Studies in the liverwort genus Fossombronia ( Metzgeriales ) from southern Africa . 8 . F \ elsieae and F . spinosa , two new Western Cape species with spinose spores

Two new species with spinose spores are described from the Western Cape. F. elsieae Perold is quite a large plant, gener­ ally with almost entire leaves, its spores on the distal face having relatively few, rather coarse spines, which often appear bro­ ken and are occasionally linked to form abbreviated ridges. F spinosa Perold is a smallish plant with irregularly lobed leaves, its spores on the distal face having numerous, fine spines, which sometimes coalesce to form short ridges. A table which com ­ pares F. glenii Perold, F. leucoxantha Lehm. and F. montaguensis S.W.Amell with the two new species, is supplied. 1. F ossom bronia elsieae Perold, sp. nov. Plantae dense coarctatae, apicem versus virides, cito alheseentes et translucentes. Folia late patentia, imbricata, obovata vel suboblonga. Dioicae. Antheridia bracteis forma irregulari subtensa. Pseudoperianthium campanulatum; ore undulato cum lobis ± 6 rotundatis. Sporae 47.5-57.5 pm diamelro, superfieie distali convexa, cum 9 vel 10 seriebus spinarum grossarum conicarum, interdum in cristis irregularibus concatenatarum; superfieie proximali sine nota triradiata, cum cristis brevibus et spinis nonnuliis. Elateres 125-165 pm longi. 7.5-12.5 pm lati, omnino ter spirales vel alii bis spirales, alii apicem versus bis spirales, alii ter, raro quater apices versus ter, spirales. TYPE.— Western Cape, 3318 (Cape Town): Table Mountain, top of Nursery Gorge, (-CD ), E. Esterhuysen 24885 (BOL, holo.). Plants in crow ded stands, apically green, soon bleached and translucent, rather fragile; shoots medium­ sized to quite large, 6 -12 mm long, ± 2.5 mm high, 3.5-4.5 mm wide; simple (Figure 2A) or apically furcate, segments 2-3 mm long, moderately divergent, some­ times with lateral buds toward base. Stems prostrate, fresh apical shoots often arising from somewhat with­ ered base (Figure 2B), in recent collections dorsally chlorophyllose, vcntrally purple, in cross section plano­ convex, in male plants at apices (Figure IN) 300-350 pm (10 cell rows) high, 500-600 pm wide and at bases (Figure lO) 220-350 x 300-430 pm; in female plants at apices (Figure IP) 300-400 pm (12 cell rows) high, 450-500 pm wide, tapering proximally anti basally (Figure IQ). 250-300 x 400-450 pm. Leaves spreading widely, overlapping, succubously inserted on stem, quite large, except for young apical ones, plane and rounded above, mostly longer than wide and narrowing toward base, obovate to almost oblong, in male plants (Figure I A-E) 1125-20ÍK) x 1125-1500 pm, in female plants (Figure 1F-K ) somewhat larger, 1700—2125 x 1625-2125 ^National Botanical Institute, Private Bag X 101, 0001 Pretoria. MS received: 199S-09-23. pm, margins nearly entire, with few (1-3) papillae, most­ ly at slight angulations. Leaf cells thin-walled, in male plants tending to be slightly smaller, but not significant­ ly so, in female plants at upper margin (Figure 1L) sub­ quadrate to rectangular across. 45-50 x 55.0-62.5 pm, at lower lateral margins long-rectangular, 90.0-112.5 x 27.5-47.5 pm; upper laminal cells 5or 6-sided, 62.5-80.0 x 50-55 pm; middle laminal cells 87.5-132.5 x 57.5-62.5 pm; basal cells 75-125 x 37.5-50.0 pm. Oil bodies present in fresh material only (Figure 1M), up to 34 per cell, shiny, globular to slightly irregular in shape, ± 5 pm diam.; chloroplasts similar in size. Dioicous. Antheridia dorsal on stem, in I or 2 rows, short-stalked, globose, ± 270 pm diam., subtended by perigonial bracts (Figure 1R-U), single or paired (Figure 2C, D), shape irregular, body 400-500(-600) x 180—28()(—3 10) pm, upper margin with low, blunt or tall, spinous projections, up to 300 pm long and 160 pm wide at base, tapering to a pointed tip topped with a papilla, cells in interior 5or 6-sided, 60.0-82.5 x 3 7 .5 ^ 5 .0 pm. Archegonia (Figure 2E) in 1 ,2 or more rows, dorsally along stem. Pseudoperianth (Figure IX, Y; Figure 2F) campanulate, near stem apices, 1500-1800 pm long, sometimes slit along the side, one margin overlapping the other, base narrow, ± 875 pm wide, upwardly widely daring, 2250-2625 pm across undulating mouth, with ± 6 rounded lobes, marginal papillae few, cells comparable in shape and size to those of leaves. Capsules globose, ± 625 pm diam., cells in inner layer of bistratose capsule wall (Figure IV) ± rectangular, roughly triangular or irregularly shaped, 52.5-65.0 x 30-50 pm, each cell wall with 1-3 nodular and some semi-annular thickenings. Seta from very short to 3.25 mm long, 150 pm or 9 cell rows wide (Figure IW). Spores brown to dark brown, hemispherical, 47.5-57.5 pm diam. including spines pro­ jecting around periphery; distal face convex (Figure 3A-D), covered with 9 or 10 rows of coarse, often coni­ cal spines (or a total of ± 39), ± 5 pm long, basally joined by fine lines, sides grooved, apices sometimes ‘broken', apparently hollow inside, occasionally linked to form irregular ridges; proximal face (Figure 3E) almost flat, lacking a distinct triradiate mark, with short, slightly sin­ uous or occasionally branched ridges and some blunt 26 Bothalia 29,1 (1999) FIGURE 1.— Fossombronia elsieae. A -E, leaves o f male plant; F-K, leaves o f female plant; L, detail of marginal area o f leaf; M, median leaf cells with oil bodies (solid lines) and chloroplasts (dotted lines); N, c/s apical part of male stem; O, c/s basal part o f male stem; P. c/s api­ cal part o f female stem; Q, c/s basal part o f female stem; R -U , perigoniaJ bracts; V, cells in capsule wall with thickenings; W, c/s seta; X, pseudoperianth from side; Y, opened pseudoperianth A -L, N-V, X, Y, Esterhuysen 24885: M, W, Braggins 97/356A. Scale bars: A -K , X, Y, 500 Mm; L, W, 100 ym; M, V, 50 ym; N -Q , R -U , 250 Mm. Bothalia 29,1 (1999) 27 FIGURE 2 .— Fossombronia elsieae. A, simple shoot; B, fresh apical shoot from rather withered base; C, D, male stem with perigonial bracts; E, female stem with archegonia from above; F, female stem with pseudoperianth and capsule from above. A, E, S.M. Perold 3475\ B -D , F, Esterhuysen 24885. A, x 7.8; B, x 12.8; C, x 10.5; D, x 16; E, x 17; F, x 17. spines in between, 21-25 spines projecting around periphery. Elaters (Figure 3F) yellow, smooth, 125-165 pm long, 7.5-12.5 pm wide in centre, often tapered toward tips and ending in a loop, 5 jam wide; 3-spiral along entire length, occasionally 2-spiral only at tips, remainder 3-spiral; some wholly 2-spiral; rarely loosely 4-spiral, tapering tips 3-spiral, or only exceptionally branched into 2 near apex, both tips 2-spiral, the lower, unbranched part 3-spiral. Fossombronia elsieae has been named in honour of Ms Elsie Esterhuysen, the doyenne of southern African plant collectors, who collected it in October 1951 at the top of Nursery Gorge, Table Mountain. This specimen was FIGURE 3.— Fossombronia elsieae. Spores and elaters. A, B, distal face; C, side view o f distal face; D, detail o f spines and lamellae at margin o f distal face; E, proximal face; F, elaters. A, S.M. Perold 3475\ B-F, Esterhuysen 24885. A, x 761; B, x 711; C, x 876; D, x 1560; E, x 715;

Dioicous.Antheridia dorsal on stem, in I or 2 rows, short-stalked, globose, ± 270 pm diam., subtended by perigonial bracts (Figure 1R-U), single or paired (Figure 2C, D), shape irregular, body 400-500(-600) x 180-28()(-3 10) pm, upper margin with low, blunt or tall, spinous projections, up to 300 pm long and 160 pm wide at base, tapering to a pointed tip topped with a papilla, cells in interior 5-or 6-sided, 60.0-82.5 x 3 7 .5^5 .0pm.Archegonia (Figure 2E) in 1 ,2 or more rows, dorsally along stem.Pseudoperianth (Figure IX, Y; Figure 2F) campanulate, near stem apices, 1500-1800 pm long, sometimes slit along the side, one margin overlapping the other, base narrow, ± 875 pm wide, upwardly widely daring, 2250-2625 pm across undulating mouth, with ± 6 rounded lobes, marginal papillae few, cells comparable in shape and size to those of leaves.Capsules globose, ± 625 pm diam., cells in inner layer of bistratose capsule wall (Figure IV) ± rectangular, roughly triangular or irregularly shaped, 52.5-65.0x 30-50 pm , each cell wall with 1-3 nodular and some sem i-annular thickenings.Seta from very short to 3.25 mm long, 150 pm or 9 cell rows wide (Figure IW).Spores brown to dark brown, hemispherical, 47.5-57.5 pm diam.including spines pro jecting around periphery; distal face convex (Figure 3A -D ), covered with 9 or 10 rows of coarse, often coni cal spines (or a total of ± 39), ± 5 pm long, basally joined by fine lines, sides grooved, apices sometimes 'broken', apparently hollow inside, occasionally linked to form irregular ridges; proximal face (Figure 3E) almost flat, lacking a distinct triradiate mark, with short, slightly sin uous or occasionally branched ridges and some blunt "w> s.In my paper re-exam ining F. leucoxantha Lehm.(Perold 1997c), I referred to this collection, which 1 have now selected as the holotype of this new species, stating that it has 'almost entire leaves'.I ilius distinguished it from F. leucoxantha, which has 'denticulate it) incised-dentate leaves', in spite of both species having spinose spores.The spores in F leucoxan tha almost invariably have papillae between the at least 90 spines covering the distal face, whereas in F. elsieae there are no papillae and only up to 39, rather coarse, often bro ken spines on the distal face.Clearly, there are subtle dif ferences between these two species (and F. spinosa) in the spore ornam entation, but much more obvious differences vegctatively.In the recently described F. glenii (Perold 1997a) from the northern summer rainfall region, the spines on the distal face of the spores are often double and the ± undulating leaf margins have shallow angular lobes.

.tr
In Table 1, some characters o f F. elsieae and F spinosa are com pared to those o f F. glenii Perold, F leucoxantha and F. montaguensis S.W.Arnell.
In addition to the m ethodology em ployed by Scott & Pike (1988), and others, who relied 'alm ost completely on spore characters-surface sculpturing, often colour and, much less, spore s i/e -for discrim inating species', I have tried to use vegetative characters also.Extensive use of LM and SEM m icrographs o f the spores and thalli has, of course, m ade com parison between specim ens easier.At times, however, it rem ains extrem ely difficult to establish species param eters in this genus.Accordingly, I am still undecided where to place the other specimen, Arnell 6 /6 .w hich I also excluded from F leucoxantha in my treatm ent of that species (Perold 1997b).Other species with coarse spines on the distal spore face have been described from elsew here in the world, notably F caespitiform is De Not., which has a total of only ± 25 spines on the distal face and ± 29 around the periphery.Vegctatively it does not appear to be closely related to F elsieae. nor do the A ustralian species with coarsely spin ose spores, F. cultriform is, F m agnaspora and F truncata.which were described by Scott & Pike (1987).
Fossom bronia elsieae (and F. spinosa), with spores ripening in spring (Septem ber/O ctober), are hitherto known from only a few localities in the Western Cape (Figure 4), which is in the winter rainfall area o f south ern Africa.This is referred to as the Fynbos Biome (sclcrophyllous, m icrophyllous vascular plant vegetation) (Cow ling et al. 1997;Rutherford 1997) tively few, rather coarse spines which are occasionally linked to form short ridges.
TYPE.-Western Cape, 3320 (M ontagu): K ogm an's Kloof, 5 km from Montagu, immediately beyond tunnel on road to Ashton, at Loftus Bridge, along disused road along the left side of the river; on soil on vertical rock face, (-C C ), S.M. Perold 3835 (PRE,holo.).

SPECIMENS EXAMINED
Held at PRE, unless otherwise indicated.Bracketed numbers after collectors' name and number refer to the species in the text in alphabetical order, namely: F. elsiei (1) and F. spinosa (2).11).In Perold (1997b: figs 8C, D; 9A, B), the figures purporting to illustrate F. mon-ings, the photographer, Mrs A. Romanowski for devel oping and printing the photographs and Ms D. Maree for typing the manuscript.Dr J. Braggins of the University of Auckland, New Zealand, is kindly thanked for making available to me the specimens which he collected locally during his visit here in 1997.The curator of BOL is thanked for the loan of specimens.

FIGURE 1 .-FIGURE 2
FIGURE 1.-Fossombronia elsieae.A -E , leaves o f male plant; F -K , leaves o f female plant; L, detail of marginal area o f leaf; M, median leaf cells with oil bodies (solid lines) and chloroplasts (dotted lines); N, c/s apical part of male stem; O, c/s basal part o f male stem; P. c/s api cal part o f female stem; Q, c/s basal part o f female stem; R -U , perigoniaJ bracts; V, cells in capsule wall with thickenings; W, c/s seta; X, pseudoperianth from side; Y, opened pseudoperianth A -L , N-V, X, Y, Esterhuysen 24885: M, W, Braggins 97/356A.Scale bars: A -K , X, Y, 500 Mm; L, W, 100 ym; M, V, 50 ym; N -Q , R -U , 250 Mm.
as F crispa.
diam.Spores light brown to darker brown, hemispherical, 35-45 pm diam., including spines projecting around periphery; distal face convex (Figure 8A-C), densely covered with 11-15 rows of fine spines, (or a total of ± 95), almost smooth or faintly grooved, ± 5 pm long, apices quite often truncate (Figure 8D), occa sionally 'broken' and apparently hollow inside, some times linked to form several short ridges; proximal face (Figure 8E) slightly concave or almost flat, lacking a dis tinct triradiate mark, finely to coarsely granular and with some thin, irregular lamellae, numerous (± 40) spines projecting around periphery.Elaters (Figure 8F) yellow, mostly smooth, 100-150 pm long, ± 7.5 pm wide in the centre and tapering slightly toward tips, ending in loops, loosely 3-or 2-spiral throughout, or 3-spiral in the mid dle and 2-spiral at one or both tips.The name Fossombronia spinosa refers to its finely spinose spores and the species is so far known from only three localities in the Western Cape (Figure 4), which is in the winter rainfall area of southern Africa.The holo type specimen o f F. spinosa and Perold 3834 grow on pockets o f sandy, rather coarse soil derived from sand stone, on a vertical rock face, kept damp by seepage water.They are mixed with Targionia hypophylla, Riccia species, Goniomitrium africanum and Bryum species.Some o f A m ell's collections of this newly isolated species, S.W. Arnell CH4037, 793, from Bath Kloof and Kogman's Kloof, Montagu, as well as my own sterile specimens, S.M. Perold 3453 and 3454 p.p., from Bath Kloof have regrettably been wrongly identified as F. montaguensis (see Table

TABLE I
.-Comparison of some characters of F. elsieue, F. spinosa, F glen ii, F. leucoxantha and F m ontaguensisBothalia 29,1 (1999) , or else the area is called the C ape R egion-phytogeographic region based on the distribution o f vascular plants (Cowling & H ilton-T aylor 1997).T he holotype specim en of F. elsieae.as already m entioned, was collected on Table M ountain.Ii grew on black, hum us-rich soil containing quart/, grains derived from Table M ountain sandstone.Other specim ens referred here are Braggins 97/354 from Silverm ine R eserve, near Kalk Bay as well as Braggins 97/356A from Stinkw ood Trail, Kirstcnbosch.Their spore ornam entation is closely sim ilar to that of the holotype specim en, but som e o f the leaves are apically divid ed into 2 ± triangular lobes.Schelpe 6368 is from B ain's Kloof, Witte River and Perold Hi K oekem oer 3475 is from Tradouw Pass.F. elsieae is distinguished by its fairly large size, by its obovate to oblong, almost entire leaves, as well as by its spores with Ihe distal face having rela-