.

quadriseta vs. only 2-4 cells in T. lindenbergii,

with a terminal slime papilla.

For comparison of T. quadriseta with T. gran-

ulata, another species with a scabrous cuticle, see

under that species.

Stephani's illustration (Icones, Lepidozia, 106)

depicts a leaf with a disc 2 cells high and lobes

with a uniseriate row 7-9 cells long. A young

perianth and a $ bract and bracteole are also il-

lustrated.

DISTRIBUTION AND ECOLOGY — Known only from

the types of L. quadriseta and L. quadricilia, and

from a few collections from Queensland. At the

Paluma site (NE of Townsville, 18° 59' S, 146°

13' E), the species occurs over rotten logs in a

montane notophyll (Cardwellia) rain forest. The

plant from Browns Creek (26° 32' S, 152° 56' E)

was collected at 40 m on bare wood of a rotting

log in a wet sclerophyll forest. Stephani (1922)

described T. quadricilia as caespitose, growing in

tufts on bark. The label of the type of T. quadri-

seta bears the notation, "Swamp, Heath."

NOMENCLATURE — Lepidozia quadricilia Steph.

(1922) was transferred to Telaranea by Engel and

Merrill (1999), however Lepidozia quadriseta

Steph. (1909) proves to be an older name for this

species. Grolle (1963, p. 175) listed L. quadriseta

Steph. as a synonym of T. tetradactyla. Grolle's

Kurzia quadriseta (1963), also a Telaranea, is

treated here as T. pallescens, based on Kurzia

96

FIELDIANA: BOTANY

FIG. 28. Telaranea quadriseta (Steph.) Engel & Merr. 1. Leaf, in situ, with base of Frullania-lype branch (= FB)

and half-leaf (= HL); dorsal view. 2. Leaf (cuticular detail not shown). 3. Leaf lobes, cuticular detail shown for lobe

at left. 4. Sector of main shoot showing underleaf and base of terminal branch (FBU = first branch underleaf; sp =

slime papilla). 5. Lobe of 6 bract, roughened cuticular shown in part. 6. Antheridium. 7. Distal portion of perianth.

8. Lobe of perianth mouth. 9, 10. Innermost 9 bract and bracteole, respectively (after Stephani Icones, Lepidozia no.

106). (All from type of Lepidozia quadricilia.)

ENGEL & MERRILL: AUSTRAL HEPATICAE TELARANEA

97

iiuilcscens Grolle, a name he published simulta-

neously.

NOTE — East Ballina, the type locality, is located

at 28° 52' S, 153° 34' E. The type locality of L.

quadricilia, "Clarence," most likely refers to the

Clarence River region (ca. 29° 34' S, 152° 43' E)

in north-eastern New South Wales (David Meagh-

er, Surry Hills, VIC, Australia, in lit.).

SPECIMENS SEEN— AUSTRALIA. QUEENSLAND: H

Track, Paluma, NE of Townsville, Meagher 01064 — c.

6 (F); Browns Creek, Dam road, W of Paluma, Meagher

1128— c. young 9 (F); Birthday Creek, 3.5 km W of

Paluma, Leaver & Fagerberg s.n. — c. $ (F); Yandina,

40 m, Windolf655B as T. Idispar (MELU).

Telaranea sect. Telaranea

Kurzia subg. Nanolepidozia Schust., J. Hattori Bot.

Lab. 48: 353. 1980. Type: Lepidozia bisetula Steph.

Telaranea martinii (Hodgs.) Schust.

Lepidozia martinii Hodgs., Trans. Roy. Soc. New

Zealand 83: 602. pi. 1, f. II. 1956 ("martini").

Telaranea martinii (Hodgs.) Schust., J. Hattori Bot.

Lab. 26: 256. 1963. Holotype: New Zealand, North

Is., road to Dawson Falls, Mt. Egmont, 2000-3000

ft., Jan., 1955, Hodgson 10220 (MPN!— c. <J +

sporo.).

Plants delicate, flexuous yet firm, loosely pro-

cumbent, pale green, nitid; plants small, the main

shoots (595)600-840 |xm wide. Branching spar-

ingly and irregularly 1 -pinnate, the branches of

the Frullania type, rarely flagelliform; branch

half-leaf undivided and subulate to ± narrow-lin-

ear, very rarely bilobed, usually obliquely insert-

ed; first branch underleaf at base of branch, ±

symmetrically or somewhat asymmetrically bi-

lobed, typically consisting of a slender, divergent

lobe with a uniseriate row of 3 rather elongate

cells, the tip cell lacking a slime papilla, and a

shorter, often ± appressed lobe aligned with the

branch and consisting of (1)2 rather short cells,

the tip cell with a slime papilla. Ventral-interca-

lary branches occasional, leafy or flagelliform.

Stems with cortical cells distinctly differentiated,

thin walled, in 9 rows of cells much larger than

those of the medulla; medullary cells thin-walled,

consisting of an outer row of 6(7) cells and, with-

in, a core of 3 cells that are smaller than those of

the outer ring. Rhizoids sparsely developed, from

either tier of underleaf disc cells. Leaves on main

shoot widely spreading, contiguous to feebly im-

bricate both on main stem and branches, plane or

nearly so, the insertion distinctly incubous both

on main stems and branches; leaves 175-525 u,m

wide X 330-455 u,m long, ± symmetric to weak-

ly asymmetric (the ventral lobe sometimes weak-

er), 3-lobed to 0.5-0.7 (exceptionally with some

4-lobed leaves, 2-lobed on branches and weaker

sectors of main shoot), the lobes divergent, longer

than disc height. Lobes narrowly attenuate to cil-

iiform, consisting of a uniseriate row of (2)3-4(5)

cells inserted on a triangular base composed of 2-

3(4) laterally juxtaposed cells, often with 1(2) ad-

ditional biseriate tiers between lobe base and uni-

seriate row in 1 or more lobes (particularly the

dorsal and middle lobes), the cells of uniseriate

portion thin-walled, not or weakly constricted at