Table Of ContentPRIMARY RESEARCH PAPER | Philippine Journal of Systematic Biology
Amphi-Pacific tropical disjunctions in the bryophyte floras of
Asia and the New World
S. Robbert Gradstein
ABSTRACT
Disjunctions between tropical America and tropical Asia, commonly called amphi-Pacific tropical disjunctions,
have frequently been discussed among flowering plants but have received very little attention in bryology. A
screening of the literature revealed nine species and sixteen genera or infrageneric taxa of bryophytes with
amphi-Pacific tropical (or subtropical) ranges. They include Austinia tenuinervis, Diphyscium chiapense, D.
longiflorum, Elmerobryum, Fissidens sect. Sarawakia, Ganguleea angulosa, Hydrogonium arcuatum,
Hymenostyliella, Hymenostylium aurantiacum, Luisierella barbula, Mniomalia, Rozea, Sphaerotheciella and
Sorapilla among the mosses and Ceratolejeunea grandiloba, Drepanolejeunea subg. Rhaphidolejeunea,
Lejeunea sect. Echinocolea, Lobatiriccardia, Myriocoleopsis sect. Myriocoleopsis, Phycolepidozia,
Pictolejeunea, Rectolejeunea, Southbya organensis and Vitalianthus among liverworts. All of them occur in
tropical or subtropical Asia and the Neotropics but are not known from Africa. The causes of the amphi-Pacific
tropical disjunctions in bryophytes are still unclear. In flowering plants, molecular analyses indicate that amphi-
Pacific tropical ranges frequently resulted from past migration across Eurasia and the northern Atlantic Ocean,
followed by local extinction. This scenario may also have operated in amphi-Pacific bryophytes but some might
have reached South America via the southern Pacific migration route. The possibility of direct long-range
dispersal across the Pacific Ocean cannot be ruled out and this scenario seems likely for Southbya
organensis, which occurs on Hawaii and freely produces spores and small gemmae. The possibility that the
disjunctive ranges reflect insufficient collecting and that some taxa also occur in Africa should also be taken
into account. There is no strong evidence for human introduction of amphi-Pacific tropical bryophytes. The
new combinations Lejeunea sect. Echinocolea (R.M.Schust.) Gradst. comb. nov. and Myriocoleopsis sect.
Protocolea (R.M.Schust.) Gradst. comb. nov. are proposed.
KEYWORDS: Biogeography, Disjunctive range, Liverworts, Long-range dispersal, Migration, Mosses
INTRODUCTION the genus, with two species occurring in Southeast Asia
and one in Guatemala, Central America. The distribution of
In 2007, the late Benito Tan together with William R. Buck Elmerobryum is noteworthy because taxa occurring in
published a taxonomic account of Elmerobryum, a small tropical Asia and tropical America are usually also found in
tropical genus in the moss family Hypnaceae with three Africa. However, Elmerobryum has never been found in
species, E. philippinense (type), E. wilhelmense and E. Africa.
guatemalense. Oddly, the type species was described three
times under a different name, each name being based on a Disjunct geographical ranges have long fascinated plant
collection from the same locality (!), Baguio in northern Luzon. geographers and there is much literature on the subject.
Even more remarkable was the highly disjunct distribution of The disjunctions between tropical Asia and tropical
America, as exemplified by Elmerobryum, are commonly
referred to as "amphi-Pacific tropical" disjunctions (Thorne,
Muséum National d'Histoire Naturelle, Institut Systématique,
1972). Over 100 genera and higher taxa of vascular plants
Evolution, Biodiversité / Grand Herbier, Case Postale 39, 12 rue
exhibiting this type of distribution were enumerated by
Buffon, 75005 Paris. France
Thorne and more examples could probably be added (e.g.,
*Corresponding author: [email protected] Fritsch et al., 2015). These disjunctive ranges are
commonly explained by past migration across Eurasia and
Date Submitted: 09 October 2017 the northern Atlantic Ocean, followed by local extinction, or
Date Accepted: 07 February 2018 by direct long-distance dispersal (LDD) across the Pacific
© Association of Systematic Biologists of the Philippines Volume 12 Issue 1 - 2018
Philippine Journal of Systematic Biology | Gradstein, S. R.: Amphi-Pacific tropical disjunctions in bryophyte floras
Ocean. However, only very few examples have been records were old, 19th century reports.
analyzed with modern molecular-phylogenetic and
biogeographic methods (Li & Wen, 2013; Yang et al., 2017). A first, preliminary review of amphi-Pacific tropical
disjunctions in bryophytes, using data from taxonomic
In this paper, I shall briefly review amphi-Pacific tropical revisions and verified floristic reports, was presented by
(including subtropical) distributions among bryophytes. Gradstein et al. (2014). Sixteen taxa with amphi-Pacific
Bryophytes are very interesting organisms from the tropical distribution, seven of mosses and nine of liverworts,
standpoint of biogeography, showing notable differences with were enumerated. Since the appearance of this paper,
flowering plants. They usually have wider ranges than several further amphi-Pacific tropical bryophyte taxa were
flowering plants and much fewer species are endemic. For added to the list while others had to be removed based on
example, endemism among vascular plant species of the results of molecular-phylogenetic analysis. For example,
Colombia is about 25% while only 4.5% of Colombian molecular analysis of Cololejeunea subg. Chlorolejeunea
bryophytes are endemic (Bernal et al., 2016). In Japan, Benedix, a small amphi-Pacific group of specialized
endemism is almost 32% in flowering plants and 8.4% in rheophytes (2 spp.), showed that the group was polyphyletic
bryophytes (Kato & Ebihara, 2011; Higuchi, 2011) The low and that the two species were only distantly related (Ye et
endemism rate in bryophytes is believed to be due to their al., 2013). Their similarity was apparently the result of
excellent dispersal capability, via spores or asexual devices parallel evolution. A further example is Leptolejeunea
such as gemmae. Van Zanten (1978) found that widely elliptica (Lehm. & Lindenb.) Schiffn., which had sometimes
distributed bryophyte species have more durable spores than been considered an amphi-Pacific tropical disjunct based on
endemic species, illustrating the important role that spores the avowed conspecificity with the Asiatic Leptolejeunea
may play in shaping distributions, whereas Laenen et al. subacuta Evans (Schuster, 1968). Bischler (1969), however,
(2016) demonstrated the importance of small gemmae for in her monographic treatment of Leptolejeunea in tropical
long-distance-dispersal of bryophytes. Disjunct occurrence of America, considered L. elliptica a purely neotropical species
bryophyte species in different continents or on remote and this has recently been confirmed by DNA analysis
oceanic islands are clearly suggestive of their excellent (Bechteler et al., 2016; L. Shu, pers. com.). The amphi-
dispersal capabilities. Continental disjunctions in the Tropics Pacific genus Spruceanthus Verd. (ca. 10 spp.), on the other
were reviewed by Pócs (1976, 1992), Gradstein et al. (1983), hand, proved to be more widespread based on molecular
Delgadillo (1993), Tan (1978) and Gradstein (2013, 2017), analysis and was newly detected in Africa where it had been
whereas island diversities were studied by Patiño et al. assigned to the wrong genus (Sukkharak & Gradstein,
(2013). By comparing the floras of 67 oceanic islands across 2017).
12 archipelagos, the latter authors could show that oceanic
barriers were not a major impediment for immigration of The current paper is an updated and expanded version of
bryophyte species. Moreover, they found that bryophyte the above-mentioned review. The paper is dedicated to the
diversity of remote oceanic islands is essentially determined memory of the late Benito Tan, world specialist of the
by habitat availability, not by island size, age, elevation or mosses of Southeast Asia and a dear friend.
geographic isolation.
RESULTS AND DISCUSSION
Very little attention has been paid in the literature to amphi-
Pacific tropical disjunctions in bryophytes. Schuster (1983) in A perusal of the literature revealed nine species and sixteen
his review of bryophyte phytogeography mentioned a single genera or infrageneric taxa of bryophytes with amphi-Pacific
example, the genus Tuyamaella S.Hatt. (Lejeuneaceae) with tropical distribution (see below: Notes on amphi-Pacific
a wide range in Southeast Asia and a single record on the taxa). The list does not pretend to be exhaustive and further
west coast of South America (Peru), and considered this bryophyte taxa exhibiting this type of range may exist. The
distribution "wholly erratic" and "inexplicable". Gradstein et al. species list includes the mosses Austinia tenuinervis,
(2018), however, found that the Peruvian specimen was Diphyscium chiapense, D. longiflorum, Ganguleea angulosa,
mislabeled and had originated from Asia. A similar case of Hydrogonium arcuatum, Hymenostylium aurantiacum and
erroneous labeling is the Peruvian record of the Asiatic Luisierella barbula, and the liverworts Ceratolejeunea
Cololejeunea inflectens (Mitt.) Benedix (Pócs et al., 2014). grandiloba and Southbya organensis (Fig. 1). All of them
Misidentifications also lead to phytogeographic errors. For occur in tropical or subtropical Asia and the Neotropics but
example, Söderström et al. (2010) enumerated 23 neotropical are not known from Africa. Among them, Ganguleea
liverwort species erroneously recorded from Java and angulosa and Luisierella barbula represent monospecific
Thouvenot et al. (2011) listed five wrong reports of neotropical genera while A. tenuinervis belongs to a small genus of two
species from New Caledonia. Most of these erroneous species, the second one occurring in northern Australia. The
© Association of Systematic Biologists of the Philippines Volume 12 Issue 1 - 2018 | 2
Philippine Journal of Systematic Biology | Gradstein, S. R.: Amphi-Pacific tropical disjunctions in bryophyte floras
Figure 1. Distribution of amphi-Pacific tropical bryophyte species. A, Austinia tenuinervis. B, Diphyscium longifolium. C, Ganguleea
angulosa. D, Luisierella barbula. E, Ceratolejeunea grandiloba. F, Southbya organensis.
disjunctive species are usually known from several (or many) which contains two subspecies: one from tropical America
localities in both continents with exception of Southbya and one from Asia (Magombo, 2003).
organensis and Ceratolejeunea grandiloba. Southbya
organensis is known from many localities in Southeast Asia Amphi-Pacific tropical genera and infrageneric taxa are more
but is extremely rare in tropical America where it is known numerous and include Elmerobryum, Fissidens sect.
from only two sites (Fig. 1F). In one of these sites (Serra dos Sarawakia, Hymenostyliella, Mniomalia, Rozea,
Orgãos, Rio de Janeiro; type locality of S. organensis) the Sphaerotheciella and Sorapilla among the mosses, and
species is probably extinct (Gradstein & Costa, 2003). Drepanolejeunea subg. Rhaphidolejeunea, Lejeunea sect.
Ceratolejeunea grandiloba, on the other hand, is rather Echinocolea, Lobatiriccardia, Myriocoleopsis sect.
widespread in the tropical Andes but is known in Asia from Myriocoleopsis, Phycolepidozia, Pictolejeunea,
only one locality (Fig. 1E). The Asian origin (Java) of the latter Rectolejeunea and Vitalianthus among liverworts (Figs. 2, 3).
specimen is unambiguous but the Asian plant is not fully Several of them are very rare and unusual taxa such as
identical to the South American ones and is placed in a Fissidens sect. Sarawakia, Phycolepidozia and Sorapilla.
different subspecies (Gradstein, 2013a). Dissimilarity of the Each of these has one species in Asia and one in the New
disjunct populations is also seen in Diphyscium chiapense World, all of them known only from a single or very few
localities. Moreover, they are morphologically highly
© Association of Systematic Biologists of the Philippines Volume 12 Issue 1 - 2018 | 3
Philippine Journal of Systematic Biology | Gradstein, S. R.: Amphi-Pacific tropical disjunctions in bryophyte floras
specialized taxa and Sorapilla is placed in a family of its own. the amphi-Pacific tropical bryophyte taxa have been carefully
The remaining amphi-Pacific genera and infrageneric taxa analyzed by modern molecular-phylogenetic and
include some that are very unevenly represented in the two biogeographic methods. Such studies are necessary in order
continents while others are more evenly spread. For example, to better understand their biogeographic histories. Before the
Drepanolejeunea subg. Rhaphidolejeunea, Hymenodon, age of molecular-phylogenetic research, intercontinental
Hymenostyliella and Lobatiriccardia are mostly Asiatic while disjunctions were often explained as ancient vicariance
Lejeunea sect. Echinocolea, Myriocoleopsis sect. patterns that dated back to the early Mesozoic, before the
Myriocoleopsis, Pictolejeunea and Rectolejeunea are mainly break-up of Pangaea (e.g., Schuster, 1983). Recent
neotropical. Elmerobryum, Mniomalia, Rozea and molecular studies, however, have shown that the ages of
Vitalianthus, one the other hand, are rather evenly extant bryophyte species, and of many of the genera, are too
represented in both continents. young for a vicariance scenario of evolution. Consequently,
dispersal scenarios are currently proposed more frequently
The causes of the amphi-Pacific tropical disjunctions in as explanations of intercontinental ranges (Heinrichs et al.,
bryophytes are still unclear. Among flowering plants, 2009; Gradstein, 2013, 2017).
molecular analyses indicate that the amphi-Pacific tropical
ranges frequently resulted from past migration across Eurasia The possibility of LDD across the Pacific cannot be ruled out
and the northern Atlantic Ocean, followed by local extinction and this scenario seems likely for Southbya organensis,
(Li & Wen, 2013). This scenario may have operated also in which occurs on Hawaii (Fig. 1F). Although being dioicous,
amphi-Pacific bryophyte genera but some, such as S. organensis is often found with sporophytes (Váňa et al.,
Hymenodon and Lobatiriccardia, may have reached South 2012) and commonly produces small gemmae, which are
America via the southern Pacific migration route (e.g., considered very suitable for LDD (Laenen et al., 2016).
Schuster, 1979). It should be stressed, however, that none of Small lacking gemmae are lacking in the other amphi-Pacific
Figure 2. Distribution of amphi-Pacific tropical genera and infrageneric taxa of mosses. A, Elmerobryum. B, Fissidens sect.
Sarawakia. C, Hymenostyliella. D, Mniomalia. E, Rozea. F, Sorapilla.
© Association of Systematic Biologists of the Philippines Volume 12 Issue 1 - 2018 | 4
Philippine Journal of Systematic Biology | Gradstein, S. R.: Amphi-Pacific tropical disjunctions in bryophyte floras
Figure 3. Distribution of amphi-Pacific tropical genera and infrageneric taxa of liverworts. A, Drepanolejeunea subg.
Rhaphidolejeunea. B, Myriocoleopsis sect. Myriocoleopsis. C, Phycolepidozia. D, Pictolejeunea. E, Rectolejeunea. F, Vitalianthus.
tropical taxa but about two third of them are monoicous and insufficiently explored. An interesting case is the specialized
produce bisexual spores (often quite small ones), which are occurrence of the moss genus Hymenodon on tree fern
also considered an important condition for successful LDD stems (e.g., Koponen et al., 1986; Vital & Prado, 2006).
(e.g., van Zanten & Pócs, 1981; Gradstein et al., 1983; More intensive inventory of their habitats may reveal
Gradstein, 2013, 2017; but see Laenen et al., 2016). additional localities, including occurrences in Africa, of the
amphi-Pacific tropical taxa.
There is no strong evidence for human introduction of amphi-
Pacific tropical bryophytes. Nevertheless, the distribution of NOTES ON AMPHI-PACIFIC TROPICAL TAXA
the pottiaceous taxa Ganguleea angulosa and Luisierella
barbula might have been influenced by man in view of their 1. MOSSES (Bryophyta s.str.)
occurrence in ruderal habitats. The possibility that the disjunct
amphi-Pacific ranges reflect insufficient collecting should also Austinia tenuinervis Müll.Hal. (Myriniaceae) Figure 1A
be taken into account. The moss genus Campylopodiella Distribution and habitat: scattered in tropical America
Card., for example, was long known only from the Neotropics (Cuba, Mexico, Peru, SE Brazil) and Southeast Asia
and the Himalayan region but was recently collected in Africa (Thailand, Sabah, Philippines: Mindanao); on bark of trees at
(Townsend, 2009). Undercollecting is likely the case in the low elevation. A second species in the genus occurs in
genus Phycolepidozia due to its minute size, and may northern Australia (Queensland).
furthermore hold for the epiphyllous Drepanolejeunea subg. Sexuality and dispersal: monoicous; spores 20-25 µm;
Rhaphidolejeunea, the rheophytic Myriocoleopsis and the specialized asexual reproduction not observed.
canopy specialists Ceratolejeunea grandiloba and Note: Buck & Crum (1978) recognized two varieties, var.
Rectolejeunea, all of which grow in habitats that have been tenuinervis in tropical America and var. micholitzii in tropical
© Association of Systematic Biologists of the Philippines Volume 12 Issue 1 - 2018 | 5
Philippine Journal of Systematic Biology | Gradstein, S. R.: Amphi-Pacific tropical disjunctions in bryophyte floras
Asia, based on quantitative differences in the sporophyte. fertile; spores 17-25 µm; specialized asexual reproduction
However, the sporophyte of plants from Thailand illustrated by not observed.
Printarakul et al. (2013) is more similar to var. tenuinervis Note: sect. Sarawakia stand out by unusually long (to 10
than to var. micholitzii. This suggest that distinction of the two cm or more), pinnately branched stems with numerous
varieties may not be warranted. sporophytes on short branches. The monophyly of the
Reference: Buck & Crum (1978), Printarakul et al. (2013). section should be tested; the possibility that the similarity of
the two Sarawakia species is due to parallel evolution (like in
Diphyscium chiapense D.H.Norris (Diphysciaceae) Cololejeunea subg. Chlorolejeunea; Yu et al., 2013) cannot
Map: Magombo (2003: Figures 3, 5). be ruled out.
Distribution and habitat: subsp. chiapense in Central Reference: Pursell et al. (1988), Pursell & Nannenga-
America (Costa Rica, Mexico: Chiapas) and subsp. Bremekamp (2004), Gradstein et al. (2012).
unipapillosum (Deguchi) T.Y.Chiang & S.H.Lin in eastern Asia
(China, Taiwan, Japan, Philippines: Mindanao); on bark, Ganguleea angulosa (Broth. & Dix.) R.H.Zander (Pottiaceae)
rotten wood, rock and soil in montane forests, (100-)1000- Figure 1C
2500 m. Distribution and habitat: northern India (Darjeeling,
Sexuality and dispersal: dioicous or monoicous; spores 8- Sikkim, Uttar Pradesh: Lucknow) and southeastern Brazil
13 µm; specialized asexual reproduction not observed. (Itatiaia municipality); on soil over rock, shaded, at low
Reference: Magombo (2003), Karén et al. (2010). elevations.
Sexuality and dispersal: monoicous; spores 10-13 µm;
Diphyscium longifolium Griff. (Diphysciaceae) Figure 1B specialized asexual reproduction not observed.
Distribution and habitat: scattered in tropical South Note: Ganguleea is monospecific and characterized by
America (Colombia, Venezuela, Suriname, Peru, Bolivia, SE pleurocarpous habit and monoicy.
Brazil) and Panama, widespread in Southeast Asia and Reference: Schäfer-Verwimp (1992), Zander (1993),
Melanesia (India to Samoa); on rock and soil, sometimes Bansal et al. (2015).
submerged, from sea level to 2500 m.
Sexuality and dispersal: dioicous (sometimes Hydrogonium arcuatum (Griff.) Wijk & Margad. (= Barbula
pseudautoicous); spores 8-14 µm; specialized asexual arcuata Griff.) (Pottiaceae)
reproduction not observed. Distribution and habitat: widespread in tropical and
Reference: Magombo (2003). subtropical Southeast Asia and tropical America; on rock,
damp soil and tree bases; in Asia mainly below 1500 m, in
Elmerobryum Broth. (Hypnaceae) Figure 2A the Neotropics up to 2000 m.
Distribution and habitat: two species in Southeast Asia (E. Sexuality and dispersal: dioicous; spores small; asexual
philippinense Broth.: Luzon, Taiwan; E. wilhelmense reproduction by rhizoidal tubers.
[E.B.Bartram] W.R.Buck & B.C.Tan: Papua New Guinea) and Reference: Eddy (1990), Allen (2002), Sollman &
one, E. guatemalense J.R.Rohrer, in Central America Koponen (2017).
(Guatemala to Costa Rica); on bark, rock and humic soil in
moist but usually rather open habitats, 1200-4000 m. Hymenodon Hook.f. & Wilson (Rhizogoniaceae)
Sexuality and dispersal: dioicous; spores 22-26 µm (E. Distribution and habitat: four to eight species in Southeast
philippinense); specialized asexual reproduction not Asia and Australasia, extending eastwards to New Caledonia
observed. and Fiji and southwards to southern Australia; more rare in
Note: the genus name is sometimes spelled the Neotropics with one species in southeastern Brazil and a
"Elmeriobryum" but according to the ICN the spelling second one in the West Indies (Jamaica, Dominican
Elmerobryum is correct. Republic); on tree trunks, especially tree ferns in
Reference: Buck & Tan (2007). submontane and lower montane areas to ca. 2000 m.
Sexuality and dispersal: dioicous; spores small; asexual
Fissidens sect. Sarawakia (Müll.Hal.) Pursell & Brug.-Nann. reproduction rare, by axillary hairs in H. reggaeus Kartt. &
(Fissidentaceae) Figure 2B S.Bäck.
Distribution and habitat: F. beccarii (Hampe) Broth. in Reference: Koponen et al. (1986), Karttunen & Bäck
Sarawak (known from only one locality) and F. hydropogon (1988), Pócs (2007b)
Spruce in Ecuador (known from two localities); on branches
and rock in streams with strongly fluctuating water levels, Hymenostyliella Bartr. (Pottiaceae) Figure 2C
frequently submerged; below 1000 m. Distribution and habitat: one or two species in Asia
Sexuality and dispersal: monoicous and usually highly (Luzon, C India) and one in southeastern Brazil (São Paulo
© Association of Systematic Biologists of the Philippines Volume 12 Issue 1 - 2018 | 6
Philippine Journal of Systematic Biology | Gradstein, S. R.: Amphi-Pacific tropical disjunctions in bryophyte floras
state); on rock and soil near rivers. specialized asexual reproduction not observed.
Sexuality and dispersal: dioicous; spores ca. 10-13 µm; Note: the family contains only the genus Sorapilla.
specialized asexual reproduction not observed. Reference: Norris & Koponen (1987), Catcheside (2012).
Reference: Zander (1993), Alam (2015).
Sphaerotheciella M.Fleisch. (Cryphaeaceae)
Hymenostylium aurantiacum Mitt. (Pottiaceae)
Distribution and habitat: widespread in tropical and Distribution and habitat: three species in subtropical and
subtropical Asia and the Neotropics, in tropical America warm-temperate eastern Asia (Himalayas, China) and two in
restricted to montane areas, above 500 m (Cano & Jimenez, tropical America (Mexico, C America, Bolivia); epiphytic,
2013); on calcareous rock near streams. Sexuality and dispersal: monoicous; spores multicellular,
Sexuality and dispersal: dioicous; spores ca. 11-16 µm; to 65 × 160 µm in mature capsules; specialized asexual
specialized asexual reproduction not observed. reproduction not observed.
Reference: Cano & Jimenez (2013), Sollman & Koponen Note: Sphaerotheciella differs from other genera of
(2017). Cryphaeaceae in the large, multicellular spores due to
endosporous development of the prothallium. This is an
Luisierella barbula (Schwaegr.) Steere (Pottiaceae) Figure 1D adaptive feature of epiphytes (Nishida, 1978).
Distribution and habitat: rather widespread in Southeast Reference: Manuel (1981), Rao (2000).
Asia (Japan, Hong Kong, Malaysia, Java), in tropical America
occurring disjunctly in the north (southeastern USA, Mexico, 2. LIVERWORTS (Marchantiophyta)
Cuba and Jamaica) and along the Atlantic coast of Brazil
(Ceara, Bahia); on rather dry limestone rock and old, Ceratolejeunea grandiloba J.B.Jack & Steph.
plastered walls, at low elevations. (Lejeuneaceae) Figure 1E
Sexuality and dispersal: monoicous or dioicous; spores ca. Distribution and habitat: subsp. grandiloba in tropical
8-10 µm; specialized asexual reproduction not observed. South America (Andes, Guyana Highland) and subsp. inflata
Note: Luisierella is monospecific. (Mizut.) Gradst. in Java (Gunung Pangrango; only known
Reference: Yano (1981), Crum & Anderson (1981), from the type specimen); epiphytic in forest canopy and
Deguchi (1987), Zander (1993), Deguchi et al. (1994). rather open, humid montane habitats, 1400-3000 m.
Sexuality and dispersal: dioicous; spores multicellular,
Mniomalia Müll.Hal. (Phyllodrepaniaceae) Figure 2D 20-40 x 45-70 µm; specialized asexual reproduction not
Distribution and habitat: M. semilimbata (Mitt.) Müll.Hal. in observed.
Southeast Asia, northeastern Australia and the Pacific region Reference: Gradstein (2013).
(China to Samoa) and M. viridis (Mitt.) Müll.Hal. in C America,
the Guianas and the Amazon basin; epiphytic in rather open Drepanolejeunea subg. Rhaphidolejeunea (Herzog) Grolle &
habitats. R.L.Zhu (Lejeuneaceae) Figure 3A
Sexuality and dispersal: dioicous, sporophytes rare; Distribution and habitat: ten species in Southeast Asia
asexual reproduction by caducous rhizoids. and one in tropical America (rainforests of the Amazon
Reference: Norris & Koponen (1987), Florschütz-de Waard basin, Guianas and Chocó region); usually epiphyllous, from
(1987), TROPICOS. sea level to 2500 m.
Sexuality and dispersal: monoicous or dioicous; spores
Rozea Besch. (Brachytheciaceae) Figure 2E multicellular; asexual reproduction by cladia.
Distribution and habitat: three species in subtropical Reference: Bischler (1968), Grolle & Zhu (2000), Dey &
eastern Asia (NW India, Myanmar, China) and two in Mexico Singh (2012), Pócs et al. (2013).
and Central America (Mexico to Venezuela); on bark, rotten
wood and rock at high elevations, ca. 2000-4000 m. Lejeunea sect. Echinocolea (R.M.Schust.) Gradst. comb.
Sexuality and dispersal: monoicous or dioicous; spores nov. (Lejeuneaceae)
small; specialized asexual reproduction not observed. Echinocolea R.M.Schust., Beih. Nova Hedwigia 9: 125.
Reference: Buck & Crum (1976). 1963.
Map: Ilkiu-Borges (2005: Figures 11, 12).
Sorapilla Spruce & Mitt. (Sorapillaceae) Figure 2F Distribution and habitat: four species in tropical America
Distribution and habitat: S. sprucei Mitt. in Ecuador and S. and one, L. reinerae Ilkiu-Borges, in eastern Malesia
papuana Broth. & Geheeb in eastern Malesia (Ceram, Papua (Sulawesi, Papua New Guinea); on bark, rotten wood and
New Guinea) and Queensland; epiphytic, montane. living leaves in the understory and canopy of virgin lowland
Sexuality and dispersal: monoicous; spores ca. 10-12 µm; and montane rain forest, from sea level to 2650 m (Ilkiu-
© Association of Systematic Biologists of the Philippines Volume 12 Issue 1 - 2018 | 7
Philippine Journal of Systematic Biology | Gradstein, S. R.: Amphi-Pacific tropical disjunctions in bryophyte floras
Borges, 2005). The diagnostic features of sect. Myriocoleopsis are
Sexuality and dispersal: monoicous or dioicous; spores considered adaptations to its rheophytic growth (sect.
multicellular; asexual reproduction by caducous branches, Protocolea is not rheophytic).
leaf fragmentation and strap-shaped gemmae. Reference: Pócs (2010), Ye et al. (2013, 2014).
Note: Ilkiu-Borges (2005) monographed Echinocolea
and showed that the group is part of the large genus Phycolepidozia R.M.Schust. (Cephaloziellaceae) Figure 3C
Lejeunea; the group has since been treated as a synonym of Distribution and habitat: P. exigua R.M.Schust. in the
Lejeunea. However, a recent molecular analysis of Lejeunea West Indies (Dominica) and Venezuela (Mt. Duida), and P.
resolved two common Echinocolea species (L. asperrima indica Gradst. et al. in the Western Ghats, India; on bark,
Spruce, L. subspathulata Spruce) in a strongly supported rock and soil, 450-1850 m.
separate lineage within Lejeunea (Heinrichs et al., 2013). Sexuality and dispersal: monoicous; spores 11-15 µm;
Morphologically, Echinocolea is a quite homogeneous group specialized asexual reproduction not observed.
characterized by asperulate leaves. The group is therefore Reference: Gradstein et al. (2014).
resurrected here as a section of Lejeunea. A broader
molecular sampling of sect. Echinocolea including more Pictolejeunea Grolle (Lejeuneaceae) Figure 3D
species is recommended. Distribution and habitat: five species in tropical America
(Central America, West Indies, northern South America) and
Lobatiriccardia (Mizut. & S.Hatt.) Furuki (Aneuraceae) one in North Borneo (Sabah); on bark, rock and rotten wood,
Map: Preußing et al. (2010: Figure 7). sea level to 1350 m.
Distribution and habitat: six species in Southeast Asia Sexuality and dispersal: monoicous or dioicous; spores
and Australasia, two in the high Andes of Ecuador; on damp multicellular; asexual reproduction by long, ribbon-like
soil, rock and rotten wood, lowlands to 4000 m. gemmae.
Sexuality and dispersal: dioicous; spores 10-21 µm; Reference: Grolle (1977), Pócs (2007a).
specialized asexual reproduction not observed.
Note: the preliminary record of Lobatiriccardia from Africa Rectolejeunea A.Evans (Lejeuneaceae) Figure 3E
(Reeb & Bardat, 2014) belongs to the new genus Distribution and habitat: four species in tropical America
Afroriccardia Reeb & Gradst. (Rabeau et al., 2017). and one in NE Queensland; sun epiphytes, occurring
Reference: Preußing et al. (2010), Nebel et al. (2013). preferably on twigs and living leaves, sea level to 2700 m.
Sexuality and dispersal: monoicous or dioicous; spores
Myriocoleopsis Schiffn. sect. Myriocoleopsis Figure 3B multicellular; asexual reproduction by caducous leaves,
Distribution and habitat: two species in South America frequently very small and produced on erect, flagelliform
(Ecuador, SE Brazil, N Argentina) and one in Vietnam; shoots.
rheophytic on periodically submerged rocks, twigs and living Reference: Reiner-Drehwald & Grolle (2012).
leaves in rivers with strongly fluctuating water levels, 100-
1300 m. Southbya organensis Herzog (Southbyaceae) Figure 1F
Sexuality and dispersal: monoicous; spores multicellular; Distribution and habitat: scattered in Southeast Asia (Sino
asexual reproduction by multicellular, disciform gemmae. -Himalayan region, Srí Lanka, Thailand, New Guinea),
Note: based on the results of morphological and Hawaii; very rare in South America (SE Brazil, Peru); on wet
molecular-phylogenetic analysis (Ye et al., 2013, 2014), the rock and soil, occasionally on bark, at high elevation, ca.
genus Myriocoleopsis may be divided into two sections, the 1700-3400 m.
amphi-Pacific M. sect. Myriocoleopsis (M. fluviatilis [Steph.] Sexuality and dispersal: dioicous; spores 18-23 µm;
M.E.Reiner & Gradst., M. gymnocolea [Spruce] M.E.Reiner & asexual reproduction by gemmae, the gemmae 1-2-celled,
Gradst., M. vuquangensis [Pócs & T.N.Ninh] Pócs) and the elliptical, 28-50 µm long.
pantropical M. sect. Protocolea (R.M.Schust.) Gradst. comb. Reference: Váňa et al. (2012).
et stat. nov. (= Cololejeunea subg. Protocolea R.M.Schust.,
Beih. Nova Hedwigia 9: 171. 1963) with one species, M. Vitalianthus R.M.Schust. & Giancotti Figure 3F
minutissima (Sm.) R.L.Zhu et al. The members of sect. Distribution and habitat: V. aphanella (Spruce) Bechteler
Myriocoleopsis stand out by rigid leafy shoots lacking a et al. in the Rio Negro and Vaupés region of Amazonia
hyalodermis and arising from a creeping stoloniform stem, (Brazil, Colombia), V. bischlerianus (K.C.Porto & Grolle)
very long androecial spikes and clustered gynoecia. In R.M.Schust. & Giancotti in the Atlantic coastal rainforests of
contrast, sect. Protocolea has fragile leafy shoots that are not Brazil (Pernambuco to Paraná), and V. guangxianus R.L.Zhu
arising from a creeping stoloniform stem and possess a et al. in southern China (Guangxi).
hyalodermis, short androecial spikes and gynoecia singly. Sexuality and dispersal: monoicous; spores multicellular;
© Association of Systematic Biologists of the Philippines Volume 12 Issue 1 - 2018 | 8
Philippine Journal of Systematic Biology | Gradstein, S. R.: Amphi-Pacific tropical disjunctions in bryophyte floras
asexual reproduction by fragmentation. Deguchi, H., 1987. Luisierella (Pottiaceae, Musci) a moss
Note: according to R.L. Zhu (pers. com.) further species genus with a disjunctive distribution in Neotropics and
might be attributable to Vitalianthus. Japan. Journal of Japanese Botany, 62: 7-14.
Reference: Gradstein & Costa (2003), He et al. (2012), Deguchi, H., S. Sulastri, H. Sujadmiko & T. Seki, 1994.
Bechteler et al. (2016). Three pottiaceous mosses from Indonesia. Hikobia, 12:
37-41.
ACKNOWLEDGMENTS Delgadillo, C., 1993. The Neotropical-African moss
disjunction. The Bryologist, 96: 604-615.
I am grateful to Professor Hironori Deguchi (Hiroshima Dey, M. & D.K. Singh, 2012. Epiphyllous liverworts of
University) for references, to Dr. Masanobu Higuchi (National Eastern Himalaya. Botanical Survey of India, Kolkata.
Museum for Nature and Science, Tsukuba, Japan) for data on 415 pp.
endemism in Japan, to Didier Geffard-Kuriyama (UMS2700, Eddy, A., 1990. A handbook of Malesian mosses. Natural
CNRS, MNHN Paris) for preparing the maps for publication, History Museum Publications, London. 256 pp.
and to an anonymous reviewer for constructive comments Florschütz–de Waard, J., 2011. Flora of the Guianas, Musci,
and corrections on the manuscript. IV: 1-432.
Frahm, J.P., 1984. A review of Campylopodiella Card. The
LITERATURE CITED Bryologist, 87: 249-250.
Fritsch, P.W., S.R. Manchester, R.D. Stone, B.C. Cruz & F.
Alam, A., K.K. Rawat, P.K. Verma, V. Sharma & D.S. Gupta, Almeda, 2015. Northern Hemisphere origins of the amphi
2015. Moss flora of Central India. Plant science today, 2: -Pacific tropical plant family Symplocaceae. Journal of
159-171. Biogeography, 42: 891-901.
Allen, B., 2002. Moss flora of Central America. Part 2. Gradstein, S.R., 2013. Afro-American hepatics revisited.
Missouri Botanical Garden, St. Louis. 699 pp. Polish Botanical Journal, 58: 149-177.
Gradstein, S.R., 2017. Amphitropical disjunctive species in
Bechteler J., A. Schäfer‐Verwimp, G.E. Lee, K. Feldberg,
the complex thalloids (Marchantiidae). Journal of
O.A. Pérez‐Escobar, T. Pócs, D. Peralta, M.A.M. Renner
Bryology, 39: 66-78.
& J. Heinrichs, 2017. Geographical structure, narrow
species ranges, and Cenozoic diversification in a Gradstein, S.R., S.P. Churchill & N. Salazar Allen, 2001.
pantropical clade of epiphyllous leafy liverworts. Ecology Guide to the Bryophytes of Tropical America. Memoirs of
and Evolution, 7: 638-653. the New York Botanical Garden, 86: 1-577.
Gradstein, S.R. & D.P. Costa, 2003. The Liverworts and
Bernal, R., S.R. Gradstein & M. Celis, 2016. Catálogo de
Hornworts of Brazil. Memoirs of the New York Botanical
plantas y líquenes de Colombia. Instituto de Ciencias
Garden, 87: 1-317.
Naturales, Universidad Nacional de Colombia, Bogotá.
Gradstein, S.R., A.L. Ilkiu-Borges & A. Vanderpoorten, 2011.
3058 pp.
Habitat specialization triggers the evolution of unusual
Bischler, H., 1968. Monographie du genre Rhaphidolejeunea
morphologies: the case of Cololejeunea stotleriana sp.
Herzog. Revue Bryologique et Lichénologique, 36: 56-
nov. from Ecuador. The Bryologist, 114: 9-22.
104.
Gradstein, S.R., B. Laenen, J.P. Frahm, U. Schwarz, B.J.
Bischler, H., 1969. Le genre Leptolejeunea (Spruce) Steph.
Crandall-Stotler, J.J. Engel, M. von Konrat, R.E. Stotler,
en Amérique. Nova Hedwigia, 17: 265-350.
B. Shaw & A.J. Shaw, 2014. On the taxonomic status of
Buck, W.R. & H.A. Crum, 1976. Revision of the genus Rozea the enigmatic Phycolepidoziaceae (Marchantiophyta:
(Musci). The Bryologist, 79: 406-421. Jungermanniales) with description of a new species,
Buck, W.R. & H.A. Crum, 1978. A re-interpretation of the Phycolepidozia indica. Taxon, 63: 498-508.
Fabroniaceae with notes on selected genera. Journal of Gradstein, S.R., T. Pócs & J. Váňa, 1983. Disjunct
the Hattori Botanical Laboratory, 44: 347-369. Hepaticae in Tropical America and Africa. Acta Botanica
Buck, W.R. & B.C. Tan, 2007. A review of Elmeriobryum Hungarica, 29: 127-171.
(Hypnaceae). Telopea, 12: 251-256. Gradstein, S.R., R.L. Zhu, L. Shu & A.L. Perez, 2018.
Cano, M.J. & J.A. Jimenez, 2013. A taxonomic revision of the Reinerantha foliicola, a new genus and species of
tribe Pleuroweisiae (Pottiaceae, Bryophyta) in South Lejeuneaceae subtribe Cololejeuneinae
America. Phytotaxa, 143: 1-42. (Marchantiophyta) from Ecuador. Journal of Systematics
Catcheside, D.G., 2012. Sorapillaceae. Australian mosses and Evolution, 56: 67-75.
online, 5: 1-2.
Grolle, R., 1977. Pictolejeunea – eine neue Gattung der
Crum, H.A. & L.E. Anderson, 1981. Mosses of North America
Lejeuneoideae aus der Neotropis und Borneo.
(Vol. 1). Columbia University Press, New York, pp. 1-663.
Feddes Repertorium, 88: 247-256.
© Association of Systematic Biologists of the Philippines Volume 12 Issue 1 - 2018 | 9
Philippine Journal of Systematic Biology | Gradstein, S. R.: Amphi-Pacific tropical disjunctions in bryophyte floras
Grolle, R. & Zhu, R.L. 2000. A study of Drepanolejeunea Phytotaxa, 81: 10-11.
subg. Rhaphidolejeunea (Herzog) Grolle & R.L. Zhu, Nishida, Y., 1978. Studies on the sporeling type in mosses.
stat. nov. (Hepaticae, Lejeuneaceae) in China with Journal of the Hattori Botanical Laboratory, 44: 371-454.
notes on its species elsewhere. Nova Hedwigia, 70: Norris, D. H. & T. Koponen, 1987. Bryophyte flora of the
373-396. Huon Peninsula, Papua New Guinea. XX.
He, Q., Y.M. Wei & R.L. Zhu, 2012. Vitalianthus guangxianus Fissidentaceae, Mitteniaceae, Phyllodrepaniaceae,
(Lejeuneaceae), sp. nov. from Guangxi, China. Journal of Phyllogoniaceae and Sorapillaceae. Annales Botanici
Bryology, 34: 32-36. Fennici, 24: 177–219.
Heinrichs, J., S. Dong, A. Schäfer-Verwimp, T. Pócs, K. Patiño, J., F. Guilhaumon, R.J. Whittaker, K.A. Triantis, S.R.
Feldberg, A. Czumaj, A.R. Schmidt, J. Reitner, M.A.M. Gradstein, L. Hedenäs, J.M. González-Mancebo &
Renner, J. Hentschel, M. Stech & H. Schneider, 2013. A.Vanderpoorten, 2013. Accounting for data
Molecular phylogeny of the leafy liverwort Lejeunea heterogeneity in patterns of biodiversity: an application of
(Porellales): Evidence for a Neotropical origin, uneven Linear Mixed Models to the oceanic island biogeography
distribution of sexual systems and insufficient taxonomy. of spore–producing plants. Ecography, 36: 904-913.
PloS ONE, 8: e82547. Pócs, T., 1976. Correlation between the tropical African and
Heinrichs, J., J. Hentschel, K. Feldberg, A. Bombosch & H. Asian bryofloras. I. Journal of the Hattori Botanical
Schneider, 2009. Phylogenetic biogeography and Laboratory, 41: 95–106.
taxonomy of disjunctly distributed bryophytes. Journal of Pócs, T., 1992. Correlation between the tropical African and
Systematics and Evolution, 47: 497-508. Asian bryofloras. II. Bryobrothera, 1: 35–47.
Higuchi, M., 2011. Endemic species of bryophytes in Pócs, T., 2007a. A sixth species of Pictolejeunea
Japan. Bulletin of the National Museum of Natural (Jungermanniopsida) from the Venezuelan Guyana. Acta
Sciences, Ser. B, 37: 117–126. Botanica Hungarica, 40: 109-119.
Ilkiu-Borges, A.L., 2005. A taxonomic revision of Echinocolea Pócs, T., 2007b. Bryophytes from Fiji Islands, I. Hymenodon
(Lejeuneaceae, Hepaticae). Nova Hedwigia, 80: 45-71. chenianus Pócs, sp. n. (Rhizogoniaceae) and
Karén, V., J. Enroth & T. Koponen, 2010. Bryophyte flora of Ephemeropsis tjibodensis Goebel. Chenia, 9: 25‒38.
Hunan Province, China. 12. Diphysciaceae (Musci). Pócs, T., 2010. Myriocoleopsis in Southeast Asia. Tropical
Annales Botanici Fennici, 47: 208-214. Bryology, 31: 123-125.
Karttunen, K. & S. Bäck, 1988. Taxonomy of Hymenodon Pócs, T., A. Bernecker-Lücking & P. Tixier, 2014. Synopsis
(Musci, Rhizogoniaceae). Annales Botanici Fennici, 25: and key to species of neotropical Cololejeunea
89-95. (Lejeuneaceae). Acta Botanica Hungarica, 56: 185-226.
Kato, M. & A. Ebihara, (eds.), 2011. Endemic Plants of Japan. Pócs, T, T.T. Luong & B.C. Ho, 2013. New or little known
A book series from the National Museum of Nature and epiphyllous liverworts, XVIII. Cryptogamie, Bryologie, 34:
Science, 11: 1-503. [Tokai University Press, Hatano]. 287-298.
Koponen, T., A. Touw & D. Norris, 1986. Bryophyte flora of Preußing, M., A. Olsson, A. Schäfer-Verwimp, N.J. Wickett,
Papua New Guinea. XIV. Rhizogoniaceae (Musci). Acta S. Wicke, D. Quandt & M. Nebel, 2010. New insights in
Botanica Fennica, 133: 1-24. the evolution of the liverwort family Aneuraceae
Laenen, B., A. Machac, S.R. Gradstein, B. Shaw, J. Patiño, (Metzgeriales, Marchantiophyta) with an emphasis on
A. Desamoré, B. Goffinet, C. Cox, A. Vanderpoorten & the genus Lobatiriccardia. Taxon, 59: 1424-1440.
A.J. Shaw, 2016. Range size in liverworts: does sex really Printarakul, N., B.C. Tan, K. Santanachote & H. Akiyama,
matter? Journal of Biogeography, 43: 627–635. 2013. New and noteworthy records of mosses from Doi
Li, R. & J. Wen, 2013. Phylogeny and biogeography of (Mt.), Inthanon, Chiang Mai, Chom Tong District,
Dendropanax (Araliaceae), an amphi-Pacific disjunct Northern Thailand. Polish Botanical Journal, 58: 245-
genus between tropical/subtropical Asia and the 257.
Neotropics. Systematic Botany, 38: 536-551. Pursell, R.A. & M.A. Nannenga-Bremekamp, 2004. A
Magombe, Z.L.K., 2003. Taxonomic revision of the moss revision of the infrageneric taxa of Fissidens. The
family Diphysiaceae M. Fleisch. Journal of the Hattori Bryologist, 107: 1-20.
Botanical Laboratory, 94: 1-86. Pursell, R.A., M.A. Nannenga-Bremekamp & B.H. Allen,
Manuel, M., 1981. Studies in Cryphaeaceae V. A revision of 1988. A taxonomic revision of Fissidens subgenus
the family in Mexico, Central America and the Caribbean. Sarawakia (Bryopsida: Fissidentaceae). The Bryologist,
Journal of the Hattori Botanical Laboratory, 49: 115-140. 91: 202-213.
Nebel, M., L. Söderström, A. Hagborg & M. von Konrat, 2013. Rabeau, L., M. Nebel, J.Y. Dubuisson, C. Krause, R.
Notes on Early Land Plants Today 28. Transfers of some Gradstein, D. Quandt & C. Reeb, 2017. New insights into
taxa to Lobatiriccardia (Aneuraceae, Marchantiophyta). the phylogeny and relationships among the worldwide
© Association of Systematic Biologists of the Philippines Volume 12 Issue 1 - 2018 | 10
