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181 FERN GAZ. 17(4). 2005
British Pteridological Society
with
Royal Botanic Garden Edinburgh
Linnean Society of London
Ferns for the 21st Century
Proceedings of the International Pteridophyte Symposium
at the
Royal Botanic Garden Edinburgh, Scotland, UK
12-16 July 2004
Part 2
Editors: M. Gibby, A. Leonard, H. Schneider & J.C. Vogel
MISSOURI BOTANICAL
MAR 0 8 2006
GARDEN LIBRARY
ROY AL
BOTANIC
GARDEN
EDINBURGH
182 FERN GAZ. 17(4). 2005
THE DELEGATES
1)Marina Romanova 2) John Mickel 3) Eric Schuettpelz 4) Alan Smith 5) Judy
Jernstedt 6) Marcus Lehnert 7) Klaus Mehltreter 8) Germinal Rouhan 9) Kunio
Iwatsuki 10) Michael Kessler 11) Jens Pahnke 12) T. Boonkerd 13) Leon Perrie 14)
Louis Chinnery 15) Stephen Blackmore 16) Niklas Wikstrom 17) Eka Iskandar 18)
Jennifer Winther 19) Julie Barcelona 20) Barbara Parris 21) Paul Kenrick 22) Elena
Estrelles 23) Michael Barker 24) Jonathan Krieger 25) Adrian Dyer 26) Ana Ibars 27)
Peter Crane 28) Ruth Stockey 29) Elizbieta Zenkteler 30) Heather McHaffie 3 1) Gar
Rothwell 32) Ruth Kirkpatrick 33) Joanne Sharpe 34) Peter Hovenkamp 35) Graham
Ackers 36) Harald Schneider 37) Paulo Windisch 38) Daniel Ballesteros 39) Thomas
Janssen 40) Carol Gibb 41) Alison Paul 42) Mary Gibby 43) Istvan Pintér 44) Irina
Gureyeva 45) Tony Braithwaite 46) Royoko Imaichi 47) Mitsuyasu Hasebe 48)
Johannes Vogel 49) Alasdair Wardlaw 50) Carl Taylor 51) Chris Page 52) Chie
Tsutsumi 53) Frank Katzer 54) Wen-Liang Chiou 55) Patrick Acock 56) Yves Krippel
57) Masahiro Kato 58) Gregor Kozlowski 59) Maarten Christenhusz 60) Kathleen
Pryer 61) Sunniva Aagaard 62) Jurgen Kluge 63) Hanna Tuomisto 64) Harriet Hunt 65)
Atsushi Ebihara 66) Fred Rumsey 67) Virag Krizsik 68) Luis Quintanilla 69) Ruth
Aguraiuja 70) Kai Runk 71) Layne Huiet 72) Nathalie Nagalingum 73) Mirkka Jones
74) Chan-Ho Park 75) Susan Klimas 76) Raymond Cranfill 77) Petra Korall 78) Tom
Ranker 79) Sabine Hennequin 80) Michel Boudrie
FERN GAZ. 17(4): 183-203. 2005 183
GRAMMITIDACEAE (PTERIDOPHYTA) OF MOUNT JAYA, NEW
GUINEA AND OTHER MONTANE MALESIAN LOCALITIES
B.S. PARRIS
Fern Research Foundation, 21 James Kemp Place, Kerikeri, Bay of Islands, New
Zealand 0470 (Email: bsparris@igrin.co.nz)
Key words: Grammitidaceae, Pteridophyta, Mt Jaya, Mt Kinabalu, Seram, Gunung
ulu, Malesia
ABSTRACT
Mount Jaya (5000 m) in Papua (Indonesian New Guinea) is the highest
mountain in Malesia and 56 species of Grammitidaceae are known from it. They
are listed, together with their altitude ranges, habitat types and geographic
distribution. Comparisons are made with Grammitidaceae of Murkele Ridge in
Seram, Moluccas, Indonesia (3000 m, 55 species), Mount Kinabalu in Sabah,
East Malaysia, Borneo (4100 m, 76 species) and Gunung Mulu, Sarawak, East
Malaysia, Borneo (2400 m, 43 species). A Malesia-wide core of 23 grammitid
species is identified.
INTRODUCTION
The limestone massif of Mount Jaya (c. 5000 m) in Papua (Indonesian New Guinea) ts
the highest peak in Southeast Asia. It is the only peak in the region high enough to
support glaciers and has long been a magnet for biologists. Previous names for the
mountain are Mount Carstensz and Punjak Sukarno. The Royal Botanic Gardens, Kew,
has organised five botanical expeditions to the mountain during 1998 to 2000, during
which many collections were made along an altitudinal transect on the southern side. A
detailed description of the alpine flora of Mount Jaya (above 3000 m) is in preparation
at the Royal Botanic Gardens, Kew (Johns et al. in prep.). While writing up the
treatment of Grammitidaceae for this account, all relevant material collected on Mount
Jaya was identified, not just that from high altitudes, and included specimens gathered
by Kloss on the Wollaston expedition on the mountain during 1912-1913 and those
collected by Miller in 1991 and 1992. Although nearly all of the available material has
been collected from only the southern side of the mountain, it is of sufficient interest to
document here in the following terms: |) how many species?. 2) what are they?, 3) what
are their altitude ranges?, 4) what vegetation types are they found in?, 5) where else do
they occur?
Comprehensive collections of pteridophytes from specific localities are rare in
Malesia, because many botanical collectors have specialised in gathering only
flowering plants and have tended to overlook the smaller and more obscure species of
pteridophytes, including most Grammitidaceae. There are only three other localities in
Malesia where the fern flora, including Grammitidaceae, has been relatively
well-collected over a wide altitudinal range: Mount Kinabalu Park in Sabah, East
Malaysia, Borneo, Gunung Mulu National Park in Sarawak, East Malaysia and Murkele
Ridge in Seram, Moluccas, Indonesia. More than 40 species are known trom each of
these localities and they provide a useful comparison with Mount Jaya. The following
questions are relevant: 1) how much do the floras of the three areas have in common
184 FERN GAZ. 17(4): 183-203. 2005
€
with Mount Jaya?, 2) is there a distinct East Malesian element present on Mount Jaya
and Murkele Ridge?, 3) is there a distinct West Malesian element on Mount Kinabalu
and Gunung Mulu?, 4) is there a Malesia-wide group of species of predictable
occurrence?, 5) do other pteridophytes show the same geographical distribution patterns
and do they also have Malesia-wide species of predictable occurrence?
Map | shows the location of Mount Jaya, Murkele Ridge, Mount Kinabalu and Gunung
Mulu.
Map 1. Malesia.
Philippines
ie
PACIFIC OCEAN
10°N
Peni Mt Kinehety
Malaysia
‘ Gunung Mulu re : M
O’N
New Guinea
Borneo Murkele Ridge ?
Seram Mt Jaya
Lesser Sunda Is.
10°S
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Most species of Grammitidaceae are epiphytes on trunks and branches of a variety of
trees and shrubs, or on the trunks of Cyathea in alpine communities, often associated
with bryophytes, and a few species, particularly those of subalpine habitats, are
lithophytes.
Mount Jaya
56 species of Grammitidaceae are known from Mount Jaya. They are listed, together
with their altitude range, vegetation types and distribution, in Table 1.
Tropical lowland rainforests are generally poor in members of Grammitidaceae,
Which are far more numerous above the average daily lower limits of cloud. On Mount
Jaya only five (9%) species occur in lowland rainforest, which contains the tree genera
Chisocheton, Ficus, Mvristica, Octomeles, Pometia and Vatica, with undergrowth of
ferns including tree ferns, Arthropteris and Marattia, and Gesneriaceae. All five species
are also found in heath forest. Heath forest, the other lowland forest habitat on Mount
Jaya, supports a surprisingly large number of species, 16 (29%). It occupies old
Pleistocene outwash terraces at the foot of the mountain (R.J. Johns pers. comm.) and
includes the trees Alstonia spectabilis R. Br. Calophyllum, Casuarina, Dacrydium,
Metrosideros and Vatica, together with Nepenthes, Davallia, Lecanopteris,
Hymenophyllaceae and Polypodiaceae. Two of the four Mount Jaya endemics,
Ctenopteris hymenophylloides (also in lowland rainforest) and Grammitis velutina,
occur here, together with two other species (Crenopteris nutans, Grammitis
scabristipes) disjunct from much higher altitudes in midmontane forest, upper montane
PARRIS: GRAMMITIDACEAE OF MOUNT JAYA, NEW GUINEA
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188 FERN GAZ. 17(4): 183-203. 2005
forest and subalpine communities. Crenopteris serrata, Grammitis pleurogrammoides,
Prosaptia burbidgei, Scleroglossum pusillum, S. sulcatum and oa ies en setulifera
are known only from heath forest on Mount Jaya, but occur in other communities
elsewhere in New Guinea and further afield in Malesia. The aie: may exist
elsewhere in New Guinea, but apparently no members of Grammitidaceae have been
collected from it. Heath forest, also called kerangas, is known from Peninsular
Malaysia, Borneo, and elsewhere in Malesia. It develops at low altitudes on very poor
or thin soils over rock or in areas of impeded drainage and is characterised by straight,
pole-like small to medium sized trees and an understorey including pitcher plants
(Nepenthes). Usually it contains very few, if any, species of Grammitidaceae.
Apart from heath forest, the most species-rich habitats on Mount Jaya are
midmontane forest (28, 50%), and subalpine communities, with 19 species (21%).
Lower montane and upper montane forests are less rich, each with 9 species (16%).
Lower montane forest contains the fern Se//iguea (no other information concerning the
vegetation of lower montane forest is available on the specimen labels). The
midmontane forest is dominated by Nothofagus; Casuarina, Dacrydium, Fagraea,
Loranthaceae, Macaranga, Papuacedrus, Rhododendron and Timonius are also present
together with the sedge Mapania, the ferns Asplenium, Belvisia, Cyathea,
Elaphoglossum and Loxogramme, and shrubby Urticaceae. Grammitis clavata is the
only Mount Jaya endemic in midmontane forest. Only eight species are restricted to
midmontane forest, with another eight extending downward to lower montane forest
and sometimes further, to heath forest and lowland rainforest, and 13 extending
upwards to upper montane forest and sometimes to the subalpine communities. Upper
montane forest is dominated by Papuacedrus papuana (F. Muell.) Li and Podocarpus
brassii Pilg., with Dicksonia, Drimys, Elaeocarpus, Nepenthes, Rapanea,
Rhododendron, Sloanea and Utricularia. Grammitis interrupta, Scleroglossum
Juncifolium and Themelium graminifolium are restricted to upper montane forest on
Mount Jaya, but occur in other vegetation types elsewhere. Another five species occur
in midmontane forest as well as upper montane forest, and two, Crenopteris
bipinnatifida and Grammitis padangensis, extend to the subalpine zone. The subalpine
grasslands are dominated by Cyathea tree ferns, while the subalpine shrubland contains
Cyathea, Coprosma, Drimys, Rhododendron and Vaccinium, and the subalpine forest
contains Dacrycarpus compactus (Wassch.) de Laub. and Podocar- pus. Nine species are
restricted to subalpine habitats: Calymmodon fragilis, C. ramifer, Ctenopteris fusca, C.
subulatipinna, Grammitis ceratocarpa, G. debilifolia, G. locellata, G. meijer-dreesii and
Prosaptia davalliacea. With the exception of G Jocellata and P. davalliacea, all are
New Guinea endemics. While the nine species all occur on Mount Jaya at altitudes
higher than that of the nearest subalpine site to New Guinea, Murkele Ridge (3000 m)
in Seram, the two species that are widespread through Malesia, G /ocellata and P.
davalliacea, occur at lower altitudes on Murkele Ridge.
Several species found on Mount Jaya are infrequently collected and are known from
very few localities. They include medium to high altitude Papua endemics known
elsewhere only from Lake Habbema - Grammitis hispida, Themelium allocotum, Mount
Trikora - Ctenopteris fusca, G. meijer-dreesii, and/or Mount Doorman, C.
subulatipinna, T. allocotum, and Mount Goliath and Hellwig Mounts - Xiphopteris
pseudospiralis. Xiphopteris govidjoaensis (syn. X. exilis Parris) is known from two
na localities, Gunung Belumut in Peninsular Malaysia and Govidjoa in Papua New
uinea.
PARRIS: GRAMMITIDACEAE OF MOUNT JAYA, NEW GUINEA 189
Four species (7%) are endemic to Mount Jaya; Crenopteris hymenophylloides,
Grammitis clavata, G. papuensis and G. velutina. Surprisingly, none occurs in the
subalpine communities, as might be expected when considering that several other
mountains in New Guinea have high altitude endemics, e. g. Grammitis dictvmioides
Copel, G habbemensis Copel., Xiphopteris antipodalis Copel. (all Lake Habbema), G.
reptans Parris (Mount Trikora), G montana Parris, G. trogophylla Copel. (both Mount
Sarawaket), G murrayana (C.Chr.) Copel. (Wharton Range) and G. salticola Parris
(Mount Giluwe) (Parris, 1983).
A number of other high altitude New Guinea species found on Mount Trikora and/or
Lake Habbema, and elsewhere in Papua New Guinea, may be expected on Mount Jaya,
including Ctenopteris pendens (Rosenst.) Copel., C. whartoniana (C.Chr.) Copel. and
Grammitis tomaculosa Parris.
New Guinea endemic species are important on Mount Jaya, comprising 36% (20
species) of the total. While they occur in all of the vegetation types they are best
represented in the high altitude subalpine communities, where they form 20% (11
species) of the total number of species. The largest number of species, 24 (43%) are
Malesia-wide in the sense that they occur in both West Malesia and East Malesia, on
both sides of Zollinger’s line (see later) between Borneo and Sulawesi, sometimes
extending into Thailand, Indochina and Taiwan to the north-west and into Melanesia
(Solomon Islands, Vanuatu, New Caledonia and Fiji) to the south-east. A small group
of species (4, 7%) is very wide-ranging, from South India to the Society Islands
(Ctenopteris blechnoides), South India to the Marquesas Islands (Prosaptia contigua),
Sri Lanka to the Marquesas Islands (Sc/eroglossum sulcatum) and from South India to
New Guinea (P. obliquata). These four species are amongst the most common and
widespread members of Grammitidaceae in the Asia-Malesia-Pacific region. Another
four (7%) species are restricted to East Malesia; Crenopteris serrata ranges from
Sulawesi to New Guinea, Calymmodon mniodes is in the Moluccas and New Guinea,
Grammitis pleurogrammoides extends from the Moluccas to the Solomon Islands and
Scleroglossum juncifolium occurs in New Guinea and the Solomon Islands.
The grammitid flora of Mount Jaya inevitably invites comparison with that of
Mount Kinabalu (4100 m) in Sabah, Borneo, East Malaysia, the highest peak between
the Himalayas and Papua, and the two other areas sufficiently well-known to permit
comparison of their grammitid floras: the Murkele Ridge, a limestone massif including
Gunung Binaiya (3000 m) in Seram, Moluccas, Indonesia, and Gunung Mulu National
Park (including Gunung Mulu, sandstone, 2400 m and Gunung Api, limestone, 1700 m)
in Sarawak, Borneo, East Malaysia. It is useful to examine the composition of these
montane grammitid floras with respect to Zollinger’s line, which divides West Malesia,
comprising Peninsular Malaysia, Sumatra, Java, Borneo, the Philippines and the Lesser
Sunda Islands, from East Malesia, consisting of Sulawesi, the Moluccas and New
Guinea, and separated by the Makassar Strait between Borneo and Sulawesi. Wallace’s
line is similar, but includes Bali, the westernmost of the Lesser Sunda Islands, in West
Malesia, and the remaining Lesser Sunda Islands in East Malesia. As the Lesser Sunda
Islands are considered a discrete botanical region, Zollinger’s line is preferred to
Wallace’s line. Mount Kinabalu and Gunung Mulu National Park (subsequently
referred to as Gunung Mulu) are in West Malesia, while Murkele Ridge and Mount Jaya
are in East Malesia. Both Mount Kinabalu and Murkele Ridge have subalpine areas
above the treeline, but Gunung Mulu is forested to its summit.
The Moluccas have been of biogeographical interest ever since Wallace (1869)
190 FERN GAZ. 17(4): 183-203. 2005
suggested that the region has been constantly receiving immigrants from New Guinea.
Kato (1989a) also points out that the affinity of the fern flora of Seram at species level
is with New Guinea to the east rather than with Sulawesi to the west. Parris (1993) notes
that the endemic pteridophytes of Seram are derived on the whole from wide-ranging
taxa or those from East Malesia, and that Seram is the end of the line for numerous
species moving east from West Malesia and moving west from New Guinea.
Grammitidaceae of Seram collected by the Japanese-Indonesian expeditions of 1983,
1985 and 1986 (but not those collected by Parris in 1987) have been listed (Kato &
Parris, 1992), the pteridophyte flora of Mount Kinabalu has been documented (Parris ef
al., 1992), and a checklist of pteridophytes of Gunung Mulu National Park has been
compiled (Parris, unpublished). Numerous changes have been made to the taxonomy of
Grammitidaceae in Seram and on Mount Kinabalu since the publications of Kato &
Parris (1992) and Parris et al. (1992).
Murkele Ridge
55 species of Grammitidaceae are known from Murkele Ridge. They are listed, together
with their altitude range, vegetation types and distribution, in Table 2.
It is useful to compare the grammitid floras of Mount Jaya and Murkele Ridge, the
two East Malesian localities, which have very similar numbers of species, 56 in the
former and 55 in the latter. 25 species are shared between the two, and nine of these are
restricted to East Malesia. The remaining 16 are more widespread, occurring in East and
West Malesia and sometimes beyond. Only Ctenopteris serrata extends throughout
East Malesia from Sulawesi to New Guinea, so a well-defined widespread East
Malesian endemic element is lacking in Grammitidaceae. Only one species, Acrosorus
Jriderici-et-pauli, reaches its eastern limit of distribution on Murkele Ridge in Seram,
but eight others, Calymmodon mniodes, Ctenopteris whartoniana, Grammitis
ahenobarba, G. collina, G. parva, G. pseudolocellata, G. subfasciata and Prosaptia
engleriana, reach their western limit in Seram, either on Murkele Ridge or slightly
further to the north-west on Gunung Kobipoto (G. collina) or Gunung Roihelu (G
parva). They amply demonstrate Wallace’s point (1869) that the Moluccas receive
immigrants from New Guinea. In addition, some of the Murkele Ridge endemic species
have obvious vicariants in New Guinea, e. g. Ctenopteris themelioides with C.
bipinnatifida, and Themelium pseudallocotum with T. allocotum. The similarity in
species numbers between Mount Jaya and the much lower Murkele Ridge is surprising,
and the reasons for it are not obvious.
Comparisons between Murkele Ridge and the two West Malesian localities, Mount
Kinabalu and Gunung Mulu, are interesting. 33 species are shared with Mount
Kinabalu, more than are shared with Mount Jaya. They include seven widespread
species collected on Gunung Mulu and known from New Guinea that are absent from
Mount Jaya: Calymmodon gracilis, C. pectinatus, C. reconditus, C tenopteris
brevivenosa, Grammitis adspersa, G. impressa, G. reinwardtii. All are expected on
Mount Jaya. 19 species are shared with Gunung Mulu and, with the exception of
Calymmodon conduplicatus, all are also known from Mount Kinabalu.
31 species (56%) on Murkele Ridge are found in both East and West Malesia and
another 5 (9%) extend well beyond Malesia, while 9 species (16%) occur in East
Malesia and another 9 (16%) are endemic to Murkele Ridge. The amount of local
endemism is much higher than on Mount Jaya, which has only four endemics. One
might expect local endemism at high altitudes on Murkele Ridge, the highest part of the
19]
PARRIS: GRAMMITIDACEAE OF MOUNT JAYA, NEW GUINEA
BOUIND) MIN O} ISAMP[NS
DWAPUd ISPly IOxINJA|
BOUIND) MOK O} BISARIRIAY “Ud
BOUIND MON O} BRUINS
BOUIND MAN, O} BULYDOPU|
njenued 0} purypliey
“S] AJOIDOG 0} BIPUT
njyenurA 0} BIsayey “VISY JS
BOUIND MIN 0} OOUIOg
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BOUINL) MON “SBdONO||
BOUIND MON O} RISYV AS
"S| UOWO]OS 0} BISARTRI “Ud
BOUND) MAN 0} OdUIOG
BOUIND MAN O} BVBWINS
dWAapuUds dIspry sayin
dIWapuds dIspry aon
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uOHNgLySIC
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‘jadoy (auinyg) eyosaqpu "DS |
NdIpPIeL WAY D (oun |
UNVJOH CMIOply) BSOUDSATAAIG “DE ]
—
Bye|NOQUap "DO ‘pI
JoyIasH 2 AUseA\’ HM (AIID) Saplouysayq sluaidoualD ‘Z|
SLUR (IDV) BPURIARIDSNU SILULURIBOSAIY) *| |
‘paul sued SNIpUuodal “Dd ‘0
‘pour sued snyeunsad *y
‘jado.y sapotuuwt “>
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[SXq’) (OuIN|g WZ SdaN) sniBT[NINd * +)
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DOO [| (YOO) JOJIAB]S “+
‘pour sLueg sisuddreulg UOpoUWAye)
aa tu 6K wo oO
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‘yadod (syd H) yned-ja-19apLy snsoso1ay
_
saiadg
‘ISpPly apayn JO avaowpniwweIyH “7 9qUL
FERN GAZ. 17(4): 183-203. 2005
192
BOUIND MON O} BQBUINS
BOUIND) MAN “SBOON[O|y
njenue, Oo} BQeUINS
‘S] UOWIOJOS 0} BYU] LIS
BOUIND) MAN “SsBdoNjO|W
BOUIND) MON ‘sedoNjop]
BOUIND MON O} BAVRUINS
SRIIN|O|] 0} OAUIOG
d1WapUd dSpry spayany
BOUINDH MAN OF BVBUINS
if1.J 0} BISARTRIY Ae[NSUIUDY
dIWapUd dSpry sjayINy
BOUIND) MON 0} UBMIB]
BOUIND MANY O} BRUINS
BOUIND) MON O} ODIO
BOUIND MAN ‘sBdoN[O|
BOUIND MON O} BIZRUINS
BOUIND MON ‘seoonjoyy
fly 01 BIS JS
BOUIND) MON ‘sKdoNjoYy
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BIUOPs[BD MAN O1 BALLS
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* ado (suasoy) eIVIOSRIQNS TD “GE
‘yadoy (4ayeg) sadnsuqess ‘ry “g¢
aun] MpaweMuUlal FH “LE ¢
SLUR BIL]]AOJOpNasd ‘FH ‘g¢
‘jadoD (asneig) eared tp “¢¢
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SLUR (ISUYD'H) BSSIWONUI TD “gz
‘jado.y essaidun 1 “47
‘Jado (‘jado,)) vsosoysijop "DH ‘gz
SLUR BULL[OS ‘Ty ‘SZ
auinyg vsoudsaes ty “pz
SLUR BqIeqoudye T “FZ
auwinyg (owinyg) esusdspe sniutuPIin *7z
‘Jado, (4YyD*D) BuriuoEYM * “1 Zz
‘Poul SLUBY SAPlOlaWay) “J ‘QZ
‘}adod (]]07Z) vdasstIpopunsasqns ‘61
193
PARRIS: GRAMMITIDACEAE OF MOUNT JAYA, NEW GUINEA
‘(WE QOOE-O0PT) SAH
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BOUIND MON O} BISARTRI “Ud
BOUIND) MON O} BISARIRI] “Udd
‘S] UOWO]OS 0} sourddiyiyd
BOUIND MON 0} OOUIOG
dWIapUda dBpry san]
sesanbiey 0} ByURT] LIS
“S| UOLUOJOS 0} pueypeyL
BOUIND MON O} BALL
dIWIApUd sBdON]O||
d1WdpUd ddply sa4ANJA|
BOUIND MON O} BIPU] S
BOUIND MAN ‘SBdoN|O/
BOUIND MON 0} BISARTRIY “Ud
"s] sesonbieyy 0} eIpUuy S
otwapud aspry spoyINy|
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WwW OOET-0001
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WO00EI
‘jadoy (aun) epyyneuurdgns "x “¢¢
Sued CAOpPlY) elayipnias suaydoydry “ps
sled (Jadod) MApoAd “| ES
SIIB (DAVE) SOPIOIPOXR] “| ZS
‘pour siueg WNnjosojjepnasd wintyawayy |S
“MIOPTY (UyNy) WUNRO[NS “S "OS
“MIOppy (aunyg) tunypisnd winssojSo19[9g “Gp
IUD’ W (azuny xo awn] g) BSO[NUDIA ‘g ‘Rp
sluvg (sLuRg 2 OY) BIqe{sqns ‘gq “Lp
‘POU SILUIBY SISUSWIRIAS ‘gf ‘OP
NOW (ounpg) eienbijgo “g “Sp
‘jadoy (asneig) euBLio|sud ‘g “pr
‘Jodo (oyeg 7 ‘[ONW4) B9ORTT[BARP ‘g “EP
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194 FERN GAZ. 17(4): 183-203. 2005
Moluccas and the only part of the region to be above the tree line, but only Calymmodon
binaiyensis is restricted to subalpine habitats. Crenopteris themelioides ranges from
lower montane forest to subalpine habitats while the other local endemics are all found
in lower montane forest. Only one species, Prosaptia subglabra, is endemic to the
Moluccas.
Mount Kinabalu
76 species of Grammitidaceae are known from Mount Kinabalu. They are listed,
together with their altitude range, vegetation types and distribution, in Table 3.
Mount Kinabalu and Mount Jaya make an interesting contrast. The former has far more
species of Grammitidaceae, 76, than the latter, with 56. Mount Kinabalu has a long
history of botanical collecting, however, over more than a century, which may explain
its species richness in part (Parris ef al., 1992). It should be noted, however, that nine
species have not been collected on the mountain in the last 60 years (Parris, 2001) and
may well be locally extinct. 23 species occur on both mountains; all of them, except
Themelium fasciatum, are also found on Murkele Ridge, and 10 of them are also found
on Gunung Mulu. Mount Kinabalu has nine endemic species of Grammitidaceae (16%),
more than double the number found on Mount Jaya. As with Murkele Ridge, one might
expect the endemic species to be associated with the high altitude vegetation
communities above the tree line, but only Grammitis kinabaluensis extends above the
tree line, and it also occurs in lower montane and mid-montane forest. Grammitis
graniticola, G. muscicola and G nubicola are restricted to upper montane forest, while
Calymmodon innominatus and Grammitis ramicola are found only in lower montane
forest and Calymmodon kinabaluensis, Grammitis havilandii and G. ultramaficola
occur in both lower montane and upper montane forest. 44 species (58%) are
Malesian-wide, a higher number and percentage than for either Mount Jaya or Murkele
Ridge. Another eight species (11%) extend well beyond Malesia. 10 species are
restricted to West Malesia, six of which are also found on Gunung Mulu. The West
Malesian Crenopteris fuscata occurs at high altitudes on Mount Kinabalu (3000-3800
m) and is not found on Gunung Mulu (2400 m), but Grammitis congener (1100-2500
m) and Themelium curtisii (1400-2400 m) are absent from Gunung Mulu, but may be
expected. Xiphopteris nudicarpa occurs on ultramafic rocks on Mount Kinabalu and on
Palawan Island in the Philippines. Ultramafic rocks, and_X nudicarpa, are absent from
Gunung Mulu. Five species are endemic to Borneo and all are shared with Gunung
Mulu.
Gunung Mulu
43 species of Grammitidaceae are known from Gunung Mulu. They are listed, together
with their altitude range, vegetation types and distribution, in Table 4.
Gunung Mulu is lower than Mount Jaya, Murkele Ridge and Mount Kinabalu, and
lacks vegetation communities above the tree line. It has no endemic species, and 36 of
its 43 species are also found on Mount Kinabalu, 22 species (51%) are Malesian-wide
and five (12%) range well beyond Malesia. The nine West Malesian species (21%) all
occur on Mount Kinabalu. Seven species are endemic to Borneo: five also occur on
Mount Kinabalu, but Scleroglossum crassifolium and Themelium sarawakense are not
known there.
In Table 5 the geographic distribution of Grammitidaceae on Mount Jaya, Murkele
Ridge, Mount Kinabalu and Gunung Mulu is summarised.
199
GRAMMITIDACEAE OF MOUNT JAYA, NEW GUINEA
PARRIS
BOUIND) MON 0} BISARTRIY “Ud
BOUIND MON 0} BISY FS
soulddiyiyg 0} eiskeyeyy ‘Udd
ROUIND MAN O} B.BUINS
soulddiyjiyg 0} Biskeyeyy ‘udd
sourddiyiyg 0} purprey,|
BOUIND MON 0} BUIYyDOpUT
nyenue, 0} puryieyy,
“s] AJOI90g 0} BIPUT “Ss
nyenurd 0} BISy 4S
sourddipiyg 0) eyurT] Ls
BOUINL) MANY 0} OSUIOG
ROUIND) MON 0} BARS
soulddiiyg 0} eave
"S] UOLUOTOS 0} BISY 4S
o1luapus nyjeqeury JUNO
dUdpUs Njeqeury JUNO
BOUIND) MAN O} BALL
BOUIND) MAN O} BISY FS
"S] UOWO]OS 0} BISARRIY ‘Ud
BOUIND MAN O} BBUINS
dIWINpUsa OAUIOg
BOUIND MON 0} OOWIOgG
soutddiyiyg 0} pueyrey.|
SRdONJO|| 0} BISY FS
woHnqLysig
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WwW 000-008 I
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apnynty
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AZUNY (AWIN|g A SAIN) BWIOTIJOUL "-y “PTZ
wiINyjOH (awinyg) enue > °
‘yadoy (auinyjg) eloyaypru “>
‘jado,y (duinjg) ensidsuosut °°
azuny (auin|g) BIBdIsNy ° ‘OZ
naipsep Wu’ D (auinjg) erepnonuap *>y 6]
wINOH] (Maply) BSOUDATADIG "5 ‘81
JOYINID w 1dUseAA HA ('AdID) saprouysayg siuaidouany */ |
sued (Jayeg) eueiaridsnu “> “9|
sued ((Wg'f) esojnpurys sntuuUeIBOsAIY) “S|
‘poul sLueg snyipuodsal “> “P|
‘paul stueg smeunoad 9 “¢]
‘jadoy (Jado) sapioosnu “5 “Z|
‘jado,y (ultu0g) snuelUassiany “> “||
‘Poul Sie” Sisuon[eqeuly *) ‘Ol
‘poul sLueg smeuluouul y “6
‘jadon snaidossoisAy 8
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AOOW'L (CYOOH) JAPAR]I“ *¢
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oral
sapods
“nyeqeury juno) Jo avacepHiUUeID “¢ afquy
FERN GAZ. 17(4): 183-203. 2005
196
dIlUAapUd OdUOg
‘S| UOWOJOS 0} ByURT LIS
dlWdpud Nyeqeury JUNO
soutddiiyg 0) enewng
BOUIND MAN 0} BIeUINS
‘S] UOLUOJOS 0} BsyRUINS
dWdpUd Njeqeury JUNO
dIWIapUd Nyeqeury iUNO|
SROON|O|Y 0} OOWIOG
ROUIND MON 0} BRUINS
ify 01 eIsAeyeyy sepnsurusg
dIWAapUs Njeqeuly JUNOPy
‘S] UOWO[OS 0} BISY 4S
BOUIND MON O} URATE],
BOUIND) MAN 0} B.YBUUINS
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d1WapUd Nyeqeury JUNO
dWapus Njeqeury JUNO]
ISAMBINS ‘OOUIOg
BOUIND MANY 0} OAUIOg
sourddiiyd 01 vIsy 4S
ROUIND MON 01 OOWIOg
ROUINDH MON 0} BRUINS
d1uIapus OsUI0g
ify 01 BISy AS
BIUOpa[eD MON 0} BARS
OaWOg 0} BAyEUINS
‘S| UOWO]OS 0} BYU] US
AW
vs AWN
AWN AW
VS AWN AW
JW dW
dW
AW
AWA AW
dW AW
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AWN AWT
AWN AWT
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‘jadod saplonpieMulal ty “¢¢
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‘jadoy (s9yRq) stsuasueped ‘F “Gp
‘jadoy (s94eg) BIL[ODOULIGO "DH ‘gp
‘pout sled Bjoorqnu DH “Lp
‘pour SLUeg BJOSIOSNU ‘DH ‘Op
“POUT SIUB_ SI[JOU fT) “Sp
‘Jodo (4axeg) BIEI[P00] ‘+H “pp
‘jadoy (4oyeg) vuLIpsoysinUy "DH “Ep
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BME] (UYNY Xd “WoOY)) PUBLIOTEL ty “|p
SLU (SY D"H) BSSIWONUI T “Op
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‘}edoD (suy9"H) yned-19-19L19pLy °D “SE
‘Jodo (‘jedoD) erosoysijop 'D “PE
auIn|g 19udTUOd ‘FD ‘EE
siivdg (‘[adod) ovtsuouiajo 1p “zZE
auinjg vsoydsavs ry “| ¢
‘jadoy (‘jedop) stsuv08u0g Dy “Y¢
owing (ounjg) essodspe siptuUBsn “67
‘edo ([9Z) Bessipopunsasqns *> *gz
wN}OH (Maply) BynuILUgns *> °2Z
naipsel, FAY" (HE) enpuedas *> “gz
197
PARRIS: GRAMMITIDACEAE OF MOUNT JAYA, NEW GUINEA
‘(WIQORE-OOSE) Son
-IUNLULUOD dUIdyEgns “WS “(QOS E-009Z) IS2404 SURIUOUL Joddn “4A (LUQO9T-OOT 1) ISAIOJ BULIVOW J9MO] ALT “(LUT 1-008) 805 |]!Y “AH
BOUIND) MIN O} BISAR[RIY ‘Ud
soutddijiyg ‘oow0g
OoWOg 0} puRyey]
ROUIND MAN 0} OOWIOg
souiddijiyg ‘oauw0g
BOUIND) MAN O} BRUINS
ooUOg ‘eyRUINS
sesonbieyy 0) Bye] US
‘S] UOWIOJOS 0} puRpeYyy
d1Wapus OdUIOg
BOUIND) MAN O} BISALRY “Ud
BOUIND) MON O} BARS
BOUIND) MAN 0} BIPUT S
BIUIND) MIN, 0} CAWOg
BOUIND) MOXj 0} BISALIRI “Ud
"S| sesanbseyy 0} eIpuy §
BOUIND MAX O} puRyeY
d1LUapUd OdUIOg
sourddijiyd 01 BIsy 4S
BOWS 0} BIPUT S
o1luapus nyeqeury jNop
BOUIND MAN O} BILLINGS
nyenur, 0} eayeung
AW)
AWN AWT
AWN AWT
Wt (OTC-OOS |
W (06T-OOLT
W OOLT-O0TI
W QOOE-OOET
W OOIT
W OOFE-OOLT
WwW OOPCT-O0F |
W QOOE-O0S I
W YOTE-OONS |
WwW OOO!
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W (06T-00SC
Ut OORT-O0S |
WOO1C-O0S I
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W QOTC-OOS I
W OOIT
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W008
Ww OOOE-O0FT
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W O00E-008 I
‘jodo,y (owing) epyneuurdgns "yx *97
SLUvd (OD 3 BOW’) edivoipnu "x ‘G/
wnyyoH (ayo’a) usnuAuoaty "Xx “PZ
SLueg (oyeg) SOploiIpoxey "| EL
stueg (‘jadod) asusuosjey “fp *7Z
sued (Suing) Wingelosey pT L
sluvg (Joyeg) USNS WUINTPOWY Lf ‘OL
“MAIOPLY (UYNy) LUINIwOINS “S "69
“MIOpLY (atuN}E) wuNyjisnd “g “g9
“pour slueg wUNUUTTLOBIs “"S “79
‘MIAPLY (UYNS) a]IGop LUNssO[TOIIIIS “99
dd DW (azuny Xd dwn] g) BSO]NUDA ‘gq ‘S9
HOW (owing) eyenbiygo “q “p9
sued (jadod) eyepneonjnw “g “¢9
Jado) (1oxeE 7 “TONA) BOdBI|[BARP “qd *79
[SAld’ (Iss94'yH) ensyuod ‘g “19
‘SIRM|'Y ZW BMBBKY (QWIN]E]) BIIGI]99 ‘gq ‘09
‘POU SLUIBY SISUDDUIOG ‘qd *6S
IUD W (aye) vypAydosyyeseg “g 8
IsLYyD'H (ouinyg) viepe endesosg *L¢
‘poul SLUR BfOOTRUTRYIN "Fy ‘OCG
‘jadod (s94eq) BURBUINS "FD ‘SS
‘jadog (sayvg) sadnsiuqeos tH “pS
a
FERN GAZ. 17(4): 183-203. 2005
198
"S| UOWO]OS 0} vISy AS
ROUIND MON 0} BBN
ISOMB[NG ‘ooulOg
o1WIapud CoUIOg
ift,y 0} eISY JS
OoulOg 0} ByBUUINS
BOUIND MON 0} BISAReI “Ud
BOUIND MAN 0} BISY FS
ooulog ‘eyeunsg
nyenue, 0} pueypreyy
“S| AJOI90§ 0} vIPU]
nyenue, 0} vIsy AS
ROUIND MON 0} OdUIOg
BOUND) MON O} BALL
‘S] UOWO]OS 0} BISY 4S
BOUIND MIN 0} BISY AS
‘S] UOWIOJOS 0} BISAR] RIA ‘Udg
BOUIND MON 0} OoUIOg
dIWIApUs OSUIOg
saurddiyiyg 07 pueprey,y
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199
PARRIS: GRAMMITIDACEAE OF MOUNT JAYA, NEW GUINEA
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FERN GAZ. 17(4): 183-203. 2005
198
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199
GRAMMITIDACEAE OF MOUNT JAYA, NEW GUINEA
PARRIS
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PARRIS: GRAMMITIDACEAE OF MOUNT JAYA, NEW GUINEA 201
DISCUSSION
Compared with Murkele Ridge, Mount Kinabalu and Gunung Mulu, Mount Jaya has
many more country endemics. This reflects the explosive speciation of Grammitidaceae
in New Guinea (together with many other groups of plants) and their distribution along
the central ranges of the island. The wide distribution of species along the central ranges
of New Guinea is probably also an explanation for the low number of endemic species
compared with Mount Kinabalu and Murkele Ridge. While Mount Jaya is the highest
peak in a series of high altitude peaks on the ranges that form the backbone of New
Guinea, Murkele Ridge and Mount Kinabalu are isolated from other areas at high
altitudes. Mount Jaya and Gunung Mulu have fewer species that are widespread in
Malesia and beyond than do Murkele Ridge and Mount Kinabalu. Gunung Mulu, being
relatively low altitude, does not have the array of habitats or even comparable areas of
the same habitats as do the higher Murkele Ridge and Mount Kinabalu. The reason for
Mount Jaya having fewer of the widespread species is less obvious. In the discussion of
Murkele Ridge (see above), seven widespread species are mentioned that are absent
from Mount Jaya, but present in New Guinea and thus to be expected. Another two
species that occur on Mount Kinabalu and are disjunct with Papua New Guinea are also
expected on Mount Jaya. They are Calymmodon atrichus and Grammitis clemensiae.
There is a group of 10 grammitid species that have been collected on all of the four
montane Malesian localities documented here: they are Calymmodon cucullatus,
Chrysogrammitis musgraviana, Ctenopteris blechnoides, C. nutans, Grammitis
knutsfordiana, G. padangensis, G. sumatrana, Prosaptia contigua, Scleroglossum
pusillum and S. sulcatum. Two other widespread high altitude subalpine species are not
found on Gunung Mulu (2400 m alt.) because it is too low: Grammitis locellata
(2800-3800 m) and Prosaptia davalliacea (2800-3800 m). Another group of species,
Calymmodon clavifer, Ctenopteris denticulata ,C. millefolia, C. subsecundodissecta,
Grammitis dolichosora, G. intromissa, G. scabristipes, Prosaptia obliquata, P. venulosa
and Xiphopteris subpinnatifida occur on Mount Jaya, Murkele Ridge and Mount
Kinabalu within the altitudinal range of Gunung Mulu and may be expected there. The
widespread Grammitis reinwardtii is absent from Mount Jaya, but occurs in the other
three localities. The species that are absent from Gunung Mulu and from Mount Jaya,
together with the 10 listed above, form a core of 23 grammitid species that are
widespread and common throughout Malesia. Examination of herbarium collections
during preparation of the account of Grammitidaceae for Flora Malesiana has provided
much detailed data on species distribution that has been collated for over 500 localities.
The core group of 23 grammitid species is well-represented in the following localities,
each of which has 20 or more species of Grammitidaceae. The altitude range over which
Grammitidaceae have been collected and the number of species currently known is
given for each locality. Peninsular Malaysia: Cameron Highlands, 1400-2000 m, 23
species; Gunung Tahan, 600-2200 m, 21 species; Sumatra, Gunung Kerinci. 300-3800
m, 22 species: Java, Gunung Gede, 1200-3000 m, 39 species; Sulawesi, Mount Roroka
Timbu, 1200-2400 m, 25 species: Philippines, Mount Apo, 1000-3000 m, 25 species:
Papua, Mount Trikora, 2800-4200 m, 36 species, Idenberg River, 700-2200 m, 22
species; Papua New Guinea, Aseki, 1300-1800 m, 20 species, Ekuti Range at
Watut-Aseki Divide, 2000-2100 m, 22 species, Mount Giluwe, 2100-4100 m, 38
species, Mount Hagen, 2400-4000 m, 34 species, Rawlinson Range, 1500-4000 m, 33
species, Saruwaged Range, 1200-4000 m, 40 species, Star Mountains, 1400-4000 m, 24
species, Mount Suckling, 400-3700 m, 20 species, Mount Wilhelm, 2700-4500 m, 26
202 FERN GAZ. 17(4): 183-203. 2005
species.
The identification of pteridophytes below the subalpine and alpine areas of Mount
Jaya is not yet complete (R.J. Johns pers. comm.), so a complete evaluation of the
relationships of the pteridophyte flora is not yet possible. Ridley (1916) records a
number of pteridophytes from Mount Jaya, however, and this is sufficient to establish
that there is a core of widespread pteridophytes i in families other than Grammitidaceae
that are also present on Mount Kinabalu (Parris ef a/., 1992), Murkele Ridge (Parris,
1993) and Gunung Mulu (Parris, unpublished). They include Asplenium tenerum G.
Forst., Diplazium cordifolium Blume, Lindsaea rigida J. Sm. (Kato, 1992),
Lycopodiella cernua (L.) Pic. Serm., Ophioglossum pendulum L. (Kato, 1989b),
Selliguea enervis (Cav.) Ching and Trichomanes pallidum Blume.
CONCLUSIONS
Mount Jaya has some unique features compared with the other Malesian mountains
considered here: the presence of a rich grammitid flora in the heath forests between 200
and 600 m alt. is unexpected, and there is a good representation of widespread New
Guinea endemic Grammitidaceae. The absence of high altitude endemics in the family
is surprising, given that several other high New Guinea mountains have one or more
endemic species. Mount Jaya is much less species-rich than Mount Kinabalu, and this
may be explained at least in part by the much longer history of collecting on the latter
and the paucity of collections from the largely inaccessible upper montane forest of the
former. Compared with Mount Jaya, the Murkele Ridge, 2000 m lower, supports a rich
flora of Grammitidaceae, but the reason for this is not evident.
Zollinger’s line does not present an insuperable barrier to the distribution of
Grammitidaceae, as 59 (45 %) of the 130 species listed in Tables | to 4 occur on both
sides of it.
The concept of a core Malesian-wide pteridophyte flora present in all suitable areas
has not been suggested previously, but there is ample evidence for it in 23 species of
Grammitidaceae and numerous species in other pteridophyte families.
ACKNOWLEDGEMENTS
I wish to thank Robert Johns and Peter Edwards (K) for organising the loans of Kew
specimens, Alison Paul (BM) for seeking out Kloss’ collections, the Keepers of BM and
K for the loan of material and Ewen Cameron (AK) for facilitating the loans.
REFERENCES
JOHNS, R.J. et al. in prep. A guide to the subalpine and alpine flora of Mount Jaya,
New Guinea. Royal Botanic Garden, Kew.
KATO, M. 1989a. The fern flora of Seram. In: BAAS, P., KALKMAN, K. &
GEESINK, R. (Eds) The Plant Diversity of Malesia, pp. 225-234. Kluwer,
Dordrecht.
KATO, M. 1989b. Taxonomic Studies of Pteridophytes of Ambon and Seram
(Moluccas) Collected by Indonesian- Japanese Botanical Expeditions. III.
Eusporangiate and Some Lower Leptosporangiate Families. Acta Phytotax. Geobot.
0: 77-92.
KATO, M. 1992. Taxonomic Studies of Pteridophytes of Ambon and Seram (Moluccas)
Collected by Indonesian-Japanese Botanical Expeditions. IX. Woodsiaceae,
Lindsaeaceae, and Adiantaceae. J. Fac. Sci. Univ. Tokyo Sec. III, 15: 135-152.
PARRIS: GRAMMITIDACEAE OF MOUNT JAYA, NEW GUINEA 203
KATO, M. & PARRIS, B.S. 1992. Taxonomic Studies of Pteridophytes of Ambon and
Seram (Moluccas) Collected by Indonesian-Japanese Botanical Expeditions. VII.
Grammitidaceae. J. Fac. Sci. Univ. Tokyo Sec. III, 15: 111-133.
PARRIS, B.S. 1983. A taxonomic revision of the genus Grammitis Swartz
(Grammitidaceae: Filicales) in New Guinea. Blumea 29: 13-222.
PARRIS, B.S. 1993. Studies on the Phytogeography and Altitudinal Zonation of the
Pteridophyte Flora of Seram. In: EDWARDS, I.D., MACDONALD, A.A. &
PROCTOR, J. (Eds) Natural History of Seram, Maluku, Indonesia, pp. 75-89.
Intercept, Andover.
ARRIS, B.S. 2001. Taxonomy of Malesian Grammitidaceae in relation to ecology and
phytogeography. In SAW, L.G., CHUA, L.S.L. & KHOO, K.C. (Eds) Taxonomy, the
Cornerstone of Biodiversity, pp 155-160. Forest Research Institute Malaysia, Kuala
Lumpur.
PARRIS, B.S. unpublished. A Checklist of Pteridophytes from Gunung Mulu National
Park.
PARRIS, B.S., BEAMAN, R.S. & BEAMAN, J,H. 1992. The Plants of Mount
Kinabalu. 1. Ferns and Fern Allies. Royal Botanic Gardens, Kew.
RIDLEY, H.N. 1916. Report on the botany of the Wollaston Expedition to Dutch New
Guinea. Trans. Linn. Soc. London (Bot.) 2, 9: 1-269.
WALLACE, A.R. 1869. The Malay Archipelago. Macmillan, London.
204 FERN GAZ. 17(4). 2005
MORPHOGENETIC EVENTS IN THE CERATOPTERIS RICHARDIT
(PARKERIACEAE: PTERIDOPHYTA) SHOOT APEX
M. ROMANOVA! & J. JERNSTEDT”
Department of Agronomy and Range Science, University of California, David,
CA 95616, USA. (Email: 'maromanova@ucdavis.edu; “jernstedt(@davis.edu)
Keywords: Shoot apex meristem, shoot apical cell, leaf apical cell, leaf primordium,
plasmodesmata, morphogenesis, development, dictyostele, protostele
The discovery of developmentally important genes, combined with remarkably regular
cell division patterns, has made the Ceratopteris richardii sporophyte an advantageous
system for developmental studies. Here we report results of an investigation of the
ontogenetic changes in shoot apical meristem (SAM) structure associated with leaf
initiation and histogensis in C. richardii. As described for a number of other ferns, the
shoot apical cell (AC) and leaf apical cell (LAC) of the first leaf of C. richardii embryos
appear simultaneously; all subsequent sporophyte leaves originate from the SAM. An
LAC originates in every immediate derivative of the tetrahedral AC. After this segment
is displaced from the AC by two, more recent derivatives, it undergoes two anticlinal
divisions and one transverse division, setting up the LAC. The pattern of LAC origin
does not change during sporophyte ontogeny, but the rhythm of leaf development does.
For the first leaves of a young sporophyte, the subsequent division of the AC is
correlated with further proliferation of cells of the incipient leaf primordium (LP), and
the AC does not undergo further divisions until the LP is established. Development of
the first 6-9 leaves is followed by a developmental pause of 12-20 days, during which
the shoot apex changes its structure and morphogenetic pattern. LPs are arrested at the
4- or 5-cell stage until displaced some distance from the AC, which continues to
segment and creates an increasing “pool” of early stage LPs. This results in an
elongated shape for the shoot apex. These changes are correlated with changes in the
shoot vascular system. A protostele is found in the young sporophytes with simple
leaves developing without pause; each leaf has a unibundle leaf trace (LT). Dictyosteles
occur in older sporophytes bearing compound leaves; these leaves start to develop when
separated from the AC by a number of incipient LPs, and have 2-5 bundles per LT.
Procambium differentiates simultaneously with LP development: no signs of vascular
differentiation are observed in either the juvenile or the adult shoot apex. Ontogenetic
changes in stelar type might be explained by the loss of meristem identity and
competence to differentiate into procambium by the central cells of the apex. This is in
agreement with Sano et a/. (unpublished observation) that KNOX class | genes are
expressed in the AC and procambium of C. richardii adult sporophytes (but not in the
other cells of the elongated apex). Because plasmodesmata (PD) provide selective
routes for signaling within the shoot apex, we examined PD architecture and
distribution in the C. richardii SAM. All PD in the C. richardii SAM are primary. PD
density in the AC and its immediate derivatives does not change significantly in
sporophyte ontogeny but is about ten-fold higher than reported for dicot SAMs. High
PD density and the resulting intercellular connectedness may compensate for the lack
of ability to form secondary PD. Only the AC and its two immediate derivatives, the
latter being constantly displaced, are interpreted to be undifferentiated in the C.
richardii SAM.
FERN GAZ. 17(4): 205-215. 2005 205
REPRODUCTIVE STRATEGIES AND POPULATION STRUCTURE
IN THE ENDANGERED PTERIDOPHYTE TRICHOMANES
SPECIOSUM (HYMENOPHYLLACEAE: PTERIDOPHYTA)
FJ. RUMSEY’, J.A. BARRETT”, M. GIBBY’, S.J. RUSSELL!
& J.C. VOGEL!
'Dept. of Botany, The Natural History Museum, Cromwell Road, London, UK.
“Dept. of Genetics, University of Cambridge, Cambridge, UK;
*Royal Botanic Garden Edinburgh, 20A Inverleith Row, Edinburgh, UK.
(Email: F.Rumsey@nhm.ac.uk)
Keywords: Glacial refugia, gametophyte, historical biogeography, dispersal, filmy-
ABSTRACT
Trichomanes speciosum Willd. (Hymenophyllaceae) is unique amongst
European ferns in that the gametophyte generation can survive indefinitely in the
absence of the sporophyte, propagate vegetatively and disperse locally.
Reproductive success, both in terms of spore production and sporophytic
recruitment currently vary widely across the species’ broad, disjunct range,
declining from south to north and west to east. Reproductive success would
appear to be controlled largely by climatic factors, although genetic components
also must be considered. Detailed study of populations throughout the species’
range, over a 15 year period, have led to a greater understanding of growth rates,
powers of dispersal and the reproductive strategies currently operating. Using
this knowledge, the extent and pattern of genetic variation regionally and locally,
i.e within sites, can be used to infer the routes and mechanisms of colonisation
and subsequent reproductive history.
Many sites for this species have been considered as glacial refugia, 1.e.,
supporting relictual populations through cycles of glaciation throughout the
Tertiary. The validity of these claims is tested using molecular and other data.
INTRODUCTION
Trichomanes speciosum Willd. (syn. Vandenboschia speciosa (Willd.) Kunkel), the
Killarney Fern, is the sole native European representative of the genus 7richomanes
sensu lato (Tutin et al., 1993) and the largest of the region’s filmy-ferns
(Hymenophyllaceae). It has been considered one of the continent’s most vulnerable
lant species, threatened both by habitat destruction and long the victim of
over-collection, and has been accorded legal protection throughout its range under the
Bern Convention (Anon, 1979) and E.C. Habitats Directive (Anon, 1992).
Trichomanes speciosum is unique amongst European ferns in that its gametophytic
generation is perennial, gemmiferous and capable of persisting and dispersing in the
absence of a sporophyte. This “gametophytic independence’ is apparently a very rare
condition amongst homosporous ferns, and known in three families, all widespread as
epiphytes in tropical regions: the Vittariaceae, the Grammitidaceae, and the
Hymenophyllaceae. However, gametophytic independence is apparently only well
developed in those rare species found at the temperate extremes of these families’
206 FERN GAZ. 17(4): 205-215. 2005
ranges (Rumsey & Sheffield, 1996).
Independent gametophytes of a Trichomanes species were first tentatively identified
from eastern North America by Wagner & Evers (1963) and confirmed by Farrar (1967,
1985, 1992) who identified a distinct taxon, 7 intricatum Farrar (Farrar, 1992)
represented solely by gametophyte populations.This taxon was found to exist far to the
north of the line of the last glacial maximum, indicating it clearly had the ability to
disperse over some distance, even if sporophytes and hence spores were currently not
produced. The other widespread North American species of Trichomanes have been
found as gametophyte populations but always in close proximity to sites in which
sporophytes are present (Farrar, 1993). The situation in Europe falls between these
extremes and differs in scale, in that both generations of 7. speciosum are known, but
the gametophyte has an extensive range (hundreds of kilometres) beyond that of the
sporophyte.
Reproductive Potential and Strategies
The sporophyte of 7 speciosum has a long-running, branching rhizome, at intervals
bearing fronds which are functional for 4 or more years, and may persist for longer.
Rhizome growth rates are heavily dependant on the plant’s nutrient status and climatic
environment but main axes may elongate by over 30 cm per annum, although usually
by much less. Suppressed side branches borne in the sporophyll axils allow for steady
patch formation should the leader not find suitable habitat and die. This growth form
allows a guerrilla-type growth strategy similar to that employed by species of
Lycopodium to exploit patchy environments (Callaghan et al.,1986). Specialised
mechanisms of vegetative propagation are absent (viviparous plantlets have been
recorded twice in cultivated material over two centuries of observation (Druery, 1910;
Ensoll, pers comm.)). Rhizome fragmentation, dislodgement and re-growth are likely to
be infrequent but significant events. Extensive, intricate and usually dense patches form
with time. Within these it is generally impossible to determine the number of
individuals present without recourse to molecular techniques. Individual clones
covering several hundred square metres have been recorded by the authors in the
Azores, obviously indicative of considerable antiquity. However, colony size is not
necessarily an accurate indication of age. Several British colonies known for over a
century are very restricted in size, their maximal extent presumably determined by the
extent of the micro-features that they grow in.
Unlike many European ferns Trichomanes speciosum is not fertile annually once
mature. Indeed Ratcliffe er al., (1993) reported that the majority of British sporophytes
had never been seen to be fertile. Detailed investigation of fertility is difficult and in
many cases not desirable because of the nature of the sites occupied. However, during
the last decade four Welsh and two English sporophytes known to the authors have at
least attempted to produce spores. Involucres have formed but the receptacular bristle
upon which the sporangia are spirally borne has, in at least two of the sites, not extended
and no spores have formed as a consequence. This receptacular bristle structure has a
basal meristem and presumably continues to elongate and produce new sporangia until
a certain trigger, perhaps desiccation causing sporangial dehiscence, causes it to cease.
Its failure to develop in marginal British sites may have climatic cause, such as
exposure to cold. This and the factors determining the onset of fertility are almost
certainly complex and have not been established for 7. speciosum. Anecdotal evidence
would seem to suggest that sporophytes in extreme shade are less regularly fertile and,
RUMSEY et al.: REPRODUCTIVE STRATEGIES IN T: SPECIOSUM 207
as most plants at the more northern and marginal extremes of the species range are
progressively restricted to more sheltered and hence usually darker microsites, this
alone may account for the reduction in northern fertility. Considerable variation exists
in the degree of fertility, i.e. the number of involucres produced — dictated by frond
stature and dissection, and the number of sporangia produced per involucre, as outlined
above. Sporophytes in northern and more easterly British sites are invariably small and
relatively poorly dissected, reducing sporing potential. Calculations based on
observation of the number of fertile fronds known in all British populations for one
year, an estimated mean number of involucres per frond and numbers of sporangia per
involucre, suggested that the total British spore production per decade would be
considerably less than that produced by a single healthy adult Athyrium filix-femina in
one year (Rumsey, 1994) ! The species is thus likely to be a poor coloniser by spore or
vegetative means.
Sporophyte recruitment parallels the pattern of spore production in that there is a
marked reduction in the production of sporophytes northwards, 1.e. into northern France
and the British Isles and even more markedly, eastwards into continental Europe. Thus
within the wetter portions of Macaronesia the species functions as a “normal”
pteridophyte, completing its life cycle from spore to spore, whereas elsewhere the cycle
progressively breaks down, until over much of the species extensive distribution in
northern and continental Europe the gametophyte is effectively functionally
independent. Immature sporophytes have rarely been noted within the British Isles and
show high levels of mortality (Rumsey, 1994), although recent years have perhaps
shown an increased incidence in their production (“Sentinel”, 2003; Rumsey, unpubl.)
which may be a reflection of the run of warm but wet summers and milder winters
experienced in the British Isles. Immature sporophytes have also been reported in
central Europe, eg. Rasbach er al., 1994 which indicates that gametangial production
and normal function are occurring in at least some “independent” gametophytic clones.
Given a potentially indefinite gametophytic lifespan the probability must be that
sporophytes would form, although perhaps extremely infrequently, in the vast majority
of populations.
Sporophytic production and recruitment almost certainly is predominantly a
reflection of the rate and extent of gametangial production. The archegoniophore is a
comparatively massive, differentiated, multicellular structure unlike the simple
irregular branching filamentous structure it is borne on. The processes triggering its
initiation are unknown but rates of cell growth and light/nutrient availability might be
implicated. The smaller antheridia are more regularly produced and may persist in a
non-dehisced state for many months. Rumsey & Sheffield (1996) summarised
knowledge on the in situ and ex situ production of gametangia by different British
gametophyte clones over a 5 year period. They identified consistent sterility, or
super-fertility, of some clones, even when sub-divided and grown apart under similar
but not rigorously controlled growth conditions, suggesting a significant genetic
component to gametangial production, but with no clear geographical, ecological or
discernible historical basis. Locally this may complicate considerations of the effect of
climatic and other external factors on gametangial production but broad regional
differences, which are reflected in sporophyte recruitment, are still apparent.
Understanding the Current Distribution- Refugia or Recent Colonisation?
First detected as recently as 1989 (Rumsey ef al., 1990; 1991). subsequent survey has
208 FERN GAZ. 17(4): 205-215. 2005
revealed the gametophyte to be widespread but extremely localised within the British
Isles, significantly occurring in many areas from which the sporophyte had never been
reported (Rumsey et al., 1998). Similarly, as the distinctive habit and habitat of the
gametophyte became known, discoveries across northern continental Europe rapidly
followed, e.g. Vogel et a/., 1993; Rasbach ef al., 1994; Bennert et al, 1994; Bujnoch &
Kottke, 1994: Kirsch & Bennnert, 1996; Horn & Elsner, 1997; Huck, 1997; Reichling,
1997; Boudrie, 2001; Krippel, 2001, Loriot et a/., 2002, Kottke, 2003, etc.
Initial reports were from deeply dissected sandstone massifs, such as the Petite
Suisse of Luxembourg, the northern Vosges and the Elbsandsteingebirge, all areas noted
for the occurrence of other rare, disjunct ‘Atlantic’ taxa (sensu Ratcliffe, 1968) but
particularly the filmy-fern Hymenophyllum tunbrigense (L.) Sm. (Richards & Evans,
1972). In these disjunct areas H. tunbrigense has been considered to be a Tertiary relict
(Drude, 1902; Klein, 1926). Subsequent examination has revealed the gametophyte of
Trichomanes speciosum to be more widely distributed and in less “exceptional”
landforms and habitats in intervening regions in western and Central Europe, such that
the recorded distribution of the species is now much less discontinuous and more
extensive than previously realised. This raises interesting questions as to when this
broader gametophyte distribution was achieved and from where?
The broad continental and northern-European distribution of Trichomanes
speciosum may have been achieved as a result of a wave of post-glacial migration from
southern and western refugia, or conversely from limited diffusion from a range of
more northerly peri-glacial refugia, or indeed result from an amalgam of both. Sites
containing gametophytes alone may thus represent recent arrivals, or the tenacious
survivors of a species uniquely suited to coping with a sub-optimal climate. Sadly, few
relevant sub-fossil or palynological finds are known, or are realistically to be expected.
Comparison with the distribution of other taxa can be informative, but elucidation of
the species quarternary history is largely reliant on molecular studies of the pattern,
extent and structure of genetic variation throughout the species range. Knowledge of the
comparative reproductive and dispersive capabilities of the different generations can
then be used to infer whether sporophytes were likely to be present historically in areas
from which they are now unknown.
Generally our expectation must be that sites which have been suitable for the
peri-glacial survival of a particular species are likely to have allowed the survival of a
Suite of taxa, and that this community, or many of its distinctive components may well
recur in other disjunct refugial sites. The peculiar biology of Trichomanes speciosum
and its ability to grow in conditions that are not conducive to the survival of all other
vascular plants may, however, mean that it is a special case.
We also expect refugial sites to support greater genetic diversity than those which
have been recently colonised. A potential confounding factor is that of severe
bottlenecking — low or no diversity may result from loss over time/genetic drift, as well
as by founder effects. We must also consider that an_ efficiently
vegetatively-reproducing organism of narrow ecological amplitude might effectively
block niches and thus prevent the arrival of other individuals (“the first in wins”).
Variation may then be reliant on the rates of mutation in the various ramets, although
these would be expected to accrue with time, meeting our original expectation. Ancient
refugial sites, if isolated, might be expected to show private variation, i.e. unique to
them, although post-glacial dispersion, if efficient, will obscure such patterns.
RUMSEY et al.: REPRODUCTIVE STRATEGIES IN T. SPECIOSUM 209
Refugia or Not?
Several disjunct areas are now known to support very small populations of Trichomanes
speciosum, many of which have been discovered comparatively recently. Within these
the relative proportion of sporophytes vs. gametophytes differs but many show
comparably low levels of sporophytic presence and association of a suite of Atlantic
species. Do they and their associated species show similar levels and patterns of genetic
diversity and what can this reveal as to their history?
Southern Spain
Trichomanes speciosum was first discovered in Southern Spain by Richards (1934) and
is now known from a very limited range of sierras leading inland from the coast near
Algeciras, northwards to near Cortes de la Frontera (Rumsey & Vogel, 1998; Garrido
Diaz & Hidalgo Maqueda, 1998). Currently nine populations are known to be extant,
more than half (5) of which are thought to support fewer than 10 discrete sporophyte
clumps each. Only one site (Sierra de Montecoche) currently contains gametophytes in
the absence of sporophytes. The species main concentration is in the Sierra de Ojén and
its westward continuation, the Sierra de Saladavieja. Here a total of approaching 200
distinct mature regularly fertile sporophytic clumps, closely associated with small
localised areas of gametophyte and numerous juvenile sporophytes, occur in 12 distinct
sub-populations by parallel small watercourses. High levels of mortality, with extreme
reductions, at least in sporophyte populations, linked with the cycles of cork-oak
harvesting and climatic variation, have been documented in the recent past
(Molesworth-Allen, 1977).
Allozymic study of material from 3 sites: Sierra de Ojén (4 sub-populations), the
Valle del Rio de la Miel and the Sierra de Montecoche, indicates that each site ha
alleles not shared with the other sites (Table 1) and that the large Sierra de Ojeén site
contained at least 16 different multilocus phenotypes (MLPs), 10 of which were present
in the largest sampled sub-population. The observed differences between closely
adjacent watercourses may indicate low levels of gene-flow even over such restricted
ranges but patterns indicate the likelihood of intergametophytic mating within these
sub-populations, with the fern fully completing its life cycle and behaving ina “normal”
pteridophytic fashion, unlike populations further north in its range.
These small populations co-exist with a limited range of Macaronesian bryophytes
unknown elsewhere in Mainland Europe (Rumsey & Vogel, 1999) as well as other
pteridophyte taxa, including Culcita macrocarpa, Diplazium caudatum, Pteris
incompleta and previously Drvopteris guanchica — the latter suggesting a closest
affinity with the Canarian flora. This floristic enclave, with fragmentary representation
on the adjacent North African coast and into the Rif mountains, either represents the
vestige of a once wider coastal African/C anarian/Iberian distribution fragmented by
aridification and the increase of the Saharan desert area, or, and less likely, represents
independent post-glacial (re-)colonisation by the many and varied Macaronesian floral
elements. All potentially are highly dispersible given the small size of their diaspores.
Alpi Apuane, Italy
Trichomanes speciosum was found, new to the Italian flora, only as recently as 1976
(Ferrarini, 1977). Subsequently a total of five “populations” have been detected
growing within the stream gorge of the Valle de Serra and its side valleys, close to the
town of Serravezza (Marchetti, in Ferrarini et al., 1986). Few discrete individual mature
210 FERN GAZ. 17(4): 205-215. 2005
Table 1. Distribution of multilocus phenotypes (MLPs) in southern Spanish
Trichomanes speciosum populations (figures represent the number of apparently
discrete individuals possessing that MLP).
Allozyme Phenotype Site
IDH | 6-PGD| MDH | Miel | Ojén 1 | Ojén 2 | Ojén 3 | Ojén 4 sina
-cocne
Z
O
=
>
og too 4 (Ml of ne 3) Fa > fo] oo a OO
bo
No
Ww
A ee = |
wl) wl] ml] wm] wm} Pm] PS] SY] SY] SE] YP] PS S| S| CY CF
Co] we me) me] el yO me Om Oe Oe Oe
>| A] DW] Wl S| wl] Se] UO] WW] wD] W| BD! S| | &
| _
sporophytes are known but at least one is extensive and regularly highly fertile. The
known sporophytes share a distinctive and apparently unique morphology.
Initial researches indicate a wider distribution of the gametophyte (Rumsey &
Vogel, unpubl.), perhaps paralleling that of another disjunct Atlantic element, the
filmy-fern Hymenophyllum tunbrigense, for which ca. 70 sites are known in the
communes of Massa, Montignoso, Stazzema, Pietrasanta, as well as Serravezza
(Pichi-Sermolli, 1936; Marchetti, 1992). Sporophytic recruitment has been observed in
sites containing mature sporophytes but not in sites currently only known to support
gametophyte populations.
Unfortunately only a limited range of material has been available for analysis. Both
localities from which multiple samples have been taken support more than one MLP (3
) with patterns consistent with the occurrence of intergametophytic mating.
RUMSEY et al.: REPRODUCTIVE STRATEGIES IN T. SPECIOSUM 211
Sporophytes are currently unknown in the most genetically diverse of these sites. One
allele in the enzyme PGM has not to date been recovered in any other Trichomanes
speciosum population throughout its global range.
All the evidence would thus tend to argue that this area must be considered refugial.
Southwest Scotland
In contrast to the Iberian and Italian sites this area was heavily glaciated and the species
occurrence here must result from post-glacial (re-) colonization. It is thus instructive to
compare the patterns of distribution and extent of genetic variation with the almost
certainly refugial areas above and more equivocal areas.
Fewer than 20 discrete sporophyte plants have been detected in Scotland since its
first discovery in 1863 (Babington, 1863) and less than half this number are known to
be extant. Only one of these colonies has ever been seen to produce spores and most
are, or were, very limited in size and extent. However, the gametophyte generation is
widespread and may be locally abundant. Recent recruitment of sporophytes has been
seen to occur both in sites containing mature sporophytes and also in at least one site
were these were absent.
Genetic diversity of the species at this the northern-most limit of its distribution was
studied by Rumsey et al., (1999). 121 samples from 37 populations in 22 sites were
examined and 7 allozyme MLPs were detected. Two of these MLPs covered large areas
(>30 km in total extent), while the others were restricted to one, or rarely few, localities.
No site was found to contain more than one MLP. Whether this is evidence for blocking
of available niches by the first arrival, or merely a reflection of the extreme rarity of
propagules of other MLPs arriving in occupied sites is unclear and may require
experimental verification through deliberate introduction. The extent of MLPs was not
correlated with presence /absence of sporophytes. Where they extend over many
kilometres, in some cases on separate islands, the only credible means of dispersal is by
spore. Where sporophytes are currently unknown such a distribution indicates their past
presence, and a greater fertility than in the recorded past. Sporophytes may have
occurred as long ago as in the sub-atlantic period, or during brief warm/wet phases
subsequently — such as we appear to be experiencing currently. Very broad distributions
must indicate considerable age given the poor dispersal powers shown by both
generations of this organism. It is tempting to speculate that geographical extent could
be correlated with arrival time but many conflicting factors have also to be considered.
Communities in glaciated and un-glaciated areas share many rare and disjunct taxa
and their presence alone is un-informative as to refugial status, but the population
structure and genetic make-up of these taxa is much more revealing. Many of the
Macaronesian bryophytic taxa show disjunctions to southwest Scotland, but the
majority of these diocious species are represented by one sex alone in the British Isles,
yet exist in mixed sex populations and fruit in southern (refugial?) areas. Such a pattern
might be expected given extremely rare long-range post-glacial dispersal events to the
nort
Central Europe
Rumsey er al., (1998) examined allozymic diversity in 35 sites from the then
easternmost known extent of the species distribution in the Elbsandsteingebirge on the
Czech- German border, through Germany to the French border in the northern Vosges
mountains. The extensive distribution of some MLPs, as in Scotland, indicated that
212 FERN GAZ. 17(4): 205-215. 2005
sporophytes almost certainly must have been present in the past. The majority of sites
supported single MLPs ; however some sites in the pfalzerwald supported multiple
MLPs, some of which are indicative of past inter-gametophytic mating and thus by
inference sporophyte production. From the allozyme data alone Rumsey ef al., (1998)
suggested that post-glacial spread was more likely but were unable to exclude the
possibility that some at least of these locations may have acted as refugia for the
species. The levels of intra-population variation, even though low, and the presence of
unique simple banding patterns for two enzyme systems (ACON, 6-PGD), gave support
to this notion but the proximity of glacial forelands to many of the current localities
suggested survival was less likely (Vogel ef al., 1993). Subsequent work by Rumsey ef
al. (in prep.), sequencing of the chloroplast DNA haplotypes first reported by Rumsey
et al. (1996) has provided additional evidence which would appear to strengthen the
case for, at least limited, peri-glacial survival. Macaronesian island refugia show a
major dichotomy between the Azores in the north, and Madeira and the Canaries to the
south. Minor haplotype variants link some Canarian populations with those from
southern Iberia, supporting the broad floristic links already known. Investigated areas
of Europe on the western Atlantic seaboard from Brittany northwards show a mix of the
haplotypic variants found in the southern refugial sites, although not an intimate one
(i.e. sites contain single haplotypes in all cases). The overall picture created is in
contrast with the usual pattern of Northern Purity — Southern Richness reported by
Hewitt (1999) and others. Interestingly, central Europe is uniformly of the Azorean,
believed to be ancestral, type, in spite of the fact that all of the nearest spore sources
(Spain, Italy) are of the contrasting major haplotype. If the distribution were the result
of post-glacial immigration one might expect a similar pattern to that shown in
northwest Europe, with all of the refugia and particularly the most proximal to be
involved. Unique minor variants also exist within central Europe, for example in
Luxembourg, where they cover an extensive area indicative of a long period of
occupation and/or past sporophyte presence. Extremely severe conditions in proximity
to sites and the growth and reproductive strategies of Trichomanes speciosum may
explain the lack of expected diversity in these putatively refugial areas.
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RUMSEY, F. J., HEADLEY, A. D., FARRAR, D. R. & SHEFFIELD, E. 1991. The
Killarney Fern (Trichomanes speciosum) in Yorkshire. Naturalist 116: 41-43.
RUMSEY, F. J., JERMY, A. C. & SHEFFIELD, E. 1998. The independent
gametophytic stage of Trichomanes speciosum Willd. (Hymenophyllaceae), the
Killarney Fern, and its distribution in the British Isles. Watsonia 22: 1-19.
RUMSEY, F. J., RAINE, C. A. & SHEFFIELD, E. 1992. The reproductive capability of
‘independent’ Trichomanes gametophytes. In IDE, J. M., JERMY, A. C. & PAUL,
A. M., (eds.) Fern a Past, Present and Future Objectives, pp.299- 304.
Intercept, Andov
RUMSEY, F. J, RUSSELL, S. J., JI, J.. BARRETT, J. A. & GIBBY, M. 1996. Genetic
variation in the endangered filmy fern Trichomanes speciosum Willd. In Camus, J.
M., Gibby, M. and Johns, R. J., (eds.) Pteridology in Perspective, pp.161-165. Royal
Botanic Gardens, Kew.
RUMSEY, F. J., SHEFFIELD, E. & FARRAR, D.R. 1990. British filmy-fern
gametophytes. Pteridol. 2: 40-42.
RUMSEY, F. J. & SHEFFIELD, E. 1996. Inter-generational ecological niche separation
and the "independent gametophyte" phenomenon. In Camus, J. M., Gibby, M. and
Johns, R. J., (eds.) Pteridology in Perspective, pp.563-570. Royal Botanic Gardens,
Kew .
RUMSEY, FJ. & VOGEL, J.C. (1998) Trichomanes speciosum willd.
(Hymenophyllaceae) in southern Spain. Fern Gaz. 15: 197-203.
RUMSEY, F.J., VOGEL, J.C., RUSSELL, S.J., BARRETT, J.A. & GIBBY, M. 1998.
Climate, colonisation and celibacy: population structure in Central European
Trichomanes speciosum (Pteridophyta). Botanica Acta 111; 481-489.
RUMSEY, F. J. & VOGEL, J. C. 1999. Tetrastichium fontanum (Mitt.) Card. new to
mainland Europe. J. Bryol. 21: 72-73.
RUMSEY, F. J., VOGEL, J. C., RUSSELL, S. J., BARRETT, J. A. & GIBBY, M. 1999.
Population structure and conservation biology of the endangered fern Trichomanes
speciosum Willd. (Hymenophyllaceae) at its northern distributional limit. Biol. J.
Linn. Soc. 66 333-344.
“SENTINEL” 2003. Killarney fern conservation. Pteridologist 4: 58-61.
TUTIN, T. G, BURGES, N. A., CHATER, A. O., EDMONDSON, J. R., HEYWOOD,
RUMSEY et al.: REPRODUCTIVE STRATEGIES IN T. SPECIOSUM 215
V. H., MOORE, D. M., VALENTINE, D. H., WALTERS, S. M. & WEBB, D. A.
1993. Flora Europaea Vol. 1, 2nd. Ed. Cambridge University Press, Cambridge.
VOGEL, J. C., JESSEN, S., GIBBY, M., JERMY, A. C. & ELLIS, L. 1993.
Gametophytes of 7richomanes speciosum (Hymenophyllaceae: Pteridophyta) in
Central Europe. Fern Gaz. 14: 227-232.
WAGNER, W. H. JR. & EVERS, R.A. 1963. Sterile prothallial clones (7richomanes)
locally abundant on IIlinois sandstones. Amer. J. Bot. 50: 623
216 FERN GAZ. 17(4). 2005
PTERIDOPHYTE DIVERSITY IN LUXEMBOURG
Y. KRIPPEL
Research associate of the Natural History Museum of Luxembourg
Rue de Rollingen, 18A; L - 7475 Schoos, Luxembourg
(Email: yves.krippel@mnhn.lu)
The Grand-Duchy of Luxembourg is internationally known for its small relictual
population of Tunbridge filmy-fern (Hymenophyllum tunbrigense (L.) Sm.), first
discovered here in 1821. In recent decades, however, ferns and allied plants - especially
subspecies and hybrids - have often been neglected by botanists in Luxembourg, so that
reliable data about distribution and status of pteridophyte species is often lacking. Over
the last few years, the study of the pteridophytes of Luxembourg has been relaunched,
and previous deficiencies in information are being reduced progressively.
Despite the small geographic area of Luxembourg (2586 km’), the diversity of
pteridophytes is remarkable. A great number of geological substrates, special
geomorphological features with particular microclimatic conditions and a great variety
of habitats, ranging from acidic to rather base-rich, have favoured the luxuriant
development of a certain number of ferns and fern allies. Not less than 57 pteridophytes
(species, subspecies, nothosubspecies and hybrids) are known for Luxembourg.
Regrettably a certain number of species have not been observed in the recent decades
and can be considered as extinct in wild. Fortunately new taxa have been discovered
recently, such as Trichomanes speciosum Willd. (gametophytes), Asp/lenium
trichomanes L. subsp. pachyrachis (Christ) Lovis & Reichstein, Asplenium
trichomanes L. nothossp. staufferi Lovis & Reichstein, Asplenium x murbekii Dorfler
and Equisetum x litorale Kiihl. ex Rupr. At present the pteridophyte flora of
Luxembourg includes five clubmosses, seven horsetails and 45 ferns. The Red List,
based on the checklist of the pteridophytes of Luxembourg, applies the revised IUCN
Red List categories (IUCN 2001) at a national level. Currently 14.7% of the taxa fall in
the category Regionally Extinct (RE), 6.5% are Critically Endangered (CR), 6.5%
Endangered (EN), 8.2% Vulnerable (VU) and 18% Extremely Rare (R). Only c. 40% of
all pteridophyte taxa are considered not to be threatened (*) for Luxembourg.
Despite the recent discoveries of new taxa, the pteridophyte flora of Luxembourg
has experienced a significant loss of species. This is predominantly due to habitat
changes and resulting climatic variations (habitat destruction, deforestation, changes in
drainage patterns etc.). Management plans for rare and endangered species may prevent
further loss. For critical species and extremely threatened sites, however, it is advisable
to envisage securing the collection through ex-situ conservation and storage of
representative samples for future scientific purposes.
REFERENCES
IUCN 2001. - IUCN Red List Categories and Criteria: Version 3.1. IUCN Species
Survival Commission. IUCN, Gland, Switzerland and Cambridge, UK. ii + 30 pp.
FERN GAZ. 17(4): 217-222. 2005 217
SOME NOTES ON THE TAXONOMY, BIOGEOGRAPHY AND
ECOLOGY OF DANAEA (MARATTIACEAE)
M.J.M. CHRISTENHUSZ' & H. TUOMISTO?’
Department of Biology, University of Turku, FI-20014 Turku, Finland,
(Email: 'maarten.christenhusz@utu.fi, *hanna.tuomisto@utu.fi)
Key words: Biogeography, Danaea, ecology, Marattiaceae, Neotropics, phylogeny,
taxonomy.
ABSTRACT
The neotropical genus Danaea is in revision and some notes on the genus are
presented here. We find that the genus can be divided into three subgroups,
based on morphological characters of the rhizome, stipe articulation and pinna
margin serration. The tree groups are discussed and some taxonomic issues are
addressed. The occurrence of bipinnate D. nodosa is reported from Jamaica, and
trifoliate specimens of D. simplicifolia are reported from French Guiana. The
identities of D. alata, D. jamaicensis, D. jenmanii, D. mazeana, D. media, D.
nodosa, D. trifoliata and D. ulei and their synonyms are discussed. Further
studies on the phylogeny, taxonomy and ecology of Danaea are needed.
INTRODUCTION
The genus Danaea Sm. is one of the few genera of ferns confined to the Western
Hemisphere. It has been repeatedly observed that species limits within the genus are
poorly understood, and that the genus is in great need of monographic study (e.g.,
Morton, 1951, Kramer, 1978). De Vriese & Harting (1853) monographed the
Marattiaceae but excluded Danaea, which was at the time, placed in a separate family:
the Danaeaceae. Underwood (1902) revised the North American species, and the
Ecuadorian species were recently revised by Tuomisto and Moran (2001), but many
taxonomic problems remain. In the recent monograph by Rolleri (2004) a large degree
of synonymization was suggested, with which we do not agree.
In recent years, it has become obvious that information on characters such as plant
habit, rhizome morphology and leaf colour are useful, and often necessary, in
identifying Danaea specimens to the correct species (Tuomisto & Moran, 2001).
Unfortunately, characters like habit and colour are not easily preserved through the
process of pressing and drying, which complicates the identification of herbarium
specimens of Danaea. Furthermore, the rhizome is cumbersome to collect and difficult
to divide between several duplicates. Collecting the rhizome kills the plant in any case.
Consequently, many herbarium specimens, especially those of the larger species, do not
include enough rhizome to allow reconstructing its morphology. Especially historical
specimens, many of which are types, do not include sufficient information about these
characters, either on the specimen itself or on the accompanying label. This has made
it difficult to establish the identity of various species with certainty.
To solve this problem, we have carried out field work in different parts of the
neotropics. As many type localities as possible were visited to establish the identity of
the type specimens. Observing the populations in the field also yields information about
the natural variation in morphological characters, and hence helps in circumscribing the
218 FERN GAZ. 17(4): 217-222. 2005
species. So far, field work has been carried out in Colombia, Costa Rica, Ecuador,
French Guiana, Jamaica, Peru, Puerto Rico, Suriname and the Lesser Antilles. Future
phylogenetic and systematic studies will hopefully give a greater insight in the diversity
and relationships between the species of Danaea.
DISTRIBUTION AND ECOLOGY OF DANAEA
Danaea occurs solely in the New World tropics. It can be found from Mexico, through
all of Central and South America, south to Bolivia. It also occurs on all Greater Antilles,
and the wetter, higher islands of the Lesser Antilles. There are isolated populations in
Atlantic south-eastern Brazil, southern Paraguay and Cocos Island (Costa Rica). The
biogeographical patterns of Danaea are presently being studied.
Danaea occurs mainly in permanently wet forests on well drained soils. Several
species seem to prefer steep slopes or creek banks. Danaea is mostly found in
undisturbed rainforests, although some species tolerate disturbance, and some species
even seem to favour recently disturbed places like shady wet roadside banks. Danaea
species can be found in lowland rainforests, up to an elevation of about 2000 m in
montane cloud forest and elfin woodland, but most species are found between about
300 and 1000 m elevation. In Amazonian lowland rain forests, it has been found that
different species grow on soils with different nutrient status (Tuomisto & Poulsen
1996).
THE THREE MAJOR GROUPS IN DANAEA
Based on morphological characters, we divide Danaea into three major groups. This
division is still tentative, as the groups overlap in some of the characters, and some
species cannot be unambiguously allocated to a group. This division does not follow
any of the sections proposed by Presl (1845), and the phylogenetic relationships
between the species are not yet known although they are being studied.
The first group consists of species that resemble Danaea nodosa and are often
identified as such in herbaria. These are large-statured species whose adults often
exceed 1.5 m in height. Most species are characterized by a creeping rhizome (except
D. erecta Tuomisto & R.C.Moran), the absence of nodes on the stipes (except D. media
Liebm.), and the presence of dentations at pinna apices (except D. erecta and D.
grandifolia Underw.).
The second group consists of species supposedly related to Danaea leprieurit
Kunze. This group of species has previously been united under the name D. elliptica
(see below). These are species of intermediate stature, 0.4-1.5 m in height. This group
is characterized by erect, radially symmetric rhizomes, stipes articulated with swollen
nodes (nodes sometimes lacking in D. bipinnata Tuomisto) and entire pinna apices. The
apical pinna is always present, never replaced by an apical bud.
The third group consists of species resembling Danaea alata Sm, D. moritziana
C.Presl and D. mazeana Underw.. These are species of small to intermediate stature,
0.1-1.5 m in height, and are characterized by radially symmetric (rarely dorsiventral)
rhizomes that can be erect, creeping or ascending. The stipes are nodose, the blade
usually bares many crowded pinnae, and the pinna apices are (coarsely) serrate. Some
of the smaller species have translucent pinnae (D. trichomanoides T.Moore ex Spruce,
D. imbricata Tuomisto & R.C.Moran, D. tenera C.V.Morton), and many species have
bicolorous fronds, with the abaxial side pale green or whitish. The apical pinna can be
replaced by a proliferous bud, but this does not seem to be a distinguishing feature
CHRISTENHUSZ & TUOMISTO: SOME NOTES ON DANAEA Pah
between species. To distinguish species within this group, Underwood used mostly
characters of the venation, which is, in most cases, also highly variable.
Notes on the Danaea nodosa group
Danaea nodosa, the type species of the genus, has always been considered to be
widespread, occurring throughout the range of the genus. However, recent field studies
and close study of herbarium material have revealed, that the name has been applied to
several clearly separate species. Often these species have different ecological
preferences.
The lectotype of Danaea nodosa is a plate in Plumier (1705), which is based on
material either collected “between Morne Rouge and Saint Pierre, Martinique” or ‘‘Port-
de-Paix, Haiti’. The latter locality was selected as the type locality by Underwood
(1909). This was a fortunate choice, because the material we have seen from Haiti
agrees with the type illustration very well: the plants have slender, dorsiventral
rhizomes with two rows of leaf scars, and the pinnae are long and parallel-sided. In
contrast, the plants that grow on Martinique do not match the type illustration: they
have very bulky rhizomes with fronds arranged in several rows, and their pinnae are
narrower and more tapering. It is obvious that the populations on the two islands
represent two different species (Christenhusz, in press).
In Jamaica, most of the plants we have seen agree very well with the material from
Haiti, and can thus be considered true Danaea nodosa. However, we found that leaf
dissection in Jamaican plants varied from the normal once pinnate to fully bipinnate
(Christenhusz & Tuomisto 3194 (IJ, TUR, UCWI). In some populations, the tendency
to bipinnate fronds was stronger than in others, but in all populations where bipinnate
fronds were found, once pinnate fronds were also present. No other differences were
found between the once and twice pinnate fronds than the degree of leaf dissection. It
seems obvious, therefore, that this is a case of within-species variation. An interesting
question concerns the identity of the South American material. We have observed some
differences in pinna shape between the Jamaican and South American populations, but
their rhizomes and habit are similar. Even though we have never observed any tendency
towards bipinnate leaf dissection in any South American population, for the time being
we consider these as D. nodosa.
Field work in Western Amazonia has shown that Danaea nodosa grows on relatively
nutrient-rich, clayey soils. Its juveniles have simple fronds only when very small, and
the fronds become pinnate usually when they are still less than 10 cm long. A very
similar plant but with cartilaginous pinna margins has been found on poorer. more
loamy soils. This species has larger pinnae, especially at the juvenile stage when fronds
often remain simple until they exceed 30 cm in length (Tuomisto & Groot, 1995,
Tuomisto & Moran, 2001). On the basis of a photograph of one of the isotypes, the
latter species was identified as D. u/ei H.Christ. Unfortunately, other isotypes of D. ulei
show clearly that the rhizomes in this species are erect, indicating that it belongs to the
D. leprieurii group. Its pinna margins are not cartilaginous, either. Consequently, the
poor-soil segregate of D. nodosa needs a new name (Christenhusz & Tuomisto, in
press).
In Costa Rica, populations of supposed Danaea nodosa were found to be variable
in whether the stipe is nodose or not. Juveniles usually had one or two nodes, but older
fronds more often had none. The presence of nodes was variable even in fronds on the
same rhizome, which shows that the two forms belong to the same species. These plants
220 FERN GAZ. 17(4): 217-222. 2005
have previously been identified either as D. nodosa or as D. elliptica, depending on
whether nodes were present or not, as this has traditionally been the most important
diagnostic character between the two species. Since true D. nodosa never has nodes, the
names that can be applied to this species are D. media Liebm. and D. elata Liebm.
Danaea media was described on the basis of small plants (with nodes) and D. e/ata on
the basis of a large specimen (without nodes) from the same population in Mexico.
These names have earlier been synonymized with D. elliptica and D. nodosa,
respectively, but our observations indicate that they represent another taxon worthy of
recognition at species level. We propose that the name D. media is used as the valid
name for this species, because this epithet refers to the species having intermediate
characters between the D. nodosa and D. leprieurii groups.
Notes on the Danaea leprieurii group
Most of the species in this group have at some stage been identified as Danaea elliptica
Sm. in Rees, which has been considered a wide-spread and common species. The
identity and typification of D. elliptica has however long been in turmoil. The lectotype
designated by Proctor (1977) is: Jamaica, Mount Diablo, Herb. Sloane 1: 85 (BM-SL).
It has been observed repeatedly that this specimen is actually a juvenile of D. nodosa
(L.) Sm (e.g., Lellinger 2000, Tuomisto & Moran 2001). We have confirmed this by
visiting Mount Diablo, where we only found individuals of D. nodosa. The issue of the
synonymization of D. elliptica is discussed in more detail in Christenhusz & Tuomisto
(in press).
As mentioned above, Underwood selected Haiti rather than Martinique as the type
locality of Danaea nodosa. \t is worth mentioning that since the two islands have
different species of the D. nodosa group, the situation would now be very different had
Underwood selected the other way round. If Martinique were the type locality, D.
nodosa would be represented by a type illustration that matches the species occurring
on that island rather poorly, and D. elliptica would become the valid name for the
species that has so far been known as D. nodosa in the Greater Antilles and South
America, which would be very confusing.
Due to the problems with the name D. elliptica, we call the present group of species
the D. leprieurii group, as this is one of the oldest accepted names in this group. This
group includes many apparently closely related species that are morphologically quite
similar, but grow on different soil types and/or altitudes.
Danaea leprieurii was described from French Guiana, but recently also material
from western Amazonia has been allocated to this species. (Tuomisto & Moran 2001).
In western Amazonia, D. leprieurii grows on relatively poor substrates, whereas the
recently described and morphologically rather similar D. bipinnata grows on richer
soils. The two can be distinguished by size (D. /eprieurii is smaller), the number of
nodes on the stipes (2-3 in D. leprieurii, 0-1 in D. bipinnata), and usually the degree of
lamina dissection. Danaea bipinnata was described from lowland Amazonia, but it is
very similar to D. elliptica Sm. var. crispula Rosenst, which was described from lower
montane elevations in the Peruvian Andes. However, D. elliptica var. crispula is never
bipinnate, and it is usually larger than D. bipinnata.
Danaea simplicifolia Rudge is confined to the Guiana shield, and can easily be
recognized by its simple fronds. The only other species with simple fronds is D.
carillensis H.Christ, an endemic of Costa Rica, with a nodose stipe, creeping rhizome
and apical dentations, and therefore belonging to the D. alata group. D. simplicifolia
CHRISTENHUSZ & TUOMISTO: SOME NOTES ON DANAEA #21
only has a single node below its simple blade. From this node, sometimes one or two
lateral pinnae are produced (French Guiana: Christenhusz 2325 (CAY, TUR),
Christenhusz 2415 (CAY, TUR)). It then resembles D. trifoliata Rchb., a species that
can be distinguished by its nodose stipe. Another character that distinguishes D.
simplicifolia from juvenile Danaea that have not developed lateral pinnae yet, are the
white abaxial surfaces of the blades, an uncommon feature in this group. It is only found
in some representatives of the D. alata group and in the Ecuadorian D. bicolor
Tuomisto & R.C.Moran.
Notes on the Danaea alata group
Danaea alata Sm. is a species confined to the Lesser Antilles (lectotype from
Martinique, Plumier 1705). It is recognized by simple, widely spaced veins, nodose
stipes, creeping, radial rhizomes and coarse apical dentations. Its synonyms are D.
fendleri Underw. (Trinidad) and D. stenophylla Kunze (Guadeloupe).
At a first glance the closely related Danaea mazeana Underw. appears to be
confined to Guadeloupe, but material from other Antilles is very variable. D. mazeana
fits well within the variation of D. jamaicensis Underw., and cannot be consistently
separated from it. However, D. jamaicensis sometimes has apical proliferations, which
are never found in D. mazeana. This might be due to possible hybridization with D.
jenmanii Underw. on Jamaica. D. mazeana differs in ecology from D. alata, as the
former is confined to cloud forests above 600 m, whereas the latter is found only in
rainforests at lower altitudes.
On present evidence, Danaea jenmanii and D. wrightii Underw. are conspecific.
The type of D. wrightii is somewhat scalier, but all other characters overlap. Field work
in Jamaica showed that the scalyness of D. jenmanii is variable. Because D. jenmanii
has been applied more widely than D. wrightii, the first is proposed here for
conservation over the latter. Both species were described in the same publication.
CONCLUSIONS
Field work and herbarium studies have shown that the diversity of Danaea is much
greater than previously assumed. The first indication of this tendency was the
Ecuadorian treatment (Tuomisto and Moran 2001) where 18 species were included,
eight of which were described as new. The same tendency is continuing in other areas,
and we now have several species in publication (Christenhusz, in press; Christenhusz
& Tuomisto, in press). It seems obvious that the concepts of the geographical
distribution ranges of the species will have to be changed once the taxonomy of the
genus has been sorted out. More species than expected seem to have rather narrow
ranges.
REFERENCES
CHRISTENHUSZ, M.J.M. in press. Three new species of Danaea (Marattiaceae) from
French Guiana and the Lesser Antilles. Ann. Bot. Fennici.
CHRISTENHUSZ, M.J.M. & TUOMISTO, H. in press. Five new species of Danaea
(Marattiaceae) from Peru and a new status for D. elliptica. Kew Bulletin.
KRAMER, K.U. 1978. The Pteridophytes of Suriname, an enumeration with keys of the
ferns and fern-allies. Uitgaven “Natuurwetenschappelijke studiekring voor
Suriname en de Nederlandse Antillen” 93:17-18, Utrecht.
LELLINGER, D.B. 2000. On the lectotypification of Danaea elliptica. Amer. Fern J.
poe FERN GAZ. 17(4): 217-222. 2005
90(3): 100-103
MORTON, C.V. 1951. A new fern of the genus Danaea from Colombia. J. Wash. Acad.
Sci. 41(8): 276.
PLUMIER, C. 1705. Lingua cervina nodosa major. Traité des Fougéres de
l’ Amériques, t. 108, page 90
PRESL, C.B. 1845. Genera Filicacearum. Supplementum Tentaminis Pteridographiae:
34-39. Praha.
PROCTOR, GR. 1977. Marattiaceae. In: HOWARD, R.A. Flora of the Lesser Antilles,
Leeward and Windward Islands, vol. 2, Pteridophyta: 45-49. Arnold Arboretum,
Harvard University, Jamaica Plain, MA.
ROLLERI, C.H. 2004. Revisién del Género Danaea (Marattiaceae-Pteridophyta).
Darwiniana 42 (1-4): 217-301
ROSENSTOCK, E. 1907. Beitrage zur Pteridophytenflora Siidbrasiliens. Hedwigia 46:
162-164
SMITH, IE. 1808. Danaea elliptica, in REES, Cycl. 11: Danaea No. 2.
TUOMISTO, H. & GROOT, A. T. 1995. Identification of the juveniles of some ferns
from western Amazonia. Amer. Fern J. 85(1): 1-28.
TUOMISTO, H. & MORAN, R.C. 2001. Marattiaceae. In: HARLING, G &
ANDERSON, L. (eds.) Fl. Ecuador 66: 23-68.
TUOMISTO, H. & POULSEN, A. D. 1996. Influence of edaphic specialization on
pteridophyte distribution in neotropical rain forests. Journal of Biogeography 23:
283-293.
UNDERWOOD, L.M. 1902. ens Ferns — V. A Review of the Genus Danaea. Bull.
Torrey Bot. Club 29 (12): 669-679.
UNDERWOOD, L.M. 1909. Family 1. Marattiaceae. In: BRITTON, N., MURRILL, W.
& BARNHART, J. (eds.) N. Amer. Fl. 16 (1): 17-23.
VRIESE, W.H. DE & HARTING, P. 1853. Monographie des Marattiacées. Noothoven
van Goor, Leiden & Arnz, Diisseldorf.
FERN GAZ. 17(4): 223-234. 2005 223
TEMPORAL VARIATION IN SPOROPHYTE FERTILITY IN
DRYOPTERIS INTERMEDIA AND POLYSTICHUM ACROSTICHOIDES
(DRYOPTERIDACEAE: PTERIDOPHYTA)
J.M. SHARPE
SHARPLEX Services, PO Box 499, Edgecomb, Maine 04556 USA
Key words: annual variation, demography, fern, Dryopteris intermedia, crown fertility,
plant fertility, growth, Polystichum acrostichoides, pteridophyte, temperate zone
ABSTRACT
A long-term demographic study to assess temporal variation in two common
New England ferns, Dryopteris intermedia (Evergreen wood fern) and
Polystichum acrostichoides (Christmas fern) focused on fertile leaf production
by the reproductively mature sporophyte. Over an eight-year period only 29.9%
of the leaves in a D. intermedia crown were fertile, while for P. acrostichoides
significantly more (48.0%) of the leaves in the crown were fertile. Annual values
of the percentage of plants with fertile leaves for D. intermedia ranged from 42%
to 77% and for P. acrostichoides from 57% to 93% reflecting significant annual
variation, possibly related to winter weather conditions. Only 16% of D.
intermedia and 14% of P. acrostichoides sporophytes were fertile every year of
the study. Transitions from fertile plant status to sterile plant status occurred in
15% of the D. intermedia observations and 14% of the P acrostichoides
observations. For both species, when a sterile year followed a fertile year the
number of leaves in the crown decreased by approximately half a_ leaf.
Determining the causes of such high levels of variability in plant fertility will be
necessary before the role of ferns in any ecosystem can be fully understood.
INTRODUCTION
Long-term monitoring of marked individuals can provide insight into the life history of
a species (Harper, 1977) as well as providing a solid basis for future experimental
research (Noss, 1999). Werth & Cousens (1990) emphasized the importance of
long-term studies of ferns in order to understand mortality and differential fitness, and
noted that very little information is available on the life spans and generation times of
ferns. There have been several short-term studies of the life history of sporophytes of
specific temperate understory fern species. Field research on various aspects of the
temperate fern life cycle has been reported for Blechnum spicant (L.) Smith (Cousens,
1973, 1981), Thelypteris dentata (Forsskal) E. P. St. John, Woodwardia virginica (L.)
Smith, Osmunda regalis L. (Bartsch & Lawrence, 1997) and Polystichum
acrostichoides (Michaux) Schott (Greer & McCarthy, 2000). Studies exceeding 12
years in length have been conducted on selected species of Botrychium (Montgomery,
1990: Johnson-Groh, 1999). Sato (1990) followed leaf production for more than nine
years in Polystichum braunii (Spenner) Fée, P. tripteron (Kunze) Pr.and Drvopteris
crassirhizoma Nakai. Few other long-term studies have considered annual variation in
temperate fern life history characteristics.
Of the seven fern life history classes identified by Cousens ef al. (1988) the
reproductively mature sporophyte class has the greatest impact on ecosystem processes.
Spore production, controlled by the mature sporophyte, is a significant element of the
224 FERN GAZ. 17(4): 223-234. 2005
reproductive effort for ferns (Greer & McCarthy, 2000) and may also limit
establishment of the next generation (Peck, 1980). Though few quantitative studies
have addressed temporal variation in fertile leaf production it is assumed that most
mature ferns reliably release spores every year (Sheffield 1996). Siman ef al. (1999)
state that fertility is the rule for mature individuals of most ferns but note that the genus
Pteridium is an exception. A few short-term studies have also noted variation from this
generally accepted assumption. Cousens (1988) reported that marked adult fertile plants
had been observed reverting to adult sterile plant status. This was based on a study of
Woodwardia areolata (L.) T. Moore (= Lorinseria areolata (L.) C. Presl.) which lasted
30 months (Cousens, ef al., 1988). Sato (1990) used counts of number of veins off the
leaf midrib (NV) to assess chronological age. He observed a year-to year decrease in
NV in about 20% of comparisons of overwintered leaves to the newly emerged leaf
cohort in individuals of D. crassirhizoma and P. tripteron, but not in P. braunii. Greer
& McCarthy (2000) found that not all reproductively mature individuals of P.
acrostichoides produced at least one fertile leaf in each of the two years of their study.
This exploratory study of the reproductively mature sporophyte of two temperate
species was undertaken to extend our knowledge of annual variation in plant fertility in
a long-term study lasting eight years. The goals were to 1) determine what percentage
of the leaves produced by an individual each year were fertile, 2) assess the number of
plants in a sample population that produced at least one fertile leaf from year to year, 3)
examine levels of consistency in the annual sequence of fertile and sterile leaf
production, 4) compare patterns of temporal variation for two different species, and 5)
compare any observed temporal patterns to relatively short term local climate
parameters.
STUDY SPECIES
Two temperate ferns common in northeastern forests of the USA were chosen for
comparative study: Dryopteris intermedia (Willdenow) A. Gray, the Evergreen wood
fern, and Polystichum acrostichoides, the Christmas fern. Leaves of D. intermedia are
described as monomorphic by Montgomery & Wagner (1993) while those of P
achrostichoides exhibit strong sterile-fertile leaf dimorphism, with the fertile leaf
exhibiting a distinctive terminal segment of spore-bearing leaflets that are greatly
reduced in size compared to the proximal sterile leaflets (Wagner, 1993). Both species
have short internodes so that the leaves form a distinctly tufted crown around the shoot
apex.
MATERIALS AND METHODS
Sporophyte fertility was observed over an eight-year period from 1993 to 2000 in a
secondary, mixed-hardwood forest approximately 15 m above sea level in the town of
Dresden, Lincoln County, Maine (USA) using methods developed for long-term studies
of fern demography in a tropical forest at the Luquillo Long Term Ecological Research
site in Puerto Rico (Sharpe, 1997). The sample population consisted of 31 individuals
of D. intermedia and 14 individuals of P acrostichoides which were considered
reproductively matures as they had produced spores at least once during the eight-year
study period. Of those monitored, 24 individuals of D. intermedia and nine individuals
of P. acrostichoides were fertile in 1993, the first year of the study. Each plant was
identified by a numbered plastic stake and a small segment of colored, plastic-covered
wire attached to a leaf base and transferred annually to one of the leaves in the next
SHARPE: TEMPORAL VARIATION IN SPOROPHYTE FERTILITY 225
year’s cohort of leaves. Annual observations of sporophytes were made during the last
week in August to insure that all leaves had fully expanded.
All leaves in the current year cohort were counted and each leaf was classified as
sterile or fertile. To calculate a crown fertility perecentage for each sporophyte, the
number of fertile leaves was compared to total number of leaves in the crown.
Percentage of the plants fertile each year was determined by comparing the number of
plants with at least one fertile leaf present in the crown to the total number of plants
observed each year. Temporal patterns of fertile leaf production were further evaluated
by noting the sequence of changes in plant fertility status, observed seven times within
the eight-year monitoring period for each plant. Year-to-year transitions in plant fertility
status were classified as FF (fertile to fertile), SS (sterile to sterile), FS (fertile to sterile)
and SF (sterile to fertile). These categories are illustrated with an example of an
individual plant which happened to experience all four types of plant fertility transitions
(Figure 1). The number of continuous years a plant was sterile following and preceding
a fertile year was also noted.
In order to evaluate whether relatively short term weather patterns could have an
effect on plant growth, values for mean daily minimum and maximum temperatures,
precipitation and snowfall were taken from records of the NOAA weather station
located within 100 km of the study site in Portland Maine. Climate data were analyzed
separately for summer (May through October) and winter (November through April).
These analyses did not show significant variation in mean daily summer temperatures
and precipitation. However, winter temperatures had significantly lower mean daily
minimums in 1994 and 1996. Snowfall was significantly higher in the winter of 1996.
Furthermore, there was a major cyclic disturbance in the form of an ice storm in
4
INDIVIDUAL PLANT FERTILITY: DRYOPTERIS INTERMEDIA #17
6 =
| FERTILE LEAVES
5 | M STERILE LEAVES | __ —— =
>
n
NUMBER OF LEAVES
No Ww
1993 1994 1995 1996 1997 1998
YEAR
Figure 1. Profile of the fertile and sterile leaves present in the crown of a sample
individual (#17) of Drvopteris intermedia for each of the eight years the plant was
monitored. Plant fertility status transitions are noted between bars (SS = Sterile Sterile,
SF = Sterile/Fertile, FF ~ Fertile/Fertile, FS = Fertile/Sterile) for this sporophyte which
happened to experience all four of the possible year-to-year transitions.
226 FERN GAZ. 17(4): 223-234. 2005
February 1998 (Darwin ef al., 2004).
Data analysis
Means and standard error of the means (SE) were calculated for crown fertility for each
plant using data collected each year (N = 8). Standard error of the means were chosen
to show variation because of the small sample sizes. All percentages were arcsine
transformed before statistical tests were applied. To factor out variability among
marked individuals, Repeated Measures Analysis of Variance (RMA) was used to
evaluate year-to-year differences in the percent of crown fertile. A one sample t-test was
used to compare plant fertility and transition type percentages from year to year for each
species. Paired t-tests were used to compare the different fertility characteristics for the
two species. Data were analyzed using Statistix 8 (Analytical Software, 2003).
RESULTS AND DISCUSSION
Crown fertility
In eight years of monitoring these populations, observations of crowns that were
composed entirely of fertile leaves were unusual for both species. Of 360 individual
annual flushes of leaves observed in this study, only eight individual leaf cohorts of
Dryopteris intermedia and 18 of Polvstichum acrostichoides were completely fertile.
Fewer than half the leaves produced by these reproductively mature plants were fertile
and the mean percentage of leaves in the crown that were fertile is significantly lower
for D. intermedia than for P. acrostichoides (Table 1). Neither species in this study
approached 100% fertile leaf production. This could be a function of the chronological
age of these plants, which is unknown. If they are all relatively young plants, perhaps
the ability to produce a full cohort of fertile leaves had not yet occurred. If they are all
relatively old plants, then possibly the ability to produce a full cohort of fertile leaves
had been lost. Although no increasing or decreasing pattern can be seen by comparing
the percentages from year to year, perhaps eight years is too small a segment of the total
life span to observe such patterns. It is also possible that full fertile leaf production is
not possible in the specific habitat where these plants were growing. It is possible that
each species has an intrinsic limit on fertile leaf production by reproductively mature
plants which integrates its habitat requirements and need for vegetative support tissue.
While there are reports of high percentages of crown fertility in some temperate ferns,
for example in Leptopteris hvmenophylloides (Sussex, 1958), low crown fertility has
also been observed in Matteuccia ol si (Von Aderkas & Green, 1986). In a
study undertaken in southern eria, Gureyeva (2001) found that environmental
factors may severely limit spore nigra ae by ferns and Cousens (1973) noted
population differences in fertile leaf production by Blechnum spicant.
There were year-to-year differences in crown fertility. The annual mean percentage
of leaves that were fertile within the crown ranged from 21.9% to 38.9% for D.
intermedia and from 37.6% to 60.6% for P. acrostichoides with little concordance
between species in the pattern of annual increases and decreases (Table 1). Dryvopteris
intermedia had the highest total fertile leaf production in 1994 following one of the two
coldest winters of the study and had the lowest total fertile leaf production following
the ice storm in 1998. In contrast, P acrostichoides had the lowest fertile leaf
production in 1996 following one of the two coldest winters of the study and exhibited
no unusual response after the ice storm. Investigations into the specific triggers for
fertile leaf production have been recommended by Wagner & Wagner (1977) and could
identify potential environmental factors involved. For example, it has been
SHARPE: TEMPORAL VARIATION IN SPOROPHYTE FERTILITY 22]
TABLE 1. Summary of annual variation in crown and plant fertility for reproductively
mature sporophytes of Dryopteris intermedia and Polystichum acrostichoides. N is the
number of plants measured. One-sample t-tests are used to compare years for
percentage of plants and transition type percentages. Repeated measures analysis of
variance is used to compare years for individual percentage of crowns fertile. Paired
t-test is used to compare the year-to-year differences between species. Significant
differences are indicated by * (P < 0.0500), ** (P < 0.0100) and *** (P < 0.0010).
Year Fertile leaves in crown Percentage — Transition type percentages
N_ Percentage of of plants
crown (mean) fertile SS SF FF FS
Dryopteris intermedia
1993 1 35.1+4.87% 77% - - - -
1994 31 38.945.88% 71% 10% 13% 58% 19%
1995 31 33.9+5.09% 71% 13% 16% 55% 16%
1996 31 22.945.11% 48% 23% 7% 42% 29%
1997 31 35.1+4.63% 77% 23% 29% 48% 0%
1998 31 21.945.35% 42% 23% 0% 42% 36%
1999 31 23.8+4.58% 55% 39% 19% 36% 7%
2000 31 27.344.12% 68% 26% 19% 48% 7%
All 248 29.9+1.68% 64% 22% 15% 47% 15%
F/t F = 1.88 t= 11.52 t=6.11 4.10 1449 3.28
df 7 7 6 6 6 6
P 0.0748 <0.0001 0.0009 0.0063 <0.0000 0.1637
EK RK *K RK
Polystichum acrostichoides
199 49 .6+10.64% 64% - ~ as ~
1994 14 41.4+9.73% 71% 23% 23% 46% 23%
1995 14 44.2+10.85% 71% 15% 15% 54% 15%
1996 14 37.6£11.99% 57% 33% 0% 58% 8%
1997 14 50.4411.28% 71% 17% 25% 50% 8%
1998 14 52.7+10.27% 79% 14% 21% 57% 7%
1999 14 60.6+9.14% 93% 0% 21% 71% 7%
2000 14 47.4+9.01% 71% 0% 7™% 64% 29%
All 112 48.0+1.71% 72% 12% 16% 58% 14%
F/t F = 0.56 t= 12.97 t=284 457 322 4.19
df 7 | 6 6 6 6
F 0.7891 <0.0001 0.0295 0.0038 <0.0000 0.0057
kK * *x RK **
Species comparisons
t -4.57 =].22 1.22 -0.31 -1.76 0.38
df 6 6
| ey 6 6
P 0.0026 0.2603 0.2696 0.7702 0.1298 0.7186
**
228 FERN GAZ. 17(4): 223-234. 2005
demonstrated in greenhouse studies (Siman & Sheffield, 2002) that under field
conditions individuals do not necessarily produce the maximum number of fertile
leaves possible each year. In a year-long study of Polypodium vulgare L. in the United
Kingdom, field-grown and greenhouse-grown plants from identical genetic stock were
compared by Siman & Sheffield (2002). The indoor population showed a four-fold
increase in new leaves, mostly fertile, emerging in a wave pattern thought to represent
three annual cohorts for field grown plants as compared to the outdoor population.
There do appear to be limits to fertile leaf production which may be consistent within
species over an extended period of time. Experiments similar to this one done by Siman
& Sheffield (2002) would confirm these limitations.
Plant fertili
Of 248 individual leaf cohorts of D. intermedia observed during the eight years of the
study, 64% included at least one fertile leaf, while for 112 cohorts of P. acrostichoides
the percentage was 72% (Table 1). The percentage of plants in the population that
produced at least one fertile leaf each year ranged from a low of 42% in 1998 to a high
of 77% in 1993 for D. intermedia, a range from the lowest year to the highest of 35%,
reflecting significant year-to-year variation. For P. acrostichoides there were also
significant year-to-year differences, with a range from the lowest year (1996) to the
highest year (1999) of 36% (Table 1). For D. intermedia, the lowest percentage of plant
fertility occurred following the ice storm. A paired comparison of annual plant fertility
percentages did not detect a significant difference between the two species (Table 1).
Annual sequence of plant fertility status
There are some elements of life history than can only be determined by making repeated
observations of marked individuals over a long period of time. One of those elements
is the year-to-year change in an individual sporophyte’s fertility status. Overall, a plant
was sterile one year as well as the next (SS) 48 times in the 217 observations of
transition for D. intermedia (Table 1). Of these SS transitions, 9 (3 plants) occurred
before a plant produced its first fertile leaf while for all others (81%) the SS transition
occurred after the plant had produced at least one fertile leaf. For P acrostichoides, SS
transitions occurred 11 times out of 98 transition observations, with one plant (of 14)
accounting for all of the SS transitions that occurred before a fertile leaf was produced.
Thus 89% of the SS transitions for P. acrostichoides occurred after a fertile leaf had
been produced. Sterile to fertile (SF) transitions occurred in 15% of the D. intermedia
observations and 16% of the P. acrostichoides observations (Table 1).
Fertile to fertile (FF) transitions occurred in 47% of the D. intermedia observations
and 58% of the P. acrostichoides observations (Table 1). A small percentages of all D.
intermedia plants observed produced fertile leaves only once or twice, while all P
acrostichoides individuals produced at least one fertile leaf during three or more years
(Table 2A). Although all plants in this study were reproductively mature, only 16% of
the individuals of D. intermedia and 14% of the individuals of P. achrostichoides
produced at least one fertile leaf during each of the eight years of the study (Table 2A).
Of the sporophytes of D. intermedia which were fertile at the start of the study in 1993,
75% were fertile in 1994 while only 21% were consistently fertile through 2000, the last
year of the study (Table 2B). Of the sporophytes of P. acrostichoides which were fertile
in 1993, 67% were fertile in 1994, compared to the 60.8% sequential reproducers
observed in a two-year study of this species in southeastern Ohio by Greer & McCarthy
(2000). By 2000, only 22% of the P. acrostichoides the individuals that were fertile in
1993 could be called sequential reproducers. Thus, as the number of years a plant is
SHARPE: TEMPORAL VARIATION IN SPOROPHYTE FERTILITY Zao
monitored increased, the percentage of sequential reproducers decreased at
approximately the same rate for both species (Table 2B).
Fertile to sterile transitions (FS) reverse the more common trend of fertile-to-fertile
transitions in the chronological maturation of a reproductively mature sporophyte
described by Sheffield (1996). However, plants that were fertile one year and produced
TABLE 2. Comparison of fertility and sterility frequencies for reproductively mature
sporophytes of D. intermedia and P. acrostichoides observed for an eight-year period
from 1993 to 2000. Table A includes all plants which became fertile during the
eight-year monitoring period. Table B includes only plants which were fertile in 1993.
Years D. intermedia P. acrostichoides
A. Number of years that individuals are fertile
l 7% 0%
2 3% 0%
a 16% 21%
+ 12% 7%
a 16% 21%
6 19% 14%
7 9% 21%
8 16% 14%
B. Number of consecutive years that fertile individuals remain fertile
l 100% 100%
= 75% 67%
5 63% 67%
4 38% 56%
5 38% 56%
6 29% 44%
7 25% 44%
8 21% 22%
C. Number of fertile/sterile transitions per individual
l 54% 50%
2 32% 37%
E 14% 13%
D. Span of sterile years after and before a fertile year
Ap WN
xe
oS
S
=
230 FERN GAZ. 17(4): 223-234. 2005
no fertile leaves the next year (FS) were seen in 15% of the D. intermedia observations
and 14% of the P. acrostichoides transition observations. For D. intermedia, 70% of the
individuals reverted to sterile status from fertile status (FS) at least once during the
eight-year monitoring period, while 57% of P. acrostichoides individuals had FS
transitions. For approximately half of the plants of both species an FS transition
occurred only once (Table 2C). Given that a plant must transition back to fertile status
in order to become sterile again, the maximum number of FS transitions possible in an
eight-year study is three. This maximum number of FS transitions was noted for 14%
of the observations for D. intermedia and 13% of the FS observations for P.
acrostichoides. The span of years that a previously fertile sporophyte remained sterile
ranged from one to five for D. intermedia and one to four for P. acrostichoides (Table
2D). For both species the majority (60%) remained sterile for only one year (Table 2D).
Of the FS transitions for Drvopteris intermedia (Table 1), most occurred in 1998 (36%),
following the ice storm and in 1996 (29%), following a winter with very low daily
minimum temperatures (Table 1). For P. acrostichoides most FS transitions (29%)
occurred in 2000 (Table 1).
Crown size and changes plant fertility status
Changes in crown size can be related to the annual transitions to and from fertile leaf
production. Plants maintained a stable crown size following FF transitions (Table 3).
Crown size was also stable for SS transitions for D. intermedia, though it increased for
P. acrostichoides. However, following FS transitions, individual plants produce a
significantly smaller sterile crown by over half a leaf (Table 3). Following SF
transitions, the increase in the size of the fertile crown is also significant for D.
intermedia, with an increase in P. acrostichoides as well (Table 3). Thus, even for the
same individuals, the number of leaves in the crown increases with a shift from plant
sterility to fertility. On a smaller scale than that observed experimentally by Siman &
Sheffield (2002), it appears that when field conditions favor above-average fertile leaf
production for D. intermedia and P. acrostichoides, more leaves are produced.
TABLE 3. Comparison of changes in crown sizes for transitions in plant fertility sta-
tus from one year to the next. Significant differences are indicated by * (P < 0.0500)
and ** (P< 0.0100).
Transition Dryopteris intermedia Polystichum acrostichoides
ean t F N ls
difference difference
Fertile/Fertile 101 -0.04040.186 -0.21 0.8314 53 -0.07640.294 0.26 0.7982
Sterile/Sterile 48 -0.02140.210 -2.25 0.0308* — 11 +0.727+40.304 -1.13 0.2793
Fertile/Sterile 35 -0.57140.254 3.20 0.0031** 13 -0.53940.475 1.16 0.2654
Sterile/Fertile 33 +0.727+0.227 -0.10 0.9212 15 +0.33340.287 2.39 0.0379*
Min/Max -4/+6 -7/+5
F 3.57 bore
P 0.0149* 0.3438
SHARPE: TEMPORAL VARIATION IN SPOROPHYTE FERTILITY 231
CONCLUSIONS
The most striking observations to emerge from this temporal study of annual variation
in individual plant fertility are 1) the overall low level of fertile leaf production in both
species with very few observations of whole crown fertility, 2) the inconsistency of the
percentage of the crown which is fertile from year to year, 3) significant differences in
the number of plants that are fertile from year to year for each species, a phenomenon
possibly linked to winter weather conditions, 4) the difference between species in the
overall percentage of plants that are fertile 5) the similarities between species with
respect to the annual patterns in changes of the percentages of fertile plants and
transitions in plant fertility status 6) the significant year-to-year variation in the percent
of the population that experiences a fertile to fertile plant status transition, 7) expansion
of individual crown size when fertile leaves are produced, and reduction of crown size
when a fertile plant fails to produce fertile leaves the following year. Most of these
characteristics of fertile leaf production could not have been known without annual
monitoring of marked individuals. Though eight years seems to be a relatively short
time compared to the life span of long-lived sporophytes, it was possible to quantify
several previously untested assumptions about sporophyte growth.
The original intention of this study was to determine if annual variability in plant
fertility could be detected in field populations of sporophytes of temperate species. With
the exception of general observations relating to significant local weather patterns, no
attempt was made to determine the cause of any observed variation. It is certainly clear
from these results that there can be a high level of annual variability in fertile leaf
production in temperate ferns. It is also clear that observing a sporophyte only once can
lead to a number of erroneous assumptions about its past life history status. A plant with
no fertile leaves cannot automatically be assumed to be an immature plant. A plant with
one fertile leaf may or may not have been producing fertile leaves longer than one with
a 100% fertile crown. A plant that is fertile may or may not be fertile the following year.
A population that has the same proportion of fertile plants from one year to the next may
still reflect numerous changes due to independent changes in the fertility status of
individual plants. While the two species chosen for observation in this study may differ
in the overall number and timing of fertile leaves produced, there was a striking
similarity in the frequency of the different types of year-to-year transitions in plant
fertility status.
Fundamental to understanding of all of these observations is a knowledge of the
specific process that determines whether an emergent leaf will be fertile or sterile
(White 1971). The current exploratory study, limited as it was to simply observing the
reproductive sporophyte, has provided few clues about the stimuli behind the observed
patterns. The differences between species detected in this long-term field study of just
two species indicate that for each species the fertility triggers may be different.
Significant year-to-year variation in plant fertility suggests that environmental factors
may be important. Annual variation may be related to climate factors but the relatively
short time span of the study and the many potential microhabitat influences require
experiments to properly assess these observations.
More long-term field studies are needed as the differences between these two
species suggest that the patterns of leaf and plant fertility can vary widely. Because the
limitations on observing plants in the field were unknown at the start of this study,
sample sizes of plants for which data was available for all eight years were small by the
aan FERN GAZ. 17(4): 223-234. 2005
end of the study, especially for P acrostichoides. The number of plant fertility
transitions from fertile to sterile taking place after a plant had become reproductively
mature could only be observed a maximum of three times in an eight-year period,
limiting the potential for identifying underlying causes. Only one large cyclic event
occurred during the study, limiting the potential for generalizations about disturbance
effects.
The length of study was limited because of the choice of study site. It was conducted
on private land which changed hands after eight years and was no longer accessible for
research. For long-lived perennials, a long-term study should encompass a large part of
the life span of a sporophyte, and should ideally extend to 100 years or more. As
indicated by Werth & Cousens (1990), a long-term study should be conducted at a site
guaranteed to be in place for the long-term. The Long Term Ecological Research
(LTER) program (Van Cleve & Martin 1991) which has extended world-wide from its
original designation of sites in the United States can provide such stable study locations
for future long-term studies. The LTER sites also provide for maintenance of
standardized data sets long-term which allows for continual monitoring of marked
individuals beyond the lifetime of a single individual researcher. The availability of
LTER data on the internet also facilitates supplemental short-term studies and
experiments based on earlier observations, another recommendation made by Werth &
Cousens (1990). Long-term natural experiments such as this one rely on inference from
uncontrolled climate data. Although this approach does not have the rigor of a
controlled experiment, it does document variation under real environmental conditions.
Another advantage of conducting both long and short term studies at an LTER is the
continual monitoring of climate parameters as well as other abiotic and _ biotic
environmental characteristics which can be related to an individual researcher’s
observations about ferns.
This long-term study of fertility in two wintergreen fern species from the same
forest showed different patterns of annual variation, suggesting that any future
ecological assumptions about ferns must account for differences among fern species.
Studies of populations of both D. intermedia and P. acrostichoides throughout their
ranges are needed and other life history stages could exhibit such amplitudes of annual
variability as well. Long-term field monitoring and additional experimentation with
other fern species will be necessary before the ecological generalizations hoped for by
Harper (1982) can be made.
ACKNOWLEDGMENTS
I am grateful for the encouragement given to me by Mike Cousens many years ago. His
pioneering work in fern demographic studies inspired many of the questions I have
addressed. Thought-provoking discussions of fern ecology with Herb Wagner and
Charlie Werth further stimulated my enthusiasm for field studies of ferns. This paper is
dedicated to the memory of these three pteridologists whose insights based on field
studies were incomparable.
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FERN GAZ. 17(4): 235-242. 2005 230
NEW ZEALAND ASPLENIUM (ASPLENIACEAE: PTERIDOPHYTA)
REVISITED — DNA SEQUENCING AND AFLP FINGERPRINTING
L.R. PERRIE' & P.J. BROWNSEY
Museum of New Zealand Te Papa Tongarewa, P.O. Box 467, Wellington, New Zealand
(Email: 'leonp@tepapa.govt.nz)
Key words: New Zealand, Asplenium, A. hookerianum, A. bulbiferum, A. gracillimum,
A. richardii, chloroplast DNA sequence, AFLP DNA-fingerprinting, polyploid,
hybridisation.
This paper summarises our continuing study of the evolutionary history of
Asplenium in New Zealand. Chloroplast DNA sequence data and AFLP
DNA-fingerprinting have been used to examine the origins and relationships of
the New Zealand Asplenium taxa, to test species boundaries, and to further
investigate the ancestry of the octoploid Asp/enium taxa belonging to the Austral
group.
INTRODUCTION
In 1863, after three years in New Zealand, Samuel Butler’s “we have one very stupid
white gentian” (Butler, 1863) was assumedly a frustrated reflection of the difficulty of
recognising distinct entities within New Zealand Gentaniella Moench (there are
actually 30 species, and only half have white flowers; Glenny, 2003). Analogous
sentiments are apparent from those who first dealt taxonomically with Asp/enium L. in
New Zealand. Joseph Hooker (1855, p.33) wrote “The New Zealand kinds have defied
all attempts to be limited by words”. Similarly, “[some] are extremely inconstant and
pass into one another, thus rendering their classification a work of considerable
difficulty” (Thomson, 1882, p.72), and “The New Zealand species present exceptional
difficulties to the student, on account of their extreme variability and the manner in
which several of them are connected by intermediate forms” (Cheeseman, 1906, p.987).
Allan (1961, p.75) stated that the species of New Zealand Asplenium were “very ill-
defined”, and that while many appeared to respond “markedly to environmental
conditions...[,] there is also no doubt that hybridism plays an important part”.
Indeed, hybridisation has proved to be frequent amongst (some) New Zealand
Asplenium, and although these hybrids blur the morphological discontinuities between
taxa, they are sterile and present little opportunity for introgressive gene flow
(Brownsey, 1977a). The identification of hybrids was an integral part of delimiting taxa
in Brownsey’s (1977b) taxonomic revision of New Zealand Asplenium, in which 14
species and four subspecies were recognised. Since that treatment three species have
been newly described (Brownsey & Jackson 1984; Brownsey, 1985; Brownsey & de
Lange, 1997), some names have changed (Brownsey, 1979; Brownsey, 1999), and the
appropriate rank for some taxa has been debated (Ogle, 1985; Brownsey, 1998; Perrie
& Brownsey, 2005a), such that a present treatment (Table 1) might recognise 18 species
and four subspecies; comprising approximately 10% of the New Zealand fern flora
(Brownsey & Smith-Dodsworth, 2000).
This paper will summarise some of our recent discoveries in New Zealand
Asplenium, but we emphasise that our understanding of the evolution of New Zealand
236 FERN GAZ. 17(4): 235-242. 2005
TABLE 1: A present taxonomic treatment of Asplenium in New Zealand compared
with that of Brownsey (1977b).
Present Treatment As treated by Brownsey (1977b)
A. appendiculatum (Labill.) C.Presl A. terrestre Brownsey subsp. ferrestre
subsp. appendiculatum
subsp. maritimum (Brownsey) subsp. maritimum Brownsey
Brownsey
A. bulbiferum G.Forst. A. bulbiferum G.Forst. subsp.
bulbiferum
A. gracillimum Colenso subsp. gracillimum (Colenso)
Brownsey
A. chathamense Brownsey -
A. cimmeriorum Brownsey et de Lange
A. flabellifolium Cav. A, flabellifolium Cav.
A. flaccidum G.Forst. subsp. flaccidum A. flaccidum G.Forst. subsp. flaccidum
subsp. haurakiense Brownsey subsp. haurakiense Brownsey
A. hookerianum Colenso A. hookerianum Colenso
A. lamprophyllum Carse A. lamprophyllum Carse
A. lyallii (Hook.f.) T.Moore A, lyallii (Hook.f.) T.Moore
A. oblongifolium Colenso A. lucidum Forst.f.
A. obtusatum G.Forst. subsp. obtusatum | A. obtusatum G.Forst. subsp.
obtusatum
subsp. northlandicum Brownsey subsp. northlandicum Brownsey
A. pauperequitum Brownsey et P.J.Jacks. | -
A. polvodon G.Forst. A, polyodon G.Forst.
A, richardii (Hook.f.) Hook.f. A. richardii (Hook.f.) Hook.f.
A. scleroprium Hombr. A. scleroprium Hombr.
A. shuttleworthianum Kunze A. shuttleworthianum Kunze
A. trichomanes L. subsp. quadrivalens A. trichomanes L.
D.E.Mey emend Lovis
subsp. nov. -
PERRIE & BROWNSEY: NEW ZEALAND ASPLENIUM REVISITED za7
Asplenium is far from complete
RELATIONSHIPS AND BIOGEOGRAPHY
Brownsey (1977a) considered the Asp/enium species hybridising in New Zealand to
comprise a closely related group, dubbed the ‘Austral’ group, and this has been
supported by phylogenetic analyses of chloroplast DNA sequence data (Perrie &
Brownsey, 2005a). The species in New Zealand that do not hybridise there - A.
flabellifolium, A. trichomanes, A. pauperequitum, and A. polvodon - have closer
affinities elsewhere (the former three with a primarily temperate, northern hemisphere
group, and 4. polyodon with a group that includes 4. aethiopicum). Interestingly, the
Austral group is closely related to the bird’s nest ferns (e.g., 4. australasicum (J.Sm.)
Hook.) and the finely dissected A. theciferum (Kunth) Mett., which, as Loxoscaphe
thecifera (Kunth) T.Moore, is the type of Loxoscaphe T.Moore.
Amongst the New Zealand members of the Austral group, three chloroplast sub-
groups are clearly delimited: the Bulbiferum, Flaccidum, and Obtusatum chloroplast
groups (Perrie & Brownsey, 2005a). However, it remains unclear how these groups are
related to one another, as different DNA regions support conflicting relationships:
{{Bul,Obt}Fla} with 71% bootstrap support from the rbcL gene, versus
{{Bul,Fla}Obt} with 85% bootstrap support from the trnL-trnF intergenic spacer.
Molecular dating, using penalised likelihood of rbhcL DNA sequence data and a
calibration of 140 million years ago for the divergence of Asplenium and
Hymenasplenium Hayata, recovers dates of 45 million years ago or younger for the
divergence between each New Zealand Asp/enium species and its closest non-New
Zealand relative in the sample set analysed (Perrie & Brownsey, 2005a). The calibration
age is probably conservatively old (for instance, Schneider e¢ al. (2004) calculated the
Asplenium-Hymenasplenium split at just 55 million years ago), so it is likely that the
estimated divergence ages are (considerably) younger than actually calculated. In any
case, the estimated ages of the divergence are much more recent than the geological
separation of New Zealand from Gondwana (c.80 million years ago), implicating
multiple events of long-distance dispersal in the origins of New Zealand Asplenium
(Perrie & Brownsey, 2005a). The entire Austral group appears to have arisen after New
Zealand’s isolation. Therefore, the disjunct distributions of each of the seven Austral
Asplenium taxa that occur in New Zealand and elsewhere requires inference of at least
one dispersal event (unless the taxa are actually polyphyletic). Dispersal across the
Tasman Sea (c.2000 km) is potentially so common that it would be of considerable
interest to investigate the degree of genetic isolation between Australian and New
Zealand populations of shared fern species.
Figure 1. Plants from the ec Ranges of rece ry ianum n that exhibit the
broad-pinnuled and narrow-pinnuled morphology that has previously been referred to
as A. hookerianum and A. colensoi, respectively.
238 FERN GAZ. 17(4): 235-242. 2005
ASPLENIUM HOOKERIANUM
AFLP DNA-fingerprinting has been used in conjunction with chloroplast DNA
sequencing to investigate some groups in more detail. The morphologically distinct
Asplenium hookerianum and A. colensoi were described by Colenso (1845) in the same
publication; the ultimate segments are broad in the former, and narrow in the latter
(Figure 1). It is quite common to find sites where both broad and narrow-pinnuled
plants grow together, with few if any intermediates, thereby creating the impression of
two morphologically discrete groups growing in sympatry. However, genetic variation
as assayed by AFLP DNA-fingerprinting and chloroplast DNA sequencing is not
concordant with the morphological variation (Perrie & Brownsey, 2005b), and indicates
that only one species should be recognised (i.e., A. hookerianum).
ASPLENIUM BULBIFERUM
In part because of the apparent origins of Asplenium (bulbiferum subsp.) gracillimum
discussed below, we believe that the taxa previously recognised as A. bulbiferum subsp.
bulbiferum and A. bulbiferum subsp. gracillimum (Brownsey, 1977b) should be treated
at the specific level as A. bulbiferum s.s. (hereafter simply A. bulbiferum) and A.
gracillimum (Perrie & Brownsey, 2005a). Circumscribed as such, the name A.
A. hookerianum
>
>
rs)
3
>
oO
=,
pst)
3
J
3
. - ae Qe
A. richardii >
A. gracillimum $
2
Jee ane eR CER PPS eat =
3
©
3
ba A. richardii
A. bulbiferum
A B
a A. bulbiferum
Figure 2. A: Maximum parsimony analysis of 469 AFLP characters for the tetraploids
Asplenium bulbiferum and A. hookerianum, together with the octoploid A. richardii, a
putative autopolyploid of A. hookerianum. Representatives from one of the A.
gracillimum lineages are included for comparison. One of 55 most parsimonious trees,
which differ in the internal arrangement of samples within 4. bu/biferum and A.
hookerianum. Thickened branches indicate those with >80% bootstrap support. B: One
of three most parsimonious trees, differing in the arrangement of the A. hookerianum
and A. gracillimum samples, from chloroplast trnL-trnF region DNA sequence data.
ae: using samples from the Flaccidum and Obtusatum chloroplast groups (not
shown).
PERRIE & BROWNSEY: NEW ZEALAND ASPLENIUM REVISITED 239
cae ila is then restricted to tetraploid plants, and is endemic to New Zealand.
urther, it is quite distinct from the plant common in cultivation that is frequently
hare “A. bulbiferum” (Perrie et al., 2005). Asplenium bulbiferum appears to be less
genetically variable than its closest known tetraploid relative, A. hookerianum. Only
one trnL-trnF region haplotype is presently known from A. bulbiferum, as opposed to
eight in A. hookerianum. Also, only 38% of AFLP loci were polymorphic within A.
bulbiferum, compared to 52% within A. hookerianum, despite more populations being
sampled from the former.
ASPLENIUM GRACILLIMUM
The octoploid Asplenium gracillimum (previously A. bulbiferum subsp. gracillimum)
appears to be an allopolyploid between the tetraploids A. bulbiferum and A.
hookerianum. Asplenium gracillimum produces bulbils like A. bu/biferum (albeit with
much less frequency), but has the chloroplast DNA sequence of A. hookerianum (Perrie
& Brownsey, 2005a). Further, A. gracillimum is intermediate between its two putative
app.app,
obt.nor lya scl app.mar shu
=
ee
ae a
. . Pg
. . ao”
‘ ae
;
r we .
{ |
bul obt.obt obl flc.flc, flc.hau, lam
cha
Figure 3. Hypothesised origins for the New Zealand species of the Asp/enium Austral
group. Octoploid taxa subtended by a single arrow are thought to be autopolyploids.
Probable allopolyploids are subtended by two arrows. Solid arrows re flect linkages
indicated by chloroplast DNA sequences; dashed arrows are linkages inferred from
morphology. Arrows not pointing directly to a tetraploid taxon reflect uncertainty. Taxa
are represented by the first three letters of their name in Table 1, except “fle” =
flaccidum. Fronds are not to scale.
240 FERN GAZ. 17(4): 235-242. 2005
parents with regards to frond architecture, scale shape, and habitat.
Reminiscent of the results of Trewick ef al. (2002) with Asplenium ceterach L.,
chloroplast DNA sequence haplotypes shared by Asplenium hookerianum and A.
gracillimum indicate that the latter has had multiple polyploid origins. Although this
work is still preliminary, the different chloroplast DNA sequence haplotypes found in
A. gracillimum appear to correspond to distinct groups detected with AFLP (Perrie &
Brownsey, in prep.). Some of these apparently independently derived lineages of A.
gracillimum appear to be sympatric, but it is unknown what happens reproductively
where they meet.
ASPLENIUM CIMMERIORUM
The recently described Asplenium cimmeriorum has a conservation status of ‘sparse’,
being known only from limestone regions in the north-west of the South Island and
around Waitomo in the North Island (Brownsey & de Lange, 1997). Chloroplast DNA
sequences indicate that the north and south populations of 4. cimmeriorum have
independent derivations; one with an 4. hookerianum-like chloroplast and the other
with an A. bu/biferum-like chloroplast (Perrie, de Lange, & Brownsey, in prep.). AFLP
also recovers them as distinct groups (with both also distinct from 4. gracillimum). This
suggests a need to provide independent conservation assessments, with the northern
Waitomo plants being much less common than those in the south.
Both groups of Asplenium cimmeriorum appear to be allopolyploids between A.
bulbiferum and A. hookerianum (like A. gracillimum). This situation presents the
problem of how to deal taxonomically with multiple lineages of independent but
effectively equivalent origins, some of which are ecologically and/or morphologically
recognisable (e.g., A. cimmeriorum from A. gracillimum) while others are not (e.g., the
cryptic lineages within A. gracillimum).
ASPLENIUM RICHARDII
The octoploid Asplenium richardii is possibly an (old) autopolyploid of a narrow-
pinnuled plant of 4. hookerianum. In maximum parsimony analysis of AFLP data, A.
richardii falls just outside the diversity found in extant 4. hookerianum, which contrasts
with the putatively allopolyploid lineages of A. gracillimum that lie intermediate
between 4. bulbiferum and A. hookerianum (Figure 2; see Perrie et a/. (2003a) for
discussion of AFLP data and type of polyploidy). The chloroplast DNA sequences of A.
richardii are allied to, but nevertheless distinct from, those found in extant 4.
hookerianum (Figure 2). This suggests that if A. richardii is an autopolyploid from a
narrow-pinnuled, A. hookerianum-like plant, then it is perhaps not of recent origin. That
is, sufficient subsequent time has passed since its origin that the A. richardii chloroplast
DNA sequences have (a) been lost from the extant populations of A. hookerianum
and/or (b) acquired apomorphic changes.
AUSTRAL GROUP OCTOPLOID ORIGINS
Chloroplast DNA sequences have revealed the origins of the octoploid, New Zealand
taxa from the Austral group with varying degrees of precision (Perrie & Brownsey,
2004, 2005a), and our present knowledge is summarised in Figure 3. For instance,
Asplenium hookerianum and A. gracillimum actually share trnL-trnF region haplotypes
(Perrie & Brownsey, 2005a; Perrie & Brownsey, in prep.), while more than 2000 base-
pairs from four chloroplast loci are unable to differentiate between A. oblongifolium and
PERRIE & BROWNSEY: NEW ZEALAND ASPLENIUM REVISITED _ 241
A. obtusatum subsp. obtusatum as the chloroplast parents of A. /vallii and A.
scleroprium (Perrie & Brownsey, 2004). Some of the octoploids are thought to have
autopolyploid origins: A. appendiculatum (from an A. flaccidum-like or A.
chathamense-like progenitor), A. obtusatum subsp. northlandicum (possibly from A.
obtusatum subsp. obtusatum), and A. richardii (see above). Others are probably
allopolyploids: A. cimmeriorum, A. gracillimum (see above), A. lvallii (between
something in the Obtusatum chloroplast group and possibly 4. hookerianum), and A.
scleroprium (between something in the Obtusatum chloroplast group and something
like A. flaccidum). Asplenium shuttleworthianum has chloroplast DNA sequences that
nest within the Flaccidum chloroplast group, but no close tetraploid is known.
CONCLUSION
The extant Asplenium of New Zealand are derived from disparate groups within the
genus. Long-distance dispersal has seemingly played a major role in shaping the
distributions of Asplenium in the south-west Pacific, as is the case for much of the flora
of New Zealand (Winkworth et al. 2002: Perrie et al. 2003b). Genetic data has allowed
the further unravelling of the histories of some of the Austral octoploids and a resolution
of some taxonomic problems (e.g., 4. hookerianum), but it also has unveiled new
avenues for investigation (e.g., the multiple, cryptic groups within 4. gracillimum).
Many issues within New Zealand Asp/enium remain to be tackled, such as clarifying the
boundaries between some taxa (e.g., Perrie & Brownsey, 2004) and establishing the
most appropriate ranks for others (Ogle, 1987).
ACKNOWLEDGEMENTS
We are grateful to Peter de Lange and Lara Shepherd who have collaborated with us on
some aspects of this work, and to the organisers for the opportunity to contribute to the
Ferns for the 21St Century conference and its proceedings. This work was funded by the
Foundation for Research, Science and Technology (contract MNZX0201).
REFERENCES
ALLAN, H.H. 1961. Flora of New Zealand, Vol. 1. Government Printer, Wellington.
BROWNSEY, P.J. 1977a. Asplenium hybrids in the New Zealand flora. New Zealand J.
Bot. 15: 601-637.
BROWNSEY, P.J. 1977b. A taxonomic revision of the New Zealand species of
Asplenium. New Zealand J. Bot. 15: 39-86.
BROWNSEY, P.J. 1979. Asplenium lucidum Forst.f., an illegitimate name for the New
Zealand shining spleenwort. New Zealand J. Bot. 17: 217-218.
BROWNSEY, P.J. 1985. Asplenium chathamense - a new fern species from the
Chatham Islands, New Zealand. New Zealand J. Bot. 23: 135-140
BROWNSEY, P.J. 1998. Aspleniaceae. Fl. Australia 48: 295-327.
BROWNSEY, P.J. 1999. A new combination in Asplenium. New Zealand J. Bot. 37:
BROWNSEY, P.J., JACKSON, P.J. 1984. Asplenium pauperequitum — a new fern
species from the Poor Knights Islands, New Zealand. New Zealand J. Bot. 22:
315-321.
BROWNSEY, P.J., DE LANGE, P.J. 1997. ig ou cimmeriorum, a new fern species
from New Zealand. New Zealand J. Bot. 35: 283-292.
BROWNSEY, P.J., SMITH-DODSWORTH, 1c. 2000. New Zealand ferns and allied
242 FERN GAZ. 17(4): 235-242. 2005
plants. 2nd eq. David Bateman Ltd., Auckland.
BUTLER, S. 1863. A first year in Canterbury Settlement. Butler, London.
CHEESEMAN, T.F. 1906. Man. New Zealand Fl. Government Printer, Wellington.
COLENSO, W. 1845. A classification and description of some newly discovered ferns,
collected in the Northern Island of New Zealand, in the summer of 1841-42.
Tasmanian J. Nat. Sci. 2: 161-189.
GLENNY, D. 2003. Understanding the New Zealand gentians. Te Taiao 2: 4-5.
HOOKER, J.D. 1855. Flora Novae Zelandiae, Vol. Il. Reeve, London
OGLE, C.C. 1987. Taxonomic changes in Asplenium (Aspleniaceae; Filicales) in New
Zealand. New Zealand J. Bot. 25: 591-593.
PERRIE, L.R., BROWNSEY, P. J., LOCKHART, P.J., LARGE, M.F. 2003a. Evidence
for an allopolyploid complex in New Zealand Polystichum. New Zealand J. Bot. 41:
189-215.
PERRIE, L.R., BROWNSEY, P. J., LOCKHART, P.J., BROWN, E.A., LARGE, M.F.
2003b. Biogeography of temperate Australasian Polystichum ferns as inferred from
chloroplast sequence and AFLP. J. Biogeogr. 30: 1729-1736.
PERRIE, L.R., BROWNSEY, PJ. 2004. #nL intron variation in New Zealand taxa of
the Asplenium obtusatum Chloroplast Group. Tuhinga - Records of the Museum of
New Zealand Te Papa Tongarewa 15: 1-5.
PERRIE, L.R., BROWNSEY, P.J. 2005a. Insights into the biogeography and polyploid
evolution of New Zealand Asplenium from chloroplast DNA sequence data. Amer.
Peo J, 95: 1-21,
PERRIE, L.R., BROWNSEY, P.J. 2005b. Genetic variation is not concordant with
merevalonien! variation in the fern Asplenium hookerianum sensu lato
(Aspleniaceae). Amer. J. Bot. 92: 1559-1564.
PERRIE, L.R., SHEPHERD, L.D., BROWNSEY, P.J. 2005. Asplenium x lucrosum
nothosp. nov.: a sterile hybrid widely and erroneously cultivated as "Asplenium
Le ely a Syst. Evol. 250: 243-257.
SCHNEIDER, H., SCHUETTPELZ, E., PRYER, K.M., CRANFILL, R.,
verter S., LUPIA, R. 2004. Ferns divenittad in the shadow of angiosperms.
Nature 428: 553-557.
THOMSON, GM. 1882. The ferns and fern allies of New Zealand. Wise, Dunedin.
TREWICK , S.A.,. MORGAN-RICHARDS, M., RUSSELL, S.J., HENDERSON, S.,
RUMSEY, F.J., PINTER, I., BARRETT, J.A., GIBBY, M., VOGEL, J.C. 2002.
Polyploidy, phylogeography and Pleistocene refugia of the rockfern Asplenium
ceterach: evidence from chloroplast DNA. Molec. Ecol. 11: 2003-2012.
WINKWORTH, R.C., WAGSTAFF, S.J., GLENNY, D., LOCKHART, P.J. 2002. Plant
dispersal N.E.W.S from New Zealand. Trends Ecol. Evol. 17: 514-520.
243 FERN GAZ. 17(4). 2005
MOLECULAR EVIDENCE FOR MULTIPLE ORIGINS OF THE
APOMICTIC TRIPLOID FERN CORNOPTERIS CHRISTENSENIANA
(WOODSIACEAE)
C.-H. PARK! & M. KATO”
‘Laboratory of Ecology Science, Department of Biology, Faculty of Science,
Kyushu University, 6-10-1, Hakozaki, Higashi-ku, Fukuoka 812-8581, Japan
Corresponding author; (Email: cparkscbh@mbox.nc.kyushu-u.ac.jp)
“Department of Biological Sciences, Graduate School of Science,
University of Tokyo, 7-3-1 Hongo, Tokyo 113-0033, Japan
Key words: apogamy, apomixis, Cornopteris christenseniana, hybrid fern, multiple
origins, triploid
Apomixis (apogamy in a broad sense, agamospermy, agamospory) is a reproductive
pathway via chromosomally unreduced spores and gametophytes and the subsequent
apogamous reproduction (in a strict sense), i.e. asexual production of a sporophyte from
vegetative cells of a gametophyte. Here we use the term apomixis in a broad sense and
apogamy in a strict sense to avoid confusion. In pteridophytes, about 10% of the world’s
species and about 15% of Japanese species are apomictic. Apomixis is often associated
with polyploidy, and three quarters of apomictic pteridophytes are triploid. It is often
argued that apomixis, like polyploidy, plays a significant role as an escape from sterility
caused by hybridization.
Although Cornopteris christenseniana had been believed to be a sterile triploid
interspecific hybrid, our previous study showed that it produces viable spores at various
frequencies and forms apogamous sporophytes at low or moderate frequencies in
culture (Park & Kato, 2003). It also suggested that apomictic reproduction (in a broad
sense) occurs naturally in an artificial environment in the Fern Garden of the Botanical
Gardens, University of Tokyo. We have analyzed molecular variation of the nuclear and
cpDNA in C. christenseniana and its closely related species. The results strongly
support the suggestion that many plants of C. christenseniana propagate by apomixis in
the artificial environment and not by in situ hybridization. We also found that plants of
C. christenseniana that had been collected from various wild populations, had arisen
independently on a markedly recurrent basis. We suggest that Cornopteris
christenseniana is an incipient apomictic species of multiple and polytopic origin, and
its apomixis is not so strongly regulated as in obligate apomicts. The multiple origins
of incipient apomicts imply that the evolution of apomixis in C. christenseniana began
with unreduced sporogenesis with a pleiotropic effect leading to subsequent apogamy.
REFERENCES
PARK, C.-H. & KATO, M. 2003 Apomixis in the interspecific triploid hybrid fern
Cornopteris christenseniana (Woodsiaceae). J. Plant Res. 116(2): 93-104.
244 FERN GAZ. 17(3). 2005
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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-
122.
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 Hymenophy llum, 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.
aS each year) <e Full Personal
THE BRITISH PTERIDOLOGICAL SOCII
Registered Charity No. 1092399
Patron: HRH The Prince of Wales
Officers aid Committee from Marth 1005
President: Dr A.F. Dyer, 499 Lanark Road West, Balerno, Edinburgh EH14 TAL.
E-mail: President@eBPS. org.uk
Vice-Presidents : M.H. Rickard, Prof. B.A. Thomas
Honorary General Secretary: Miss J.M. i: 42 Crown Woods tes Eltham, London SE9 2NN
ail: Secretary@eBPS.org. uk
Treasurer: A. Leonard, |! Victory Roa, taka Hants.PO13DR
il: mail@andrew-leonard.co.uk
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il: Meetings@eBPS.org.uk _
Conservation Officer/Recorder: Dr H. Ss: seseimig 180 ee Ra Edinburgh, EHS [AH ©
= . Rumsey, Department of Botany, The Natural History
Museum, Cromwell Road, Condon. $9 SW7 SBD; E-mail: Conservation@eBPS. org.uk
Editor of the Bulletin: | Miss A.M. Paul, Department of Botany, The Natural History M fuseum,
“Cromvell pak London SW7 SBD; E-mail: Bulletin@eBPS. ae uk
Editor of the Fern Gazett Prof. M. Gibby, Royal Botanic Garden ae
20A Tnverteith Row, Edinburgh EH3 SLR; E-mail: FernGazette@eBPS. org. ak %
Editor of the Pteridologist: Dr J.W. Merryweather, "The Whins', Auchtertyre,
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The BRITISH PTERIDOLOGICAL SOCIETY was founded i in ike and today continues asa s focus a fae
fern enthusiasts. It provides a wide range of informatio blications cae
me once literature. It also organises formal talks, informal discussions, field ‘meetings, oe visits. :
The S ; has a wide membership ' which sikaded ee
ae ers, nurserymen and | botanists, both amateur and professi eelteeal The Society's journals, the Fern Gazette,
Preridologist and ee are published annually. The Fern Gazette pee matter chiefly of specialist
interest on inter dology, the Preridologist, topics of. wet serial catty |, and the e Bulletin, Society
_ ses and ets eps www Sos restuc’y wEBrSo org.uk :
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THE FERN GAZETTE :
Rel oat aes
2005
eta ts 5
; and Population structure in
Racer pers Periph)
205-215
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