Effects of chelated iron on oogenesis and vegetative growth of kelp gametophytes (Phaeophyceae)

The supply of iron has been reported to affect gametogenesis in the gametophytes of some species of kelps (order Laminariales). Spores of the kelps Alaria marginata Postels & Ruprecht, Dictyoneurum californicum Ruprecht, Egregia menziesii (Turner) Areschoug, Laminaria setchellii Silva, and Macrocystis pyrifera (L.) C. Agardh were cultured in enriched seawater with and without added chelated iron (Fe‐ethylenediaminetetraacetate) to determine the effects of iron on oogenesis. All species showed a decrease in oogenesis without added Fe‐ethylenediaminetetraacetic acid (EDTA). Gametophytes of E. menziesii showed predominant gametogenesis with or without supplied iron, resulting in all cells being converted to gametes so that vegetative growth did not continue. Vegetative gametophytes were obtained in the other species used. Gametophytes of M. pyrifera did not show any oogenesis without added Fe‐EDTA, while those of L. setchellii, A. marginata and D. californicum were intermediate in their response, showing some gametogenesis without added Fe‐EDTA. When Fe‐EDTA supply was delayed by 6, 13 and 20 days with spores of M. pyrifera, the gametophytes produced fewer eggs, with a greater decrease as the delay grew longer. A range of Fe‐EDTA concentrations was investigated using isolated female gametophytes of two strains of M. pyrifera and one of Macrocystis integrifolia Bory. None of these three strains produced gametes without the addition of Fe‐EDTA. Gametophytes of M. integrifolia required the least amount of added Fe‐EDTA to achieve gametogenesis while gametophytes of M. pyrifera required higher amounts, with the two strains showing somewhat different responses. Iron nutrition appears to be an essential factor for gametogenesis in several species of kelps.     

100 Effects of chelated iron on gametogenesis in kelp gametophytes

Gametogenesis in several species of kelp gametophytes was investigated in relation to the supply of chelated iron (Fe-EDTA) in the culture medium. Spores of Laminaria setchellii, Egregia menziesii, Alaria marginata, Dictyoneurum californicum , and Macrocystis pyrifera were grown in Provasoli's Enriched Seawater with and without added Fe-EDTA. All species showed a decrease in gametogenesis without added Fe-EDTA. Gametophytes of E. menziesii showed predominant gametogenesis even without supplied iron, resulting in all cells being converted to gametes so that vegetative growth did not continue. Vegetative gametophytes were obtained in the other species. M. pyrifera did not show any gametogenesis without added Fe-EDTA. L. setchellii, A. marginata and D. californicum were intermediate in their response. In addition, Fe-EDTA supply was delayed by 1, 2 and 3 weeks with spores of M. pyrifera , resulting in an intermediate degree of gametogenesis which decreased as the delay grew longer. Fe-EDTA supply was also investigated using isolated female gametophytes of two strains of M. pyrifera and one of Macrocystis integrifolia . None of the three strains produced gametes without the addition of Fe-EDTA. Gametophytes of M. integrifolia required the least amount of added Fe-EDTA to achieve gametogenesis while gametophytes of M. pyrifera required higher amounts, with the two strains showing different responses. Thus, by varying the supply of chelated iron, it is possible to control vegetative growth or gametogenesis in most of these kelp species. In addition, different species and individuals show varied responses to added chelated iron.     

ROLE OF NUTRIENT FLUCTUATIONS AND DELAYED DEVELOPMENT IN GAMETOPHYTE REPRODUCTION BY MACROCYSTIS PYRIFERA (PHAEOPHYCEAE) IN SOUTHERN CALIFORNIA1

Organisms occurring in environments subject to severe disturbance and/or periods of poor environmental quality that result in severe adult mortality can survive these periods by relying on alternate life stages that delay their development in a resistant state until conditions improve. In the northeast Pacific, the forest‐forming giant kelp Macrocystis pyrifera (L.) C. Agardh periodically experiences widespread adult mortality during extended periods of extremely low nutrients and high temperatures, such as those associated with El Niño. Recovery following these periods is hypothesized to occur from microscopic life stages that delay their development until the return of favorable conditions. In the laboratory, we experimentally examined the environmental conditions responsible for regulating delayed development of the microscopic stages of M. pyrifera from Southern California, USA. Nutrients controlled the delay and resumption of gametophyte growth and reproduction, perhaps linked to the large fluctuations in nutrients occurring seasonally and interannually in this region. Although growth of gametophytes proceeded in the virtual absence of nitrate, both nitrate and other trace nutrients were necessary for gametogenesis. Upon exposure to elevated nutrients, delayed gametophytes produced sporophytes more quickly (5–20 d) and at smaller sizes (10–200 μm) than gametophytes that had never been delayed (18–80 d, 80–400 μm, respectively), reducing negative density‐dependent effects. This finding demonstrates that delayed gametophytes of M. pyrifera rapidly utilize increased resources to consistently produce sporophytes. Further work is needed to assess their potential role in population recovery following periods of poor environmental quality.     

Responses of vegetative gametophytes of Undaria pinnatifida to high irradiance in the process of gametogenesis

The population of Undaria pinnatifida in its ecologic niche sustains itself in high temperature summer in the form of vegetative gametophytes, the haploid stage in its heteromorphic life cycle. Gametogenesis initiates when seawater temperature drops below the threshold levels in autumn in the northern hemisphere. Given that the temperature may fall into the appropriate range for gametogenesis, the level of irradiance determines the final destiny of a gametophytic cell, either undergoing vegetative cell division or initiating gametogenesis. In elucidating how vegetatively propagated gametophytes cope with changes of irradiance in gametogenesis, we carried out a series of culture experiments and found that a direct exposure to irradiance as high as 270 μmol photons m^?2 s^?1 was lethal to dim‐light (7–10 μmol photons m^?2 s^?1) adapted male and female gametophytes. This lethal effect was linearly corelated with the exposure time. However, dim‐light adapted vegetative gametophytes were shown to be able tolerate as high as 420 μmol photons m^?2 s^?1 if the irradiance was steadily increased from dim light levels (7–10 μmol photons m^?2 s^?1) to 90, 180 and finally 420 μmol photons m^?2 s^?1, respectively, at a minimum of 1–3 h intervals. Percentage of female gametophytic cells that turned into oogonia and were eventually fertilized was significantly higher if cultured at higher but not lethal irradiances. Findings of this investigation help to understand the dynamic changes of population size of sporophytic plants under different light climates at different site‐specific ecologic niches. It may help to establish specific technical details of manipulation of light during mass production of seedlings by use of vegetatively propagated gametophytes.     

LIFE HISTORY AND GROWTH OF LAURENCIA BRACHYCLADOS (RHODOPHYTA,CERAMIALES)1

Laurencia brachyclados Pilger from Hawai'i completed a “Polysiphonia-type” tri-phasic life history in 21 weeks in laboratory culture. Tetraspores developed into gametophytes in a nearly 1:1:1 ratio of females: males: non-reproductive. Carpospores were released as early as 21 days after mixing virgin female and male gametophytes. Cultured thalli showed a “guerilla type” growth form. Other Hawaiian Laurencia species in culture had longer maturation times or remained non-reproductive. Variation in life history schedules may influence Laurencia species coexistence and algal community structure.     

Life history and growth in culture of the endemic New Zealand kelp <Emphasis Type="Italic">Lessonia variegata</Emphasis> J. Agardh in response to differing regimes of temperature,photoperiod and light

Lessonia variegata J. Agardh (Laminariales, Phaeophyta) is endemic to New Zealand, where it occurs in subtidal kelp forests on wave exposed coasts in the North, South and Stewart Islands. This is the first account of the growth in culture and life history of L. variegata. Microscopic gametophytes alternate with macroscopic sporophytes, characteristic of members of the order Laminariales. The life history was completed in culture within 14 days under growth conditions of 12 °C, 12:12 (L:D) and 15 °C, 15:9 (L:D). Maximum growth of sporophytes occurred at 15 °C, 15:9, and slowest growth at 10 °C, 9:15. Under low light conditions (8–9 μmol photon m⁻² s⁻¹) filamentous growth of gametophytes predominated, and both the proportion of gametogenesis and the growth of sporophytes after 30 days was much reduced from equivalent cultures grown under conditions of higher light (16–17.5 μmol photon m⁻² s⁻¹). Interest in this species relates both to its potential for commercial utilisation as well as in the role it plays in coastal rocky reef ecosystems.     

Cultivation of the red alga Chondracanthus chamissoi: sexual reproduction and seedling production in culture under controlled conditions

The red alga Chondracanthus chamissoi (Gigartinales) is endemic to the southern-central region of South America. In the Pacific Ocean, it is distributed from north-central Peru to Chiloe Island. This species is of economic importance because it is edible and used for carrageenan production. The tetrasporophyte phase was grown in the laboratory, obtaining male and female gametophytes that were incubated under different photoperiod, pH, salinity and temperature conditions. These gametophytes developed and generated reproductive structures that led to in vitro maturation. Subsequently, fertilisation occurred and formation of cystocarps was observed. Finally, carpospores were released and the formation of sporophytes completed the life history of this species under laboratory conditions. Reproductive phase growth rates were recorded for each of the different culture conditions used. Sporophytes reached the highest daily growth rate (22%), while gametophyte’s daily growth rate was slower (9%). This research confirms, in vitro, the assumption that C. chamissoi has a sexual triphasic life history Polysiphonia type with isomorphic gametophytes and tetrasporophytes. The development of the complete life history took 20 months in the laboratory.     

PHOTOSYNTHETIC RESPONSES TO LIGHT AND TEMPERATURE OF THE HETEROMORPHIC MARINE ALGA MASTOCARPUS PAPILLATUS (RHODOPHYTA)1

Physiological differentiation of the heteromorphic life-history phases of the red alga Mastocarpus papillatus (C. Agardh) Kützing was assessed. Photosynthetic responses to light and temperature of the erect, foliose gametophyte were compared to those of the crustose tetrasporophyte. Plants of both phases were collected from four locations on the Pacific coast of Baja California, Mexico, and California, USA, between 32–4l° N latitude. Within each location, the chlorophyll-specific, light-saturated photosynthetic rates of gametophytes were generally five times greater than those of tetrasporophytes. Initial slopes of photosynthesis: irradiance curves were greater for gametophytes than for tetrasporophytes. The crust and the blade from each location were similar with respect to dark respiration rates. For tetrasporophytes from all locations, the photosynthetic temperature optima were between 12–15° C. The photosynthetic temperature optima for gametophytes ranged from 15–17° C for plants from Trinidad Head (41° N) to 22–25° C for plants from Punta Descanso (32° N). Both gametophytes and tetrasporophytes from the northernmost location had significantly higher photosynthetic rates than the same phase from the other three locations. The photosynthetic responses to light support models for the life history in which life history phases have different ecological roles. The gametophyte, thought to be specialized for rapid growth and competition, may allocate more resources to photosynthetic machinery, hence the higher maximum photosynthetic rates. The tetrasporophyte, thought to be specialized for resistance to herbivores, may allocate more resources to structural or chemical defenses in preference to photosynthetic machinery. Consequently, the tetrasporophyte has lower photosynthetic capacity.     

High throughput culture and gametogenesis induction of <Emphasis Type="Italic">Laminaria japonica</Emphasis> gametophyte clones

In anticipation of the application of a new sporeling-raising method using gametophyte clones to Laminaria commercial cultivation in China, techniques of mass culture and gametogenesis induction of L. japonica gametophyte clones were developed, as a mass of fertile gametophytes is a prerequisite for sporeling-raising with the new method. Gametophyte clones which were subjected to fed-batch culture exhibited a classical logistic growth curve. Growth rates decreased gradually after 2 months of culture, and were negatively correlated to cell density. UNOVA also showed that only cell density has a significant effect on the growth of gametophyte clones under the experimental conditions. Based on the dynamics models revealed, a culture strategy only directed at the control of cell density was adopted. By this strategy, a total of 36 kg wet weight from an initial weight of 0.75 kg was achieved after 3 months culture in 100 20-L bottles. The final average density reached 24 g L⁻¹. For the subsequent gametogenesis induction, amplificatory male and female gametophyte clones were cut, mixed and cultured in bottles under the same conditions used in amplification except for a change of photoperiod from continuous irradiance to 10 h light: 14 h dark cycle. Egg discharge occurred 10 days after the mixed culture and increased gradually with the culture duration. Most gametophytes gave rise to sporophytes 20 days after induction. Large-scale culture of gametophyte clones and gametogenesis induction for commercial cultivation in 2003–2005 have been conducted successfully.     

MORPHOLOGY AND REPRODUCTION OF THE RED ALGA ACROCHAETIUM PECTINATUM IN CULTURE12

The life history of the marine red alga Acrochaetium pectinatum (Kylin) Hamel was studied in unialgal culture using supplemented natural seawater media. The tetrasporophytes are larger than the gametophytes, have a compact filamentous basal system, and produce monosporangia and tetrasporangia. Mono-spores give rise to tetrasporophytes. Tetraspores develop into small gametophytes with unicellular bases. The gametophytes are heterothallic when small (usually less than 500 μ) but as some females become larger (2-3 mm) they produce spermatangia as well as carpogonia. Gametophytes may bear mono-sporangia in addition to carpogonia or spermatangia. These monospores give rise to gametophytes. Fertilization of the carpogonia has not been observed. The tetrasporophytes produce only monosporangia in day-lengths of 12-16 hr, but both tetrasporangia and monosporangia are formed in daylengths of 6–10 hr. Tetrasporangial production is reduced at 15 C compared with 10 C. Light intensity in the range of 5-200 ft-c (cool white fluorescent lighting) has no apparent influence on induction of tetrasporangia. Induction of tetrasporangia is not a photoperiodic response because their development is not inhibited by a brief light break in the middle of the dark period in short daylengths. Plastid morphology, origin, and frequency of sporangia and vegetative branching are variable during the ontogeny, and consequently are somewhat unreliable as taxonomic criteria. Differences in basal systems between gametophytes and tetrasporophytes also indicate that this feature, which is used to distinguish major subgeneric groupings in Acrochaetium, may not be as useful as previously thought.     

Somatic meiosis in the life history of Bonnemaisonia asparagoides and Bonnemaisonia clavata (Bonnemaisoniales,Rhodophyta) from the Iberian peninsula

Carpospores isolated from Bonnemaisonia asparagoides and Bonnemaisonia clavata (Bonnemaisoniaceae, Rhodophyta) and grown in culture developed into their respective ‘Hymenoclonium’ prostrate phases. In both species, young gametophytes were initiated directly on the prostrate phase from tetrasporangia-like protuberances. Comparison of the relative fluorescence area (rfa) of nuclear DNA over the sequence of life-history stages indicated that the lowest ploidy levels (1–2C) occurred in the gametophytes, whereas the lowest ploidy levels in the prostrate phases were 2–4C. Rfa data demonstrated that meiosis occurred in the tetrasporangia-like protuberances where 1C values were recorded. The present observations establish that B. asparagoides and B. clavata have a heteromorphic diplohaplontic life history, which involves a haploid gametophyte produced directly on a diploid prostate phase after somatic meiosis. We conclude that the life history of these taxa corresponds to the Lemanea-type. This indicates that the life history of several Bonnemaisoniales with a ‘Hymenoclonium’ phase but lacking tetrasporangia requires re-investigation.     

Growth and reproductive responses of Laminaria longicruris (Laminariales,Phaeophyta) to nutrient enrichment

A series of comparative culture experiments were conducted in order to determine responses of Laminaria longicruris male and female gametophytes and juvenile sporophytes to several temperatures (5, 10, 15, 20 °C), light levels (10, 35, 75 µmol m^–2 s^–1) and media nitrogen concentrations (0, 20, 100 µM ammonium-nitrogen). Responses were measured as numbers of male and female gametophytes producing gametangia and number of sporophytes produced following fertilization. Both male and female gametogenesis was reduced at 5 and 20 °C versus 10 and 15 °C. At 20 °C gametogenesis inhibition was greater with higher levels of ammonium-nitrogen concentration (100 µM). Sporophyte production was more sensitive to light, temperature and nitrogen concentration than gametogenesis. Production of sporophytes was inhibited completely at 20 °C. At lower temperatures, increasingly higher nutrient concentrations produced greater inhibition of production of sporophytes.     

How light and biomass density influence the reproduction of delayed Saccharina latissima gametophytes (Phaeophyceae)

Kelp life-cycle transitions are complex and susceptible to various (a)biotic controls. Understanding the microscopic part of the kelp's lifecycle is of key importance, as gametophytes form a critical phase influencing, among others, the distributional limits of the species. Many environmental controls have been identified that affect kelp gametogenesis, whose interactive effects can be subtle and counterintuitive. Here we performed a fully factorial experiment on the (interactive) influences of light intensity, light quality, and the Initial Gametophyte Density (IGD) on Saccharina latissima reproduction and vegetative growth of delayed gametophytes. A total of 144 cultures were followed over a period of 21 d. The IGD was a key determinant for reproductive success, with increased IGDs (≥0.04 mg DW · mL⁻¹) practically halting reproduction. Interestingly, the effects of IGDs were not affected by nutrient availability, suggesting a resource-independent effect of density on reproduction. The Photosynthetically Usable Radiation (PUR), overarching the quantitative contribution of both light intensity and light quality, correlated with both reproduction and vegetative growth. The PUR furthermore specifies that the contribution of light quality, as a lifecycle control, is a matter of absorbed photon flux instead of color signaling. We hypothesize that (i) the number of photons absorbed, independent of their specific wavelength, and (ii) IGD interactions, independent of nutrient availability, are major determinants of reproduction in S. latissima gametophytes. These insights help understand kelp gametophyte development and dispersal under natural conditions, while also aiding the control of in vitro gametophyte cultures.     

Temperature requirements for growth and maturation of the warm temperate kelp Eckloniopsis radicosa (Laminariales,Phaeophyta)

The annual kelp Eckloniopsis radicosa is distributed along Japanese coasts and occurs within the area with a February isotherm ranging 15–18°C and August isotherm ranging 25–28°C. In this study, the effects of temperature on the gametophyte growth and maturation, and the young sporophyte growth of E. radicosa were examined and the results are discussed in relation to the distribution of other warm‐adapted kelp species and the potential effects of climate change. The optimal temperature ranges for growth of male and female gametophytes were 23–27°C and 20–26°C, respectively. The upper survival temperature for gametophyte growth was 31°C for males and 30°C for females, respectively. The optimal temperature range for maturation of female gametophytes was ≤23°C. The optimal temperature range for growth of young sporophytes was 14–22°C. It was clarified that E. radicosa has the most warm‐adapted characteristics for growth and maturation of gametophytes among members of the Laminariales studied so far. The natural seawater temperature ranges during the growth and maturation seasons for gametophytes of E. radicosa, as well as the growth season for young sporophytes near to the northern and southern distribution limits (Izu‐Oshima: 14.9–24.5°C, Ichiki‐kushikino: 17.1–29.6°C), agreed with the experimentally determined temperature requirements. The warm‐adapted gametophyte stage and annual lifecycle are major factors enabling survival of E. radicosa in warm waters near tropical regions along the Japanese coast.     

Gametogenesis in Amphimictic and Parthenogenetic Populations of Aphelenchus avenae

The fetnale reproductive system of Aphelenchus avenae, studied in orcein-stained material, showed a peculiar structural pattern not yet reported in other nematodes. Chromosome morphology and behavior during gametogenesis could be studied in more detail than in other tylenchid or aphelenchid species investigated to date. In a bisexual population from Australia, gametogenesis was by normal meiosis and reproduction by amphimixis. The haploid chromosome number was n=8 in both males and females, and no sex chromosomes were detected. Three monosexual populations from Australia, California, and North Carolina underwent oogenesis by meiosis but reproduced hy parthenogenesis. The haploid chromosome number was n=8 in the Australia and the North Carolina populations, but n=9 in the California population. Spermatogenesis in temperature-induced males of the California population was by normal meiosis, and sperm had n=9 chromosomes. Most chronmsomes consisted of a central euchromatic section and two characteristic heterochromatic ends. No centromere was observed in any chronmsome. The relationship hetween the California population with n=9 and all the other populations with n=8 chromosomes is not well understood.     

Effects of light intensity and nutrients on gametophytes and gametogenesis of Lessonia nigrescens Bory (Phaeophyta)

Meiospores of Lessonia nigrescens Bory collected in Central Chile, were cultivated under different light and nutrient conditions to test for their influence on the development of gametophytes and gametogenesis. The microscopic stages of the life cycle are described. Under 45 μE·m^?2 ·s^?1 (12 h daily) and sea water enriched with nitrate and phosphate, development of few-celled gametophytes, gametogenesis and sporophyte differentiation occur within 35 days. At 10 μE ·m^?2 · s^?1, most gametophytes are multicellular and remain vegetative up to 65 days. Only enriched sea water or enriched sea water with double concentrations of phosphate lead to fertile few-celled gametophytes under 45 μE · m^?2 · s^?1. The addition of double concentrations of nitrate allows for the development of few-celled gametophytes but inhibits gametogenesis. The average gametophyte size progressively decreases when increasing amounts of nitrate and phosphate are added to the media.     

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.     

A MULTISPECIES LABORATORY ASSESSMENT OF RAPID SPOROPHYTE RECRUITMENT FROM DELAYED KELP GAMETOPHYTES1

Recent work suggests that the ability to delay reproduction as resistant haploid gametophytes may be important for seaweeds that experience unpredictable disturbances or seasonal periods of poor conditions that result in adult sporophyte absence. Further, delayed gametophytes of some kelp species (order Laminariales) may produce sporophytes more rapidly than if they had never experienced a delay, conferring a competitive advantage when conditions improve or after disturbance events. Here, it was determined that the gametophytes of the canopy‐forming kelp Macrocystis pyrifera (L.) C. Agardh could delay reproduction in a one‐ to two‐cell state (<50 μm) for at least 7 months when grown under nutrient‐limiting conditions. These stages retained reproductive viability and produced sporophytes within 5 d once nutrients were increased. This finding suggests that gametophytes could potentially promote recovery of M. pyrifera populations after extended periods of sporophyte absence. In addition, the time required for sporophyte production between gametophytes of the four most conspicuous kelp species in Southern California that had delayed reproduction and gametophytes that had not was compared. For these four kelp species, a delay of at least 30 d conferred a 40%–76% reduction in the time required for sporophyte production once nutrients were received. Fecundity did not decrease with delay duration, suggesting there is no apparent cost of delayed development for kelps as has been observed in other organisms. Thus, delayed development may be a viable strategy for surviving and initially dominating in environments with variable quality.     

Morphology and reproduction of Bostrychia pinnata (Rhodomelaceae, Ceramiales) in laboratory culture

In unialgal culture, isolates of vegetative plants of Bostrychia pinnata often developed tetrasporangial stichidia and released viable tetraspores. Most tetra‐sporelings developed normal branching before reproduction, however, some sporelings developed procarps, and later, spermatangia on juvenille unbranched stages (< 1 mm). Most normally branched gametophytes (> 2 mm) were initially female before becoming bisexual when older and larger (> 5 mm). Unisexual male gametophytes were not seen in culture. Carposporophytes developed slowly (40–70 days) and were sometimes abortive or produced reduced numbers of carpo‐sporangia (10–15 in cultures compared with 40–50 in field material). Carpospores germinated more slowly than tetraspores and the tetrasporophytes required up to 6 months to reach reproductive maturity. The Polysi‐phonia‐type life‐history was completed in 9–12 months in the laboratory. Peripherohaptera were absent in cultured plants. The isolates from Florida, Guatemala and Peru did not reproduce sexually, but regularly underwent lower branch abscission as a means of vegetative reproduction. After several years in culture, most individual isolates became self‐incompatible. In the female, 1 or 2 procarps formed on each axial segment and most were 4‐celled with a few having 3 cells. A single branched sterile group of 3–7 cells was also present. After fertilization, the diploid nucleus in the carpogonium divided twice, isolating the capping element and trichogyne and establishing the connecting element adjacent to the auxiliary cell. In mature normal cystocarps the sterile group persisted and secreted mucilage into the central cavity. The mature pericarp was four layers thick (one layer of axial filaments and three cortical cell layers). Pseudocystocarps were common, produced no carpo‐sporangia, contained elongate sterile cells, and were enclosed with a partially developed pericarp. Carpogonia in which 4 nuclei were usually seen may indicate developmental failure resulting in pseudocystocarps.