Related Genera Research Articles

Cragandhara, a new genus for two new species from India (Lepidoptera: Erebidae: Arctiinae: Lithosiini)

A new genus Cragandhara gen. n. is erected to include two new species described herein from India: Cragandhara himalaya sp. n. (type species of the new genus), and Cragandhara khasia sp. n. Adults as well as male and female genitalia of the new and the related genera and species are illustrated.

Genome sequence and characterization of Streptomyces phages Vanseggelen and Verabelle, representing two new species within the genus Camvirus

Despite the rising interest in bacteriophages, little is known about their infection cycle and lifestyle in a multicellular host. Even in the model system Streptomyces, only a small number of phages have been sequenced and well characterized so far. Here, we report the complete characterization and genome sequences of Streptomyces phages Vanseggelen and Verabelle isolated using Streptomyces coelicolor as a host. A wide range of Streptomyces strains could be infected by both phages, but neither of the two phages was able to infect members of the closely related sister genus Kitasatospora. The phages Vanseggelen and Verabelle have a double-stranded DNA genome with lengths of 48,720 and 48,126 bp, respectively. Both phage genomes contain 72 putative genes, and the presence of an integrase encoding protein indicates a lysogenic lifestyle. Characterization of the phages revealed their stability over a wide range of temperatures (30–45 °C) and pH values (4–10). In conclusion, Streptomyces phage Vanseggelen and Streptomyces phage Verabelle are newly isolated phages that can be classified as new species in the genus Camvirus, within the subfamily Arquattrovirinae.

Comprehensive Analysis of Chloroplast Genome of Hibiscus sinosyriacus: Evolutionary Studies in Related Species and Genera

The Hibiscus genus of the Malvaceae family is widely distributed and has diverse applications. Hibiscus sinosyriacus is a valuable ornamental tree, but it has not been extensively researched. This study aimed to complete the chloroplast genome of H. sinosyriacus and elucidate its evolutionary relationship with closely related species and genera. The complete chloroplast genome of H. sinosyriacus was found to be 160,892 bp in length, with annotations identifying 130 genes, including 85 coding genes, 37 tRNAs, and 8 rRNAs. Interspecific variations in the Hibiscus spp. were explored, and H. sinosyriacus has species-specific single-nucleotide polymorphisms in four genes. Genome structure analysis and visualization revealed that in the Abelmoschus genus, parts of the large single-copy region, including rps19, rpl22, and rps3, have been incorporated into the inverted repeat region, leading to a duplication and an increase in the number of genes. Furthermore, within the Malvales order, the infA gene remains in some genera. Phylogenetic analysis using the whole genome and coding sequences established the phylogenetic position of H. sinosyriacus. This research has further advanced the understanding of the phylogenetic relationships of Hibiscus spp. and related genera, and the results of the structural and variation studies will be helpful for future research.

Genkalia aravindia sp. nov.: a new diatom (Bacillariophyta) associated with wet wall habitats from Eastern Himalayas and comments on the related genera

Genkalia aravindia sp. nov.: a new diatom (Bacillariophyta) associated with wet wall habitats from Eastern Himalayas and comments on the related genera

Genetic diversity in Stryphnodendron adstringens (Mart.) Coville (Leguminosae, Caesalpinioideae, Clade Mimosoid) and its correlation with species richness of Stryphnodendron Mart. and related genera

Genetic diversity in Stryphnodendron adstringens (Mart.) Coville (Leguminosae, Caesalpinioideae, Clade Mimosoid) and its correlation with species richness of Stryphnodendron Mart. and related genera

Complete Mitochondrial Genome Assembly of an Upland Wild Rice Species, Oryza granulata and Comparative Mitochondrial Genomic Analyses of the Genus Oryza.

Wild upland rice species, including Oryza granulata, possess unique characteristics that distinguish them from other Oryza species. For instance, O. granulata characteristically has a GG genome and is accordingly classified as a basal lineage of the genus Oryza. Here, we deployed a versatile hybrid approach by integrating Illumina and PacBio sequencing data to generate a high-quality mitochondrial genome (mitogenome) assembly for O. granulata. The mitogenome of O. granulata was 509,311 base pairs (bp) with sixty-seven genes comprising two circular chromosomes, five ribosomal RNA (rRNA) coding genes, twenty-five transfer RNA (tRNA) coding genes, and thirty-seven genes coding for proteins. We identified a total of 378 simple sequence repeats (SSRs). The genome also contained 643 pairs of dispersed repeats comprising 340 palindromic and 303 forward. In the O. granulata mitogenome, the length of 57 homologous fragments in the chloroplast genome occupied 5.96% of the mitogenome length. Collinearity analysis of three Oryza mitogenomes revealed high structural variability and frequent rearrangements. Phylogenetic analysis showed that, compared to other related genera, O. granulata had the closest genetic relationship with mitogenomes reported for all members of Oryza, and occupies a position at the base of the Oryza phylogeny. Comparative analysis of complete mitochondrial genome assemblies for Oryza species revealed high levels of mitogenomic diversity, providing a foundation for future conservation and utilization of wild rice biodiversity.

Taxonomy, Ecology, and Cellulolytic Properties of the Genus Bacillus and Related Genera

Bacteria of the genus Bacillus and related genera (e.g., Paenibacillus, Alicyclobacillus or Brevibacillus) belong to the phylum Firmicutes. Taxonomically, it is a diverse group of bacteria that, to date, has not been well described phylogenetically. The group consists of aerobic and relatively anaerobic bacteria, capable of spore-forming. Bacillus spp. and related genera are widely distributed in the environment, with a particular role in soil. Their abundance in the agricultural environment depends mainly on fertilization, but can also depend on soil cultivated methods, meaning whether the plants are grown in monoculture or rotation systems. The highest abundance of the phylum Firmicutes is usually recorded in soil fertilized with manure. Due to the great abundance of cellulose in the environment, one of the most important physiological groups among these spore-forming bacteria are cellulolytic bacteria. Three key cellulases produced by Bacillus spp. and related genera are required for complete cellulose degradation and include endoglucanases, exoglucanases, and β-glucosidases. Due to probable independent evolution, cellulases are encoded by hundreds of genes, which results in a large structural diversity of these enzymes. The microbial degradation of cellulose depends on its type and environmental conditions such as pH, temperature, and various substances including metal ions. In addition, Bacillus spp. are among a few bacteria capable of producing multi-enzymatic protein complexes called cellulosomes. In conclusion, the taxonomy of Bacillus spp. and related bacteria needs to be reorganized based on, among other things, additional genetic markers. Also, the ecology of soil bacteria of the genus Bacillus requires additions, especially in the identification of physical and chemical parameters affecting the occurrence of the group of bacteria. Finally, it is worth adding that despite many spore-forming strains well-studied for cellulolytic activity, still few are used in industry, for instance for biodegradation or bioconversion of lignocellulosic waste into biogas or biofuel. Therefore, research aimed at optimizing the cellulolytic properties of spore-forming bacteria is needed for more efficient commercialization.

The complete chloroplast genome sequences of six Hylotelephium species: Comparative genomic analysis and phylogenetic relationships.

To evaluate the phylogenetic relationships between Hylotelephium and Orostachys, and to provide important information for further studies, we analyzed the complete chloroplast genomes of six Hylotelephium species and compared the sequences to those of published chloroplast genomes of congeneric species and species of the closely related genus, Orostachys. The total chloroplast genome length of nineteen species, including the six Hylotelephium species analyzed in this study and the thirteen Hylotelephium and Orostachys species analyzed in previous studies, ranged from 150,369 bp (O. minuta) to 151,739 bp (H. spectabile). Their overall GC contents were almost identical (37.7-37.8%). The chloroplast genomes of the nineteen species contained 113 unique genes comprising 79 protein-coding genes (PCGs), 30 transfer RNA genes (tRNAs), and four ribosomal RNA genes (rRNAs). Among the annotated genes, fourteen genes contained one intron, and two genes contained two introns. The chloroplast genomes of the nineteen Hylotelephium and Orostachys species had identical structures. Additionally, the large single copy (LSC), inverted repeat (IR), and small single copy (SSC) junction regions were conserved in the Hylotelephium and Orostachys species. The nucleotide diversity between the Hylotelephium chloroplast genomes was extremely low in all regions, and only one region showed a high Pi value (>0.03). In all nineteen chloroplast genomes, six regions had a high Pi value (>0.03). The phylogenetic analysis showed that the genus delimitation could not be clearly observed even in this study because Hylotelephium formed a paraphyly with subsect. Orostachys of the genus Orostachys. Additionally, the data supported the taxonomic position of Sedum taqeutii, which was treated as a synonym for H. viridescens in previous studies, as an independent taxon.

Revision of the genus Regulaneuria (Ephemeroptera, Heptageniidae,Ecdyonurinae), with description of a new species.

Regulaneuria Zhou, a genus of Ecdyonurinae endemic to China, is revised. Regulaneuria chongchongae sp. nov., a new species from Hainan Island of southern China, is described and illustrated at all stage. The differentiated characters that distinguish Regulaneuria from its closely related genus Asionurus are also clarified.

Unravelling phylogenetic relationships of the tribe Cereeae using target enrichment sequencing.

Cactaceae are succulent plants, quasi-endemic to the American continent, and one of the most endangered plant groups in the world. Molecular phylogenies have been key to unravelling phylogenetic relationships among major cactus groups, previously hampered by high levels of morphological convergence. Phylogenetic studies using plastid markers have not provided adequate resolution for determining generic relationships within cactus groups. This is the case for the tribe Cereeae s.l., a highly diverse group from tropical America. Here we aimed to reconstruct a well-resolved phylogenetic tree of tribe Cereeae and update the circumscription of suprageneric and generic groups in this tribe. We integrated sequence data from public gene and genomic databases with new target sequences (generated using the customized Cactaceae591 probe set) across representatives of this tribe, with a denser taxon sampling of the subtribe Cereinae. We inferred concatenated and coalescent phylogenetic trees and compared the performance of both approaches. Six well-supported suprageneric clades were identified using different datasets. However, only genomic datasets, especially the Cactaceae591, were able to resolve the contentious relationships within the subtribe Cereinae. We propose a new taxonomic classification within Cereeae based on well-resolved clades, including new subtribes (Aylosterinae subtr. nov., Uebelmanniinae subtr. nov. and Gymnocalyciinae subtr. nov.) and revised subtribes (Trichocereinae, Rebutiinae and Cereinae). We emphasize the importance of using genomic datasets allied with coalescent inference to investigate evolutionary patterns within the tribe Cereeae.

Pollen morphology of Matthiola and the related genus Dvorakia (Brassicaceae) in Iran

Our study focused on the pollen morphology of 13 species of Matthiola and the closely related genus Dvorakia. We used light and scanning electron microscopes to examine the importance of pollen characteristics in classifying these genera. Pollen grains were acetolysed, measured, photographed, and described based on qualitative and quantitative characteristics. Here, we present statistical analyses and multivariate statistics for quantitative data. Pollen in both genera are tricolpate and varies in shape from subprolate to prolate. We distinguish two pollen types based on lumen width. Matthiola pollen has both reticulate and macroreticulate ornamentation, while Dvorakia alyssifolia pollen is reticulate. The thickness of the exine and the width of the mesocolpium are important traits in addition to exine ornamentation. Iranian Dvorakia and Matthiola species’ pollen size can be divided into four types, one of which is Type I (Dvorakia alyssifolia type). Among the examined species, M. tomentosa has the smallest pollen grains, while D. alyssifolia has the largest grains. Pollen morphology confirms the transfer of D. alyssifolia from Matthiola to Dvorakia. Additionally, we propose a species key for the genera.

Characteristics and Comparative Analysis of Seven Complete Plastomes of Trichoglottis s.l. (Aeridinae, Orchidaceae).

Trichoglottis exhibits a range of rich variations in colors and shapes of flower and is a valuable ornamental orchid genus. The genus Trichoglottis has been expanded by the inclusion of Staurochilus, but this Trichoglottis sensu lato (s.l.) was recovered as a non-monophyletic genus based on molecular sequences from one or a few DNA regions. Here, we present phylogenomic data sets, incorporating complete plastome sequences from seven species (including five species sequenced in this study) of Trichoglottis s.l. (including two species formerly treated as Staurochilus), to compare plastome structure and to reconstruct the phylogenetic relationships of this genus. The seven plastomes possessed the typical quadripartite structure of angiosperms and ranged from 149,402 bp to 149,841 bp with a GC content of 36.6-36.7%. These plastomes contain 120 genes, which comprise 74 protein-coding genes, 38 tRNA genes, and 8 rRNA genes, all ndh genes were pseudogenized or lost. A total of 98 (T. philippinensis) to 134 (T. ionosma) SSRs and 33 (T. subviolacea) to 46 (T. ionosma) long repeats were detected. The consistent and robust phylogenetic relationships of Trichoglottis were established using a total of 25 plastid genomes from the Aeridinae subtribe. The genus Trichoglottis s.l. was strongly supported as a monophyletic group, and two species formerly treated as Staurochilus were revealed as successively basal lineages. In addition, five mutational hotspots (trnNGUU-rpl32, trnLUAA, trnSGCU-trnGUCC, rbcL-accD, and trnTGGU-psbD) were identified based on the ranking of PI values. Our research indicates that plastome data is a valuable source for molecular identification and evolutionary studies of Trichoglottis and its related genera.

Taxonomic Status and Composition of the Genera Caulaster, Porcellanaster and Eremicaster (Asteroidea) Based on Juvenile Morphology and Molecular Phylogenetic Data

The genus Porcellanaster Thomson 1878 has hitherto included two species, the type species P. ceruleus Thomson, 1878 and P. ivanovi Belyaev, 1969. According to WoRMS, the full list of junior synonyms of P. ceruleus consists of 15 specific names, including porcellanasterids of the genera Caulaster Perrier 1882 and Albatrossia Ludwig 1905. In this paper, the morphology of juvenile and adult Porcellanaster and synonymized species from the Atlantic, Pacific, Indian and Southern Oceans are studied. As a result, the composition of the genus Porcellanaster has been revised. The genus Caulaster is re-established and now includes two species, and two new morphological forms of Caulaster are described. The genera Albatrossia and Damnaster H.E.S Clark & McKnight, 1994 are considered junior synonyms of Caulaster. The number of species of the genus Porcellanaster has increased to five, including the recovered P. caulifer, P. fragilis and P. sladeni. The study of representatives of the closely related genus Eremicaster Fisher, 1905 has not led to a change in its species composition. The main diagnostic feature of Caulaster is the presence of adambulacral and dorsal pedicellariae both in adults and juveniles with a major radius (R) greater than 3.5 mm. Pedicellariae have never been found in Porcellanaster or Eremicaster at any stage of postlarval development. The species of Caulaster and some species of Porcellanaster are the smallest porcellanasterids, with the most simplified morphology and reduced papulae and paxillae responsible for respiration. These genera are located on the phylogenetic tree in different clades, which demonstrates the parallel evolution of these taxa and confirms the validity of the genus Caulaster. One of the reasons for miniaturization is likely the burrowing mode of life with feeding on subsurface sediment of low nutritional value. The simplification of morphology can be traced by many morphological charactercs in many ways, including the reduction in papulae and paxillae.

Phylogeny of Rhynchium and Its Related Genera (Hymenoptera: Eumeninae) Based on Universal Single-Copy Orthologs and Ultraconserved Elements.

The subfamily Eumeninae is a large group of fierce predatory insects that prey mainly on the larvae of Lepidoptera pests. Because of the highly similar morphologies of the genus Rhynchium and its related genera in the subfamily, including Rhynchium Spinola, Allorhynchium van der Vecht, Anterhynchium de Saussure, Pararrhynchium de Saussure, it is essential to delineate their relationships. A previous phylogenetic analysis based on mitochondrial genomes suggested the inconsistent relationships of these genera under traditional classification based on morphological characters. In this study, we first used single-copy orthologs [USCO] and ultraconserved elements [UCE] extracted from 10 newly sequenced low-coverage whole genomes to resolve the phylogenetic relationships of the above genera. The newly sequenced genomes are 152.99 Mb to 211.49 Mb in size with high completeness (BUSCO complete: 91.5-95.6%) and G + C content (36.31-38.76%). Based on extracted 5811 USCOs and 2312 UCEs, the phylogenetic relationships of Rhynchium and its related genera were: ((Allorhynchium + Lissodynerus) + (Pararrhynchium + (Anterhynchium + (Dirhynchium + Rhynchium)))), which was consistent with the mitochondrial genome results. The results supported the genus Rhynchium as monophyletic, whereas Anterhynchium was recovered as paraphyletic, with Anterhynchium (Dirhynchium) as a sister to Rhynchium and hence deserving generic status; In addition, in the genus Pararrhynchium, P. septemfasciatus feanus and P. venkataramani were separated, not clustered on a branch, just as P. septemfasciatus feanus was not together with P. striatum based on mitochondrial genomes. Since Lissodynerus septemfasciatus, the type species of the genus Lissodynerus, was transferred to Pararrhynchium, it is considered that the genus Lissodynerus should be restituted as a valid genus, not a synonym of Pararrhynchium.

On the taxonomic status of Dina ratschaensis Kobakhidze, 1958 with a description of two new species – Dina imeretiensis sp. nov. and D. samegreloensis sp. nov. (Annelida, Hirudinida: Erpobdellidae)

Two species of leeches were described from Georgia in the past, Dina ratschaensis Kobakhidze, 1958 from the Racha-Lechkhumi and Kvemo Svaneti region and Trocheta ariescornuta Grosser, Barjadze & Maghradze, 2021 from the Samegrelo-Zemo Svaneti region. Both species were the only known typical representatives of cave leeches in Georgia. Recently, two more species of the genus Dina R. Blanchard, 1894 have been found in karst caves in this country. These leeches are morphologically similar to D. ratschaensis, from which they differ in the shape of the reproductive system, primarily the shape of the cornua of the genital atrium and the shape and extension of the vasa deferentia and ovisacs, justifying the description of two new species, Dina imeretiensis Grosser, Barjadze & Maghradze sp. nov. from the Imereti region and Dina samegreloensis Grosser, Barjadze & Shavadze sp. nov. from the Samegrelo-Zemo Svaneti region. Data on the cave dwelling invertebrate communities in the studied caves are provided. Molecular data are provided, and a phylogenetic tree based on Cox1 sequences of Dina spp. and related genera is provided and discussed.

Two new species of Varicus from Caribbean deep reefs, with comments on the related genus Pinnichthys (Teleostei, Gobiidae, Gobiosomatini, Nes subgroup).

Tropical deep reefs (~40-300 m) are diverse ecosystems that serve as habitats for diverse communities of reef-associated fishes. Deep-reef fish communities are taxonomically and ecologically distinct from those on shallow reefs, but like those on shallow reefs, they are home to a species-rich assemblage of small, cryptobenthic reef fishes, including many species from the family Gobiidae (gobies). Here we describe two new species of deep-reef gobies, Varicusprometheussp. nov. and V.roatanensissp. nov., that were collected using the submersible Idabel from rariphotic reefs off the island of Roatan (Honduras) in the Caribbean. The new species are the 11th and 12th species of the genus Varicus, and their placement in the genus is supported by morphological data and molecular phylogenetic analyses. Additionally, we also collected new specimens of the closely-related genus and species Pinnichthysaimoriensis during submersible collections off the islands of Bonaire and St. Eustatius (Netherland Antilles) and included them in this study to expand the current description of that species and document its range extension from Brazil into the Caribbean. Collectively, the two new species of Varicus and new records of P.aimoriensis add to our growing knowledge of cryptobenthic fish diversity on deep reefs of the Caribbean.

Macroevolutionary trends of the Neotropical genus Ameroglossum (Linderniaceae) in rocky outcrop environments

Ameroglossum is a rare plant genus endemic to northeastern of Brazil, initially monospecific (A. pernambucense) and recently expanded by the description of eight new species and two related genera. The genus was initially placed in the family Scrophulariaceae, but this has never been phylogenetically tested. This group is ecologically restricted to rocky inselberg habitats that function as island-like systems (ILS) with spatial fragmentation, limited area, environmental heterogeneity, temporal isolation and low connectivity. Here we use a phylogenetic perspective to test the hypothesis that Ameroglossum diversification was related to island-like radiation in inselbergs. Our results support that Ameroglossum is monophyletic only with the inclusion of Catimbaua and Isabelcristinia (named here as Ameroglossum sensu lato) and this group was well-supported in the family Linderniaceae. Biogeographic analyses suggest that the ancestral of Ameroglossum and related genus arrived in South America c.a. 15 million years ago by long-distance dispersal, given the ancestral distribution of Linderniaceae in Africa. In rocky outcrop habitats, Ameroglossum s.l. developed floral morphological specialization associated with pollinating hummingbirds, compatible with an island-like model. However, no increase in speciation rate was detected, which may be related to high extinction rates and/or slow diversification rate in this ecologically restrictive environment. Altogether, in Ameroglossum key innovations involving flowers seem to have offered opportunities for evolution of greater phenotypic diversity and occupation of new niches in rocky outcrop environments.

Genetic mapping of the LOSS OF PARTHENOGENESIS locus in Pilosella piloselloides and the evolution of apomixis in the Lactuceae.

Pilosella piloselloides var. praealta (syn. P. praealta; Hieracium praealtum) is a versatile model used to study gametophytic apomixis. In this system apomixis is controlled by three loci: one that controls the avoidance of meiosis (LOA), one that controls the avoidance of fertilization (LOP) and a third that controls autonomous endosperm formation (AutE). Using a unique polyhaploid mapping approach the LOP locus was mapped to a 654 kb genomic interval syntenic to linkage group 8 of Lactuca sativa. Polyhaploids form through the gametophytic action of a dominant determinant at LOP, so the mapped region represents both a functional and a physical domain for LOP in P. piloselloides. Allele sequence divergence (ASD) analysis of the PARTHENOGENESIS (PAR) gene within the LOP locus revealed that dominant PAR alleles in Pilosella remain highly similar across the genus, whilst the recessive alleles are more divergent. A previous report noted that dominant PAR alleles in both Pilosella and Taraxacum are modified by the presence of a class II transposable element (TE) in the promoter of the gene. This observation was confirmed and further extended to the related genus Hieracium. Sufficient differences were noted in the structure and location of the TE elements to conclude that TE insertional events had occurred independently in the three genera. Measures of allele crossover amongst the polyhaploids revealed that P. piloselloides is an autopolyploid species with tetrasomic inheritance. It was also noted that the dominant determinant of LOP in P. piloselloides could transmit via a diploid gamete (pollen or egg) but not via a haploid gamete. Using this information, a model is presented of how gametophytic apomixis may have evolved in several members of the Lactuceae, a tribe of the Asteraceae.

Plastid phylogenomics of the Eriostemon group (Rutaceae; Zanthoxyloideae): support for major clades and investigation of a backbone polytomy

Most of Australia’s sclerophyllous Rutaceae belong to a clade informally known as the ‘Eriostemon group’ (including 16 genera, ~209 species). We investigated generic relationships in this group using analyses of complete plastome sequence data for 60 species and analyses of a supermatrix including sequences of four plastome spacer regions for 22 additional species. Maximum likelihood, Bayesian inference, and shortcut coalescent phylogenetic analyses produced congruent phylogenies that were highly supported, except for a series of short unsupported branches in the backbone of the Eriostemon group. We found high support for four major clades branching from this polytomy and discuss evolutionary inferences of generic relationships in each lineage. In an effort to resolve the polytomy, we analysed gene tree topologies in tree space, phylogenetic informativeness with likelihood mapping, and conducted topology tests to assess support for all possible topological resolutions of the polytomy. These approaches did not clarify the polytomy, which may be caused by insufficient data, features of plastome evolution, or rapid radiation. Results from analyses of the combined supermatrix dataset suggest that Philotheca section Philotheca is paraphyletic with regards to Drummondita and Geleznowia. In all phylogenies, Philotheca sections Corynonema and Cyanochlamys were not placed with other members of Philotheca.

Exochorion in the tribe Nymphalini (Lepidoptera: Nymphalidae): the genus Hypanartia Hbner, [1821] and comparison with related genera.

We describe and compare the exochorion of six species of Hypanartia (H. paullus, H. bella, H. lethe, H. godmanii, H. dione disjuncta, and H. trimaculata autumna) using specimens collected in the field and deposited in entomological collections. We used a standard staining technique and scanning electron microscopy to visualize and describe the main exochorionic characters, especially the ridges and the differentiation in the micropylar, perimicropylar, and transition zones in the apical region. We included plates with photographs, drawings, and schemes for clearer visualization of these structures, which are tabulated for comparison. For our characterization, we considered the ridges in colonnade as the main exochorionic feature in the Nymphalini. Thus, we made brief comparisons of these features among several species of the tribe, particularly with the genus Antanartia. The exochorionic characters agree with the separation of groups (paullus group and dione group). By examining the characteristics of H. paullus, the basal species of the paullus group, we found that ridges with conspicuous columns (e.g., in one section only) appears to be the plesiomorphic state, while those with elongated shafts and walls only in the intercolumn represent the apomorphic state.