Bryologie 2023 ● 44 ● 1

Directeur de la publication / Publication director: Bruno DAVID

Président du Muséum national d’Histoire naturelle

Rédacteur en chef / Editor-in-chief: Denis LAMY

Assistante de rédaction / Assistant editor: Chris LE COQUET-LE ROUX (bryo@[Link])

Mise en page / Page layout: Chris LE COQUET-LE ROUX

Rédacteurs associés / Associate Editors

Biologie moléculaire et phylogénie / Molecular biology and phylogeny
Bernard GOFFINET
Department of Ecology and Evolutionary Biology, University of Connecticut (United States)
Mousses d’Europe / European mosses
Isabel DRAPER
Centro de Investigación en Biodiversidad y Cambio Global (CIBC-UAM), Universidad Autónoma de Madrid (Spain)
Francisco LARA GARCÍA
Centro de Investigación en Biodiversidad y Cambio Global (CIBC-UAM), Universidad Autónoma de Madrid (Spain)
Mousses d’Afrique et d’Antarctique / African and Antarctic mosses
Rysiek OCHYRA
Laboratory of Bryology, Institute of Botany, Polish Academy of Sciences, Krakow (Pologne)
Bryophytes d’Asie / Asian bryophytes
Rui-Liang ZHU
School of Life Science, East China Normal University, Shanghai (China)
Bioindication / Biomonitoring
Franck-Olivier DENAYER
Faculté des Sciences Pharmaceutiques et Biologiques de Lille, Laboratoire de Botanique et de Cryptogamie, Lille (France)
Écologie des bryophytes / Ecology of bryophyte
Nagore GARCÍA MEDINA
Department of Biology (Botany), and Centro de Investigación en Biodiversidad y Cambio Global (CIBC-UAM), Universidad Autónoma de Madrid (Spain)

Couverture / Cover:
Photographie de Heteroscyphus deplanchei prise par L. Thouvenot en 2020 / Photograph of Heteroscyphus deplanchei taken by L. Thouvenot in 2020

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 A taxonomic revision of the
Lophocoleaceae Vanden Berghen
(Marchantiophyta) of New Caledonia

Louis THOUVENOT
11 rue Saint Léon, 66000 Perpignan (France)
[Link]@[Link]

Submitted on 21 February 2022 | Accepted on 29 July 2022 | Published on 16 January 2023

Thouvenot L. 2023. — A taxonomic revision of the Lophocoleaceae Vanden Berghen (Marchantiophyta) of New
Caledonia. Cryptogamie, Bryologie 44 (1): 1-60. [Link]
[Link]

ABSTRACT
Prior to this study, 51 species of Lophocoleaceae Vanden Berghen were reported from New Caledonia.
Evolving taxonomic concepts have resulted in major recent changes in the taxonomy of Lophoc-
oleaceae at generic level. This paper presents the results of taxonomic revision of the Lophocoleaceae
KEY WORDS
New Caledonia, of New Caledonia based on study of the types and further historical as well as recent collections. All
liverworts, accepted species are described and illustrated, with citation of types and synonyms. A key to the New
Lophocoleaceae, Caledonian species and an index of species names, including those excluded from Lophocoleaceae, are
Acrobolbaceae,
Brevianthaceae, provided. As a result of this revision, 27 species of Lophocoleaceae are accepted for New Caledonia.
determination key, Heteroscyphus Schiffn. is the largest genus with 18 species, followed by Lophocolea (Dumort.) Dumort.
lectotypifications,
new synonyms, (5 spp.), Chiloscyphus Corda (1), Cryptolophocolea L.Söderstr. (2) and Otoscyphus [Link], Bardat &
new combinations. Thouvenot (1). Seven species are excluded from the territory and one species is a doubtful taxon.

RÉSUMÉ
Révision taxonomique des Lophocoleaceae Vanden Berghen (Marchantiophyta) de Nouvelle-Calédonie.
Avant cette étude, 51 espèces de Lophocoleaceae Vanden Berghen étaient connues en Nouvelle-Calé-
donie. L’évolution des idées en taxonomie a conduit à d’importants changements dans la conception
des genres de Lophocoleaceae. Cet article présente les résultats de la révision taxonomique des Lopho-
coleaceae en Nouvelle-Calédonie à partir de l’examen des types et des exemplaires disponibles dans les
MOTS CLÉS
Nouvelle-Calédonie, collections historiques ou contemporaines. Toutes les espèces acceptées sont décrites et illustrées avec
hépatiques, la citation des types et des synonymes. Une clé de détermination des espèces de Nouvelle-Calédonie et
Lophocoleaceae, un index des noms d’espèces, incluant celles qui ont été récemment exclues des Lophocoleaceae, sont
Acrobolbaceae,
Brevianthaceae, fournis. Il résulte de cette révision une liste de 27 Lophocoleaceae acceptées pour la Nouvelle-Calé-
clé de détermination, donie. Heteroscyphus Schiffn. est le genre le plus riche avec 18 espèces, suivi par Lophocolea (Dumort.)
lectotypifications,
synonymes nouveaux, Dumort. (5 spp.), Chiloscyphus Corda (1), Cryptolophocolea L.Söderstr. (2) et Otoscyphus [Link],
combinaisons nouvelles. Bardat & Thouvenot (1). Sept espèces sont exclues du territoire et une garde un statut douteux.

CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1) © Publications scientifiques du Muséum national d’Histoire naturelle, Paris. [Link]/bryologie 1
 Thouvenot L.

INTRODUCTION Tetracymbaliella Grolle in Heteroscyphus Schiffn. Since then,

Lophocolea has been reinstated as a genus in the classification
The taxonomy of the liverwort family Lophocoleaceae Van- of the Marchantiophyta (Crandall-Stotler et al. 2009) or the
den Berghen has been reviewed by Engel & Schuster (1984), world checklist of hornworts and liverworts (Söderström et al.
He-Nygrén & Piippo (2003), Hentschel et al. (2006a, b), 2016), Cryptolophocolea and Otosycphus [Link], Bardat &
Crandall-Stotler et al. (2009), Söderström et al. (2013a) and Thouvenot (Engel et al. 2012) have been added, and Tetra-
others. Patzak et al. (2016) discussed the status of two import- cymbaliella has been transferred to Brevianthaceae (Söder-
ant genera of this family, Lophocolea (Dumort.) Dumort. and ström et al. 2013a) and Conoscyphus Mitt. to Acrobolbaceae
Chiloscyphus Corda, which were treated in the “Checklist of (Dimon et al. 2018).
the liverworts and hornworts of New Caledonia” (Thouvenot This study deals with 45 species names listed by Thouvenot
et al. 2011) and in many other publications as one single broad et al. (2011) together with four species that were subsequently
genus Chiloscyphus (with Lophocolea included as a subgenus), described as new to science (Thouvenot & Price 2020; Thou-
following Engel & Schuster (1984), but have recently again venot & Engel 2021; Engel et al. 2021), one species new to
been reinstated as two separate genera based on molecular New Caledonia (Thouvenot & Müller 2021), two unpublished
evidence (Hentschel et al. 2006a, b, 2007; Glenny et al. occurrences, and four new combinations published by Engel
2009), with recognition of a third genus, Cryptolophocolea et al. (2012) and Thouvenot et al. (2018).
L.Söderstr. (e.g. Söderström et al. 2013a, 2016).
Since Nees (1836), the most widely accepted criteria seg-
regating the genera of Lophocoleaceae are the position and MATERIAL AND METHODS
shape of the gametangia (Table 1; see also Engel & Schuster
1984). In addition, vegetative features like connate vs free For the purpose of updating the checklist of the liverworts
underleaves are also of interest both at regional and wider of New Caledonia (Thouvenot et al. 2011), New Caledonian
scale. The recent clarification of the generic classification species of Lophocoleaceae were checked from type specimens
using a molecular approach (Hentschel et al. 2006b, 2007; and a few additional historic collections in G, PC, REN, BM,
Glenny et al. 2009) has confirmed the opinion of Grolle L, Z+Zt together with recent materials collected by Frank
(1995) and others that Chiloscyphus is a small Laurasian Müller (2001-2003) and the author (2008-2019) and kept
genus (Söderström et al. 2013a). But a few species initially in DR, PC and the author’s private herbarium. In addition,
reported for this genus in New Caledonia are still pending the author had the chance to observe the single specimen of
further studies since the specimens available including types Lophocolea muricata (Lehm.) Nees hitherto reported from
lack gametoecia of either one or both sexes. This is of critical New Caledonia (Hürlimann 1998), and kept at Z, thanks to
importance to assign them to Chiloscyphus genus, as well as photos and comments from Heike Hofmann. The original
molecular analysis, but if that is confirmed, the range of this diagnoses together with personal observations allowed us
genus will be broadened to southern hemisphere. to check the discriminant characters and to provide concise
This study deals with Lophocoleaceae in New Caledo- descriptions and drawings from fresh material using a drawing
nia with the aim of establishing a reliable and updated list tube. As to typification, original specimens were rarely fully
of species present in the territory, and with removal of all cited by Stephani, Pearson and Mitten, but was usually done
unnecessary names (synonyms and erroneously reported by Herzog. The loan of original specimens from BM allowed
taxa). None of the modern taxonomic studies have included to select lectotypes for Pearson’s new species. With regards to
New Caledonian species, so this work cannot benefit from species described by Stephani, the citations in Bonner (1963,
recent data as to generic assignment. Because of the limited 1966) of original material present in the Stephani herbarium
scope of the paper, the author follows the most recent generic (G) have been taken into account. In view of the plasticity
classification of the family (Söderström et al. 2016). Dif- of the species, many specimens must be studied for their cir-
ficulty in circumscribing the species due to their plasticity cumscription. Unfortunately, several species are only known
have been underlined by many authors (e.g. Jones 1953). from the type or few specimens. Nevertheless, the synonymy
In New Caledonia, a good example may be Heteroscyphus of many species names has been established. The possibility
grandiflorus (Steph.) Hürl. whose variability is commented remains, however, that some of the species names accepted in
below (see comments in corresponding paragraph). Many this treatment will turn out to be synonyms of species described
species were described under several names based on small from other countries. When samples of related extraterritorial
morphological differences which turned out to be variable in species could not be checked, the author has conservatively
single specimens or within a range of samples. For example, kept the New Caledonian species name, pending further
Heteroscyphus coalitus (Hook.) Schiffn. currently includes five study. In addition, a few species known only from limited
heterotypic synonyms in Japan (Yamada & Iwatsuki 2006) type material unsuitable for detailed morphological analysis
and four in Java (Söderström et al. 2010). have been included under their original name.
As mentioned, the Lophocoleaceae treatment in the checklist Engel & Glenny (2019) highlighted the low reliability of
of the liverworts of New Caledonia (Thouvenot et al. 2011) taxonomic criteria used to define the genera of Lophocoleoideae.
essentially followed Engel & Schuster (1984), so that all Here, the generic attribution of the New Caledonian species
species of Lophocolea were included in Chiloscyphus, and of is largely based on characters of the gametangia, including

2 CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1)

 Lophocoleaceae of New Caledonia

Table 1. — Discriminant features of Lophocoleaceae Vanden Berghen genera occurring in New Caledonia.

Heteroscyphus Chiloscyphus Lophocolea Cryptolophocolea Otoscyphus [Link],

Schiffn. Corda (Dumort.) Dumort. L.Söderstr. Bardat & Thouvenot
Androecia position On short lateral On long normal On long normal On long normal On long normal leaved
branches lacking leaved shoots leaved shoots leaved shoots shoots
normal leaves
Male bracts Much smaller than Like the leaves but Smaller than leaves, Like the leaves but Smaller than leaves,
leaves, fully saccate with small dorsal saccate at dorsal with dorsal saccate dorsal saccate base
saccate base base, ventral lamina base, shorter than ½ bract length
variously developed the lamina
Antheridia stalk Biseriate to Uniseriate Uniseriate Biseriate Unknown
oligoseriate
Gynoecial position On short lateral On short lateral On normal leaved On normal leaved On normal leaved
branches lacking branches lacking shoots, more or less elongate shoots elongate shoots and
normal leaves normal leaves elongate short leafless ventral
branches
Perianth Cyathiform. Cupulate. Oblong. 3-keeled Oblong. 3-keeled Cyathiform.
Obscurely plicate Obscurely plicate Obscurely plicate
Calyptra Hidden deep within Reaching perianth Hidden deep within Hidden deep within Hidden deep within
perianth apex to exserted perianth perianth perianth
Leaf insertion Mainly subopposite Mainly alternate Alternate Subopposite Alternate
Underleaf connation Typically bilateral Free or unilateral Free or unilateral Bilateral. Free
and wide and narrow and narrow Typically wide
Cell walls Thin, rarely thick Thin Thin Thin Thin
Trigones Trigones typically Trigones small Trigones small Trigones small Trigones large
large or lacking or lacking or large

their shape and their location on main shoots or on abbrevi- Engel & Schuster (1984), most of the former Chiloscyphus spe-
ated branches, associated or not with normal vegetative leaves. cies in New Caledonia belong in Heteroscyphus. This includes
This approach is marred, however, by the frequent absence of specimens which have gametangial characters of Heteroscyphus,
gametangia in historic specimens (Engel 2015). Additional but with “lophocoleoid” vegetative features, e.g. underleaves
relevant vegetative characters include leaf ornamentation and deeply bifid and free from the leaves or connate on one side
insertion, underleaf shape and connation to the leaves, as well only, small trigones or none, and leaves dorsally free. How-
as cell wall thickenings (Table 1). ever, a handful of former Chiloscyphus species included in
When gametoecia are lacking, the original species names the checklist (Thouvenot et al. 2011) seem to be nearer to
are kept in the incertae sedis group. Chiloscyphus than to Lophocolea or Heteroscyphus so that the
Collections by the author in New Caledonia were made on strictly Laurasian range of Chiloscyphus can be questionable.
the basis of the following permits: Province Sud, nr 1238- Besides members of Heteroscyphus, Lophocoleaceae of New
2012, 794-2016, 2825-2019; Province Nord, nr 609012- Caledonia include five species of Lophocolea, one of Chilos-
1275-2016, 609011-45/2019. Unless otherwise specified, cyphus, two of Cryptolophocolea and the monospecific genus
specimens cited are kept in the author’s private herbarium. Otoscyphus. A single further species is kept as incertae sedis,
in the absence of fertile specimen available for examination.
In addition, seven species names are excluded from the New
RESULTS AND DISCUSSION Caledonian flora. In comparison, 41 Lophocoleaceae spe-
cies were listed by Thouvenot et al. (2011) (apart from four
This treatment accepts 27 species of Lophocoleaceae in New that were recently classified in other families). Seventeen
Caledonia, plus one species whose generic assignation is doubt- of these have been reduced to synonymy, among them five
ful, including seven that were described as new to science or under Heteroscyphus coalitus, five under Lophocolea convexula
reported as new to New Caledonia since the publication of the Sande Lac. and one (Chiloscyphus novae-caledoniae [Link] &
checklist (Thouvenot et al. 2011): Chiloscyphus parapilistipulus [Link].) under Goebelobryum unguiculatum (Hook.f. &
Thouvenot (here treated as Heteroscyphus), Heteroscyphus assur- Taylor) Grolle, a member of the family Acrobolbaceae. In
gentissimus [Link], Thouvenot & Frank Müll., H. diestianus the index (Appendix 1), the synonyms are listed with the
(Sande Lac.) Piippo, H. kanakensis Thouvenot, H. supinopsis accepted names in bold face. The key to the species given
[Link], Thouvenot & Frank Müll., H. succulentus (Gott.) below includes the genera recently transferred towards differ-
Schiffn. and Lophocolea bidentata (L.) Dumort. The genus ent families. To allow for identification of sterile specimens,
Heteroscyphus is, by far, the richest Lophocoleaceae genus in the key uses primarily vegetative features.
New Caledonia with 18 species. In comparison, Papua New Distribution maps of the Lophocoleaceae species in New
Guinea has 15 species (Piippo 1985, 1992). As explained by Caledonia are available on [Link]

CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1) 3

Thouvenot L.

Key to New Caledonian species of Lophocoleaceae Vanden Berghen

and selected taxa recently transferred to other families

1. Basal parts of the leaves with pouches made of expanded or incurved parts of the lamina .............................. 2
— Leaves without pouches ................................................................................................................................. 3
2. Each pouch formed by the incurved basal margin of a single leaf or underleaf; underleaves without linear lobes;
leaves widely ovate with shortly toothed margins ................................Tetracymbaliella Grolle (Brevianthaceae)
— Each pouch formed by two successive leaves, the dorsally enlarged base of a leaf set above the previous one;
underleaves with spreading linear lobes; leaves rectangular in outline, entire margined, the apices deeply bifid
............................................. Otoscyphus crassicaulis (Steph.) [Link], Bardat & Thouvenot (Lophocoleaceae)
3. Plants brownish; leaf cells mammillose, with large contiguous spheroid trigones; underleaves reniform-rounded
superficially toothed ........................... Conoscyphus trapezioides (Sande Lac.) Mitt. ex Schiffn. (Acrobolbaceae)
— Plants green, rarely red tinged; leaf cells smooth, rarely bulging or papillose, with trigones null to large and rarely
contiguous; underleaves of various shapes ...................................................................................................... 4
4. Plants light green usually red tinged; underleaves vestigial or very small, narrower than the stem, and bifid with
linear lobes, made of up to three short cells; fertile plants with a long cylindrical marsupium at the end of the
fertile shoots .................................... Goebelobryum unguiculatum (Hook.f. & Taylor) Grolle (Acrobolbaceae)
— Plants green when fresh, without red pigment; underleaves usually well developed, at least as wide as the stem;
fertile plants without a marsupium ............................................................... Lophocoleaceae Vanden Berghen
5. Leaves subopposite and underleaves connate to both nearest leaves, connation usually wide .......................... 6
— Leaves alternate, rarely subopposite, with underleaves at most narrowly connate to one of the nearest leaves or
free on both sides ........................................................................................................................................ 23
6. Gynoecia and androecia terminal on main shoots or long leaved branches; leaves asymmetrically oblong with
the ventral base widened, leaf apices with 1-2 sharp lobes, lateral margins sharply toothed or not; underleaves
ovate or reniform, shortly bifid and more or less toothed all around .............................................................. 7
— Gynoecia and androecia terminal on short leafless ventral-lateral branches; leaves and underleaves of various
shapes ........................................................................................................................................................... 8
7. Normal leaves canaliculate, gradually narrowing from base to apex; lateral margins of leaves and underleaves
usually toothed; leaf cells with strong trigones ......................... Cryptolophocolea subcostata (Steph.) Thouvenot
— Normal leaves convex, ovate oblong to lingulate, the apex somewhat narrower; lateral margins of leaves usually
naked or with a few teeth; lateral margins of underleaves with at most a single tooth on both sides; leaf cells
without trigones ........................................................ Cryptolophocolea explanata (Mitt.) Váňa & Crand.-Stotl.
8. Leaves entire or with the rounded apices superficially retuse but in a few shoots some of the leaves may be shortly
bifid .............................................................................................................................................................. 9
— Leaves lobed, toothed or ciliate-laciniate, or at least with a small tooth at both angles of truncate apices ..... 16
9. Underleaves widely reniform, the apices entire, margins with a few scattered small teeth; leaves entire except
small teeth at the base of the dorsal margins ................................. Heteroscyphus diestianus (Sande Lac.) Piippo
— Underleaves reniform or not, the apices at least superficially bifid, margins variously ornamented; leaves never
with teeth restricted to the base of the dorsal margin ................................................................................... 10
10. Leaves asymmetrically oval acuminate .............. Heteroscyphus subacuminatus (Herzog) Thouvenot, comb. nov.
— Leaves of various shapes, rounded, ovate or oblong, never acuminate .......................................................... 11
11. Underleaves shortly bifid with margins densely toothed all around ..................................................................
..........................................................................................Heteroscyphus splendens (Lehm. & Lindenb.) Grolle
— Underleaves more conspicuously bifid with margins remotely toothed or laciniate ...................................... 12
12. Leaves oblong; cells without trigones or with nodulose trigones .................................................................. 13
— Leaves widely ovate to rounded; cells with large nodulose or truncate trigones ............................................ 15
13. Plants large, up to 5 mm wide; underleaves reniform in outline with lateral margins laciniate ..................... 14
— Plants smaller, up to 3.8 mm wide; underleaves ovate to sub-reniform with lateral margins remotely toothed
........................................................................... Heteroscyphus supinopsis [Link], Thouvenot & Frank Müll.
14. Plants fleshy; leaves dorsally free; cells without trigones .....................Heteroscyphus succulentus (Gott.) Schiffn.
— Plants rigid; leaves dorsally connate; cells with nodulose trigones ..... Heteroscyphus deplanchei (Steph.) Schiffn.

4 CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1)

 Lophocoleaceae of New Caledonia

15. Plants medium sized; leaves ovate, entire, up to 1.6 mm long; underleaves bifid with acute triangular lobes and
shortly toothed margins ............................................................. Heteroscyphus kanakensis Thouvenot & Engel
— Plants large; leaves rounded, apices retuse, 1.5-3 mm long; underleaves bifid with lanceolate lobes and laciniate
margins ..................................................................................................Heteroscyphus giganteus (Steph.) Hürl.

16. Leaves rounded, bifid, lobes short, narrow, acute, margins entire to superficially toothed; underleaves wide,
rounded, bifid to 4-fid, margins toothed; cells with large nodulose trigones ................................................ 17
— Leaves either oblong to rectangular with various apices, or rounded to oval with long toothed-ciliate margins;
underleaves various; cells with minute to large trigones ............................................................................... 18

17. Leaves asymmetrically bifid with dorsal margins straight or slightly convex, usually strongly recurved, ventral
margins widely rounded, bifid apices shifted toward the upper dorsal margins; leaf and underleaf lobes usually
acuminate with long linear apices ...........................Heteroscyphus aselliformis (Reinw., Blume & Nees) Schiffn.
— Leaves symmetrically bifid with both margin curvatures similar and sinus set on the top of the rounded apex;
leaf and underleaf lobes triangular acute, if acuminate, then with shorter apices ..............................................
.......................................................................................................Heteroscyphus confertus (Steph.) Thouvenot

18. Leaves rounded to ovate, apices rounded or widely triangular with three long ciliate processes, triangular at base;
margins usually with few to numerous cilia restricted to subapical and ventral parts; underleaves transversely
elongate, 2-3 times as wide as the stem, bifid, lobes like the leaf appendages single or furcate, each lateral mar-
gins variously toothed to lobate ciliate ................................................Heteroscyphus grandiflorus (Steph.) Hürl.
— Leaves elongate, oval-oblong to ovate trapezoid or rectangular, apices various, if rounded then with small teeth,
if triangular then teeth spinose, not ciliate; margins entire except near apices or with a tooth more on ventral
margins; underleaves various ....................................................................................................................... 19

19. Shoots canaliculate with the leaves dorsally assurgent, the abaxial surface of opposite leaves facing one another;
leaves ovate trapezoid; leaf cell walls strongly thickened, with one thin pore on each side of the cells ...............
.................................................................. Heteroscyphus assurgentissimus [Link], Thouvenot & Frank Müll.
— Shoots complanate with the leaves spreading out, the abaxial surfaces flat or convex, not facing to the opposite
ones; leaves oblong to lingulate; leaf cell walls thin, with or without trigones .............................................. 20

20. Underleaves transversally elongate in outline, bifid with triangular lobes acuminate and widely divergent, one
or two small teeth on both lateral margins ................................................................................................... 21
— Underleaves either reniform with 4-6 subequal teeth or bifid with margins heavily lobate-laciniate ............. 22

21. Leaf cells without trigones; leaf apices widely rounded to truncate, rarely acute, with 4-6 or more sharp teeth,
roughly equal; underleaf lateral margins with one tooth on both sides .............................................................
.................................................................................... Heteroscyphus argutus (Reinw., Blume & Nees) Schiffn.
— Leaf cells with large nodulose trigones; leaf apices truncate, concave, with a single tooth at both angles; underleaf
lateral margins with two teeth on both sides ....................................................... Chiloscyphus longifissus Steph.

22. Underleaves strongly narrowed below and so narrowly decurrent and connate to the leaves by a band up to five
cells wide; leaf cells with large nodulose trigones; leaves ovate-lingulate with apices variously shaped in a same
shoot, entire rounded, or acute to bifid, or acute with 1-2 subapical teeth so that the leaf apices seem bifid or trifid
................................................................................................................................... Heteroscyphus deplanchei
— Underleaves not narrowed below and so widely decurrent and connate to both adjacent leaves by a band
of cells usually more than six cells wide; leaf cells with minute to inconspicuous trigones; leaves evenly
shaped along the same shoot, leaf apices truncate with a single short to medium tooth at both angles
............................................................................................................. Heteroscyphus coalitus (Hook.) Schiffn.

23. Leaf surfaces appearing rough because of many spinose projections .............Lophocolea muricata (Lehm.) Nees
— Leaf surfaces smooth, rarely with cells at most bulging ................................................................................ 24

24. Plants small, shoots c. 1.5 mm wide; leaf and underleaf margins densely and sharply toothed; leaf cells less than
20 µm wide with relatively large trigones .............................................................. Lophocolea caledonica Steph.
— Plants larger, shoots usually more than 1.5 mm wide, or, if smaller, with entire or emarginate leaves; underleaf
lateral margins with at most two teeth on both sides; leaf cells more than 20 µm wide with trigones small or null
.................................................................................................................................................................... 25

25. Leaves fully entire with rounded apices; plants medium to large .................................................................. 26
— At least some leaves toothed or lobed or emarginate at apices, plants small to medium ................................ 27

CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1) 5

 Thouvenot L.

26. Leaves widely ovate, wider or as wide as long, 0.6-0.7 mm long, dorsally assurgent-convex, giving an inflate
appearance to the shoots; shoots c. 2 mm wide when flattened; underleaves 1/2 bifid, 2-2.5 times the stem width,
lobes erect ....................................................................................................... Chiloscyphus trigonifolius Steph.
— Leaves ovate-oblong, 1.6-2.2 mm long, horizontally spreading, shoots somewhat complanate, 3.5-4 mm wide;
underleaves more than 1/2 bifid, 4-5 times the stem width, lobes widely spreading ...........................................
........................................................................... Heteroscyphus cornutistipulus (Steph.) Thouvenot, comb. nov.
27. Leaf apices widely rounded to truncate, with 4-6 or more sharp teeth, almost equal; cell walls evenly thickened
without trigones; plants medium, shoot width 1.5-2.0 mm ............................................. Heteroscyphus argutus
— Leaf apices various, teeth if any restricted to distal angles; cell walls thin, trigones minute to absent; shoot width
ranging from 1 to 6 mm .............................................................................................................................. 28
28. All leaf apices bifid, lobes triangular acuminate, with piliform apices usually 3-5 cells long .............................
................................................................................................................... Lophocolea bidentata (L.) Dumort.
— Leaf apices various, when bilobate unevenly so in the same shoot and lobe apices not piliform ................... 29
29. Underleaves reniform, lateral margins heavily toothed-laciniate; leaf apices usually shallowly triangular with
spinosely toothed angles .................................................................. Heteroscyphus caledonicus (Steph.) Schiffn.
— Underleaves reniform or not, lateral margins with 0-2 small teeth; leaf apices truncate, rounded or emarginate
.................................................................................................................................................................... 30
30. Leaves rectangular, contiguous to distant, apices transversally concave, with distal angles sharply toothed or
less frequently rounded, a third subapical tooth eventually present on the ventral margins; leaf cells without
trigone; gynoecia and androecia on short latero-ventral leafless branches .........................................................
...................................................................................Heteroscyphus etesseanus (Steph.) Thouvenot, comb. nov.
— Leaves ovate to oblong, more or less strongly imbricate, apices straight or varying from rounded to shallowly
emarginate-bilobate often on the same shoot, angles or lobes rounded to acute; leaf cells with or without trigo-
nes; gynoecia or androecia or both on main shoots or long branches with normal vegetative leaves ............. 31
31. Leaf cells with large nodulose trigones; underleaves very wider than the stems with lobes triangular and widely
divergent ....................................................................................................................... Chiloscyphus longifissus
— Leaf cells with small trigones or trigones lacking; underleaves not or hardly wider than the stems with lobes
narrow, roughly parallel ............................................................................................................................... 32
32. Plants small, shoots 1-2 mm wide; leaves ovate, leaf apices conspicuously narrower that the bases, rounded
to narrowly emarginate, margins crenulate by the bulging marginal cells; underleaf lobes lanceolate, rigid
....................................................................................................................... Lophocolea convexula Sande Lac.
— Plants medium sized, shoots more than 2.5 mm wide; leaves oblong, leaf apices quite as wide as the bases, from
widely convex or emarginate to shortly bifid, margins smooth; underleaf lobes linear, curved ..................... 33
33. Gynoecia and androecia on short latero-ventral leafless branches; perianths cupulate, without keels, bracts strongly
differentiated from normal leaves, bracteole deeply bifid; underleaf insertions concave, discs more than 2 cells
long, lobes with triangular bases ................. Heteroscyphus parapilistipulus (Thouvenot) Thouvenot, comb. nov.
— Gynoecia and androecia at the end of main shoots or long leafy branches, perianths oblong and 3-angled, bracts
slightly differentiated from normal leaves, bracteole oblong, toothed, not deeply bifid; underleaf insertions in
an inverted U, discs very short, two cells long, lobes sublinear ................................Lophocolea savesiana Steph.

TAXONOMIC TREATMENT branches, bracts and bracteoles variously toothed to laciniate,

the latter usually bifid; 7) perianths well developed, oblong-
All New Caledonian species of Lophocoleaceae are described cylindrical to cupulate in outline, from tri-carinate in the
below. Their shared characters include essentially: 1) plants distal part to shallowly plicate, narrowly winged or smooth,
small to large, leaves succubous, obliquely inserted, alternate mouth more or less deeply trifid, lobes possibly bifid, toothed
or subopposite, sometimes dorsally decurrent or fused, never to laciniate; and 8) androecia position either intercalary on
duplicate; 2) leaf apices entire to incised or toothed, margins leading shoots or vegetative branches, bracts smaller than the
smooth or toothed to laciniate; 3) underleaves conspicuous, vegetative leaves, or in a spike at the apices of short almost
usually wider than the stem, connate or not with one or both leafless ventral-lateral branches with smaller inflated bracts,
nearest leaves; 4) cells hexagonal, with usually thin walls, or, in paroecious species, at the base of gynoecia set at the
rarely strongly thickened, trigones inconspicuous to large, end of normal-leaved shoots.
acute or bulging, sometimes confluent; 5) plants usually The list of accepted species of Lophocoleaceae is followed
dioecious, less commonly autoecious, rarely paroecious; 6) by a commented dubious species, whose generic classification
gynoecia terminal either on leading shoots or long branches remains unresolved (incertae sedis), a list of species transferred
with normal leaves, or on short almost leafless ventral-lateral to other families and a list of excluded taxa.

6 CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1)

 Lophocoleaceae of New Caledonia

A C

Fig. 1. — Chiloscyphus trigonifolius Steph.: A, leaf cells; B, leaves and underleaf in ventral view; C, shoot portion in ventral view; D, androecium; E, leaves and
underleaf in dorsal view. A, C, drawn from the lectotype (G00069423); B, D, E, drawn from the isolectotype (PC0167659). Scale bars: A, 20 µm; B-E, 500 µm.

New Caledonian species of Lophocoleaceae Further specimen examined. — New Caledonia. South Province,
in the vicinity of Nouméa, 1909, Le Rat s.n., determinavit Stephani
as “Chiloscyphus cornutistipatus” (PC[PC0167669]).
Genus Chiloscyphus Corda
Distribution in New Caledonia. — Only known from the type
locality, a remote place hard to reach, and one further historic speci-
Chiloscyphus trigonifolius Steph. men without precise location.
(Fig. 1)
Total range. — Endemic.
Species Hepaticarum 6: 316 (Stephani 1922). — Type: New Cal-
edonia. “In summo Mt. Dent de St Vincent, 1425 m”, VII.1909, Description
L. Le Rat 101 (lecto-, here designated, G[G00069423]!; isolecto-, Dioecious.
PC[PC0167659]!, REN[herb. E. G. Paris]!).

Chiloscyphus trigonifolius Steph. ex Paris, Revue bryologique 37: 129 Habit

(Paris 1910), nom. inval. (no description). — Material: REN[herb. Plant medium, moist shoots 1.00-2.00 mm wide, 2.50 mm
E. G. Paris] syn. nov. wide when flattened; leaf bases dorsally assurgent, strongly

CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1) 7

 Thouvenot L.

convex, masking the stem, the leaves distally curving downward, Heteroscyphus cornutistipulus is close to Chiloscyphus
leaves second, imbricate, alternate, dorsally free, separated by trigonifolius, with leaves and underleaves being superficially
a two ranked band of cortical cells. similar. It differs essentially by: 1) shoots complanate; 2)
leaves slightly convex and horizontally spreading; 3) larger
Leaves sizes in all dimensions: shoot width, leaf length and width,
Ovate-triangular to rounded in outline, wider than long, cell diameters, trigones width; and 4) androecia on very
0.80-0.90 mm long, 1.10 mm wide at base, entire margined, short ventral branches. Furthermore, the leaves are ovate
apices rounded, margins convex, slightly crenulate by the oblong, longer than wide, instead of widely ovate, wider
prominent marginal cells. than long in the latter. Out of New Caledonia, Heteroscy-
phus furcistipulus, endemic to New Zealand, is similar to
Cells C. trigonifolius in all vegetative characters (Engel & Glenny
Leaf cells 20-25 µm wide, thin walled with small acute 2019), but the former has very distinctive androecia of
trigones. heteroscyphoid type, with 3-8 pairs of ventricose bracts on
leafless ventral branches that are hidden under the normal
Underleaves leaves, while male bracts of C. trigonifolius are relatively
Small, 2 times the stem width, 0.60 mm long, 0.40-0.60 mm shortly ventricose at the base of large rounded flat lamina
wide, connate to one adjacent leaf, bifid up to mid-length, and are set in spikes at the end of normal leaved branches
sinus lunate, lobes lanceolate, acuminate to obtuse, upward or main shoots, or inserted inside main shoot portions;
directed, disc margins with 1-2 short teeth. androecia of heteroscyphoid type are lacking. Furthermore,
H. furcistipulus is calcicolous since C. trigonifolius was found
Gametangia in ultramafic massifs.
Androecia in series of numerous bracts and antheridia (10 pairs)
inserted between normal-leaved sections of the main stems or
at the end of long leafy branches; male bracts similar in size Genus Cryptolophocolea L.Söderstr.
to vegetative leaves but with inflated dorsal base including
antheridia; gynoecia not seen. Cryptolophocolea explanata (Mitt.) Váňa & Crand.-Stotl.
(Fig. 2)
Comments
The alternate leaves, unilateral connation of underleaves Phytotaxa 202: 69 (Söderström et al. 2015). — Lophocolea explanata
Mitt., Flora Vitiensis: 404 (Mitten 1871 [1873]). — Chiloscyphus
with leaves, little differentiated male bracts and androecia explanatus (Mitt.) [Link] & [Link]., Nova Hedwigia 39:
on elongate shoots separate Chiloscyphus trigonifolius from 414 (Engel & Schuster 1984 [1985]). — Type: Samoa. Powell s.n.
Heteroscyphus while the entire leaves and, again, little differ- (lecto-, selected by Váňa et Crandall-Stotler [in Söderström et al.
entiated male bracts with saccate dorsal base distinguish it 2015], NY[NY00965738], not seen; isolecto-, G[G00121761]!).
from most Lophocolea. Unfortunately, gynoecia are unknown Further specimens examined. — New Caledonia. South Province,
in C. trigonifolius and definitive evidence for its placement Mts. Koghis, in forest, V-VI.1905, Etesse s.n., ex herb. E. G. Paris
in Chiloscyphus vs Lophocolea is therefore lacking. But, since nr 13 (G s.n.).
all known characters match Chiloscyphus, we conservatively
Distribution in New Caledonia. — South Province. Known
keep the species in this latter genus pending further collec- from a single collection in Mts Koghis by Etesse in 1905.
tions of female plants.
In the original publication of Chiloscyphus trigonifolius, Total range. — Pacific Islands, occurs also in Samoa (type locality).
Stephani mentions Franc as collector, but the label of the
original material of C. trigonifolius in G mentions Louise Description (based on the New Caledonian specimen)
Lerat as the collector. In the checklist of the New Caledonian Autoecious.
liverworts, Thouvenot et al. (2011) stated that the identity
of Chiloscyphus trigonifolius Steph ex Paris was unclear since Habit
Paris did not give any description of this taxon. The speci- Plants medium sized, wet shoots up to 2.10 mm wide when
men kept in the herbarium of E. G. Paris at REN have been flattened. Leaves obliquely spreading, convex to nearly explanate.
checked; its label is identical to the one in G; moreover, the
plant exactly matches Stephani’s diagnosis and drawing of Leaves
C. trigonifolius and therefore constitutes a duplicate of the In medium parts of stems, leaves 0.70-0.90 mm long, 0.45-
material used by Stephani to define his new species. A letter 0.65 mm wide at base, 0.20-0.25 at apex, long ovate to
from Stephani (9 Febr. 1910) (Rennes 1 University Library) narrowly trapezoid when flattened, both margins nearly sym-
reports Chiloscyphus trigonifolius as a new species, cited as metrically convex above the base becoming nearly straight to
“nr 101”, among the material collected by Mr. and Mrs. Le slightly concave subapically, apices usually obliquely truncate,
Rat and sent to Stephani by Paris. This confirms that the with a sharp tooth at both angles and rare additional short
type is not from Franc and that the specimen at REN must teeth on ventral margins, dorsal margins entire, decurrent,
be an isolectotype. not connivent.

8 CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1)

 Lophocoleaceae of New Caledonia

B
A

E
F G

H K

J
I

Fig. 2. — Cryptolophocolea explanata (Mitt.) Váňa & Crand.-Stotl.: A, shoot in ventral view; B, D, E, G, leaves; C, leaf apex; F, cells; H, female bract; I, female
bracteole; J-L, underleaves. All drawn from the specimen Etesse s.n. at G (s.n.). Scale bars: A-E, G-K, 500 µm; L, 200 µm; F, 50 µm.

Cells margins convex, recurved near base, with one tooth on both
Leaf cells hexagonal, 24-36 µm wide, 24-50 µm long, thin sides, teeth shorter or like apical lobes.
walled, trigones small, acute.
Gametangia
Underleaves Gynoecia terminal on normal-leaved branches or main shoots,
Hardly wider than the stem, 0.30-0.40 mm, usually as wide bracts ovate lanceolate, asymmetric, 2.20 mm long, 0.90 mm
as long, narrowly connate to the nearest leaves on both sides, wide, canaliculate, margins laciniate, bracteoles oblong,
rounded-quadrate to slightly obovate, bifid up to ¼ down 1.60 mm long, 1.20 mm wide, margins laciniate; perianths
with rather short triangular acuminate lobes and lunate sinus, subcylindrical, 3-carinate, keels broadly winged, ciliate along

CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1) 9

 Thouvenot L.

the whole length, mouth densely laciniate; androecia in long Underleaves

spikes with more than 15 pairs of bracts, intercalary or terminal Large, widely connate to the leaves on both sides, rounded,
on normal leaved branches, bracts 1.00 mm long, 0.50 mm short bifid with connivent lobes and lunate sinus, margins
wide, smaller than the normal leaves and saccate at dorsal sharply toothed with many long teeth like the apical lobes,
base, antheridia stalk biseriate. disc 0.40-0.60 mm wide.

Comments Gametangia
Lophocolea explanata was reported from New Caledonia by Paris Gynoecia terminal on normal leaved branches, bracts much
(1906) based on a specimen collected by Etesse. No voucher tongue like, canaliculate, margins unevenly ciliate-toothed,
has been found in the herbarium of E. G. Paris at REN, but bracteoles shorter, margins ciliate-toothed; perianths subcy-
a duplicate is kept in G (see also Stephani’s correspondence lindrical, 3-carinate, keels broadly winged, ciliate along the
to Paris, [Link].1905 [Rennes 1 University Library]) match- whole length, mouth shortly trilobate, lobes oval, laciniate;
ing the isolectotype of L. explanata in G and confirms the androecia in long spikes, intercalary to subterminal on leaved
presence of this species in New Caledonia. Stephani (1906) branches, bracts like the vegetative leaves but smaller and
described the species as being dioecious, likely owing to the dorsally saccate at base.
absence of male shoots in the type, but the New Caledonian
specimen is autoecious. Comments
As mentioned by Thouvenot et al. (2018), this species is
separated from the two additional Cryptolophocolea species
Cryptolophocolea subcostata (Steph.) Thouvenot reported from New Caledonia by leaves strongly canaliculate
(Fig. 3) (separating it from C. explanata (Mitt.) Váňa & Crand.-
Stotl.) and autoecious sexuality and short oval perianth
Cryptogamie, Bryologie 39 (3): 364 (Thouvenot et al. 2018). —
Lophocolea subcostata Steph., Species Hepaticarum 6: 295 (Stephani lobes (separating it from C. levieri (Schiffn.) L. Söderstr.).
1922). — Chiloscyphus subcostatus (Steph.) [Link] & [Link]., From the closely related C. costata (Nees) L.Söderstr. it is
Nova Hedwigia 39: 423 (Engel & Schuster 1984 [1985]). — Type: separated mainly by the subtruncate to shortly trilobed peri-
New Caledonia. Le Rat s.n. (lecto-, here designated, G[“In jugo anth mouths, smaller size with leaves less than 2 mm long
Dogny, 1050 m”, VII.1909, L. Le Rat 424, G00112445]!; isolecto-, (vs 2-3 mm), rounded underleaves and dorsal leaf margins
REN[herb. E. G. Paris s.n.]).
entire or with a few small teeth. As molecular evidence for
Further specimens examined. — New Caledonia. South Prov- a possible synonymy with the latter is lacking, C. subcostata
ince, ridge between the path to Dzumac and Mt. Ouin, 1100 m, is kept as a separate species.
17.V.1951, Hürlimann 2599 (PC[PC167677]); Païta, Humboldt
massif, 1205 m, cloud forest, [Link].2008, Thouvenot NC1799;
North Province, Hienghène, Panié massif, between Bwa Téan and
Payolé, 1000 m, 9.X.2012, Thouvenot NC1879. Genus Heteroscyphus Schiffn.
Distribution in New Caledonia. — Scattered in North and
South Provinces, in cloud forest above 1000 m. Heteroscyphus argutus (Reinw., Blume & Nees) Schiffn.
(Fig. 4)
Total range. — Endemic.
Oesterreichische Botanische Zeitschrift 60: 172 (Schiffner 1910). —
Description Jungermannia arguta Reinw., Blume & Nees, Nova Acta Physico-
Autoecious. medica Academiae Caesareae Leopoldino-Carolinae Naturae Curiosorum
Exhibentia Ephemerides sive Observationes Historias et Experimenta
12: 206 (Reinwardt et al. 1824). — Chiloscyphus argutus (Reinw.,
Habit Blume & Nees) Nees, Synopsis Hepaticarum: 183 (Gottsche et al.
Plants medium sized, moist shoots 3.00 mm wide. 1845). — Type: Java. “In montibus Sadjra […] Iavae insulae, prope
Rawayan…”, Blume s.n. (G[G00280112]).
Leaves
Chiloscyphus acutus Steph., Species Hepaticarum 6: 302 (Stephani
In medium parts of stems, leaves 0.90-1.00 mm long, 1922). — Type: New Caledonia. “Lerat” s.n. (lecto-, here desig-
0.60-0.70 mm wide at bases, 0.30-0.40 mm at apices, nated, G[“Île des Pins, Forêt de Condo”, V.1909, L. Le Rat 44,
obliquely spreading, dorsally convex, canaliculate, when G00069505]!; isolecto-, REN[herb. E. G. Paris]!; syn-, G[“Île des
flat long ovate, asymmetrical, ventral bases widened, dorsal Pins, Grotte d’Aoupéna”, V.1909, L. Le Rat 72, G00282946]!)
margins nearly straight, apices acute, truncate or bilobate, syn. nov.
with additional irregular sharp teeth at apices and ventral Further specimens examined. — New Caledonia. South Province,
margins, dorsal margins entire or with a few smaller teeth, Farino, Grandes Fougères Nature Park, Houé river, 430 m, [Link].2008,
decurrent. Thouvenot NC435; 370 m, [Link].2016, Thouvenot NC2450; La Foa,
Mts. Koghis, 500 m, IV.2013, Coulerie COU143; Mt. Dogny, 540 m,
9.V.2015, Metoyer MET061; Mt. Dzumac, IV.1907, Le Rat s.n. as
Cells “Chiloscyphus jackii” (REN[herb. E. G. Paris]); Mts. Koghis, on bark
Leaf cells round, 25-35 µm wide, trigones large, basal cells and rocks near stream, 1000 ft, forest, [Link].1914, Compton 801 as
larger. Chiloscyphus argutus (BM[BM013409505]).

10 CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1)

 Lophocoleaceae of New Caledonia

A B

D
E

Fig. 3. — Cryptolophocolea subcostata (Steph.) Thouvenot: A, B, E, leaves and underleaves, ventral view; C, G, leaves; D, perianth; F, cells; H, female bracteole;
I, androecium. All illustrated from Hürlimann 2599a (PC0167677). Scale bars: F, 20 µm; A-E, G, H, 500 µm; I, 1 mm.

Distribution in New Caledonia. — Frequent in North and South Leaves

Provinces, including Isle of Pines, in moist conditions on ground,
dead wood, trees or tree ferns, collected between 300-1200 m. Ovate-oblong, 1.20-1.40 mm long, 1.00-1.20 mm wide at
bases, 0.60-0.80 mm below apices, apices widely rounded,
Total range. — Southern Asia, Indonesia, Oceania, Australasia,
Africa. with usually four linear teeth, up to eight uniseriate cells
long, the upper ones acute to narrowly rounded, sometimes
description
with one or two additional teeth, lateral margins naked but
Based on the lectotype of Chiloscyphus acutus. Further descrip- usually with one subapical tooth on both dorsal and ventral
tions and illustrations in Piippo (1985). sides and the ventral often with a tooth more.
Dioecious.
Cells
Habit
Plants medium, wet shoots 1.50-2.50 mm wide; leaves alternate Leaf cells hexagonal, their size progressively increasing up to
to subopposite, dorsally obliquely assurgent in a wide angle down, (20-)25-35 µm wide, cell walls evenly a little thick,
c. 130°, spreading at a right angle when flattened. without trigones.

CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1) 11

 Thouvenot L.

A B C

Fig. 4. — Heteroscyphus argutus (Reinw., Blume & Nees) Schiffn.: A, portion of shoot in ventral view; B, leaf apices; C, cells; D, underleaf. All drawn from the
lectotype of Chiloscyphus acutus Steph. (G00069505). Scale bars: A, B, 500 µm; C, 20 µm; D, 200 µm.

Underleaves Heteroscyphus aselliformis

Free on both sides or unilaterally narrowly connate to the (Reinw., Blume & Nees) Schiffn.
nearest leaf, overall dimensions 0.40-0.50 mm long, 0.80- (Fig. 5)
1.00 mm wide, disc transversally elongate, narrow, 0.10-
0.15 mm wide, 0.40 mm long, sinus wide, flat or shallowly Oesterreichische Botanische Zeitschrift 60: 172 (Schiffner 1910). —
Jungermannia aselliformis Reinw., Blume & Nees, Nova Acta Physico-
convex, bifid with lobes widely spreading, curved frontward, medica Academiae Caesareae Leopoldino-Carolinae Naturae Curiosorum
horn like, narrowly triangular at base, evenly acuminate, end- Exhibentia Ephemerides sive Observationes Historias et Experimenta
ing in linear points, lateral margins short, each one with a 12: 412 (Reinwardt et al. 1824). — Chiloscyphus aselliformis (Reinw.,
linear segment backward turned. Blume & Nees) Nees, Synopsis Hepaticarum: 176 (Gottsche et al.
1845). — Type: Java. Blume s.n., not seen.
Gametangia Chiloscyphus theriotii Steph., Species Hepaticarum 6: 316 (Stephani
At the end of very abbreviated leafless branches; gynoecia 1922). — Type: New Caledonia. Franc s.n. (lecto-, here des-
with bracts and bracteoles unevenly toothed, perianths ignated, G[“Mts. Koghis, 1/11/1909”, Franc s.n., hb. The-
cyathiform, deeply 3-lobate, lobes spinose-toothed; androe- riot 169, G00069428]!; isolecto-, PC[as Chiloscyphus aselliformis,
PC0767992]!).
cia in long narrow spikes made of 3-9 pairs of bracts,
bracts deeply bifid with narrow segments ending in 4-6 Further specimens examined. — New Caledonia. North Province,
uniseriate cells. Pouébo, Diahoué, Tooliwök pass, on bark in rain forest, 700 m,
[Link].2019, Thouvenot NC2827; Mandjélia, on a tree fern base, 701 m,
[Link].2019, Thouvenot NC2896 (PC[PC0779855]); South Province,
Comments Mts. Koghis, Mt. Bouo ridge, 700 m, [Link].1951, Guillaumin et Bau-
Based on Stephani’s diagnoses and drawings, Chiloscyphus mann-Bodenheim 12670 (PC[PC0767993]); Mts. Koghis, Mt. Moné,
acutus is separated from C. argutus by the leaf apices, which south side, 750 m, [Link].1951, Hürlimann 2779 (PC[PC0767988]);
are acute instead of truncate. In the type specimens, how- Nouméa, along the pathway from the Koghis lodge towards Mt. Bouo,
ever, the leaf apices are truncate to convex, rarely acute, with epiphyte, c. 850 m, F. Müller NC277; Yaté, Rivière Bleue Provincial
Park, Pourina trail, 480 m, on bark in an open wet forest, [Link].2016,
4-6 short teeth like Heteroscyphus argutus. All other features Thouvenot NC2482; Mts. Koghis, [Link].1909, Franc s.n., det. Stephani
are identical. as “Chiloscyphus hebridensis” (G, PC[PC0767992]).
Both specimens of Chiloscyphus acutus in G were col- Papua New Guinea. Morobe Prov., Cromwell Mts., c. 2250 m,
lected by Mrs Louise Le Rat on Île des Pins and sent by [Link].1981, Koponen 31350, det. S. Piippo (PC[PC0767989]).
E. G. Paris to Stephani for identification. One of them, Java. Res. Pasoeroean, 1200 m, [Link].1930, Verdoorn 19, det. W.E. Ni-
cholson (PC[PC0767990]).
collected in Condo Forest, was annotated “[Link].”, “original”
and “cum flore sterili” by Stephani, and can therefore be Distribution in New Caledonia. — Frequent in North and South
selected as the lectotype. The second, collected in Aoupéna Provinces on tree trunks and fern tree rhizomes in rainforests and meso-
cove, was first identified by Paris as Plagiochila; Stephani philous forests at medium elevations (collected between 300-750 m).
subsequently added the name Chiloscyphus acutus. As it Total range. — Southeast Asia, Indonesia, Melanesia.
was collected in a nearby locality at the same date by the
same collector, it may reasonably be regarded as a syntype Description
of C. acutus. Dioecious.

12 CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1)

 Lophocoleaceae of New Caledonia

B
A C

F
E

H
I
G

Fig. 5. — Heteroscyphus aselliformis (Reinw., Blume & Nees) Schiffn.: A, leaf and underleaf; B, E, leaves; C, male bract with antheridium; D, G, H, underleaves;
F, perianth; I, cells. All except F (from Guillaumin et Baumann-Bodenheim 12670 [PC0767993]) drawn from the following types of Chiloscyphus theriotii Steph.:
A-D, H, from PC0767992; E, G, I, from G00069428. Scale bars: A, B, D-H, 1 mm; C, 500 µm; I, 50 µm.

Habit apical lobe, both lateral margins entire or the ventral sparsely
Plants medium to large, soft textured, 4-5 mm wide when toothed, leaf variations involve smaller size and symmetry,
flattened; leaves dorsally assurgent, concave, widely imbricate, then the overall shape is oval, and margin ornamentation
subopposite, dorsal leaf margins confluent, lateral-intercalary varying from 0 to +/- 5 small teeth, one celled, or with a few
branches sparse, flagelliform branches occasional to frequent. piliform processes.

Leaves Cells
Leaf shapes variable with normal leaves typically 1.25-2.00 mm 30-50 µm, cell walls thin with large trigones at most contiguous.
long, 1.25-2.50 mm wide, asymmetrically quadrate-rounded,
ventral margins rounded, dorsal margins nearly straight, apices Underleaves
asymmetrical, shortly bifid, sinus lunate, shifted toward the Large, 0.80-1.70 mm wide, rounded to slightly reniform, shortly
dorsal margin, two short triangular lobes erect to connivent, bifid, sinus lunate, lobes erect, narrowly triangular acuminate,
acute to acuminate, ending in long piliform apices, fragile, apices piliform, lateral margins with many small teeth, some-
often broken and absent, sometimes with one additional sub- times with 1-2 subapical additional long teeth, lobe-like.

CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1) 13

 Thouvenot L.

Gametangia Distribution in New Caledonia. — Only known from the type

At the end of short leafless branches, lateral-intercalary; locality in South Province.
gynoecia with two innermost pairs of bracts and bracteoles, Total range. — Endemic.
free, margins fimbriate all around, bracteoles 2.0 mm long,
1.30 mm wide, deeply bifid, bracts 2.00 mm long, 2.50 wide, description
3(-4)-fid, perianth cyathiform to subcylindrical, 2.00 mm Further description and illustrations in Engel et al. (2021).
long, split to mid length in three lobes toothed-fimbriate; Dioecious.
androecia in short spikes of one or two pairs of bracts, bracts
cochleariform, margins minutely toothed with two sharper Habit
teeth at apices, antheridia stalks bi-seriate. Plants rigid, up to 5 mm wide when flattened, the shoot
tips sharply dorsally assurgent; ventral-intercalary branching
Comments frequent; stems narrow for plant size, 0.20-0.30 mm wide;
Heteroscyphus aselliformis is a very variable species (Stephani leaves subopposite, stiffly dorsally assurgent, often forming a
1907) and has been described under several names, depend- gentle bow-like arch, the adaxial face of opposing leaves fac-
ing on their geographic origin, i.e., as Chiloscyphus theriotii in ing one another and forming a deep narrow channel; leaves
New Caledonia. Some of the specimens from New Caledonia, closely imbricate, dorsally connate and forming an elevated,
including the type material of C. theriotii, are rather stable laminar lip to c. 4 cells wide; leaves moderately convex in
with a large size, leaves 3.25 mm long and 2.5 wide, oblong- dorsal sector, moderately abaxially concave in ventral sector.
rounded, asymmetrically bifid with apices shifted toward the
dorsal margins which is loosely recurved (vs tightly so, or Leaves
duplicate), ventral leaf margins eventually with a short tooth 1.50-2.50 mm long, 0.90-1.30 mm wide at widest point, elon-
beneath the apex, underleaves connate to both nearest leaves gate ovate-trapezoid to lingulate, distinctly narrowed toward
by a medium to narrow band, wide rounded, apices bifid, the apex, moderately to distinctly dilated in ventral-basal
lobes narrowly triangular acute to acuminate, margins evenly sector; apex bifid to 0.05-0.10, the lobes divergent, occasion-
and sharply denticulate all around. Other New Caledonian ally parallel, the ventral lobe often larger than the dorsal, the
collections, however, and specimens from other countries, dorsal lobe at times reduced to a small tooth or lacking, the
are more variable. Specimens from Papua New Guinea, for lobes narrowly acute to acuminate, the sinus mostly lunate,
example, are smaller and with entire underleaf margins while occasionally narrowly rounded; lamina margins entire.
those from Java seem to be intermediate in size and with
crenulate underleaf margins. Cells
In the field, H. aselliiformis resembles H. confertus and 30-47 µm wide, 37-57 µm long, somewhat larger near leaf
H. giganteus by its pale colour, light green to brownish, large bases, median leaf cells with strong secondary thickenings of
size and leaves dorsally assurgent, somewhat cochleariform. the radial walls, each thickening interrupted by a single, large,
It differs from those two species by its longer and sharper clearly defined pit, the wall thickenings converging at the cell
leaf lobes, and the presence of additional sharp teeth on junctures to form massive nodose trigones.
the leaf margins. Furthermore, the rounded to reniform
underleaves, which are bifid with narrowly triangular erect Underleaves
lobes and lateral margins copiously denticulate, differ from 2.3-3.5 times the stem width when flattened, connate with
those of H. giganteus which has laciniate underleaf margins, the leaves on both sides, connate portion 3-6 cells wide,
and from those of H. confertus which has widely triangular imbricate, stoutly ovate to subreniform, 0.80-1.30 mm long,
underleaf lobes. In addition, the latter species is smaller. 0.80-1.10 mm wide; apex 2-4-lobed, the median pair of lobes
The species also resembles H. hebridensis and some New often larger, the lobes ± parallel, acute-apiculate to acuminate,
Caledonian specimens of H. aselliformis collected by Franc entire or with a tooth toward the outer margin, lamina margin
(G, PC) were misidentified as H. hebridensis by Stephani. on each side with 1-5 small teeth.
Heteroscyhus hebridensis, however, has rather widely triangular
ovate leaves, more strongly laciniate underleaves and small Gametangia
to inconspicuous trigones. Gynoecia not seen; androecia small for plant size, hidden by
leaves in dorsal aspect, on abbreviated, determinate, ventral-
intercalary branches, the androecia foliose, with 2-5 pairs of
Heteroscyphus assurgentissimus bracts, the bracts strongly dorsally assurgent, suberect, similar
[Link], Thouvenot & Frank Müll. to leaves in shape and form except the bracts are more concave
(Fig. 6) and the apex is notably more narrowed, the apex bidented,
the base concave ventricose, the dorsal pocket more strongly
Nova Hedwigia 113: 67 (Engel et al. 2021). — Type: New Cal-
edonia. South Province, Parc Provincial de la Rivière Bleue, c. 200- inflated, with a several-celled tooth distally; the antheridia
380 m, lowland rain forest, on a rockface, [Link].2001, Müller NC327 stalk biseriate; bracteole stoutly ovale, bifid to c. 0.5, the
(holo-, DR; iso-, F, hb. Thouvenot). lamina margins each with a small tooth.

14 CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1)

 Lophocoleaceae of New Caledonia

A B E

G I
F

K L

Fig. 6. — Heteroscyphus assurgentissimus [Link], Thouvenot & Frank Müll.: A-D, leaves; E, shoot habit in dry condition, lateral view; F-H, K, L, underleaves;
I, shoot portion, ventral view; J, leaf cells. All illustrated from the holotype. Scale bars: A-D, F-H, K, L, 500 µm; E, I, 1 mm; J, 20 µm.

Comments narrowly pitted. It could be regarded as close to Cryptolophoc-

Heteroscyphus assurgentissimus clearly stands out by: 1) dorsally olea explanata by the leaf and underleaf shapes, but the latter
assurgent leaves and shoot apices; 2) strictly ventral-intercalary species can be easily separated from H. assurgentissimus by
branching; 3) ovate trapezoid to lingulate, bifid leaves; and 4) its complanate habit, thin-walled cells without trigones and
massive trigones extending into strongly thickened cell walls gametangia on elongate shoots.

CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1) 15

 Thouvenot L.

Heteroscyphus caledonicus (Steph.) Schiffn. Comments

(Fig. 7) Heteroscyphus caledonicus can be distinguished from the vari-
able H. grandiflorus by: 1) leaves mainly oblong-rectangular
Oesterreichische Botanische Zeitschrift 60: 172 (Schiffner 1910). — instead of ovate; 2) leaves contiguous or narrowly overlapping
Chiloscyphus caledonicus Steph., Species Hepaticarum 3: 216 instead of imbricate; 3) apical teeth spinose, shorter, not end-
(Stephani 1907). — Type: New Caledonia. Deplanche s.n. ing in piliform tips; and 4) cells with small trigones, if any.
(holo-, G[G00069500]!). The type in G is sterile and scanty, but fresh material col-
Further specimens examined. — New Caledonia. South Province,
lected by the author contained gynoecia, which are described
Mé Areimbo, X.1909, L. Le Rat 341 (G); Mts. Koghis, in forest, here for the first time and are similar to those of Heteroscyphus
300 m, [Link].1909, Franc s.n. (G); North Province, Touho, Mas- and Chiloscyphus. The species is provisionally maintained
sif des Lèvres, Tipelei upper valley, on sedimentary rocks, 315 m, in Heteroscyphus pending the discovery of androecia. Like
12.X.2012, Thouvenot NC2426 (PC[PC0712113]). H. etesseanus and H. cornutistipulus (see below) as well as
Distribution in New Caledonia. — Reported from South further Heteroscyphus species in Australasia (Engel 2015),
Province according to the labels of historic collections, but recently H. caledonicus lacks typical vegetative characters of Het-
found only in North Province, in remote places of the north-eastern eroscyphus such as connate underleaves, large trigones and
coastal massifs, at low elevations. opposite leaves.
Total range. — Endemic.

Description Heteroscyphus coalitus (Hook.) Schiffn.

Dioecious. (Fig. 8)

Habit Oesterreichische Botanische Zeitschrift 60: 172 (Schiffner 1910). —

Plants fragile, minute to medium sized, very variable in width, Chiloscyphus coalitus (Hook.) Dumort., Recueil d’Observations sur les
usually up to 3.00 mm, but many shoots less than 1 mm Jungermanniacées 1: 19 (Dumortier 1835). — Jungermannia coalita
Hook., Musci Exotici 2: tab. 123 (Hooker 1820). — Type: New
wide; leaves subopposite, horizontally spreading at right angle, Zealand. Dusky Bay, A. Menzies s.n. (G[G00283086]!).
slightly imbricate to contiguous, dorsally free, the homologous
leaves separated by a strip of 3-4 epidermal cells on the dorsal Chiloscyphus confertifolius Steph., Species Hepaticarum 6: 304 (Stephani
midline of the stem; stems 0.15-0.20 mm wide. 1922). — Type: New Caledonia. Lerat s.n. (lecto-, here desig-
nated, G[“Nov. Caledon. Inter Farino et Table Unio. 07/1909”,
Le Rat 95, G00069493]!) syn. nov.
Leaves
1.10-1.40 mm long, 0.85-1.20 mm wide, rectangular to Lophocolea latistipula Steph., Species Hepaticarum 6: 281 (Stephani
oblong, apices truncate to widely triangular, with a spinose 1922). — Chiloscyphus latistipulus (Steph.) [Link] & [Link].,
Nova Hedwigia 39: 418 (Engel & Schuster 1984 [1985]). — Type:
tooth on each three angles, teeth at most 3 cells long, dorsal New Caledonia. Franc s.n. (lecto-, here designated, G[“Lophocolea
and ventral margins entire. latistipa (sic) [Link]., Nlle Calédonie, forêt de Tao, 600 à 800 m”,
I.1910, Franc s.n., G00121768]!) syn. nov.
Cells
Chiloscyphus latistipus Steph., Species Hepaticarum 6: 309 (Stephani
Leaf cells hexagonal, 60-70 µm long, 40-50 µm wide, longer 1922). — Type: New Caledonia. Lerat s.n. (lecto-, here desig-
near bases, without trigones or trigones small and acute. nated, G[“Nova Caledonia, Me Areimbo. s.d., Dna L. Le Rat 171,
mit Balantiopsis, mit Lepidozia”, G00069461]!; isolecto-,
Underleaves REN[herb. E. G. Paris, s.n.]!) syn. nov.
Inserted in an inverted U, narrowly decurrent on one or both Chiloscyphus similis Steph., Revue bryologique 35: 28 (Paris 1908),
sides, sometimes narrowly connate to the ventral bases of the non Chiloscyphus similis Steph., Kongliga Svenska Vetenskaps
nearest leaves, otherwise free and set away from the correspond- Academiens Handlingar, Ny Följd 46: 56 (Stephani 1911). —
ing leaves to the middle of the next ones; underleaves wider Chiloscyphus subsimilis Steph., Species Hepaticarum 6: 314 (Stephani
than long, 0.30-0.40 mm long, 0.40-0.60 mm wide, usually 1922), nom. illeg. — Type: New Caledonia. Lerat s.n. (lecto-,
here designated, G[“Chiloscyphus subsimilis. Nov. Caledon. Sine
1-2 times the stem width, bifid rarely trifid, discs narrowly schedule”, 1907, Le Rat 195, G00069431]!) syn. nov.
transverse, only two cells high at sinus insertion, lobes linear,
above 2-celled bases or not, erect, sinus widely lunate or flat, Chiloscyphus francanus Steph., Species Hepaticarum 6: 306 (Stephani
lateral margins with one to three linear segments, lobes and 1922). — Type: New Caledonia. Franc s.n. (lecto-, here des-
segments fragile, often broken, sometimes furcate. ignated, G[“Mt. Dzumac, 900 m, tronc d’arbre”, [Link].1907,
Franc s.n., G00283066]!; syn-, G[“Nlle Calédonie, Mts. Koghis,
versant ouest, 300 m, bords d’un torrent”, 20.X.1907, Franc s.n.,
Gametangia G00069483, G0069484! s.l., s.d., Franc s.n., G00283067]!;
Gynoecia at the end of short leafless lateral-intercalary branches, isolecto-, PC[PC0101950]!; isosyn-, PC[PC101948, PC0101949]!)
bracts asymmetrically bifid, laciniate-toothed, bracteoles sym- syn. nov.
metrically bifid up to mid length, margins toothed; perianths Further specimens examined. — New Caledonia. South Prov-
cyathiform, deeply trilobed, lobes ovate with laciniate-toothed ince, Sarraméa, Dogny plateau, on ground in rain forest, 650 m,
margins; androecia not seen. 24.X.2019, Thouvenot NC2602 (PC[PC0779853]); Yaté, Dzumac

16 CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1)

 Lophocoleaceae of New Caledonia

A B

G
H

Fig. 7. — Heteroscyphus caledonicus (Steph.) Schiffn.: A, shoot portion, ventral view; B, two adjacent leaves, dorsal view; C, leaf; D, E, underleaves; F, cells
from upper third of leaf; G, cells from median part; H, cells from lower part. A, D-G, drawn from Thouvenot NC2426; B, C, H, from the lectotype of Chiloscyphus
caledonicus Steph. (G00069500). Scale bars: A-C, 500 µm; D, E, 100 µm; F-H, 50 µm.

massif, on ground in rain forest, 900 m, 26.X.2012, Thouvenot Distribution in New Caledonia. — Frequent in rain forest
NC2392; North Province, Poindimié, Amoa valley, Tipwadabwé, on and creek banks in both provinces of Grande Terre, collected from
damp rock in a creek, 163 m, 13.X.2019, Thouvenot NC2726; South 150 to 1150 m, on soils or rocks, occasionally at the base of trees.
Province, Mé Amméri, 700 m, [Link].1950, Guillaumin & Baum.
Bod. 9150, det. Hürlimann, as Chiloscyphus francanus (G[G045946]); Total range. — South-East Asia, Indonesia, Melanesia, Australa-
sia, (Argentina?).
“Mts. Koghis, forêt”, Franc s.n. as Chiloscyphus similis (sensu 1908)
(PC[PC0167662], G s.n.); “in jugo Dogny 1040 m”, VII.1909,
L. Le Rat s.n. as Chiloscyphus latistipus (G s.n.); “in jugo Dogny”,
Description
X.1909, L. Le Rat s.n. as Chiloscyphus latistipus (PC[PC0101941,
PC0150605]); “Mts. Koghis”, 1.X.1909, Franc s.n. as Chiloscyphus Dioecious.
latistipus (G s.n.).
Chiloscyphus similis (sensu Stephani 1911): Chili. Fuegia, Scotts- Habit
berg s.n. (G[G00069415]). Plants large with shoots 2.50-4.00 mm wide.

CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1) 17

 Thouvenot L.

Leaves The name Chiloscyphus similis was given to two different

Spreading at nearly right angle, longitudinally convex, oval- type specimens a few years apart by Stephani. In 1908, a New
oblong to subrectangular, 1.30-1.60 mm long, 1.00-1.60 mm Caledonian specimen collected by Le Rat was validly pub-
wide at bases, 0.50-0.80 mm at apices, usually as wide as lished under this name in Revue bryologique and the original
long at base, not dorsally confluent, margins entire, truncate materiel can be identified with the number 195 according to
apices straight to slightly convex, with a single tooth at both a letter to E. G. Paris dated 2 January 1908 (Rennes 1 Uni-
angles, teeth acuminate to linear, acute, 3-6 uniseriate cells versity Library). But, in 1911, Stephani described another
long above and 0-2 biseriate ranks at base. C. similis based on a voucher collected by Scottsberg from
Tierra del Fuego. The drawing and protologue show a very
Cells different plant, confirmed by the examination of the cor-
Median leaf cells hexagonal, somewhat elongate, 30-60 µm responding specimen from Tierra del Fuego kept at G. No
long, 30-75 µm wide, trigones acute, small to inconspicuous. New Caledonian specimen labelled C. similis and match-
ing the diagnosis published in 1908 has been found at G,
Underleaves REN or PC. On another hand, in 1922, Stephani published
Reniform, 2-3 times the stem width, 0.20-0.50 mm long, Chiloscyphus subsimilis based on a type specimen kept at G
0.70-1.20 mm wide, inserted on the stem in a deep sinus and marked with the same number (195) than C. similis
and connate to both adjacent leaves by a very wide band of (1908), according to the correspondence from Stephani to
cells, the apical margin widely convex with 4-6 straight teeth, E. G. Paris. Consequently, C. subsimilis is an illegitimate
sublinear acute, the innermost erect, the lateral spreading, name, superfluous against C. similis (1908) (International
like the leaf ones. Code of Nomenclature, art. 52.1, Turland et al. 2018; Loiseau
et al. 2019), which was validly published with an identified
Gametangia type, while C. similis (1911) is invalid as a later homonym.
Gynoecia at the end of short leafless ventral lateral branches, Fortunately, C. similis (1911) from Tierra de Fuego is fur-
bracts small, rounded, 2-toothed, perianths 1 mm long, cupu- thermore synonym of Heteroscyphus valdiviensis (Mont.)
late with trilobate mouth, lobes rounded, toothed, surface Schiffner (Fulford 1976).
mammillose; androecia not seen. A lot of duplicates of the original material used by Stephani
for Chiloscyphus francanus were examined, raising the oppor-
Comments tunity to recognise a set of syntypes. But a further specimen
Heteroscyphis coalitus is morphologically highly variable and kept at PC as an isosyntype (PC0101947), cannot be a type
a large number of forms and varieties have been described since it was collected one year later than the specimens kept
in this species (Tropicos 2021). According to Piippo (1985), at G and lacks annotation by Stephani.
the species especially varies in the shape of the underleaves
and their connation to the leaves. As it is a species widely
distributed, it is not surprising that there are so many names Heteroscyphus confertus (Steph.) Thouvenot
for this taxon. (Fig. 9)
The original materiel of Chiloscyphus confertifolius at G is
limited to a single packet handwritten by Stephani which is Cryptogamie, Bryologie 39 (3): 365 (Thouvenot et al. 2018). —
therefore selected as the lectotype. Regarding Lophocolea latis- Chiloscyphus confertus Steph., Species Hepaticarum 6: 305 (Stephani
tipula, two specimens are kept as types at G, both collected by 1922). — Type: New Caledonia. Franc s.n. (lecto-, here designated,
Franc in the same locality. The specimen with the mark “[Link].” is G[“Mts. Koghis, 600 m”, [Link].1909, Franc 210, G00069492]!;
syn-, PC[“Mts. Koghis, 300 m”, 1911, Franc s.n., PC0101951]!).
selected despite the orthographic error “latistipa” which may not
lead to confusion, since the epithet latistipus is in fact assigned Heteroscyphus rotundiphyllus ([Link].) Thouvenot, Cryptogamie,
to Chiloscyphus latistipus with a type collected by Mrs L. Le Bryologie 39 (3): 365 (Thouvenot et al. 2018). — Chiloscyphus
Rat. In addition, the drawing by Stephani matches with this rotundifolius Steph., Species Hepaticarum 6: 313 (Stephani
1922), nom. illeg. — Chiloscyphus rotundiphyllus [Link].,
specimen. In contrast, a second specimen labelled Lophocolea Phytologia 47: 321 (Miller 1981). — Type: New Caledo-
latistipula in G (G00112472, duplicate PC0102424) turned nia. “Lerat” s.n. (lecto-, here designated, G[New Caledonia,
to be Chiloscyphus longifissus Steph. (see below). “Dent de St Vincent, VII 1909”, Le Rat 189 (herb. Thériot),
Among the specimens marked as types of Chiloscyphus latisti- G00069432]!) syn. nov.
pus at G, the specimen G00069461 is selected as lectotype since
F urther specimens examined . — New Caledonia. North
it contains the collecting number (171) and the annotation Province, Hienghène, Ouaième Rocks, on twigs in mountain
“[Link].” in a letter from Stephani to E. G. Paris dated 31 March scrubland, on metamorphic bed rock, 982 m, [Link].2019, Thou-
1910 (Rennes1 University Library). Furthermore, “original” is venot NC3285; South Province, “near Nouméa”, s.d., Le Rat s.n.
written on the label in the author’s handwriting. In contrast, (PC[PC0167665]); Yaté, Rivière Bleue natural Park, Pourina
the specimens kept at PC (PC0101941 and PC0150605) are trail, on bark in a wet forest clearing, 670-695 m, [Link].2016,
Thouvenot NC2477; Païta, Mont Mou, on twigs in cloud forest,
in packets with the handwriting of E. G. Paris without any 1219 m, [Link].2016, Thouvenot NC1971; Humboldt massif, along
annotation by Stephani, so that they cannot be considered the trail to the hut, on bark in cloud forest, 1205 m, [Link].2008,
with certainty as parts of the original material. Thouvenot NC1832; NC1851.

18 CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1)

 Lophocoleaceae of New Caledonia

A B C

F H
E G

J M
K

Fig. 8. — Heteroscyphus coalitus (Hook.) Schiffn.: A, leaves and underleaf, dorsal view; B, C, G, J, M, leaves and underleaves, ventral view; D, H, I, underleaves;
E, K, L, leaf cells; F, tooth at leaf apex. A, B, D, H, E, drawn from the isotype of Jungermannia coalita Hook. (G00283086); C, G, I, from the syntype of Chiloscy-
phus francanus Steph. (G00283067); J, K, from a specimen Franc s.n. of Chiloscyphus similis Steph. (PC0167662); F, L, M, from the lectotype of Chiloscyphus
subsimilis Steph. (G00069431). Scale bars: A-D, G-J, M, 1 mm; E, F, K, L, 50 µm.

Distribution in New Caledonia. — Scattered in both provinces Habit

of the main island, Grande Terre, collected from 300 to 1200 m, Plants large, shoots 3.50-4.00 mm wide when flattened, soft
where it grows usually on barks in wet scrublands and open forests, and fragile; shoots as a whole convex in ventral view and
more rarely on rocks in creek banks.
dorsally canaliculate, at most tube like and the ventral leaf
Total range. — Endemic faces exposed with conspicuous underleaves; leaves dorsally
assurgent, undulate from back to front with the antical half
Description convex, facing the corresponding antical half of the opposite
Dioecious. leaf and the postical half concave, leaves densely imbricate,

CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1) 19

 Thouvenot L.

subopposite, the dorsal margins of opposite leaves confluent instead of clearly bifid, and the underleaf lobes are thinner
on stem back. and forward directed. A specimen of H. confertus collected
in New Caledonia by Le Rat and kept in PC (PC0167665)
Leaves is a doubtful type, contrary to the assumption of Thouvenot
Orbicular, symmetrical, up to 2.50 mm as wide as long, api- et al. (2018), since the locality differs from that of the type
ces shortly bifid, at times entire, sinus lunate, lobes erect to in G. Moreover, the date of collecting is lacking.
connivent, triangular acuminate, margins entire, crenulate
or minutely toothed.
Heteroscyphus cornutistipulus
Cells (Steph.) Thouvenot, comb. nov.
Leaf cells rounded hexagonal, 25-50 µm wide, with medium (Fig. 10)
to large bulging trigones, becoming longer towards bases.
Chiloscyphus cornutistipulus Steph., Species Hepaticarum 6: 303
(Stephani 1922). — Type: New Caledonia. Franc s.n. (lecto-, here
Underleaves designated, G[“Nlle Calédonie, Mts. Koghis”, [Link].1909, Franc 166,
Large, up to 5 times the stem width, widely connate to the G00069490]!).
nearest leaves on both sides, rounded to slightly reniform,
0.70-1.40 mm as wide as long, 1/10-1/3 bifid, sinus obtuse to Further specimen examined. — New Caledonia. South Province,
Sarraméa, Dogny, 855 m, on dead wood in wet forest, [Link].2016,
right angled, lobes triangular acuminate to lanceolate-linear, Thouvenot NC3283 (PC[PC0779841]).
both margins with few to numerous small teeth.
Distribution in New Caledonia. — Only known from scat-
Gametangia tered localities in South Province, on dead wood in wet forest at
medium elevation.
At the end of short leafless lateral-intercalary branches; gynoecia
with bracts lobate, toothed, half long as the perianth, brac- Total range. — Endemic.
teoles small, toothed, perianths cyathiform, 3.00 mm long,
1.70-2.50 mm wide, with 5-6 rounded plicae, upper parts Description
lobate, margins toothed-ciliate, calyptrae small, 0.6-0.7 times Dioecious.
the perianth length; androecia in curved spikes, small, 2-4
pairs of bracts, bracts oval with retuse apices and 1-2 unicel- Habit
lular teeth, margins with a few slime papillae, antheridium Plants medium sized, branching lateral-intercalary; wet shoots
stalk cells two ranked. 3.50-4.00 mm wide, somewhat complanate, with leaves alternate,
horizontally spreading, slightly convex, antical margins free,
Comments separated by 2 cortical cell files, shortly decurrent on the stem.
Stephani (1922) published Chiloscyphus confertus and C. rotun-
difolius with nearly the same description. Checking the type Leaves
specimens at G, it was not possible to find clear distinctive 1.60-2.20 mm long, 1.20-2.00 mm wide, oval to oblong,
features and the differences between the corresponding draw- slightly longer than wide, apices entire to slightly retuse,
ings by Stephani (icones nr 001925 and 001978) are parts of widely rounded, margins entire, hardly crenulate.
the variability in leaf and underleaf marginal ornamentations.
Fresh specimens also show variations in leaf apices, leaf and Cells
underleaf ornamentations in the same plants so that they Median leaf cells rounded-hexagonal 40-80 µm wide, with thin
could be equally assigned to both species. As a result, they walls and small to medium acute trigones up to 9 µm wide.
are considered as synonyms. As both names were published
at the same date and as the name Chiloscyphus rotundifolius Underleaves
was illegitimate (Turland et al. 2018: art 11.4), H. confertus Narrowly to conspicuously connate on single or both sides to
is the correct name. the closest leaves, connation 1-6 cells wide, underleaves 3-5
This species can be separated from other New Caledonian times as wide as the stem, bifid, lobes narrowly triangular, set
species with orbicular leaves, such as Heteroscyphus aselliformis wide apart, sinus lunate, disc transversely elongate, 1-2 short
and H. kanakensis, by the homomallous shoots. In addition, teeth on both lateral margins.
H. aselliformis is readily separated from H. confertus by asym-
metrical leaves and leaf apices with two long filiform lobes, Gametangia
and H. kanakensis by stronger texture (not fragile), smaller Androecia and gynoecia on short leafless lateral branches;
plant size with shoots up to 3 mm wide and leaves to 1.6 mm androecia in spikes made of 6 pairs of bracts very smaller than
long, and by the entire or crenulate leaf margins and rounded the normal leaves and inflated; female bracts not seen, female
apices. It also shares many features with Heteroscyphus men- bracteoles 1/3-1/2 bifid, with 1 tooth on both sides; perianths
ziesii (Mitt.) [Link], from New Zealand, but the latter is a cyathiform, 2.20 mm long, 1.20 mm wide, 3-plicate, plicae
smaller species, the leaves have a conspicuous lip like connation rounded, mouth wide, lobate-laciniate, margins roughly
at back, leaf apices are 2-toothed with sublinear short teeth toothed; calyptrae 2/3 shorter than the perianths.

20 CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1)

 Lophocoleaceae of New Caledonia

A B

E
G

D
H

Fig. 9. — Heteroscyphus confertus (Steph.) Thouvenot: A, shoot portion, ventral view; B, E, G, underleaves; C, D, F, leaves from different shoot segments.
A, H, I, drawn from a syntype of Chiloscyphus confertus (PC0101951); B, C, from Thouvenot NC2477; D-G, from the lectotype of Chiloscyphus rotundiphyllus
Steph. (G00069432). Scale bars: A-G, 1 mm; H, I, 50 µm.

Comments 1.5-2 mm, strongly involute leaves when dry and the ovate-
The type specimen has no gametangium, although the packet is triangular leaf shape. Furthermore, the androecia are in spikes
annotated “c. per.”. A recently collected specimen has androe- on normal-leaved shoots, male bracts being quite similar to
cia and gynoecia (Thouvenot NC3283) and indicates that the the normal leaves. Heteroscyphus succulentus differs by its larger
species belongs in Heteroscyphus. The fertile plant has smaller size, subopposite leaves, underleaves margins long laciniate
trigones but is otherwise vegetatively identical to the type of instead of short toothed, bulging leaf cells without trigones,
Chiloscyphus cornutistipulus. Like in H. caledonicus (see under and crenulate leaf margins.
that species), some vegetative characters of H. cornutistipulus
(Steph.) Thouvenot, comb. nov. are chiloscyphoid, such as the
alternate leaves and the narrow connation of the underleaves. Heteroscyphus deplanchei (Steph.) Schiffn.
Heteroscyphus cornutistipulus (Steph.) Thouvenot, comb. nov. (Fig. 11)
is close to Chiloscyphus trigonifolius and H. succulentus, but
C. trigonifolius is easily separated from H. cornutistipulus Oesterreichische Botanische Zeitschrift 60: 172 (Schiffner 1910). —
(Steph.) Thouvenot, comb. nov. by its smaller size with shoots Chiloscyphus deplanchei Steph., Species Hepaticarum 3: 203

CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1) 21

 Thouvenot L.

A B

H
G

Fig. 10. — Heteroscyphus cornutistipulus (Steph.) Thouvenot, comb. nov.: A, C, shoot portions, ventral view; B, leaf; D-G, underleaves; H, mid-leaf cells. All drawn
from the lectotype of Chiloscyphus cornutistipulus Steph. (G00069490). Scale bars: A-E, G, 500 µm; F, 500 µm; H, 50 µm.

(Stephani 1907). — Type: New Caledonia. Deplanche s.n. (holo-, Province, Dumbéa, Pic des Sources, s.d., Le Rat 265a (G); Mt. Mou,
G[G00069487]!). VII.1904, Le Rat s.n. (G); Pourina valley affluent, 150 m, on dead branch
in mesophilous forest, [Link].1951, Hürlimann 2655 (GOET); Yaté, Wé
Lophocolea heteromorphis Steph., Species Hepaticarum 6: 275 (Stephani Toa, on stump bark in rain forest, 500 m, [Link].2019, Thouvenot NC2852
1922). — Chiloscyphus heteromorphus (Steph.) [Link] & (PC[PC0779849]); Plaine des Lacs, on rocks in lowland wet forest,
[Link]., Nova Hedwigia 39: 416 (Engel & Schuster 1984 260 m, 6.X.2016, Thouvenot NC2441; Pic du Grand Kaori, on rocks
[1985]). — Type: New Caledonia. “Lerat” s.n. (holo-, G[“Lophocolea covered with humus in lowland wet forest, 480 m, 4.X.2016, Thouvenot
heteroformis (sic) [Link]., Dent de St Vincent, juillet 1909”, Le Rat s.n., NC2433; Lac Chakéké, on damp soil in creek bank, 425 m, 4.X.2019,
G00112505]!) syn. nov. Thouvenot NC2729; “Rivière Bleue, forêt vallicole, épixyle”, 160 m,
15.V.1970, Schmid 148 as “Chiloscyphus physanthus” (PC[PC0146342]);
Further specimens examined. — New [Link] Province, Mont Dore, Mouirange, on dead wood in rain forest, 550 m, [Link].2019,
bank of the upper Diahot, on siliceous sand, 430 m, [Link].1951, Thouvenot NC2818; Rivière Blanche, Rivière Bleue Natural Park, sentier
Hürlimann 2901 (G); Hienghène, Pwé Hwa Wéc river, on rocks in creek des Cochons, 556 m, [Link].2019, Thouvenot NC2859; MacKee 30813
bank, 630 m, [Link].2019, Thouvenot NC2773 (PC[PC0779850]); South as “Chiloscyphus caledonicus” (PC[PC0737581]).

22 CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1)

 Lophocoleaceae of New Caledonia

C
A
B

G
I
J

ca

Fig. 11. — Heteroscyphus deplanchei (Steph.) Schiffn.: A, shoot portion, ventral view; B, leaf cells; C, K, leaf apices diversity; D, leaf; E, shoot portion, dorsal
view; F, female bract; G, androecium, dorsal view; H, pair of male bracts and bracteole, dorsal view; I, leaf and underleaf; J, perianth open with calyptra (ca).
A, B, drawn from the holotype of Chiloscyphus deplanchei Steph. (G00069487); C-F, J, from Thouvenot NC2852; G, H, from Thouvenot NC2773; I, K, from the
type of Lophocolea heteromorphis Steph. (G00112505). Scale bars: A, C-F, I-K, 1 mm; B, 50 µm; G, 400 µm; H, 200 µm.

Distribution in New Caledonia. — Fairly frequent in the ultra- Total range. — Endemic.
mafic massif of South Province, rare in siliceous massifs of North
Province, at low to medium elevation (collected from 150 to 1000 m) Description
on dead wood or tree stumps in rain forest. Dioecious.

CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1) 23

 Thouvenot L.

Habit describes Lophocolea heteromorphis with leaves “inaequaliter

Large plants, shoots 3.00-5.00 mm wide, usually single, lateral- bifida vel solum apiculata”. The type specimen seen at G
intercalary branching rare; leaves firms, obliquely assurgent shows mostly such leaves but leaves with entire rounded
dorsally, canaliculate-convex, spreading at right angle to the apices occur on some shoots. Underleaves may be smaller,
stem, subopposite, reaching the dorsal mid-line of the stem less wide and less divided, but here too there are underleaves
and usually confluent. like in Chiloscyphus deplanchei. In fact, all other features are
very similar: habit, shoot width, stem anatomy, cell size
Stem and shape, gametangia and most of the underleaf charac-
300 µm wide, with 60-100 homogeneous thin-walled cells ters. As the leaf apex shape is the only diverging character,
on transverse section. varying along a continuum between two extreme patterns
represented by Chiloscyphus deplanchei and Lophocolea het-
Leaves eromorphis, it can be asserted that they represent variants
Mostly ovate-lingulate, wider at bases, 1.80-2.70 mm long, of the same species.
1.20-2.00 mm wide at base, margins entire as well as the apices Heteroscyphus deplanchei superficially looks like H. suc-
that are widely rounded to obliquely truncate to retuse with culentus (Gottsche) Schiffn. (see below), which has a similar
rounded angles, in some secondary shoots a few or many leaves leaf shape, but the latter can be distinguished by the fleshy,
may be emarginate to shortly bifid with 1-2 acute short lobes. translucent, dorsally free leaves with a more lax areolation,
and underleaves long toothed rather than laciniate and the
Cells cells without trigones. Furthermore, the male bracts of H. suc-
Leaf cells 40-60 µm wide, thin walled with strong nodulose culentus are rounded truncate and the perianth is 3-plicate
trigones, not confluent. with a toothed mouth (Piippo 1985). Heteroscyphus deplanchei
might also be confused with H. supinopsis. For the distinctive
Underleaves characters, see below the relevant paragraph.
Reniform, 8-10 times wider than the stem, 0.80-1.50 mm Although Stephani described the gynoecia, the type material
long, 1.80-2.50 mm wide, connate on both sides to the in G is sterile. The above account of the gametangia, which
ventral bases of the nearest leaves, apices bifid to quadrifid, contains the first description of the androecia, is based on
lobes narrowly acuminate from triangular bases, sinus widely material collected by the author in 2019.
lunate, margins long toothed-laciniate all around, teeth nar-
rowly acuminate, at times furcate.
Heteroscyphus diestianus (Sande Lac.) Piippo
Gametangia (Fig. 12)
Androecia and gynoecia at the apices of very short lateral-
intercalary leafless branches; androecia small, up to 1 mm long, Acta Botanica Fennica 131: 165 (Piippo 1985). — Chiloscyphus
diestianus Sande Lac. in Miquel, Annales Musei Botanici Lugduno-
in spikes made of four pairs of bracts, bracts inflate concave Batavi 1: 296 (Sande Lacoste 1864). — Type: Indonesia. Banca,
with truncate apices, 3-toothed, 400 µm long, 300 µm wide in in m. Maras, 2000’; van Diest s.n., not seen.
situ, bracteoles shortly bifid, 250 µm as wide as long; gynoecia
with two series of female bracts and bracteoles, more or less Specimen examined. — New Caledonia. South Province, Mont
Humboldt, c. 1600 m, [Link].2003, Müller NC803 (DR and au-
similar, obovate in outline, apices lobate, more or less deeply thor’s private herbarium).
incised, margins toothed to laciniate, bracts 3.00 mm long,
2.50 mm wide, bracteoles smaller, 2.25 mm long, 2.00 mm Distribution in New Caledonia. — Only found in South Prov-
wide, perianths cyathiform, without plicae or shallowly plicate ince at the highest elevation.
in upper part, 3.50 mm long, 2.80 mm wide, mouth deeply Total range. — Also occurring in Borneo, Indonesia, Malaysia
laciniate, calyptrae very shorter than the perianths. and Papua New Guinea (Piippo 1993).

Comments Description
Stephani (1907) described Chiloscyphus deplanchei with leaf Based on the New Caledonian specimen. Further descriptions
apices “truncato-rotundo vel emarginatulo”. The scarce shoots and illustrations in Piippo (1993).
of the holotype at G fit this description, but examination of
additional material shows that this character is rather vari- Habit
able. The largest forms have leaves with essentially entire, Plants fleshy, rather swollen, medium sized, shoots 3.00 mm
rounded apices, and strongly laciniate underleaves with the wide, usually single, lateral-intercalary branching rare; leaves
apical lobes hardly distinguishable from the lateral processes. subopposite, thin, imbricate, dorsally obliquely assurgent
But mostly, the plants have long shoot portions with acute at angle of ± 45°, laterally spreading at right angle to the
or shortly bifid leaf apices and other portions, mainly ter- stem, the dorsal segments convex, the ventral strongly con-
minal, with entire, rounded leaves. In rare cases almost all cave, dorsal margins not reaching the mid-line of the stem,
leaves are acute or bifid, with a few entire, rounded leaves not fused to the opposite leaf margin, the dorsal bases not
being restricted to the shoot apices. Later, Stephani (1922) decurrent.

24 CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1)

 Lophocoleaceae of New Caledonia

B
A

H
G

Fig. 12. — Heteroscyphus diestianus (Sande Lac.) Piippo: A, shoot portion, ventral view; B, dorsal face of leaf, with reflexed ventral margin; C, underleaf, adax-
ial face (dorsal view); D, F, H, leaves, ventral view; E, cells; G, appendices in basal segment of dorsal leaf margin. All drawn from Müller NC803. Scale bars:
A-D, F, H, 500 µm; E, 50 µm; G, 100 µm.

Leaves entire or with a few teeth near base, ventral margins entire,
Ovate, 1.20-1.40 mm long, 0.90-1.20 mm wide at bases, arched and recurved in part making a conspicuous concavity
0.45-0.60 mm wide below apices, dorsal bases expanded, on dorsal surface, apices rounded, often sinuous to angulate,
usually toothed, dorsal margins nearly right above the bases, apical margin entire; teeth small to conspicuous, from linear,

CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1) 25

 Thouvenot L.

1-3 cells long, to triangular, 2-3 cells wide at base, 4-5 cells Leaves
long, the teeth cells rounded. Rectangular, 0.60 mm long, 0.50 mm wide, apices truncate,
concave, both distal angles triangular acute, usually with a
Cells short linear tooth three cells long, some leaves pseudo-trilobate,
Leaf cells rounded to oval, 33-60 µm long, 25-37 µm wide, sometimes with a further tooth, apical or subapical on the
very thick-walled with one pit on each side, trigones fused to ventral margin, margins otherwise entire.
the walls, marginal cells smaller, cells often bulging.
Cells
Underleaves Leaf cells hexagonal, 24-40 µm wide, firm walled without
Inserted in a deep sinus, 5-9 times wider than the stems, trigone.
widely reniform, 0.55-0.80 mm long, 1.10-1.80 mm wide,
very narrowly connate on both sides to the nearest leaves, Underleaves
free margins abaxially recurved giving a swollen appearance Small, hardly wider than the stem, obconical, free on both
to the underleaves, apices essentially entire but somewhat sides, deeply bifid, the discs sometimes strongly reduced,
undulate, with a few remote teeth or none, teeth like on the lobes divergent, setaceous, sinus variable, V-like or lunate or
leaves but smaller. nearly flat, both lateral margins with a single tooth at most,
small to linear.
Gametangia
Not seen. Gametangia
On short leafless, lateral-intercalary branches; gynoecia with
Comments laciniate bracts, bifid bracteoles with a single small tooth on
Heteroscyphus diestianus is easily recognized by the large swol- both lateral margins; perianths about 1.5 mm hight, cam-
len underleaves, widely reniform, as wide as the leaves, the panulate, deeply trilobate, lobes laciniate-toothed, calyptrae
teeth restricted to the base of leaf dorsal margins and the very shorter than the perianths; androecia in spikes of up to
strongly thickened cell walls. Its presence in New Caledonia six pairs of bracts.
is documented by a single specimen collected in 2003, in the
vicinity of one of the highest summits of the country. Comments
Some vegetative features of the species such as the alternate leaves,
the leaf cells without trigones and the small, free underleaves
Heteroscyphus etesseanus are suggestive of Chiloscyphus. But the spicate androecia on
(Steph.) Thouvenot, comb. nov. abbreviated ventral branches are typical of the genus Heteroscy-
(Fig. 13) phus. Recent studies on Heteroscyphus in Australasia treat species
Chiloscyphus etesseanus Steph., Species Hepaticarum 3: 217 (Stephani with similar vegetative features and highlight the importance of
1906). — Type: New Caledonia. Etesse s.n. (lecto-, here designated, androecia and perianth types to discriminating these two gen-
G[“ex Herb. E. G. Paris, Nova Caledonia, in ditione Noumeana, era (e.g. Engel & He 2010; Engel 2013, 2015). These studies
1904”, Etesse 12, G00061024]!). warrant the transfer of Chiloscyphus etesseanus to Heteroscyphus.
Further specimen examined. — New Caledonia. North Province, Heteroscyphus etesseanus (Steph.) Thouvenot, comb. nov. is
Poindimié, Amoa valley, Tipwadabwé, on rocks in wet mesophilous superficially similar to H. argutus, but the latter species can
forest, 163 m, 13.X.2019, Thouvenot NC2752 (PC[PC0779851]). be easily distinguished from H. etesseanus (Steph.) Thouvenot,
Distribution in New Caledonia. — Hitherto only known
comb. nov. by its larger size and the rounded instead of retuse
from North Province, rarely collected on rocks at low altitude leaf apices with at least four teeth.
(160-250 m) in wet forest with tall trees. The mention “Nouméa”
on the label of the type specimen is not reliable since Etesse’s
specimens were not precisely located and were commonly labelled Heteroscyphus giganteus (Steph.) Hürl.
as having originated from the capital of New Caledonia. In fact,
his bryophyte gatherings were made along the Tipindjé river in (Fig. 14)
North Province (Morat 2010), about 30 km from the locality of
the recent collection. Bauhinia 12: 114 (Hürlimann 1998). — Chiloscyphus giganteus
Steph., Species Hepaticarum 6: 307 (Stephani 1922). — Type: New
Total range. — Endemic to North Province, on the north-eastern Caledonia. Franc s.n. (lecto-, here designated, G[“Mt. Mou, 1200 m”,
versant of the central range. [Link].1908, Franc s.n., G00069481]!; isolecto-, G[G00283069]!).
Chiloscyphus giganteus fo. minor Herzog, Arkiv för Botanik, n.s. 3 (3):
Description 46 (Herzog 1953). — Type: New Caledonia. [Link].1949, Selling
Dioecious. B126 p.p. (holo-, S).
Further specimens examined. — New Caledonia. North Prov-
Habit ince, Hienghène, Ouaïème rocks, on trunks in mountain shrubland,
Plants small, wet shoots 1.60 mm wide, stems rigid, leaves 854 m, [Link].2019, Thouvenot NC2798; South Province, Mont
horizontally spreading, slightly convex, contiguous, alternate. Dore, Prony, Baie du Carénage, [Link].1950, Guillaumin 8579a

26 CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1)

 Lophocoleaceae of New Caledonia

A B C D

E G
F

H
J
L

K
I

P
N

M
O

Fig. 13. — Heteroscyphus etesseanus (Steph.) Thouvenot, comb. nov.: A, shoot portion, ventral view; B-D, leaves; E-I, underleaves; J, median leaf cells; K, tooth
at leaf apex; L, androecium; M, female bracteole; N, perianth tearing showing the inserted calyptra; O, female bract; P, perianth. A, L, M-P, drawn from the
lectotype of Chiloscyphus etesseanus Steph. (G00061024); B-D, K, from Thouvenot NC2752; E-J, from Thouvenot NC2753. Scale bars: A, L, 1 mm; B-D, M-P,
500 µm; E-I, 200 µm; J, K, 50 µm.

(GOET); Yaté, Forêt du Mois de Mai, 235 m, 23.X.2012, Thou- Total range. — Endemic.
venot NC1008; Yaté, Goro, 200-310 m, [Link].2016, Metoyer 122;
Yaté, Pic du Grand Kaori, on bark, 472 m, 4.X.2016, Thouvenot
2435 (PC[PC0779852]); Bouloupari, Mt. Do, on bark, 960 m, Description
[Link].2016, Thouvenot NC2470. Dioecious.
Distribution in New Caledonia. — Frequently found in South Habit
Province, rarer in North Province, at low to medium altitudes (col-
lected from 0 to 1250 m), Heteroscyphus giganteus grows on barks in Plants light green, large, fragile, 4.00 mm wide in fresh con-
various vegetation types, from photo-xerophilous to cloud forests, dition, 6.00-7.00 mm when flattened, simple or sparsely
primary or secondary mountain scrublands, gallery forests. furcate, branching lateral-intercalary, leaves subopposite, dor-

CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1) 27

 Thouvenot L.

sally assurgent, longitudinally undulate with the ventral half Chiloscyphus quadricilius Steph., Species Hepaticarum 6: 312 (Stephani
concave and the dorsal convex, thus the shoots are swollen. 1922). — Type: New Caledonia. “Lerat” s.n. (lecto-, here designated,
G[“Nova Caledonia”, s.l., s.d., Lerat 21, G00283105]!) syn. nov.
Leaves Further specimens examined. — New Caledonia. North Prov-
Large, rounded, more than 3.00 mm wide, margins entire, ince, Poindimié, Tango plateau, Napapwa, on the ground in rain
forest, 467 m, [Link].2019, Thouvenot NC2895; South Province,
apices emarginate with short lunate sinus and two short lobes Bouloupari, Mt. Do, on peridotite rocks in cloud forest, 995 m,
triangular to rounded. 25.X.2012, Thouvenot NC752; Paita, Dzumac massif, on rocks in
mesophilous forest, 915 m, [Link].2008, Thouvenot NC331; Yaté,
Cells Nooti, on laterite rock, in photo-xerophyte forest, 443 m, 5.X.2019,
Thouvenot NC2750; Yaté road, col des deux Tétons, [Link]. 2014,
Leaf cells 30-56 µm wide, thin walled, hexagonal with rounded Métoyer NC1686 (NOU); MET027 (NOU); Yaté road, col des deux
lumina and large truncate trigones. Tétons, [Link].2014, Métoyer NC1687 (NOU); NC1688 (NOU);
MET055 (NOU); MET056 (NOU).
Underleaves
Distribution in New Caledonia. — One of the most frequent
Reniform, 1.00-1.60 mm long, 1.00-2.00 mm wide, bilaterally species of Lophocoleaceae in both provinces of Grande Terre, Het-
connate to the nearest leaves, deeply bifid, the lobes lanceolate, eroscyphus grandiflorus grows in usually wet conditions on the ground
simple or furcate, widely divergent, lateral margins usually where it occupies a range of substrates, including bare soil, humus,
with at last two long narrow teeth or laciniae. plant debris, dead wood, rock, etc. at low to medium elevations
(collected from sea level to 1000 m).
Gametangia Total range. — Endemic.
At the end of very short leafless lateral shoots; gynoecia
2.75 mm long, involucres strongly inflate, bracts 2.00 mm Description
long, 1.75 mm wide, ovate in outline, deeply and asymmetri- Dioecious.
cally bifid, margins coarsely toothed, teeth large and triangular,
bracteoles 2.00 mm long, 1.50 mm wide symmetrically bifid, Habit
sinus half-length deep, lateral margins coarsely toothed, young Variable (see comments); plants medium to large, light green
perianths cyathiform, deeply multifid, lobes ovate-triangular to red-brown or olive-brown, wet shoots 2.00-4.00(-5.00) mm
sparsely and coarsely toothed. Androecia in curved spikes, wide, leaves subopposite, densely imbricate, dorsally free, when
made of about 5 pairs of bracts. dry dorsally concave to canaliculate, patent at 60-90° angle.
Comments Leaves
This species is easily separated from all other New Caledonian The most typical leaves are 1.00-2.00 mm long, 0.80-2.00 mm
Lophocoleaceae by the combination of the following characters: wide at base, widely oval-ovate, apices rounded to widely
1) large shoots, 6-7 mm wide; 2) large leaves rounded concave triangular, usually with 3 long cilia made of 3-6 uniseriate
and dorsally assurgent so that the shoots seem inflate; 3) mar- elongate cells narrower from base to tip, ventral margins with
gins entire; 4) apices emarginate with small lunate sinus; 5) leaf 0-3(-4) cilia identical to the apical ones.
cells with large truncate trigones; and 6) underleaves bifid with
lobes widely divergent, and 2-3 laciniae on each lateral margin. Cells
Fresh fertile specimen shows the dioecious condition and allows Leaf cells 37-65 µm long, 32-42 µm wide, trigones medium
to describe the gynoecium. Heteroscyphus giganteus superficially to large, rounded to truncate.
looks like H. aselliformis and H. confertus; to distinguish charac-
ters see under these species. The size of C. giganteus fo. minor is Underleaves
like in H. confertus, but underleaves and trigones match those of Connate with the ventral bases of both nearest leaves, the most
C. giganteus. Consequently, it is not necessary to separate this form. typical underleaves transversely elongate, 2-4 times wider than
the stem, discs 0.35-0.50 mm high, 0.60-1.00 mm wide, apices
bifid-quadrifid, sinus lunate, lobes narrowly linear lanceolate,
Heteroscyphus grandiflorus (Steph.) Hürl. single or furcate, lateral margins with one to several long narrow
(Fig. 15) teeth sometimes furcate, at most the entire margins appearing
laciniate all around, minor forms with only four apical teeth
Bauhinia 12: 114 (Hürlimann 1998). — Chiloscyphus grandiflorus or two apical and one small marginal on both sides.
Steph., Species Hepaticarum 6: 307 (Stephani 1922). — Type: New
Caledonia. Lerat s.n. (lecto-, here designated, G[“Original”, Lerat
265-03, G00069506]!; syn-, G[Pic des Sources, “moitié”, Le Rat 259, Gametangia
G00283070!, Mt. Mou, VII.1909, Le Rat 133, G00283071]!; iso- On short lateral-intercalary branches lacking normal vegeta-
syn-, REN[Mt. Mou, VII.1909, Le Rat s.n., herb. E. G. Paris ]!). tive leaves, rarely with one pair of small vegetative leaves at
Chiloscyphus grandiflorus var. latifolius Herzog, Arkiv för Botanik, base; gynoecia with bracts large, lobate-laciniate, bracteoles
n.s. 3 (3): 49 (Herzog 1953). — Type: New Caledonia. [Link] smaller, deeply bifid and laciniate; perianths cyathiform,
of Yaté River, 400 m., [Link].1949, Selling B12 (holo-, S) syn. nov. 3.00 mm long, 2.00 mm wide, deeply lobate-laciniate at

28 CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1)

 Lophocoleaceae of New Caledonia

A B

F G
E

H I

J
K

Fig. 14. — Heteroscyphus giganteus (Steph.) Hürl.: A, shoot portion, lateral view; B, D, F, G, leaves; C, leaf cells; E, I-K, underleaves; H, androecium, dorsal
view; I, habit, dorsal view. A, B, H-J, drawn from Coulerie 196; C-E, from the lectotype of Chiloscyphus giganteus Steph. (G00069481); F, G, K, from Thouvenot
NC1008; L, photo from Thouvenot NC2823. Scale bars: A, B, D-K, 1 mm; C, 50 µm; L, 5 mm.

mouth; androecia in short spikes, 0.40 mm long, with 2-5 mentations, even in the same plant, so that some specimens
pairs of bracts cochleariform-canaliculate, toothed-ciliate, might be thought to belong to different species or distinct
bracteoles shortly bifid with a few unicellular teeth, anther- varieties. Usually, main shoots have the largest leaves, with
idium stalk three ranked. many cilia around the margins and underleaves with two apical
lobes often furcate and lateral margins lobate ciliate, whereas,
Comments on the secondary branches, leaf margins may be naked except
As underlined for many species of the genus, this frequent the apical three cilia and underleaf cilia may be simple; mini-
species has variable looks, mainly in leaf and underleaf orna- mal forms of underleaves have only two apical cilia and one

CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1) 29

 Thouvenot L.

short cilia on both margins. But, in all observed specimens, For the distinction from Heteroscyphus caledonicus, see
the overall leaf shape and areolation are constant despite the under this species.
size and ornamentation diversity, except the single sample The antheridium characters have not been published by
from semi-aquatic habitat with oblong-oval leaves, usually Stephani in his diagnosis of Chiloscyphus grandiflorus. It can
biciliate leaf apices, olive brown colour (Thouvenot NC2894). newly be described from vouchers of recently collected male
Other characters varying case to case are colour, from light shoots (e.g. Metoyer NC1688, Thouvenot NC2750).
green to red brown or olive green, leaf cell trigones truncate
to rounded, underleaf cell trigones which may be like the
leaf ones to nearly absent. The well-developed populations Heteroscyphus kanakensis Thouvenot & Engel
include usually a large morphological variability. But we are (Fig. 16)
often faced with quite homogeneous populations with a pecu-
liar set of characteristics raising the question of the variety Nova Hedwigia 112: 166, 1-2 (Thouvenot & Engel 2021). —
Type: New Caledonia. South Province, Yaté, Rivière Bleue Natural
status. Notably, the variety latifolius defined by Herzog from Park, La Tranchée, on tree bark in river side, ultramafic massif,
differences in leaf size and number of leaf segments falls in 200 m, 19.X.2016, Thouvenot NC2487 (holo-, PC[PC0779856];
synonymy to the typical forms since such characters can be iso-, F, hb. Thouvenot).
observed in some shoots of the types kept at G: the observed
Further specimens examined. — New Caledonia. North Prov-
variability in the leaf base width ranges from 0.70 to 1.00 mm ince, Pouébo, Diahoué, on wet rocks in the creek Wé Wayat, 400 m,
and ornamentation from 3 to 8 fringe segments. This is a fre- [Link].2019, Thouvenot NC2771; NC2829; Touho, Pombei, Tiwaka
quent character in Lophocoleaceae where secondary shoots and valley, on rocks in high bush of Melaleuca quinquenervia (Cav.)
ultimate branches exhibit less process numbers and smaller [Link] and Gymnostomum sp., volcano-sedimentary massif, 415 m,
sizes than primary stems. Old primary parts may be destroyed 12.X.2016, Thouvenot NC2488 (para-, NOU, F); South Province,
Lac Chakéké, on silt-laden deposits on barks along a temporary
or broken and inconspicuous in some type specimen so that creek, in gallery forest, 411 m, 4.X.2019, Thouvenot NC2727 (para-,
it could give rise to different diagnoses. At the other end, PC[PC0779848], F, hb. Thouvenot).
populations might be gathered into a variety “minor” when
the following forms of morphological characters are evenly Distribution in New Caledonia. — In a few localities at low
altitudes in North and South Province, from 200 to 400 m. It grows
expressed: leaves without marginal processes except the three, on various substrates, barks, rocks, or silt laden deposits on trunks
rarely two, apical ones, underleaves of the less developed type, along creeks and was found in wet forest in sedimentary massif as
hardly twice as wide as the stem, 2-4-fid, lobes short linear, well as gallery forest inside ultramafic scrubland.
one short tooth on both margins (e.g. Metoyer NC1687,
Total range. — Endemic.
Thouvenot NC2895). Some were collected in peculiar habitats,
namely very shaded, semi-aquatics or disturbed habitats. In
the latter cases, the author hypothesise they could be mats Description
of secondary shoots regenerating from broken stems, since Further description and illustrations in Thouvenot & Engel
it was sometimes possible to find a few old shoot segments (2021).
with typical wide laciniate underleaves. Dioecious.
Among the two type specimens of Chiloscyphus quadri-
cilius kept at G, only number G00283105 matches the Habit
Stephani’s protologue and, therefore, is selected as the Plants rather rigid, fragile, pale green, pale brown in older
lectotype. The selection of the specimen G00069435 sectors, somewhat water repellent, the shoots to 3 mm wide
“sur les arbres, 1905, Le Rat 302” as a type specimen of when flattened; stems narrow for plant size; leaves suboppo-
C. quadricilius fide Bonner (1963: 767) is not supported by site, typically obliquely dorsally assurgent (the basal sector of
the observation of its morphological characters since leaf dorsal half angling upward at c. 45° from stem and never facing
apices are widely rounded with four small apiculate teeth dorsal base of opposing leaf ), the leaves densely imbricate, free
at leaf apices and small bifid underleaves with only one dorsally; leaves somewhat longitudinally undulate in dorsal
marginal tooth on both margins, so that it differs highly aspect, moderately convex in dorsal sector, moderately abaxially
from the diagnose and drawing of Stephani and turns to concave in ventral sector, the concave ventral sector overlayed
be Heteroscyphus argutus. The type specimen of Chiloscy- by the convex dorsal half of the leaf immediately above.
phus quadricilius here selected is sterile, but the vegetative
and male characters are convenient for a transfer into the Leaves
genus Heteroscyphus, underleaves widely connate on both Subsymmetrical, up to 1.60 mm long, ovate and up to c. 1.3
side with the neighbouring leaves, leaves subopposite, leaf times longer than wide; apex broadly rounded, entire; margins
cells with large bulging trigones, androecia at the end of broadly and ± evenly arched, entire to faintly crenulate, the
short leafless lateral branches. Furthermore, the features dorsal margin not decurrent.
described by Stephani together with our observation on
the type perfectly match Heteroscyphus grandiflorus, even Cells
if Stephani states the leaf cells are lacking trigones. In the Leaf cells with nodulose trigones separated by narrow, thin-
type specimen, the cells show large nodulous trigones. walled places, the trigones sometimes confluent, the median

30 CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1)

 Lophocoleaceae of New Caledonia

A B C

M
H
I
L

J K

Fig. 15. — Heteroscyphus grandiflorus (Steph.) Hürl.: A, E, H, shoot portions from different parts of a single specimen; B, D, G, underleaves; C, I, K, leaves;
F, ventral margins of two adjacent leaves showing their imbrication; J, leaf and underleaf; L, cilia from leaf apex; M, cells. A, E, H, drawn from the specimen
Thouvenot NC2745; B, from Thouvenot NC1486; C, D, from Thouvenot NC0752; F, from the syntype of Chiloscyphus grandiflorus Steph. (G00283070); G, from
Thouvenot NC2895; I, J, from the lectotype of Chiloscyphus grandiflorus (G00069506); K, from the syntype of Chiloscyphus grandiflorus (G00283071); L, M, from
Thouvenot NC2750. Scale bars: A-K, 1 mm; L, M, 20 µm.

leaf cells uneven in size, 8-12 μm wide, 10-20 μm long, sinus narrowly to widely rounded; lamina margins on both
somewhat larger near the base; surface with a granular, scurffy sides with a large tooth, the lamina margins otherwise entire
appearance throughout. or with 1-2(3), smaller, accessory teeth below.

Underleaves Gametangia
4-5 times the stem width when flattened, symmetrically Gynoecia on short lateral-intercalary, leafless branches; bracts
connate with the leaves on both sides, the connate portion deeply laciniate-lobed, the margins toothed toward the base;
1-3 cells wide; underleaves imbricate, rotund to oblate, 0.50- bracteole deeply bifid, the margins sparsely laciniate or toothed;
0.80 mm long, 0.50-0.90 mm wide, usually 1.15 times wider perianths seen only in juvenile state, cyathiform, eplicate,
than long; apex bifid to c. 0.35, the lobes ± parallel to weakly the mouth lobulate, the lobules ending in a uniseriate row
divergent, widely triangular, acute to stoutly acuminate, the and with small teeth below; androecia small for plant size,

CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1) 31

 Thouvenot L.

nearly hidden by leaves in dorsal aspect, fully exposed in Distribution in New Caledonia. — Scattered in North and
ventral aspect, on abbreviated, determinate, lateral-intercalary South Provinces, rarely collected.
branches, the androecia usually strongly arched, with 2-8 pairs Total range. — Endemic.
of bracts, the bracts dorsally convex, strongly ventricose, half-
spherical-helmet shaped, the dorsal pocket scarcely defined, Description
the inflated upper faces of the bracts elevated above the level Further description and illustrations in Thouvenot & Price
of the axis so that the spike is furrowed, the apices rounded (2020).
to retuse, entire, the margins sparingly crenulate by projec- Dioecious.
tions of a single cell, with slime papillae few to numerous;
bracteoles oblong, short bifid, the margins entire, rarely with Habit
a small tooth, with slime papillae few to numerous; anther- Plants olive green, medium to large, shoots up to 4.5 mm
idial stalk biseriate. wide; shoots often attenuated with leaves becoming progres-
sively smaller toward shoot tops; branching latero-ventral;
Comments leaves spreading at a wide angle from the stem, slightly convex,
Heteroscyphus kanakensis superficially resembles H. menziesii alternate, not dorsally decurrent.
(Mitt.) [Link] described from New Zealand (Engel &
Glenny 2019). They share many characters, among them: 1) Leaves
a habit with light colour, medium size 2-3 mm wide when Normal leaves 1.40-2.00 mm long, 0.90-1.50 mm wide,
flattened, canaliculate shoots with subopposite leaves more oval-oblong, transversally to obliquely truncate; apices usu-
or less assurgent; 2) rounded leaves densely imbricate, lon- ally concave with a single acute tooth at both distal angles,
gitudinally undulate; 3) large trigones, cells bulging so that sometimes shallowly bifid with short lobes widely triangular,
the leaf surfaces are bumpy; 4) bifid rounded underleaves, minutely apiculate, some leaves with rounded apices; lateral
relatively large, connate on both sides with the nearest leaves. margins entire and smooth.
However, Heteroscyphus kanakensis is characterized by: 1)
rather rigid shoots; 2) leaves obliquely assurgent, the dorsal Cells
insertion of both opposite leaves separated by one file of stem Leaf cells hexagonal, isodiametric to slightly elongate, 44-66 µm
cells; 3) leaf margins and apices entire; 4) underleaves 4-5 long, 25-52 µm wide, walls thin without trigone.
times the stem, with two triangular acute teeth at apices and
1-3 smaller teeth on both lateral margins; and 5) androecia Underleaves
longer, in arched spikes of 2-8 pairs of bracts, conspicuous Small, not or hardly wider than the stem, overall oval, deeply
in ventral view. In contrast, typical forms of H. menziesii bifid, disc wider than long, 3-4 cells high, insertion line
has soft and spongy shoots, leaves vertically assurgent, the semi-circular, lateral margins with a small obtuse tooth on
dorsal base facing the dorsal base of the opposite leaf, dor- both sides, sometimes linear, apices with sinus lunate, both
sal leaf margins conspicuously connate by way of a laminar lobes erect or crescent shaped, narrowly lanceolate acumi-
strip several cells wide, leaf apices 2-toothed, underleaves nate to linear.
up to 3 times the stem width, margins with many denti-
form or ciliiform teeth, androecia shorter with 1-2 pairs of Gametangia
bracts, hidden in both ventral and dorsal views (Engel & Gametangia of both sexes terminal on short leafless branches,
Glenny 2019). lateral ventral; only juvenile gynoecia seen, bracts oval, 1.20-
1.80 mm long, bifid, lobes ovate acute, a lanceolate segment
For differences from the similar species Heteroscyphus con- usually developed on the upper part of a single lateral margin
fertus, see the comparison under that species. otherwise entire or with rare small teeth, bracteole deeply
bifid, 1.50 mm long, lobes narrowly lanceolate, more or less
convergent, margins entire, perianth 2.0-2.5 mm long, cyathi-
Heteroscyphus parapilistipulus form, smooth, tri-lobate, lobes deeply laciniate; androecia in
(Thouvenot) Thouvenot, comb. nov. thin spikes of ± 6 pairs of ventricose bracts.
(Fig. 17)
Comments
Chiloscyphus parapilistipulus Thouvenot, Candollea 75: 286, This plant resembles Lophocolea convexula (synonym: L. pil-
figs 1; 2 (Thouvenot & Price 2020). — Type: New Caledonia.
South Province, La Foa, Dogny plateau, on wet rock in the creek istipula, see below) but differs from the latter species by:
Dogny, 918 m, Thouvenot NC2451 (holo-, PC[PC0763751]!; iso-, 1) a larger size of shoots, leaves and cells; 2) the absence of
author’s private herbarium; para-, G[New Caledonia. “In jugo Dogny trigones; 3) outer cell walls flat thus leaf margins smooth,
(1050 m)”, VII.1909, L. Le Rat s.n., G00051491!, G00051492!]; not crenulate; 4) a dioecious condition; 5) gynoecia on very
PC[PC0102406!, PC0150609!]; REN s.n.!). short branches without vegetative leaves; and 6) androecia
Further specimens examined. — New Caledonia. North Prov- in spikes of six pairs of bracts on ventral lateral abbrevi-
ince, St Paul, Pwâ Yogac river, on damp emerged rock in a creek ated branches. Among the three syntypes of L. pilistipula
bed, 438 m, [Link].2019, Thouvenot NC2786. Steph. at G, only the tiniest specimen, from Isle of Pines

32 CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1)

 Lophocoleaceae of New Caledonia

A B

D
E

J
G K

F
H
I

P
N

M
L R
O

T
Q

V U

S
W

Fig. 16. — Heteroscyphus kanakensis Thouvenot & [Link]: A, leaf pair and attached underleaf; B, portion of shoot (habit in dorsal view); C, median leaf cells;
D, H, leaves; E-G, I, L, M, underleaves; K, marginal cells at leaf apex; J, stem transverse section; O, U, male bracteoles; P, male bract; T, androecium habit
(ventral view); N, shoot transverse section showing dorsally assurgent antical part of leaves; V, top part of perianth (expanded); Q, S, female bracteoles; R, gy-
noecium habit (ventral view); W, X, female bracts. A, F, L, drawn from the paratype Thouvenot NC2488; B, J, N-X, from the holotype; C, D, E, G, H, I, K, M, from
the paratype Thouvenot NC2727. Scale bars: A, D-I, Q, S, T, V-X, 500 µm; B, R, 1 mm; C, J, K, 20 µm; L, M, 200 µm; N-P, U, 100 µm.

(G00112487), matches the morphological features of this near base, and cells 44-66 µm long and 33-50 µm wide.
species. The other two, both from Dogny (G00051491 and Furthermore, the leaf margins are smooth instead of crenu-
G00051492) are sterile, have larger dimensions and belong late. Duplicates of these collections are in PC. The initial
to the new species Heteroscyphus parapilistipulus (Thouvenot) placement of the species in Chiloscyphus was warranted by
Thouvenot, comb. nov. with shoots being 3.00-4.50 mm the discovery of a female plant (Thouvenot & Price 2020),
wide, leaves 1.40-2.00 mm long and 0.90-1.50 mm wide but then, another specimen collected in North Province

CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1) 33

 Thouvenot L.

A B

C
F

androecium
K

Fig. 17. — Heteroscyphus parapilistipulus (Thouvenot) Thouvenot, comb. nov.: A, shoot portion (ventral view); B, median leaf cells; C, F, leaves; D, E, I, underleaves;
G, H, female bracts; J, female bracteole; K, top part of perianth (expanded). All drawn from the holotype. Scale bars: A, 400 µm; B, 50 µm; C-K, 200 µm; L, 1 mm.

(Thouvenot NC2786) has proven to be male, with a few old limited growth, despite some vegetative leaves were seen at
antheridia of heteroscyphoid type hidden amongst oldest the end of a single spike. The combination of both sexual
parts of shoots. They consist of elongate spikes of six pairs characters brings evidence for placing this species in the
of bracts at the end of short ventral lateral branches of genus Heteroscyphus.

34 CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1)

 Lophocoleaceae of New Caledonia

A
B

E
D
C

Fig. 18. — Heteroscyphus splendens (Lehm. & Lindenb.) Grolle: A, shoot portion, ventral view; B, underleaf; C-E, leaves. All drawn from the New Caledonian
specimen Hürlimann 2830 (GOET) of Chiloscyphus decurrens Nees. Scale bars: 1 mm.

Heteroscyphus splendens Habit

(Lehm. & Lindenb.) Grolle Plants large, moist shoots c. 3.00 mm wide when flattened;
(Fig. 18) leaves dorsally assurgent when dry, spreading when moist,
subopposite, dorsal margins confluent.
Acta Botanica Fennica 125: 68 (Grolle & Piippo 1984). — Junger-
mannia splendens Lehm. & Lindenb., Novarum et Minus Cognitarum Leaves
Stirpium Pugillus 4: 22 (Lehmann 1832). — Type: India (‘India
orientalis’). Wallich (lecto-, selected by Grolle [1984], S), not seen. Oblong, c. 1.60 mm long, 1.40 mm wide at base, apices
rounded-truncate, entire, ventral margins entire or with 1-2
Chiloscyphus decurrens Nees, Synopsis Hepaticarum: 173 (Gottsche et al. small teeth near base, dorsal margin entire.
1845). — Jungermannia decurrens Reinw., Blume & Nees, Nova Acta
Physico-medica Academiae Caesareae Leopoldino-Carolinae Naturae Cells
Curiosorum Exhibentia Ephemerides sive Observationes Historias
et Experimenta 12: 206 (Reinwardt et al. 1824). — Type: Java. Median leaf cells 20-36 µm wide, lumina rounded to oval,
Blume s.n., not seen. trigones medium to large, acute.
Specimens examined. — New Caledonia. North Province, Poin- Underleaves
dimié, Amoa Valley, Twipadabwé, mixed with other liverworts
on a rock in a river bank, volcano-sedimentary bedrocks, 211 m, Connate to the ventral margins of both nearest leaves, rounded
13.X.2019, Thouvenot NC2730 (PC[PC0779854]); Hienghène, to reniform, 0.80-1.00 mm long, 1.00-1.20 mm wide, 4.5
Pwé Hwa Wéc river, on rocks on creek bank, 630 m, [Link].2019, times the stem width, apices rounded, shortly bifid, lobes
Thouvenot NC2773; Gomen road, east slope of Mt. Ignambi, 600 m, narrowly triangular, sinus narrow, margins spinose toothed
[Link].1951, Hürlimann 2830 as Chiloscyphus decurrens (GOET). to shortly ciliate all around.
Java Occidental. Priangan, Gede-Prangango, c. 2800 m, VIII.1930,
Verdoorn 2375 (PC[PC00035416]).
Gametangia
Distribution in New Caledonia. — In New Caledonia, this very Gynoecia at the end of short lateral-intercalary branches,
distinctive species is only known from two localities in North Province. perianths cyathiform; androecia not seen.
Total range. — Paleotropical: from East Africa to Indonesia,
Malaysia, Melanesia, South Pacific Islands (Hürlimann 1998). Comments
Heteroscyphus deplanchei and H. supinopsis superficially resem-
ble H. splendens but can easily be separated from the latter
Description by the ornamentation of the underleaf margins, which are
Based on the New Caledonian specimens. Further descrip- strongly laciniate in H. deplanchei and more remotely toothed
tions and illustrations in Piippo (1985). in H. supinopsis (see below).

CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1) 35

 Thouvenot L.

A B

D
E
F

Fig. 19. — Heteroscyphus subacuminatus (Herzog) Thouvenot: A, shoot portion, ventral view; B, C, leaf apices; D, F, half underleaves; E, mid-leaf cells; G, male
bract with antheridium; H, leaf. All drawn from Thouvenot NC2740. Scale bars: A, D, F, H, 1 mm; B, C, 500 µm; E, 50 µm; G, 200 µm.

Heteroscyphus subacuminatus Leaves

(Herzog) Thouvenot, comb. nov. Leaf shape asymmetrically ovate acuminate, 2.00-2.50 mm
(Figs 19; 20) long, 1.30-1.70 mm wide, ventral margins rounded in lower
2/3-3/4, dorsal margin nearly right, apices widely acuminate,
Chiloscyphus subacuminatus Herzog, Arkiv för Botanik, n.s. 3 (3): acute to narrowly rounded, rarely bifid, frequently broken
49 (Herzog 1953). — Type: New Caledonia. Slope above N. branch
of the Yaté River, near the Forêt du Mois de Mai, [Link].1949, Sell- (Fig. 20C, D).
ing B144 p.p., not seen.
Cells
Specimen examined. — New Caledonia. North Province, Poin- Leaf cells hexagonal, 30-50 µm wide, walls thin with large
dimié, Amoa valley, Tipwadabwé, on rocks along the trail when
crossing a flooding creek, 211 m, 13.X.2019, Thouvenot NC2740. subacute trigones, cell lumina round to ovate.

Distribution in New Caledonia. — Rarely collected in both Underleaves

provinces of the main island, Grande Terre, at low elevation. Narrowly connate to both nearest leaves, 0.75-1.00 mm long,
Total range. — Endemic. 1.20-1.50 mm wide, the disc rounded with widely wedge-
shaped bases, insertion in a deep sinus, apices 1/3 bifid, sinus
Description narrowly lunate or acute, segments narrowly triangular, end-
Dioecious. ing in piliform acumen, margins unevenly spinose-toothed
all around.
Habit
Plants fragile, light olive green, brown in older parts, large Gametangia
shoots 3.50-4.50 mm wide, leaves subopposite, secund, dor- Androecia at the end of very short lateral-intercalary branches
sal margins conspicuously connate over the stem, dorsally lacking normal vegetative leaves, arranged in small spikes with
assurgent at least in upper parts of the shoots, the lowest up to 4 pairs of bracts, bracts very smaller than the vegetative
horizontally spreading. leaves, imbricated, hemispheric, margins with a few short

36 CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1)

 Lophocoleaceae of New Caledonia

A B

C D

Fig. 20. — Heteroscyphus subacuminatus (Herzog) Thouvenot, comb. nov.: A, shoot in dorso-lateral view; B, antheridium; C, androecia on shoot portion, ventral
view; D, shoot portion, dorsal view. All from Thouvenot NC2740. Scale bars: A, C, D, 1 mm; B, 100 µm.

teeth, bracteoles oblong-rectangular, shortly bifid, margins Heteroscyphus succulentus (Gott.) Schiffn.
entire or with rare teeth, slime papillae rare to numerous on (Fig. 21)
the margins and tooth apices of bracts and bracteoles, anther-
idia stalks biseriate; gynoecia not seen.
Oesterreichische Botanische Zeitschrift 60: 171 (Schiffner 1910). —
Chiloscyphus succulentus Gött., Natuurkundig Tijdschrift voor Neder-
Comments landsch-Indië 4: 574, 576 (Gottsche 1853). — Type: Java. Zollinger
3513 ex parte (iso-, G[G00114825]!).
According to Herzog (1953), the type specimen of Chiloscyphus
subacuminatus is sterile. The author collected male plants of Specimens examined. — New Caledonia. North Province, Ouégoa,
the species in 2019, which showed the presence of androecia Diahot upper valley, 430 m, on siliceous sandy ground on the river-
in short spikes on very short lateral branches and allowed the bank, [Link].1951, Hürlimann 2899a (G); 2901 (G); Poindimié,
transfer of the species to Heteroscyphus. The transfer is also Tipwadabwé, Amoa valley, 244 m, on dead wood in river side wet
forest, in volcano-sedimentary bedrock, 13.X.2019, Thouvenot
supported by the underleaves being connate with the nearest NC3286 (PC[PC0779844]); South Province, Koghis, SE ridge of the
leaves on both sides, the subopposite, dorsally connate leaves, Mt. Bouo, 830 m, on dead wood in mountain forest, [Link].1950,
and the biseriate antheridial stalks. Hürlimann 2088 (PC[PC0763233]).

CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1) 37

 Thouvenot L.

A B

Fig. 21. — Heteroscyphus succulentus (Gottsche) Schiffn.: A, shoot portion, ventral view; B, leaf; C, atypical leaf apex; D, F, underleaves; E, leaf cells. All drawn
from the isotype of Chiloscyphus succulentus Gött. (G00114825). Scale bars: A-D, F, 1 mm; E, 50 µm.

Distribution in New Caledonia. — New to New Caledonia, in

North and South Province. Underleaves
Narrowly connate to both nearest leaves, reniform in outline, 0.75-
Total range. — Reported from Malaysia, Indonesia, Papua New
Guinea and Tahiti (Piippo 1985, GBIF accessed 29 Sept. 2021). 1.25 mm long, 1.25-2.50 mm wide, deeply bifid, lobes widely
divergent, decurved, long lanceolate acuminate with linear apices,
sinus widely lunate to flat, disc transversally rectangular, lateral
Description margins with 2(-3) laciniae similar to the lobes; cells bulging.
Habit
Plant large, light green, glossy, fleshy; shoots creeping, 4.00- Gametangia
4.50 mm wide, leaves subopposite, dorsally free and slightly Not seen.
decurrent, spreading, dorsally assurgent with a convex base,
then somewhat secund. Comments
Heteroscyphus succulentus looks superficially like H. deplanchei
Leaves but can easily be distinguished by: 1) a fleshy appearance; 2)
Thin, translucent, ovate-oblong to tongue-like, 1.80-2.00 mm large, thin-walled, leaf cells without trigones; 3) dorsal margins
long, 1.30-1.50 mm wide at bases, 0.60 wide at apices, typi- of paired leaves not connate; and 4) the leaf surface appearing
cally with apices widely convex and entire, or truncate, with uneven and glossy because of the strongly bulging cells. For dif-
or without 1-celled teeth at angles, atypical leaves obliquely ferences with H. cornutistipulus see above the relevant paragraph.
truncate or shortly bifid with triangular lobes separated by The species was never formally reported from New Cal-
a wide sinus, obtuse or lunate; margin crenulate by the pro- edonia, although Hürlimann collected a specimen in South
truding bulging cells. Province in 1950 labelled Heteroscyphus cf. succulentus (PC,
GOET, G) (unpublished). Parts of this material in PC and
Cells G were checked, together with further specimens, and the
Hexagonal, 40-60 µm in upper half of the leaves, becoming presence of this species in both provinces of the main island,
elongate in basal part, up to 100 µm long, walls thin, without Grande-Terre, can be confirmed. The whole range of the spe-
trigones, bulging so that leaf surface looks rough. cies in New Caledonia remains under-documented.

38 CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1)

 Lophocoleaceae of New Caledonia

A B

D E

Fig. 22. — Heteroscyphus supinopsis [Link], Thouvenot & Frank Müll.: A, G, leaves; B, pair of leaves with associated underleaf, ventral view; C, F, underleaves;
D, cells from leaf median portion; E, shoot portion, ventral view; H, dorsal leaf connation, dorsal view. All drawn from the holotype. Scale bars: A-C, F, G, 500 µm;
D, 50 µm; E, 1 mm; H, 200 µm.

Heteroscyphus supinopsis Distribution in New Caledonia. — Only known from the type
[Link], Thouvenot & Frank Müll. specimen in South Province.
(Fig. 22) Total range. — Endemic.

description
Nova Hedwigia 113: 62 (Engel et al. 2021). — Type: New Caledo-
nia. South Province, Mt. Ouin, epiphytic in mossy forest, c. 900 m, Further description and illustrations in Engel et al. (2021).
[Link].2003, Müller NC797 (holo-, DR; iso-, F, hb. Thouvenot). Dioecious.

CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1) 39

 Thouvenot L.

Habit Genus Lophocolea (Dumort.) Dumort.

Plant medium sized, rather rigid, 2.8-3.8 mm wide when
flattened; ventral intercalary branching infrequent; stem Lophocolea bidentata (L.) Dumort
narrow for plant size, 0.15-0.20 mm wide; leaves subop- (Fig. 23)
posite, horizontal grading to obliquely dorsally assurgent,
weakly to moderately convex in dorsal sector, moderately Recueil d’Observations sur les Jungermanniacées 1: 17 (Dumortier
1835). — Jungermannia bidentata L., Species Plantarum: 1132 (Lin-
abaxially concave in ventral sector, loosely imbricate, the naeus 1753). — Type: Great Britain. Zollinger 3613 (OXF), not seen.
dorsal leaf bases connate and forming a laminar lip to c. 6
cells wide. Specimen examined. — New Caledonia. South Province, Hum-
boldt Massif, 1200 m, on rocks, [Link].2008, Thouvenot NC2490
(PC[PC0779842]).
Leaves
1.30-1.80 mm long, 0.80-1.40 mm wide, subsymmetrical, Distribution in New Caledonia. — Newly reported, its distri-
mostly lingulate occasionally ovate to trapezoid, widest near bution in New Caledonia is under-documented.
the insertion; apex variable: broadly rounded to truncate, Total range. — Subcosmopolitan. In the austral Indo-Pacific region,
often retuse, sporadically short bilobed with lobe apices Lophocolea bidentata is reported from the Mascarenes (Ah-Peng & Bar-
rounded to broadly acute, otherwise entire; margins straight dat 2005), Australia (McCarty 2006 as Chiloscyphus latifolius [Nees]
or nearly so, entire. [Link] & [Link].), New Zealand (Glenny 1998 as C. chlo-
rophyllus [Hook.f. & Taylor] Mitt.), Java (Söderström et al. 2010 as
C. coadnatus [Hook f. & Taylor] Mitt.) and New Guinea (Piippo 1985).
Cells
25-35 µm wide, 30-40 µm long, somewhat larger near the
base; in median portion of leaves with knot-like trigones that Description
are separated by narrow, thin-walled places. Based on the New Caledonian specimen.

Underleaves Habit
3.8-6.5 times the stem width, connate with the leaves on both Plant medium sized, in loose mats of interwoven stems with
sides, imbricate, stoutly ovate to subreniform, 0.6-0.9 mm many branches, lateral-intercalary; main stems 0.20 mm wide,
long, 0.6-0.8 mm wide; apex bifid to (0.2)0.35-0.55, the shoots complanate, main shoots 2.00-2.50 mm wide, branches
lobes ± parallel to weakly divergent, acuminate from a broad variously narrower, sometimes strongly attenuated; leaves dis-
base, entire or with a tooth toward base of outer margin; tichous, spreading at right angle, plane, contiguous to slightly
lamina 9-12 cells high and c. 2-5 times wider than long, the imbricated, not connate dorsally, brownish in older main shoots.
margins on each side with 1-5 dentiform to subciliiiform to
occasionnaly laciniiform processes. Leaves
Ovate to oblong-rectangular, 1.20-1.40 mm long, 0.75-0.90 mm
Gametangia wide, margins smooth, entire, apices truncate, bilobed, lobes tri-
Gynoecia seen only in juvenile state, at the end of short leaf- angular at base with acute to piliform apices 1-3 cells long, sinus
less branches, lateral-intercalary, bracts ciliate to laciniate; wide, lunate to right-angled, branch leaves contiguous to remote,
androecia not seen. smaller, with lobes narrower and piliform lobe apices longer.

Comments Cells
Heteroscyphus supinopsis resembles H. supinus of New Zea- Leaf cells rounded to oval, 25-30 µm x 25-45 µm, with small
land and Tasmania and H. deplanchei of New Caledonia. to large nodulous trigones.
Like these species it is easily separated from other New
Caledonian Lophocoleaceae by: 1) leaves usually lingulate Underleaves
with the apices and lateral margins entire; 2) cells with Deeply bifid, disc small, 3 cells high at mid-insertion, 0.15-
conspicuous trigones; 3) large underleaves wider than long 0.20 mm long, 0.20-0.25 mm wide, lobes narrowly lanceolate-
with apices deeply bifid and margins armed with processes linear, piliform apices 3-7 cells long, overall size 0.35-0.50 mm
laciniate to dentiform. Heteroscyphus supinopsis is separated long, 0.40-0.60 mm wide, circa 2 times the stem width, not
from the former by: 1) leaves distinctly connate dorsally; connate to the ventral margins of the leaves.
and 2) underleaves less deeply bifid (to 0.35-0.55 vs 0.8),
underleaf discs longer (9-12 cells high at insertion vs 4-7) Gametangia
and less transversally elongate (2-5 times wider than long vs Not seen.
7). It can be distinguished from the latter by: 1) its smaller
size with shoots up to 3.8 mm when flattened vs 5 mm; Comments
2) the absence of shoot segments with emarginate to bifid This species is newly reported from New Caledonia. Its wide
leaves; 3) shorter lateral processes on the underleaf margins, range might be further extended since several austral species
toothed to ciliate vs long laciniate; and 4) less robust stems have already turned to be synonyms (see above total range)
with c. 40 cells in diameter vs up to 100. and many countries of the Pacific region are still understudied.

40 CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1)

 Lophocoleaceae of New Caledonia

A B C

E
F
G

Fig. 23. — Lophocolea bidentata (L.) Dumort.: A, main shoot, ventral view; B, secondary shoot, ventral view; C, leaf cells; D, G, underleaves; E, leaf apex; F, leaf.
All drawn from Thouvenot NC2490. Scale bars: A, B, 1 mm; D-G, 200 µm; C, 50 µm.

The New Caledonian specimen is sterile and shows a large Schuster 1984 [1985]). — Type: New Caledonia. “Lerat” s.n. (lecto-,
variation in shoot width. It somewhat differs from typical here designated, G[“prope summo Mt. Mou”, XI.1908, Le Rat 210,
L. bidentata only in the small to medium-sized trigones. The G00112967]!; isolecto-, REN[herb. E. G. Paris]!).
material differs from L. bispinosa, a New Zealand species Further specimen examined. — New Caledonia. South Province,
which has similar overall shape and conspicuous trigones but Païta, Humboldt massif, on the trail toward the hut, on rock in
possesses heterogeneous areolation with larger cells scattered cloud forest, [Link].2008, Thouvenot NC3338.
inside the lamina and along the margins. Furthermore, the Distribution in New Caledonia. — Lophocolea caledonica was
latter species has a deeper sinus between longer piliform lobes found at highest elevations in the peridotite massifs of South Province,
and leaf margins with ciliate processes. only known from the type specimen and a single recent collection.
The rarity of Lophocolea bidentata in New Caledonia could be Total range. — Endemic.
linked to the ecological conditions since it was found in one of the
highest localities of the island where it grows with Cryptolophoc-
olea subcostata (Steph.) Thouvenot, Telaranea bisetula (Steph.) Description
[Link]., Trichocolea pluma (Reinw., Blume & Nees) Mont. Dioecious.
and Cheilolejeunea trapezia (Nees) [Link]. & Kachroo.
Habit
Plant small, shoots 1.50 mm wide, leaves narrowly imbricate,
Lophocolea caledonica Steph. spreading, alternate, dorsally free.
(Fig. 24)
Leaves
Species Hepaticarum 6: 267 (Stephani 1922). — Chiloscyphus Widely oval to oblong, 0.70-0.80 mm long, 0.50-0.60 mm
leratii [Link] & [Link]., Nova Hedwigia 39: 418 (Engel & wide, apices usually truncate to retuse, the margins densely

CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1) 41

 Thouvenot L.

spinose-toothed around apices and distal parts of ventral and Lophocolea kurzii Sande Lac., Annales Musei Botanici Lugduno-Bat-
dorsal margins, both leaf surfaces smooth. avi 1: 296 (1864). — Chiloscyphus kurzii (Sande Lac.) [Link] &
[Link]., Nova Hedwigia 39: 417 (Engel & Schuster 1984
[1985]). — Type: Java. Tjiliwong valley, Bogor, 800’, Kurz s.n.
Cells (holo-, L[L0060979, fide Kitagawa in sched. 1970]!) syn. nov.
Leaf cells small, 16-20 µm, with conspicuous nodulose trigones.
Lophocolea fragillima Steph., Species Hepaticarum 6: 273 (Stephani
1922). — Chiloscyphus fragillimus (Steph.) [Link] & [Link].,
Underleaves Nova Hedwigia 39: 415 (Engel & Schuster 1984 [1985]). — Type:
Narrowly decurrent on one side but free from connation with New Caledonia. “Lerat” s.n. (lecto-, here designated, G[“Lerat,
the leaves, obconical, apices deeply bifid with lunate sinuses, Herb. Thériot”, G00112413]!) syn. nov.
lobes triangular lanceolate, lobes and lateral margins with Lophocolea parva Steph., Species Hepaticarum 6: 287 (Stephani
teeth like the leaves. 1922). — Chiloscyphus parvus (Steph.) [Link] & [Link].,
Nova Hedwigia 39: 420 (Engel & Schuster 1984 [1985]). —
Gametangia Type: New Caledonia. Franc (lecto-, here designated, G[“Nova
Caledonia, 1906”, Franc s.n., “Herbier Lacouture”, G00061334!];
Androecia at the end of normal leafy shoots or abbreviated isolecto-, PC[PC0102407!, PC0102408!]) syn. nov.
leafless branches, ventral-lateral, with 3-5 pairs of bracts,
rectangular, spinose-toothed at apices, the bases of the dorsal Lophocolea papulimarginata [Link], Phytologia 47: 323 (Miller
margins recurved and saccate; gynoecia at the end of normal 1981). — Lophocolea papulosa Steph., nom. illeg. Species Hepati-
carum 6: 286 (Stephani 1922). — Chiloscyphus papulimargina-
leafy shoots, 2 pairs of bracts similar in shape to normal tus ([Link].) [Link] & [Link]., Nova Hedwigia 39: 420
vegetative leaves but larger, 1.10 mm long, 0.60 mm wide; (Engel & Schuster 1984 [1985]). — Type: New Caledonia.
bracteoles smaller, 0.50-0.70 mm long, 0.35-0.40 mm wide, “Lerat” s.n. (lecto-, here designated, G[“Ad arbores in summo
ovate, 1/2 or more bifid, sinus narrow, lobes oblong to lan- Mt. Mou (1224 m)”, I.1906, Le Rat 44, G00112490!]; isolecto-,
ceolate, often divided in secondary lobes, the upper margins REN[herb. E. G. Paris]!; syn-, G[Pic des Sources, s.d., Le Rat 237,
G00051485!, Mt. Dzumac, IV.1907, Le Rat 225, G00051484!,
toothed-laciniate; perianth elongate, basal half cylindrical, idem, Le Rat 232, G00051487!, “Île des Pins, Forêt de Uapan”,
upper half trigone, not winged, mouth trilobed, sinus depth V.1909, L. Le Rat 68, G00051486!], isosyn-, REN[“Île des Pins”,
1/3 the length of the whole perianth, lobes oval to lanceolate, V.1909, L. Le Rat s.n., herb. E. G. Paris]!) syn. nov.
subdivided by several deep cuts, margins sharply and densely
toothed, the surface of the perianth sparsely bristled with Lophocolea pilistipula Steph., Species Hepaticarum 6: 288 (Stephani
1922). — Chiloscyphus pilistipulus (Steph.) [Link] & [Link].,
short spikes, usually 1 (-2) cell long. Nova Hedwigia 39: 421 (Engel & Schuster 1984 [1985]). — Type:
New Caledonia. “Lerat” s.n. (lecto-, here designated, G[“Île des
Comments Pins, forêt de Watchia, à la baie de Oupe”, V.1909, L. Le Rat 55,
Lophocolea caledonica is a very distinct, elegant species with G00112487]!; isolecto-, REN[herb. E. G. Paris]!) syn. nov.
all leaves and underleaves finely spinose-toothed. Unfortu- Further specimens examined. — India. Shambaganur, II.1932,
nately, the type specimen is sterile as underlined by Stephani Foreau 3075 as Lophocolea kurzii (PC[PC0167666]).
(1922). In the herbarium of E. G. Paris (REN), the author Java. “Ad Arengae sacchariferae truncos in agro Buitenzorgensis”, 250 m,
found part of the original specimen examined by Stephani [Link].1908, Schiffner 696 as Lophocolea kurzii (PC[PC0763155]).
with well-developed gametangia. The position of the gynoecia New Caledonia. South Province, Sarraméa, on the treck from
Sarraméa to the Dogny Plateau, epiphyte, c. 170 m, [Link].2003,
at the end of normal leafy shoots confirms its assignment F.Müller NC799 (DR); Nouméa, Tina, border between dry forest
to Lophocolea, together with vegetative characters, i.e., and mangrove, 0-2 m, [Link].2012, Thouvenot NC2395; Farino,
underleaves free, bifid, leaf apices emarginate. This species Grandes-Fougères Natural Park, Creek Houé, on dead wood in a
resembles the Australasian species of the Lophocolea section mesophylous forest, 370 m, [Link].2016, Thouvenot NC2448; Mont
Microlophocolea Spruce (Engel 2010 as Chiloscyphus subg. Dore, Demazures forest, on dead wood in wet forest, 330-420 m,
[Link].2016, Thouvenot NC2466; Mont Dore, Yahoué, on trunk in
Microlophocolea) especially L. muricata (Lehm.) Nees (see the mesophilous forest along the river, 122 m, 10.X.2019, Thouvenot
below). They share: 1) sizes of shoots, leaves, underleaves and NC2756; North Province, Pouhembout, Forêt Plate, near Ouendé
cells; 2) leaf and underleaf shapes; and 3) margin ornamenta- Falls, 300 m, road embankment, [Link].2003, F.Müller NC815;
tions. Lophocolea caledonica differs from all the Australasian South Province, Mt. Dzumac, IV.1907, Le Rat s.n. as Lophocolea
papulosa (G[G00051484, G00051487]); Pic des Sources, Le Rat s.n.
species of this section in the leaves smooth on both faces “fragments” as Lophocolea papulosa (G[G00051485]); “Île des Pins,
instead of armed with numerous conspicuous teeth on one forêt de Wapan”, V.1909, L. Le Rat s.n. as Lophocolea papulosa
or both surfaces. (G[G00051486]); Nouméa, Ouen Toro, [Link].1950, Hürlimann
2038a as Lophocolea papulosa (PC[PC0167657]); Mé Aoui, 500 m,
[Link].1951, Guillaumin & Baumann-Bodenheim 10375 as Lophocolea
papulosa (PC[PC0167656], GOET); Boulari, 200 m, [Link]. 1950,
Lophocolea convexula Mitt. Hürlimann 2015 as Lophocolea parva (G[G060814]); Hürlimann 2017
(Figs 25; 26) as Lophocolea parva (G[G060826]); ex herbarium Bonati, Franc s.n.
determinavit Stephani as Lophocolea fragillima (PC[PC0167654]);
Flora Vitiensis: 405 (Mitten 1871 [1873]). — Chiloscyphus con- North Province, “ad radices Mt. Panié”, II.1910, Le Rat s.n. “frag-
vexulus (Mitt.) [Link] & [Link]., Nova Hedwigia 39: 413 ments” as Lophocolea autoica (G[G00128109]); South Province,
(Engel & Schuster 1984 [1985]). — Type: New Caledonia. Isle of Mts. Koghis, [Link].1914, Compton 801 as “Lophocolea levieri”
Pines, Strange s.n. (holo-, NY[NY00965719!]). (BM[BM013409501, BM013409504]); Mt. Mou, on stone and

42 CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1)

 Lophocoleaceae of New Caledonia

A C
B

F
E

I
G
H

J L

K
M

Fig. 24. — Lophocolea caledonica Steph.: A, shoot portion, ventral view; B, midleaf cells; C, leaf; D, androecium, lateral view; E, H, underleaves; F, leaf angle with
teeth; G, male bract; I, perianth (torn); J, female bract; K, female bracteole; L, M, spines on the perianth surface. A, E, H, drawn from the lectotype (G00112967);
B-D, F, G, I-M, from the isotype (REN). Scale bars: A, I-K, 1 mm; B, F, L, M, 50 µm; C-E, G, H, 0.5 mm.

mud in stream, [Link].1914, Compton 443 as “Lophocolea levieri” secund, dorsally free; both terminal and lateral-intercalary
(BM[BM013409502]). branching present.
Distribution in New Caledonia. — The most frequent member
of Lophocoleaceae at low altitude, in both provinces of the main Leaves
island, including Isle of Pines; it grows on various substrates on the Ovate-oblong, 0.55-1.00 mm long, 0.40-0.80 mm wide, leaf
ground, rarer on barks of living trees, at low to medium elevations margins crenulate due to bulging marginal cells, leaf apices
(collected from 0 to 900 m). uneven on a same shoot, ranging from widely rounded or
Total range. — Java (Söderström et al. 2010 as Lophocolea kur- obtuse to retuse with widely concave sinus and rounded angles,
zii), Fiji (Söderström et al. 2011 as Chiloscyphus parvus), Malaysia possibly with one short acute tooth, other times leaf apices
(Chuah-Petiot 2011 as Lophocolea kurzii), Thailand (Lai et al. 2008 very shortly bifid with widely triangular lobes.
as Lophocolea kurzii).
Cells
Description Hexagonal, 16-36 µm, thin-walled, trigones very small, acute,
Paroecious, more rarely autoecious. free walls bulging.

Habit Underleaves
Plants small to medium, shoots 1.00-2.00 mm wide; leaves Small, free or very narrowly connate to one of the nearest
alternate, patent, dorsally assurgent, individually convex and leaves, 1-2 times the stem width, insertion in a deep sinus,

CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1) 43

 Thouvenot L.

A B C

G
E F H

I K

O
N
M P

R
S

Fig. 25. — Lophocolea convexula Mitt.: A, shoot portion, dorsal view; B, C, K, L, leaves; D, median cells; E, G, N-P, underleaves; F, leaf lobe; H, marginal cells; I,
J, shoot portions, ventral view; M, perianth with a bract; Q, leaf apex, strongly crenulate margin; R, female bract; S, female bracteole. A-H, drawn from the holo-
type of Lophocolea kurzii Sande Lac. (L0060979); I-L, N-S, from the lectotype of Lophocolea fragillima Steph. (G00112413); M, from the lectotype of Lophocolea
papulimarginata [Link] (G00112490). Scale bars: A, I, J, M, R, S, 1 mm; B, C, E, G, K, L, N-P, 200 µm; D, F, H, 50 µm; Q, 100 µm.

apices deeply bifid, lobes narrowly lanceolate, discs shorter on both sides, more rarely similar to the lobes, then with one
than wide, lateral margins with a single short to filiform tooth more tooth.

44 CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1)

 Lophocoleaceae of New Caledonia

A B C D E

H J
F
G
I

L
M
N O

U
S
Q
P T

R
V

W
Y

Fig. 26. — Lophocolea convexula Mitt.: A, G, P, shoot portion, ventral view; B, leaf and underleaf; C-E, J, S-U, leaf apices; F, H, I, N, O, R, V, underleaves; K,
marginal cells; L, autoicous gametangia: perianth terminal with one pair of female bracts and four pairs of male bracts below, dorsal view; M, Q, leaves; W,
adjacent leaves on stem; X, apex margin; Y, median cells. A-F, drawn from the lectotype of Lophocolea pilistipula Steph. (G00112487); G-K, from the lectotype
of L. papulimarginata [Link] (G00112490); M-U, from the lectotype of Lophocolea parva Steph. (G00061334); L, V-Y, from the isotype of Lophocolea parva
(PC0102408). Scale bars: A, G, L, P, W, 1 mm; B, H, I, M-O, Q, R, 200 µm; C-F, J, S-V, 100 µm; K, X, Y, 50 µm.

Vegetative reproduction Gametangia

Gemmae lacking, but regenerants occur on perianth mouth Gynoecia at the apices of main shoots or long branches with
and bract margins in some specimens. normal vegetative leaves, bracts oblong, variously and shortly

CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1) 45

 Thouvenot L.

toothed, bracteoles bifid to the middle, lateral margins vari- of E. G. Paris (REN), but a lot of new species names, here
ously and shortly toothed; perianths oblong, 2-3 mm long, considered as synonyms, were given by Stephani to the
with cupulate-ovate base and trigone upper part, plicae nar- New Caledonian material sent by E. G. Paris, on the basis
row, acute, not winged, mouth deeply 3-fid to half the whole of different forms of characters, such as leaf and underleaf
perianth length or more, the lobes narrowly ovate-triangular, apices, cell width, leaf and perianth sizes. Careful examina-
strongly toothed to laciniate; androecia beneath the gynoecia, tion shows that these characters are variable even in a same
in series of 3-5 pairs of male bracts smaller than the female specimen. Consequently, Stephani invented new species
and saccate at dorsal base. which turned to be the same, when examined together. All
the New Caledonian names, being published after the name
Comments of L. convexula, fall in synonymy with the latter, so that, the
In addition to the plant described by Mitten as Lophocolea author assume that it might be more widely distributed in
convexula, this group of taxa includes other New Caledonian southern tropics.
plants described by Stephani (1922) as distinct species but The main differences seen in Stephani’s diagnoses and draw-
very similar in most respects and similar to the South Asian ings or found in the type material of the New Caledonian taxa
L. kurzii. They share the following common features: 1) small concern the perianth shapes, oblong in L. parva, cupulate in
size and dorsally convex secund leaves; 2) ovate leaves wider L. papulosa, cyathiform in L. convexula and obovate-conical
at base or at small distance above, decreasing to the apices in L. fragillima. This aspect is difficult to evaluate on some
in curve lines; 3) apices variable in a same shoot, typically old specimens, hardly fertile or with broken organs. However,
entire or retuse, rarely or frequently shortly bifid with wide all other gynoecial characters being similar or included in a
triangular lobes; 4) leaf margins crenulate; 5) underleaves short range of continuous variation, this diversity may be
small, hardly wider than the stems, deeply bifid, usually with resumed in a length-width ratio ranging from 4 (L. parva),
one small tooth on both sides; 6) usually paroecious; and 7) 3-2.5 (L. fragillima) to 2 (L. papulosa). The conservation
perianths deeply 3-fid, triangular lobes sharply toothed or requirements of old specimens, especially if they are types,
laciniate. As stressed by many authors (Sande Lacoste 1864; prevent to provide sufficient data on size and ornamenta-
Schiffner 1900; Pearson 1922; Kitagawa 1973; Gradstein tion of the gynoecia and to assume a continuous variability
2011) these plants look like Lophocolea heterophylla (Schrad.) in perianth shapes. However, the observed differences seem
Dumort., a species widely distributed in the Northern likely the result of various stages of maturity. In vegetative
Hemisphere and very rare in the tropics where it has been features, the main changes are the underleaves and leaf orien-
reported from Cuba and ranges southwards to Brazil in tation. Underleaves are variable in a same specimen or even
the Southern Hemisphere (Gradstein & Pinheiro da Costa in a same shoot, the lobes being sometimes shorter without
2003). Besides this geographic separation, L. heterophylla filiform apex, their orientation varying from erect incurved
may be distinguished by: 1) a leaf shape rather rectangular, to spreading at an angle of 30-45°. Leaf arrangements are
not so decreasing to the apex; 2) some or many leaf apices contiguous to slightly imbricate and obliquely patent in
bilobed, the lobes longer and sharper, separated by a deeper L. fragillima, or conspicuously imbricate and patent at right
and wider sinus; 3) leaf margins entire, free cell walls flat; angle in L. parva, less conspicuously so in further species.
4) perianths less deeply 3-fid, the lobes truncate, rounded However, dimensions are comparable in all the types and
to widely triangular, unevenly toothed; and 5) gemmae fresh specimens measured. Thus, morphological differences
occasional and regenerants unknown. It would have been are insufficient to separate New Caledonian specimens as
interesting to compare Brazilian material (not seen) with distinct taxa.
L. convexula. Schuster (1980) describes L. heterophylla subsp. For lectotypification purpose of Lophocolea pilistipula, the
cladogyna [Link]., a subspecies with some specific specimen G00112487 from Pines Island is selected here since
characters of L. convexula namely the rounded leaves usually it perfectly matches the original description and belongs to
entire. But the subspecies of L. heterophylla usually produce the original materiel examined by Stephani as shown by the
gemmae, never regenerants (Paton 1999); in contrast, such handwritten name and mentions “[Link].” and “c. per.”. The
cell clusters (regenerant) are found on leaves, bracts and original description includes perianth characters and the
perianth margins in some specimens of L. convexula col- selected lectotype is the single fertile specimen among the
lected in New Caledonia (e.g. NC2756), but gemmae have three type specimens kept at G. The remaining material at G,
never been observed. from Dogny, is sterile and larger in all the dimensions. They
The holotype of Lophocolea convexula is fertile and shows belong to another species described as Heteroscyphus parapil-
paroecious sexual condition. As it is the earlier name among istipulus (see above). The original materials of L. fragillima
the many species described with this set of vegetative and and L. parva at G each comprise a single packet handwrit-
sexual characters, we assume this name must be used for these ten “[Link].” by Stephani and are the only ones available for
plants in New Caledonia as well as in the Asian and South lectotypification. Among five packets of L. papulosa available,
Pacific regions where L. kurzii was reported. the lectotype selected have the label marked “original”, and
In New Caledonia, a single report of Lophocolea kurzii is “autoica, c. per.” handwritten by Stephani. This is also the type
from Paris (1908) based on a specimen collected by Le Rat designated by Miller (1981) when he replaced its epithet by
in Loyalty Islands. This specimen is lacking in the herbarium papulimarginata.

46 CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1)

 Lophocoleaceae of New Caledonia

B
A

Fig. 27. — Lophocolea muricata (Lehm.) Nees: A, cells from lobe apex (top) to median portion of leaf; B, leaf margin in dorsal view, with focus on the echinae;
C, shoot portion, dorsal view. All drawn from Hürlimann 2090 (Z007-1972) (from photos by H. Hofmann). Scale bars: A, B, 20 µm; C, 100 µm.

Lophocolea muricata (Lehm.) Nees Habit

(Fig. 27) Plants soft, shoots up to 1 mm wide, branches common,
stems smooth; leaves alternate or subopposite, usually sub-
Synopsis Hepaticarum: 169 (Gottsche et al. 1845). — Jungermannia
muricata Lehm., Linnaea 4: 363 (Lehmann 1829). — Chiloscyphus horizontal ± dorsally assurgent in basal portions, imbricate,
muricatus (Lehm.) J.J. Engel & [Link]., Nova Hedwigia 39: the base of dorsal margins reaching the midline of the stem
419 (Engel & Schuster 1984 [1985]). — Type: South Africa. In but not fused.
parietibus saxorum in vertice montis Tafelberg, Cape of Good Hope,
Ecklon s.n., not seen.
Leaves
Reference specimen examined. — New Caledonia. South Province, Ovate-oblong, shortly bifid to 0.15-0.20, the lobes trian-
Mts. Koghis, SE ridge of Mt. Bouo, on bark in meso-hygrophilous
forest, 830 m, [Link].1950, Hürlimann 2090 (Z[007-1972]).
gular, medium acute; dorsal leaf surface hispid, except near
ventral base, armed with 1-3-celled erect, acute echinae,
Distribution in New Caledonia. — South Province. Hitherto one per cell, half ventral leaf surface armed with scattered
only known from a single collection by Hürlimann (1998). 2-4-celled similar echinae; lateral and lobe margins copi-
Total range. — Pantropical-South temperate. ously spinose-dentate to ciliate-dentate, the teeth 2-3(4)
cells long.
Description
Further description and illustrations in Fulford (1976) or Cells
Engel (2010 as Chiloscyphus muricatus). Leaf cells 14-18 µm wide, 17-20 µm long, walls moderately
Autoecious. thickened, trigones small to medium.

CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1) 47

 Thouvenot L.

Underleaves right angle, slightly convex, alternate but slightly so, seem-
1.4-1.6 times the stem width, narrowly connate on one side, ing subopposite, dorsally free, dorsal insertion separated by
mostly cuneate to long rectangular, bifid to 0.74 or more, lobes four cell rows.
diverging or subparallel, long linear, margins and lobes ciliate.
Leaves
Gametangia Oblong to trapezoid, margins nearly parallel below middle
Gametangia terminal or intercalary on main shoots or ± long then width progressively decreasing in upper part or ventral
normal leaved branches, androecia in short spikes, bracts margin nearly orthogonal to the stem and dorsal margin
much smaller than leaves, strongly ventricose; except the oblique, 1.00-1.20 mm long, 0.60-0.90 mm wide near base,
reflexose distal portion, bifid and ornamented as in the leaves; 0.35-0.45 below apex, leaf apices truncate, flat to slightly
antheridial stalk uniseriate; gynoecia with innermost bracts concave with a single tooth at both angles, acute, (1-)3(-4)
much larger than leaves, elliptic convex, short bifid, with both cells long, leaf margins entire.
surfaces armed as in the leaves, lobes broadly acute, margins
copiously spinose-dentate or ciliate-dentate; innermost brac- Cells
teoles smaller, oblong to narrowly ovate, bifid to ± 0.4-0.5, Hexagonal, (25-)30-40 µm, thin-walled, without trigones.
margins ciliate; perianth long exserted, nearly cylindrical,
somewhat dorsally compressed, ± trigonous when young, the Underleaves
surface densely hispid with echinae as in the leaves, perianth Small, as wide as stems, narrowly connate to both nearest
mouth 3-lobed, lobes shallowly bifid, the segments acute to leaves, connation 1-3 cells wide, insertion in an inverted U,
subacuminate, lobe margins spinose dentate; calyptra small, apices deeply bifid with lunate sinus, discs arched, narrow,
included within periant at maturity. usually two cells high, nine cells wide, lobes linear lanceolate,
each lateral margins with a single short to filiform tooth.
Comments
Lophocolea muricata is easily distinguished from all other New Gametangia
Caledonian species by the acute echinae set on both dorsal Male and female gametangia set at apices of main shoots or
and ventral leaf surfaces. Otherwise, it could be confused long branches with normal leaves; gynoecia with two pairs of
with Lophocolea caledonica with which it shares size, colour, bracts, bract shapes like the normal leaves but larger, 1.8 mm
leaf and underleaf shapes, but the latter has both leaf surfaces long, ventral margins with one tooth, bracteoles oblong, mar-
smooth, the spinose papillae being restricted to the perianth gins toothed with 3-4 strong teeth in upper part; perianths
surface, and leaf cells with conspicuous medium trigones vs oblong, cupulate at base, trigone in upper part, not winged,
trigones lacking or very small in the former. The single refer- mouth shallowly 3-lobed, the lobes strongly toothed, acute
ence specimen reported from New Caledonia (Hürlimann teeth triangular to lanceolate; androecia with up to eight
1998) could be checked thanks to photos and observations pairs of bracts.
from Heike Hofmann (Z+Zt). According to the original
diagnosis (Lehmann 1829) and further descriptions (Fulford Comments
1976; Engel 2010), the organisation and morphology of the The type specimen at G is copiously fertile with both kinds
gametangia are characteristic of the genus Lophocolea. of gametangia and the dioecious condition is likely since
no connection between the male and female shoots could
be noted. Superficial view could lead to assign this species
Lophocolea savesiana Steph. to the genus Heteroscyphus considering the shape and nearly
(Fig. 28) subopposite insertion of the leaves and it may be closer to
Heteroscyphus etesseanus which has similar leaves, underleaves
Species Hepaticarum 3: 122 (Stephani 1906). — Chiloscyphus and cells, but the gametoecia locations and shapes separate
savesianus (Steph.) [Link] & [Link]., Nova Hedwigia 39: 422
(Engel & Schuster 1984 [1985]). — Type: New Caledonia. Savès s.n. them as well as some vegetative characters, H. etesseanus
(lecto-, here designated, fide Kitagawa 1969 in sched. G[Nouméa, having smaller size, with shoots 1.50 mm wide when flat-
1887, Savès s.n., G00061342]!). tened, and leaves contiguous to distant. Lophocolea savesi-
ana has gametangia as in Lophocolea, set at the end of long
Distribution in New Caledonia. — Only known from the type
(“Nouméa”). normal-leaved shoots, oblong perianths that are conspicu-
ously trigonous in upper part, and androecial bracts smaller
Total range. — Endemic. than normal leaves, in series of 8 pairs. The species may be
compared with L. convexula, but the latter is smaller with
Description ovate leaves that are strongly narrowed to the apex, and
Dioecious. with bulging leaf cells making the leaf margins crenulate.
The species also resembles Heteroscyphus parapilistipulus but
Habit the latter species is a larger plant with shoots up to 4.5 mm
Medium plants with shoots 2.50 mm wide when flattened; wide, leaves rather oval and gynoecia and androecia on short
leaves imbricate, 1/3 overlapping, horizontally spreading at leafless lateral branches.

48 CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1)

 Lophocoleaceae of New Caledonia

A B

E G

Fig. 28. — Lophocolea savesiana Steph.: A, shoot portion, ventral view; B, adjacent leaves, dorsal view; C, underleaves; D, top part of a male shoot, dorsal view;
E, perianth; F, H, teeth at leaf apex angles; G, median cells. All illustrated from the lectotype (G00061342). Scale bars: A, B, D, E, 500 µm; G, 50 µm; C, F, H, 100 µm.

1.X.2008, Thouvenot NC1730; Mt. Ouin, on dead wood in moun-

Genus Otoscyphus [Link], Bardat & Thouvenot tain wet forest, 1000-1025 m, [Link].2016, Thouvenot NC2098;
Dzumac massif, on dead wood in mesophilous mountain forest,
Otoscyphus crassicaulis 915 m, [Link].2008, Thouvenot NC486; Yaté, base of Pic du Grand
Kaori, on peridotite rock, 250 m, 22.X.2012, Thouvenot NC736;
(Steph.) [Link], Bardat & Thouvenot Wé Toa, on dead wood, in rain forest, 500 m, [Link].2019, Thouvenot
(Fig. 29) NC2872 (PC[PC0779846]); Mont Dore, Mouirange, on dead wood
in Nothofagus rain forest, 546 m, [Link].2019, Thouvenot NC2883.
Cryptogamie, Bryologie 33 (3): 280 (Engel et al. 2012). — Lophocolea
crassicaulis Steph., Species Hepaticarum 6: 268 (Stephani 1922). — Distribution in New Caledonia. — Fairly frequent on dead hard
Chiloscyphus crassicaulis (Steph.) [Link] & [Link]., Nova wood, more rarely on rocks, in various woodland habitats on ultra-
Hedwigia 39: 413 (Engel & Schuster 1984 [1985]). — Type: New mafic soils, from lowland photo-xerophytic forests to wet mountain
Caledonia. Franc s.n. (lecto-, G[Mt. Mou, 1200 m, [Link].1909, forests or scrublands, at all altitudes (collected from 250 to 1400 m).
Franc s.n., G00017607]!).
Total range. — Endemic to the southern ultramafic massif of
Further specimens examined. — New Caledonia. South Province, New Caledonia, South Province.
Dumbéa, Montagne des Sources strict nature reserve, on dead wood
in mountain wet forest with Araucaria rulei, 950 m, [Link].2016,
Thouvenot NC1992; Païta, Mt. Humboldt, on peridotite rocks Description
in mountain scrubland with Araucaria humboldtensis, 1400 m, Further description and illustration in Engel et al. (2012).

CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1) 49

 Thouvenot L.

Dioecious. dorsally assurgent, densely imbricate, strongly ventricose in

basal ± 0.5, the bracts deeply bilobed, the lobes similar to
Habit those of the leaves except shorter; antheridia solitary, the
Plants delicate, shoots up to 0.6-1.0 mm wide; branching stalk not seen; gynoecia terminal on main shoot, or on short
occasional, exclusively of ventral-intercalary type, Frullania- ventral-intercalary branches lacking normal vegetative leaves;
type branches rare; stems zig-zag in growth, markedly narrow bracts bifid to ± 0.35-0.5, the dorsal lobe smaller or reduced
for shoot size; leaves alternate, with basal sector vertically dor- to a small rounded projection; bracteoles of innermost series
sally assurgent and defining an adaxial concavity, the leaf then somewhat smaller than bracts, symetrically bifid to ± 0.25-
strongly convex-arched, in cross section forming a ± inverted 0.5; perianths long exserted, 1.7-2.2 mm long, subterete
U or, with further curvature of the lobes, even a semicircle; at toward base, obscurely trigonous above, oblong-elliptic, not
the base, the vertical sectors of opposing leaves fused. or slightly narrowed toward the deeply 3-lobed mouth, the
lobes subequal, free for ± 0.30-0.35 the perianth length; lobes
Leaves deeply bifid with several often long ciliiform processes; keelar
0.37-0.38 (0.40) mm long, bilobed, each leaf with dorsal base wings common, 1-2 per perianth; calyptra extending ± 0.4
abruptly and strongly dilated resulting in a conspicuous flange the perianth length.
of tissue extending towards shoot base at least to the level of
the median sector of the leaf immediately below; leaves with Comments
a pronounced adaxial concavity toward base that is totally This small species is easily separated from all other New
covered by the gently convex decurrent flange of tissue, the Caledonian liverworts by the combination of the following
concavity pocket-like; the exposed, main portion of the leaf characters: 1) leaves rectangular with apices deeply bifid, lobes
convex, narrowly elongate sub-rectangular, the sides from the long linear; 2) dorsal leaf margins widely expanded, forming
base to lobe tip gently incurved, the leaves bifid to 0.40-0.45 a pouch together with the underlying ventral margins of the
of leaf length; lobes subparallel to ± divergent, subequal in adjacent leaves; 3) underleaves transversely elongate, bifid,
size, long ciliform-setaceous; leaf disc 15-17 cells wide at base, with sinus at a flat angle and long linear lobes widely spread-
the median sector 4-6 cells wide, the dorsal margin plane, ing, lateral margins very short, entire; 4) stems thin; and 5)
entire, the ventral margin plane, with a conspicuous tooth branching nearly exclusively ventral-intercalary. As discussed
near the base, this tooth asymmetrical, curved towards the by Engel et al. (2012), this species is close to Chiloscyphus
stem and forming an orbicular mouth-like opening for the but differs from that genus by the above-mentioned set of
pocket-like leaf base. morphological traits. Notably, the underleaves of Otoscyphus
crassicaulis are very unusual and resemble those found in
Cells Drepanolejeunea and Leptolejeunea (Lejeuneaceae), while the
Median leaf cells 25-30 μm wide, 35-50 μm long; cells of the pouches made of superimposed bases of leaves are unique in
exposed main portion of the leaf with massive trigones that liverworts.
are confluent or separated by narrow thin-walled places; cells Following Engel et al. (2012), who described the type at G,
in basal 4-5 rows of cells and flange with trigones minute to the lectotype is formally designed here. The relationships of
small; marginal cells with exposed wall notably thick; surface the genus Otoscyphus are unclear and await molecular study
of disc finely striate, of lobes finely papillose. (Söderström et al. 2013a).
Underleaves Incertae sedis
Free, distant, the stem broadly exposed; disc a little wider
than stem but with lobes laterally extending at least to level
Genus Chiloscyphus Corda
of sinus bases of the adjacent leaves; disc cunate, bifid nearly
to base, small, with a distal row of 2-3 larger cells with the
Chiloscyphus longifissus Steph.
exposed wall distinctly thickened, basal to this row is a feebly
(Fig. 30)
dome-like cluster of c. 8 smaller, quadrate, rhizoid initials, the
summit of disc truncate or (often) broadly rounded, entire, Species Hepaticarum 6: 310 (Stephani 1922). — Type: New
the lateral margins entire or at times with a 1-2-celled tooth, Caledonia. Franc s.n. (lecto-, here designated, G[env. de Tao,
the disc margins decurrent; lobes diverging by an angle of 100 m, I.1910, Franc s.n., hb. Thériot 225, G00069456]!; isolecto-,
c. 180° or nearly so, the line formed by lobes and disc apex PC[PC0101940]!).
maximally lunate to straight, the lobe shape much like those
Chiloscyphus longifissus Steph. ex Paris, Revue bryologique 37: 129 (Paris
of the leaves but comprised of 1-3 biseriate tiers and a uni- 1910), nom. inval. (no description). — Reference specimen: New
seriate row of 3-6 cells, the lobe margins uniformly entire. Caledonia. “In jugo Dogny (1050 m)”, VII.1909, L. Le Rat s.n.
(REN[herb. E. G. Paris]!; duplicate, PC[PC0103808]!) syn. nov.
Gametangia
Chiloscyphus beesleyanus Pearson, Journal of the Linnean Society, Botany
Androecia terminal but becoming intercalary in position 46: 22 (Pearson 1922). — Type: New Caledonia. Ignambi, on rocks
on main shoot or rather long ventral-intercalary branches, by creek, 3000 ft., Compton 1530 (holo-, BM[BM013409500]!)
somewhat narrower than sterile sectors; bracts in 2-4 pairs, syn. nov.

50 CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1)

 Lophocoleaceae of New Caledonia

B C D

E G
F
ddb

c
lb

ul

Fig. 29. — Otoscyphus crassicaulis (Steph.) [Link], Bardat & Thouvenot: A, habit; B, shoot portion, ventral view; C, shoot portion, dorsal view showing dilated
dorsal leaf bases overlapping the previous leaf ventral bases (SEM); D, gynoecium; E, leaf; F, underleaf; G, shoot cross section showing connation (c) of dilated
dorsal leaf bases (ddb), the concave sector of leaf base (lb), the stem (s) and the underleaf (ul). All illustrated from the specimen Thouvenot NC486 (photos
J. Bardat, drawings J.J. Engel). Scale bars: B, C, 200 µm; D, 500 µm; E-G, 100 µm.

Distribution in New Caledonia. — Scattered in North and Further specimens examined. — New Caledonia. North Province,
South Provinces, in wet forest from 100 to 1200 m, rarely collected. “inter Panié et Hienghène”, II.1910, Le Rat s.n. (REN); South Province,
“in jugo Dogny (1050 m)”, VII.1909, L. Le Rat s.n. (REN); North
Total range. — Endemic. Province, Tao, forest, 600-800 m, I.1910, Franc s.n. as “Lophocolea

CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1) 51

 Thouvenot L.

latistipula” (PC[PC0102424]); South Province, Pic des Mousses, checked again and confirmed to be identical to Chiloscyphus
Mt. Mou, 1200 m, [Link].1950, Baumann-Bodenheim 5704 longifissus Steph. During the examination of the samples present
(GOET); South Province, “Inter Farino et Table Unio”, VII.1909,
Le Rat s.n. as “Chiloscyphus Le Rati” Steph. nom. herb. (REN).
in the three herbariums, the author found some more misiden-
tifications. Some specimens of C. longifissus, collected by Franc
in Tao in January 1910, likely part of the type material, were
Description labelled Lophocolea latistipula Steph., and one of them kept in
Habit G as a type of the latter (G00112472), with a duplicate in PC
Plants large, with shoots up to 6.00 mm wide; leaves spread- (PC0102424) (see comments under Heteroscyphus coalitus).
ing, subopposite to moderately staggered, dorsally free. The checked type of Chiloscyphus beesleyana is not different
from the type of C. longifissus.
Leaves
Oval-oblong to trapezoid, sizes uneven in successive segments Species transferred to other families
of a same shoot, 1.50-3.00 mm long, 1.00-1.50 mm wide
near the base; leaf apex c. 1/2 the base width, truncate to widely Family Acrobolbaceae [Link].
concave, with a single tooth at both angles, teeth short and Genus Conoscyphus Mitt.
acute, margins otherwise entire.
Conoscyphus trapezioides
Cells (Sande Lac.) Mitt. ex Schiffn.
Hexagonal, 40-80 µm with strong trigones bulging to truncate.
Conspectus Hepaticarum Archipelagi Indici 125 (Schiffner 1898). —
Underleaves Chiloscyphus trapezoides Sande-Lac., Nederlandsch Kruidkundig
Archief. Verslangen en Mededelingen der Nederlandsche Botanische
Asymmetrically connate to the adjacent leaves, one side nar- Vereeniging 3: 417 (Sande Lacoste 1855). — Type: Java. s. dat.,
rowly connate, the opposite side connate or narrowly decur- Junghuhn s.n., not seen.
rent onto the respective leaf, underleaves widest than long,
deeply bifid, 1/2-3/4 the whole length at stem insertion, sinus Specimen examined. — New Caledonia. South Province, near
Sunshine Mine, on trunk in rain forest, 700 m, [Link].1951, Hür-
lunate to acute, lobes triangular acuminate, spreading at right limann 2426 (G).
to obtuse angle from each other, discs transversely elongate,
lateral margins with 1-2 sharp teeth. Distribution in New Caledonia. — Known from a few col-
lections in South Province where it grows on barks in rain forests
(collected from 700 to 950 m).
Gametangia
Androecia in sets of up to nine pairs of bracts, terminal or Total range. — Paleotropical: Africa, Indomalaya, Melanesia,
intercalary on main shoots or on long leafy branches; gynoe- South Pacific.
cia not seen.
Description and illustrations
Comments See Piippo (1985).
Gametangia are lacking in the type specimen at G which
seems sterile even though Stephani underlines the presence of
androecia (“androecia parva cauligena…”). The isolectotype Genus Goebelobryum Grolle
in PC, however, contains several male shoots with series of
bracts terminal or intercalary. Therefore, the genus Chiloscyphus Goebelobryum unguiculatum
might be appropriate if the shape of the bracts would not be (Hook.f. & Taylor) Grolle
so different from the leaves, as in the genus Lophocolea, and
the subopposite leaves ventrally connate to the underleaves Journal of the Hattori Botanical Laboratory 25: 137 (Grolle 1962). —
which are conspicuously wider than the stem, reminiscent Jungermannia unguiculata Hook.f. & Taylor, London Journal of
of Heteroscyphus. The combination of these features is more Botany 5: 279 (Taylor 1846). — Type: New Zealand. Hooker 258
consistent with Cryptolophocolea, as defined in Söderström (FH), not seen.
et al. (2013a) but the lack of gynoecium and the impossibility Lophocolea purpurea Steph., Species Hepaticarum 6: 289 (Stephani
of checking the thickness of antheridia stalks prevent alloca- 1922). — Chiloscyphus novae-caledoniae [Link] & [Link].,
tion to a genus and it is necessary to keep the current name Nova Hedwigia 39: 419 (Engel & Schuster 1984 [1985]). —
pending the availability of fertile female material. Type: New Caledonia. “Lerat” s.n. (lecto-, here designated,
G[“In jugo Dogny (1040 m)”, L. Le Rat 211 bis, “General Paris
On the other hand, original specimens collected by Le Rat are misit”, G00112486]!; isolecto-, PC[PC0102405]!; REN[herb.
kept in the herbarium of E. G. Paris (REN) and can be reference E. G. Paris]!) syn. nov.
vouchers for Chiloscyphus longifissus Steph. ex Paris. The dupli-
cate seen at PC was previously misidentified as Heteroscyphus Further specimen examined. — New Caledonia. South Prov-
ince, Mt. Mou, on shady bare soil in trail side across shrubland in
grandiflorus so that Chiloscyphus longifissus Steph. ex Paris was ultramafic massif, 750-850 m, [Link].2016, Thouvenot NC2105; “in
said a synonym of the latter in the checklist (Thouvenot et al. jugo Dogny (1050 m), Catene centralis”, VII.1909, L. Le Rat s.n.
2011). Both REN specimens and their duplicate at PC were as “Lophocolea defectistipula” (REN[herb. E. G. Paris]).

52 CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1)

 Lophocoleaceae of New Caledonia

A
B C

Fig. 30. — Chiloscyphus longifissus Steph.: A, male shoot end with androecium, dorsal view; B, leaf cells; C, D, shoot portions, ventral view; E, underleaf; F, shoot
portion, dorsal view. A, D, F, from the isotype (PC0101940); B, C, E, from the lectotype (G00069456). Scale bars: A, D, 1 mm; B, 50 µm; C, E, F, 0.5 mm.

Distribution in New Caledonia. — Goebelobryum unguiculatum Cells

is known from a few collections in both provinces where it grows
on bare soils in mountain shrublands and forests (collection data Leaf cells large, 40-80 µm wide, cell walls thin, red tinged as
from 700 to 1000 m). well as the medium globose trigones, external wall of mar-
ginal cells thicker.
Total range. — Australasia.
Underleaves
Description Very small to vestigial, associated with a large bulging rhizoi-
Based on the type material of Lophocolea purpurea. Further dal plate, underleaves not connected to the leaves, at most
descriptions and illustrations in Grolle (1962), Engel & 80-160 µm long, 80-120 µm wide, narrower than the stem,
Glenny (2012). sometimes inconspicuous, reduced to a few cells, when devel-
oped bifid, with two short lobes erect linear, up to three cells
Habit long, separated by a “v” shaped sinus; the lamina null or, at
Plant medium sized, light green and purple tinged to wine- most, small rectangular, 3-4 cells long or wide, lateral margins
red coloured, blackish in older parts, shoots 3.00 mm wide, with a single small unicellular tooth if any.
stem relatively thick, 0.30 mm wide; leaves alternate, dorsally
assurgent-convex, spreading at right angle when flattened, Marsupium
dorsally free; rhizoids thin, hyaline, dense, obscuring the Developed at the arched end of some shoots, long cylindrical,
ventral face of the stems and the underleaves, inserted on hidden in the substrate; gynoecia and androecia not seen.
bulging rhizoidal plates at the bases of the underleaves and
leaf ventral margins. Comments
The type specimen of Lophocolea purpurea seen by Stephani
Leaves was sterile so that he missed the marsupium-bearing condition
1.40-2.00 mm long, 1.50-2.50 mm wide, rounded to reni- of this species and, although the specimen exhibits vegetative
form, apices somewhat angular or emarginate, margins characters unusual in the family Lophocoleaceae, namely
long ciliate, more distantly on the lower parts, with up the small or vestigial underleaves, Stephani described it as
to 10 rigid whitish cilia, made of 4-5 elongate, uniseriate, Lophocolea purpurea. Later, when Engel & Schuster (1984)
thick-walled cells. downgraded Lophocolea to a subgenus of Chiloscyphus, they

CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1) 53

 Thouvenot L.

replaced this name by Chiloscyphus novae-caledoniae (the Tetracymbaliella decipiens (Gottsche) Grolle
blocking name Chiloscyphus purpureus Steph. is from South
America), likely without reviewing the type. There is a part of Nova Hedwigia 3: 49-50 (Grolle 1961). — Chiloscyphus decipiens
the original type material sent to Stephani in the herbarium Gottsche, Synopsis Hepaticarum: 176 (Gottsche et al. 1845). —
Heteroscyphus decipiens [Link] & [Link]., Nova Hedwigia
of E. G. Paris (REN). This specimen has some shoots with 39: 402 (Engel & Schuster 1984 [1985]). — Type: New Zealand.
marsupia so that it belongs to Goebelobryum. Goebelobryum Menzies, not seen.
unguiculatum was already known from New Caledonia
(Thouvenot et al. 2011). Comments
The checklist of the liverworts of New Caledonia (Thou- The only New Caledonian report of this species, otherwise
venot et al. 2011) mentions a doubtful record of Plagiochila known as endemic to New Zealand, was by Furuki & Higuchi
blepharophora (Nees) Lindenb., according to a specimen (1996). It was impossible to check the sample the record was
reported by Paris (1910) under the name Lophocolea defectis- based on and its presence in New Caledonia needs confirmation.
tipula Steph. The relevant specimen at REN has been checked
and has proven to be a poorly coloured Lophocolea purpurea. Taxa excluded from the New Caledonian
Therefore, Plagiochila blepharophora must be removed from bryophyte flora
the New Caledonian bryophyte flora.
Genus Cryptolophocolea L.Söderstr.

Family Brevianthaceae [Link] & [Link]. Cryptolophocolea levieri (Schiffn.) L.Söderstr.

Genus Tetracymbaliella Grolle
Phytotaxa 112: 21 (Söderström 2013b). — Lophocolea levieri
Tetracymbaliella comptonii (Pearson) Grolle Schiffn., Die Hepaticae der Flora von Buitenzorg 182 (Schiffner
1900). — Chiloscyphus paroicus [Link] & [Link]., Nova
Hedwigia 39: 420 (Engel & Schuster 1984 [1985]). — Type: Indo-
Revue bryologique et lichénologique 32: 164 (Grolle 1963). — nesia. Schiffner 1207-1210 (isosyn-, PC[PC0102563]!).
Chiloscyphus comptonii Pearson, Journal of the Linnean Society,
Botany 46: 23 (Pearson 1922). — Heteroscyphus comptonii (Pear- Further specimens examined. — Java. Preanger province, Tjibur-
son) [Link] & [Link]., Nova Hedwigia 39: 402 (Engel & rum near Tjibodas, in primary cloud forest, 1575 m, [Link].1894,
Schuster 1984 [1985]). — Type: New Caledonia. Pearson 740, Schiffner 1207 as Lophocolea levieri (PC[PC0102563]).
not seen. New Caledonia. Mts. Koghis, [Link].1914, Compton 801 as Lophoc-
olea levieri (BM[BM013409501, BM013409504]); Mt. Mou, on
Tetracymbaliella ratiana Grolle, Nova Hedwigia 3: 51-52 (Grolle
stone and mud in stream, [Link].1914, Compton 444 as Lophocolea
1961). — Chiloscyphus grossitextus Steph. ex Paris, nom. inval. —
levieri (BM[BM013409502]).
Type: New Caledonia. Pic des Sources, Le Rat s.n. (holo-,
M[fide Grolle 1961]; iso-, REN[“Chiloscyphus grossitextus,
Nov. Caledon. Pic des Sources”, VII.1909, Le Rat s.n., herb.
E. G. Paris]!). Comments
This species was reported from New Caledonia by Pearson
Further specimens examined. — New Caledonia. South Province, (1922) as Lophocolea levieri and included as Chiloscyphus paro-
Yaté, Wé Toa, on dead wood in wet forest in ultramafic massif, 500 m, icus in the checklist (Thouvenot et al. 2011). The description
[Link].2019, Thouvenot NC2857; Mt. Mou, VII.1909, Le Rat s.n. as
“Chiloscyphus cymbaliferus” (REN); Mt. Dzumac, “ad arbores in given by Pearson does not match the original diagnosis of
silvat., 1000 m alt”, VII.1905, Le Rat s.n. (REN). L. levieri (Schiffner 1900) in the following features: 1) entire
leaf margins; and 2) deeply bilobed underleaves, free or nar-
Distribution in New Caledonia. — Scattered in South Province rowly connate on a single side, with a single tooth on each
where it grows on dead wood in rain forest at medium elevations lateral margin. Furthermore, Pearson stated that his specimen
(collected from 500 to 1000 m).
was close to Lophocolea heterophylla (Schrad.) Dumort. The
Total range. — Endemic. three checked parts of the Compton collection named by
Pearson as L. levieri (BM) turned to be Lophocolea convexula,
Description and illustration which fits Pearson’s remarks on the likeness to L. heterophylla.
Grolle (1961) as Tetracymbaliella ratiana. Therefore, Cryptolophocolea levieri (as Chiloscyphus paroicus)
must be removed from the bryophyte flora of New Caledonia.
Comments
This species is very distinctive by the conspicuous pouches
on leaf and underleaf margins. This character is shared Genus Heteroscyphus Schiffn.
with Tetracymbaliella cymbalifera (New Zealand) that might
introduce some misidentification (see below). T. ratiana is Heteroscyphus hebridensis (Steph.) Schiffn.
distinguished from the latter by: 1) larger trigones almost
Oesterreichische Botanische Zeitschrift 60: 172 (Schiffner 1910). —
as wide as the cell lumina; 2) pouches longer than wide; 3) Chiloscyphus hebridensis Steph., Hedwigia 32: 323 (Stephani
dorsal leaf margins not connivent; and 4) leaf cells strongly 1893). — Type: Vanuatu. “Novae Hebridae”, comm. K.Müller,
mammillose. Hal. (G[G00283079, G00283073, G00069477]!).

54 CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1)

 Lophocoleaceae of New Caledonia

Further specimens examined. — New Caledonia. South Province, be Heteroscyphus deplanchei (as H. heteromorphis). Therefore,
Mts. Koghis, [Link].1909, Franc s.n. (G[G005125], PC[PC0167675]). this species must be removed from the New Caledonian flora.

Comments
Unpublished specimens labelled “Chiloscyphus hebridensis, from Genus Lophocolea (Dumort.) Dumort.
Mts. Koghis, Franc s.n., 1/11/1909, determinavit Stephani”,
have been found at PC and G, and turned to be Heteroscy- Lophocolea autoica Steph.
phus asellifomis. The features distinctive from H. hebridensis
are: 1) leaves rounded with margins dorsally confluent vs Species Hepaticarum 6: 262 (Stephani 1922). — Chiloscyphus
widely triangular-ovate and dorsally free; 2) underleaf mar- autoicus (Steph.) [Link] & [Link]., Nova Hedwigia 39:
411 (Engel & Schuster 1984 [1985]). — Type: Haïti. Faurie s.n.!
gins toothed vs deeply laciniate; and 3) leaf cells with large
trigones vs minute trigones. The presence of this species in Further specimen examined. — New Caledonia. Lerat s.n.
New Caledonia remains unconfirmed. (G[G00128109]).

Comments
Heteroscyphus jackii (Steph.) Schiffn. Stephani (1922) published Lophocolea autoica based on two
Oesterreichische Botanische Zeitschrift 60: 172 (Schiffner 1910). — syntypes, from Hawaï (Faurie) and New Caledonia (Lerat). The
Chiloscyphus jackii Steph., Botanisches Centralblatt. 60: 102 (Jack & type specimen from Haïti in G matches the protologue well. The
Stephani 1894). — Type: Samoa. Graeffe 1636 (G[G00069467]!). New Caledonian specimen, however, is Lophocolea convexula.
Since this is the only mention of this species in New Caledonia,
Further specimens examined. — New Caledonia. North Prov-
ince, Mt. Dzumac, IV.1907, Le Rat s.n. (REN); “sine schedule”, it must be removed from the bryophyte flora of the territory.
1907, Le Rat s.n. (G).

Genus Plagiochila (Dumort.) Dumort.

Comments
Hürlimann (1998) mentions New Caledonia as part of the
Plagiochila blepharophora (Nees) Lindenb.
range of this species, likely following Paris (1908) who reports
a specimen of Chiloscyphus jackii Steph. collected by Le Rat, Species Hepaticarum 2-4: 102 (Lindenberg 1840). — Type: Java.
also reported by Miller et al. (1983). In G, only one specimen (G[G00115176]), not seen.
collected in New Caledonia by Le Rat is labelled Chiloscyphus
jackii by Stephani. This specimen turned to be Heteroscyphus Lophocolea defectistipula Steph., Species Hepaticarum 3: 130 (Stephani
1906). — Type: Caroline Islands. Parkinson s.n. (G[G00061183]),
argutus. In the herbarium of E. G. Paris (REN), a duplicate of not seen.
the original sent to Stephani has the same misidentification.
All the Hürlimann’s specimens in G are from Tonga. As the Specimen examined. — New Caledonia. South Province, “in jugo
only voucher from New Caledonia is not this species, Heteros- Dogny (1050 m), Catene centralis”, VII.1909, L. Le Rat s.n. as
“Lophocolea defectistipula” (REN[herb. E. G. Paris]).
cyphus jackii must be removed from the New Caledonian flora.
Comments
Genus Leptoscyphus Mitt. Treated as doubtful record in Thouvenot et al. (2011), this
species must be removed from the New Caledonian flora
Leptoscyphus physanthus (Hook.f. & Taylor) [Link] since the reference specimen turned to be Goebelobryum
unguiculatum (see above).
Nova Hedwigia 99: 168 (Engel 2014). — Jungermannia physantha
Hook.f. & Taylor, London Journal of Botany 3: 561 (1844). — Het-
eroscyphus physanthus (Hook.f. & Taylor) [Link]., The Hepaticae
and Anthocerotae of North America 4: 248 (1980). — Type: New
Genus Tetracymbaliella Grolle
Zealand. Not seen.
Tetracymbaliella cymbalifera
Specimen examined. — New Caledonia. South Province, Rivière (Hook.f. & Taylor) Grolle
Bleue valley, on dead wood in forest, 160 m, 15.V.1970, Schmid 148
as “Chiloscyphus physanthus” (PC[PC0146342]).
Nova Hedwigia 3: 50 (Grolle 1961). — Jungermannia cymbalifera
Hook.f. & Taylor, Flora Antarctica 1: 151 (Hooker 1845). — Chilos-
Comments cyphus cymbaliferus (Hook.f. & Taylor) Gottsche, Lindenberg &
Heteroscyphus physanthus (Hook.f. & Taylor) [Link]. is Nees, Synopsis Hepaticarum 5: 711 (Gottsche et al. 1847). — Type:
reported as doubtful record in Thouvenot et al. (2011). The only New Zealand. Auckland Islands, not seen.
report in New Caledonia of this species, regarded as endemic
Specimen examined. — New Caledonia. South Province, Mt. Mou,
to New Zealand, was from Tixier (1972) as Chiloscyphus phy- VII.1909, Le Rat s.n. as “Chiloscyphus cymbaliferus” (REN); Mt. Dzu-
santhus Taylor. The relevant voucher in the Muséum national mac, “ad arbores in silvat., 1000 m alt”, VII.1905, Le Rat s.n. as
d’Histoire naturelle of Paris (PC[PC0146342]) turned out to “Chiloscyphus cymbaliferus” (REN).

CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1) 55

 Thouvenot L.

Comments Engel J. J. 2013. — Studies on Lophocoleaceae XXII. New taxa

The only report of this species in New Caledonia is from and combinations in New Zealand Heteroscyphus Schiffn. Pol-
ish Botanical Journal 58 (1): 95-106. [Link]
E. G. Paris (1906, 1910) as Chiloscyphus cymbaliferus. The pbj-2013-0011
two specimens in his herbarium kept at Rennes University Engel J. J. 2014. — Studies on Lophocoleaceae XXIII. Novelties in
(REN) are not different from Tetracymbaliella comptonii. Heteroscyphus Schiffn. together with refinements in Cryptolophocolea
Therefore, T. cymbalifera must be removed from the New L.Söderstr., Crand.-Stotl., Stotler & Váňa and Leptoscyphus Mitt.
Caledonian flora. Nova Hedwigia 99 (1-2): 157-170.
Engel J. J. 2015. — Studies on Lophocoleaceae. XXV. A conspec-
tus of Heteroscyphus Schiffn. in temperate Australia together
with nomenclatural changes in Chiloscyphus Corda and Leptos-
Acknowledgements cyphus Mitt., refinements and a range extension of Clasmato-
The author warmly thanks the curators and staff members colea Spruce, and a range extension in Stolonivector [Link].
at G, PC and REN for their kind availability and assistance Nova Hedwigia 100 (3-4): 553-582. [Link]
nova_hedwigia/2015/0260
during working visits: A. Chambet, A. Gautschi, L. Kervran, Engel J. J. & Glenny D. 2012. — Austral Hepaticae 48. Goebelo-
S. Leblond, M. Lemaire, M. J. Price, I. Valette; the curators bryum Grolle (Acrobobaceae). Nova Hedwigia 95 (3-4): 319-336.
of BM, GOET, L, Z+Zt for the loan of type specimens; [Link]
H. Hofmann (Z) for kindly providing photos and com- Engel J. J. & Glenny D. 2019. — A Flora of the liverworts and
ments of a rare specimen; G. Briand (Rennes 1 University hornworts of New Zealand. Vol. 2. Monographs in Systematic
botany from the Missouri Botanical Garden 134: 1-750.
Library) for the correspondence of Paris with Stephani;
Engel J. J. & He X. 2010. — Studies on Lophocoleaceae. XIX.
S. R. Gradstein for many kind supports, especially by con- The systematic identity of Cyanolophocolea R.M. Schust., an
siderably improving a previous version of the manuscript intriguing liverwort from New Zealand and Australia, based
and making available Hürlimann’s specimens; Frank Müller on morphological and molecular evidence. The Bryologist 113
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help in terms of documentation and advice; D. Glenny and Engel J. J. & Schuster R. M. 1984 [1985]. — An overview and
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presidents and environment managers of Province Nord and Lophocoleaceae XXI. Otoscyphus [Link], Bardat & Thouvenot,
Province Sud of New Caledonia for collecting permits; Edith a new liverwort genus from New Caledonia with an unusual
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Submitted on 21 February 2022;

accepted on 29 July 2022;
published on 16 January 2023.

58 CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1)

 Lophocoleaceae of New Caledonia

APPENDIX

Appendix 1. — Index of names. The names of taxa included in Lophocoleaceae Vanden Berghen according to the checklist of the New Caledonian liverworts
(Thouvenot et al. 2011) are listed below with their currently accepted name in bold. The new species, unknown in New Caledonia at the time of the checklist
publication, are marked with an asterisk.

Chiloscyphus acutus Steph. = Heteroscyphus argutus

C. autoicus (Steph.) [Link] & [Link]. (excluded name)
C. beesleyanus Pearson = C. longifissus
C. confertifolius Steph. = Heteroscyphus coalitus
C. confertus Steph. = Heteroscyphus confertus
C. convexulus (Mitt.) [Link] & [Link]. = Lophocolea convexula
C. cornutistipulus Steph. = Heteroscyphus cornutistipulus
C. crassicaulis Steph. = Otoscyphus crassicaulis
C. etesseanus Steph. = Heteroscyphus etesseanus
C. explanatus (Mitt.) [Link] & [Link]. = Cryptolophocolea explanata
C. fragillimus (Steph.) [Link] & [Link]. = Lophocolea convexula
C. francanus Steph. = Heteroscyphus coalitus
C. heteromorphus (Steph.) [Link] & [Link]. = Heteroscyphus deplanchei
C. kurzii (Sande Lac.) [Link] & [Link]. = Lophocolea convexula
C. latistipulus (Steph.) [Link] & [Link]. = Heteroscyphus coalitus
C. latistipus Steph. = Heteroscyphus coalitus
C. leratii [Link] & [Link]. = Lophocolea caledonica
C. longifissus Steph. (incertae sedis)
C. muricatus (Lehm.) [Link] & [Link]. = Lophocolea muricata
C. novae-caledoniae [Link] & [Link]. = Goebelobryum unguiculatum (Acrobolbaceae)
C. papulimarginatus ([Link].) [Link] & [Link]. = Lophocolea convexula
*C. parapilistipulus Thouvenot = Heteroscyphus parapilistipulus
C. paroicus [Link] & [Link]. = Cryptolophocolea levieri (excluded name)
C. parvus (Steph.) [Link] & [Link]. = Lophocolea convexula
C. pilistipulus (Steph.) [Link] & [Link]. = Lophocolea convexula
C. quadricilius Steph. = Heteroscyphus grandiflorus
C. rotundiphyllus [Link]. = Heteroscyphus confertus
C. savesianus (Steph.) [Link] & [Link]. = Lophocolea savesiana
C. similis Steph. (1908) non C. similis Steph. (1911) = Heteroscyphus coalitus
C. subacuminatus Herzog = Heteroscyphus subacuminatus
C. subcostatus (Steph.) [Link]& [Link]. = Cryptolophocolea subcostata
C. subsimilis Steph. = Heteroscyphus coalitus
C. trigonifolius Steph.
Conoscyphus trapezioides (Sande Lac.) Mitt. ex Schiffn. (moved to Acrobolbaceae)
Heteroscyphus argutus (Reinw., Blume & Nees) Schiffn.
H. aselliformis (Reinw., Blume & Nees) Schiffn.
*H. assurgentissimus [Link], Thouvenot & Frank Müll.
H. caledonicus (Steph.) Schiffn.
H. coalitus (Hook.) Schiffn.
H. comptonii (Pearson) [Link] & [Link]. = Tetracymbaliella comptonii (moved to Brevianthaceae)
H. cymbalifer (Hook.f. & Taylor) [Link] & [Link]. = Tetracymbaliella cymbalifera (excluded name)
H. decipiens (Gottsche) [Link] & [Link]. = Tetracymbaliella decipiens (Brevianthaceae) (presence in New
Caledonia needs confirmation)
H. deplanchei (Steph.) Schiffn.
*H. diestianus (Sande Lac.) Piippo
H. giganteus (Steph.) Hürl.
H. grandiflorus (Steph.) Hürl.
H. jackii (Steph.) Schiffn. (excluded name)
*H. kanakensis Thouvenot & [Link]
H. splendens (Lehm. & Lindenb.) Grolle
*H. succulentus (Gott.) Schimp.

CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1) 59

Thouvenot L.

*H. supinopsis [Link], Thouvenot & Frank Müll.

*Lophocolea bidentata (L.) Dumort.

60 CRYPTOGAMIE, BRYOLOGIE • 2023 • 44 (1)