Asplenium sylvaticum (Aspleniaceae), a new fern species from Yungas and Paraná forests, Neotropical region.

A new species of Asplenium L. (Aspleniaceae) is described and illustrated. Asplenium sylvaticum is endemic of the Yungas and Paraná biogeographic provinces in the Southern Cone. The main diagnostic characters for this taxon are: apical pinnae with one to four caudate lobes at the base, similar in size to lateral pinnae, smooth pinnate blades with 18-36 pairs of caudate pinnae and spores with reticulate perispore, winged folds with scarce equinulaes on the margins and the laesurae wide with smooth margin. The new species belongs to the A. serra species complex group, characterised by xeromorphic habit, creeping rhizome and coriaceous fronds with branched scales on both surfaces, and resembles the other species inhabiting in Argentina, A. achalense and A. serra, in overall leaf architecture and shape but differs of the aforementioned species (and the other species of the A. serra complex) by the unique combination of characters of its rhizomatic scales, fronds and spores. A key with the morphological and palynological characters that differentiate A. sylvaticum from the other taxa belonging to the A. serra complex in the South American Cone and Bolivia is given.

Germination and gametophyte development of Cyathea corcovadensis (Raddi) Domin (Cyatheaceae) from spores stored at low temperatures / Germinação e desenvolvimento gametofítico de Cyathea corcovadensis (Raddi) Domin (Cyatheaceae) com esporos armazenados em baixas temperaturas

The tree fern Cyathea corcovadensis (Raddi) Domin is an endangered species in the state of Rio Grande do Sul, Brazil. It currently occurs only in the northern segment of the coastal region. Spore storage would help in conservation programs since it maintains genetic variability and provides material for in vitro cultures. Current study evaluates the effect of low temperatures combined to different spore storage times on the germination and initial gametophyte development of C. corcovadensis. Spores were divided into two groups: spores of the first group were sowed immediately in Meyer culture medium with nystatin, at pH 4.0, while spores of the second group were stored at 7, -20 and -196C during 60, 120, 180, and 365 days and then sowed in the same medium. Spore storage at 7 and -196C for 365 days not only provided higher germination percentages than those reported for recently-collected spores but also stimulated gametophytic development. The latter was demonstrated by the higher percentages of laminar gametophytes in these treatments. The possibility of storing spores provides material for in vitro experiments, which is of special interest for C. corcovadensis due to its ornamental potential and conservation status.(AU)

A samambaia arborescente Cyathea corcovadensis (Raddi) Domin está em perigo de extinção no Estado do Rio Grande do Sul, Brasil, ocorrendo somente no norte da região costeira. O armazenamento de esporos pode auxiliar programas de conservação, uma vez que oferece a possibilidade de manutenção da variabilidade genética e fornece material para culturas in vitro. O objetivo deste estudo foi avaliar o efeito de temperaturas baixas combinadas com diferentes tempos de armazenamento de esporos sobre a germinação e o desenvolvimento inicial de gametófitos de C. corcovadensis. Os esporos foram divididos em dois grupos: os do primeiro grupo foram semeados imediatamente em meio de cultura Meyer com nistatina, com pH 4,0 e os do segundo grupo foram armazenados a 7, -20 e -196C durante 60, 120, 180 e 365 dias e então semeados no meio supra mencionado. O armazenamento de esporos a 7 e -196C durante 365 dias propiciou porcentagens de germinação superiores àquelas observadas para esporos recém coletados e também estimulou o desenvolvimento gametofítico, o que foi demonstrado pelos elevados valores de porcentagem de gametófitos laminares observados nestes tratamentos. A possibilidade de armazenar esporos oferece material para experimentos in vitro, o que é de especial interesse para C. corcovadensis, considerando seu potencial ornamental e estado de conservação.(AU)

[Arborescent species of the genus Blechnum (Blechnaceae: Pteridophyta)]. / Especies arborescentes del género Blechnum (blechnaceae: Pteridophyta).

Although the genus Blechnum has been largely studied, problems of species characterization, nomenclature, classification and taxonomy remain unsolved. For these reasons, in this study we intend to deepen in the characterization of the arborescent group of Blechnum with a review of eight species: B. brasiliense, B. columbiense, B. cycadifolium, B. magellanicum, B. moritzianum, B. schomburgkii, B. tabulare and B. werckleanum.Type material and several collections from diverse herbaria were studied. For morphological studies we used both herbarium and fresh material collected by the authors in fieldwork between 2005 and 2010. Rhizomes, stipes, vascularization of costae, as well as several characters of the lamina, such as outline, division, venation, margin structure, epidermal patterns, stomata, presence and types ofhypodermis, and mesophyll ofpinnae were studied. Important aspects of the morphology of the sporophyte have been analyzed, and external characters have been correlated with internal diagnostic ones to use the most reliable external traits in keys and descriptions. Taxonomy has been updated, and corrections have been introduced to the species nomenclature and comments were included to summarize the current knowledge of the geographical distribution and ecology. The presence of B. tabulare in America is confirmed, and the taxonomy of the species is updated, since external and internal morphology indicated that other related, American and African species, such as B. spannagelii and B. madagascariense must be included under that name. Similar studies carried out on B. schomburgkii indicated that the name is a synonym of B. auratum subsp. auratum, while B. auratum subsp. columbiense is treated here as a well characterized species. Detailed study of numerous specimens allowed to characterize B. moritzianum, being the correct name to apply to the recently described B. yungense. A refreshed description of the species is presented together with a list of new, selected materials to illustrate current known geographical distribution. Expanded data on ecology, geographical distribution and affinities are given, and a key for species identification is presented.

Especies arborescentes del género Blechnum (Blechnaceae: Pteridophyta) / Arborescent species of the genus Blechnum (Blechnaceae: Pteridophyta)

Although the genus Blechnum has been largely studied, problems of species characterization, nomenclature, classification and taxonomy remain unsolved. For these reasons, in this study we intend to deepen in the characterization of the arborescent group of Blechnum with a review of eight species: B. brasiliense, B. columbiense, B. cycadifolium, B. magellanicum, B. moritzianum, B. schomburgkii, B. tabulare and B. werckleanum.Type material and several collections from diverse herbaria were studied. For morphological studies we used both herbarium and fresh material collected by the authors in fieldwork between 2005 and 2010. Rhizomes, stipes, vascularization of costae, as well as several characters of the lamina, such as outline, division, venation, margin structure, epidermal patterns, stomata, presence and types of hypodermis, and mesophyll of pinnae were studied. Important aspects of the morphology of the sporophyte have been analyzed, and external characters have been correlated with internal diagnostic ones to use the most reliable external traits in keys and descriptions. Taxonomy has been updated, and corrections have been introduced to the species nomenclature and comments were included to summarize the current knowledge of the geographical distribution and ecology. The presence of B. tabulare in America is confirmed, and the taxonomy of the species is updated, since external and internal morphology indicated that other related, American and African species, such as B. spannagelii and B. madagascariense must be included under that name. Similar studies carried out on B. schomburgkii indicated that the name is a synonym of B. auratum subsp. auratum, while B. auratum subsp. columbiense is treated here as a well characterized species. Detailed study of numerous specimens allowed to characterize B. moritzianum, being the correct name to apply to the recently described B. yungense. A refreshed description of the species is presented together with a list of new, selected materials to illustrate current known geographical distribution. Expanded data on ecology, geographical distribution and affinities are given, and a key for species identification is presented.

La bibliografía sobre el género Blechnum L. es muy abundante tanto en lo que respecta a las especies neotropicales como paleotropicales. Sin embargo, aún faltan estudios anatómicos completos, una caracterización actualizada de su morfología externa e interna, taxonomía, citología y afinidades. En este estudio se revisaron ocho especies de Blechnum de hábito arborescente: B. brasiliense, B. columbiense, B. cycadifolium, B. magellanicum, B. moritzianum, B. schomburgkii, B. tabulare y B. werckleanum. Se analizaron caracteres de rizomas, estípites y costas, división de la lámina, y venación, estructura de márgenes, modelos epidérmicos, presencia y tipos de hipodermis y mesofilo de las pinnas. Se estudió por primera vez la hipodermis, un área dermatoide adaxial presente en el mesofilo de todas las especies. Se correlacionaron caracteres externos e internos con el fin de utilizar los externos más confiables en claves y descripciones. Los estudios realizados confirmaron nuevamente la presencia de B. tabulare en América, que incluyen en su sinonimia a B. spannagelii de América tropical y B. madagascariense de África. También se concluyó que el nombre B. schomburgkii es un sinónimo de B. auratum subsp. auratum, en tanto que B. auratum subsp. columbiense se trata aquí con rango de especie. Se caracterizó a B. moritzianum, el nombre que corresponde aplicar a la recientemente descrita B. yungense. Se presentó una descripción actualizada de las especies, se resolvieron varios problemas nomenclaturales pendientes; se presentaron datos amplios sobre la ecología, distribución geográfica y afinidades entre las especies, así como una clave para determinarlas.

Influence of plant size on the ecophysiology of the epiphytic fern Asplenium auritum (Aspleniaceae) from Costa Rica.

PREMISE OF THE STUDY: A central goal of plant ecophysiological studies is to generate patterns of physiological behavior that are applicable to a species, which can be complicated when plant size is considered. Studies indicate that plant size can influence numerous ecophysiological parameters, especially in vascular epiphytes. The few studies that have included ferns in their analyses suggest that plant size is less important in ferns than angiosperms. This study investigates this apparent disparity by examining the relationship between plant size and an array of ecophysiological parameters in the epiphytic fern Asplenium auritum, especially the role of plant size in determining responses to water stress. METHODS: Plants were classified according to size and measured for a variety of functional traits, including maximum photosynthetic rate, stomatal conductance, water-use efficiency, stomatal density, chlorophyll content, chlorophyll a/b ratio, specific leaf area, whole plant drying rates, and desiccation tolerance. Results for all traits were compared across size classes to determine size-related differences. KEY RESULTS: Plant size significantly influenced most traits examined, most notably photosynthetic rate, stomatal density, stomatal conductance, whole plant drying rates, and recovery from desiccation. We report the first evidence of size-mediated shifts in desiccation tolerance in plants: small individuals tolerated water loss, whereas larger individuals avoided desiccation. CONCLUSIONS: Our findings indicate that size-mediated ecophysiological shifts are more important than previously thought in ferns, particularly in regard to water relations. Desiccation tolerance may allow young fern sporophytes to establish in stressful environments and is subsequently lost in older individuals.

Biometry of stomata in Blechnum species (Blechnaceae) with some taxonomic and ecological implications for the ferns.

Morphological stomatal traits, such as size, form and frequency, have been subject of much literature, including their relationships with environmental factors. However, little effort have focused on ferns, and very few in the genus Blechnum. Stomatal length, width and frequency (as stomatal index) of a number of specimens of fourteen Neotropical species of Blechnum were measured in adult pinnae. The aim of the work was to find biometrical relationships between stomatal traits and between stomatal traits and habit, habitat and ecosystem of the plants. Statistical analyses of data were conducted using Exploratory Data Analysis and Multivariate Statistical Methods. Stomatal length and width showed a very high correlation, suggesting an endogenous, genetic control, thus giving these traits a considerable diagnostic utility. With respect to the relationships between stomatal traits and environment, we found significant statistical relationships between altitude and stomatal index. We also addressed the interpretation of the ecological-selective significance of various assemblages of stomatal traits in a diverse conjunction of habits, habitats and ecosystems.

Biometry of stomata in Blechnum species (Blechnaceae) with some taxonomic and ecological implications for the ferns

Morphological stomatal traits, such as size, form and frequency, have been subject of much literature, including their relationships with environmental factors. However, little effort have focused on ferns, and very few in the genus Blechnum. Stomatal length, width and frequency (as stomatal index) of a number of specimens of fourteen Neotropical species of Blechnum were measured in adult pinnae. The aim of the work was to find biometrical relationships between stomatal traits and between stomatal traits and habit, habitat and ecosystem of the plants. Statistical analyses of data were conducted using Exploratory Data Analysis and Multivariate Statistical Methods. Stomatal length and width showed a very high correlation, suggesting an endogenous, genetic control, thus giving these traits a considerable diagnostic utility. With respect to the relationships between stomatal traits and environment, we found significant statistical relationships between altitude and stomatal index. We also addressed the interpretation of the ecological- selective significance of various assemblages of stomatal traits in a diverse conjunction of habits, habitats and ecosystems. Rev. Biol. Trop. 59 (1): 403-415. Epub 2011 March 01.

Los caracteres morfológicos estomáticos, tales como tamaño, forma y frecuencia, han sido objeto de abundante investigación, incluyendo su relación con los factores ambientales. Sin embargo, poco esfuerzo se ha realizado en esta materia en helechos y menos todavía en el género Blechnum. En este trabajo se midieron la longitud, anchura y frecuencia (como índice estomático) de estomas de pinnas adultas de un número de individuos en catorce especies de Blechnum neotropicales. El objetivo fue encontrar relaciones biométricas entre los caracteres estomáticos, y entre los caracteres estomáticos y el hábito, hábitat y ecosistema de las plantas. Se realizaron análisis estadísticos como Análisis Exploratorios de Datos y Métodos Estadísticos Multivariantes. La longitud y la anchura de los estomas mostraron una muy fuerte correlación, sugiriendo un control genético endógeno que otorga a estos caracteres un considerable valor diagnóstico. Con respecto a las relaciones entre los caracteres estomáticos y el ambiente, encontramos una relación estadísticamente significativa entre la altitud y el índice estomático. También se incluyen interpretaciones de la significación ecológico- selectiva de un conjunto de caracteres estomáticos en diferentes conjuntos de hábitos, hábitats y ecosistemas.

Intraspecific trait variation and covariation in a widespread tree species (Nothofagus pumilio) in southern Chile.

• The focus of the trait-based approach to study community ecology has mostly been on trait comparisons at the interspecific level. Here we quantified intraspecific variation and covariation of leaf mass per area (LMA) and wood density (WD) in monospecific forests of the widespread tree species Nothofagus pumilio to determine its magnitude and whether it is related to environmental conditions and ontogeny. We also discuss probable mechanisms controlling the trait variation found. • We collected leaf and stem woody tissues from 30-50 trees of different ages (ontogeny) from each of four populations at differing elevations (i.e. temperatures) and placed at each of three locations differing in soil moisture. • The total variation in LMA (coefficient of variation (CV) = 21.14%) was twice that of WD (CV = 10.52%). The total variation in traits was never less than 23% when compared with interspecific studies. Differences in elevation (temperature) for the most part explained variation in LMA, while differences in soil moisture and ontogeny explained the variation in WD. Traits covaried similarly in the altitudinal gradient only. • Functional traits of N. pumilio exhibited nonnegligible variation; LMA varied for the most part with temperature, while WD mostly varied with moisture and ontogeny. We demonstrate that environmental variation can cause important trait variation without species turnover.

Changes in sexual organ reciprocity and phenotypic floral integration during the tristyly-distyly transition in Oxalis alpina.

Although the 6 magnitude and pattern of correlation among floral traits (phenotypic integration) is usually conceived as an adaptation for successful pollination and reproduction, studies on the evolution of plant reproductive systems have generally focused on one or a few characters. If evolutionary transitions between reproductive systems involve morphological floral adjustments, changes in the magnitude and pattern of phenotypic integration of floral traits may be expected. In this study, we focused on the evolutionary dynamics of a complex adaptive trait, the extent of reciprocity (reciprocal placement) among sexual organs in a heterostylous species, and explored the associated changes in phenotypic floral integration during the transition from tristyly to distyly. The extent of reciprocity and both the magnitude and pattern of floral integration were characterized in 12 populations of Oxalis alpina representing the tristyly-distyly gradient. Although the extent of reciprocity increased along the tristyly-distyly transition, the flower size diminished. These adjustments did not affect the magnitude, but did affect the pattern, of floral integration. *Changes in the pattern of floral integration suggested that allometric, functional and pleiotropic relationships among floral traits were affected during this evolutionary transition.

Nuevos registros y observaciones de algunas especies de Elaphoglossum (Elaphoglossaceae) del Perú / New records and observations of some species of Elaphoglossum (Elaphoglossaceae) of Peru

Elaphoglossum cardioglossum Mickel y Elaphoglossum molle (Sodiro) C. Chr. son reportados para la flora peruana, de colecciones realizadas en Cusco y Pasco. Por primera vez es descrito material fértil de Elaphoglossum alipes Mickel, Elaphoglossum atropunctatum Mickel, Elaphoglossum cardioglossum Mickel y Elaphoglossum haynaldii (Sodiro) I. Losch. También se actualizan sus distribuciones.

Elaphoglossum cardioglossum Mickel and Elaphoglossum molle (Sodiro) C. Chr. are reported for the fi rst time for the Peruvian flora, from collections made in Cusco and Pasco. We also describe for the fi rst time fertile material of Elaphoglossum alipes Mickel, Elaphoglossum atropunctatumMickel, Elaphoglossum cardioglossum Mickel and Elaphoglossum haynaldii (Sodiro) I. Losh, and we update their current distributions.

The influence of life form on carbon and nitrogen relationships in tropical rainforest ferns.

Tropical ferns are characterized by a high diversity of plant life forms, yet there have been few large-scale studies on the functional ecology of these different forms. We examined epiphytic, hemiepiphytic, and terrestrial ferns, and asked whether there are differences in the mineral nutrition and water relations across different growth forms of a diverse assemblage of species. We measured specific leaf area, leaf nitrogen concentrations, and natural abundance of the stable isotopes delta(15)N and delta(13)C of 48 fern species from 36 genera across a wide range of habitats at La Selva Biological Station in Costa Rica. We found that epiphytes were significantly different in all measured variables from hemiepiphytic and terrestrial species, and that terrestrial and soil-rooted hemiepiphytes were indistinguishable in all variables excluding SLW. A multivariate analysis revealed that aspects of N nutrition were the most reliable at separating epiphytic species from other life forms. Our study demonstrates that the natural abundance of both C and N as well as N relations and leaf morphology are useful when segregating different plant life forms, and that the N cycle of epiphytic and terrestrial habitats function independently from each other.

Una nueva especie de helecho del género Danaea (Marattiales: Marattiaceae) endémica de Costa Rica / A new species of fern, genus Danaea (Marattiales: Marattiaceae) endemic to Costa Rica

Danaea tuomistoana A. Rojas (Marattiaceae) is described and illustrated as a new species endemic to Costa Rica; it differs from D. crispa Endres et Richb. by a truncate blade, and by the pinnae, which are 2/5-1/2 incised, cuneated at the base and with a flat margin. The range of D. erecta H. Tuomisto & R.C. Moran in Costa Rica is expanded with a new locality in Puntarenas (8 degrees 57'15[quot ] N-82 degrees 50'1[quot ] W, 1,500-1,580 masl).

Se describe e ilustra una especie nueva endémica de helecho de Costa Rica, Danaea tuomistoana A. Rojas (Marattiaceae), caracterizada porque las pinnas son incisas 2/5-1/2 entre la costa y el margen, y son cuneadas en la base. Se amplia el ámbito geográfico de D. erecta H. Tuomisto & R.C. Moran a Puntarenas, Costa Rica.

[A new species of fern, genus Danaea (Marattiales: Marattiaceae) endemic to Costa Rica]. / Una nueva especie de helecho del género Danaea (Marattiales: Marattiaceae) endémica de Costa Rica.

Danaea tuomistoana A. Rojas (Marattiaceae) is described and illustrated as a new species endemic to Costa Rica; it differs from D. crispa Endres et Richb. by a truncate blade, and by the pinnae, which are 2/5-1/2 incised, cuneated at the base and with a flat margin. The range of D. erecta H. Tuomisto & R.C. Moran in Costa Rica is expanded with a new locality in Puntarenas (8 degrees 57'15" N-82 degrees 50'1" W, 1,500-1,580 masl).

New taxa, new records and redefined concepts in the Elaphoglossum sect. Elaphoglossum subsec. Pachyglossa (Lomariopsidaceae) from Mexico and Central America.

Twelve new species are described in the taxonomically difficult Elaphoglossum (sect. Elaphoglossum): Elaphoglossum angustifrons A. Rojas, E. delgadilloanum A. Rojas, E. ellipticifolium A. Rojas, E. incognitum A. Rojas, E. mesoamericanum A. Rojas, E. nicaraguese A. Rojas, E. polypodium A. Rojas, E. tejeroanum A. Rojas, E. reptans A. Rojas, E. terrestre A. Rojas, E. variabile A. Rojas and E. zavale A. Rojas. Also, E. latifolium (Sw.) J. Sm., E. sartorii (Liebm.) Mickel and E. viride (E. Fourn.) C. Chr. are amended, E. andicola (Fée) T. Moore and E. sporadolepis (Kunze ex Kuhn) T. Moore are reported.

Notes on Elaphoglossum (Lomariopsidaceae) section Polytrichia subsection Hybrida in Mexico and Central America.

In Elaphoglossum sect. Polytrichia subsect. Hybrida six new species are described: E. angustioblongum A. Rojas, E. baquianorum A. Rojas, E. cotoi A. Rojas, E. jinoteganum A. Rojas, E. neeanum A. Rojas and E. silencioanum A. Rojas. New combination is made for Elaphoglossum mexicanum (E. Fourn.) A. Rojas. Two species are reported: E. barbatum (H. Karst.) Hieron. and E. scolopendrifolium (Raddi) J. Sm. Two species are redefined: E. erinaceum (Fée) T. Moore and E. tambillense (Hook.) T. Moore. E. pallidum (Baker ex Jenman) C. Chr. is eliminated for Mexico and Central America. Of the new species only E. neeanum is present outside of the region. A key is given to those species in Mexico and Central America.

New species, new combinations and new distributions in neotropical species of Elaphoglossum (Lomariopsidaceae).

More new species, new combinations and new distributions are made in the large and difficult genus Elaphoglossum in Neotropic. Ten new species are described, this are: Elaphoglossum adrianae A. Rojas, E. betancuri A. Rojas, E. caridadae A. Rojas, E. glabromarginatum A. Rojas, E. kessleri A. Rojas, E. macdougalii A. Rojas, E. paramicola A. Rojas, E. paxense A. Rojas, E. pseudoherminieri A. Rojas and E. solomonii A. Rojas. Three species are combined, this are: Elaphoglossum beitelii (Mickel) A. Rojas, E. curvans (Kunze) A. Rojas and E. eutecnum (Mickel) A. Rojas. Finally, 86 species are reported from other countries and some species names are synonymized.

[Six new species and two new records of (Pteridophyta) from Costa Rica ]. / Seis especies nuevas y dos nuevos registros de helechos (Pteridophyta) para Costa Rica.

Six new species of Pteridophyta from Costa Rica are described: Dryopteris flaccisquama A. Rojas, Hypolepis lellingeri A. Rojas, H. moraniana A. Rojas, Melpomene alan-smithii A. Rojas, Selaginella osaënsis A. Rojas and Terpsichore esquiveliana A. Rojas. Blechnum stoloniferum (Mett. ex E. Fourn.) C. Chr. and Trichomanes micayense Hieron. are new records for the country. Only D. flaccisquama and M. alan-smithii are present outside Costa Rica.

[New species, newly used names and new ranges of tree ferns (Filicales: Cyatheaceae) in the Neotropics]. / Nuevas especies, nombres nuevamente utilizados y nuevas distribuciones en los helechos arborescentes (Filicales: Cyatheaceae) para el Neotrópico.

Two new species are described for the Neotropics, four species are newly accepted, and four species and one variety are reported. The new species are: Cyathea grayumii A. Rojas and C. panamensis A. Rojas. Cyathea alfonsiana L. D. Gómez, C. holdridgeana Nisman & L. D. Gómez, C. onusta H. Christ and C. squarrosa (Rosenst.) Domin are recognized; Cnemidaria coclena Stolze, Cyathea andina (H. Karst.) Domin, C. caracasana var. meridensis (H. Karst.) R. M. Tryon, C. macrosora (Baker) Domin and C. pseudonanna (L. D. Gómez) Lellinger are reported from Costa Rica and Panama.

[Eight new species and new ranges of Tectariaceae (Felicales) in the Neotropics]. / Ocho nuevas especies y nuevos ámbitos geográficos de helechos de la familia Tectariaceae (Filicales) en el Neotrópico.

The following species of Tectariaceae (Filicales) from Nicaragua, Costa Rica, Panamá, Colombia and Perú are described as new: Ctenitis sotoana A. Rojas, Megalastrum ctenitoides A. Rojas, Tectaria darienensis A. Rojas, T. faberiana A. Rojas, T. longipinnata A. Rojas, T. murilloana A. Rojas, T. pascoensis A. Rojas and T. subdimorpha A. Rojas. The new combination of M. karstenianum (Klotzsch) A. Rojas is proposed. The distributions of C. bullata A. R. Sm., C. microchlaena (Fée) Stolze, C. submarginalis (Langsd. et Fisch.) Ching, T. andina (Baker) C. Chr. and T. sodiroi (Baker) Maxon are enlarged.