A fossil hominoid proximal femur from Kikorongo Crater, southwestern Uganda

The external morphology of a fragmentary right proximal femur from southwestern Uganda is described here. Discovered in the Kikorongo Crater of Queen Elizabeth National Park in 1961, this specimen was informally assigned to Homo sapiens (although never described) and tentatively dated to the late Pleistocene. However, because aspects of the external morphology of the femur align the fossil with the African great apes, we suggest that the Kikorongo femur may be the first postcranial fossil of the genus Pan. Like the African apes, the Kikorongo specimen lacks both an obturator externus groove and an intertrochanteric line. It has a short femoral neck with a circular cross section, and a narrow and deep superior notch. Using resampling statistics and discriminant function analysis, the Kikorongo femur clustered with the genus Pan, as opposed to Gorilla or Homo. However, if the specimen is from Pan, it would be large for this taxon. Furthermore, features that clearly distinguish the external morphology of Plio-Pleistocene hominin proximal femora from African ape femora, such as the shape of the femoral neck in cross section and femoral neck length, have converged in Holocene humans and African apes. Unfortunately, the internal morphology of the femoral neck of the Kikorongo fossil was not discernable. Although we hypothesize that the Kikorongo femur is from the genus Pan, there is such variability in the proximal femora of modern humans that, although it would be an unusual human, it remains possible that this fossil represents H. sapiens.     

The Belohdelie frontal: new evidence of early hominid cranial morphology from the Afar of Ethiopia

Study of the Belohdelie frontal has demonstrated that this four-million-year-old specimen belongs to a very generalized hominid that may be close to the divergence point of the hominid and African ape clades. Features associated with the temporalis muscle in the Belohdelie frontal and other new hominids from Hadar (AL 333-125) and West Turkana (KNM-ER 17000) suggest that the earliest hominids shared a large anterior component of this muscle relative to the extinct and extant apes. Results of this study support the phylogenetic hypothesis put forward by many workers that A. afarensis gave rise to the “robust” Australopithecus and A. africanus clades.     

Model of a hypothetical protohominid dentition and its implications for hominid phylogeny

The complete dentition of the common ancestor ofAustralopithecus andHomo, intermediate between that of a pongid and a hominid, is virtually unknown. The maxillary dentition (P3-M2) ofRamapithecus brevirostris Lewis, 1934, a pongid from the Early Pliocene, and that of hominids from the Late Pliocene and Plio/Pleistocene is known. SinceR. brevirostris is probably ancestral to the hominids, a model of intermediate maxillary dentition (P3-M2) is extrapolated and described. The model represents a hypothetical protohominid dentition. It does not conform with the teeth ofAustralopithecus, but shows greater morphological affinity to hominine dentition and to 5 myo hominids. TheHomo lineage, therefore, may go back to the Middle Pliocene. According to the normal sequence of evolution, it is most unlikely thatAustralopithecus gave rise toHomo, but much more probable that a very early, generalizedHomo evolved into an advanced, specializedAustralopithecus.     

Significance of palynology for phylogeny ofAnnonaceae: Experiments with removal of pollen characters

Phylogenetic analyses based on morphology placeAnaxagorea and other taxa with granular monosulcate pollen, as in otherMagnoliales, at the base ofAnnonaceae. Taxa with columellar tetrads, granular tetrads, and inaperturate monads form a derived clade. To test the systematic importance of palynology, we analyzed the data set with pollen characters removed. The result was lower resolution and a different rooting of the family, betweenUvariopsis and other groups with columellar tetrads.Anaxagorea and other monosulcates are higher in the tree, implying that granular monosulcate pollen, laminar stamens, and irregular endosperm ruminations are reversals. This rooting is highly unparsimonious when pollen characters are included, and only weakly supported over theAnaxagorea rooting when pollen is excluded. Together with preliminary molecular analyses, these experiments confirm the special value of palynology in systematics ofAnnonaceae. This paper is dedicated to emer. Univ.-Prof. DrFriedrich Ehrendorfer on the occasion of his 70th birthday     

Allometry, sexual dimorphism, and phylogeny: A cladistic analysis of extant African papionins using craniodental data

This study conducts a phylogenetic analysis of extant African papionin craniodental morphology, including both quantitative and qualitative characters. We use two different methods to control for allometry: the previously described narrow allometric coding method, and the general allometric coding method, introduced herein. The results of this study strongly suggest that African papionin phylogeny based on molecular systematics, and that based on morphology, are congruent and support a Cercocebus/Mandrillus clade as well as a Papio/Lophocebus/Theropithecus clade. In contrast to previous claims regarding papionin and, more broadly, primate craniodental data, this study finds that such data are a source of valuable phylogenetic information and removes the basis for considering hard tissue anatomy “unreliable” in phylogeny reconstruction. Among highly sexually dimorphic primates such as papionins, male morphologies appear to be particularly good sources of phylogenetic information. In addition, we argue that the male and female morphotypes should be analyzed separately and then added together in a concatenated matrix in future studies of sexually dimorphic taxa. Character transformation analyses identify a series of synapomorphies uniting the various papionin clades that, given a sufficient sample size, should potentially be useful in future morphological analyses, especially those involving fossil taxa.     

The hominoid proximal radius: re-interpreting locomotor behaviors in early hominins

Studies of fossil hominins are traditionally taxonomically narrow and often exclude comparisons with hylobatids. Hence, results of functional analyses of postcrania, interpreted as indicating that early hominins are "African-ape-like" in their postcranial skeletons and positional behaviors, may reflect an artifact of inadequate taxonomic and morphological breadth of the comparative sample. To address this problem and better understand early hominin positional behaviors, this study included hylobatids in a comparative analysis, focusing on the hominoid elbow joint. Specifically, morphometric variables of the proximal radius were derived from measurements from a sample of all genera of extant hominoids and casts of extinct hominin species. Univariate and multivariate analyses were performed on these data. Results show that early hominins are morphologically diverse and are not, as a group, similar to any one extant group. Instead, the fossils resemble Pan, Gorilla, and Hylobates, and are not like modern Homo sapiens or Pongo. This suggests that the morphology of Hylobates may reflect a morphotype for all later hominoids, thus complicating the functional interpretations of fossil hominins. The implications of these results are that the proximal radius is not a sensitive indicator of locomotor behavior among hominoids since the morphology in hylobatids and Gorilla and Pan is similar despite widely varying positional repertoires. Furthermore, inferences of function from form in extinct hominins can be drastically affected by the comparative outgroup selection. A re-evaluation of the functional morphology of the proximal radius in early hominins is addressed.     

The context of Stw 573, an early hominid skull and skeleton from Sterkfontein Member 2: taphonomy and paleoenvironment

The reconstructed taphonomic and paleoenvironmental contexts of a ca. 4 million-year-old partial hominid skeleton (Stw 573) from Sterkfontein Member 2 are described through presentation of the results of our analyses of the mammalian faunal assemblage associated stratigraphically with the hominid. The assemblage is dominated by cercopithecoids (Parapapio and Papio) and felids (Panthera pardus, P. leo, Felis caracal, and Felidae indet.), based on number of identified specimens, minimum number of elements and, minimum number of individuals. In addition, the assemblage is characterized by a number of partial skeletons and/or antimeric sets of bones across all taxonomic groups. There is scant indication of carnivore chewing in the assemblage. These observations, in addition to other taphonomic data, suggest that the remains of many animals recovered in Member 2 are from individuals that entered the cave on their own-whether accidentally by falling through avens connecting the cave to the ground surface above or by intentional entry-and were then unable to escape, rather than primarily through systematic collection by a biotic, bone-accumulating agent. The taphonomic conclusion that animals with climbing proclivities (i.e., primates and carnivores) are preferentially preserved over other taxa, ultimately because of those proclivities, urges caution in assessing the fidelity of the assemblage for reconstruction of the Member 2 paleoenvironment. With that caveat, we infer that the Member 2 paleoenvironment was typified by rolling, rock-littered and brush- and scrub-covered hills, indicated by the abundant F. caracal and cercopithecoid fossils recovered and the identified presence of the extinct Caprinae Makapania broomi. In addition, the valley bottom may have retained standing water year-round, perhaps supporting some tree cover--a setting suitable for the well-represented ambush predator P. pardus and suggested by the presence of Alcelaphini. Finally, the reconstructed taphonomic and paleoenvironmental settings of Sterkfontein Member 2 are compared to penecontemporaneous sites in South and East Africa.     

Molecular phylogeny of Allograpta (Diptera, Syrphidae) reveals diversity of lineages and non-monophyly of phytophagous taxa

Phylogenetic relationships of genera Allograpta, Sphaerophoria and Exallandra (Diptera, Syrphidae) were analyzed based on sequence data from the mitochondrial protein-coding gene cytochrome c oxidase subunit I (COI) and the nuclear 28S and 18S ribosomal RNA genes. The three genera are members of the subfamily Syrphinae, where nearly all members feed as larvae on soft-bodied Hemiptera and other arthropods. Phytophagous species have recently been discovered in two subgenera of Allograpta, sg Fazia and a new subgenus from Costa Rica. Phylogenetic analyses of the combined datasets were performed using parsimony, under static alignment and direct optimization, maximum likelihood and Bayesian inference. Congruent topologies obtained from all the analyses indicate paraphyly of the genus Allograpta with respect to Sphaerophoria and Exallandra. Exallandra appears embedded in the genus Sphaerophoria, and both genera are placed within Allograpta. The distribution of phytophagous taxa in Allograpta indicates that plant feeding evolved at least twice in this group.     

Variation in enamel development of South African fossil hominids

Dental tissues provide important insights into aspects of hominid palaeobiology that are otherwise difficult to obtain from studies of the bony skeleton. Tooth enamel is formed by ameloblasts, which demonstrate daily secretory rhythms developing tissue-specific structures known as cross striations, and longer period markings called striae of Retzius. These enamel features were studied in the molars of two well known South African hominid species, Australopithecus africanus and Paranthropus robustus. Using newly developed portable confocal microscopy, we have obtained cross striation periodicities (number of cross striations between adjacent striae) for the largest sample of hominid teeth reported to date. These data indicate a mean periodicity of seven days in these small-bodied hominids. Important differences were observed in the inferred mechanisms of enamel development between these taxa. Ameloblasts maintain high rates of differentiation throughout cervical enamel development in P. robustus but not in A. africanus. In our sample, there were fewer lateral striae of Retzius in P. robustus than in A. africanus. In a molar of P. robustus, lateral enamel formed in a much shorter time than cuspal enamel, and the opposite was observed in two molars of A. africanus. In spite of the greater occlusal area and enamel thickness of the molars of both fossil species compared with modern humans, the total crown formation time of these three fossil molars was shorter than the corresponding tooth type in modern humans. Our results provide support for previous conclusions that molar crown formation time was short in Plio-Pleistocene hominids, and strongly suggest the presence of different mechanisms of amelogenesis, and thus tooth development, in these taxa.     

Size- and locomotion-related aspects of hominid and anthropoid pelves: An osteometrical multivariate analysis

Two multivariate methods — the logarithmic principal component analysis (LPCA), and the logarithmic factorial analysis (LFA) — have been used tocompare the hip bone proportions of hominoids biometrically. The results have shown that size effects among apes and hominids interact to a centain extent with locomotor specializations, which are related to the attainment of more or less terrestrial behaviors. The pelvic morphology of great apes (Pongo, Pan, Gorilla) has retained numerous morphological traits — such as a gracile and elongated hip bone —, which were inherited from common adaptations to arboreal locomotion. In spite of these common traits, the African pongids (Pan, Gorilla) present two very different pelvic morphologies corresponding to two adaptative modes of terrestrial quadrupedalism. The hip bone of humans is proportionnally short and robust, most particularly at the level of its axial part. These characteristics, as well as the whole pelvic proportions, clearly indicate that gravitational forces exert a strong pressure on the pelvic walls during bipedalism. Among hominids, the transition from an australopithecine-like pelvic pattern to a human-like one corresponds to an increase of loading constraints on the hip jiont. This seems to indicate an evident change in locomotor behavior. Progression apparently became exclusively terrestrial with the genusHomo.     

Vegetative diversification and radiation in subtribeDendrobiinae (Orchidaceae): Evidence from chloroplast DNA phylogeny and anatomical characters

The data derived from a chloroplast DNA restriction site analysis of subtribeDendrobiinae (Orchidaceae) indicate that extreme vegetative diversification is concentrated in two limited parts of this group. Overlaying the vegetative character states onto the chloroplast DNA cladogram suggests that several xeromorphic, vegetative characters evolved in the lines leading to the above-mentioned clades. Several anatomical characters are also associated with xeromorphy. These vegetative and anatomical characters facilitated the establishment of this group in various dry habitats. On the other hand, the modifications of size and number of parenchymatous cells substantially contributed to the vegetative diversification. This fact implies that a simple structural adjustment can result in a major modification of growth habits in theDendrobiinae.     

Freshwater gomphonemoid diatom phylogeny: preliminary results

A cladistic analysis of eleven freshwater gomphonemoid diatom taxa yielded three equally most parsimonious cladograms. The three cladograms suggest Gomphoneis Cleve is non-monophyletic as originally and recently circumscribed. Gomphoneis elegans and G. transsilvanica (Pant.) Krammer are more closely related to Gomphopleura Reichelt ex Tempère than to other members of the genus. Gomphoneis geitleri Kociolek & Stoermer appears to represent a monotypic lineage. Among the taxa with differentiated apical pore fields Gomphonema kaznakowi Mereschkowsky appears to be primitively astigmate.The three cladograms vary with regard to systematic affinities among the Gomphoneis herculeana (Ehrenberg) Cleve group, Miocene gomphonemoid species from Idaho (USA), gomphonemoid taxa from East Africa and Gomphonema sensu stricto. Despite variability in number of puncta rows per stria and presence of an axial plate in the Miocene species from North America, inclusion of these taxa in the analysis provided better resolution of relationships than if they were excluded.     

Archaeological predictions for hominid land use in the paleo-Olduvai Basin, Tanzania, during lowermost Bed II times

We present a preliminary predictive model of Oldowan stone artefact and scavenged larger mammal bone assemblages for 11 landscape facets modeled earlier to occur across a large portion (>300 km²) of the paleo-Olduvai Basin during lowermost Bed II times. This second phase of model-building is based on our earlier characterizations of the basin's landscape ecostructure and the inter-facet distribution of key resources and hazards probably encountered by Late Pliocene hominids (Peters & Blumenschine, 1995, 1996). Our current extension of the model of hominid–landscape interactions specifies additional theoretical components, including: (1) the assumed capabilities of Oldowan hominids (presumablyHomo habilis, primarily); (2) the landscape-facet-specific tasks they carried out; (3) the immediate stone and bone task residues they produced; and (4) the predicted composition, condition, density, and clustering of stone artefact and butchered and unbutchered bone assemblages for each facet. We develop ecological linkages between these new and formerly reported modeling components, the most fundamental of which is the facet-specific degree of tree/shrub cover abundance, and the correlated degree of competition among larger carnivores and hominids for scavengeable larger mammal carcasses. These factors condition variability among landscape facets in scavenging opportunities encountered by hominids, which in our model is the major predictor of bone and stone artefact assemblage composition. The predictive value of scavenging reflects the bias of paleoanthropological traces toward technology and butchery in their landscape context, but the model is surprisingly insensitive to what are usually thought to be critical social components of hominid land use. The predictions for the traces of hominid–landscape interactions modeled herein can be tested in the future against the landscape archaeological sample being excavated from lowermost Bed II by the Olduvai Landscape Paleoanthropology Project.     

Stratigraphy, artefact industries and hominid associations for Sterkfontein, member 5

A revised stratigraphy for the early hominid site of Sterkfontein (Gauteng Province, South Africa) reveals a complex distribution of infills in the main excavation area between 2.8 and 1.4 m.y.a, as well as deposits dating to the mid to late Pleistocene. New research now shows that the Member 4 australopithecine breccia (2.8-2.6 Ma) extends further west than was previously thought, while a late phase of Member 4 is recognized in a southern area. The artefact-bearing breccias were defined sedimentologically as Member 5, but one supposed part of these younger breccias, the StW 53 infill, lacks in situ stone tools, although it does appear to post-date 2.6 Ma when artefacts first appear in the archaeological record. The StW 53 hominid, previously referred to Homo habilis, is here argued to be Australopithecus. The first artefact-bearing breccia of Member 5 is the Oldowan Infill, estimated at 2-1.7 Ma. It occupies a restricted distribution in Member 5 east and contains an expedient, flake-based tool industry associated with a few fossils of Paranthropos robustus. An enlarged cave opening subsequently admitted one or more Early Acheulean infills associated in Member 5 west with Homo ergaster. The artefacts attest to a larger site accumulation between ca. 1.7 and 1.4 Ma, with more intensive use of quartzite over quartz and a subtle but important shift to large flakes and heavier-duty tools. The available information on palaeoenvironments is summarized, showing an overall change from tropical to sub-tropical gallery forest, forest fringe and woodland conditions in Member 4 to more open woodland and grassland habitats in the later units, but with suggestions of a wet localized topography in the Paranthropus -bearing Oldowan Infill.     

When genes meet nomenclature: tortoise phylogeny and the shifting generic concepts of Testudo and Geochelone

We used a five-gene data set (mtDNA: 12S rRNA, 16S rRNA, cyt-b; nDNA: Cmos, Rag2) comprising approximately two-thirds of all extant testudinid species and, for the first time, including all five Testudo species to investigate the question of whether all western Palaearctic testudinids are monophyletic. Further, we examined whether the recently suggested allocation of the African Geochelone pardalis in the otherwise exclusively South African genus Psammobates and of the Malagasy G. yniphora in the monotypic genus Angonoka is justified in the face of considerable morphological evidence against such placements. Our phylogenetic analyses do not support the paraphyly and generic break-up of Testudo, as suggested by previous papers using a smaller taxon sampling and mtDNA data only. We propose a continued usage of the generic name Testudo for all five western Palaearctic tortoise species. Within Testudo, two monophyletic subclades are present, one containing T. hermanni+T. horsfieldii, and the other comprising (T. kleinmanni+T. marginata)+T. graeca. Nomenclaturally, we demonstrate that Eurotestudo Lapparent de Broin et al., 2006, which was recently erected with the type species T. hermanni, is an objective junior synonym of Chersine Merrem, 1820 and Medaestia Wussow, 1916. Recognition of a monotypic genus Angonoka for G. yniphora is unwarranted according to both our re-analysis of sequence data and morphological data. Acknowledging the strong morphological similarity between G. yniphora and G. radiata, we suggest placing both species into the genus Astrochelys. Although sequence data for only one of the three Psammobates species was available for analysis, there is currently no cause to challenge the monophyly of this genus as established on the basis of morphological evidence. Thus, we hypothesize that G. pardalis is sister to a monophyletic Psammobates. In light of the clear morphological gap between G. pardalis and Psammobates species, the recognition of a distinct genus Stigmochelys for the former seems justified.     

Molecular phylogeny of Phebalium (Rutaceae: Boronieae) and related genera based on the nrDNA regions ITS 1+2

Parsimony analyses of the internal transcribed spacer regions of nuclear ribosomal DNA (ITS 1 & ITS 2) for 38 taxa sampled from the Phebalium group (Rutaceae: Boronieae) and two outgroups confirm that, with the exception of Phebalium sensu stricto and Rhadinothamnus, six of the currently recognised genera within the group are monophyletic. The data indicate that Phebaliums. str. is paraphyletic with respect to Microcybe, and Rhadinothamnus is paraphyletic with respect to Chorilaena. Rhadinothamnus and Chorilaena together are the sister group to Nematolepis. Drummondita, included as an outgroup taxon, clustered within the ingroup as sister to Muiriantha and related to Asterolasia.The phylogeny suggests that the evolution of major clades within a number of these genera (e.g. Phebalium) relates to vicariance events between eastern and south-western Australia. Leionema is an eastern genus, with the most basal taxon being the morphologically distinct Leionema ellipticum from northern Queensland. Leionema also includes one species from New Zealand, but this species (as with some others) proved difficult to sequence and its phylogenetic position remains unknown. Taxonomic changes at the generic level are recommended.The authors wish to thank Paul G.Wilson, PERTH, for advice and discussion, and Paul Forster, BRI, for collecting and providing material of Leionema ellipticum. The project was supported by a Melbourne University Postgraduate Award (to BM), the Australian Biological Resources Study (ABRS), Australian Systematic Botany Society and Wolf Den (Australia) Investments.     

A multi-gene phylogeny of Clavicipitaceae (Ascomycota, Fungi): identification of localized incongruence using a combinational bootstrap approach

Multi-gene phylogenetic analyses were conducted to address the evolution of Clavicipitaceae (Ascomycota). Data are presented here for approximately 5900 base pairs from portions of seven loci: the nuclear ribosomal small and large subunit DNA (nrSSU and nrLSU), beta-tubulin, elongation factor 1alpha (EF-1alpha), the largest and second largest subunits of RNA polymerase II (RPB1 and RPB2), and mitochondrial ATP Synthase subunit 6 (mtATP6). These data were analyzed in a complete 66-taxon matrix and 91-taxon supermatrix that included some missing data. Separate phylogenetic analyses, with data partitioned according to genes, produced some conflicting results. The results of separate analyses from RPB1 and RPB2 are in agreement with the combined analyses that resolve a paraphyletic Clavicipitaceae comprising three well-supported clades (i.e., Clavicipitaceae clade A, B, and C), whereas the tree obtained from mtATP6 is in strong conflict with the monophyly of Clavicipitaceae clade B and the sister-group relationship of Hypocreaceae and Clavicipitaceae clade C. The distribution of relative contribution of nodal support for each gene partition was assessed using both partitioned Bremer support (PBS) values and combinational bootstrap (CB) analyses, the latter of which analyzed bootstrap proportions from all possible combinations of the seven gene partitions. These results suggest that CB analyses provide a more consistent estimate of nodal support than PBS and that combining heterogeneous gene partitions, which individually support a limited number of nodes, results in increased support for overall tree topology. Analyses of the 91-taxa supermatrix data sets revealed that some nodes were more strongly supported by increased taxon sampling. Identifying the localized incongruence of mtATP6 and analyses of complete and supermatrix data sets strengthen the evidence for rejecting the monophyly of Clavicipitaceae and much of the current subfamilial classification of the family. Although the monophyly of the grass-associated subfamily Clavicipitoideae (e.g., Claviceps, Balansia, and Epichlo?) is strongly supported, the subfamily Cordycipitoideae (e.g., Cordyceps and Torrubiella) is not monophyletic. In particular, species of the genus Cordyceps, which are pathogens of arthropods and truffles, are found in all three clavicipitaceous clades. These results imply that most characters used in the current familial classification of Clavicipitaceae are not diagnostic of monophyly.     

Horned lizard (Phrynosoma) phylogeny inferred from mitochondrial genes and morphological characters: understanding conflicts using multiple approaches

The genus Phrynosoma includes 13 species of North American lizards characterized by unique and highly derived morphologies and ecologies. Understanding interspecific relationships within this genus is essential for testing hypotheses about character evolution in this group. We analyzed mitochondrial ND4 and cytochrome b gene sequence data from all species of Phrynosoma in conjunction with a previously published dataset including 12S and 16S rRNA gene sequences and morphological characters. We used multiple phylogenetic methods and diagnostic tests for data combinability and taxonomic congruence to investigate the data in separate and combined analyses. Separate data partitions resulted in several well-supported lineages, but taxonomic congruence was lacking between topologies from separate and combined analyses. Partitioned Bremer support analyses also reveals conflict between data partitions in certain tree regions. When taxa associated with well-supported clades were removed from analyses, phylogenetic signal was lost. Combined, our results initially suggest conflict between data partitions, but further tests show the data are only appropriate for phylogenetic reconstruction of those parts of the topology that were well resolved. Nonetheless, our data analyses reveal five well-supported clades: (1) Phrynosoma ditmarsi and Phrynosoma hernandesi, (2) P. ditmarsi, P. hernandesi, and Phrynosoma douglasii, (3) P. ditmarsi, P. hernandesi, P. douglasii, and Phrynosoma orbiculare, (4) Phrynosoma mcallii and Phrynosoma platyrhinos, and (5) Phrynosoma braconnieri and Phrynosoma taurus.     

A molecular phylogeny of Hebeloma species from Europe

In order to widen the scope of existing phylogenies of the ectomycorrhizal agaric genus Hebeloma a total of 53 new rDNA ITS sequences from that genus was generated, augmented by sequences retrieved from GenBank, and analysed using Bayesian, strict consensus and neighbour joining methods. The lignicolous Hebelomina neerlandica, Gymnopilus penetrans, and two species of Galerina served as outgroup taxa. Anamika indica, as well as representatives of the genera Hymenogaster and Naucoria, were included to test the monophyly of Hebeloma, which is confirmed by the results. Hebeloma, Naucoria, Hymenogaster and Anamika indica cluster in a strongly supported monophyletic hebelomatoid clade. All trees largely reflect the current infrageneric classification within Hebeloma, and divide the genus into mostly well-supported monophyletic groups surrounding H. crustuliniforme, H. velutipes, H. sacchariolens, H. sinapizans, and H. radicosum, with H. sarcophyllum being shown at an independent position; however this is not well supported. The section Indusiata divides with strong support into three groups, the position of the pleurocystidiate Hebeloma cistophilum suggests the possible existence of a third subsection within sect. Indusiata. Subsection Sacchariolentia is raised to the rank of section.