The Ultrastructure of Chaenea teres and an Analysis of the Phylogeny of the Haptorid Ciliates

Chaenea teres has typical haptorid ultrastructure. The somatic monokinetid has two transverse microtubular ribbons, an overlapping postciliary microtubular ribbon, and a laterally directed kinetodesmal fiber. The evered cytopharynx forms a dome at the apical end of the cell. The base of the dome is surrounded by oral dikinetids. The left, anterior kinetosome of the oral pair is not ciliated and has a transverse microtubular ribbon, a nematodesmata and a single postciliary microtubule. The right, posterior kinetosome is ciliated and has only postciliary microtubules. The kinetosomes at the anterior ends of the somatic kinetics are close together and their transverse microtubules and nematodesmata contribute to the support of the cytopharynx. The transverse microtubules of these oralized somatic kinetosomes, together with those from the oral dikinetids, line the cytopharynx. Accessory or bulge microtubules arise perpendicular to the transverse microtubules. A dorsal brush of three kineties of clavate cilia is found on the cell surface just posterior to the oral region. Mucocysts and a single type of toxicyst are present. The toxicysts are confined to the oral region. There are multiple ovoid macronuclei that stain weakly. Micronuclei were not observed. Cladistic analysis indicates the Chaenea may be most closely related to Fuscheria and Acropisthium. The cladistic analysis also suggests that existing taxonomies of the subclass Haptoria need to be revised. We propose some modifications to Foissner & Foissner's classification that include transferring Helicoprorodon, Actinobolina, the buetschiliids, and the balantidiids to the order Haptorida and recognizing the close relationship between pleurostomes and spathidiids.     

A Reexamination of the Ultrastructure of Didinium nasutum and a Reanalysis of the Phylogeny of the Haptorid Ciliates

ABSTRACT. The cilia of Didinium nasutum are restricted to two girdles encircling the cell. Each row of cilia in both girdles is made up of two to three anterior pairs of kinetosomes followed by several single kinetosomes. Each single kinetosome has two sets of transverse microtubules, an overlapping postciliary microtubular ribbon, and a laterally directed kinetodesmal fiber. The pairs of kinetosomes are homologous to the oral dikinetids of other haptorians: the nonciliated kinetosome of the pair has a transverse microtubular ribbon that extends to line the membrane of the proboscis, a single short postciliary microtubule, and a nematodesma; the ciliated kinetosome has a ribbon of postciliary microtubules and two sets of transverse microtubules. The presence of these characters in Didinium invalidates Leipe & Hausmann's conclusion that the Didiniidae should be removed from the subclass that contains the other haptorians (Leipe, D. D. & Hausumann, K. 1989. Somatic infraciliature of the haptorid ciliate Homalozoon vermiculare (Kinetofragminophora, Gymnostomata) Ditransversalia n. subcl. and phylogenetic implications. J. Protozool. , 36 :280–289). In light of this, the justification for a subclass Ditransversalia is challenged and shown to be unnecessary.     

Ultrastructure of Prorodon (Ciliophora, Prostomatida) I. Somatic Cortex and Some Implications Concerning Kinetid Evolution In Prostomatid and Colpodid Ciliates

ABSTRACT. This study describes the ultrastructure of the somatic cortex of Prorodon aklitolophon and Prorodon teres. the meridionally arranged somatic kineties of both species can be separated into two parts: a short anterior part, which consists of a few somatic dikinetids (in which both kinetosomes are ciliated), and a longer posterior consisting of monokinetids. the somatic monokinetids are associated with a convergent postciliary microtubular ribbon, a transverse microtubular ribbon flatly inserted in front of the kinetosome, a short and steeply extending kinetodesmal fibre attached to kinetosomal triplet 5 and 7, and a desmose anterior to triplet 3. From this desmose, two to five prekinetosomal microtubules originate and extend anteriorly. the posterior kinetosome of the somatic dikinetids is associated with the same microfibrillar and microtubular structures as the somatic monokinetid, except that no prekinetosomal microtubules originate from the desmose. the anterior kinetosome has a single postciliary microtubule and a tangentially oriented transverse microtubular ribbon. the permanent collecting canals of the unique contractile vacuole system extend parallel and adjacent to the somatic kinetics of Prorodon . the collecting canals are supported by the prekinetosomal microtubules. A similarly organized contractile vacuole system is not yet known from any other ciliate group. One of the most surprising results of this investigation was finding a significant similarity between the somatic dikinetid pattern of Prorodon and the colpodid dikinetid pattern. A hypothesis is presented to illustrate the evolution of the somatic kinetid patterns in colpodid and prostomatid ciliates.     

Ultrastructural aspects of the somatic and buccal infraciliature ofColeps amphacanthus Ehrenberg 1833

Summary Somatic and buccal infraciliature ofColeps amphacanthus Ehrenberg 1833 were studied by light and electron microscopy. The somatic kineties are composed of monokinetids and 2 dikinetids at the anterior end of each kinety. The monokinetids are associated with postciliary microtubules at triplet 9, a kinetodesmal fiber at triplet 5 and 7 and nearly radially arranged transverse microtubules at triplet 4. The associated fibrillar systems of the posterior kinetosome of the dikinetids are like those of the monokinetids. The anterior kinetosome is associated with transverse microtubules at triplet 4 and one or few postciliary microtubules at triplet 9. The anterior kinetosome bears only a short cilium.The oral ciliature is composed of a kinety of nearly circumorally arranged paroral dikinetids and 3 adoral organelles at the ventral left side of the oral opening. Nematodesmata arising from the oral ciliature form the major component of the cytopharyngeal apparatus which is lined by microtubular ribbons of postciliary origin. The buccal cavity is surrounded by oral papillae which often contain toxicysts.     

Etude Ultrastructurale du CiliéKuklikophrya dragescoi gen. n., sp. n.*

SYNOPSIS The cortical infraciliature of Kuklikophrya dragescoi gen. n., sp. n. is composed of double kinetosomes. Each kinetosome has transverse fibers. The anterior transverse fibers are associated with a sheet of dense material and the posterior transverse fibers are directed toward the posterior part of the body. The posterior kinetosome of a pair has only a short protuberance in the position of the kinetosomal fiber. The cortex has a well developed alveolar layer and a thick ecto-endoplasmic boundary. A distinctive characteristic of the buccal ciliature is the circumoral ciliature whose infraciliature is made up of pairs of cilia-bearing kinetosomes. The antero-posterior polarity of the paroral segment is in inverse relationship to that of the remaining ciliature of the organism. The adoral and preoral ciliary organelles consist of 2 rows of kinetosomes, each of which bears postciliary fibers. A frame of nematodesmata surrounds the cytopharynx which is supported by microtubular bands which impart to it a very specific laminated appearance. The “phagoplasm” is formed by “vermicelli”-like vesicles. The micronucleus is found in the perinuclear area of the macronucleus.     

Ultrastructure of Prorodon (Ciliophora, Prostomatida) Ii. Oral Cortex and Phylogenetic Conclusions

ABSTRACT. The ultrastructure of the oral region and the ultrastructural architecture of the basket of Prorodon aklitolophon and Prorodon teres are described. the oral region of Prorodon consists of: 1) A circumoral kinety at the anterior pole of the cell surrounding the typically slit-shaped cytostomial funnel. This kinety is composed of inversely oriented dikinetids in which both kinetosomes are ciliated and are associated with a very short postciliary microtubular ribbon and a few transverse microtubules; 2) Three brush rows aligned in parallel and extended meridionally in the anterior part of the cell. the individual brush rows consist of dikinetids, but in contrast to the dikinetids around the cytopharynx they are not inverted and only the anterior kinetosomes bear specialized short brush cilia and are associated with a divergent-tangential transverse microtubular ribbon. the posterior kinetosome is non-ciliated and bears a prominent convergent postciliary microtubular ribbon. Schematized dikinetid patterns of both oral regions of Prorodon are provided. In addition, a three-dimensional reconstruction of the basket of the genus Prorodon based on serial thin sections is presented. A phylogenetic tree, mainly based on stomatogenic data, is given to show the phylogenetic relationships of some prostomatid genera as well as the hypothesized sistergroup relationship of colpodid and prostomatid ciliates.     

Ultrastructure of the cytoplasm of the lower holotrichous infusorian Tracheloraphis prenanti]

The ciliature of T. prenanti Dragesco 1960 (forma oligocineta Raikov et Kovaleva, 1968) consists of 14-18 ventral and lateral longitudinal kineties with paired kinetosomes, carrying either two cilia or one cilium per kinetosome pair (in the latter case, the nonciliated kinetosome is always the posterior one). The ectoplasmic fibrillar system belongs to the postciliary type. A pair of kinetosomes shares a common basal plate. The anterior kinetosome gives rise to a short ribbon of transverse microtubules, the posterior one, to a poorly developed kinetodesmal filament and to a strong ribbon of postciliary microtubules. The latter proceeds backwards along 8 to 12 kinetosome pairs, being incorporated into a laminated postciliodesma which accompanies each kinety on its right side. Rows of Golgi elements, sending secretory vesicles and channels towards the body surface, exist beneath the kinetosome bases. Each kinety is accompanied on its left by a microfibrillar myoneme, surrounded by perimyary vesicles and underlain by a row of mitochondria. The median part of the dorsal surface is nonciliated; the cytoplasm here is rich of membrane systems, contains peripheral, electron-dense, extrusible inclusions and sometimes also bacteria. The electron-dense inclusions develop in the endoplasm, in close contact with mitochondria. The endoplasm contains also large microfibrillar spheres of unknown nature.     

Ultrastructure of the Somatic Cortex of the Gymnostome Ciliate Spathidium spathula (O. F. M.)1

The somatic cortex of Spathidium spathula is described ultrastructurally. The pellicle consists of an outer membrane and an underlying alveolar system. Numerous membrane-bound mucocysts and spherical electron-opaque bodies have similar circular sites of attachment to the outer membrane. Below these are a microfibrillar zone and an underlying region of rough ER. Mitochondria are lined up under the rough ER in longitudinal cortical ridges. Parasomal sacs are found near the basal bodies and are associated with cytoplasmic membranous sacs. Various microtubular and fiber systems are associated with single basal bodies: (1) a short kinetodesmal fiber; (2) two transverse microtubular ribbons and a transverse fiber; (3) a postciliary microtubular ribbon, initially sandwiched by two fibers, which gives rise to longitudinal subpellicular microtubules extending posteriorly for a distance of some four or five basal bodies; and (4) a system of overlapping subkinetal microtubules. A three-dimensional reconstruction is included. The somatic cortex of S spathula is similar to that reported for other Haptorida of the ciliate subclass Gymnostomata.     

Fine Structure and Molecular Phylogeny of Parametopidium circumlabens (Ciliophora: Armophorea), Endocommensal of Sea Urchins

Metopid armophoreans are ciliates commonly found in anaerobic environments worldwide; however, very little is known of their fine structure. In this study, the metopid Parametopidium circumlabens (Biggar and Wenrich 1932) Aescht, 1980, a common endocommensal of sea urchins, is investigated for the first time with emphasis on transmission electron microscopy, revealing several previously unknown elements of its morphology. Somatic dikinetids of P. circumlabens have a typical ribbon of transverse microtubules, an isolated microtubule near triplets 4 and 5 of the anterior kinetosome, plus two other microtubules between anterior and posterior kinetosomes, a short kinetodesmal striated fiber and long postciliary microtubules. In the dikinetids of the perizonal stripe, the kinetodesmal fiber is very pronounced, and there is a conspicuous microfibrillar network system associated with the kinetosomes. A new structure, shaped as a dense, roughly cylindrical mass surrounded by microtubules, is found associated with the posterior kinetosome of perizonal dikinetids. The paroral membrane is diplostichomonad and the adoral membranelles are of the “paramembranelle” type. Bayesian inference and maximum‐likelihood analysis of the 18S‐rDNA gene unambiguously placed P. circumlabens as sister group of the cluster formed by ((Atopospira galeata, Atopospira violacea) Metopus laminarius) + Clevelandellida, corroborating its classification within the Metopida.     

The Cytology of Sheep Rumen Ciliates. II. Ultrastructure of Eudiplodinium maggii1

The anterior adoral zone of syncilia (AZS) of Eudiplodinium maggii is mounted on an extrusible peristome within a vestibulum. The peristome contains cytopharyngeal components derived from the infraciliature. These components include a crescent-shaped palisade of nematodesmata, two types of sub-membrane cytopharyngeal ribbons, and an ensheathing fibrous layer enclosing a phagoplasmic zone containing the other components. A convoluted esophagus is continuous with and extends from the posterior of the cytopharynx adjacent to the macronucleus. A posterior cytoproct has specialized cytoplasm around it and associated myoneme-like elements. The skeletal plate is composed of finely granular platelets and lies under the cortex ventral to the macronucleus. The endoplasm is separated from the ectoplasm by a fibrous boundary layer. The cortex has an external glycocalyx, a membranous layer, epiplasm, and microtubular and microfilament layers. The AZS infraciliature is of the usual cntodiniomorph type, kinetosomes linked by a sub-kinetosomal rod and with associated bifurcated kinetodesma, postciliary and transverse microtubules-the latter extending into the cytopharynx—nematodesmata, and a fibrous reticulum. A possible vestigial, somatic infraciliature consisting of short, barren kinetosomes with associated basal and cortex-directed microtubules and a periodic incomplete fiber, is found subcortically throughout the cell.     

Oral Monokinetids in the Free-Living Haptorid Ciliate Enchelydium polynucleatum (Ciliophora,Enchelyidae): Ultrastructural Evidence and Phylogenetic Implications1

Special ultrastructural characteristics of the haptorid soil ciliate Enchelydium polynucleatum Foissner, 1984 are the restriction of the parasomal sacs to the area of the “brush” and finger-like projections of the food vacuole membrane into the lumen of the vacuole. The general organization of the infraciliature is similar to that of Spathidium and some buetschliids because the anterior ends of the somatic kineties are condensed and obliquely bent. Enchelydium is similar to haptorids and buetschliids in possessing monokinetid somatic fibrillar structures with the classical fibrillar associates: 1) a short kinetodesmal fiber; 2) two transverse microtubular ribbons; 3) a long postciliary microtubular ribbon; and 4) a system of overlapping subkinetal microtubules, which seems to be absent in the buetschliids. Unlike Spathidium and all other haptorids so far investigated ultrastructurally, serial sections show that there are no oral dikinetids, as in the endocommensal buetschliids and balantidiids. Instead, three to six anterior kinetids in each ciliary row have nematodesmal bundles extending into the cytoplasm and surrounding the cytopharynx. These kinetids lack cilia and all fibrillar associates except enlarged transverse ribbons, which extend anteriorly and inwards to support the cytopharynx. Other similarities between the buetschliids and Enchelydium are the conspicuous rough endoplasmic reticulum and abundant sausage-like vesicles in the oral region. As in other haptorids, Enchelydium has two types of toxicysts and one type of mucocyst. These observations strongly suggest that Enchelydium belongs to the ancestral stock of both the Haptorida and the Archistomatida. The similarities in the somatic and oral infraciliature and ultrastructure of the Haptorida and the Archistomatida suggest that they belong to the same subclass, Haptoria Corliss, 1974.     

Buccal and Somatic Infraciliature of Lagynus elegans (Engelmann, 1862) Quennersted, 1867 (Ciliophora, Prostomatea): Remarks on its Systematic Position

The somatic and buccal infraciliature of Lagynus elegans are described, and aspects of its division and conjugation are reported. Its somatic infraciliature is made up of 37–46 meridianal kineties composed of isolated kinetosomes that have thick and long kinetodesmal fibers. In the anterior zone of the cell, the circumoral infraciliature can be observed: it is composed of short, slightly oblique kinetal segments, which are formed of three kinetosomes each. The brosse of this species consists of 3 or 4 groups that possess 4 to 6 ciliated kinetosomes each; these kinetosomes lack kinetodesmal fibers. On the apical pole of the cell, surrounding the oral opening, a crown of nematodesmata is observed; these nematodesmata are connected to each other by a fibrillar structure. Taking into account these features, we propose that this genus be transferred from the order Prostomatida to a new family, Lagynidae, of the order Prorodontida.     

The Cytology of Sheep Rumen Ciliates. I. Ultrastructure of Epidinium caudatum Crawley1

Epidinium caudatum has an anterior vestibulum containing the adoral zone syncilia (AZS) on an extrusible peristome. The cytopharyngeal structures include a funnel-shaped arrangement of nematodesmata, longitudinal and transversely oriented microtubular ribbons all of which are located in the peristome, a structure which also contains filamentous phagoplasm. The origins of the microtubular ribbons indicate affinities to the rhabdos type of cytopharynx. The peristomal base is continuous with the tubular esophagus, the region connecting the two being ensheathed by a fibrous layer and low density cytoplasm. The esophagus has a microtubular/microfilamentous wall. A distinct cytoproct with associated myonemal structures occurs posteriorly. The skeletal plates consist of a large number of interconnected, variably shaped platelets and may have dual skeletal and storage functions. The endoplasm is more vesicular than the ectoplasm, the two separated by a fibrous boundary layer. The five-layered cortex has an external glycocalyx, a plasma membrane with two subtending membranes, homogeneous, microtubular, and microfilamentous layers. The syncilia of the AZS are mounted in a U-shaped band on the peristome with transversely oriented kinetics consisting of kinetosomes linked by a sub-kinetosomal rod. There is a bifurcated kinetodesma, dense support material forming a lateral spur with associated transverse microtubules, and postciliary, interkinetal, and occasional basal microtubules, nematodesmata, and a subciliary reticulum. A barren, possibly vestigial, somatic infraciliature consists of non-ciliated kinetosomes and a basal striated fiber with associated basal and perpendicular (cortical) microtubules.     

Cortical ultrastructure of Coleps bicuspis Noland, 1925 and the phylogeny of the class Prostomatea (Ciliophora)

The ultrastructure of Coleps bicuspis Noland, 1925 is described. The ciliate is a typical prostomate: the somatic kinetid is a monokinetid with a postciliary ribbon at triple 9, a kinetodesmal fibril originating near triplets 5, 6, 7 and an apparently radial transverse ribbon at triplet 4. The oral area is circular and has three brosse kineties associated with it. The brosse kineties are composed of dikinetids whose anterior kinetosome bears a tangential transverse ribbon and whose posterior kinetosome bears the fibrillar associates typical of a somatic monokinetid. The oral dikinetids are oriented parallel to the circumference of the oral cavity, which is surrounded by oral papillae and oral ridges. Pairs of nematodesmata, originating from oral dikinetid kinetosomes, are typically triangular in transection. A phylogeny of rhabdophoran ciliates is presented using the mixed parsimony algorithm and is discussed with reference to the systematic revisions of the phylum Ciliophora.     

Ultrastructural Aspects of the Somatic Cortex and Contractile Vacuole of the Ciliate,Ichthyophthirius multifiliis Fouquet1

The trophont stage in the life cycle of Ichthyophthirius multifiliis was studied in the electron microscope. Surface ridges contain up to 24 ridge microtubules, disposed as a ribbon. Kinetosomes show the classic morphology of 9 triplets of microtubules. Associated with each kinetosome is a kinetodesmal fibril, originating in proximity to triplets 5, 6, and 7, and having a 30 nm periodicity; 3 to 5 postciliary microtubules, originating between triplets 8 and 9; and up to 3 transverse microtubules, originating at triplet 4, as well as a parasomal sac. Each cell is partially enclosed by a system of 3 “unit” membranes: the outer limiting membrane, and the outer and inner alveolar membranes. The last two membranes define the alveolar sac. Mucocysts, each with a dense core, are present in large numbers. The contractile vacuole system includes the contractile vacuole, associated tubules and vesicles, injection canals, a discharge canal, and a pore. Microtubules abound in the walls of the contractile vacuole, injection and discharge canals, and in the region of the pores, where both ring and radial microtubular arrangements are noted. The ultrastructure suggests that I. multifiliis is more closely related to Tetrahymena pyriformis than to Paramecium aurelia.     

Light and Electron Microscopic Observations on the Heterotrich Ciliate Climacostomum virens

SYNOPSIS. Alveolar membranes and an epiplasm exist under the cell membrane of the noncontractile heterotrich ciliate Climacostomum virens. Postciliary microtubular ribbons join at the right of each somatic kinety to form a Km fiber. Two transverse microtubular fibers occur per kinetosomal pair. A myonemal network interconnects the kinetosomal bases intrakinetally and interkinetally. Ultrastructural comparisons are made between the contractile and noncontractile heterotrichs.
The buccal cortex consists of an adoral zone of membranelles, a peristomal field, a buccal tube, the apical membranelles, and a haplokinety. The kineties of the peristomal field and buccal tube are rows of paired kinetosomes, with a postciliary ribbon of microtubules arising from the posterior kinetosome of each pair, and a transverse ribbon and an oblique ribbon from the anterior kinetosome. No Km fibers exist in this region. The haplokinety is a collar of paired kinetosomes surrounding the cytostome; a postciliary microtubular ribbon descends from each kinetosomal pair into the cytostomal region. Ultrastructural details of the buccal cortex of C. virens and other heterotrichs are compared. The nemadesmata which lie under the membranelles are implicated in the body bending of C. virens.
Algae endosymbiotic in the cytoplasm of C. virens are described.     

The Cytology of Sheep Rumen Ciliates. III. Ultrastructure of the Genus Entodinium (Stein)1

This study examines previously undescribed general and cytopharyngeal features of the genus Entodinium. The cytopharynx contains three types of microtubular ribbons underlying the cytostomal membrane as well as a loose palisade of nematodesmata. A protoesophagus composed of microtubular bundles associated with a fibrous wall lies internally to one side of an extrusible peristome on which the adoral zone of syncilia (AZS) is mounted. Macronuclear structures are very similar to those of other ophryoscolecids. The micronucleus has chromatin bodies forming a compact mass but lacks the thick wall found in other species. A tubular spongiome surrounds the contractile vacuole and the cytoproct is relatively undifferentiated. Cortical structure follows the usual five-layered ophryoscolecid pattern with subcortical barren kinetosomes arranged into indistinct kineties. The infraciliature of the AZS has kinetosomes set upon a subkinetal rod and with associated bifurcated kinetodesmata and transverse microtubules, some of which extend into the cytopharynx. Components newly described for Entodinium are the one to three postciliary microtubules and the interkinetosomal centro-lateral strand, all of which are present in other species of ophryoscolecid ciliates. The infraciliature of the paralabial ciliary tuft shows similar components to that of the main AZS, but lacks the subkinetal rod. The microtubular components of the cytopharynx are discussed in relation to the “alimentary” structures in other ophryoscolecids, and a relationship of these structures to dietary differences is suggested.     

The Resorption of Cilia in Cyathodinium piriforme*

SYNOPSIS. The fine structure of the cilium, kinetosome, kinetodesmal fiber, and basal microtubules has been described in Cyathodinium piriforme. The ciliary axoneme is encased in an electron-dense jacket termed the axonemal jacket. This jacket surrounds the axoneme and is found midway between the axoneme and the ciliary membrane when viewed in cross section. Before division or reorganization the cilia are withdrawn into the cell. Intact cilia surrounded by their jackets are found in the cytoplasm during the early phases of retraction. Degradation of the axonemal microtubules precedes the dissolution of the axonemal jacket. Profiles of the jackets are observed after the microtubules have been resorbed. The cilia appear to detach from the kinetosomes. Barren kinetosomes are seen below the cell surface frequently with kinetodesmal fibers still attached. Whether all or some of these barren kinetosomes contribute to the formation of the new ciliary anlage cannot be ascertained.     

Somatic Infraciliature in the Haptorid Ciliate Homalozoon vermiculare (Kinetophragminophora, Gymnostomata) Ditransversalia N. Subcl. and Phylogenetic Implications

The ultrastructure of the cortex of Homalozoon vermiculare is described. The ventral side bears 13–15 iongitudinal kineties composed of monokinetids. On the dorsal surface, there are 3 kineties, 2 of which are composed of dikinetids in the anteriormost part of the cell. Consequently there exist 3 different kinds of kinetids within the somatic cortex: 1) The monokinetids on the ventral side are associated with a kinctodesmal fibril, 2 transverse microtubular ribbons and 7 postciliary microtubules in a double-row configuration; 2) The monokinetids on the dorsal side are very similar but they are associated with just 3 very 'short postciliary microtubules; 3) The posterior kinetosome of the dorsal dikinetids bears the same fibrillar associates as the dorsal monokinetid, but it lacks the second transverse ribbon. The anterior kinetosome of each pair is associated with a single postciliary' microtubule. The kinetid organization of Homalozoon is compared to that of other members of the Haptorida. Their phylogeny is discussed. A monophyleiic taxon within the litostomate ciliates is characterized by data on the somatic kinetids, and the new subclass Ditransversalia n. subcl. is constituted. The new subclass comprises the genera Balantidium, Bryophyllum, Enchelydium, Homalozoon, Isotricha, Lacrymaria, Lepidolrachelophyllum, Spathidium and Vestibulongum .