PHYLOGENETIC AFFINITIES OF “CHANTRANSIA” STAGES IN MEMBERS OF THE BATRACHOSPERMALES AND THOREALES (RHODOPHYTA)1

This study evaluated the phylogenetic relationship among samples of “Chantransia” stage of the Batrachospermales and Thoreales from several regions of the world based on sequences of two genes—the plastid‐encoded RUBISCO LSU gene (rbcL) and the nuclear SSU ribosomal DNA gene (SSU rDNA). All sequences of “Chantransia macrospora” were shown to belong to Batrachospermum macrosporum based on both molecular markers, confirming evidence from previous studies. In contrast, nine species are now associated with “Chantransia pygmaea,” including seven species of the Batrachospermales and two of the Thoreales. Therefore, the presence of “C. macrospora” in a stream can be considered reliable evidence that it belongs to B. macrosporum, whereas the occurrence of “C. pygmaea” does not allow the recognition of any particular species, since it is associated with at least nine species. Affinities of “Chantransia” stages to particular taxa were congruent for 70.5% of the samples comparing the rbcL and SSU analyses, which were associated with the same or closely related species for both markers. Sequence divergences have been reported in the “Chantransia” stage in comparison to the respective gametophyte, and this matter deserves further attention.     

Phylogenetic placement of the Tasmanian spider Acrobleps hygrophilus (Araneae, Anapidae) with comments on the evolution of the capture web in Araneoidea

This paper studies the family‐level phylogenetic placement of the conflicting Tasmanian spider genus Acrobleps using both morphological and behavioral data. We also provide a formal taxonomic revision of Acrobleps, including information on its web architecture and natural history, as well as detailed morphological information for A. hygrophilus, its only species. Acrobleps hygrophilus lacks the typical mysmenid features. Furthermore A. hygrophilus does have all typical and diagnostic characteristics of Anapidae, except for the labral spur. We also discuss two noteworthy morphological features of Acrobleps: the pore bearing depressions of the carapace and the granulated cuticle of the spinnerets. Variation in the latter feature might provide a useful phylogenetic character. Based on the results of cladistic analyses we propose the transfer of Acrobleps from the Mysmenidae to its original placement within the Anapidae. We also propose a new lineage, informally labeled as the “clawless female clade”, which includes synaphrids, cyatholipids and “symphytognathoids.” The secondary absence of the female palpal claw provides support for the “clawless female clade.” We discuss the evolution of the orb web within anapids and other symphytognathoids based on the results of our cladistic analyses. The identical bi‐dimensional webs of the anapid Elanapis and of symphytognathids have evolved independently. Finally, we comment on the implications of one of our analyses regarding araneoid web evolution. We conclude that the taxon sample included in the previous orbicularian data matrix (modified and used in this study) is adequate to test the phylogenetic placement of Acrobleps in Anapidae but insufficient to significantly assess web evolution within Araneoidea. © The Willi Hennig Society 2007.     

Morphological clines across a karyotypic zone of house mice in Central Italy

Multivariate morphometric differentiation between chromosomal races of the mouse Mus domesticus in Central Italy was investigated using a population of 2n = 22 “CB” karyotype, three populations of standard 2n = 40 karyotype, five populations of 2n = 22 “CD” karyotype and three populations from the hybrid zone between the latter two karyotypes. Whilst populations of different karyotype generally have significantly different morphometry, canonical analysis does not reveal that the populations ordinate into distinct aggregations based on karyotype, largely because the 2n = 22 “CD” populations are so diverse. Nevertheless, canonical analysis does reveal a significant cline in morphology across the contact zone between the 2n = 40 and 2n = 22 “CD” mice. The nature of this transition, i. e. a cline 1. within the 2n = 40 range, 2. within the hybrid range (but unrelated to chromosome number) and 3. within the 2n = 22 “CD” range, tends to indicate that the morphometric divergence is due to adaptation to the different ecological regimes across which these mice are distributed rather than the phylogenetic divergence of the karyotypic races.     

Phylogenetic relationships in the Sceloporus variabilis (Squamata: Phrynosomatidae) complex based on three molecular markers,continuous characters and geometric morphometric data

The monophyly of the Sceloporus variabilis group is well established with five species and two species complexes, but phylogenetic relationships within species complexes are still uncertain. We studied 278 specimens in 20 terminals to sample all taxa in the “variabilis group,” including three subspecies in the “variabilis complex,” and two outgroups (Sceloporus grammicus and Sceloporus megalepidurus). We assembled an extensive morphological data set with discrete and continuous characters (distances and scale counts), including geometric morphometric data (landmark coordinates of three shapes), and a three‐marker molecular data set as well (ND4, 12S and RAG1). We conducted parsimony and Bayesian phylogenetic inferences on these data, including several partitioning and weighting schemes. We suggest elevating three subspecies to full species status. Therefore, we recommend recognition of nine species in the “variabilis group.” First, S. variabilis is sister to Sceloporus teapensis. In turn, Sceloporus cozumelae is sister to Sceloporus olloporus. These four species are a monophyletic group, which is sister to Sceloporus smithi. Finally, Sceloporus marmoratus is sister of the clade of five species. The other species in the “variabilis group” (Sceloporus chrysostictus, Sceloporus couchii and Sceloporus parvus) are a paraphyletic grade at the base of the tree. Our analyses reject the existence of the “variabilis complex.” We conducted a parsimony‐based ancestral reconstruction on body size (snout–vent length), femoral pores and dorsal scales and related morphological changes to geographic distribution of the species. Our phylogenetic hypothesis will allow best designs of comparative studies with species in the “variabilis group,” one of the earliest divergent lineages in the genus.     

Mitochondrial gene rearrangements suggest a new genus in the subfamily Cantharinae (Coleoptera)

The Cephalomalthinus semifumatus species group, referred to as the “semifumatus” group henceforth, is interesting because of its heterogeneous morphology resembling either Cephalomalthinus Pic, 1921 or Rhagonycha Eschscholtz, 1830. To elucidate its phylogenetic status, mitochondrial genomes of four species of the “semifumatus” group, 11 Cephalomalthinus species, and 11 Rhagonycha species were sequenced and examined. All analysed mitogenomes were similar with respect to genome size, nucleotide composition, and AT content. Surprisingly, a rearrangement of the trnW-trnC and trnY genes was detected in the “semifumatus” group, presumably caused by tandem duplication and random loss events. Furthermore, genetic distance analyses showed that the proximity of the “semifumatus” group to Cephalomalthinus and to Rhagonycha was comparable to that between the latter two. Moreover, the produced phylogeny strongly supported the monophyly of the “semifumatus” group, and molecular clock analyses dated its divergence from Cephalomalthinus to 32.52 Ma. Thus, the new genus Amphimorphus gen. nov. is suggested to comprise the “semifumatus” group, in which the observed gene rearrangement was a synapomorphy. Moreover, morphological evidence regarding the unique structure of the aedeagus supported this separation. These results indicate that mitochondrial gene rearrangement provide important phylogenetic implications for revising Cephalomalthinus, a speciose genus that is puzzling in the morphology-based taxonomy.     

ACARYOCHLORIS MARINA GEN. ET SP. NOV. (CYANOBACTERIA), AN OXYGENIC PHOTOSYNTHETIC PROKARYOTE CONTAINING CHL D AS A MAJOR PIGMENT1

The phylogenetic position of an oxygenic photosynthetic prokaryote containing chl d as a major pigment, which have been tentatively named “Acaryochloris marina,” was analyzed using small subunit rDNA sequences. Phylogenetic relationships inferred among A. marina, selected strains from the Cyanobacteria, and plastids showed that A. marina was within the cyanobacterial radiation. The A. marina lineage diverged independently from other phylogenetic subgroups of the Cyanobacteria. No organism was found to be identical or related closely to A. marina by a similarity search and phylogenetic analysis. Based on these results, in addition to the reported characteristics of the cell morphology, pigment composition, and photosynthesis, a new taxon, Acaryochloris marina Miyashita et Chihara gen. et sp. nov., is formally proposed for the oxy‐genic photosynthetic prokaryote.     

Comprehensive phylogenetic analyses of the Ruppia maritima complex focusing on taxa from the Mediterranean

Recent molecular phylogenetic studies reported high diversity of Ruppia species in the Mediterranean. Multiple taxa, including apparent endemics, are known from that region, however, they have thus far not been exposed to phylogenetic analyses aimed at studying their relationships to taxa from other parts of the world. Here we present a comprehensive phylogenetic analyses of the R. maritima complex using data sets composed of DNA sequences of the plastid genome, the multi-copy nuclear ITS region, and the low-copy nuclear phyB gene with a primary focus on the Mediterranean representatives of the complex. As a result, a new lineage, “Drepanensis”, was identified as the seventh entity of the complex. This lineage is endemic to the Mediterranean. The accessions included in the former “Tetraploid” entity were reclassified into two entities: an Asia–Australia–Europe disjunct “Tetraploid_α” with a paternal “Diploid” origin, and a European “Tetraploid_γ” originating from a maternal “Drepanensis” lineage. Another entity, “Tetraploid_β”, is likely to have been originated as a result of chloroplast capture through backcrossing hybridization between paternal “Tetraploid_α” and maternal “Tetraploid_γ”. Additional discovery of multiple tetraploidizations as well as hybridization and chloroplast capture at the tetraploid level indicated that hybridization has been a significant factor in the diversification of Ruppia.     

Morphological Identities of Two Different Marine Stramenopile Environmental Sequence Clades: Bicosoeca kenaiensis (Hilliard, 1971) and Cantina marsupialis (Larsen and Patterson, 1990) gen. nov., comb. nov.

Although environmental DNA surveys improve our understanding of biodiversity, interpretation of unidentified lineages is limited by the absence of associated morphological traits and living cultures. Unidentified lineages of marine stramenopiles are called “MAST clades”. Twenty‐five MAST clades have been recognized: MAST‐1 through MAST‐25; seven of these have been subsequently discarded because the sequences representing those clades were found to either (1) be chimeric or (2) affiliate within previously described taxonomic groups. Eighteen MAST clades remain without a cellular identity. Moreover, the discarded “MAST‐13” has been used in different studies to refer to two different environmental sequence clades. After establishing four cultures representing two different species of heterotrophic stramenopiles and then characterizing their morphology and molecular phylogenetic positions, we determined that the two different species represented the two different MAST‐13 clades: (1) a lorica‐bearing Bicosoeca kenaiensis and (2) a microaerophilic flagellate previously named “Cafeteria marsupialis”. Both species were previously described with only light microscopy; no cultures, ultrastructural data or DNA sequences were available from these species prior to this study. The molecular phylogenetic position of three different “C. marsupialis” isolates was not closely related to the type species of Cafeteria; therefore, we established a new genus for these isolates, Cantina gen. nov.     

Revised classification of the Cyanidiophyceae based on plastid genome data with descriptions of the Cavernulicolales ord. nov. and Galdieriales ord. nov. (Rhodophyta)

The Cyanidiophyceae, an extremophilic red algal class, is distributed worldwide in extreme environments. Species grow either in acidic hot environments or in dim light conditions (e.g., “cave Cyanidium”). The taxonomy and classification systems are currently based on morphological, eco-physiological, and molecular phylogenetic characters; however, previous phylogenetic results showed hidden diversity of the Cyanidiophyceae and suggested a revision of the classification system. To clarify phylogenetic relationships within this red algal class, we employ a phylogenomic approach based on 15 plastomes (10 new) and 15 mitogenomes (seven new). Our phylogenies show consistent relationships among four lineages (Galdieria, “cave Cyanidium”, Cyanidium, and Cyanidioschyzon lineages). Each lineage is distinguished by organellar genome characteristics. The “cave Cyanidium” lineage is a distinct clade that diverged after the Galdieria clade but within a larger monophyletic clade that included the Cyanidium and Cyanidioschyzon lineages. Because the “cave Cyanidium” lineage is a mesophilic lineage that differs substantially from the other three thermoacidophilic lineages, we describe it as a new order (Cavernulicolales). Based on this evidence, we reclassified the Cyanidiophyceae into four orders: Cyanidiales, Cyanidioschyzonales, Cavernulicolales ord. nov., and Galdieriales ord. nov. The genetic distance among these four orders is comparable to, or greater than, the distances found between other red algal orders and subclasses. Three new genera (Cavernulicola, Gronococcus, Sciadococcus), five new species (Galdieria javensis, Galdieria phlegrea, Galdieria yellowstonensis, Gronococcus sybilensis, Sciadococcus taiwanensis), and a new nomenclatural combination (Cavernulicola chilensis) are proposed.     

The Dolezal affair: race,gender, and the micropolitics of identity

ABSTRACT

This article treats the pairing of “transgender” and “transracial” in the intertwined discussion of Caitlyn Jenner and Rachel Dolezal as an intellectual opportunity rather than a political provocation. I situate the Dolezal affair in the context of the massive destabilization of long taken-for-granted categorical frameworks, which has significantly enlarged the scope for choice and self-fashioning in the domains of race, ethnicity, sex, gender, and sexuality. Anxieties about opportunistic, exploitative, or fraudulent identity claims have generated efforts to “police” unorthodox claims – as well as efforts to defend such claims against policing – in the name of authentic, objective, and unchosen identities. Instead of a shift from given to chosen identities, as posited by theories of reflexive modernity, we see a sharpened tension between idioms of choice, autonomy, subjectivity, and self-fashioning on the one hand and idioms of givenness, essence, objectivity, and nature on the other.     

Phylogenetic placement of DethawiaMeum,and Rivasmartinezia (Apioideae,Apiaceae): Evidence from nuclear and plastid DNA sequences

The flora of Western Europe is rich in endemic species of Apiaceae, many of which have been poorly investigated and whose phylogenetic relationships are poorly known. To investigate relationships among three endemic European genera (Dethawia, Meum, and Rivasmartinezia gen. nov.) and to ascertain their higher-level phylogenetic placements within the subfamily Apioideae, we examined nuclear ribosomal DNA ITS sequences and the plastid trnL-trnF region. Phylogenies estimated using parsimony and Bayesian inference reveal that (1) the historically known “Conioselinum chinense” Clade (Conioselinum chinense; C. scopulorum; Ligusticum canadense; L. porteri; Meum athamanticum; Mutellina purpurea; and Trochiscanthes nodiflora) comprise a strongly supported monophyletic group (100% BS); (2) the genera Dethawia and Meum comprise a strongly supported monophyletic group also included in the “Conioselinum chinense” Clade; and finally (3) a new genus (Rivasmartinezia) with one species (R. vazquezii) from the Northwestern of the Iberian Peninsula, and placed in the basal position in the “Conioselinum chinense” Clade, is described for the family Apiaceae subfamily Apioideae.     

Bony labyrinth morphology in early neopterygian fishes (Actinopterygii: Neopterygii)

Endocasts of the osseous labyrinth have the potential to yield information about both phylogenetic relationships and ecology. Although bony labyrinth morphology is well documented in many groups of fossil vertebrates, little is known for early Neopterygii, the major fish radiation containing living teleosts, gars and the bowfin. Here, we reconstruct endocasts of the bony labyrinth and associated structures for a sample of Mesozoic neopterygian fishes using high‐resolution computed tomography. Our sample includes taxa unambiguously assigned to either the teleost (Dorsetichthys, “Pholidophorus,” Elopoides) and holostean (“Aspidorynchus,” “Caturus,” Heterolepidotus) total‐groups, as well as examples of less certain phylogenetic position (an unnamed parasemionotid and Dapedium). Our models provide a test of anatomical interpretations for forms where bony labyrinths were reconstructed based on destructive tomography (“Caturus”) or inspection of the lateral wall of the cranial chamber (Dorsetichthys), and deliver the first detailed insights on inner ear morphology in the remaining taxa. With respect to relationships, traits apparent in the bony labyrinth and associated structures broadly support past phylogenetic hypotheses concerning taxa agreed to have reasonably secure systematic placements. Inner ear morphology supports placement of Dapedium with holosteans rather than teleosts, while preserved structure in the unnamed parasemionotid is generalized to the degree that it provides no evidence of close affinity with either of the crown neopterygian lineages. This study provides proof‐of‐concept for the systematic utility of the inner ear in neopterygians that, in combination with similar findings for earlier‐diverging actinopterygian lineages, points to the substantial potential of this anatomical system for addressing the longstanding questions in the relationships of fossil ray‐finned fishes to one another and living groups. J. Morphol. 279:426–440, 2018. © 2016 Wiley Periodicals, Inc.     

Aphyly: identifying the flotsam and jetsam of systematics

Many taxon names in any classification will be composed of taxa that have yet to be demonstrated as monophyletic, that is, characterized by synapomorphies. Such taxa might be called aphyletic, the flotsam and jetsam in systematics, simply meaning they require taxonomic revision. The term aphyly is, however, the same as, if not identical to, Hennig's “Restkörper” and Bernardi's merophyly. None of these terms gained common usage. We outline Hennig's use of “Restkörper” and Bernardi's use of merophyly and compare it to aphyly. In our view, application of aphyly would avoid the oft made assumption that when a monophyletic group is discovered from within an already known and named taxon, then the species left behind are rendered paraphyletic. By identifying the flotsam and jetsam in systematics, we can focus on taxa in need of attention and avoid making phylogenetic faux pas with respect to their phylogenetic status.     

Multiple origins of Hawaiian drosophilids: Phylogeography of Scaptomyza Hardy (Diptera: Drosophilidae)

Scaptomyza is a highly diversified genus in the family Drosophilidae, having undergone an explosive radiation, along with the Hawaiian‐endemic genus Idiomyia in the Hawaiian Islands: about 60% of 269 Scaptomyza species so far described are endemic to the Hawaiian Islands. Two hypotheses have been proposed for the origin and diversification of Hawaiian drosophilids. One is the “single Hawaiian origin” hypothesis: Scaptomyza and Idiomyia diverged from a single common ancestor that had once colonized the Hawaiian Islands, and then non‐Hawaiian Scaptomyza migrated back to continents. The other is the “multiple origins” hypothesis: Hawaiian Scaptomyza and Idiomyia derived from different ancestors that independently colonized the Hawaiian Islands. A key issue for testing these two hypotheses is to clarify the phylogenetic relationships between Hawaiian and non‐Hawaiian species in Scaptomyza. Toward this goal, we sampled additional non‐Hawaiian Scaptomyza species, particularly in the Old World, and determined the nucleotide sequences of four mitochondrial and seven nuclear genes for these species. Combining these sequence data with published data for 79 species, we reconstructed the phylogeny and estimated ancestral distributions and divergence times. In the resulting phylogenetic trees, non‐Hawaiian Scaptomyza species were interspersed in two Hawaiian clades. From a reconstruction of ancestral biogeography, we inferred that Idiomyia and Scaptomyza diverged outside the Hawaiian Islands and then independently colonized the Hawaiian Islands, twice in Scaptomyza, thus supporting the “multiple origins” hypothesis.     

Phylogeny of the large extinct South American Canids (Mammalia,Carnivora, Canidae) using a “total evidence” approach

South America currently possesses a high diversity of canids, comprising mainly small to medium‐sized omnivorous species, but in the Pleistocene there were large hypercarnivorous taxa that were assigned to Protocyon spp., Theriodictis spp., Canis gezi, Canis nehringi and Canis dirus. These fossils have never been included in phylogenies based on quantitative cladistics, but hand‐constructed cladograms published in the 1980s included some of them in the South American canine clade and others in the Canis clade. In this work, the phylogenetic position of the large extinct South American canids was studied using a large sample of living and extinct canids, as well as different sources of characters (e.g. DNA and 133 osteological characters). The phylogenetic analysis corroborates the inclusion of Theriodictis and Protocyon in the “South American clade”, where C. gezi is also included. In addition, the position of C. dirus as a highly derived Canis species is confirmed. The simultaneous analysis supports hypercarnivory having arisen at least three times in Caninae and once in the “South American clade”. The combination of the phylogenetic analyses, the fossil record and divergence dates estimated in previous works suggests that at least three or four independent lineages of the “South American clade” invaded South America after the establishment of the Panama bridge around 3 million years ago, plus other events corresponding to the immigration of Urocyon and Canis dirus.
© The Willi Hennig Society 2009.     

Evolutionary trajectory of fish Piscine novirhabdovirus (=Viral Hemorrhagic Septicemia Virus) across its Laurentian Great Lakes history: Spatial and temporal diversification

Piscine novirhabdovirus = Viral Hemorrhagic Septicemia Virus (VHSV) first appeared in the Laurentian Great Lakes with large outbreaks from 2005 to 2006, as a new and novel RNA rhabdovirus subgenogroup (IVb) that killed >30 fish species. Interlude periods punctuated smaller more localized outbreaks in 2007, 2010, and 2017, although some fishes tested positive in the intervals. There have not been reports of outbreaks or positives from 2018, 2019, or 2020. Here, we employ a combined population genetics and phylogenetic approach to evaluate spatial and temporal evolutionary trajectory on its G‐gene sequence variation, in comparison with whole‐genome sequences (11,083 bp) from a subset of 44 individual isolates (including 40 newly sequenced ones). Our results show that IVb (N = 184 individual fish isolates) diversified into 36 G‐gene haplotypes from 2003 to 2017, stemming from two originals (“a” and “b”). G‐gene haplotypes “a” and “b” differed by just one synonymous single‐nucleotide polymorphism (SNP) substitution, remained the most abundant until 2011, then disappeared. Group “a” descendants (14 haplotypes) remained most prevalent in the Upper and Central Great Lakes, with eight (51%) having nonsynonymous substitutions. Group “b” descendants primarily have occurred in the Lower Great Lakes, including 22 haplotypes, of which 15 (68%) contained nonsynonymous changes. Evolutionary patterns of the whole‐genome sequences (which had 34 haplotypes among 44 isolates) appear congruent with those from the G‐gene. Virus populations significantly diverged among the Upper, Central, and Lower Great Lakes, diversifying over time. Spatial divergence was apparent in the overall patterns of nucleotide substitutions, while amino acid changes increased temporally. VHSV‐IVb thus significantly differentiated across its less than two decades in the Great Lakes, accompanied by declining outbreaks and virulence. Continuing diversification likely allowed the virus to persist at low levels in resident fish populations, and may facilitate its potential for further and future spread to new habitats and nonacclimated hosts.     

A MOLECULAR PHYLOGENY OF THE DROSOPHILA WILLISTONI GROUP: CONFLICTS BETWEEN SPECIES CONCEPTS?

The six sibling species of the Neotropical Drosophila willistoni group have a long history in studies of evolutionary biology, yet to date only one molecular study, which used allozymes, has been published on the phylogeny of the group. Here we present a phylogeny of the siblings based on the sequences of two nuclear genes, period (per) and Alcohol dehydrogenase (Adh), as well as the mitochondrial gene Cytochrome oxidase I (COI). Taken individually, only per has a strong phylogenetic signal supporting a well-resolved phylogeny of the group, and this phylogeny is different from that obtained using allozymes. The COI dataset by itself produces trees that disagree with per, and neither that data nor the Adh data have a strong phylogenetic signal, as indicated by low bootstrap values for all analyses. Combining the Adh and COI datasets results in the same tree as per alone. Combining all three genes results in the same topology, which is strongly supported. Two problematic taxa, D. pavlovskiana and a “Carmody strain,” which were identified as potentially separate species based on reproductive isolation, clearly cluster in the phylogenetic analyses within D. paulistorum and D. equinoxialis, respectively. Thus, there appears to be a conflict between the biological species concept and the phylogenetic species concept.     

An updated classification for the hyper-diverse genus Corydalis (Papaveraceae: Fumarioideae) based on phylogenomic and morphological evidence

The genus Corydalis, with ca. 530 species, has long been considered taxonomically challenging because of its great variability. Previous molecular analyses, based on a few molecular markers and incomplete taxonomic sampling, were clearly inadequate to delimit sections and subgenera. We have performed phylogenetic analyses of Corydalis and related taxa, using 65 shared protein-coding plastid genes from 313 accessions (including 280 samples of ca. 226 species of Corydalis) and 152 universal low-copy nuclear genes from 296 accessions (including 271 samples of Corydalis) covering all 42 previously recognized sections and five independent “series”. Phylogenetic trees were inferred using Bayesian Inference and Maximum Likelihood. Eight selected morphological characters were estimated using ancestral state reconstructions. Results include: (i) of the three subgenera of Corydalis, two are fully supported by both the plastid and nuclear data; the third, subg. Cremnocapnos, is weakly supported by plastid DNA only, whereas in the nuclear data the two included sections form successive outgroups to the rest of the genus; (ii) among all 42 sections and five “series”, 25 sections and one “series” are resolved as monophyletic in both data sets; (iii) the common ancestor of Corydalis is likely to be a perennial plant with a taproot, yellow flowers with a short saccate spur, linear fruits with recurved fruiting pedicels, and seeds with elaiosomes; (iv) we provide a new classification of Corydalis with four subgenera (of which subg. Bipapillatae is here newly described), 39 sections, 16 of which are consistent with the previous classification, 16 sections have been recircumscribed, one section has been reinstated and six new sections are established. Characters associated with lifespan, underground structures, floral spur, fruit and elaiosomes are important for the recognition of subgenera and sections. These new phylogenetic analyses combined with ancestral character reconstructions uncovered previously unrecognized relationships, and greatly improved our understanding of the evolution of the genus.