an papers on all aspects of DONE

pa

, and books etc. sent for review, to: Prof. M. Gibby,
.20A Inverleith Row, Edinburgh, EH3 5LR, UK
E-mail: FernGazette@eBPS.org.uk

FERN GAZ. 18(3):81-90. 2008 81

THE CYTOTAXONOMY OF THE ASPLENIUM STUHLMANNII
COMPLEX (ASPLENIACEAE, PTERIDOPHYTA) IN AFRICA

A. F. BRAITHWAITE MISSOURI BOTANICAL

4 Kendal Drive, Beeston, Nottingham, NG9 3AW AUG ee 2008
(Email: afbraithwaite@btinternet.com)
Key words: Asplenium mantoniae, Asplenium jaundeense, new species, GARDE ¢!B RARY
numbers.

ABSTRACT
A new octoploid species, Asplenium mantoniae, intermediate between two
tetraploids species, A. stuhlmannii and A. jaundeense, is described from West
Africa. The morphology and distribution of members of the complex are
consistent with the view that the octoploid has probably arisen by hybridization
of the two tetraploids and chromosome duplication

INTRODUCTION

Asplenium stuhlmannii Hieron. was first described from tropical East Africa and is
widely distributed from northern Tanzania to the southern Sudan. Similar, though not
identical, material occurs in West Africa and was placed in A. stuhlmannii by Tardieu-
Blot (1953) and Alston (1959) and reported by Manton (1959) to be octoploid with n=
144 chromosomes, based on meiotic analysis. Subsequent cytological examination of
East African material has shown the genuine A. stuhlmannii to be tetraploid with n = 72
chromosomes (Appendix in Braithwaite 1964; Fig.l). Further studies of the two
cytotypes suggest that they are sufficiently distinct morphologically and geographically
to merit recognition as two separate species and the octoploid material from West Africa
is described here as a new species, A. mantoniae. The new species also bears some
resemblance to A. jaundeense Hieron., a tetraploid species (Fig.1) from West Africa,
and its possible relationships to the two tetraploids is discussed.

&

Figure 1. Acetocarmine meiotic squash preparations. A, Asplenium stuhlmannii n= 72,
Somalia (Brittish Somaliland; B, 4. jaundeense n = 72, arrow points to an out of focus
bivalent, Yaunde, Cameroon. Scales = 10um

Species

A. stuhlmannii Hieron

|

|
|
|
|
|

A. mantoniae sp. nov.

| A, jaundeense Hieron.

|
|
|

* means + SD (n = 100)

Locality and origin

- Somalia, collected by Mr Desmond Kelsall (O. S Dept,

| Chromosome number

t

a=Fo

St Andrews) on a hill walking expedition in 1949, ex |

| University Botanic Gardens, St Andrews.

| Kenya, living plant sent to Kew with a consignment of
orchids

| Nigeria, plant raised in Leeds from spores taken from

_ Hambler 513 (BM)

_ Ghana, locality not known, collected by Adams (see

Manton 1959), plant in this study raised from spores —

taken from herb. specimen of Manton’s plant

Cameroon, Yaunde, collected for the author as a living
plant and first established at Kirstenbosch in 1961 then
sent to Kew and subsequently transferred to Leeds

| n=72

|

n= 144
n= 144
n=72

+

-
|

Length

40-9 + 2-49
|

39-7 + 2-0
47-1 + 3°57

45:7 +243

Table 1. Locality, source, chromosome numbers and spore sizes of living plants of the Asplenium stuhlmanii complex.

3414285

Spore size*

|

23-4 + 1-69

Breadth

30:3 + 2-06

26°4 + 1:0

SPE eee,

30-44 2°13
|

8007 06-18 (E)81 “ZVD Nags

BRAITHWAITE: THE ASPLENIUM STUHLMANNII COMPLEX 83

MATERIAL AND METHODS

Live plants for cytological investigation were accumulated from a variety of sources
(see Table 1) and finally established in the Botanic Garden, University of Leeds. In
addition, all the relevant herbarium material at BM and K was examined. Voucher
specimens will be deposited in B

Chromosome counts were made on acetocarmine squash preparations of meiosis
using the standard method for ferns (Manton 1950).

Spore samples for measurement were mounted in either gum chloral or Depex and
measured using a calibrated eyepiece and <40 objective.

THE SPECIES

Asplenium stuhlmannii Hieron.

ngl. Pflanzenw. Ost.-Afr. C: 83, 1895; type: Kassesse, Towalio in Western des Vict.-
Njansa, Stuh/man 936 (B!—lectotype), 953!, 1179, 3246, 3981a (B—paratypes); non
Tardieu-Blot, in Mem. Inst. fr. Afr. noire 28: 190, t. 36, fig. 3, 1953; Alston in The Ferns
and Fern Allies of West Tropical Africa, suppl. 2 ed. Flora of West Tropical Africa, 59,
1959.

Asplenium stuhlmannii var. laciniata Hieron., in Engl. Pflanzen. Ost-Afr. C: 83,

1895; type: Gumango, Lande der Niamniam, Schweinfurth 3915 (B!—holotype;

BM!, K!—isotypes).

Asplenium amoenum C. H. Wright, in Johnstone, The Uganda Prot. 1: 326, 1902,

non Pres! 1836; type: Uganda, E. Toro, R. Kiawiume, alt. 5000’. Doggett s.n. (K!—

holotype).

Rhizome creeping with rather stiff loosely tufted fronds. Rhizome scales 3—S5mm long,
0-5—0:75mm wide at the base, linear subulate with a rounded base and ending in a short
hair point, clathrate with smooth cell walls; median cells dark thick-walled and
compressed with narrow, sometimes obscured, lumens, becoming thinner walled and
pellucid towards the margins. Fronds up to 24—45cm long, pinnate to deeply
bipinnatifid; stipes black becoming green near apex, sparsely scaly. Lamina bright
green, linear lanceolate, 15-37 x 2-5—9-5cm with 10-17 pairs of opposite or
subopposite spreading pinnae. Pinnae 1-8—5-0 = 1-0—2-3cm, coriaceous, broadly trullate
to trullate, base broadly cuneate; basal pinnae deeply divided, upper pinnae less so, into
3—5 lobes; largest lobes (pinnules) cuneate-obovate and further cut into 3 or 4 segments
with truncate lacerate apices, smaller lobes rectangular with truncate lacerate-crenate
apices. Veins and sori subflabelliform. Spores plano-convex, (32—) 36-44 (—50) = (20-)
24-30 (-35) um perispore costate with loosely oe low ridges or folds.
Reproduction sexual, chromosome number n = 7 44).

Ecology and distribution. Asplenium stuhlmannii is a fern of rocky outcrops often
in grassland at altitudes from 850 — 2188m. It grows in sheltered rock crevices or in
shade around the base of rocks or on top of rocks in the shade or less often in full sun.
It is widely distributed in East Africa: Tanzania, Rwanda, Uganda, Kenya, E Zaire,
Sudan and E Central African Republic (Figure 1).

Notes. The name A. stuhlmannii is here restricted to the East African material with
creeping rhizomes bearing scales made up of smooth walled cells, pinnate to deeply
bipinnatifid fronds with broadly trullate to trullate pinnae and spores with costate-alate
spores with rather low smooth folds or wings (Figure 3). All the material is tetraploid
in so far as can be judged from comparisons of spore measurements from specimens in

84 FERN GAZ. 18(3): 81-90. 2008

the herbaria with those from the cytologically authenticated material.

Asplenium stuhlmannii var laciniata with larger pinnae, more deeply dissected into
laciniate lobes probably represents a larger luxuriant shade form. The rhizome scales
and spore ornamentation and size match those of typical A. stuhlmannii.

Specimens examined. ZAIRE: Haut Zaire: Zande Dist.: entre Doruma et Niangoro,
1931, J. Lebrun 3180 (K, BM). SUDAN: Western Equatoria: Yambio Dist.: Mt
Bangenze, in rocky crevice, in shade on hillside, local, 900m, G A.. Prowse 146 (BM);
ibid. on stone outcrop near Mt Bangenze, F’ W. Andrews A1497 (BM); Rest House, on
granite outcrop near Borago, 1939, & W. Andrews A1612 (BM); Mt Ledua near Ibba, J.
G. Myers 6570 (K); Baragu-marshes, 1939, J. W. G Wyld 542 (BM). Maridi Dist..: Azza
Forest, in crevice of rocks in depression forest, tufted frond, local, 850m, 1953, Prowse
153 (BM); near Azza Forest, 1939, 2 W. Andrews A1347 (BM). Eastern Equatoria: Yei
Dist.: Lado, Yei R., F. Sillitoe 110 (K); Kapigo, in pockets of earth on open rocky
hillside especially near bushes, common, 1273m, 1953, G. A. Prowse 33 (BM); Mt
Gumbiri, in Selaginella mats on open rock surface, common, 1500m, 1953, G. A.
Prowse 17 (BM); Mt Kala, rocky crevice and under Euphorbia sp., locally common,
1200m, 1953, G A. Prowse 191 (BM); Imatong Mts, Talanga, 4° 01° N, 32° 45” E,
950m, 1980, /. Friis & K. Vollesen 577, 578 (K). UGANDA: Northern Prov.: West Nile
Dist.: Koboko Hill, damp crevices in granite rocks, 4400ft, R. J. Chancellor 111 (K);
Mt Otzi,, 500ft below summit, 5000 feet, fairly common, 1951, E. M. Cherry 5 (K,
BM). Acholi Dist.: Chua, fern on rocky outcrop at Ukuti, 1935, W. J. Eggeling 2411,
2413 (K); Lotuturu, Chua, Maxwell-Forbes 94 (PRE). Lango Dist.: Maruzi, ground fern
growing in interstices of rocks of Kibuji Hill, 1937, R. G Sangster 321 (BM). Buganda
Prov.: Mubende Dist., Bugangadzi, grows on granitic rocks, 4200ft, Dawe 137 (K).
Mengo Dist., Entebbe, Loe, 1909, R. Fyffe 71 (K). Western Prov.: Toro Dist.: Kyaka,
growing on tree trunk, 4500ft, 1913, J. D. Snowden 150 (K), 1914 (BM); Mwenge,
Oruha Hill, fern in crack of gneiss rock, 1951, H. A. Osmaston 1221 (BM); Kyegegwa,
granitic outcrop, 4200ft, 1933, 4. S. Thomas (K). Ankole Dist.: Igara, rocky outcrop in

+
i ”
H “
a ;
of :

Figure 2. Distributions of Asplenium stuhlmannii (@), A. jaundeense (WB) and A.
mantoniae (4). The locality of A. jaundeense in Gabon not placed.

BRAITHWAITE: THE ASPLENIUM STUHLMANNII COMPLEX 85

short grassland, 5500ft, 1939, Purseglove 577 (K); Nyakalakye-Buyenja, Kashari, 0°
22° S, 30° 31° E, growing in and around rock crevices, 1680m, 1986, PK.
Rwaburindove 2235 (K). Eastern Prov.: Teso Dist.: Kyere, crevices among rocks
forming rocky outcrops, 3650ft, P- Chandler 971 (K). Busoga Dist..: Bulimba Banga,
hill N of Mpumuda, Kibili Rd, growing in shade under large round boulders on hill top,
1950, G A. T. Wood Y2 (K). KENYA: Rift Valley Prov.: Trans-Nzoia Dist.: NE Elgon
foothills, Simpsons rocks, rock crevice in outcrop rock in savannah, 6200ft, Tweedie
1832 (K). Eldoret Dist.: Oldan Sapuk, c. 7000ft, 1951, P J. Greenway 8538 (BM).
Yanza Prov.: North Kavirondo Dist.: S foothills of Elgon above Kimilili, growing in
rock crevices, 5900’, Tweedie 2921 (K). Uasin-Gishu Dist.: Oldan Sapuk, c. 7000ft,
1951, P J. Greenway 8538 (K). Nandi Dist.: Kaimosi Farm, 5800ft, G R. Williams 571
(K). Central Prov.: Fort Hall Dist..: Fort Hall, 1903, R. Meinertzhagen s.n. (BM).
Coastal Prov.: Teita Dist..: Teita Hills, Mwatata R. Valley 5.8km Mwatata-Wundanye
along new alignment, 870—1000m, R. B. Faden 70/443 (K). ? Masai Prov.: Masai Dist..:
Emoli Hill ft, 5900ft, Opiko 326 (K). TANZANIA: Lake Prov.: Mwanza Dist.: coast of
Speke Gulf near Mwanza, moist cracks and hollows below great granite boulders,
3740ft, 1931, B. D. Burtt 2474 (K, BM); Mwanga, fern common on rocky hills near
Lake Mwanga, 19385 (?8), PR. E. Glover 303 (K). Ngara Dist.: Bugarama, Bushubi, on
rock face in crevice, 5000ft, 1961, R. Tanner 5829 (K). RWANDA: Biumba Territory,
Region de Mutara, env. de Mimuli, colline Nyakagenge, anfractuosités dans
affleurements granitiques, 1975, Troupin 3726 (K); ibid., creux ombragé sous roche
surplomb, 1975, Troupin 3267; ibid., affleurements rocheux, 1450 — 1500m, 1975,
Troupin 4212 (K).

Asplenium mantoniae A. Braithw. sp. nov.
Asplenium stuhlmannii et A. jaundeense similis sed differt chromosomate numero (n =
144) et sporis majoribus; a A. stuhlmannii pinnis trapezio-trullatis, basaleis acutis et
lobis paucioribus; a A. jaundeense \obatis profunde trapezioid-trullatis pinnis.
Holotypus: Nigeria, Ondo Province, insel-berg nr Ado Aiye, 7° 54’ N, 3° 30’ E, with
creeping rhizome growing under overhanging boulders, Hambler 513 (BM).
Asplenium stuhlmannii sensu Tardieu-Blot, in Mem. Inst. fr. Afr. noire 28: 190, t. 36,
fig. 3, 1953; Alston in The Ferns and Fern Allies of West Tropical Africa, suppl. and
ed., Fl. W. Trop. Afr., 59, 1959.

Figure 3. Morphology of the Asplenium stuhlmannii complex, spores. A, A
stuhlmannii, Somalia; B, A. mantoniae, Hambler 513; C, A. jaundeense, Yaunde,
Cameroon. Scales: A-C = 10 u

86 FERN GAZ. 18(3): 81-90. 2008

Rhizome creeping with stiff tufted fronds; rhizome scales dark brown, up to 6mm long,
Imm wide, linear-subulate, clathrate, cell walls thick and opaque in the middle,
becoming thinner and pellucid towards the margin, walls facing lumens often
irregularly minutely denticulate (Figure 3). Fronds 15-44cm long, pinnate to deeply
pinnatifid; stipe dark purplish brown 4-5—20cm long with small scales similar to those
of the rhizome at the base; rachis dark purplish brown, for the most part becoming green
towards the apex, sparsely scaly. Lamina 10—24cm long, 3-5—7-Scm wide, narrowly
ovate with 8-13 pairs of subopposite spreading pinnae merging into a lobed apex.
Pinnae 2-0-6°5 x 0-75—2:5cm, narrowly parallelogram-shaped to trapezoid-trullate,
base narrowly cuneate then divided into 3-4 lobes merging into lacerate-serrate,
sometimes attenuate, acute apex (Figure 2); largest basal acroscopic lobe obtriangular
to rectangular and further cut into 3 lobes with irregularly crenate apices, remaining
lobes rectangular with irregularly crenate apices, becoming progressively smaller
towards apex of pinna. Veins and sori subflabellate. Spores plano-convex, (40—) 46 — 48
(-—55) um x (28—) 31-32 (-37) um, costate-alate with much anastomosing high wavy
(undulating) ridges or wings. Reproduction sexual, chromosome number n = 144 (2n =
288).

Ecology and distribution. This new species is generally associated with rocky
outcrops, usually but not exclusively granite, e.g. the inselbergs of Nigeria, where it is
found in the shade on rock faces, in damp rock crevices or growing under overhanging
boulders from 100 — 400m. It has a predominantly West Africa distribution (Guinea,
Sierra Leone, Ivory Coast, Ghana, Nigeria, Cameroon) with outliers in the Central
African Republic (Figure 2).

The new species resembles A. stuh/mannii in its general growth habit, frond
architecture, colour and texture, but is distinguished by its slightly larger rhizome scales
made up of cells with thicker and often dentate cell walls, generally larger but less
divided and more trapezioid or parallelogram-shaped pinnae with narrower cuneate
bases and by its larger costate-alate spores with broader more frequently anastomosing
wavy (undulating) perispore folds or wings. In these respects it resembles the following
species, A. jaundeense, despite being traditionally confused with A. stuhlmannii. (see
Figures 2 & 3).

Etymology. Dedicated to the late Professor Irene Manton F. R. S. in recognition of
her contribution to knowledge of the cytology of the fern flora of West Tropical Africa.
specimens examined. GUINEA: Haut Niger (Fouta Djallon): Mt Bambaya, vers 100m,
1945, P. Jaeger 2113 (K); Macenta, Adams 5513 (P); Gueckedou, Adams 5581 (P).
SIERRA LEONE: Northern Prov., Bumba (?Bumban), 1200ft,1932, R. Glanville 436
(BM). IVORY COAST: Séguéla, rocher granitique a 15km E. sur la route de Beoul,
sous un surplombe humide, 1948, H. des Abbayes 619 (BM); entre Seguela et
Maukono, fissure rocheuse, 1954, R. Schell (K); Rocher d’Issia, in groove on granitic
rock, 250m, 1962, 4. J. M. Leeuwenberg 4140 (K); Pays des Ayoles, sommet du Mont
Kouan prés Aanamé (manelog granitique), 400m, 1909, 4. Chevalier 21269 (K,P); Mt
Dou, Man, Porteres s.n. (P); Mt Semelebou, 4. Chevalier 22095 (P). GHANA:
Bhonso Akai Prov.: Ebaw Rock near Mim, growing in moist crevices on granite
outcrop near thicket margin, 1973, Hall & Abbiw 44559 (K). NIGERIA: Oyo Prov.: hill
3 km N of Iseyin, 7° 58’ N, 3° 34’ E, 350m, rather shady crevice in granitic rocks, 196?,
J. B. Gillett 13399 (K); Okeho, group of low hills about 3 miles from Iseyin road, in
savannah country, 1959, R. W. J. Keay FHI 37748 (K); Ibadan, Oje Rock, sheltered rock
crevices near summit, 1967, D. P- M. Guile 3018 (K). Ondo Prov.: Erio near Aramoko-

BRAITHWAITE: THE ASPLENIUM STUHLMANNII COMPLEX 87
A B -

Figure 3. Morphology of the Asplenium stuhlmannii complex showing silhouettes of
fresh juvenile fronds (top, scale = 5cm), pinnae from lower part of mature dried fronds
(middle, scale = 2 cm) and sections (middle to margin, scale = 0-1mm) of the rhizome
scales (bottom). A, A. stuhlmannii, Somalia; B, A. mantoniae, Hambler 513; C, A.
jaundeense, Yaunde, Cameroon.

88 FERN GAZ. 18(3): 81-90. 2008

Ekiti, base of Hildegardia on steep rock faces, 1968, J. B. Hall 67 (K); ibid., shade of
rock on inselberg, 1968, J. B. Hall 71 (K); Idanre, Jones s.n. (FHI 14846) (BM); among
granite boulders on the lower more or less forested slopes of granite hills, 1946, 4. P
D. Jones 14844 (BM); Idanre, behind Rest House, in rock fractures, texture coriaceous,
dark green, 1968, D. Gledhill 957 (K); inselberg NE of Ado Rock, 7° 51° N, 3° 30’ E,
amongst rubble under overhanging ledge — small quantity only growing with Pellia
doniana, 1958, D. J. Hambler 419 (BM); inselberg near Tapa, 7° 34’ N, 3° 30’ E, under
overhanging boulders growing with Pellia doniana and Sanseviera sp., 1958, D. J.
Hambler 526 (BM). Plateau Province: near Farm Rua between Willsani Camp and
Marhai, soil filled crevices in shady side of rock, 1968, /. B. Hall 659 (K); Sha, 3600ft,
rock crevice in deep shade, scarp slope, dark green erect fern, D. W. Lawlor & J. B. Hall
570 (K). Taraba Prov.: Gembu, Mambilla Plateau, fern up to 25 cm high with grooved
rachis, found growing up between rocks on a wet rocky outcrop behind the Vet. Rest
House, c. 5500ft, 1973, H. M. Chapman 102 (K). CAMEROON: Nord Prov.: 7km S of
Poli, Mango, 1974, G Fotius 2190 (K). CENTRAL AFRICAN REPUBLIC: Bamingui-
Bangoran Prov., Manova-Gounda- St Floris National Park, WWF Inter. Elephant Cons.
Project, 2 km w of Camp Koumbala on Koumbala Creek, 8° 30’ N, 21° 12’ E, 570m
sandstone plants with some outcropping of sandstone ‘buttes’, under shaded overhang
in sandstone, 1983, J. M. Fay 5615 (K). Haut Kotto(Obangui-Chari): Wadda, sur les
rocheurs near Wadda, 1922, G le Testu 4106 (BM).

A, jaundeense Hieron.
Engl. Bot. Jahrb. Syst. 46: 369, 1911; Tardieu. Mem. Inst. fr. Afr. noire 28: 192, t. 36,
fig. 4, 1953; type: Cameroon, an Felsen und Baustammen des Urwaldes bei Jaunde in
800 — 1000 Hohe ii. M., in dem Jahren 1890 — 1892, Zenker 214 (B!—lectotype,
selected here), 1894 — 1895, Zenker 214a (K!, BM!— isolectotypes); 23 Okt 1894,
Zenker & Staudt 526 (B! K!—paratypes); 31 Juli 1897, Zenker 1492 (B, K!—
paratypes); in lichten Waldungen an steinigen abhangen der Lagoo-Berge in 300 Hohe
i. M., 25 Juni 1909, Ledermann 4396 (B—paratype).

Asplenium dimidiatum var. zenkeri Hieron., in Engler Veg. der Erde 9: 28, fig. 24,

908.

Rhizome creeping giving rise to fronds a few millimetres apart; rhizome scales dark
brown, up to 6 x Imm, subulate, clathrate with median cells dark thick-walled and
compressed, often with narrow occluded lumens, marginal cells thin-walled and
pellucid, walls facing lumens distinctly minutely dentate. Fronds up to 66 cm long;
stipes up to 26cm, matt black with scales at the base similar to those of the rhizome.
Lamina up to 41 x 8cm, oblong to linear-oblong with 10 — 20 opposite-subopposite
pairs of spreading pinnae merging progressively into a lobed apex; rachises similar to
stipe but glabrous and becoming green towards the apex. Pinnae up to 6 = 3cm,
trapezoid-rhomboid to cuneiform hastate, base narrowly cuneate, inaequilateral,
acroscopic side up to 1-5cm long, basiscopic side up to 3cm long; largest pinnae
subtrilobed to trilobed, lateral lobes truncate irregularly inciso-dentate, terminal lobe
long, elongate-deltoid, sparsely alternately inciso-dentate, ending in an acute point;
trilobing tending to disappear and pinnae becoming trapezoid-rhomboid towards apex
and in juvenile fronds. Costae not evident, veins and sori subflabellate. Spores plano-
convex, (28—) 33-36 (40) ym x (19-) 22-24 (27) um, perispore costate-alate with
anastomosing high wavy ridges or folds (wings). Reproduction sexual, chromosome

BRAITHWAITE: THE ASPLENIUM STUHLMANNII COMPLEX 89

number n = 72 (2n = 144).

Ecology and distribution. The limited ecological information available suggests that
this species is associated with rocky outcrops or boulders at altitudes ranging from 300
— 1000m. It is known only from West Africa: Gabon (locality not placed), Cameroon
and eastern Nigeria (Figure 2).

Notes. Distinguished from Asp/enium mantoniae by its trapezioid- rhomboid three
lobed pinnae with often long attenuate apical lobe and smaller spores, (28—) 33 — 36
(—40) um x (19-) 22 — 24 (-27) um.

The morphologically similar A. megalura Hieron. differs by its erect rhizome, rather
brittle wiry fronds with fewer and often long-attenuate pinnae, a large 3-lobed apical
pinna and spores with sparsely anastomosing shallow ridges or wings. It also differs
ecologically being usually epiphytic in forest and is widely distributed in tropical and
subtropical Africa.

Specimens examined. NIGERIA: Ogoja Prov.: [kom and Obudu Divs, nr top of 2500ft
hill about pillar no. 30 on the E boundary of Boje enclave, Jones & Onochie s.n. (FHI
No. 18757) (BM). CAMEROON: Milbraed, III Reise nach Afrika 1913-14. Ubergangs
—und Kampfgebiet gegen die Savanna an der Nord-grenze der Hylea siidlich des Sanaga
zwischen Yaunde und Dangdeng umwelt der Vereinigung von Lom (Sanaga) und
Djerem Etwa 68 Km N. O. Jaunde, Feb 1914, Sommer 8718 (K). GABON: Andoun,
rocheres de tete, Jestu 238 (locality not placed) (BM).

DISCUSSION
The three species of the Asplenium stuhlmannii complex are united by the presence of
a creeping rhizome clothed in clathrate scales with dark median band and translucent
margins, and bearing rather stiff once- pinnate narrowly lanceolate or narrowly ovate
fronds. The rhizome scales, pinnae and spores of the three species are shown in Figures
2 & 3.

The clathrate rhizome scales of all three species are made up of dark thick-walled
cells with compressed lumens in the middle parts becoming thinner-walled and
translucent towards the margins. Apart from differences in the size of the cells related
to ploidy, there are subtle differences in the inner surface of the cell walls bordering the
lumens. Those of A. stuhlmanni are smooth while those of A. jaundeense are minutely
denticulate. Those of A. mantoniae are often, but not always, sparsely denticulate
(Figure 2), and thus somewhat intermediate.

The pinnae and spore ornamentation of A. mantoniae are also intermediate between
those of the two tetraploids. The pinnae possess a narrower acute base and are more
trapezoid-rhomboid and less dissected than the broadly trullate pinnae of A. stuhlmannii
and possess a broader cuneate base and are more trulllate with deeper lobes than the
trapezio-rhomboid almost entire pinnae of A. jaundeense. The spores of A. mantoniae
are larger than those of the two tetraploids on account of their higher level of
polyploidy. Their higher more anastomosing wavy perispore folds or wings contrast
with the rather low sparingly anastomosing straight perispore wings of A. stuhlmannii.
In this respect they resemble the spores of A. jaundeense.

The morphological analysis suggests a close relationship between the three
members of the complex and is consistent with the view that A. mantoniae has most
likely arisen by hybridisation between the two tetraploids and subsequent doubling of
the chromosome number of the hybrid to produce the octoploid. The present

90 FERN GAZ. 18(3): 81-90. 2008

distributions of A. jaundeense and A. stuhlmannii are widely separated in West and East
Africa respectively so that such a hypothesis would imply that in the past their
distributions must have either been sympatric or in close contact. The distribution of A.
mantoniae partly falls between those of the two tetraploids, but has also extended the
distribution of the complex anlage in West Africa where it may now be found as
far west as Sierra Leone and Guin

REFERENCES

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BRAITHWAITE, A. F. 1964. A cytotaxonomic investigation on the Asplenium
aethiopicum complex in Africa. Unpublished Ph. D. Thesis, University of Leeds.

MANTON, I. 1950. Problems of cytology and evolution in the pteridophyta.
Cambridge University Press, Cambridge.

MANTON, I. 1959. Cytological information on the ferns of West Tropical Africa. In
ALSTON, A H. G. The ferns and fern allies of West Tropical Africa. Suppl to 2nd
ed. of Flora of West Tropical Africa, pp. 75-81. Crown Agents, London.

TARDIEU-BLOT, M. L. 1953. Les Pteridophytes de |’ Afrique Intertropicale Francaise.
Mem. Inst. Fr. Afr. Noire 28: 1-241.

FERN GAZ. 18(3):91-97. 2008 91

ASPLENIUM DELAVAYI (PTERIDOPHYTA, ASPLENIACEAE)
A FERN SPECIES NEW TO NEPAL

ARUN RIJAL

P.O. Box 4326, Kathmandu, Nepal
(E-mail: arunrijal@yahoo.com)

Key words: Asplenium delavayi, Pteridophyta, west Nepal, disjunct

ABSTRACT
Asplenium delavayi (Franch.) Copel., a Sino-Himalayan species, described from
S.W. China and previously known only as far west as Sikkim, has been
discovered in the Baitadi District of west Nepal. This is much further west than
might be expected and emphasises the possibility of more species being
discovered in Nepal with increased collection and study.

INTRODUCTION

Although Nepal is only c. 885 km from west to east and 145 to 241 km from north to
south, it contains extreme differences in climatic and geographic situation, as a result of
the great range of altitude, from c. 60 m. in the southern terai zone to more than 8000
m. in the high Himalaya. Plant diversity ranges from stunted alpines surviving in harsh
environments in the frozen mountains, to mighty trees of the steamy lowland jungles.
This diverse environment provides habitats for a surprisingly large number of plant
species and more than 6000 species of flowering plants have been recorded so far from
Nepal. It has been estimated that over 6600 species will be listed from the country when
the poorly known remote regions are fully explored (RBGE 2007).

Nepal also has a rich fern-flora with 532 species of pteridophytes reported so far
(Thapa 2002; Fraser-Jenkins & Thapa in prep.). Nepalese pteridophytes belong to two
main phytogeographical elements, with a smaller third element. The dominant group of
species are Sino-Himalayan species, with their centre of diversity in S.W. China, which
have spread westwards along the Himalayan chain and are typically plants of the main
and inner ranges, occurring from mid to higher altitudes, though some may occur in the
lower, outer ranges nearer the plains. Such species usually occur more-or-less
throughout Nepal, though the number of species is higher in the higher rainfall areas of
the east. The second group of species are the S.E. Asian elements, which require more
tropical conditions and high rainfall. They occur predominantly in E. Nepal from lower-
mid to mid altitudes, and although they also extend westwards through Nepal, their
numbers decrease rapidly further westward, particularly towards far-west Nepal. There
is a small second enclave of S.E. Asian species in the locally high-rainfall area around
Pokhara in W.C. Nepal. The third element is a small group of either European or W.
Himalayan species, the latter a special part of the Sino-Himalayan group. These extend
eastwards from the W. Indo-Himalaya into the far-west part of Nepal, with several
occurring mainly behind the Himalayan line in C. Nepal.

Asplenium is one of the larger genera in the flora and contains 32-34 species and an
additional subspecies in Nepal (Thapa 2002, Fraser-Jenkins & Thapa in prep.) of which
19-21 species may be considered Sino-Himalayan elements, eight S.E. Asian elements
and five European elements (Fraser-Jenkins pers. comm., Dec. 2007). Most new

92 FERN GAZ. 18(3):91-97. 2008

records for Nepal concern what is presumably the richest part of the flora in eastern
Nepal. But the central region is by far the best collected due to its greater accessibility
and, as currently known, more species are recorded from central Nepal than further east.
This is likely to be an artificial anomaly resulting from the greater attentions of
collectors there. The least known area is western Nepal, especially the far west. There
is a higher proportion of semi-arid and European and W. Himalayan elements here, but
it is clear that fewer species occur in W. Nepal due to the lower rainfall. However
because the area is so under-collected some surprises occasionally turn up there, with
some species extending further west than might be expected.

In 1997, the author came across an unusual species in far W. Nepal which was not
familiar to him. It had the long, indusiate sori of an Asplenium, but a uniquely rounded,
entire and simple lamina of small size, and a black stipe. This was subsequently
identified in Dec. 2007 by C.R. Fraser-Jenkins as the very rare and little known species,
Asplenium delavayi (Franch.) Copel., which was not previously known from Nepal,
even in the east.

TAXONOMIC TREATMENT
Asplenium delavayi (Franch.) Copel., Genera Filicum: 165 (1947).

Basionym: Scolopendrium delavayi Franch., Bull. Soc. Bot. France, sér. 2, 32: 28
(1885); synonyms: Phyllitis delavayi (Franch.) C.Chr., Index Fil. 2: 492 (1906);
Schafjneria delavayi (Franch.) Tardieu, Naturalist recipe gee 9(1): 30 (1957);
Sinephropteris delavayi (Franch.) Mickel, Brittonia 28(3): 327 (1976).

Type: China, Yunnan: “in montibus, ad Chouang-che-teou, supra Ta-pin-tze, prope
Tali [now Dali]”. Abbé PJM. Delavay 67, 19 Aug. 1883 (P, K).

Distribution: W. Nepal, Sikkim, Bhutan, N.E. India (Manipur), N. Myanmar, S.W.
and S.C. China (Yunnan, Sichuan, Guizhou and Guangxi).

Asplenium delavayi was illustrated by Clarke (1888), who first reported it from
India, Ching (1930), Mickel (1976) and Wu (1999) and is probably related to the
similar, but larger, Chinese and Japanese species, 4. cardiophyllum (Hance) Baker
(syn.: Boniniella cardiophylla (Baker) Tagawa). Although several splinter-genera,
including Sinephropteris, were raised at various times, these are now generally sunk

Figure 1: Map of Nepal showing the location of A. delavayi in Baitadi District.

RIJAL: ASPLENIUM DELAVAYI - A FERN SPECIES NEW TO NEPAL 93

into Asplenium as they all have a recognisably similar can morphology and are
connected by intermediate species to the main-stream of the genus.

Asplenium delavayi was collected by the author in Nepal at Nwaghargad, Baitadi
District, Far-West Development Region (Mahakali Zone); in crevices of damp rock in
open places, 1258 m. alt. (4125 ft.). A. Rijal 3251, 15 January 1997 (KATH). The
distribution of A. delavayi is that of a Sino-Himalayan species, though occurring at
lower altitude.

Description: Rhizome short, unbranched, up to c. 1cm. long, erect, bearing a tuft of
branched, dark-brown roots below, scales on the apex similar to those on the stipes but
slightly smaller, and a tuft of semi-erect, radiating stipes. Stipes cylindrical with an
adaxial groove, blackish-brown to black, glossy, 2.5-5cm. long, erect, bearing a tuft of
small, greyish, narrowly lanceolate, acuminate, slightly toothed scales towards their
bases, c. 2-3mm. long, 0.4-0.7mm. wide towards their bases. Lamina persistent in
Winter, simple, 2-2.6 x 2.1-2.6 cm., glabrous above, bearing a few scattered small
scales beneath, bright green, paler below, entire to slightly wavy at the margin,
markedly rounded-ovate with a shallow cordate base and sometimes a slightly rounded-
pointed apex, a vague darkish midrib present towards the base of the lamina, veins
immersed, inconspicuous, radiating from the midrib, several times dichotomously
forked, with the smallest forks towards the margin, but not reaching it, with occasional
anastomoses near to the margin. Sori c. 20-28 per frond, elongated, straight to slightly
curved, placed along the veins, not forked, obliquely radiating, starting near the midrib
and ending below the margin, larger main sori c. lcm. long, occasional smaller sori
borne between the larger sori from about half way along their length, indusiate;
indusium pale, very thin, becoming brown, lifting and shrivelling somewhat on soral
ripening, opening towards the adaxial (midrib) side, Spores good, regular, bilateral,
with short wings of perispore, c. 28 x 20 um.

The general habit of the fern is similar to a small, more delicate Adiantum reniforme
L. (from Macaronesia and S.W.China), but with the lamina not horizontal as in that
species and the sori elongated and radiating on the lower surface of the frond, not short
and around the edge. It is also similar to a very small Asplenium scolopendrium L.
(from N. America, Europe, Macaronesia, W. Asia, China and Japan), but with round,
not elongated ovate fronds and thin, black stipes.

CONCLUSION
The discovery of this rare species in far-west Nepal where it has been overlooked so far
is probably connected with that area being under-collected. The considerable
disjunction of this population from those further east is not likely to be an actual
botanical reality. It is also likely that further unrecorded species may be found if more
intensive collecting were made, particularly if carried out by specialists familiar with
all the Nepalese pteridophytes who could easily pick out any unfamiliar taxa. Some
other species also show similar disjunctions which may be expected to be only
temporary until further study has been made. At one time there was a considerable gap
in recording between Darjeeling/Sikkim and the W. Himalaya in Kumaon
(Uttarakhand) for the majority of species extending that far west. But since the
pteridophyte-flora of Nepal has become largely known over the latter half of the 20th
Century, to date, the “central Himalayan gap” has largely been filled and can be seen as
an artificial one. A few large disjunctions still remain, these are:
Acrophorus paleolulatus Pic.Serm. - gap from C. Nepal to Kumaon (Chamoli).

94 FERN GAZ. 18(3):91-97. 2008

2-3 mm

Single scale

Structure

Figure 2: Sketch of the general habit, a stipe-base scale and the abaxial surface of
the frond with sori.

RIJAL: ASPLENIUM DELAVAYI - A FERN SPECIES NEW TO NEPAL 95

Actiniopteris radiata (Sw.) Link - gap from E. Nepal to Lumaon (Garhwal).

a tenuisecta (Blume) Tagawa - gap from E. Nepal to Kumaon (Nainital).

Ano reichsteinii Fras.-Jenk. - gap from C. Nepal (Chapagaon, Kathmandu;

scsi saan doubtful) to Kumaon (Nainital).

Asplenium delavayi (Franch.) Copel. - gap from Sikkim to far-west Nepal (Baitadi).

Athyrium otophorum (Miq.) Koidz. - gap from N.E. India (Meghalaya) to Kumaon

(Pithoragarh).

erate roseum Christ - China; Darjeeling (Fraser-Jenkins) and gap to W.C. Nepal

(below Annapurna Base-Camp. Fraser-Jenkins

eee quadripinnatifida M.Kato - gap sn C. Nepal to Kumaon (Chamoli).
Deparia lancea (Thunb.) Fras.-Jenk. - gap from C. Nepal (Sankhu) to ?Kumaon

(Pithoragarh. Punetha & Kholia).

Plate 1 & 2: Asplenium delavayi from Baitadi, general view.

96 FERN GAZ. 18(3):91-97. 2008

Dryopteris pulvinulifera (Bedd.) Kuntze - gap from E. Nepal to Kumaon (Nainital).
Lepisorus sublinearis (Baker ex Takeda) Ching - gap from W.C. Nepal (Pokhara) to
Kumaon (Nainital).
Lindsaea commixta Tagawa - gap from N.E. India (Assam) to W.C. Nepal (Andheri
Khola, N. of Tansen; locality sometimes given on label as Mussoorie in error).
Matteuccia intermedia C.Chr. - gap from W.C. Nepal (Annapurna Base Camp. Fraser-
Jenkins) to Kumaon (Pithoragarh).
Microlepia hallbergii (d’ Almeida) C.Chr. - gap from C. Nepal (Sankhu) to Kumaon
(Garhwal).
Microsorum zippelii (Blume) Ching - gap from far E. Nepal to Kumaon (Chamoli).
Pteris kathmanduensis Fras.-Jenk., in prep. - gap from C. Nepal (Kathmandu) to
Kumaon (Pithoragarh)
Pteris vittata L. subsp. vermae Fras.-Jenk. - se from China, Tibet and ?Bhutan
(Thimphu. Fraser-Jenkins) to Kumaon (Nainital
Thelypteris Se lebeufii (Baker) bectanihi - gap from N.E. India (Assam State)
to Kumaon pur. Fraser-Jenkins).
Trichomanes peace ih (Baker) Copel. - gap from Myanmar to W.C. Nepal (Gorkha).
Trichomanes saxifragoides C.Presl - gap from Bhutan to W.C. Nepal (Pokhara).

It is to be hoped that future work, concentrating on Nepal, may continue to make the
full distribution patterns of Himalayan pteridophytes clear and other new records may
be discovered.

Plate 3: Single frond (abaxial surface) showing the arrangement of the sori.

RIJAL: ASPLENIUM DELAVAYI - A FERN SPECIES NEW TO NEPAL 97

ACKNOWLEDGEMENTS
The author wishes to thank Mr. C.R. Fraser-Jenkins, of Naya Bazaar, Kathmandu,
Research Associate of the Royal Botanic Garden Edinburgh, for helping in
identification of the collection, revision of this paper and providing information on
disjunct species.

REFERENCES

CHING, R.C. 1930. Icones Plantarum Sinicarum 1: ¢. 26.

CLARKE, C.B. 1888. On the Plants of Kohima and Muneypore. J. Linn. Soc. (Lond.),
Bot., 25: [1-] 93 [-101] er tt, t. 41.

MICKEL, J.T. 1976. Sinephropteris, a new genus of scolopendrioid ferns. Brittonia
28(3): 326-328.

ROYAL BOTANIC GARDEN EDINBURGH (RBGE) 2007. Plants of Nepal: Nepal
and its plants. Webpage of Royal Botanic Garden Edinburgh
(http://www.floraofnepal.org/, June 17 2007).

THAPA, N. 2002. Pteridophytes of Nepal. Department of Plant Resources (DPR),
Ministry of Forest and Soil Conservation (MFSC), Thapathali, Kathmandu, Nepal.

WU, S.H. (1999). Aspleniaceae, in Flora Reipublicae Popularis Sinicae etc. 4(2): [1-]
143-145 [-265].

98 FERN GAZ. 18(3). 2008

SHORT NOTE

CORRECTION TO A REVISED HANDBOOK TO THE FLORA OF
CEYLON, VOLUME XV, PARTS A & B, FERNS AND FERN-ALLIES,
2006. M.D. creer (General Editor), Monika Shaffer-Fehre (Editor).
ISBN, SET 1-57808-411-3

Monika Shaffer-Fehre, The Herbarium, Royal Botanic Gardens , Kew,
ichmond, Surre
(m.shaffer-fehre@rbgkew.org)

These volumes devoted to ferns and fern allies contribute to the Flora of Ceylon.
Unfortunately some errors have been identified.

The following corrections are required to the contents: the family Dryopteridaceae
(Vol. A) is treated in two chapters rather than one only, and Polypodiaceae begins
Volume B. Regarding the Woodsiaceae, the name of the senior author is missing in
‘Contents’; three of the four genera treated, viz.: Hypodematium, Deparia and Athyrium
have been described by C.R. Fraser-Jenkins and one, Diplazium, has been contributed
by Dr M. Zink. There is one further omission: Vol. B, page 356: PTERIDACEAE, B.
Verdcourt. The following text is missing from Hemionitis tomentosa at the very end of
the treatment:

Specimens Examined. KANDY DISTRICT: Above Kandy: Roseneath Valley, The
Heremitage, Sledge 1139 (BM, K); Abatage on Gampola-Pusselawa road, Faden &
Faden 76/246 (K); Peradeniya, Matthews s.n.; cultivated at Kew from Ceylon material
in 1954 (K).

The editor apologises for these mishaps, and a correct list of contents is given on the
next page. The editor would appreciate hearing from readers of any other mistakes. A
list of additional corrections will then be published at a later date.

FERN GAZ. 18(3). 2008

FOREWORD
ACKNOWLEDGEMENTS

99

INTRODUCTION AND SURVEY OF TAXA WITH REFERENCES AND

GENERAL LITERATURE by M. Shaffer-Fehre and P.J. Edwards

li—-xx

KEY to the FERNS and FERN-ALLIES of SRI LANKA by PJ. Edwards xxi-xxix

VOLUME 15 PART A

ASPLENIACEAE by D.Philcox
AZOLLACEAE by Jennifer M. Ide
BLECHNACEAE by P. Jayasekara
CYATHEACEAE by D. Philcox
DAVALLIACEAE by H. P. Nooteboom
DENNSTAEDTIACEAE by M. J. Zink
DRYOPTERIDACEAE | by C. R. Fraser-Jenkins
DRYOPTERIDACEAE 2 by M. J. Zink
EQUISETACEAE by B. Wadhwa
GLEICHENIACEAE by Monika Shaffer-Fehre
GRAMMITIDACEAE by B. S. Parris
HYMENOPHYLLACEAE by P. Jayasekara
ISOETACEAE by B. Wadhwa
LOMARIOPSIDACEAE by M. J. Zink
LOXOGRAMMACEAE by M. G Price
LYCOPODIACEAE by D. Philcox
MARATTIACEAE by D.M.U.B. Dhanasekara
MARSILEACEAE by Jennifer M. Ide
OLEANDRACEAE by B. Verdcourt
OPHIOGLOSSACEAE by D. Philcox
OSMUNDACEAE by P. Jayasekara
PARKERIACAE by B. Wadhwa

VOLUME 15 PART B

POLYPODIACEAE by P. H. Hovenkamp
PSILOTACEAE by B. Wadhwa
PTERIDACEAE by C. R. Fraser-Jenkins, B. Verdcourt & T. G Walker
SALVINIACEAE by Jennifer M. Ide
SCHIZAEACEAE by D. Philcox
SELAGINELLACEAE by D. Philcox
THELYPTERIDACEAE by Monika Shaffer-Fehre
VITTARIACEAE by Monika Shaffer-Fehre
WOODSIACEAE by C.R. Fraser-Jenkins and M. J. Zink

New Names published in this Volume

List of endemic species

Selective Glossary of Terms

Index to Scientific Names

305-307

100 FERN GAZ. 18(3). 2008

SHORT NOTE

TWO NEW SPECIES OF SELAGINELLA SUBGENUS HETEROSTACHYS
(SELAGINELLACEAE) FROM THE GUIANAS - A CORRECTION

In the recent paper by Cremers & Boudrie (2007), the two names are ascribed to
Valdespino ex Cremers & Boudrie. However, in referring to the descriptions, Cremers
& Boudrie state that “what follows” was extracted verbatim from Valdespino’s
monograph. As both the names and the descriptions are therefore ascribed to
Valdespino, under Art. 46.2 the correct attribution of the names should be:

Selaginella | gynostachya Valdespino in Fern Gaz. 18: 42[-46]. 2007
Selaginella karowtipuensis Valdespino in Fern Gaz. 18: 46[-51]. 2007.

I thank Professor John McNeill for advice.

CREMERS, G. & BOUDRIE, M. 2007. Two new species of Selaginella sugbenus
Heterostachys (Selaginellaceae) from the Guianas. Fern Gazette 18(2):41-52.

Mary Gibby

FERN GAZ. 18(3): 101-109. 2008 101

MORPHOTYPE AND CONFLICTING TAXONOMIES IN PTERIDIUM
(DENNSTAEDTIACEAE: PTERIDOPHYTA)

JOHN A. THOMSON

National Herbarium of NSW, Botanic Gardens Trust, Mrs Macquaries Road, Sydney,
NSW 2000, Australia.
(Email: johnt@accsoft.com.au)

Key words: Dennstaedtiaceae, bracken, genetics, morphology, morphotype,
polymorphism, Pteridium, subspecies, taxonomy.

ABSTRACT

Conflicting taxonomic treatments of the diploid bracken ferns of the Laurasian

lineage within Preridium are briefly reviewed. The infraspecific entities

recognised in this section of the genus can be separated on morphological,

phenological and physiological criteria into two main groups referred to

respectively as ‘aquilinum’ and ‘latiusculum’ morphotypes. Evidence of gene

flow between morphotypes is discussed, including the finding from DNA studies

that taxa of the ‘aquilinum’ and ‘latiusculum’ morphotypes in each major

geographic region are more closely related to the contrasting morphotype from

the same region than to the corresponding morphotype from another region. It is

proposed here that the ‘aquilinum’ and ‘latiusculum’ morphotypes may be

regarded as mediated by a simple genetic polymorphism involving alternative

pleiotropic alleles at a single major morphogenetic locus comparable to those

previously described in Athyrium and postulated in Asplenium and Pteridium,

but other possible models are discussed.

It is concluded that P. aquilinum is best treated as a single species containing at

present 11 subspecies.

INTRODUCTION

A vigorous ongoing polemic continues to cloud taxonomic treatment of the ar ]
= 104; Wolf ef al., 1987,1988) bracken ferns [Pteridium aquilinum (L.) Kuhn]
comprising a Laurasian lineage within the genus. Thomson and colleagues fame.
2004; Thomson et al., 2005; Thomson et al., 2008; contra Thomson, 2000) regard P
aquilinum as a grouping of 11 subspecies (Table 1) that in general correspond to
varieties within Tryon’s (1941) P. aquilinum subsp. aquilinum. In contrast, certain other
contemporary classifications are based on separation of these taxa into two phenotypic
groups designated here as the ‘aquilinum’ and ‘latiusculum’ morphotypes (Table 1).
Page (1997) and coauthors (Page & Mill, 1995; Gureyeva & Page 2005) regard these
‘aquilinum’ and ‘latiusculum’ morphotypic groups as multi-species complexes
represented in Europe by P. aquilinum and by P. pinetorum C.N. Page & R.R. Mill
respectively, each with several subspecies. Fraser-Jenkins (1997), on the other hand,
recognises P. Jatiusculum (Desv.) Hieron. ex R.E.Fr. at species level and accords
subspecific rank to P Jlatiusculum subsp. pinetorum (C.N. Page & R.R. Mill)
Fraser-Jenk. Stace (1997) and Karlsson (2000) group the European ‘aquilinum’
phenotypes as P. aquilinum subsp. aquilinum and those of ‘latiusculum’ as P. aquilinum
subsp. /atiusculum (Desv.) Hultén. Shorina & Perestoronina (2000) distinguish the
European ‘latiusculum’ morphotype as P. aquilinum var. pinetorum (C.N. Page & R.R.

102 FERN GAZ. 18(3): 101-109. 2008

Table 1: Geographic distribution and current subspecific nomenclature of northern
hemisphere diploid (2n = 104) bracken taxa indicating ‘aquilinum’ or ‘latiusculum’
morphotype, together with Tryon’s (1941) varietal names for reference.

Geographic region/ P. aquilinum subspecies P. aquilinum variety
h

morphotype
Europe
‘aquilinum’ aquilinum aquilinum
‘latiusculum’ pinetorum (C.N. Page & R.R. — latiusculum (Desv.) Underw.
Mill) J.A. Thomson!” ex Heller
Asia
‘aquilinum’ wightianum (Wall. ex J. wightianum (J. Agardh) R.
Agardh) Shieh* Tryon
*latiusculum’ Japonicum (Nakai) A. Love & — latiusculum (Desv.) Underw.
D. Léve'
Africa
‘aquilinum’ capense (Thunb.) C. Chr.* aquilinum
‘latiusculum’ centrali-africanum Hieron. ex africanum Bonap.
RE. Fr.’
North America
‘latiusculum’ feei (W. Schaffn. ex Fée) J.A. _feei (W.Schaffn. ex Fée)

Hawaiian Islands
‘intermediate’

Thomson, Mickel & K
Mehitreter”

latiusculum (Desv.) Hultén
auct. Thomson (2004)!
pseudocaudatum (Clute)
Hultén

pubescens (Underw.) J.A.
Thomson, Mickel& K.
Mehltreter”

decompositum (Gaudich.)

Lamoureux ex J.A. Thomson!

Maxon ex Yuncker
latiusculum (Desv.) Underw.
ex Heller

pseudocaudatum (Clute)

pubescens Underw.

decompositum (Gaudich.) R.
Tryon

' Thomson, 2004

* Also referred to as P aquilinum subsp. latiusculum (Desv.) Hultén; P. aquilinum
var. pinetorum (C.N. Page & R.R. Mill) Perestor.; P. pinetorum C.N. Page &
R.R. Mill; P. latiusculum subsp. pinetorum (C.N. Page & R.R. Mill) Fraser-Jenk.

> Also referred to as P revolutum (Blume) Nakai

* Thomson et al., 2005
> Thomson et al. 2008

THOMSON: CONFLICTING TAXONOMIES IN PTERIDIUM 103

pe fei!

dearth of useful discriminatory characters often leads to difficulties in
ents etfs taxonomy, especially in the case of low order taxa that may be separated
on as few as one single character (Perrie & Brownsey, 2005). In many fern species the
segmental units represented at successive levels by frond, pinna, pinnule, and even
pinnulet, form a repetitive hierarchical series for which a feature expressed at one level
may not be independent of a corresponding feature scored at another segmental level.
In general, such attributes should be treated as relating to a single character, and may
well share a common genetic basis. Examples are evident in the morphometric
documentation of the relative number, size, shape and spacing of pinnae, pinnules and
pinnulets in subspecies of Pteridium aquilinum (L.) Kuhn (Thomson et. al., 2005;
Thomson ef. a/., 2008). Further, many commonly used features of indumentum, false
indusium and laminal segmentation are extremely labile, being affected by age,
shading, water-logging and other environmental conditions (Ashcroft & Sheffield,
1999; Karlsson, 2000; Thomson ef a/., 2005; Thomson et al., 2008) and must be
employed quantitatively and with caution.

Three recent re-evaluations of disputed taxonomic treatments in ferns highlight the
contribution that genetic evidence, either directly from breeding experiments or
indirectly from molecular data, may make to more consistent and objective assignment
of taxonomic rank. In Athyrium distentifolium Tausch ex Opiz (Woodsiaceae),
morphotype ‘flexile’ is endemic to Scotland and is found only at localities where
‘distentifolium’ is also present (McHaffie et a/., 2001). Sporophytes of the ‘flexile’
phenotype are distinguished from ‘distentifolium’ by frond morphology, position of sori
and response to nutrients. The two phenotypes are controlled by alleles of a single major
gene with pleiotropic effects expressed in both gametophyte and sporophyte, the
‘flexile’ morphotype being recessive to ‘distentifolium’. In one natural population the
frequency of the recessive allele was about 0.4. The two morphotypes were originally
described as separate species but McHaffie and colleagues recommend a taxonomic
status not higher than variety.

In Asplenium hookerianum Colenso (Aspleniaceae), morphotype ‘hookerianum’
occurs in Australia and New Zealand and is distinguished from the endemic New
Zealand morphotype ‘colensoi’ which has _ consistently narrower pinnules. Stands in
New Zealand commonly comprise morphotype ‘hookerianum’ alone or co-occurring
with ‘colensoi’: ‘colensoi’ by itself is rare (Perrie & Brownsey, 2005). The two forms
were originally described as separate species. Neither nuclear genomic fingerprinting
by AFLP nor the chloroplast sequence results separated the specimens according to
morphotype. Geographic source accounts for much more of the total DNA-sequence
variation than pinnule morphology in both the AFLP data (29% of genetic variation
versus 0%) and the chloroplast haplotype data (58% of genetic variation versus 3%;
Perrie & Brownsey, 2005). Plants of contrasting morphotype originating from close
sites are more similar to each other genetically than to plants of the their own
morphotype from distant sites. The phenotypic stability of plants of the two
morphotypes even when growing intertwined in the field argues against a major role for
environmental variables, and ‘colensoi’ is purported to breed true from spores (Perrie &
Brownsey, 2005). It appears probable that morphotypic variation in Asplenium, like that
in Athyrium, results from a simple genetic polymorphism based on allelic variation at a
single locus. Perrie & Brownsey (2005) conclude that there is no significant genetic
discontinuity between the two morphotypes, and no justification for separating them at

104 FERN GAZ. 18(3): 101-109. 2008

ig i varietal or form rank.

n P aquilinum in North America, the status of the two eastern taxa now
nem subsp. latiusculum and subsp. pseudocaudatum (Table 1) proved
contentious until detailed analyses of morphology and isozyme complements were
made by Speer & Hilu (1999) and Speer ef al. (1999). These taxa are clearly distinct on
morphological and distributional grounds (Speer & Hilu, 1999; Thomson et al., 2008).
Over their main east-coast range, ‘latiusculum’ is more abundant in the northern states,
“‘pseudocaudatum’ in the south (Speer & Hilu, 1999), with only very few narrow zones
of overlap where intermediates suggesting introgression are seen. Detailed isozyme
analyses showed unrestricted gene flow between the two morphotypes at one such zone
of co-occurrence (Speer et al., 1999). Speer and colleagues postulate that the
‘latiusculum’ and ‘pseudocaudatum’ phenotypes may be controlled by alternative
alleles at a single genetic locus with pleiotropic effects. If the ‘latiusculum’ phenotype
is dominant and determined by an allele with high frequency in the north of its range,
while the recessive allele for the ‘pseudocaudatum’ phenotype is at high frequency or
fixed in the south, the roughly equal frequencies of the two forms observed in the North
Carolina piedmont would correspond to a frequency of about 0.7 for the recessive allele
(Speer et al., 1999). Strong selective pressure in relation to environmental features and
habitat preference are indicated, with a taxonomic ranking of subspecies for the two
morphotypes (Thomson ef a/., 2008).

aeniiaarinig testinal OF GENERALISED ‘AQUILINUM’ AND
*‘LATIUSCULUM’ MORPHOTYPES IN PTERIDIUM
Characters that reflect the relative number, relative size, spacing and shape of laminal
segments at blade, pinna, pinnule and pinnulet levels (Thomson, 2000; Thomson et ai.,
2005; Thomson et al., 2008), are particularly effective in discriminating a
‘latiusculum’-like group of taxa from an ‘aquilinum’-like phenotypic group.
The ‘aquilinum’ morphotype: Blade long, ovate to linear, typically 2-3 times the
length of the longest pinna which is generally one of the third to the fifth pair from the
lamina base towards the tip. Pinnae, pinnules (and less markedly pinnulets), relatively
long and narrow; typically 13-18 pinnules in the basal half of the longest pinna. Pinnae
and pinnules commonly inserted on rachis and costa at 70-90°. Frond expansion
gradual, progressing from base to apex. Basal 1-2 pinna pairs often marcescent before
expansion of the distal pinnae is complete. Frond axes relatively thick and fibrous;
dense stands leave heavy, often partially upright, litter in winter that is slow to collapse
next season.
The ‘latiusculum’ morphotype: Blade short, broadly triangular, typically 1-1.5 times
the length of the longest pinna which is generally one of the first or second pair from
the base towards the tip. Pinnae, pinnules (and less markedly, pinnulets) relatively short
and broad; typically 6-9 pinnules in the basal half of the longest pinna. Basal segment(s)
of each order sometimes much reduced, even vestigial. Pinnae and pinnules commonly
inserted on rachis and costa at 45-70°. Frond expansion rapid, often almost
synchronous, although progressive from base to apex. Basal pinna pairs not marcescent
before frond expansion is complete. Frond axes relatively thin, not heavily fibrous;
dense stands leave thin litter in winter that collapses readily next season
The ovate to linear form of the expanded blade and pinna es ‘aquilinum
morphotypes results from both the shorter length of the lowest pinna and pinnule pair(s)
compared with those placed more centrally and the relatively higher number of these

THOMSON: CONFLICTING TAXONOMIES IN PTERIDIUM 105

segments on the frond axis and costa respectively. The broadly triangular frond blade
and pinna of the ‘latiusculum’ morphotype could be envisaged as derived from the
‘aquilinum’ type by loss of pinna and pinnule pairs from the base of blade and pinna
upwards, a process which would at the same time increase the apparent length, relative
to more distal segments, of the pinnae/pinnules which thus become the basal or near
basal components. Overall the ‘latiusculum’ morphotype appears to reflect adaptation
to conditions providing a short growing season.

re zones of contact or parapatry occur locally, as in Europe (Rumsey ef al.,
1991; Karlsson, 2000), NE Asia (Tryon, 1941), and Africa (Verdcourt, 2000), recurrent
field observations of morphological intermediates provide evidence of recent, probably
ongoing, interbreeding between ‘aquilinum’ and ‘latiusculum’ morphotypes.
Multi-locus isozyme marker studies also reveal gene flow between multiple genets
(Bridges et al., 1998) of subsp. pinetorum in Scotland and local populations of subsp.
aquilinum (Rumsey et al., 1991).

DNA fingerprinting using the Arbitrarily Primed Polymerase Chain Reaction (AP-
PCR) and Inter Simple Sequence Repeat (ISSR) procedures show that genomic
similarity is higher between ‘aquilinum’ and ‘latiusculum’ morphotypes from the same
geographic region than either is with morphotypes of its own group from other regions
(Thomson et al., 2005; Thomson ef a/., 2008). Thus subspp. capense and centrali-
africanum from sub-Saharan Africa are more similar genomically than capense is to
aquilinum of Europe or centrali-africanum is to the N American /atiusculum (Thomson
et al., 2005; Thomson ef al., 2008). Similarly, subspp. pinetorum and aquilinum of
Europe share more genomic markers than pinetorum shares with /atiusculum of North
America (Thomson, 2000; Thomson ef al., 2005). Chloroplast genome haplotype for
the rps4-trnS region based on presence or absence of one or both of two 5-base direct
repeats is not concordant with the ‘aquilinum’ or ‘latiusculum morphotypic groupings
(Thomson et al., 2005; Thomson et al., 2008). For example, the haplotypes (Type A or
B, Thomson ef al., 2005) of three geographically contiguous ‘latiusculum’ and
‘aquilinum’ morphotype pairs are: subspp. pinetorum/aquilinum (Europe) respectively
Types A and B, subspp. centrali-africanum/capense (Africa) both Type B, and subspp.
Japonicum/wightianum (NE Asia) both Type A.

hese findings provide strong evidence against a taxonomy based on separation of
‘latiusculum’ and ‘aquilinum’ clades (Thomson ef a/., 2008), and in particular argue
against recognition of ‘latiusculum’ or ‘aquilinum’ as reproductively-separated
groupings that individually merit full species status. Within Preridium aquilinum as
recognised here (Table 1), reticulate phylogenetic relationships overlie evolutionary
trends presumably based on diversification and local adaptation.

IS THE ‘AQUILINUM’ VERSUS ‘LATIUSCULUM’ DISTINCTION A SIMPLE
GENETIC POLYMORPHISM?
Descriptively, the morphological polymorphisms analysed in Athyrium (McHaffie et
al., 2001) and postulated in both Asplenium (Perrie & Brownsey, 2005) and Pteridium
(Speer et al., 1999) behave as if based on variant alleles of major genes, with allele
frequencies being maintained by balancing selection pressures in particular
environments and/or differentially in the gametophyte or sporophyte stage. The
contention that the ‘aquilinum’ versus ‘latiusculum’ contrast also represents a genetic
polymorphism is supported by distributional data (Table 1), the genetic relationships of
the contrasting morphotypes in areas of overlap, and evidence that laminal segment

106 FERN GAZ. 18(3): 101-109. 2008

morphology at frond, pinna, pinnule and even pinnulet levels is hierarchically repetitive
and likely to be under control of a common gene or genes with alternative, apparently
pleiotropic, alleles. Definitive characterisation of the genetic basis of morphotype in
bracken must await integrated breeding and molecular analyses. The loci involved are,
however, clearly distinct from those responsible for the small-scale morphological
variants often collected and cultivated by growers (see, for example, Andersson, 1927)
that are rare in natural populations, with very low allele frequencies maintained by
recurrent mutation opposing negative selection (McHaffie er a/., 2001).

While for descriptive purposes morphotypic variation in the ferns discussed here
may be adequately treated in terms of single locus polymorphisms with pleiotropy,
functional and structural molecular analyses of the loci responsible are now required to
reveal how the underlying developmental process are controlled. In general, the more
numerous and diverse the multiple phenotypic effects involving disparate
morphological and physiological characteristics that appear to be under single gene
control, the more an explanation invoking pleiotropy becomes questionable. In the
present cases we therefore need to consider other possible models of gene structure such
as that exemplified by the complex S locus responsible for self-incompatibility
(SI)/compatibility (SC) in a number of families of flowering plants such as
Brassicaceae. This super-locus provides co-ordinated unitary genetic control over
multiple aspects of SI mediated by tightly linked co-adapted sequence segments
collectively forming an S haplotype (Charlesworth et al., 2005). Multiple
developmental, cellular, physiological and biochemical processes in different S
phenotypes are mediated by the alternative S haplotypes represented in polymorphic
populations or in different taxa (Bechsgaard et al., 2006; Sherman-Broyles et al., 2007).
The several features characterising each S$ phenotype are controlled by distinct
sequence elements and do not involve pleiotropy. Where aspects of phenotype are
controlled by a complex super-locus or through a stepwise multigene pathway, multiple
mutations may appear as a single event if a mutation acting at an early step blocks
expression of accumulated changes effective later in the pathway. If features
distinguishing allopatric morphotypes are subject to selection, the allelic variants

THOMSON: CONFLICTING TAXONOMIES IN PTERIDIUM 107

CONCLUSIONS

In most taxonomic investigations of ferns, especially where multiple taxa must be
considered, controlled breeding experiments of the kind so admirably applied in
Athyrium (McHaffie et al., 2001) are not practicable. More widely feasible are genomic
comparisons based on methodologies such as AP-PCR (RAPD), AFLP, ISSR, or
microsatellite analyses. If plants of taxon A from locality 1 are more closely related
genetically to plants of taxon B from locality 1 than to taxon A plants from localities

...n, and/or a comparable situation holds for the relationships of taxon B, there is prima
facie evidence of recent and/or ongoing gene flow between the taxa that is likely to be
inconsistent with the degree of reproductive isolation normally associated with full
specific status. Rather, such a situation strongly suggests that the taxa concerned should
be accorded infraspecific rank, for example as subspecies, varieties, or forms (Perrie &
Brownsey, 2005; Thomson ef al., 2008).

The work of McHaffie and colleagues on the ‘distentifolium’/‘flexile’
polymorphism in Athyrium represents an extraordinary contribution to our
understanding of fern morphogenetics by showing how morphological and
physiological studies of the gametophyte as well as the sporophyte may reveal at these
two life stages ‘antagonistic pleiotropy’ (Shaw, 2001) that determines allele frequencies
in natural populations through otherwise unsuspected opposing positive and negative
selective pressures. However, as pointed out above, questions remain open regarding
the structure of the genetic loci involved in the control of fern morphotype, and thus on
the strict applicability of the term pleiotropy in these cases.

P. aquilinum appears at present to be best treated as a single species containing 11
subspecies as listed in Table 1. Further morphometric and genetic studies are needed to
clarify the status of a number of named taxa described from many parts of Eurasia
which on present evidence should, if named, be ascribed varietal, form or ecotype
rankings (Stace, 1997)

ACKNOWLEDGEMENTS
I thank Elizabeth Brown (NSW) and Barbara Parris (Fern Research Foundation,
Kerikeri) for helpful discussion and comments. Miguel Garcia (NSW) provided
generous library assistance. I am indebted to Paul Wolf (Utah State University) for a
constructive critical review leading to substantial improvement of the final manuscript.

REFERENCES

ANDERSSON, I. 1927. Notes on some characteristics in ferns subject to Mendelian
inheritance. Hereditas 9: 157-179.

ASHCROFT, C.J. & SHEFFIELD, E. 1999 Rejection of Preridium aquilinum
subspecies atlanticum (C. N. Page). Bot. J. Linn. Soc. 130: 157-170.

BECHSGAARD, J.S., CASTRIC, V., CHARLESWORTH, D., VEKEMANS, X. &
SCHIERUP, M.H. 2006. The transition to self-compatibility in Arabidopsis thaliana
and evolution within S-Haplotypes over 10 Myr. Mol. Biol. Evol. 23: 1741-1750.

BRIDGES, K.M., ASHCROFT, C.J. & SHEFFIELD, E. 1998. Population analysis of
the type localities of some recently recognised taxa of British Pteridium
(Dennstaedtiaceae: Pteridophyta). Fern Gaz. 15 :205-213.

CHARLESWORTH, D., VEKEMANS, X., CASTRIC, V. & GLEMIN, S. 2005. Plant
self-incompatibility systems: an evolutionary perspective. New Phytol. 168: 61-69.

FRASER-JENKINS, C. R. 1997. New species syndrome in Indian pteridology and the

108 FERN GAZ. 18(3): 101-109. 2008

ferns of Nepal. International Book Distributors, Dehra Dun.

GUREYEVA, I.I. & PAGE, C.N. 2005. Towards the problem of bracken taxonomy in
Siberia. Taxonomy Notes of the Krylov Herbarium of Tomsk State University 95:
18-26.

KARLSSON, T. 2000. Preridium. In: JONSELL, B. (Ed.) Flora Nordica, vol. 1, pp.
43-47. Royal Swedish Academy of Sciences, Stockholm.

MCHAFFIE, H.S., LEGG, C.J. & ENNOS, R.A. 2001. A single gene with pleiotropic
effects accounts for the Scottish endemic taxon Athyrium distentifolium var. flexile.
New Phytol. 152: 491-500.

PAGE, C.N. 1997. The ferns of Great Britain and Ireland, 2nd ed. Cambridge
University Press, Cambridge.

PAGE, C.N. & MILL, R.R. (1995) The taxa of Scottish bracken in a European
perspective. Bot. J. Scotl. 47: 229-247.

PERRIE, L.R., & BROWNSEY, P.J. 2005. Genetic variation is not concordant with
morphological variation in the fern Asplenium hookerianum sensu lato
(Aspleniaceae). Amer. J. Bot. 92: 1559-1564.

RUMSEY, F.J., SHEFFIELD, E. & HAUFLER, C.H. 1991. A re-assessment of
Pteridium aquilinum (L.) Kuhn in Britain. Watsonia 18: 297-301.

SHAW, A.J. 2001. Antagonistic pleiotropy and the evolution of alternate generations.
New Phytol. 152: 365-368.

SHERMAN-BROYLES, S., BOGGS, N., FARKAS, A., LIU, P., VREBALOV, J.,
NASRALLAH, M.E. & NASRALLAH, J.B. 2007. S Locus genes and the evolution
of self-fertility in Arabidopsis thaliana. Plant Cell 19: 94-106.

SHORINA, N.I. & PERESTORONINA, O.N. 2000. Taxonomic studies of Russian
bracken I. Taxonomy of Pteridium in territories of European Russia, Crimea and
Caucasus. In: TAYLOR, J.A. & SMITH, R.T. (Eds.) Bracken fern: toxicity, biology
and control, pp. 48-51. International Bracken Group, Aberystwyth.

SPEER, W.D. & HILU, K.W. 1999. Relationships between two infraspecific taxa of
Pteridium aquilinum (Dennstaedtiaceae). I. Morphological evidence. Syst. Bot. 23:
305-312.

SPEER, W.D., WERTH, C.R. & HILU, K.W. 1999. Relationships between two
infraspecific taxa of Pteridium aquilinum (Dennstaedtiaceae). II. Isozyme evidence.
Syst. Bot. 23: 313-325.

STACE, C.A. 1997. Field flora of the British Isles, 2nd ed. Cambridge University
Press, Cambridge.

THOMSON, J.A. 2000. Morphological and genomic diversity in the genus Pteridium
(Dennstaedtiaceae). Ann. Bot. (Oxford) 85 (Suppl. B): 77-99.

THOMSON, J.A. 2004. Towards a taxonomic revision of Pteridium
(Dennstaedtiaceae). Telopea 10: 793-803.

THOMSON, J.A., CHIKUNI, A.C. & MCMASTER, C.S. 2005. The taxonomic status
and relationships of bracken ferns (Pteridium: Dennstaedtiaceae) from sub-Saharan
Africa. Bot. J. Linn. Soc. 148: 311-321.

THOMSON, J.A., MICKEL, J.T. & MEHLTRETER, K. 2008. Taxonomic status and
relationships of bracken ferns (Pteridium: Dennstaedtiaceae) of Laurasian affinity in
Central and North America. Bot. J. Linn. Soc. 157: 1-17.

TRYON, R.M. 1941. A revision of the genus Pteridium. Rhodora 43: 1-31, 37-67.

VERDCOURT, B. 2000. Pteridium. In: BEENTJE, H.J. (Ed.) Flora of tropical East
Africa, Dennstaedtiaceae, pp.5-8. A.A. Balkema, Rotterdam.

THOMSON: CONFLICTING TAXONOMIES IN PTERIDIUM 109

WOLF, P.G., HAUFLER, C.H. & SHEFFIELD, E. 1987. Electrophoretic evidence for
genetic diploidy in the bracken fern (Preridium aquilinum). Science 236: 947-949.

WOLF, P.G., HAUFLER, C.H. & SHEFFIELD, E. 1988. Electrophoretic variation and
mating system of the clonal weed Pteridium aquilinum (L.) Kuhn (bracken).
Evolution 42: 1350-1355.

110 FERN GAZ. 18(3). 2008

BOOK REVIEW

THE LIVERWORTS, MOSSES AND FERNS OF EUROPE, BY
WOLFGANG FREY (HORNWORTS AND _ LIVERWORTS),
JAN-PETER FRAHM (MOSSES), EBERHARD FISCHER AND
WOLFRAM LOBIN (FERNS AND FERN-ALLIES). Translated and
updated by the authors with illustrations prepared by H. Lunser
(Bryophytes) and E. Fischer (Ferns). Harley Books. English edition revised
and edited by T.L.Blockeel .2006. 512pp.( ferns and former fern allies pp.363-
455), 166 figs, each incorporating numerous line drawings (ferns and former
fern allies figs. 121-166). 218 X 155mm, hardback. ISBN 0 946589 70 4. £45.
German edition first published 1995.

Wide ranging titles such as this usually mean a book will be a ‘Jack of all subjects’ and
master of none, so you can imagine the pleasant surprise I had on dipping into it. Ferns
are indeed a relatively minor part filling fewer than 100 pages but the coverage is more
comprehensive than given in many field handbooks.

Apart from a couple of prefaces there are no introductory chapters. The books start
straight away with the dichotomous keys. At the end there is a glossary and
bibliography.

None but the most well read specialist will be familiar with all the taxa described here.
Recently discovered species and subspecies are included. There are no lengthy field
notes, details are precise and to the point. Most species are covered with fewer than four
lines of text. Distribution is given by country or island within Europe and more briefly
outside of Europe. As with the rest of the book the entire fern section is a generously
annotated dichotomous key. The key is supported by many line drawings which work
well for the well defined species but by the time you get down to the really critical taxa
they are not so helpful. I looked long and hard at one sketch of the base of the
lowermost pinna of Gymnocarpium robertianum but | still cannot work it out!

The fern tourist venturing anywhere in Europe, be it the Azores, Crete, Cape Verde,
Madiera, Russia or closer to home will find all the local fern species covered. A few
hybrids are also mentioned although the criteria for their inclusion is obscure to me. For
example the very rare Aspelnium x heufleri is in the key but many others, such as A. x
Sarniense, are not.

I can think of no other up to date book covering all the ferns of Europe. Remy Prelli’s
Les fougeres et plantes allies de France et d'Europe occidentale has a more restricted
range. Where the two overlap I suggest the Prelli book will be the more user friendly
even though it is in French. For full coverage of Europe in one go, however, this guide
is a useful volume.

Martin Rickard

FERN GAZ. 18(3). 2008 111

BOOK REVIEW

BIOLOGY AND EVOLUTION OF FERNS AND LYCOPHYTES. Tom A.
Ranker & Christopher H. Haufler (eds). 2008. 480pp., 66 line figures 45
halftones. ISBN 978-0521-87411-3 hardback; ISBN 978-0521-69689-0
paperback. Cambridge University Press. £75.00, £35.00.

The editors of this new book have taken up the challenge of presenting the reader with
the major research developments that have taken place over the last 10 years that
contribute to the current understanding of the biology of ferns and lycophytes. To
achieve this goal they have brought together the work of some 28 contributors all active
in research, mainly from USA but also from Canada, Europe and Japan.

The subject areas are diverse. The first section on development and morphogenesis
starts with the gametophyte and alternation of generations, before exploring the
structure of meristems and subsequent diversification into the various organs — root,
stems and leaves. The next section deals with genetics and reproduction, including
chapters on the role of antheridogens, breeding systems, and the chloroplast and nuclear
genomes. Ecology is a complex discipline in itself, and the focus here is on phenology
and habitat specificity in tropical ferns, the ecology of gametophytes and a review of
conservation biology. The final section, systematics and evolutionary biology, includes
a review of the evolution of ferns and fern-like plants from the fossil record, a chapter
on species and speciation, exploration of diversity, biogeography and floristics, and
ending with phylogeny and classification.

Each chapter presents a personal review of the subject by it author(s), with summaries
of discoveries and developments, explanations of concepts and ending with a section on
prospects for the future - all supported by an extensive list of references.

This is a scholarly work, clearly aimed at the undergraduate student market, and I can
see it being added to the essential reading list for many courses in plant evolutionary
biology. But it is more than being a standard undergraduate textbook and with such a
wide variety of topics covered, there will be sections to interest a variety of general
readers and particularly for those with a fascination for fern biology. It is good to see
that it available as a more modestly priced paperback as well in hardback format.

Mary Gibby

112 FERN GAZ. 17(3). 2005
INSTRUCTIONS FOR AUTHORS

PAPERS should not usually exceed 20 printed pages and are generally expected to be
considerably shorter. Review articles, as well as reports of original research, are
encouraged. Short notes are acceptable e.g. new records. The senior author should
supply a fax and email address to facilitate correspondence.

MANUSCRIPTS should be submitted in English (British) in electronic format
(preferably) or hard copy (two copies), in 10-point Times New Roman font and wie
spaced. Electronic versions of text and tables should be compatible with WORD, w
figures as pdf or jpg files, and sent as email attachments or CDroms. All eee
will be refereed

THE TITLE should reflect the content of the paper and be in BOLD CAPITALS (11-
point) and centrally aligned. Generic and specific names should be in italics and any
title containing a generic or specific name must be followed by the family and
Pteridophyta in brackets e.g.

TRICHOMANES SPECIOSUM (HYMENOPHYLLACEAE:
PTERIDOPHYTA) IN SOUTHERN SPAIN

AUTHOR ABBREVIATIONS should follow Pichi Sermolli's (1996) Authors of
scientific names in Pteridophyta, Royal Botanic Gardens, :
MAIN HEADINGS: should be in BOLD CAPITALS (10-point) and centrally

igned.
SUBSIDIARY HEADINGS: should be in bold, the first letter of each word in capitals,
the rest in lower case and left-aligned.

AUTHORS' NAMES AND FULL ADDRESSES: follow the title and are centrally
aligned.

KEY WORDS: up to ten.

ABSTRACT: should reflect the content of the paper.

FIGURES: there is no distinction between photographs and line drawings in
numbering. All should be presented in a form ready for reproduction, ideally in JPG
format (please contact editor with queries), with a scale bar where appropriate.
Lettering or numbers (Arabic) should be in the bottom left using uppercase Times
Roman and be sufficiently large to be legible if reduction is necessary during printing.
The number of photographs allowed in any one issue is limited by cost. Figure captions
should be on a separate sheet

TABLES: can be printed in either portrait or landscape format. Authors should consider
this when preparing tables. Authors should ensure that tables fit the printed page size in
a legible form

MEASUREMENTS: should follow the metric system.

CHECKLISTS: should follow the format of Baksh-Comeau, Fern Gaz. 16(1, 2): 11-

REFERENCES: should follow the style of a recent issue of The Fern Gazette, e.g.:-

HOOKER, W.J. 1864. Species Filicum, 5. Dulau & Co., London

MORTON, C.V. 1947. The American species of Hymenophyllum, section
Sphaeroconium. Contr. U.S. Natl. Herb. 29(3): 139-201.

STEVENSON, D.W. & LOCONTE, H. 1996. Ordinal and familial relationships of
pteridophyte genera. In: CAMUS, J.M., GIBBY, M. & JOHNS, R.J. (Eds)
Pteridology in perspective, pp. 435-467. Royal Botanic Gardens, Kew.

JOURNAL ABBREVIATIONS: should follow Botanico Periodicum Huntianum &

Supplements.

Alterations from the original text at proof stage will be charged for unless they are

minor points of detail. Twenty-five offprints will be supplied free to the senior author.

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THE FERN GAZETTE

VOLUME 18 PART 3 2008
CONTENTS
MAIN ARTICLES
The cytotaxonomy of the Asplenium stuhlmannii complex (Aspleniaceae:
Pteridophyta) in Africa
A. F. Braithwaite 81-90
Asplenium delavayi (Aspleniaceae: Pteridophyta) - A fern species new to
: A. Rijal 91-97
M photyp flicting t ies in Pteridium (Dennstaedtiaceae:
J. A. Thomson 101-109
SHORT NOTES

Corrections to A revised handbook to the Flora of Ceylon, Volume
Parts A & B, Ferns and Fern-allies
M. Shaffer-Fehre-