JUL 2 4 1984

w.o

FIELDIANA
Botany

Published by Field Museum of Natural History

Volume 36, No. 11 August 27, 1975

Austral Hepaticae III

Stolonophora, A New Genus of Geocalycaceae1

JOHN J. ENGEL
DONALD RICHARDS ASSISTANT CURATOR OF BRYOLOGY

DEPARTMENT OF BOTANY
FIELD MUSEUM OF NATURAL HISTORY

and

R. M. SCHUSTER

DEPARTMENT OF BOTANY
UNIVERSITY OF MASSACHUSETTS, AMHERST

There are three well-defined subfamilies within the Geocaly-
caceae, i.e., subf. Lophocoleoideae, subf. Geocalycoideae and subf.
Leptoscyphoideae (see Schuster and Engel, 1973). The genus
Stolonophora Engel & Schust., new genus, belongs to subf.
Lophocoleoideae, which is chfl^^e|Qf^^^^.pflssessing perianths
with three ± equally developed lobes of wnicn me^third is ventral
in position.

DEC la 1975

Stolonophora Engel &

Plantae dioicae, erectae, virides ver*!Mfttyifc<CW^M>3q^|ationes semper fere
intercalares, rami typi ventricalis atque lateralis, rami terminales rarissimi, typi
Frullaniae; stolones frequentes, prostrati vel descendentes. Cortex caulinus e cellulis
admodum incrassatis efformatus. Folia transverse vel subsuccube disposita, concava,
oblata vel rotundata vel ovata, apice et margine integris. Cellulae foliorum leviter vel
manifeste crassi-tunicatae, trigonis mediocribus vel magnis; cellulis dimorphis,
plerumque 2 (raro 3) guttis olei praeditis, paucis guttula destitutis. Amphigastria
manifesta, deminuta, 0. 15-0.55 x latitudine caulina, indivisa vel bifida. Gemmae
nullae. Androecia intercalaria vel terminalia ad axillas foliacias longas; antheridia
solitaria, pedicellus uniseriatus vel hie inde biseriatus. Gynoecia ad axillas foliaceas

1 Grateful acknowledgement is made to the National Science Foundation— grant
GV-26615 to Dr. Henry A. Imshaug, Michigan State University, which supported the
senior author's field work, and grant BMS-72-02149 AOl to Field Museum of Natural
History for support of the scanning electron microscope facility. The senior author
would also like to thank Dr. Rolf Singer for assistance with the Latin diagnosis and
Mr. Fred Huysmans for his patient and enthusiastic assistance with the scanning
electron microscope at Field Museum.

Library of Congress Catalog Card Number: 75-23065

US ISSN 0015-0746

Publication 1208 111

112 FIELDIANA: BOTANY, VOLUME 36

longas; bracteae integrae, bracteolae admodum deminutae, bifidae vel indivisae.
Perianthia trigona, clavata vel ± campanulata, ore integro. Capsula pariete 3-4-
stratoso instructa. Sporae 14-17 /i; elateres 7-8(-9) /i lati, bispirales.

Typus.-Leioscyphus (?) abnormis Besch. & Mass. Bull. Mens.
Soc. Linn. Paris 1: 629. 1886.

Plants caespitose; erect, axes 0.8-10.5 cm. tall, leafy shoots 0.5-1.2 mm. wide;
green to yellow-brown to red-brown to brown.

Branches nearly exclusively ventral- and lateral-intercalary, terminal branching
very rare, of Frullania type, leafy branches erect; stolons present, prostrate or
descending, often branched, often originating near base of microphyllous or leafless
erect axis that soon becomes leafy.

Stems ± rigid or wiry, 8-11 cells in diameter, cortex in 1-2 rows of very thick-
walled cells the same size as or smaller than the medullary; medullary cells thin to
slightly thick walled. Rhizoids on leafy axes few or none, restricted to lower underleaf
bases; on stoloniform axes at bases of both leaves and underleaves.

Leaves transverse to subsuccubously oriented, insertion narrow, transverse to
distinctly succubous, leaves commonly subvertical, entire leaf weakly to moderately
concave or moderately concave in median portions with the peripheral portions
nearly flat; oblate to rotundate to ovate; apices undivided, entire; margins entire,
broadly rounded, dorsal margin rather long decurrent.

Leaf cells with walls slightly to distinctly thickened, trigones medium to large;
cuticle smooth or finely papillose. Cells dimorphic: a majority with 2(-3) oil-bodies
per cell, 10-20 per cent of cells without oil-bodies; oil-bodies colorless, faintly
granulate, subspherical to ovoid.

Underleaves reduced, ± appressed to moderately spreading, hard to discern, 0.15-
0.55 x the stem width; undivided or bifid and with segments cilialike; lamina margins
entire or with 1 tooth.

Gemmae absent.

Plants dioecious; androecia intercalary or (at first) terminal on main axis or long
leafy branches; bracts subequal to or somewhat smaller than leaves, ovate to
rotundate, free margin of saccate portion entire or with 1 slime papilla or with 1-2
teeth, bracts otherwise entire; antheridia solitary, stalk uniseriate throughout or with
isolated cells vertically septate.

Gynoecia on main axis or long intercalary branches; subfloral innovations absent
or from axil of bract of innermost series or bract or bracteole of second series,
innovations often repeatedly producing perianths. Bracts becoming progressively
larger toward the perianth; bracts of innermost series deeply concave in median basal
portion, rotundate to subrectangular to obovate; apices undivided, entire; margins
entire. Bracteoles of innermost series free from bracts; highly reduced, hardly
modified from underleaves; very short bifid or undivided; lamina margins entire or
with several slime papillae and 1 to several small teeth. Perianth trigonous, clavate to
± campanulate, somewhat or hardly laterally compressed, ventral lobe convex to
somewhat infolded; perianth expanding or ± contracted toward the mouth; the
mouth ± 3-lobed, entire.

Seta 7-9 cells in diameter, with 25-32 rows of outer cells surrounding an inner
core of cells smaller, subequal or slightly larger than outer row. Capsule wall of 3-4

ENGEL & SCHUSTER: AUSTRAL HEPATICAE III 113

layers, outer layer of cells 2.2-3.7 X as thick as each of interior strata, outer layer
with red-brown thickenings on radial walls, the thickenings wide nodulelike to wide
spinelike, semiannular bands sparing or absent, exposed wall thickened; intermediate
and inner layers of cells subequal in thickness, intermediate layer(s) with thickenings
on radial walls, but often considerably tangentially dilated; inner layer of cells with
incomplete semiannular bands extending for varying lengths, bands occasionally
complete, nodulose and sharp spinose thickenings common. Spores 12-17 /i, light
brown, exine with short, close vermiculate ridges. Elaters 7-10 ju wide, bispiral to 2-3-
spiral.

One of the two species here placed into Stolonophora has been
previously assigned to Clasmatocolea (Grolle, 1960), and the other
species has been treated as a synonym of a Clasmatocolea taxon
(Grolle, 1972). The following ensemble of features will distinguish
Stolonophora from the genus Clasmatocolea: a) the presence of
stolons; b) the nearly exclusively intercalary branching (only a
single terminal, Frullania-type branch was observed); c) the erect
growth; d) the dimorphic leaf cells, with the majority possessing oil
bodies, but with 10-20 per cent of the cells lacking them (see fig. 5);
and e) the vertically oriented, subtransverse leaves.

Rather than possessing affinities with Clasmatocolea, it seems
more likely that Stolonophora is closer to the purely Austral
complex centering around Pachyglossa and Lophocolea boveana of
southern South America, and the L. austrigena-gunniana complex
of South America-New Zealand-Tasmania. The relationships of
Stolonophora to the New Zealand taxa Pachyglossa tenacifolia and
Lophocolea gunniana seem particularly pertinent. In both of the
latter taxa, as well as in Stolonophora perssonii, we find the leaf
cells are clearly dimorphic, with a varying percentage devoid of oil-
bodies, others bearing typically one or two, rarely three, finely
botryoidal oil-bodies of, basically, identical form. Although this
similarity may be fortuitous, it is linked with a) evolution of a stem
with a distinct, small-celled, thick-walled cortex; b) a marked
tendency for the leaves to become unlobed; c) an equally marked
tendency— very unusual in Lophocoleoideae— for the leaves to be
vertical and transversely oriented. Within the above complex of
taxa, we find clear analogies to, especially, Pachyglossa. In
Pachyglossa we also find brown-to-fuscous coloration in some taxa;
microphyllous stolons; growth away from the substrate (in some
taxa); and a combination of ventral and lateral-intercalary axillary
branching with isolated Frullania-type branching, (depending on
the species) rare or lacking. Stolonophora, however, is very distinct
from the Pachyglossa- Lophocolea boveana-austrigena-gunniana

114 FIELDIANA: BOTANY, VOLUME 36

complex in the highly reduced and appressed underleaves, and the
basically bilateral and two-ranked organization of vegetative axes.

KEY TO THE SPECIES OF STOLONOPHORA

1. Leaf margins not to weakly decurved, leaves pale to dark brown to red brown, leaf
cells with walls moderately to distinctly thickened; axes 1.8-10.5 cm. tall; stems
185-335 n in diameter; underleaves 0.15-0.35(-0.45)X the stem width; perianth ±
included, for the most part sheathed by bracts; plants submerged in streams or
pools or on stream banks. Southern South America S. abnormis.

1. Leaf margins tightly and narrowly revolute, leaves green, not developing secondary
pigments, leaf cells with walls slightly thickened; axes 0.8-1.5 cm. tall; stems 150-
170 p in diameter; underleaves 0.40-0.55 x the stem width; perianth exserted, bracts
obliquely patent; plants on cliff walls subject to inundation. North and South
Island, New Zealand S. perssonii.

1. Stolonophora abnormis (Besch. & Mass.) Engel & Schust.
comb. nov.

Leioscyphus (?) abnormis Besch. & Mass. Bull. Mens. Soc. Linn.
Paris 1: 629, 1886. Lophocolea abnormis (Besch. & Mass.) Steph.
Sp. Hep. 3: 62. 1906. Clasmatocolea abnormis (Besch. & Mass.)
Grolle, Revue Bryol. Lichen. 29: 71. 1960. Type: Chile, Prov.
Magallanes, Isla Hoste, Hyades (FH!, G!)(c. per.).

Plants gregarious with other bryophytes or in dense pure tufts; axes 1.8-10.5 cm.
tall, the leafy shoots 0.5-1.1 mm. wide, pale to dark brown to yellow-brown,
occasionally tinged with red-brown, stems red-brown or very dark brown, rather stiff,
wiry.

Branches frequent, of lateral- and ventral-intercalary types, Frullania-type
branching very rare (only 1 branch seen), leafy branches erect; stolons frequent but
not uniformly present on all axes, pinkish tinted, often in an interwoven mass in
which soil particles may be trapped, often prostrate from median or basal portion of
leafy axis; stolons occasionally branched, with scaly leaves and underleaves which are
sometimes caducous.

Steins 185-335 /t in diameter, 8-11 cells in diameter, cortex in 1-2 rows of very
thick-walled cells the same size as or smaller than the medullary; the outer row with
walls thickest and often with small lumen; medullary cells thin to slightly thick
walled, with comers thickened similarly to small to medium trigones. Rhizoids absent
from leafy shoots, common on stolons, at base of leaves and underleaves or
occasionally stolons with a few regions of scattered rhizoids.

Leaves (fully developed) (0.7-)0.9-1.7 mm. wide, 0.7-1.3 mm. long, axes commonly
with regions of small leaves intercalary between fully developed leaves; the later
transverse to weakly succubously oriented, insertion narrow, transverse to distinctly
succubous, leaves commonly subvertical, erect to weakly spreading, approximate to
weakly imbricate, entire leaf weakly to moderately concave and with margins plane
or leaves weakly to moderately concave in median portion but with peripheral
portions nearly flat and with margins plane or slightly reflexed; leaves oblate to
rotundate to ovate; apex broadly rounded to truncate, occasionally retuse.

FlG. 1. Stolonophora abnormis (Besch. & Mass.) Engel & Schust. 1, perianth-
bearing axis with basal stolons (VL - ventral lobe), x 16; 2, median leaf cells, x
450; 3, portion of leafy axis, X 11; 4-6, underleaves of main, leafy axis, X 90; 6-7,
leaves of main axis, x 23; 8, stem, cross-section through main, leafy axis, X 270.
Numbers 1, 4-6 from Engel 6092, Chile, Prov. Magallanes, Puerto Gallant; 2, 3, 7, 8
from type material.

115

116

FIELDIANA: BOTANY, VOLUME 36

Leaf cells with walls moderately to distinctly thickened, trigones medium to
large, often confluent, the sides straight to bulging, occasionally concave; median leaf
cells (12-)17-44 ^ wide, (14-)26-53(-56) jn long, median basal cells distinctly thick
walled, 18-42(-48) /* wide, (34-)41-76(-82) /i long; cuticle smooth or finely papillose.

Underleaves distinct, reduced, of very thick-walled cells, 0.15-0.35(-0.45)x the
stem width, moderately spreading; bifid to varying degrees and with segments
cilialike, or undivided and then acuminate, underleaf segments or apex terminating in
a slime papilla; lamina margins entire or with 1 single-celled tooth.

Plants dioecious; <5 bracts somewhat smaller than leaves, obovate to rotundate,
moderately concave, margins plane, free margin of saccate portion often broadly
rounded, entire, with 1 slime papilla or with 1-2 sharp teeth terminating in a slime
papilla, bract otherwise entire; antheridial stalk uniseriate throughout or with
isolated cells vertically septate.

Gynoecia on main axis or long lateral- or ventral-intercalary branches; subfloral
innovations absent or from axil of bract of innermost series or bract or bracteole of
second series, innovations often repeatedly producing perianths. Bracts in 2-3 series;

FIG. 2. Stolonophora abnormis (Besch. & Mass.) Engel & Schust. 1-2, bract and
bracteole of innermost series, x 23; 3, capsule wall, outer layer, x 450; 4, capsule
wall, cross-section, x 450; 5, capsule wall, inner layer, X 450; 6, antheridial stalk, X
450; 7, partially torn seta, cross-section, x 90. Numbers 1-5, 7 from Engel
6092, Chile, Prov. Magallanes, Puerto Gallant; 6 from Engel 5718, Chile, Prov.
Magallanes, Bahfa Tuesday.

ENGEL & SCHUSTER: AUSTRAL HEPATICAE III 117

bracts of innermost series free or with dorsal margin fused for entire length with
perianth keel; deeply concave in median basal portion; rotundate to subrectangular
to obovate, basal portion often ± clasping perianth; apices broadly rounded to
subtruncate. Bracteoles of innermost series free from bracts, rarely inserted on basal
portion of perianth and then with 1 margin partially fused with perianth; bracteole
highly reduced, 0.2-1.2 mm. long, hardly modified from underleaves; very short bifid
or undivided and then with apices narrowly rounded or short to long apiculate, apex
or segments terminating in a slime papilla; lamina margins entire or with several
slime papillae and 1 to several small teeth terminating in a slime papilla. Perianth
trigonous, clavate to ± campanulate, somewhat laterally compressed, the sides
convex, the ventral lobe convex to (more commonly) somewhat infolded, occasionally
canaliculate; perianth expanding toward the mouth; the mouth wide or subcon-
stricted, lobes broadly rounded, inflexed or occasionally straight in median portion
and reflexed near the keels.

Seta 7-8 cells in diameter, with 25-28 rows of outer cells surrounding an inner
core of scattered cells slightly to distinctly smaller than outer row and with corners
thickened similarly to medium or large trigones. Capsule wall 32-48 /t thick, of 3-4
layers, outer layer of cells 2.2-3.6 x as thick as each of interior strata, outer layer 16-
26 /i thick, with red-brown thickenings on radial walls, the thickenings wide
nodulelike to wide spinelike and often feebly tangentially dilated, with a few to
several semiannular bands which are sometimes branched; inner layer of cells 7-8(-ll)
ft. thick, with light brown thickenings on radial and tangential walls, incomplete
semiannular bands extending for varying lengths, bands only occasionally complete,
nodulose and sharp spinose thickenings common. Spores 14-17 /i, light brown, exine
with short, stout vermiculate ridges. Elaters 7-8(-9) n wide, bispiral to both ends or 1
or both apical portions with thick nonspiral walls, elater walls light brown.

The senior author has studied the spores of this species with
the aid of a scanning electron microscope. Figures 3-4 reveal the
spore exine is covered with short, stout, vermiculate ridges which
are rather close to one another or often coalesce. It may be
observed from Figure 4 that the ridges possess spherical nano-
granules which are usually distinct and separate, with intervening
spaces, but occasionally are juxtaposed or fused.

Ecology-Phytogeography.—Stolonophora abnormis occurs in
Tierra del Fuego and the wet, cold Patagonian Channels north to
52° 17' S. The species occurs submerged in rather rapidly moving
streams and occasionally on stream banks in evergreen Nothofagus
forests or submerged in small pools in the Magellanian moorland. It
may also be subalpine or alpine. It is also known from near
Concepcion (ca. 36° 50' S in Prov. Concepcion) in the Valdivian
region. Arnell (1958) reports the species from Tristan da Cunha.

Specimens seen.-CHILE. PROV. CONCEPCION: Peumo,
Dusen 186 as Jamesoniella paludosa (S-PA). PROV. MAGAL-
LANES: Isla Pacheco, Skottsberg 141 as Jamesoniella grandiflora

118 FIELDIANA: BOTANY, VOLUME 36

(S-PA); Isla Desolaci6n, Bahia Tuesday, at head of inner harbour,
Engel 5697, 5718 (MSC); Isla Desolation, Puerto Churruca, at head
of Brazo Lobo, Engel 5852, 5857 (MSC); Isla Desolation, Puerto
Angosto, ca. 200 m., Dusen 186 as L. abnormis (G), as Jamesoniella

FlG. 3. Stolonophora abnormis (Besch. & Mass.) Engel & Schust. Scanning
electron photograph of spore, x 7,500; from Engel 6092.

paludosa (L, S-PA, UPS); NE side of Puerto Gallant, Engel 6065D,
6092 (MSC); Rio Azopardo, ca. 500 m., Dusen 112 (G, S-PA); W end
of Lago Fagnano, "alpinus," Halle 144, as Jamesoniella oenops
(UPS); Isla Londonderry, Exp. Ant Belg. 397 (G).

2. Stolonophora perssonii (Schust.) Engel & Schust. comb. nov.

Calyptrocolea perssonii Schust. Revue Bryol. Lichen. 34: 699. 1967.
Holotype: New Zealand, South Island, Fiordland Nat. Park, E

FIG. 4. Stolonophora abnormis (Besch. & Mass.) Engel & Schust. Scanning
electron photograph of spore, X 14,350; from Engel 6092. Details of exine showing
ridges and nanogranules.

119

FIG. 5. Stolonophora perssonii (Schust.) Engel & Schust. 1, main axis, lateral
view, X 23; 2-3, abaxial (AB) and adaxial (AD) views of same main axis leaf, X 46;
4-6, underleaves of main, leafy axis, x 90; 7, dimorphic median leaf cells, with oil-
bodies in some cells while absent in others, X 400; 8, axis with stoloniform branching,
ventral view (LA = leafy axis base), x 9; 9, stem, cross-section through main leafy
axis, x 270. All from holotype.

120

FlG. 6. Stolonophora perssonii (Schust.) Engel & Schust. 1, shoot apex, ventral
aspect, with barely emergent dehisced capsule, X 22; 2, shoot apex with perianth,
dorsal aspect, x 22; 3, shoot sector, dorsal aspect, with lateral-intercalary branch, x
31; 4, shoot sector, lateral aspect; note appressed underleaves, x 31; 5, female bract
and, within, bracteole, to one scale, x 22; 6, bracteole of Figure 5, x 375; 7, leaf, x
31; 8, capsule wall, outer layer, x 312; 9, capsule wall, inner layer, X 312; 10,
capsule wall, cross-section, X 500; 11, portion of plant showing highly ramified base;
arrows indicate points of removal of leafy shoot apices; note branches x, y, z, from all
three merophyte rows of the primary axis, x 44. Numbers 1-10 from Schuster 48872,
New Zealand, Mt. Egmont; 11 from holotype.

121

122 FIELDIANA: BOTANY, VOLUME 36

entrance of Homer Tunnel, head of Hollyford Valley, 3000-3050
ft., Schuster 55596 (RMS!).1

Plants in nearly pure, rather dense tufts; axes 0.8-1.5 cm. tall, the leafy shoots
ca. 0.7-1.2 mm. wide, subequally leaved throughout, green to slightly olive green, with
the mature stem brownish.

Branches frequent, intertwined, of lateral- and ventral-intercalary types, the
ventral-intercalary more common, terminal branching absent, leafy branches erect;
stolons frequent, usually pinkish tinted, often descending or prostrate from base of
leafy axis; stolons often branched, with scaly, often hyaline leaves and underleaves
which are sometimes caducous.

Stems ca. 150-170 ju, in diameter, 9-11 cells in diameter, cortex in 1-2 rows of very
thick-walled cells smaller than the medullary; medullary cells thin walled, with
corners thickened similarly to small trigones. Rhizoids rather frequent, on leafy
shoots restricted to underleaf bases; on stoloniform axes at bases of leaves and
underleaves.

Leaves 0.6-0.85(-0.9) mm. wide, 0.6-0.7 mm. long, transverse to subsuccubously
oriented, insertion narrow, weakly to distinctly succubous, leaves subvertical,
moderately to rather strongly spreading, distant to weakly imbricate, moderately
concave in median portions, the peripheral portions nearly flat, but with margins
narrowly, often strikingly reflexed; leaves rotundate; apex broadly rounded to
truncate to weakly retuse.

Leaf cells with walls slightly thickened, trigones medium, occasionally confluent,
the sides concave to straight; median leaf cells 13-20 ju, wide, 22-29(-37) ju. long, median
basal cells uniformly and moderately thick walled, elongated, 13-20 ju wide, 22-29(-37)
H long; cuticle smooth. Cells dimorphic: a majority each with 2, extremely rarely 3
oil-bodies, a minority of 10-20 per cent without oil-bodies; oil-bodies colorless, faintly
granulate, almost smooth externally, subspherical to ovoid, ca. 3-5 x 5-8 ju or
spherical and 4-5 p.

Underleaves distinct, reduced, 0.40-0.55 X the stem width, appressed; with 1-2
often cilia like segments or undivided.

Plants dioecious; $ bracts hardly smaller than leaves, ovate to rotundate, deeply
concave, margins reflexed, entire; antheridial stalk uniseriate but with local biseriate
areas.

Gynoecia terminal on leading axes; subfloral innovations absent. Bracts in 3
series; bracts of innermost series rather stiffly obliquely patent, deeply concave to
cupulate, especially toward median base; rotundate, essentially identical to leaves in
size and shape; apices narrowly revolute to decurved, broadly rounded; margins
distally narrowly revolute to decurved. Bracteoles of innermost series free from
bracts; minute, at most 0.05-0.1 area of bracts; appressed. Perianth often rather
stipitate, clearly exserted, obovoid to ovoid, inflated-trigonous, the 2 lateral keels
locally weakly winged and ± sharp, the dorsal keel blunt to rounded; slightly
contracted to the shallowly trilobed mouth, lobes blunt to rounded, margins reflexed.

Seta 8-9 cells in diameter, with 31-32 rows of outer cells surrounding an inner
core of scattered cells subequal to or slightly larger than outer row and with corners

1 The acronym RMS indicates the private herbarium of R. M. Schuster.

ENGEL & SCHUSTER: AUSTRAL HEPATICAE III 123

thickened similarly to medium trigones. Capsule spherical, wall 32-40 /* thick, of 4(-5)
layers, outer layer of cells 2.8-3.7 x as thick as each of interior strata, outer layer 17-
18 p. thick, with thickenings on radial walls, the thickenings wide nodulelike to wide
spinelike; inner layer of cells 5-6 /i thick, with weak nodular thickenings on radial
walls, thickenings rarely tangentially dilated, semiannular bands rare to sporadic,
usually incomplete. Spores 12-14 j^, light brown, exine rather thin and with low
vermiculate, close, irregular anastomosing ridges. Elaters 8-10 /x wide in median
portion, medially mostly 3-spiral, becoming bispiral distally, spirals very narrow.

Taxonomic remarks.— This species was treated by Grolle (1972,
p. 365) as a synomym of Clasmatocolea fiordlandiae (Hodgs.)
Grolle. Clasmatocolea fiordlandiae, type material of which has
been studied by the senior author, rightfully belongs to Clasmato-
colea and is not conspecific with Stolonophora perssonii. For notes
regarding generic differences between Stolonophora and Clasmato-
colea see p. 113.

Schuster (1967, p. 700) assigned this species to Calyptrocolea
(thus the Adelanthaceae) and placed it there because it is
exceedingly close in habit to Calyptrocolea gemmipara Schust.
However, he noted that the position of "C." perssonii in
"Calyptrocolea is subject to possible doubt" and noted that it
differed from C. gemmipara, in "a series of good vegetative
characters," i.e.: the dimorphic leaf cells; the thick-walled cortical
cells that are short-oblong rather than long-rectangular; the
occasional lateral-intercalary branches; and the distinct, if small
underleaves. The near-identity of the two taxa in size, stolons, leaf
form and orientation, plus the absence of terminal branching, is
truly astonishing. Study of the antheridial and perianth-bearing
plants we have seen clearly shows that the species rightfully
belongs in the Geocalycaceae.

The name "Calyptrocolea dimorpha" in Schuster (1966, p. 211)
is actually an herbarium name for C. perssonii.

It should be specifically mentioned that S. perssonii has
dimorphic leaf cells, i.e., some leaf cells possess oil-bodies while
others are totally oil-body free (see fig. 5). This situation is rather
exceptional in the Hepaticae and occurs in several unrelated taxa
(cf. Schuster, 1966).

Ecology -Phytogeography.— Stolonophora perssonii is known
only from limited materials. It is an alpine species and occurs on
cliff walls subject to inundation, adjacent to steep cascades and
small waterfalls at 3000-3050 ft. in Fiordland National Park, South
Island, New Zealand, and at similar sites at 5000-5400 ft. on Mt.

124 FIELDIANA: BOTANY, VOLUME 36

Egmont, North Island, New Zealand. It may be associated with
Temnoma quadrifidum.

Specimen seen.-NEW ZEALAND. TARANAKI PROV.: Mt.
Egmont, east slope of mountain near Warwick Castle in alpine
summit zone, 5000-5400 ft., Schuster 48872 - c. <5 + sporo. (RMS).

REFERENCES

ARNELL, S.

1958. Hepatics from Tristan da Cunha. Results Norw. Sclent. Exped. Tristan da
Cunha 1937-1938, no. 42, pp. 1-76, 30 figs.

GROLLE, R.

1960. Nachtrag zur "Revision der Clasmatocolea-Arten." Rev. Bryol. Lichen., 29,
pp. 68-91, pi. 1-3.

1972. Zur Kenntnis von Adelanthus Mitt. Jour. Hattori Bot. Lab., 35, pp. 325-370,
figs. 1-11.

SCHUSTER, R. M.

1966. The Hepaticae and Anthocerotae of North America east of the hundredth
meridian, Vol. 1. Columbia University Press, i-xvii, 1-802 pp., figs. 1-84.

1967. Studies on Hepaticae, VII-X. On Adelanthus Mitten and Calyptrocolea
Schuster, gen. n. Rev. Bryol. Lichen., 34, pp. 676-703. (1966).

SCHUSTER, R. M. and J. J. ENGEL.

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UNIVERSITY OF ILLINOIS-URBANA