Did the Late Ordovician mass extinction event trigger the earliest evolution of ‘strophodontoid’ brachiopods?

‘Strophodontoid’ brachiopods represented the majority of strophomenide brachiopods in the Silurian and Devonian periods. They are characterized by denticles developed along the hinge line. The evolution of denticles correlated with the disappearance of dental plates and teeth and were already present when the clade originated in the Late Ordovician. Specimens of Eostropheodonta parvicostellata from the Kuanyinchiao Bed (early–middle Hirnantian, uppermost Ordovician) in the Hetaoba Section, Meitan, Guizhou Province, South China, display clear fossil population variation, during a process of loss of dental plates and the development of denticles. Three phenotypes of E. parvicostellata are recognized in a single fossil bed, likely heralding a speciation process. Non-metric multidimensional scaling (NMDS) based on five key characters of genera of the Family Leptostrophiidae shows a much wider morphospace for Silurian genera than for those in the Devonian. Phylogenetic analysis of the Family Leptostrophiidae supports the NMDS analysis and mostly tracks their geological history. The fossil population differentiation in E. parvicostellata discovered between the two phases of the Late Ordovician mass extinction event (LOME) linked to a major glaciation, suggests a Hirnantian origination of the ‘strophodontoid’ morphology, and links microevolutionary change to a macroevolutionary event.     

Mesozoic Trachypachidae (Insecta: Coleoptera) from China

Abstract: Four well‐preserved beetles, attributed to Eodromeinae within Trachypachidae, are described from the Mesozoic of Inner Mongolia, China. Eodromeus robustus sp. nov., E. daohugouensis sp. nov. and Unda chifengensis sp. nov. are from the Middle Jurassic of Daohugou. The diagnostic characters for the two genera are revised, and all species of these genera are keyed. A new genus and species, Sinodromeus liutiaogouensis gen. nov., sp. nov., is described from the Lower Cretaceous Yixian Formation of Liutiaogou. The current fossil records of Trachypachidae from China are reviewed, and only five species (four described here) can be convincingly attributed to this family. The morphological disparity of Mesozoic Eodromeinae suggests that they evolved a broad spectrum of locomotory lifestyles.     

Evolutionary relationships and systematics of Atoposauridae (Crocodylomorpha: Neosuchia): implications for the rise of Eusuchia

Atoposaurids are a group of small‐bodied, extinct crocodyliforms, regarded as an important component of Jurassic and Cretaceous Laurasian semi‐aquatic ecosystems. Despite the group being known for over 150 years, the taxonomic composition of Atoposauridae and its position within Crocodyliformes are unresolved. Uncertainty revolves around their placement within Neosuchia, in which they have been found to occupy a range of positions from the most basal neosuchian clade to more crownward eusuchians. This problem stems from a lack of adequate taxonomic treatment of specimens assigned to Atoposauridae, and key taxa such as Theriosuchus have become taxonomic ‘waste baskets’. Here, we incorporate all putative atoposaurid species into a new phylogenetic data matrix comprising 24 taxa scored for 329 characters. Many of our characters are heavily revised or novel to this study, and several ingroup taxa have never previously been included in a phylogenetic analysis. Parsimony and Bayesian approaches both recover Atoposauridae as a basal clade within Neosuchia, more stemward than coelognathosuchians, bernissartiids, and paralligatorids. Atoposauridae is a much more exclusive clade than previously recognized, comprising just three genera (Alligatorellus, Alligatorium, and Atoposaurus) that were restricted to the Late Jurassic of western Europe, and went extinct at the Jurassic/Cretaceous boundary. A putative Gondwanan atoposaurid (Brillanceausuchus) is recovered as a paralligatorid. Our results exclude both Montsecosuchus and Theriosuchus from Atoposauridae. Theriosuchus is polyphyletic, forming two groupings of advanced neosuchians. Theriosuchus (restricted to Theriosuchus pusillus, Theriosuchus guimarotae, and Theriosuchus grandinaris) spanned the Middle Jurassic to early Late Cretaceous, and is known from Eurasia and North Africa. Two Cretaceous species previously assigned to Theriosuchus (‘Theriosuchus’ ibericus and ‘Theriosuchus’ sympiestodon) are shown to be nested within Paralligatoridae, and we assign them to the new genus Sabresuchus. The revised phylogenetic placement of Theriosuchus has several implications for our understanding of eusuchian evolution. Firstly, the presence of fully pterygoidean choanae, previously regarded as a defining characteristic of Eusuchia, is not found in some basal members of Eusuchia. However, eusuchians can be distinguished from Theriosuchus and other basal neosuchians in that their choanae are posteriorly positioned, with an anterior margin medial to the posterior edge of the suborbital fenestra. This feature distinguishes eusuchians from Theriosuchus and more basal neosuchians. Secondly, our refined understanding of Theriosuchus implies that this taxon possessed only amphicoelous presacral vertebrae, and therefore fully developed vertebral procoely is likely to have evolved only once in Crocodylomorpha, on the lineage leading to Eusuchia. These and other findings presented herein will provide an important framework for understanding the neosuchian–eusuchian transition.     

GENERIC HOMES FOR BRITISH SILURIAN LINGULOID BRACHIOPODS

Abstract: Since the brachiopod Order Lingulida has been greatly revised in recent years, with particular emphasis on Cambrian and Ordovician genera, many well‐established Silurian linguloid species from the British Isles have been left without an appropriate genus in which to place them. This is rectified here by allocating the various species, mostly erected in the nineteenth or early twentieth centuries, to a variety of genera which have been mainly erected within the last forty years. Two new genera are erected, Mergliella, with type species Lingula bechei Davidson, 1866 , and Striatilingula, with type species Lingula? striata J. de C. Sowerby, 1839 .     

Phylogeny and taxonomic revision of all genera of Conopidae (Diptera) based on morphological data

All global genera of the fly family Conopidae are revised here. A cladistic analysis of 117 morphological characters recorded from 154 species, including representatives of 59 genera and subgenera, recovers a phylogenetic hypothesis for the family. This hypothesis is used as the basis of a new classification for the family. Both Sicini and Zodionini are removed from Myopinae and elevated to subfamilial status. A new tribe, Thecophorini, is proposed within Myopinae to accommodate Thecophora, Scatoccemyia, and Pseudoconops. Two genera, Pseudomyopa and Parazodion, are removed from Dalmanniinae and placed in Myopinae and Zodioninae, respectively. Conopinae is divided into 11 tribes, seven of which are newly described (Asiconopini, Caenoconopini, Gyroconopini, Microconopini, Neoconopini, and Siniconopini). Some examined species are transferred to different or new genera and subgenera. A new genus, Schedophysoconops gen. nov. , and subgenus Asiconops (Aegloconops) subgen. nov. within Conopinae are described. A review of character evolution and phylogeography is included in light of the new classification. A catalogue of all genus‐group names is included with new emendations noted.     

Phylogenetic classification of the Drosophilidae Rondani (Diptera): the role of morphology in the postgenomic era

Molecular sequences now overwhelm morphology in phylogenetic inference. Nonetheless, most molecular studies are conducted on a limited number of taxa, as DNA rarely can be analysed from old museum types or fossils. During the last 20 years, more than 150 molecular studies have challenged the current phylogenetic classification of the family Drosophilidae Rondani based on morphological characters. Most studies concerned a single genus, Drosophila Fallén, and included only few representative species from 17 out of the 78 genera of the family. Therefore, these molecular studies were unable to provide an alternative classification scheme. A supermatrix analysis of seven nuclear and one mitochondrial genes (8248 bp) for 33 genera was conducted using outgroups from one calyptrate and four ephydroid families. The Bayesian phylogeny was consistent with previous molecular studies including whole genome sequences and divided the Drosophilidae into four monophyletic clades. Morphological characters, mostly male genitalia, then were compared thoroughly between the four clades and homologous character states were identified. These states were then checked for 70 genera and a revised phylogenetic, family‐group classification for the Drosophilidae is proposed. Two genera –Cladochaeta Coquillett and Diathoneura Duda – of the tribe Cladochaetini Grimaldi are transferred to the family Ephydridae. The Drosophilidae is divided into two subfamilies: Steganinae Hendel (30 genera) and Drosophilinae Rondani (43 genera). A further two genera, Apacrochaeta Duda and Sphyrnoceps de Meijere, are incertae sedis, and Palmophila Grimaldi, is synonymized with Drosophila syn.n. The Drosophilinae is subdivided into two tribes: the re‐elevated Colocasiomyini Okada (nine genera) and Drosophilini Okada. The paraphyly of the genus Drosophila was not resolved to avoid affecting the binomina of important laboratory model species; however, its subgeneric classification was revised in light of molecular and morphological data. Three subgenera, namely Chusqueophila Brncic, Phloridosa Sturtevant and Psilodorha Okada, were synonymized with the subgenus Drosophila (Drosophila) Fallén syns.n. Among the 45 species groups and 5 species complexes of Drosophila (Drosophila), 22 groups and 1 complex were transferred to the subgenus Drosophila (Siphlodora) Patterson & Mainland and 6 groups, 2 species subgroups and 3 complexes are considered incertae sedis within the genus Drosophila. Different morphological characters provide different signals at different phylogenetic scales: thoracic characters (wing venation and presternal shape) discriminate families; grasping and erection‐related characters discriminate subfamilies to tribes; whereas phallic paraphyses, i.e. auxiliary intromittent organs, discriminate genera and Drosophila subgenera. The study shows the necessity of analysing morphological characters within a molecular phylogenetic framework to translate molecular phylogenies into taxonomically‐comprehensive classifications.     

Caradoc strophomenoid and plectambonitoid brachiopods from Wales and the Welsh Borderland

Abstract: Ordovician strophomenoidean and plectambonitoidean brachiopods are reviewed and partially redescribed from the Caradoc (Sandbian to Early Katian) age rocks of Wales and the Welsh Borderland of England, then forming part of the Avalonia Terrane. There are 51 nominal species available, of which 12 are synonymised here and many transferred to different genera. Strophomenoids occurred sporadically, often rarely but occasionally abundantly, particularly in mid‐shelf benthic assemblages, and 21 named species and subspecies are noted, including the new Kiaeromena (Kiaeromena) harperi, as well as six in open nomenclature. The common genus Kjaerina, with six species known from the area, is revised in detail. In contrast to the strophomenoids, plectambonitoids were more common in shallow and mid‐shelf assemblages, including Eoplectodonta (Eoplectodonta) abigailae sp. nov., with Sowerbyella in particular dominating many bedding planes, and 17 named species are reviewed, as well as three in open nomenclature. The distinctive Gunningblandella, hitherto known only from Australia and Kazakhstan, is recorded for the first time from Europe. The Avalonian strophomenides are compared with those from neighbouring terranes, particularly Laurentia and Baltica, during a time when the seas between those terranes were narrowing, and the faunas from the three terranes are found to be largely similar at the generic level, although there are few species in common. In contrast, there are few genera in common between those three terranes and the Mediterranean Province of Northwestern Gondwana.     

Revision of <Emphasis Type="Italic">Actinastrea</Emphasis>, the most common Cretaceous coral genus

The very common and species-rich Scleractinian genus Actinastrea (family Actinastraeidae, suborder Archeocaeniina) is revised on the basis of the type material of its type species and additional material from the type locality. A lectotype is designated for the type species. It was discovered that Jurassic to Early Cretaceous corals currently assigned to Actinastrea do not fit into the concept of this genus. These species belong to the genus Stelidioseris, which is also revised on the basis of the type of the type species, including designating a lectotype. These two genera are distinguished by various characteristics: septal external parts are swollen in Actinastrea but not in Stelidioseris, the costae are confluent in Stelidioseris but not in Actinastrea, the coenosteum is granulated in Actinastrea but narrow than in Actinastrea and only with costae in Stelidioseris. Actinastrea is restricted to the Late Cretaceous (Late Turonian—Maastrichtian), whereas Stelidioseris originates in the Jurassic and reaches into the Late Cretaceous, but is less common from the Turonian on.     

Implications of fossil conifers for the phylogenetic relationships of living families

Fossils have played a central role in our understanding of the evolution of conifers. Interpretation of the seed cone as a compound strobilus and the homologies of the ovuliferous scales of modern conifers with the axillary dwarf shoot of Pennsylvanian forms are based on fossils. Similarly, early evolutionary trends involving the reduction, fusion, and planation of the fertile and sterile elements of the axillary dwarf shoot, leading to structures characteristic of modern families, are documented in Late Permian and Triassic conifers. However, a phylogeny elucidating the derivation of modern families from fossil forms based on shared derived features has been elusive. The present cladistic treatment using 11 characters of ovulate cones and one of pollen grains suggests three phylogenetic groups of Late Paleozoic conifers, represented loosely by the Emporicaceae, Utrechtiaceae, and Majonicaceae of Mapes and Rothwell. The Taxaceae appears to have diverged from ancestors within the Utrechtiaceae, whereas the other modern families owe their origins to the Majonicaceae. The origin of the Taxodiaceae appears to have been biphyletic.Taxodium, Cupressus andSciadopitys are strongly linked toDolmitia of the Majonicaceae, butCryptomeria, Cunninghamia andAraucaria are grouped together and diverge basal to the former taxa.Pinus branches from a position basal to the known genera of the Majonicaceae and all modern families except the Taxaceae.Podocarpus also diverges basal toMajonica but may share an ancestor with this genus;Cepahalotaxus diverges basal to theDolmitiaPseudovoltzia subclade but distal toMajonica. Similarly, the Cheirolepidiaceae originated from basal members of the Majonicaceae and shows no close phylogenetic relationship with any modern family. Except for a strong linkage betweenCycadocarpidium and theAraucariaCunninghamia subclade, genera of the Voltziaceae appear to have branched more or less independently from within the Majonicaceae and show no strong affinity with modern conifers. Thus differences between modern conifer families are due mainly to their divergence from different Paleozoic ancestors.     

First report of craniide brachiopods in the Palaeozoic of Iran (Pseudocrania,Ordovician), and Early to Mid‐Ordovician biogeography of the Craniida

Abstract: Lower Ordovician faunas of Bohemia (Perunica), Baltica and North China include the oldest known representatives of the Order Craniida, but otherwise in Gondwana and associated terranes, the record of craniides is sparse. Pseudocrania insperata sp. nov. from the Lashkarak Formation of the Eastern Alborz Mountains is the first and as yet only record of the occurrence of craniides in the Middle Ordovician (Darriwilian) of Iran and temperate to high latitude peri‐Gondwana. Pseudocrania was known hitherto only from the Middle Ordovician of Baltoscandia and the Chu‐Ili terrane of Kazakhstan.     

Phylogeny,classification and evolution of the water scavenger beetle tribe Hydrobiusini inferred from morphology and molecules (Coleoptera: Hydrophilidae: Hydrophilinae)

The water scavenger beetle tribe Hydrobiusini contains 47 species in eight genera distributed worldwide. Most species of the tribe are aquatic, although several species are known to occur in waterfalls or tree mosses. Some members of the tribe are known to communicate via underwater stridulation. While recent morphological and molecular‐based phylogenies have affirmed the monophyly of the tribe as currently circumscribed, doubts remain about the monophyly of included genera. Here we use morphological and molecular data to infer a species‐level phylogeny of the Hydrobiusini. The monophyly of the tribe is decisively supported, as is the monophyly of most genera. The genus Hydrobius was found to be polyphyletic, and as a result the genus Limnohydrobius stat. rev. is removed from synonymy with Hydrobius, yielding three new combinations: L. melaenus comb.n. , L. orientalis comb.n. , and L. tumbius comb.n. Recent changes to the species‐level taxonomy of Hydrobius are reviewed. The morphology of the stridulatory apparatus has undergone a single remarkable transformation within the lineage, from a simple, unmodified pars stridens to one that is highly organized and complex. We present an updated key to genera, revised generic diagnoses and a list of the known distributions for all species within the tribe.     

The scleractinian genusHydnophora (revision of Tertiary species)

This report presents a systematic revision of the Tertiary species of the common reef-building coralHydnophora. Major diagnostic characters of the various Tertiary genera that have been commonly confused withHydnophora (Leptoria,Monticulastraea, Staminocoenia, Michelottiphyllia andAngeliphyllia) are defined and subsequently compared in order to establish differences and synonymies. With the exception ofLeptoria, all other genera are considered synonyms ofHydnophora. The forty-five Tertiary species ascribed toHydnophora, or to synonymous genera, have been revised and type material analyzed. Collected material from Oligocene Italian localities and Miocene localities from Somalia and Pakistan has been also analyzed in order to provide a consistent taxonomy for distinguishing species within the genus, especially for the Mediterranean Tertiary. According to this report, of the forty-five species ofHydnophora previously described, only twenty-one appear to be distinct. The Tertiary distribution of the genus, which started in the Late Paleocene, clearly shows that species richness increased significantly from Paleocene to Miocene. Originations of species were mostly concentrated during two time intervals, respectively the Chattian for the Mediterranean and the Burdigalian for Eastern Tethys. As regards the Mediterranean, the genus developed only during Chattian time with five fully described species. The genus became extinct in both Caribbean and western-central Mediterranean regions at the end of the Oligocene and subsequently developed in eastern Tethys regions during the Miocene. A new name is proposed for the only Paleocene species:Hydnophora gregoryi.     

Nektaspid arthropods from the Lower Cambrian Emu Bay Shale Lagerstätte,South Australia,with a reassessment of lamellipedian relationships

Abstract: The lower Cambrian Emu Bay Shale on Kangaroo Island, South Australia, contains the only known Cambrian Burgess Shale‐type biota in Australia. Two new lamellipedian arthropods, Emucaris fava gen. et sp. nov. and Kangacaris zhangi gen. et sp. nov., from the Emu Bay Shale Lagerstätte are described as monotypic genera that are resolved cladistically as a monophyletic group that is sister to Naraoiidae + Liwiidae and classified within the Nektaspida as a new family Emucarididae. Shared derived characters of Emucarididae involve a bipartite, elongate hypostome and elongation of the pygidium relative to the cephalic shield and very short thorax. A monophyletic Liwiidae is composed of Liwia and the Ordovician Tariccoia + Soomaspis but excludes Buenaspis, and even the membership of Buenaspis in Nektaspida is contradicted amongst the shortest cladograms. New morphological interpretations favour affinities of Kwanyinaspis with Conciliterga rather than with Aglaspidida, and Phytophilaspis with Petalopleura.     

A revision of the family Leucochloridiidae Poche (Digenea) and studies on the morphology of Leucochloridium paradoxum Carus, 1835

Summary The sporocyst and metacercaria of Leucochloridium paradoxum Carus, recovered from Succinea putris L. in Norway, are described and figured. Adults recovered from experimental infection of Taeniopygia guttata (Vieillot) with L. paradoxum metacercariae are described, using results from scanning electron microscopy, and compared with earlier information. The family Leucochloridiidae Poche is revised. Four genera are recognized within the family (i.e. Leucochloridium Carus, 1835; Urogonimus Monticelli, 1888; Urotocus Looss, 1899; Michajlovia Pojmanska, 1973). The larval ecological and morphological characteristics are tabulated for the three genera with known larval stages. The status of the genus Neoleucochloridium Kagan, 1951, is changed, on the grounds of larval similarities with Leucochloridium Carus. This means that within the genus Leucochloridium Carus three subgenera are erected, based primarily on sporocyst characteristics. These are Leucochloridium, Neoleucochloridium and Papilloleucochloridium. The diagnostic larval and adult characters are tabulated. The known larval records of species in the genus Leucochloridium are discussed and revised within the framework of this new classification. ac]19790812     

On the phylogenetic position and systematics of extant and fossil Aclopinae (Coleoptera: Scarabaeidae)

The Aclopinae is a small subfamily within the family Scarabaeidae. It currently comprises five extant genera with 28 species, and eight fossil genera with 25 species. The systematic position of Aclopinae within the family Scarabaeidae is uncertain, particularly because representative species of Aclopinae have been absent in previous phylogenetic studies. Here we performed phylogenetic analyses using morphological and molecular data to investigate the phylogenetic position of fossil and extant Aclopinae. For this objective, we expanded and revised a former morphological data matrix (composed of 68 characters) including all extant genera of Aclopinae. We complemented our morphological investigations with a molecular phylogenetic analysis based on four genes of several extant taxa of Aclopinae and a wide sample of diverse Scarabaeoidea. Our phylogenetic analyses show that all the type species of the fossil genera formerly included within Aclopinae do not belong within the extant Aclopinae clade and support both the exclusion of those fossil taxa and the monophyly of the extant genera of Aclopinae: Aclopus Erichson, Desertaclopus Ocampo & Mondaca, Gracilaclopus Ocampo & Mondaca, Neophanaeognatha Allsopp and Phanaeognatha Hope. Our results also show that the fossil taxa Prophaenognatha robusta Bai et al. and Ceafornotensis archratiras Woolley are closely related to Ochodaeidae, while the remaining type species of fossils formerly included in Aclopinae (Cretaclopus longipes (Ponomarenko), Holcorobeus vittatus Nikritin, Juraclopus rodhendorfi Nikolajev, Mesaclopus mongolicus (Nikolajev), and Mongolrobeus zherikhini Nikolajev) belong to a distinct lineage closely related to Diphyllostomatidae. Based on these results, the subfamily Aclopinae appears monophyletic and sister to the ‘pleurostict’ lineage. Consequently, we propose the following changes to the current classification of the fossil taxa: Holcorobeus monreali (Gómez‐Pallerola) belongs to Carabidae (incertae sedis) as proposed by the original author, and we place Ceafornotensis Woolley, Cretaclopus Nikolajev, Holcorobeus Nikritin, Juraclopus Nikolajev, Mesaclopus Nikolajev, Mongolrobeus Nikolajev and Prophaenognatha Bai et al. in Scarabaeoidea (incertae sedis). Furthermore, we provide an identification key to, and diagnoses of, the genera, illustrations of diagnostic characters and checklists of their included species. The evolutionary perspective presented provides new insights into the evolution of the pleurostict condition in Scarabaeoidea and the biogeography of this group, which is now regarded as Gondwanan, probably evolving during the Cretaceous and not from the upper Jurassic as previously assumed.     

PHYLOGENY OF THE RUBIACEAE AND THE LOGANIACEAE: CONGRUENCE OR CONFLICT BETWEEN MORPHOLOGICAL AND MOLECULAR DATA?

Phylogenetic analyses of 33 genera of Rubiaceae were performed using morphological and a few chemical characters. Parsimony analysis based on 29 characters resulted in eight equally parsimonious trees, with a consistency index of 0.40 and a retention index of 0.69. These results were compared to a phylogenetic analysis of the same genera based on chloroplast DNA restriction site data. There are discrepancies between the two analyses, but if we consider groupings reflected in the present classification there is much congruency. With the exception of four genera, all the genera are positioned in the same group of taxa in the two analyses. Clades of taxa representing three of the four subfamilies (~the Antirheoideae, ~the Rubioideae, and the ~Ixoroideae) are monophyletic, while the fourth subfamily Cinchonoideae is shown to be paraphyletic. Both analyses support a widened tribe Chiococceae, including the former subtribe Portlandiinae (Condamineeae). Furthermore, in both analyses the tribe Hamelieae is placed outside the subfamily Rubioideae where it is now housed. In search for the most plausible sister group to the Rubiaceae, the genus Cinchona (Rubiaceae) was analyzed together with 13 genera of the Loganiaceae, Nerium (Apocynaceae), and Exacum (Gentianaceae). Cornus (Comaceae), Olea (Oleaceae), and these two genera together were used as outgroups. The analysis, including 25 characters, 16 taxa, and with Cornus and Olea together as an outgroup, resulted in four equally parsimonious trees, with a consistency index of 0.53 and a retention index of 0.62. The non-Loganiaceae taxa Cinchona (Rubiaceae), Nerium (Apocynaceae), and Exacum (Gentianaceae) were all found to have their closest relatives within the Loganiaceae indicating that the Loganiaceae are paraphyletic and ought to be reclassified. As a result of the morphological data the most plausible sister group to the Rubiaceae is the tribe Gelsemieae of the Loganiaceae.     

Embryology of Siparunaceae (Laurales): characteristics and character evolution

Embryological characters of Siparunaceae, which are poorly understood, were studied on the basis of two constituent genera, an African Glossocalyx and a South American Siparuna, to better understand their evolution within Laurales. These two genera have many embryological characteristics in common with the other lauralean families. Noticeably, they share the multi-celled ovule archesporium (uncertain in Glossocalyx) as a synapomorphy with all the other lauralean families except Lauraceae, the anthers dehisced by valves as a synspomorphy with all the other lauralean families except Calycanthaceae and Monimiaceae, and the bisporangiate anther as a synapomorphy with Gomortegaceae and Atherospermataceae. Siparunaceae are, however, distinct from all other laularean families in having unitegmic ovules that were derived from bitegmic ovules, probably due to an elimination of the outer integument. Likewise, the lack of the testa (i.e., developed outer integument), the "endotegmic" seed coat, and the perichalazal seed at maturity are also characteristics of Siparunaceae. Within the family, Siparuna differs from Glossocalyx in having plural tetrads of megaspores and plural, starchy-rich, one-nucleate, tubular embryo sacs (autapomorphies). On the other hand, Glossocalyx is characterized by having bilaterally flattened seeds (autapomorphy). Although functional aspects of those autapomorphies are uncertain, both Glossocalyx and Siparuna show evolution in different embryological characters.