Sorophores of<Emphasis Type="Italic">Lygodium</Emphasis> Sw. (Schizaeaceae) from the Late Cretaceous of New Jersey

We report here on a series of specimens of charcoalified sorophores with characteristics of the extant fern genusLygodium (Schizaeaceae) collected from sediments of the Raritan Formation (Late Cretaceous). Each elongate lobed fertile pinnule (sorophore) is flattened and bears alternately arranged sporangia on one surface. Each sporangium is covered by an indusium continuous with the margin of the lamina. Sporangia are oblong in shape, short stalked, and have an apical annulus formed by a single ring of radiating cells that dehisces longitudinally. The sporangial cap or distal face is formed by only one cell. All of these features are characteristic of the extant genusLygodium. Small numbers of trilete, psilate spores are found in the sporangia. Megafossils assignable toLygodium are known from the Upper Cretaceous of North America and Germany with worldwide distribution during the Tertiary. The newLygodium fossils are compared with others previously referred to the genus.     

THE SPORANGIUM OF HORNEOPHYTON LIGNIERI (RHYNIOPHYTINA)

Numerous sporangia of Horneophyton lignieri from the Rhynie Chert locality in Scotland have been studied. The sporangia are branched, with two to four columellate lobes of varying length, and a continuous sporogenous zone or cavity occurs among the lobes. Unbranched sporangia, generally thought to be the typical form for the plant have not been found, and their presence is not established. Although not definitely proven, evidence suggests that the sporangia opened by means of a small apical pore or stoma. An area of thick-walled cells at the apex of each sporangial lobe probably played some role in this opening. Radial, trilete, azonate spores ranging from 39–49 μm in diam, with curvaturae perfectae are produced most commonly in tetrahedral tetrads and occasionally in isobilateral tetrads. Matters of spore preservation and possible ornamentation are discussed. The branched sporangia of this genus are unique among bryophytes and vascular plants and provide some evidence that certain synangia may have arisen from a single sporangium rather than from multiple sporangia borne singly at the tips of ultimate branches.     

SPORE MORPHOLOGY IN THE CYATHEACEAE. I. THE PERINE AND SPORANGIAL CAPACITY: GENERAL CONSIDERATIONS

The literature on cyatheaceous spore morphology relative to the presence of a perine layer is reviewed, and evidence based on a sodium-hydroxide assay is presented indicating that the outer scultpine layer in certain cyatheaceous spores is perine. Perine so defined characterizes Metaxya, paleotropical and certain neotropical species of Sphaeropteris, nearly all species of Alsophila, all species of Nephelea, and certain species of Trichipteris and Cyathea. It is lacking in Lophosoria, many species of Trichipteris and Cyathea, and all species of Cnemidaria. Two major patterns of spore number per sporangium in the family are reported. Lophosoria, Sphaeropteris, Trichipteris, Cyathea, Cnemidaria, and probably Metaxya are characterized by 64-spored sporangia, whereas most species of Alsophila and all species of Nephelea are characterized by 16-spored sporangia. The congruence of this generic distribution of sporangial-capacity types with Tryon's phyletic arrangement of cyatheaceous genera supports the naturalness of his system. The intrasporangial germination of spores retained in dehisced and dispersed sporangia supports the suggestion that decreased spore number per sporangium in Alsophila and Nephelea may relate to the role of the sporangia as dispersal units. The decreased number of spores per sporangium is associated with a trend toward increase in the number of sporangia per sores, with the highest known count approaching 1000 sporangia per sorus. The Alsophila-Nephelea evolutionary line has probably not been ancestral in the phylogeny of the more advanced groups of ferns.     

SULLITHECA DACTYLIFERA GEN. ET SP. N.: A NEW MEDULLOSAN POLLEN ORGAN AND ITS EVOLUTIONARY SIGNIFICANCE

The pteridosperm (Medullosaceae) pollen organ Sullitheca dactylifera gen. et sp. n. is described from middle Pennsylvanian coal balls. The proximally fused units of the obpyriform compound synangium separate and extend distally as finger-like projections. Each projection contains 4–6 vertically oriented cylindrical sporangia arranged in pairs along the radius of the unit; each unit extends from the outer cover wall toward the center. The distal portion of the compound synangium is hollow as a result of the lateral separation of the centripetally and distally directed synangial units. About 40 tubular sporangia are present in all and dehiscence occurs along a lateral slit in each sporangium. Vascular strands are disposed around the periphery of the organ in addition to a single strand paralleling each sporangium. Two- or three-cell trichomes and stomata are present on the organ surface. Pollen of the Monoletes type is present. A paired row of sporangia in Sullitheca composing a synangial unit is considered the homologue of a paired row of sporangia in the more compact and highly evolved genus, Dolerotheca.     

Neoleptonema yongpilii E.-Y. Lee & I.K. Lee,gen. et sp. nov. (Phaeophyceae), based on morphological characters and RuBisCO spacer sequences

An elachistacean epiphyte, Neoleptonema yongpilii E.-Y. Lee & I.K. Lee, gen. et sp. nov., is reported from Korean coasts. The plants are distinguished by having unbranched assimilatory filaments with intercalary plurilocular sporangia as well as lateral plurilocular sporangia from the cortex. The new genus differs from the genera Leptonematella P. Silva and Halothrix Reinke by having pod-shaped plurilocular sporangia on the medulla, and from Elachista Duby and Proselachista Y.P. Lee & Garbary by having intercalary plurilocular sporangia and a poorly developed medulla. The phylogenetic relationships of Neoleptonema yongpilii were inferred from the spacer sequences between the genes coding for the large and small subunits of the RuBisCO gene. The new genus is a member of a poorly resolved clade consisting of several genera within the Elachistaceae, within which it is more closely related to Halothrix and Elachista nipponica than to Leptonematella.     

ACAULANGIUM GEN. N., A FERTILE MARATTIALEAN FROM THE UPPER PENNSYLVANIAN OF ILLINOIS

Material described by Graham as Cyathotrachus bulbaceus is believed to represent a new genus that is a common constituent of Upper Pennsylvanian coal balls. The sessile synangia of Acaulangium gen. n. are borne in a row on either side of the pinnule midrib and are composed of four to six short, tapering, laterally appressed sporangia. The sporangia have extended tips which curve over the inside of the synangium distally and delimit a small open area inside the synangium. The outer facing walls of the sporangia are two to three cells thick throughout while the inner facing walls are uniseriate. During dehiscence the sporangia separate laterally and spore release results from the rupture of a row of elongate cells along the inner sporangium midline. Among species of Scolecopteris the new genus resembles S. illinoensis and S. minor var. parvifolia but differs in its sessile synangial attachment. The additional parenchyma present between sporangial cavities in the synangia of Acaulangium, and the tendency toward bilateral symmetry suggests an early stage in the evolution of a bivalve synangium such as is present in Marattia.     

BISCALITHECA KANSANA SP. N. (COENOPTERIDALES,ZYGOPTERIDACEAE), A COMPRESSION FROM THE LAWRENCE SHALE (UPPER PENNSYLVANIAN), KANSAS,U. S. A.

The organ bearing the sporangia of Biscalitheca kansana is a laminate rachis with subopposite fertile appendages, a structure previously unknown for Biscalitheca. The sporangia differ from those of the petrified Biscalitheca musata only by lacking nests of sclerotic cells. They contain spores which are called Convolutispora when found dispersed. Speculations are made that Monoscalitheca fasciculata will ultimately be regarded as a synonym of Biscalitheca musala and that some fossils called Aphlebia and Brittsia problematica may be the sterile foliage of zygopterids.     

BOTRYOPTERIS PINNULES WITH ABAXIAL SPORANGIA

Small sporangia borne abaxially on pinnules attached to Botryopteris foliar members are described from coal ball petrifactions of Early Pennsylvanian age. This is the first report of laminar sporangia in this genus. Sporangia are stalked and borne singly near lateral veins on Sphenopteris-like pinnules. Individual sporangia are of the leptosporangiate type, with a lateral annulus and a dehiscence zone of thin-walled cells immediately adjacent to the annulus. Spores are small, trilete, triangular in outline, typically have blunt spines covering the exine, and correspond to the dispersed spore genera Acanthotriletes, Leiotriletes, or Lophotriletes. These sporangia and their spores are unlike previously described globose Botryopteris fructifications from the Middle and Upper Pennsylvanian, but are similar to sporangia produced by modern members of the Osmundaceae.     

Morphology and life history of Scytosiphon canaliculatus comb. nov. (Scytosiphonales,Phaeophyceae) from Japan*

Morphology of field material and life history in culture were studied in Scytosiphon canaliculatus (Setchell et Gardner) comb. nov. from northern Japan. Erect gametophytes of S. canaliculatus are cylindrical, tubular, up to 7 mm wide and 40 cm long, and without regular constrictions. S. canaliculatus has pronounced anisogamy and ascocysts accompanied with plurilocular gametangia. The life history of S. canaliculatus showed an alternation between erect gametophytes and crustose prostrate sporophytes bearing unilocular sporangia. Since field sporophytes of S. canaliculatus were found to be identical with Hapterophycus canaliculatus Setchell et Gardner (Ralfsiaceae, Phaeophyceae), it is proposed to transfer H. canaliculatus to the genus Scytosiphon. In the field, gametophytes with plurilocular gametangia appeared in spring and disappeared in summer. Sporophytes with unilocular sporangia were collected in late autumn and winter. Unilocular sporangia were produced at 15°C in short-day culture conditions and unispores developed into erect gametophytes at 5–15°C. It is suggested that the seasonal Occurrence of gametophytes in the field is due to the seasonal formation of unilocular sporangia, which is regulated by temperature and photoperiod.     

Camellia curryana and C. longii spp. nov. (Theaceae) from Vietnam

Two species of Camellia L. (Theaceae) are described here as new: C. curryana and C. longii. The new species were discovered in the southern Annamite Mountains of Vietnam and are endemic to tropical rainforest remnants that occur at altitudes between 1500 and 1700 m a.s.l. Camellia curryana possesses almost sessile, unevenly circular, two whorled, white‐cream and proximally yellowish flowers, proximally joined outer stamens, three styles and mature fruit that dehisces into three parts, or longitudinally into two halves. Camellia longii has campanulate or almost campanulate, intensely dark orange to red flowers with uneven whitish margins, 5–6 petals and 3–2 petaloids, filaments that are united with the petals and one another, a diamond shaped ovary and compound 5–6 styles that are connate at the base.     

IDANOTHEKION GEN. N., A SYNANGIATE POLLEN ORGAN WITH SACCATE POLLEN FROM THE MIDDLE PENNSYLVANIAN OF ILLINOIS

Idanothekion glandulosum gen. et sp. n. is a synangiate pollen organ represented by approximately 30 specimens contained in coal balls from the middle Pennsylvanian of Illinois. Each synangium is composed of seven to nine elongate sporangia that are fused laterally for approximately four-fifths of their length, and are radially arranged about, and fused to, a short central column; the central column is restricted to the proximal one-third of the synangium. Distal to the column the sporangia surround a hollow central area. Dehiscence occurred by means of a longitudinal slit along the mid-line of the inner face of each sporangium. The outer walls of the sporangia have a complex histology involving an external epidermis, a middle presumably glandular layer containing scattered enlarged cells, and an inner layer made up of thin-walled parenchyma. Vascular tissue is present in the central column and outer walls of the sporangia. Each sporangium has a prominent, attenuate, multicellular tip. Large numbers of saccate pollen grains similar to those found in numerous fossil and extant coniferophytes as well as some Mesozoic pter-idosperms were produced in each sporangium. Idanolhekion resembles some synangia assignable to Paleozoic members of the Marattiales; however, the new genus compares most closely with pollen organs believed to have been produced by members of the Pteridospermales. It seems most likely that Idanothekion represents the pollen organ of some member of the Lyginopteridaceae that produced pollen of a type which up to now has not been known from Paleozoic seed ferns.     

A new species of vascular plants from the Xujiachong Formation (Lower Devonian) of Yunnan Province, China

A new species of early land vascular plants, Huia gracilis, is described from the Lower Devonian (Pragian-early Emsian) Xujiachong Formation, Qujing district, Yunnan Province, southwestern China. The plant possesses K- or H-type branching in the rhizome and a dichotomously branched erect system. An axillary tubercle is sometimes present at the branching position. Fertile axes terminate in loose spikes comprising terminal and lateral sporangia arranged helically. Sporangia with long stalks are ovate or elongate-ovate and reflex adaxially. The sporangium dehisces longitudinally in the radial plane of the fertile axis into two parts. The xylem is probably centrarch. Tracheids of G-type are characterised by annular secondary thickenings, between which is a sheet with irregular simple perforations. There may be some 'pores' in the enclosing wall of the perforation. H. gracilis differs from the type species H. recurvata mainly in the branching pattern, more slender morphology of the stem, sporangium and stalk. It is suggested that Huia may have originated from the primitive Cooksonia-like plants.     

<Emphasis Type="Italic">Pythium montanum</Emphasis> sp. nov., a new species from a spruce stand in the Bavarian Alps

During a study on soilborne Pythiaceae in the Bavarian Alps, 3 isolates of a papillate Pythium species with aplerotic oospores were found in rhizosphere soil from a declining spruce stand. The isolates showed a unique combination of sporangial and gametangial characters, and could not be assigned to any known species of the genus. They were characterised by a relatively low growth rate, internally proliferating sporangia and extremely variable, monoclinous, sessile or shortly stalked antheridia. Direct germination of sporangia frequently occurred in water culture. Internal transcribed spacer (ITS) regions of the ribosomal DNA showed very low levels of similarity to those of all known species of Pythium. This paper describes the isolates as Pythium montanum sp. nov., gives details on its morphology and provides preliminary information on its pathogenicity. Additional molecular evidence for its status as a new species is also provided. Phylogenetic analyses with a set of Pythium spp. showed that P. montanum groups in a clade together with P. vexans, a species whose taxonomical status still remains unclear, and that is considered a ‘border species’ between Pythium and Phytophthora.     

DIMENSIONAL CORRELATIONS IN DEVELOPING SELAGINELLA SPORANGIA

With length of sporangia as a developmental index, the growth relationships of sporangia during differentiation were studied in strobili of Selaginella bigelovii. The strobili usually contain two rows of megasporangia and two rows of microsporangia with a mega- opposite a microsporangium at each node. Prior to the sporocyte stage a sporangium in a megasporangiate row is larger and elongates more rapidly than a sporangium opposite it at the same node in a microsporangiate row. The number of sporogenous cells is similar in sporangia of the same length from both rows until cell multiplication ceases in sporangia of the megasporangiate row, while it continues in sporangia of the same size in the microsporangiate row. The observed growth differences between sporangia of the micro- and megasporangiate rows are interpreted as events in the differentiation of two sporangial types.     

A spectrophotometric approach for determining sporangium and zoospore viability of Plasmopara viticola

A spectrophotometric method for determining the viability of sporangia and zoospores of the oomycete Plasmopara viticola (causal agent of grapevine downy mildew) is described and evaluated to overcome the limitations of currently available methods for assessing propagule viability. Sporangia produced on leaf discs in the laboratory were harvested at different days after the initiation of sporulation (DAS) to yield differences in sporangium viability. Sporangia were suspended in sterile water, the suspensions were placed in a cuvette, and sporangium germination was monitored in a spectrophotometer (λ = 600 nm) at 2- to 3-min intervals for 5 hr. Absorbance started to increase after sporangia were suspended in water for ~30–60 min followed by major peak(s) for younger sporangia (1–3 DAS), whereas low to no increase in absorbance was observed for senescent sporangia (>7 DAS). Microscopic observation confirmed that the increase in absorbance corresponded to the release and active swimming of zoospores, whereas absorbance decreased when zoospores encysted and settled. A positive correlation (r = .839, p = .0365) was observed when the time to the initial increase in absorbance was plotted against the age of sporangia. The time to the absorbance peak (marking the time of maximum zoospore movement) was shortest for immature sporangia (0 DAS), longest for young sporangia (2 DAS) and decreased for mature and senescent sporangia. A similar pattern was observed for the standardized area under the absorbance curve (indicating the overall quantity of zoospores released), for which values were lowest for immature and senescent sporangia, highest for young sporangia and intermediate for mature sporangia. Consistent patterns obtained across two independent experiments suggest that the method is reproducible and may be further developed for other zoospore-releasing pathogens.     

CALAMOPHYTON PRIMAEVUM: THE COMPLEX MORPHOLOGY OF ITS FERTILE APPENDAGE

Although holotype material of Calamophyton primaevum Kläusel and Weyland has not been available since the Second World War, two fertile specimens of that species are in the personal collection of Professor H. Weyland. The purpose of this paper is to disclose the complexity of the organization of the fertile appendage. Formerly it was believed to consist of one segment that had dichotomized once and that had borne two pendulous sporangia, one at each branch. Numerous degagements of sporangia have revealed a complex morphology, consisting of one main segment divided into two secondary segments. Each supports three recurved side stalks which in turn bear 2 sporangia, a total of 12 sporangia. Each secondary segment terminates in two elongated projections. The morphology of the fertile organs is compared with that of two other species, C. bicephalum Leclercq and Andrews and C. forbesii Schopf. Great similarity of organization exists between C. primaevum and C. bicephalum. The possible synonymy of these two species is discussed.     

SPORANGIA OF RALFSIA FUNGIFORMIS (GUNN.) SETCHELL ET GARDNER12

Unilocular and plurilocular sporangia of Ralfsia fungiformis are, for the first time, recorded for North America. The validity of certain types of structures, considered by some authors as plurilocular sporangia, is discussed.     

DEVELOPMENT OF ENDOPHYTIC FRANKIA SPORANGIA IN FIELD- AND LABORATORY-GROWN NODULES OF COMPTONIA PEREGRINA AND MYRICA GALE

Field-collected nodules of Comptonia peregrina (L.) Coult. and Myrica gale L. (Myricaceae), infected by the nitrogen-fixing actinomycete Frankia sp., were of two types: those that lacked sporangia entirely, designated spore(-), and those that showed extensive sporangial development, designated spore(+). In spore(+) nodules of C. peregrina, sporangia began to develop after the differentiation of endophytic vesicles and the concomitant onset of nitrogenase activity. At the onset of sporangial differentiation, infected host cells appeared healthy. However, endophytic vesicles and host cell cytoplasm and nuclei began to senesce rapidly as sporangia developed. Staining of sectioned material with the fluorescent stain Calcofluor White suggested that vesicles, hyphae and young sporangia were enclosed within a host-derived encapsulation layer, but mature sporangia were no longer encapsulated. In both C. peregrina and M. gale, vesicles were more short-lived in spore(+) than in spore(-) nodules. Field-collected spore(+) M. gale nodules exhibited a pronounced seasonality of sporangial formation. Sporangia began to differentiate in June, after the formation of vesicles and became more prominent in late summer. Inter- and intraspecific cross-inoculations suggest that the ability to form sporangia in the symbiotic state is controlled by endophytic strain type rather than host genotype or host/endophyte combination. The host may, however, influence the number and seasonal appearance of sporangia formed.