Amphi-Pacific Tropical Disjunctions in the Bryophyte Floras of Asia and the New World

PRIMARY 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, Evolution, Biodiversité / Grand Herbier, Case Postale 39, 12 rue 1972). Over 100 genera and higher taxa of vascular plants Buffon, 75005 Paris. France exhibiting this type of distribution were enumerated by 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

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

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

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.

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

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

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-

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;

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