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The New Genus Benitotania (Daltoniaceae, Bryopsida) from Mt. Kinabalu
Abstract
A new monospecific genus, Benitotania, is described on the basis of B. elimbata H. Akiyama, T. Yamag, & M. Suleiman, collected in the northern part of Sabah, Malaysia. It appears to be most closely related to Adelothecium and Bryobrothera.
... Mount Tambuyukon (2570 m) in Kinabalu Park (Sabah, Malaysia) is the largest ultramafic mountain on Borneo Island and quite rich in plant species (Van der Ent et al., 2014). During a field survey carried out on this mountain in August 2008, the junior author collected a curious moss in a mossy forest looking quite similar to Benitotania elimbata H.Akiyama et al. (2003: 456), formerly reported from Kinabalu Park (Akiyama et al. 2003). The plants were collected again during a scientific expedition to Sungai Imbak Forest Reserve (Sabah, Malaysia) in April 2014. ...
... Benitotania elimbata, however, differs from the Sabah plant in a number of morphological features, such as (1) shorter leaves (ca. 2.0 mm long), (2) totally plane lamina, (3) short rectangular to hexagonal upper and median laminal cells, (4) slight mamillation on laminal cells, (5) slender costa not reaching near leaf apices, and (6) short 3–4 cells axillary hairs (Akiyama et al. 2003). While, Adelothecium bogotense differs from the Sabah plant in the features, such as (1) presence of microphyllous and gemmiferous tips of secondary stems bearing clusters of minute gemmae, (2) broadly ovate to obovate, rounded or rounded-obovate leaves, (3) totally plane lamina, and (4) nearly isodiametric upper lamina cells (Ochyra et al. 1992, Whittemore & Allen 1989). ...
We report a new species Bryobrothera tambuyukonensis, based on the specimens collected in Sabah, Borneo Island. Phylogenetic analyses with plastid (rps4 and trnL-F) and mitochondrial (nad5) genes along with a number of morphological features (for example, undulate lamina, oblong-lanceolate leaves with a single strong costa reddish brown in color, and thick-walled and porose laminal cells) confirmed its distinctiveness from B. crenulata and also close relationship to the elimbate group comprised of Adelothecium bogotense and Benitotania elimbata.
... These were excludedTable 1 Literature sources for information on sexual conditions in homocostate pleurocarpous mosses. Afonina and Ignatova (2007); Akiyama (1988); Akiyama et al. (2003); Allen and Magill (2007); Allen (1987a Allen ( ,b,c, 1990 Allen ( , 1999); Allen and Crosby (1986a,b); Allen et al. (1985a Allen et al. ( , 1986 (1986, 1987, 1994, 1996, 1997); Higuchi (1983, 1984 (1992, 1993, 1994); Buck (1997 Buck ( , 1998 (1989, 1991, 1993, 1994a (1969, 1974, 1990, 1992a,b); Buck (1994, 2009); Iwamasa (1940); Iwatsuki (1958 Iwatsuki ( , 1963 Iwatsuki ( , 1966 Iwatsuki ( , 1970 Iwatsuki ( , 1979 Iwatsuki ( , 1992 (1970, 1971, 1972, 1976, 1989, 1991, 1994 (1991a,b, 1992, 1997, 1999, 2000, 2001);Takaki ( (1943, 1962, 1966, 1971, 1972a,b, 1974); Veling (1996); Veloira (1959); Williams (1924 Williams ( , 1925 Williams ( , 1930a Williams ( ,b, 1931 Williams ( , 1932 Investigations made specifically to assess the occurrence of dwarf versus normal male plants in a selection of species. Species selection was arbitrary, with a focus on species with seemingly abundant sporophyte production and few or unknown normal males. ...
Dwarf males occur in several animal groups but are unique to mosses among the green land plants. Mosses, with more than 50% unisexual species and common fertilisation distances in the range of a few decimetres, have evolved various means to cope with potential problems to successfully achieve fertilisation and to ensure sporophyte production and sexually produced diaspores. We explore the abundance of different kinds of extreme female-biased sexual size dimorphism in homocostate pleurocarpous mosses. Based on direct observations and literature information, we investigated the large-scale geographical distribution of this phenomenon, and whether male dwarfism was associated with family position. Almost 60% of 1737 species with information about sexuality are dioecious, and males are known in 769 dioecious species. Dwarf males occur in 178 of these (23%), and are considered obligate in 113 species (63% of 178). We thoroughly examined a subset of 162 species. Among these 72 species (44%) form dwarf males, but only 18 species (25% of 72) produce obligate dwarf males. We conclude that especially facultative male dwarfism is much overlooked among dioecious mosses. The distribution of dioecious species and dwarf males among the latter was related to family membership, but showed little correlation with geographic area. Male dwarfism thus appears useful as a taxonomic character to define bryophyte families. We estimate that between roughly one quarter and almost half of dioecious pleurocarpous moss species exhibit male dwarfism, and the majority of these (75%) form dwarf males facultatively. Male dwarfism in mosses is suggested to have evolved as a result of (1) competing selective pressures of the cytoplasmic and nuclear genomes, (2) competition between the sexes, (3) selection for reduced fertilisation distances, or a combination of these. The associated niche shift of the males may also provide these with a suitably humid and nutrient-rich habitat.
... The monophyly of, and relationships among, Adelothecium, Benitotania and Bryobrothera have been assessed without disagreement (Akiyama et al., 2003;Buck et al., 2005;Gradstein & Wilson, 2008) and are re-confirmed here. However, the sistergroup relationship of this clade with the genus Ephemeropsis, found here, does not agree with the findings of Buck et al. (2005) or Gradstein & Wilson (2008). ...
Phylogenetic relationships in Daltoniaceae (~200 species in 14 genera) are inferred from nucleotide sequences from five genes, representing all genomic compartments, using parsimony, likelihood and Bayesian methods. Alternative classifications for Daltoniaceae have favoured traits from either sporophytes or gametophytes; phylogenetic transitions in gametophytic leaf limbidia and sporophytic exostome ornamentation were evaluated using ancestral state reconstruction to assess the levels of conflict between these generations. Elimbate leaves and the cross-striate exostome are reconstructed as plesiomorphic states. Limbate leaves and papillose exostomes evolved at least two and six times, respectively, without reversals. The evolution of leaf limbidia is relatively conserved, but exostome ornamentation is highly homoplasious, indicating that superficial similarity in peristomes gives unreliable approximations of phylogenetic relatedness. Our phylogenetic analyses show that Achrophyllum and Calyptrocha- eta are reciprocally monophyletic. Within core Daltoniaceae, relationships among taxa with elimbate leaves are generally well understood. However, taxa with limbate leaves form a monophyletic group, but resolved subclades correspond to biogeographical entities, rather than to traditional concepts of genera. Daltonia (~21 species), Distichophyllum (~100 species) and Leskeodon (~20 species) are polyphyletic. Seven nomenclatural changes are proposed here. As the current taxonomy of Daltoniaceae lacks phylogenetic consistency, critical generic revisions are needed.
Southeast Asia has the highest rate of deforestation among all tropical regions in the world. Depending on the number of undiscovered
species not yet known to science, a sizeable proportion of species may have gone extinct or will go extinct in the future
without record. We compiled species datasets for eight taxa, each consisting of a list of native species and their description
dates. Birds, legumes, mosquitoes, and mosses showed recent declines in species discovery rate. For these taxa, we estimated
the total species richness by applying generalized linear models derived from theory. The number of undiscovered species in
each taxon was calculated and the extent of undiscovered species among the taxa compared. Among these taxa that displayed
a species discovery decline, the legumes had the highest extent of undiscovered species while the birds had the most complete
species inventory. Although quantitative estimates of the number of undiscovered species for amphibians, freshwater fish,
hawkmoths, and mammals could not be derived, the extent of undiscovered species is likely to be high as their recent discovery
rates showed a continued increase. If these taxa are more or less representative of other Southeast Asian taxa, many species
are likely to go extinct before ever being discovered by science under the current rates of habitat loss. We therefore urge
the intensification of taxonomic and species discovery research in the taxa in which the extent of undiscovered species is
relatively high, i.e., amphibians, freshwater fish, hawkmoths, mammals, and legumes.
KeywordsBiodiversity-Conservation-Species discovery-Southeast Asia-Species richness
Classification of families of hypnobryalean mosses into the Hypnales, Leucodontales, and Hookeriales has been taxonomically difficult. Several researchers have sequenced different genes for independent phylogenetic studies of these three pleurocarp groups. Our goal is to summarize available molecular data and compile the largest data set to infer phylogenetic rela- tionships among families as basis for classification at ordinal level. Sequences of rbcL, trnL-F, and rps4 loci for 38 exemplars of most families of Hypnales, Leucodontales, and Hookeriales were analyzed to evaluate whether or not each of the three orders is monophyletic. Cladistic analyses of combined sequences, using five taxa in the Bryales as outgroups, reveal a robust clade (decay 5) including all hypnobryalean pleurocarps. Within this group, one clade (decay 2) includes only taxa of the Hookeriales, and is sister to a large monophyletic group (Hypnales sensu lato) containing all other taxa (decay 2) previously in the Leucodontales and Hypnales. These relationships suggest that the ordinal level taxonomy needs to be reconsidered since major line- ages detected do not correspond to the traditional Leucodontales or Hypnales. These two orders are not supported by any molecular evidence fromrbcL, trnL-F, or rps4, either analyzed singly or in different combinations. Additionally, present results indicate the need for changes to the current system of three suborders of Hypnales and four of the Leucodontales. Phylogenetic re- constructions based on molecular data emphasize the need for a re-examination of the taxonomic relevance of morphological characters and corroborate previous interpretations of sporophytic morphological similarities as multiple transitions to similar solutions to epiphytism among the pleurocarps.
The diplolepideous-alternate peristome, when most highly developed, has endostome segments attached to a basal membrane and positioned alternate to the outer exostome teeth, with cilia often present between the segments. This peristome type defines the Bryidae ( sensu Vitt et al. 1998), which includes four orders: the Bryales, Leucodontales, Hypnales, and Hookeriales, of which the latter three are mainly pleurocarpous in their growth form. Chloroplast (rbcL, rps4, and trnL-trnF) and nuclear (18S rRNA) gene sequences have been analyzed using the parsimony optimality criterion to elucidate relationships among the Bryidae. The analyses strongly support the paraphyly of the Bryidae, with the Splachnidae, and possibly the Orthotrichidae, having arisen from ancestors within the Bryidae. The Leucodontales, Hypnales, and Hookeriales form a monophyletic group, as do the pleurocarpous members of the Bryidae. However, the two pleurocarpous clades are not resolved as sister groups, although their non-monophyly is not supported by the bootstrap. The phylogenetic hypothesis provides a context in which to infer evolutionary transitions in some key morphological characters relating to the peristome and the transition from the acrocarpous to pleurocarpous growth forms.
sequence data (1324 bp on average). Our analyses indicate (1) that Antitrichia is distantly related to the other members of Leucodontaceae and should be excluded from the family, (2) that Cryphaeaceae
form a monophyletic clade, not with Anomodontaceae and Leptodontaceae, but with Leucodontaceae, refuting the placement of
Leucodontaceae and Cryphaeaceae in different superfamilies, (3) that Forsstroemia, variously classified in Leucodontaceae, Cryphaeaceae or Leptodontaceae, forms a monophyletic clade with Neckera (Neckeraceae), and (4) that the presumed monophyly of Anomodon and that of Anomodontaceae are not supported.
The affinities as well as the circumscription of the Orthotrichaceae (Bryopsida), one of the most diverse families of mosses, have been the focus of a controversy for much of the last century. We obtained rbcL sequences for 37 arthrodontous mosses, including 27 taxa of the Orthotrichales. The sequences were analyzed using maximum parsimony and neighbor joining in order to (1) test the monophyly of the Orthotrichales and the Orthotrichaceae; (2) determine their phylogenetic relationships; and (3) test the current subfamilial classification within the Orthotrichaceae. Both analyses suggest that the Orthotrichales are polyphyletic. The Erpodiaceae and the Rhachitheciaceae as well as Amphidium and Drummondia, two genera of the Orthotrichaceae, are shown to be of haplolepideous affinity. The Splachnales, the Bryales sensu lato, and the Orthotrichales form a monophyletic clade sister to the Haplolepideae. Both neighbor joining and maximum parsimony also suggest that the Orthotrichaceae are composed of two major lineages dominated either by acrocarpous or cladocarpous taxa. The monophyly of the family is, however, only well supported by Tamura's distances. The genera Macrocoma, Macromitrium, Orthotrichum, Ulota, and Zygodon all appear to be artificial assemblages. This study illustrates the contribution of rbcL sequence data to bryophyte systematics and, particularly, in determining the affinities of taxa lacking a peristome, whose characters are central to the classification of mosses.
An approximately unbiased (AU) test that uses a newly devised multiscale bootstrap technique was developed for general hypothesis
testing of regions in an attempt to reduce test bias. It was applied to maximum-likelihood tree selection for obtaining the
confidence set of trees. The AU test is based on the theory of Efron et al. (Proc. Natl. Acad. Sci. USA 93:13429–13434; 1996),
but the new method provides higher-order accuracy yet simpler implementation. The AU test, like the Shimodaira–Hasegawa (SH)
test, adjusts the selection bias overlooked in the standard use of the bootstrap probability and Kishino–Hasegawa tests. The
selection bias comes from comparing many trees at the same time and often leads to overconfidence in the wrong trees. The
SH test, though safe to use, may exhibit another type of bias such that it appears conservative. Here I show that the AU test
is less biased than other methods in typical cases of tree selection. These points are illustrated in a simulation study as
well as in the analysis of mammalian mitochondrial protein sequences. The theoretical argument provides a simple formula that
covers the bootstrap probability test, the Kishino–Hasegawa test, the AU test, and the Zharkikh–Li test. A practical suggestion
is provided as to which test should be used under particular circumstances.
Circumscription of the family Sematophyllaceae is controversial, and taxonomic position of the genera Brotherella, Trismegistia, Wijkia and its allies has been repeatedly discussed and is still fluctuating. A phylogenetic study was carried out to investigate the phylogenetic position of the genera Brotherella, Trismegistia, Wijkia and their allies inferred from the chloroplast ribulose 1,5-bisphosphate carboxylase/oxygenase large subunit (rbcL) gene data. The relationships between 40 species representing the Sematophyllaceae and related taxa are evaluated based on the rbcL data set using the following methods: neighbor-joining (NJ), minimum-evolution (ME), maximum-parsimony (MP) and maximum-likelihood (ML) methods. The results of the present study suggest that (1) the family Sematophyllaceae (s. lat.) are monophyletic; (2) the genera Wijkia, Trismegistia and Acanthorrhynchium do not form a single clade; and (3) the genera Brotherella, Pylaisiadelpha, Heterophyllium, Isocladiella, and some species treated as members of the Hypnaceae, such as Hypnum tristo-viride or Isopterygium tenerum should be placed in the Sematophyllaceae (s. lat.). Our research shows that the Sematophyllaceae (sensu Hedenäs & Buck 1999) are not monophyletic. Since the monophyly of the Sematophyllaceae (s. lat.) including Brotherella and its allies is well supported, we propose here that we could treat the family in a wider sense. Furthermore, the present study suggests that Entodontaceae (s. str.) are resolved as a monophyletic group, and should be placed in a clade sister to the Sematophyllaceae (s. lat.). The results show the necessity to reconsider the familial circumscription of the Sematophyllaceae and the Hypnaceae.
Distinction between the Hypnaceae and the Sematophyllaceae is controversial. Taxonomic position of the genera Brotherella, Wijkia and its allies is still fluctuating and has been repeatedly discussed. A phylogenetic study was carried out to investigate the phylogenetic position of the genera Brotherella, Wijkia and their allies, based on phylogenetic analyses of the chloroplast ribulose 1,5-bisphosphate carboxylase/oxygenase large subunit (rbcL) gene data. Chloroplast rbcL genes from eight mosses were newly sequenced. Phylogenetic trees were constructed by neighbor-joining (NJ), minimum-evolution (ME), maximum-parsimony (MP) and maximum-likelihood (ML) methods for 30 rbcL sequences. The results suggest that the genus Brotherella and its allies (Pylaisiadelpha tenuirostris, Wijkia hornschuchii and Heterophyllium nematosum) are monophyletic with high bootstrap support. Hypnum tristo-viride is also included in the Brotherella clade. The Sematophyllaceae (sensu Seki 1968) are paraphyletic to the Brotherella clade. These results support the conclusion that the Sematophyllaceae (sensu present), including Brotherella and its allies, are monophyletic. The present study also suggests that the genus Entodon should be placed in a clade sister to the Sematophyllaceae (sensu present). The results show the need to reconsider the familial circumscription of the Hypnaceae and Sematophyllaceae.
Bryophytes of Kinabalu National Park (including Poring) are reported here based on collections made by two Japan-Malaysia collaborative expeditions held in 1997. Although our research centered mainly on moss flora, some hepatics were also collected. A total of 40 families, 111 genera and 203 species of mosses, as well as 18 families, 25 genera and 31 species of hepatics were found among our collections. Twenty-five species were new to Kinabalu National Park or Sabah, 17 of which were new to Borneo Island.
To study molecular evolution of plants, the plastid encoded rbcL sequences are widely used. In most plastid DNAs, an open reading frame (ORF) designated ycf11 can be found next to the highly conserved rbcL gene. This ORF appears to be only loosely conserved and its function is a matter of debate: On the one hand it is the only gene in plastid DNA of land plants suspected to encode a regulatory zinc finger protein. On the other hand it was postulated to encode the ß-subunit of an acetyl-CoA-carboxylase. Accordingly, this ORF has been previously described as zfpA or accD, respectively. Phylogenetic analysis reveals evolutionary conservation of two ycf11-domains from bacteria to the plastids of dicots. We show that in dicots ycf11 has gained additional sequences through insertions, whereas it has been lost from the plastid DNA in monocots. These findings may reflect physiological differences between major groups of land plants. Furthermore, we show that ycf11 may be a useful molecular marker in the study of plant evolution.
Adelothecium bogotense (Hampe) Mitt. is reported for the first time from continental Africa, where it was collected in the Uluguru Mountains in eastern Tanzania. The African material is described and illustrated, and the world distribution of the species is reviewed and mapped. The African record represents a significant disjunct extension in the range of A. bogotense, and it is suggested that the occurrence of the species in Africa and Madagascar has a long historical foundation.
The monotypic genus, Bryobrothera, is transferred from the Rhizogoniaceae to the Hookeriaceae on the basis of recent collections with mature, calyptrate sporophytes. A more complete description of the species is provided and specimens are cited from northern Queensland, Fiji and the Solomon Islands.
The Hookeriales are evaluated to discern familial limits. Five families are recognized in the order: Hookeriaceae, Leucomiaceae,
Daltoniaceae, Callicostaceae, and Adelotheciaceae fam. nov. Chaetomitrium Dozy & Molk., Chaetomitriopsis. Fleisch., Dimorphocladon Dix., and Elharveya Crum are transferred to the Hypnaceae. Hookeriopsis (Besch.) Jaeg. is split into four genera: Hookeriopsis s. str., Brymela Crosby & Allen, Thamniopsis (Mitt.) Fleisch., and Trachyxiphium Buck, gen. nov. The following new combinations are proposed to coincide with the recognition of these four genera rather
than Hookeriopsis s. lat.: Brymela acuminata, B. callicostelloides, B. cuspidata, B. fissidentoides, B. fluminensis, B. obtusifolia, B. parkeriana, B.
rugulosa, B. websteri, Thamniopsis cheiloneura, T. cruegeriana, T. diffusa, T. incurva, T. langsdorffii, T. pappeana, T. purpureophylla,
T. secunda, T. sinuata, T. terrestris, T. undata, T. utacamundiana, T. versicolor, Trachyxiphium aduncum, T. guadalupense,
T. heteroicum, T. hypnaceum, T. pernutans, T. subfalcatum, T. tenue, T. vagum, and T. variable. Excluded species are treated as Isopterygium plumicaule, Schizomitrium cirrhosum, and S. subsecundum. Schizomitrium belangerianum (Besch.) Buck, comb. nov. is considered distinct from S. depressum (Hedw.) Buck & Steere, and S. pallidum (Hornsch.) Crum & Anders. is shown to have a smooth seta sometimes. The taxonomy of the aquatic species of Cyclodictyon Mitt. and Lepidopilum (Brid.) Brid. is clarified, resulting in the recognition of three species, Cyclodictyon subtortifolium (Bartr.) Buck, comb. nov., C. roridum (Hampe) Kuntze and L. tortifolium Mitt. Two new combinations are proposed in Calyptrochaeta Desv., C. setigera and C. albescens.
Distichophyllum scabrisetum H
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Four hot spots of moss diversity in Malesia
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Funariaceae: Funaria hygrometrica Hedw
- Tsubota
Leucobryaceae: Leucobryum scabrum Sande Lac.
(AB029388, Tsubota et al. 1999). Funariaceae: Funaria
hygrometrica Hedw. (AF005513, Goffinet et al. 1998);
15385 (NY) (AB103354)
- Adelothecium Bogotense
- Churchill Colombia
Adelothecium
bogotense, Colombia, Churchill et al. 15385 (NY)
(AB103354);
Annotated catalogue of the Hepaticae and Anthocerotae of Borneo
———. 1988b. Annotated catalogue of the Hepaticae and
Anthocerotae of Borneo. Journal of the Hattori Botanical Laboratory 65: 145–206.
Adelothecium bogotense (Hampe) Mitt., Colombia, Theirs 3702 (NY) (AB103352)
- Hookeriaceae
Hookeriaceae: Adelothecium bogotense (Hampe) Mitt.,
Colombia, Theirs 3702 (NY) (AB103352);
Streimann 57716 (NY) (AB103353); Hookeria acutifolia Hook
- De Luna
Bryobrothera crenulata (Broth. & Par.) Ther., Australia,
Streimann 57716 (NY) (AB103353); Hookeria acutifolia
Hook. & Grev. (AF158170; De Luna et al. 2000). Hypopterygiaceae: Hypopterygium tamarisci (Sw.) Müll.