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.