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Corresponding author: Ainun Nadhifah ( ainun.nadhifah@lipi.go.id ) Academic editor: Matt Renner
© 2021 Ainun Nadhifah, Lars Söderström, Anders Hagborg, Eka Aditya Putri Iskandar, Ida Haerida, Matt von Konrat.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Nadhifah A, Söderström L, Hagborg A, Iskandar EAP, Haerida I, von Konrat M (2021) An archipelago within an archipelago: A checklist of liverworts and hornworts of Kepulauan Sunda Kecil (Lesser Sunda Islands), Indonesia and Timor-Leste (East Timor). PhytoKeys 180: 1-30. https://doi.org/10.3897/phytokeys.180.65836
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The first ever liverwort and hornwort checklist is provided for the Kepulauan Sunda Kecil (Lesser Sunda Islands) of Indonesia and Timor-Leste (East Timor). We report 129 accepted taxa, 12 doubtful taxa and three rejected taxa previously reported for the Lesser Sunda Islands. The list is based on over 130 literature references, including monographs, regional studies, and molecular investigations. It is clear that bryophytes from this region have been overlooked historically, and under collected, compared to seed plants, birds, and other organisms, forming a remarkable gap in the flora of Indonesia. Publications dealing with liverworts of Lesser Sunda Islands are few and scattered. We predict that further fieldwork, in addition to collections unveiled from regional herbaria, will uncover a number of new records that remain to be reported, especially considering that regionally widespread species have been recorded elsewhere.
Checklist, hornworts, Indonesia, Kepulauan Nusa Tenggara, Kepulauan Sunda Kecil, Lesser Sunda Islands, liverworts, Timor-Leste (East Timor)
The Lesser Sunda Islands (LSI), known in Indonesia as Kepulauan Sunda Kecil, are an area covering a longitudinal distance of some 600 kilometres in the southeastern portion of Indonesia, extending between Java in the west and New Guinea in the east (Fig.
The island chain of LSI comprises 5.2% of endemic species based on families treated in Flora Malesiana (
The islands of LSI include seven ecoregions in two biomes, Tropical and Subtropical Moist Forests and Tropical and Subtropical Dry Forests (Fig.
The “Tropical and Subtropical Dry forests” consist of three ecoregions from Lombok in the west to Timor and Wetar in the east. All three ecoregions are critically endangered/endangered. The largest of the ecoregions is “Lesser Sunda Deciduous Forests” (https://www.worldwildlife.org/ecoregions/aa0201) on Lombok, Sumbawa, Flores and west to Alor including smaller surrounding islands. It consists mainly of semi-evergreen dry forests. With an average annual rainfall of 1,349 mm, this is the driest area but also the most seasonal in Indonesia (
Further east, two more ecoregions of “Tropical and Subtropical Moist Deciduous Forests” occur. The “Banda Sea Islands Moist Deciduous Forest” (https://www.worldwildlife.org/ecoregions/aa0102) includes all islands west of Timor/Wetar, except Aru Island. It includes evergreen rain forest (Kepulauan Kai), semi-evergreen rain forest, moist deciduous forest, and dry deciduous forest (
Bryophytes, including mosses, liverworts and hornworts, are the second largest group of land plants after flowering plants and are pivotal in our understanding of early land plant evolution (e.g.,
It is clear that bryophytes from this region have been overlooked historically, and under-collected, compared to seed plants, birds, and other organisms, forming a remarkable gap in the flora of Indonesia and/or Timor-Leste. Publications dealing with liverworts of LSI also are few and scattered. The first apparent report of liverworts from LSI was by
We here present the first-ever checklist of liverworts and hornworts for the Lesser Sunda Islands to serve as the baseline information in our study of the liverworts diversity of this archipelago. Furthermore, this checklist can serve to promote and encourage bryological research in the region. The significance of checklists is summarized by
As with many other regions in the world, given the relatively poor focus on liverworts in LSI historically, we predict that a vast number of new records are yet to be reported for the area. In this checklist we report 129 taxa, with another 12 taxa questioned and three rejected. The number of known species from individual islands varies from 101 (Bali) to 0 (Barat Daya Islands), but only Bali has more than 20 known species (Fig.
Nomenclature and taxonomy follows the world checklist of hornworts and liverworts (
This checklist covers the Lesser Sunda Islands based on reports from the literature that have been subdivided into 10 geographical units that correspond to geopolitical units. These include from west to east (Fig.
Recently, there has been some debate whether the type information by
It is not always clear if more than one collection exists for a possible type.
Anastrophyllum integerrimum Steph., Sp. Hepat. (Stephani) 6: 107, 1917 (
Chiloscyphus communis Steph., Bull. Herb. Boissier (sér. 2) 7 (10): 839 [=Sp. Hepat. (Stephani) 3: 211], 1907 (
Drepanolejeunea moluccensis Herzog, Ann. Bryol. 7: 88, 1934 (
Fimbraria latifrons Steph., Sp. Hepat. (Stephani) 6: 15, 1917 (
Frullania longispica Steph., Sp. Hepat. (Stephani) 4: 454, 1911 (
Lepidozia newtonii Steph., Sp. Hepat. (Stephani) 3: 623, 1909 (
Madotheca elbertii Steph., Sp. Hepat. (Stephani) 6: 520, 1924 (
Mastigobryum sumbavense Steph., Hedwigia 25 (6): 236, 1886 (
Folioceros
*** Folioceros fuciformis (Mont.) D.C.Bharadwaj Bali:
Acanthocoleus
*** Acanthocoleus javanicus (Steph.) Kruijt. Bali:
Acrolejeunea
*** Acrolejeunea aulacophora (Mont.) Steph. Lesser Sunda Is.: Bali:
*** Acrolejeunea fertilis (Reinw., Blume et Nees) Schiffn. Bali:
Aneura
*** Aneura maxima (Schiffn.) Steph. Bali:
*** Aneura pinguis (L.) Dumort. Bali:
Asterella
*** Asterella blumeana (Nees) Kachroo Lombok:
*** Asterella vulcanica (Schiffn.) Kachroo et Bapna Bali:
Bazzania
*** Bazzania longicaulis (Sande Lac.) Schiffn. Sumba:
*
Cheilolejeunea
** Cheilolejeunea ceylanica (Gottsche) R.M.Schust. et Kachroo Sumba:
*** Cheilolejeunea trapezia (Nees) Kachroo et R.M.Schust. Bali:
Chiastocaulon
*** Chiastocaulon dendroides (Nees) Carl. Bali:
*** Chiastocaulon oppositum (Reinw., Blume et Nees) S.D.F.Patzak, M.A.M.Renner, Schäf.-Verw. et Heinrichs. Lesser Sunda Is.:
Cololejeunea
*** Cololejeunea angustiflora (Steph.) Mizut. Bali:
*** Cololejeunea appressa (A.Evans) Benedix. Bali:
** Cololejeunea gottschei (Steph.) Pandé, K.P.Srivast. et Ahmad Bali:
*** Cololejeunea macounii (Spruce) A.Evans Bali:
*** Cololejeunea obliqua (Nees et Mont.) Schiffn. Bali:
*** Cololejeunea ocelloides (Horik.) Mizut. Bali:
** Cololejeunea subfloccosa Mizut. Bali:
** Cololejeunea triapiculata (Herzog) Tixier Bali:
*** Cololejeunea trichomanis (Gottsche) Besch. Bali:
Colura
**
Diplasiolejeunea
*** Diplasiolejeunea cavifolia Steph. Bali:
Drepanolejeunea
*** Drepanolejeunea levicornua Steph. Sumba:
** Drepanolejeunea moluccensis Herzog. Bali: Syntype,
*** Drepanolejeunea pentadactyla (Mont.) Steph. Bali:
Dumortiera
*** Dumortiera hirsuta (Sw.) Nees Bali:
Fossombronia
*** Fossombronia himalayensis Kashyap. Bali:
Frullania
** Frullania apiculata (Reinw., Blume et Nees) Nees Sumba:
*** Frullania ericoides (Nees) Mont. Bali:
*** Frullania gaudichaudii (Nees et Mont.) Nees et Mont. Bali:
*** Frullania gracilis (Reinw., Blume et Nees) Nees Bali:
** Frullania intermedia subsp. intermedia. Aru Islands:
*** Frullania junghuhniana Gottsche. Bali:
*** Frullania meyeniana Lindenb. Bali:
***
*** Frullania nodulosa (Reinw., Blume et Nees) Nees Aru Islands:
*** Frullania ornithocephala (Reinw., Blume et Nees) Nees Bali:
*** Frullania rio-janeirensis (Raddi) Ångstr. Bali:
*** Frullania serrata Gottsche Lombok:
*** Frullania ternatensis Gottsche Bali:
** — var. non-appendiculata S.Hatt. Bali:
*
*** Frullania trichodes Mitt. Kai Islands:
Gottschelia
*** Gottschelia schizopleura (Spruce) Grolle Lesser Sunda Is.:
Herbertus
** Herbertus ceylanicus (Steph.) Abeyw. Flores:
*** Herbertus dicranus (Gottsche, Lindenb. et Nees) Trevis. Bali:
** Herbertus longispinus J.B.Jack et Steph. Flores:
** Herbertus ramosus (Steph.) H.A.Mill. Bali:
*** Herbertus sendtneri (Nees) Lindb. Lesser Sunda Is.:
Heteroscyphus
*** Heteroscyphus argutus (Reinw., Blume et Nees) Schiffn. Aru Islands:
*** Heteroscyphus aselliformis (Reinw., Blume et Nees) Schiffn. Bali:
*** Heteroscyphus coalitus (Hook.) Schiffn. Bali:
*** Heteroscyphus splendens (Lehm. et Lindenb.) Grolle Sumba
Jackiella
*** Jackiella javanica Schiffn. Bali:
Lejeunea
*** Lejeunea alata Gottsche Sumba:
*** Lejeunea apiculata Sande Lac. Bali:
*** Lejeunea mimula Hürl. Bali:
Lepidozia
* Lepidozia newtonii Steph. Timor: Type,
Leptolejeunea
*** Leptolejeunea elliptica (Lehm. et Lindenb.) Besch. Sumba:
*** Leptolejeunea epiphylla (Mitt.) Steph. Bali:
** Leptolejeunea foliicola Steph. Bali:
* Leptolejeunea massartiana Schiffn. ex Herzog Bali:
*** Leptolejeunea subacuta Steph. ex A.Evans Bali:
Lopholejeunea
*** Lopholejeunea eulopha (Taylor) Schiffn. Bali:
*** Lopholejeunea horticola Schiffn. Bali:
*** Lopholejeunea nigricans (Lindenb.) Schiffn. Bali:
*** Lopholejeunea recurvata Mizut. Bali:
*** Lopholejeunea subfusca (Nees) Schiffn. Lesser Sunda Is.:
*** Lopholejeunea zollingeri (Steph.) Schiffn. Bali:
Marchantia
*** Marchantia acaulis Steph. Bali:
*** Marchantia emarginata Reinw., Blume et Nees Bali:
*** — subsp. emarginata. Bali:
*** Marchantia geminata Reinw., Blume et Nees Bali:
*** Marchantia treubii Schiffn. Lesser Sunda Is.:
Metalejeunea
*** Metalejeunea cucullata (Reinw., Blume et Nees) Grolle Bali:
Metzgeria
*** Metzgeria ciliata Raddi Bali:
*** Metzgeria consanguinea Schiffn. Sumba:
* Metzgeria foliicola Schiffn. Flores:
*** Metzgeria lindbergii Schiffn. Bali:
Pallavicinia
*** Pallavicinia lyellii (Hook.) Gray Bali:
Plagiochila
*** Plagiochila bantamensis (Reinw., Blume et Nees) Mont. Bali:
*** Plagiochila frondescens (Nees) Lindenb. Bali:
*** Plagiochila javanica (Sw.) Nees et Mont. Bali:
*** Plagiochila junghuhniana Sande Lac. Bali:
*
** Plagiochila massalongoana Schiffn. Bali:
*** Plagiochila obtusa Lindenb. Bali:
*** Plagiochila parvifolia Lindenb. Bali:
** Plagiochila propinqua Sande Lac. Bali:
*** Plagiochila renitens (Nees) Lindenb. Bali:
*** Plagiochila salacensis Gottsche Bali:
*** Plagiochila sciophila Nees Bali:
** Plagiochila semidecurrens (Lehm. et Lindenb.) Lindenb. Bali:
** Plagiochila spathulifolia Mitt. Bali:
*** Plagiochila teysmannii Sande Lac. Bali:
Pleurozia
*** Pleurozia gigantea (F.Weber) Lindb. Flores:
Porella
** Porella acutifolia (Lehm. et Lindenb.) Trevis. Bali:
** — var. acutifolia. Lombok: Lectotype of Madotheca elbertii,
Ptychanthus
*** Ptychanthus striatus (Lehm. et Lindenb.) Nees Bali:
Radula
** Radula acuminata Steph. Bali:
*** Radula campanigera Mont. Bali:
** Radula javanica Gottsche Bali:
* Radula multiflora Gottsche ex Schiffn. Aru Islands:
* Radula pinnulata Mitt. Aru Islands:
*** Radula ventricosa Steph. Bali:
Reboulia
*** Reboulia hemisphaerica (L.) Raddi Bali:
Riccardia
** Riccardia crenulata Schiffn. Bali:
Riccia
*** Riccia billardierei Mont. et Nees Bali:
*** Riccia cruciata Kashyap. Bali:
*** Riccia discolor Lehm. et Lindenb. Bali:
*** Riccia junghuhniana Nees et Lindenb. Bali:
*** Riccia mangalorica Ahmad ex Jovet-Ast Bali:
Scapania
*** Scapania javanica Gottsche Bali:
Schiffneriolejeunea
*** Schiffneriolejeunea tumida (Nees) Gradst. Bali:
*** — var. haskarliana (Gottsche) Gradst. et Terken. Bali:
Schistochila
*** Schistochila aligera (Nees et Blume) J.B.Jack et Steph. Bali:
*** Schistochila blumei (Nees) Trevis. Bali:
Solenostoma
*** Solenostoma tetragonum (Lindenb.) R.M.Schust. ex Váňa et D.G.Long Bali:
*** Solenostoma truncatum (Nees) R.M.Schust. ex Váňa et D.G.Long Bali:
Spruceanthus
*** Spruceanthus polymorphus (Sande Lac.) Verd. Aru Islands:
*** Spruceanthus semirepandus (Nees) Verd. Lombok:
Thysananthus
*** Thysananthus humilis (Gottsche) Sukkharak et Gradst. Bali:
*** Thysananthus ligulatus (Lehm. et Lindenb.) Sukkharak et Gradst. Sumba:
*** Thysananthus spathulistipus (Reinw., Blume et Nees) Lindenb. Bali:
*** Thysananthus virens Ångstr. Bali:
Wiesnerella
*** Wiesnerella denudata (Mitt.) Steph. Bali:
A couple of taxa are published from Lesser Sunda Islands as varieties of species synonymized under other names, without transferring or synonymizing the variety. We have not been able to trace any specimen that they may be based on and, thus, not been able to refer them to any valid taxon.
Chiloscyphus
? Chiloscyphus zollingeri var. subintegerrimus Schiffn. Bali:
Riccardia
? Riccardia platyclada var. leiomitra Schiffn. Bali:
Bazzania
** Bazzania ceylanica (Mitt.) Steph. Lesser Sunda Is.:
*** Bazzania erosa (Reinw., Blume et Nees) Trevis. Lesser Sunda Is.:
*** Bazzania tridens (Reinw., Blume et Nees) Trevis. Lesser Sunda Is.:
Ceratolejeunea
*** Ceratolejeunea cf. papuliflora Steph. Sumba:
Cololejeunea
*** Cololejeunea cf. lanciloba Steph. Bali:
** Cololejeunea cf. serrulata Steph. Bali:
Colura
*** Colura ari (Steph.) Steph. Lesser Sunda Is.:
** Colura imperfecta Steph. Lesser Sunda Is.:
Conoscyphus
*** Conoscyphus trapezioides (Sande Lac.) Schiffn. Lesser Sunda Is.:
Drepanolejeunea
*** Drepanolejeunea ternatensis (Gottsche) Schiffn. Lesser Sunda Is.:
Radula
*** Radula complanata (L.) Dumort. Bali:
Targionia
*** Targionia hypophylla L. Bali:
Ceratolejeunea
*** Ceratolejeunea ceratantha (Nees et Mont.) Schiffn. Sumbawa:
Frullania
* Frullania ludoviciae Steph. Sumbawa:
Thysananthus
*** Thysananthus auriculatus (Wilson et Hook.) Sukkharak et Gradst. Bali:
Anastrophyllum integerrimum Steph. = Gottschelia schizopleura
Aneura crenulata (Schiffn.) Steph. ≡ Riccardia crenulata
Aspiromitus falsinervius (Lindenb. ex Meissner) Steph. = Folioceros fuciformis
Chiloscyphus argutus (Reinw., Blume et Nees) Nees ≡ Heteroscyphus argutus
Chiloscyphus aselliformis (Reinw., Blume et Nees) Nees ≡ Heteroscyphus aselliformis
Chiloscyphus coalitus (Hook.) Nees ≡ Heteroscyphus coalitus
Chiloscyphus communis Steph. = Heteroscyphus coalitus
Chiloscyphus decurrens (Reinw., Blume et Nees) Nees = Heteroscyphus splendens
Chiloscyphus trapezioides Sande Lac. ≡ Conoscyphus trapezioides
Cololejeunea goebelii (Gottsche ex Schiffn.) Schiffn. = Cololejeunea trichomanis
Cololejeunea leonidens var. saccata Benedix = Cololejeunea ocelloides
Cololejeunea nymannii (Steph.) Benedix = Cololejeunea obliqua
Colura apiculata Steph. = Colura leratii
Fimbraria latifrons Steph. = Reboulia hemisphaerica
Frullania africana Steph. = Frullania rio-janeirensis
Frullania longispica Steph. = Frullania ericoides
Frullania moniliata subsp. breviramea (Steph.) Verd. = Frullania moniliata
Frullania secundiflora Mont. = Frullania nodulosa
Frullania squarrosa (Mont.) Nees = Frullania ericoides
Frullania tamarisci var. breviramea (Steph.) S.Hatt. = Frullania moniliata
Frullania tenuicaulis Mitt. = Frullania trichodes
Herbertus armitanus (Steph.) H.A.Mill. = Herbertus sendtneri
Heteroscyphus communis (Steph.) Schiffn. = Heteroscyphus coalitus
Heteroscyphus decurrens (Nees) Schiffn. = Heteroscyphus splendens
Jungermannia tetragona Lindenb. ≡ Solenostoma tetragonum
Jungermannia truncata Nees ≡ Solenostoma truncatum
Lejeunea ceratantha Nees et Mont. ≡ Ceratolejeunea ceratantha
Lejeunea subfusca (Nees) Nees et Mont. ≡ Lopholejeunea subfusca
Lopholejeunea sagrana var. β subfusca (Nees) Schiffn. ≡ Lopholejeunea subfusca
Madotheca elbertii Steph. = Porella acutifolia var. acutifolia
Mastigobryum sumbavense Gottsche ex Steph. ≡ Bazzania sumbavensis
Mastigolejeunea auriculata (Wilson et Hook.) Steph. ≡ Thysananthus auriculatus
Mastigolejeunea humilis (Gottsche) Schiffn. ≡ Thysananthus humilis
Mastigolejeunea ligulata (Lehm. et Lindenb.) Schiffn. ≡ Thysananthus ligulatus
Mastigolejeunea virens (Ångstr.) Steph. ≡ Thysananthus virens
Phragmicoma polymorpha Sande Lac. ≡ Spruceanthus polymorphus
Plagiochila dendroides (Nees) Lindenb. ≡ Chiastocaulon dendroides
Plagiochila jackii Schiffn. nom. illeg. = Plagiochila salacensis
Plagiochila opposita (Reinw., Blume et Nees) Lindenb. ≡ Chiastocaulon oppositum
Plagiochilion oppositum (Reinw., Blume et Nees) S.Hatt. ≡ Chiastocaulon oppositum
Porella acutifolia var. elbertii (Steph.) S.Hatt. = Porella acutifolia var. acutifolia
Riccardia tenuicostata Schiffn. nom. illeg. = Riccardia crenulata
Riccardia tenuicostata Schiffn. = Riccardia inconspicua
Riccia mangalorica Ahmad nom. inval. ≡ Riccia mangalorica
Thysananthus polymorphus (Sande Lac.) Schiffn. ≡ Spruceanthus polymorphus
We would like to thank Research and Innovation in Science and Technology Project (RISET-Pro) - Ministry of Research and Technology/National Research and Innovation Agency of Indonesia for granting the scholarship to the first author, Cibodas Botanic Garden, Research Center for Plant Conservation and Botanic Gardens - LIPI for support, and Mr. Lukman Budiman from PDDI – LIPI for his kindly assistance in the BO library. The Biodiversity Heritage Library is acknowledged for the facility they provide that has greatly accelerated our efforts. The generous support by the National Science Foundation (Award No.’s 1145898, 1146168, 1458300, 1541506, 2001509 and 0531730) is gratefully acknowledged. We also recognize the support of the Museum Collection Spending Fund, administered by Field Museum, as well as curatorial support provided by Y. Rodriguez, L. Kawasaki, and A. Balla (Field Museum). Alfons Schäfer-Verwimp kindly provided us with specimen data from his unpublished Diplasiolejeunea cavifolia. We are very grateful for the helpful suggestions and comments made by the editor and reviewers: Matt A. Renner, S.R. Gradstein, Li Zhang, and an anonymous reviewer.