Table Of ContentPolish Botanical Journal 57(1): 129–135, 2012
THE FRIENDLY ISLANDS – A CHECKLIST OF HORNWORTS
AND LIVERWORTS OF TONGA
LARS SÖDERSTRÖM, ANDERS HAGBORG & MATT VON KONRAT
Abstract. The fi rst checklist of hornworts and liverworts is provided for Tonga. Geographic coverage includes the islands
Ha’apai, Niua, Tongatapu, Vava’u, and surrounding smaller islands. It is apparent that there remain many localities in Tonga,
including entire islands, which have never or scarcely been collected for this group of plants. We report three hornwort and 47
liverwort taxa. Two additional liverwort taxa previously reported from Tonga are noted as doubtful records. The list is based
on over 35 literature references from over 5000 papers that the ELPT project has processed so far, including monographs and
regional studies. New records will undoubtedly be reported, based on regionally widespread species that have been recorded
elsewhere, especially nearby Fiji and Samoa. It is clear Tonga has received negligible attention, and almost no publications
exist that have focused solely on this group of islands. The current checklist is part of a broader study producing checklists for
islands across the South Pacifi c, which we hope will foster and encourage research of liverworts and hornworts in this globally
important biodiversity hotspot.
Keywords: Anthocerotophyta, Australasia, biodiversity, Marchantiophyta, Pacifi c region, Tonga
Lars Söderström, Department of Biology, Norwegian University of Science and Technology, N-7491 Trondheim, Norway; e-mail:
[email protected]
Anders Hagborg & Matt von Konrat, Botany Department, The Field Museum, Chicago, Illinois 60605-2496, U.S.A.; e-mail:
[email protected], mkonrat@fi eldmuseum.org
INTRODUCTION
Mittermeier et al. (2005) identifi ed all the islands fl orae hepaticarum polynesiae (Miller et al. 1983)
of Micronesia and Polynesia, including the is- provided a comprehensive checklist of liverworts
lands of Tonga, as one of 35 global biodiversity and hornworts from the tropical islands of the
‘hotspots’, often referred to as the Polynesia-Micro- Pacifi c, including those of Tonga. This work was
nesia hotspot. Conservation International (www. based on a compilation of literature and served
conservation.org, accessed 16/08/2011) considers as a fi rst step towards making knowledge about
the Polynesia-Micronesia hotspot the epicentre of Pacifi c island bryophytes more accessible and has
the current global extinction crisis. The kingdom remained a signifi cant work. Yet, the publication
of Tonga (sometimes referred to as the Friendly was largely an uncritical compilation of names
Islands because of the friendly reception of Captain from literature (Söderström et al. 2011).
Cook) comprises an archipelago of 176 islands We here present the fi rst checklist of liverworts
of varying sizes (most of them uninhabited), in and hornworts for the islands of Tonga. The current
the tropical southwest Pacifi c with a land area of paper is part of a larger series of papers focusing
714 km2. The highest elevation (on the island of on increasing our fl oristic knowledge in the South
Kao in the Ha’apai group) is 1033 m. Pacifi c including nearby Fiji (Söderström et al.
No publications have focused solely on liv- 2011; Pócs et al. 2011; von Konrat et al. 2011)
erworts from Tonga. The most important contri- and New Caledonia (Thouvenot et al. 2011). Al-
bution to the knowledge of Tongan liverworts is most three decades after Miller et al. (1983) we
the series of papers from South Pacifi c by Hans hope the series of papers will work towards the
Hürlimann (Hürlimann 1960–1998). Prodromus same goal as spirited by them, help promote further
130 POLISH BOTANICAL JOURNAL 57(1). 2012
bryological research in the region, and provide to the International Plant Names Index (http://www.ipni.
a foundation for revisionary studies and fl oristic org/index.html). Taxa doubtfully present in Tonga and
research. Undoubtedly, many new records will be synonyms are listed separately. The list of synonyms
only contains names used for Tonga.
reported for the Tongan islands, given that at least
The following abbreviations are used for the island
190 other liverwort and hornwort species occur on
groups: HA = the Ha’apai group (including Tofua, Kao,
the nearby islands of Samoa and at least another
Nomuka and the Lifuka group); NI = the Niua group
230 species in Fiji. In this checklist we report 50
(including Tafahi, Niuafo’ou and Niuatoputapu); TT =
taxa as occurring on the Tongan islands. Two ad- the Tongatapu group (including ‘Eua, Nuku’alofa and
ditional taxa are reported as doubtful records. Ata); VA = the Vava’u group (including Fonualei, Tokua
It is clear that Tongan bryophytes have been and Late); and TON = lack of precise locality informa-
overlooked historically, compared to seed plants, tion in the cited references.
birds, and other organisms forming a remarkable
gap in the knowledge of the fl ora of Tonga. More- LIST OF HORNWORTS AND LIVERWORTS
over, the state of our fl oristic knowledge of these RECORDED FOR TONGA
organisms is defi cient compared to many other
biodiversity hotspots and other ecologically signifi - ANTHOCEROTOPHYTA
cant areas of the globe. The island of ‘Eua should
Dendroceros
be especially interesting because of its fl oristic af-
*** D. javanicus (Nees) Nees
fi nities, as it is the oldest island of the archipelago
and is thought to be partly an ancient fragment of HA: Miller et al. (1983); NI: HÜRLIMANN (1960);
VA: Miller et al. (1983).
Gondwana (Mueller-Dombois & Fosberg 1998).
As an example, it has the endemic Podocarpus
Folioceros
foliolatus de Laub. and podocarps are good sub-
*** F. fuciformis (Mont.) D. C. Bharadwaj
strates for interesting bryophytes.
HA: Miller et al. (1983) as Anthoceros fuciformis;
TT: HÜRLIMANN (1960) as Anthoceros vesiculosus.
METHODS AND PRESENTATION
Megaceros
The present checklist is based on a compilation of all *** M. fl agellaris (Mitt.) Steph.
the published records of species and infraspecifi c taxa
TT: HÜRLIMANN (1960) as Megaceros grandis,
of liverworts and hornworts from Tonga. The ELPT da-
HÜRLIMANN (1960) as Megaceros vescoanus, Miller
tabase (von Konrat et al. 2010) was an important source.
et al. (1983) as Megaceros grandis and Megaceros
The format of the present checklist follows that of the
vescoanus.
database. Accordingly, confi dence levels are provided
for each taxon, as follows (Söderström et al. 2010; von
Konrat et al. 2010): * = serious doubts about the value MARCHANTIOPHYTA
of the taxon, ** = probably a good taxon, but not studied Acrolejeunea
critically in the framework of a recent revision or mono-
*** A. aulacophora (Mont.) Steph.
graph, *** = good taxon as currently understood, studied
in the framework of a recent revision or monograph. TT: GRADSTEIN (1975), Miller et al. (1983), HÜR-
As in the recent checklists in the same series a dis- LIMANN (1989).
tinction in the species lists is made between primary
references (providing details on specimens and/or exact Archilejeunea
localities) and secondary ones (lacking details). Primary *** A. planiuscula (Mitt.) Steph.
references are in small capitals and secondary ones in
TT: HÜRLIMANN (1991), VA: HÜRLIMANN (1991).
normal font.
No attempt has been made to examine the specimens
Bazzania
of the published records and verify the identifi cations.
** B. paradoxa (Sande Lac.) Steph.
Nomenclature follows the latest taxonomic opinions as
much as possible and author abbreviations are according NI: HÜRLIMANN (1985).
L. SÖDERSTRÖM ET AL: A CHECKLIST OF HORNWORTS AND LIVERWORTS OF TONGA 131
*** B. tridens (Reinw., Blume & Nees) Trevis. that more than one taxon may be involved. There
NI: HÜRLIMANN (1985) as Bazzania australis. is at least one distinct segregate in Central America
(Forrest et al. 2011) but we do not know where
Cheilolejeunea Pacifi c reports belongs.
** C. ceylanica (Gottsche) R. M. Schust.
Harpalejeunea
& Kachroo
*** H. fi licuspis (Steph.) Mizut.
TT: HÜRLIMANN (1995); VA: HÜRLIMANN (1995).
TT: HÜRLIMANN (1995); VA: HÜRLIMANN (1995).
*** C. intertexta (Lindenb.) Steph.
Heteroscyphus
VA: HÜRLIMANN (1995).
*** H. argutus (Reinw., Blume & Nees) Schiffn.
Cololejeunea
VA: HÜRLIMANN (1998).
** C. ceatocarpa (Ångstr.) Steph.
** H. jackii (Steph.) Schiffn.
TON: Pócs et al. (2011); NI: HÜRLIMANN (1987).
NI: HÜRLIMANN (1998); TT: HÜRLIMANN (1998).
*** C. cocoscola Tixier
Jackiella
TON: Eggers & Pócs (2010); VA: PÓCS ET AL.
(2011) (TYPE). *** J. javanica Schiffn.
** C. hirta Steph. TON: Piippo (1985), Váňa (1992); TT: HÜRLIMANN
(1974), Miller et al. (1983), HÜRLIMANN (1989).
TON: Pócs et al. (2011); NI: HÜRLIMANN (1987).
Lejeunea
*** C. kapingaensis H. A. Mill.
*** L. anisophylla Mont.
VA: HÜRLIMANN (1987).
TON: Zhu & So (2001), Lai et al. (2007), Pócs
* C. maritima Tixier (2010); TT: HÜRLIMANN (1993); VA: HÜRLIMANN
(1993).
TT: HÜRLIMANN (1987).
** L. caviloba (Steph.) Steph. ex Besch.
** C. pacifi ca Pócs
VA: HÜRLIMANN (1993).
TT: PÓCS (2012).
** L. connatistipula (Steph.) Steph.
*** C. raduliloba Steph.
TT: HÜRLIMANN (1993).
TT: MÜLLER & SCHÄFER-VERWIMP (1999) as
Cololejeunea pakseana.
*** L. fl ava (Sw.) Nees
** C. vidaliana Tixier TT: FULFORD (1959), Miller et al. (1983).
NI: HÜRLIMANN (1987). ** L. fl ava subsp. orientalis R. M. Schust.
*** C. wightii Steph. TT: HÜRLIMANN (1993).
TON: Pócs et al. (2011); NI: HÜRLIMANN (1987); ** L. micholitzii Mizut.
TT: HÜRLIMANN (1987), ZHU & SO (1999).
VA: HÜRLIMANN (1993) as Lejeunea parvisac-
Dumortiera cata.
*** D. hirsuta (Sw.) Nees
Leptolejeunea
TT: Fulford (1959), HÜRLIMANN (1960), Miller et *** L. elliptica (Lehm. & Lindenb.) Schiffn.
al. (1983).
NI: HÜRLIMANN (1987) as Leptolejeunea subacuta;
NOTES. This is a variable species and recent TT: HÜRLIMANN (1995) as Leptolejeunea subacuta; VA:
studies (N. Salazar-Allen, pers. comm.) indicate HÜRLIMANN (1995) as Leptolejeunea subacuta.
132 POLISH BOTANICAL JOURNAL 57(1). 2012
*** L. epiphylla (Mitt.) Steph. ** R. javanica Gottsche
TON: HÜRLIMANN (1995). TT: HÜRLIMANN (1978), Miller et al. (1983).
Lopholejeunea * R. multifl ora Gottsche ex Schiffn.
*** L. subfusca (Nees) Schiffn. NI: HÜRLIMANN (1978); VA: Miller et al. (1983).
TT: Hürlimann (1991), Zhu & Gradstein (2005); *** R. retrofl exa Taylor
VA: Hürlimann (1991).
NI: HÜRLIMANN (1978) as Radula miqueliana;
VA: Miller et al. (1983) as Radula retrofl exa var. ret-
Mastigolejeunea
rofl exa.
*** M. auriculata (Wilson) Schiffn.
Riccardia
VA: HÜRLIMANN (1989).
*** R. alba (Colenso) E. A. Br.
*** M. calcarata (Mitt. ex Steph.) Verd.
NI: HÜRLIMANN (1976) as Riccardia samoana;
TT: HÜRLIMANN (1989). VA: Miller et al. (1983) as Riccardia samoana, Brown
& Braggins (1989).
*** M. virens (Ångstr.) Steph.
*** R. graeffei (Steph.) Hewson
TT: HÜRLIMANN (1989); VA: HÜRLIMANN (1989).
TON: Miller et al. (1983), Furuki (2002, 2006);
Metalejeunea NI: HÜRLIMANN (1976); TT: HÜRLIMANN (1976); VA:
Miller et al. (1983).
*** M. cucullata (Reinw., Blume & Nees) Grolle
VA: HÜRLIMANN (1993) as Lejeunea cucullata. ** R. tenuicostata Schiffn.
NI: HÜRLIMANN (1976); VA: Miller et al. (1983).
Phaeolejeunea
*** P. amicorum (Hürl.) Pócs Schiffneriolejeunea
TT: HÜRLIMANN (1991) as Phaeolejeunea etesseana *** S. tumida (Nees) Gradst.
subsp. amicorum Hürl. (TYPE), PÓCS (2008), PÓCS et al. TT: HÜRLIMANN (1989).
(2011).
Solenostoma
Plagiochila
*** S. haskarlianum (Nees) R. M. Schust. ex Váňa
** P. chauviniana Mont. & D. G. Long
NI: HÜRLIMANN (1974) as Plagiochila novae- TON: Váňa & Inoue (1983) as Jungermannia
guineae; VA: Miller et al. (1983) as Plagiochila novae- haskarliana, Váňa & Piippo (1989) as Jungermannia
guineae. haskarliana, Váňa (1991) as Jungermannia haskar-
liana, Schuster (2002) as Solenostoma haskarlianum;
*** P. javanica (Sw.) Nees & Mont. TT: HÜRLIMANN (1974, 1989) both as Jungermannia
NI: HÜRLIMANN (1974) as Plagiochila powellii; VA: haskarliana, VÁŇA (1975) as Jungermannia haskarliana,
Miller et al. (1983) as Plagiochila powellii. Miller et al. (1983) as Jungermannia haskarliana.
** P. vitiensis Mitt. Thysananthus
TON: Miller et al. (1983), Inoue (1986); NI: HÜRLI- *** T. spathulistipus (Reinw., Blume & Nees)
MANN (1974), TT: SO (2001); VA: INOUE (1970, 1981), Lindenb
Scott & Bradshaw (1985). VA: HÜRLIMANN (1989).
Radula
DOUBTFUL RECORD
** R. acuta Mitt.
Radula
TT: HÜRLIMANN (1978) as Radula paucidens, Miller
et al. (1983) as Radula paucidens. *** R. quadrata Gottsche
L. SÖDERSTRÖM ET AL: A CHECKLIST OF HORNWORTS AND LIVERWORTS OF TONGA 133
TT: FULFORD (1959) as Radula andicola, Miller Phaeolejeunea etesseana subsp. amicorum Hürl. ≡
et al. (1983) as Radula mollis and R. andicola. Phaeolejeunea amicorum
Plagiochila novae-guineae Sande Lac. = Plagiochila
NOTE. This is an African and American taxon
chauviniana
reported only once from the Pacifi c and never re-
Plagiochila powellii Mitt. ex Steph. = Plagiochila
ported from SE Asia. javanica
Radula andicola Steph. = Radula quadrata
REJECTED RECORDS Radula miqueliana Taylor = Radula retrofl exa
Radula mollis Lindenb. & Gottsche = Radula quad-
Cololejeunea rata
Radula paucidens Steph. ex Castle = Radula
*** C. cardiocarpa (Mont.) A. Evans
acuta
TON: Pócs et al. (2011); TT: HÜRLIMANN (1987). Riccardia samoana (Steph.) H. A. Mill. = Ricca-
NOTE. The specimen that this report is based rdia alba
Solenostoma haskarlianum (Nees) R. M. Schust.
on belongs to C. pacifi ca (Pócs 2012).
nom. inval. ≡ Solenostoma haskarlianum
Lejeunea
ACKNOWLEDGEMENTS. We would like to thank John
*** L. capensis Gottsche Engel for access to his personal library and Tamás Pócs
TT: Hürlimann (1993) as Lejeunea cf. caespitose; for critical comments on the manuscript. We dedicate
VA: Hürlimann (1993) as Lejeunea cf. caespitosa. this paper to Marian Kuc for his contribution to bry-
ology, and we hope that this paper encourages research
NOTE. Only reported with some uncertainty. of liverworts and hornworts across the world. This
Lejeunea caespitosa is technically a synonym of work was supported, in part, by GBIF (Seed Money:
L. capensis (Reiner-Drehwald 2009). However, 2007-41), and is part of the Early Land Plants Today
specimens so named from the Pacifi c probably all project.
belong to L. anisophylla.
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Received 18 January 2012
