Brassicaceae phylogeny and trichome evolution

To estimate the evolutionary history of the mustard family (Brassicaceae or Cruciferae), we sampled 113 species, representing 101 of the roughly 350 genera and 17 of the 19 tribes of the family, for the chloroplast gene ndhF. The included accessions increase the number of genera sampled over previous phylogenetic studies by four-fold. Using parsimony, likelihood, and Bayesian methods, we reconstructed the phylogeny of the gene and used the Shimodaira-Hasegawa test (S-H test) to compare the phylogenetic results with the most recent tribal classification for the family. The resultant phylogeny allowed a critical assessment of variations in fruit morphology and seed anatomy, upon which the current classification is based. We also used the S-H test to examine the utility of trichome branching patterns for describing monophyletic groups in the ndhF phylogeny. Our phylogenetic results indicate that 97 of 114 ingroup accessions fall into one of 21 strongly supported clades. Some of these clades can themselves be grouped into strongly to moderately supported monophyletic groups. One of these lineages is a novel grouping overlooked in previous phylogenetic studies. Results comparing 30 different scenarios of evolution by the S-H test indicate that five of 12 tribes represented by two or more genera in the study are clearly polyphyletic, although a few tribes are not sampled well enough to establish para- or polyphyly. In addition, branched trichomes likely evolved independently several times in the Brassicaceae, although malpighiaceous and stellate trichomes may each have a single origin.     

基于28S rDNA序列的姬小蜂科(膜翅目,小蜂总科)分类

选择28S rDNA D2区基因,针对GenBank中姬小蜂科总计542条相关序列,借助Blast Align、MUSCLE及TNT等生物信息学软件进行计算分析,提出了一种基于亚科水平的姬小蜂科快速DNA分类鉴定方法。建树结果对目前分类系统中姬小蜂科4亚科分类体系(Bouek,1988)予以支持;综合分析结果基本支持对于姬小蜂亚科以及灿小蜂亚科的分族、分属方法。同时对地位不明的两属Anselmella和Ophelimus的分类学地位提出了假设。     

Placing the woody tropical genera of Polygonaceae: A hypothesis of character evolution and phylogeny

? Premise of the study: Taxonomic groups have often been recognized on the basis of geographic distinctions rather than accurately representing evolutionary relationships. This has been particularly true for temperate and tropical members from the same family. Polygonaceae exemplifies this problem, wherein the woody tropical genera were segregated from temperate members of the family and placed in the subfamily Polygonoideae as two tribes: Triplarideae and Coccolobeae. Modern phylogenetic studies, especially when inferred from many lines of evidence, can elucidate more probable hypotheses of relationships. This study builds on previous work in the family and aims to test the traditional classification of the tropical woody taxa, which have been understudied and undersampled compared to their temperate relatives. ? Methods: A phylogenetic study was undertaken with expanded sampling of the tropical genera with data from five plastid markers (psbA-trnH, psaI-accD, matK, ndhF, and rbcL), nuclear ribosomal DNA (ITS) and morphology. ? Key results: Results support the placement of nine of 12 genera of the Triplarideae and Coccolobeae within Eriogonoideae, in which these genera form a paraphyletic assemblage giving rise to Eriogoneae. The remaining woody tropical genera excluded from Eriogonoideae occur in the paleotropics. ? Conclusions: Traditional characters used to delimit Coccolobeae and Triplarideae are not useful for defining monophyletic groups. The six-tepal condition is derived from the five-tepal condition, and unisexual flowers have arisen multiple times in different sexual systems. Ruminate endosperm has arisen multiple times in the family, suggesting this character is highly plastic.     

Phylogeny of the bee family Megachilidae (Hymenoptera: Apoidea) based on adult morphology

Phylogenetic relationships within the bee family Megachilidae are poorly understood. The monophyly of the subfamily Fideliinae is questionable, the relationships among the tribes and subtribes in the subfamily Megachilinae are unknown, and some extant genera cannot be placed with certainty at the tribal level. Using a cladistic analysis of adult external morphological characters, we explore the relationships of the eight tribes and two subtribes currently recognised in Megachilidae. Our dataset included 80% of the extant generic‐level diversity, representatives of all fossil taxa, and was analysed using parsimony. We employed 200 characters and selected 7 outgroups and 72 ingroup species of 60 genera, plus 7 species of 4 extinct genera from Baltic amber. Our analysis shows that Fideliinae and the tribes Anthidiini and Osmiini of Megachilinae are paraphyletic; it supports the monophyly of Megachilinae, including the extinct taxa, and the sister group relationship of Lithurgini to the remaining megachilines. The Sub‐Saharan genus Aspidosmia, a rare group with a mixture of osmiine and anthidiine features, is herein removed from Anthidiini and placed in its own tribe, Aspidosmiini, new tribe . Protolithurgini is the sister of Lithurgini, both placed herein in the subfamily Lithurginae; the other extinct taxa, Glyptapina and Ctenoplectrellina, are more basally related among Megachilinae than Osmiini, near Aspidosmia, and are herein treated at the tribal level. Noteriades, a genus presently in the Osmiini, is herein transferred to the Megachilini. Thus, we recognise four subfamilies (Fideliinae, Pararhophitinae, Lithurginae and Megachilinae) and nine tribes in Megachilidae. We briefly discuss the evolutionary history and biogeography of the family, present alternative classifications, and provide a revised key to the extant tribes of Megachilinae.     

Resolving robust phylogenetic relationships of core Brassicaceae using genome skimming data

The Brassicaceae is an economically and scientifically important family distributed globally, including oilseed rape and the model plant, Arabidopsis thaliana. Although growing molecular data have been used in phylogenetic studies, the relationships among major clades and tribes of Brassicaceae are still controversial. Here, we investigated the core Brassicaceae phylogenetics using 222 plastomes and 235 nrDNA cistrons, including 106 plastomes and 112 nrDNA cistrons assembled from newly sequenced genome skimming data of 112 taxa. The sampling covered 73 genera from 61.5% tribes and four unassigned genera and species. Three well supported lineages LI, LII, and LIII were revealed in our plastomic analyses, with LI sister to LII + LIII. In addition, the monophyly of the newly delimitated LII was strongly supported by three different partition strategies, concatenated methods under Bayesian and Maximum Likelihood analyses. LII comprised 13 tribes, including four tribes previously unassigned to any lineage, that is Biscutelleae as the earliest diverging clade and Cochlearieae as the sister to Megacarpaeeae + Anastaticeae. Within LII, the intertribal relationships were also well resolved, except that a conflicting position of Orychophragmus was detected among different datasets. In LIII, Shehbazia was resolved as a member of Chorisproreae, but Chorisproreae, Dontostemoneae, and Euclidieae were all resolved as paraphyletic, which was also confirmed by nrDNA analyses. Moreover, the loss of the rps16 gene was detected as likely to be a synapomorphy of the tribes Arabideae and Alysseae. Overall, using genome skimming data, we resolved robust phylogenetic relationships of core Brassicaceae and shed new light on the complex evolutionary history of this family.     

Molecular phylogeny of the sawfly subfamily Nematinae (Hymenoptera: Tenthredinidae)

Abstract.  Nematinae is one of the largest subfamilies in the sawfly family Tenthredinidae, but internal relationships are unknown in the absence of any formal phylogenetic analysis. To understand the internal phylogeny of Nematinae, we sequenced a portion of the mitochondrial cytochrome oxidase I gene and the nuclear elongation factor-1α gene from thirteen outgroup taxa and sixty-eight nematine species, the ingroup taxa of which represent all major genera and subgenera within the subfamily. Maximum parsimony and Bayesian phylogenetic analyses of the DNA sequence data show that: (1) Nematinae are monophyletic in a broad sense which includes Hoplocampa , Susana and the tribe Cladiini, which have been classified often into separate subfamilies; together with Craterocercus , these taxa form a paraphyletic basal grade with respect to the remaining Nematinae, but among-group relationships within the grade remain weakly resolved; (2) the remainder of the ingroup, Nematinae s. str, is monophyletic in all combined-data analyses; (3) within Nematinae s. str, the 'Higher' Nematinae is divided into three groups, Mesoneura and the large tribes Nematini and Pristiphorini; (4) although the traditional classifications at the tribal level are largely upheld, some of the largest tribes and genera are obviously para- or polyphyletic; (5) according to rate-smoothed phylogenies dated with two fossil calibration points, Nematinae originated 50–120 million years ago. In addition, the results from all Bayesian analyses provide strong and consistent support for the monophyly of Tenthredinidae, which has been difficult to demonstrate in previous parsimony analyses of morphological and molecular data.     

Contribution to the genome size knowledge of New World species from the Heliantheae alliance (Asteraceae)

This article contributes first genome size assessments by flow cytometry for 16 species, 12 genera, and 3 tribes from family Asteraceae, mostly belonging to the Heliantheae alliance, an assembly of 13 tribes from subfamily Asteroideae with a large majority of its species in the New World. Most genome sizes are accompanied by their own chromosome counts, confirming in most cases, although not all, previous counts for the species, and revealing possible cases of unknown dysploidy or polyploidy for certain taxa. The data contribute to the pool of knowledge on genome size and chromosome numbers in the family Asteraceae and will further allow deeper studies and a better understanding on the role of dysploidy in the evolution of the Heliantheae alliance. However, we still lack data for tribes Chaenactideae, Neurolaeneae, Polymnieae, and Feddeeae (the latter, monospecific) to complete the alliance representation.     

The taxonomy and systematics of Apocynaceae: where we stand in 2012

In their most recent classification of Apocynaceae in 2000, Endress and Bruyns recognized five subfamilies of Apocynaceae (Rauvolfioideae, Apocynoideae, Periplocoideae, Secamonoideae and Asclepiadoideae). Subsequently, through various studies using molecular data, it has been shown that most tribes and subtribes of Rauvolfioideae were not monophyletic, and new tribes and subtribes have been erected to reflect improved phylogenetic understanding of the family: Aspidospermeae in Rauvolfioideae; Nerieae, Odontadenieae and Baisseeae in Apocynoideae; Fockeeae in Asclepiadoideae; and Orthosiinae in Asclepiadeae. Several genera in Rauvolfioideae have been reassigned to different tribes in order to improve the monophyly of these tribes. The sister group of Asclepiadoideae plus Secamonoideae is not Periplocoideae, as formerly assumed, but tribe Baisseeae. Periplocoideae are nested in Apocynoideae. However, tribal composition remains unclear in some parts of the family. Clade structure in Apocynaceae is now generally well understood. The principal challenges now lie in identifying characters that can reflect and articulate these clades in a formal classification. Species‐rich, recent radiations such as core Asclepiadinae in Africa and the Metastematinae in Latin America present particular problems in this regard. © 2013 The Linnean Society of London     

Phylogeny and classification of the Catantopidae at the tribal level (Orthoptera, Acridoidea)

The grasshopper family Catantopidae is a well-known group, whose members include some of the most notorious agricultural pests. The existing classifications of the family are mostly utilitarian rather than being based on phylogenetic analysis and therefore unable to provide the stability desired for such an economically important group. In the present study, we present the first comprehensive phylogenetic analysis of the family based on morphology. By extensively sampling from the Chinese fauna, we included in the present analysis multiple representatives of each of the previously recognized tribes in the family. In total, we examined 94 genera represented by 240 species and evaluated 116 characters, including 84 for external morphology and 32 for male genitalia. The final matrix consists of 86 ingroup taxa and 88 characters. Our phylogenetic analyses resulted in a high resolution of the basal relationships of the family while showed considerable uncertainty about the relationships among some crown taxa. We further evaluated the usefulness of morphological characters in phylogeny reconstruction of the catantopids by examining character fit to the shortest trees found, and contrary to previous suggestions, our results suggest that genitalia characters are not as informative as external morphology in inferring higher-level relationship. We further suggest that earlier classification systems of grasshoppers in general and Catantopidae in particular most probably consist of many groups that are not natural due the heavy reliance on genitalia features and need to be revised in the light of future phylogenetic studies. Finally, we outlined a tentative classification scheme based on the results of our phylogenetic analysis.     

A revision of the higher taxonomy of the Afrotropical freshwater crabs (Decapoda: Brachyura) with a discussion of their biogeography

The higher taxonomy of the 20 known genera of Afrotropical freshwater crabs is revised to reflect the evolutionary relationships revealed by the consensus of a series of recent morphological and molecular phylogenetic studies of the group. The Afrotropical freshwater crab genera fall into two monophyletic groups, one from Socotra with two genera (Potamidae) and another that includes the remaining 18 genera. The latter group, which includes the bulk of the region's freshwater crab fauna, forms a well-supported monophyletic clade. We recognize two monophyletic sister groups (subfamilies) within the Potamonautidae, one for seven genera from Africa (the Potamonautinae) and one for 11 genera from Africa, the Seychelles, and Madagascar (the Deckeniinae). The Deckeniinae includes two monophyletic groups (tribes), one with seven genera from Madagascar (the Hydrothelphusini), and one with four genera from Africa and the Seychelles (the Deckeniini). The Deckeniini is further divided here into two subtribes, the Deckeniina and the Globonautina. The Platythelphusidae is not recognized, and the Deckeniidae and Globonautinae are lowered in rank. There is no phylogenetic support for the continued inclusion of any genus from the Afrotropical region in the Gecarcinucidae which is treated here as an exclusively Oriental family. The Afrotropical freshwater crabs (excluding those from Socotra) form a monophyletic assemblage that has no representatives outside of the region. The wider biogeographical implications of the taxonomic revision are discussed.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 399–413.     

Generic interrelationships within the Spionidae (Annelida: Polychaeta)

The phylogenetic relationships of spionid genera are estimated from parsimony analyses of morphological characters, with Trochochaetidae, Poecilochaetidae and Uncispionidae as outgroups. A first analysis of currendy recognised genera proved inconclusive and even exclusion of six of the most polymorphic genera resulted in 13 305 equally parsimonious trees and a fully collapsed consensus tree. A second analysis using only the type species of each genus, yielded four equally parsimonious trees; reduced to two after successive weighting. The topologies of these two trees indicated division of the family into four main groups: (1) Aonidella and Xandaros; (2) Prionospio (sensu fato)-complex, Laonice, Spiophanes and Aonides; (3) a large assemblage of genera, including Polydora-{senm late), Scolelepis, Malacoceros and Spio; (4) Atherospio, Pseudatherospio and Pygospiopsis. Earlier literature classifications of the group are evaluated and compared with die new results.     

Evolutionary relationships in the showy mistletoe family (Loranthaceae)

Loranthaceae (73 genera and ca. 900 species) comprise mostly aerial hemiparasitic plants. Three monotypic genera considered relicts are root parasites. The family is diverse in tropical areas, but representatives are also found in temperate habitats. Previous classifications were based on floral and inflorescence morphology, karyological information, and biogeography. The family has been divided into three tribes: Nuytsiae, Elytrantheae (subtribes Elytranthinae and Gaiadendrinae), and Lorantheae (subtribes Loranthinae and Psittacanthinae). Nuytsiae and Elytrantheae are characterized by a base chromosome number of x = 12, whereas subtribes Loranthinae (x = 9) and Psittacanthinae (x = 8) numbers are derived via aneuploid reduction. To elucidate the phylogeny of the family, we analyzed sequences from five genes (nuclear small and large subunit rDNA and the chloroplast genes rbcL, matK, and trnL-F) representing most genera using parsimony, likelihood, and Bayesian inference. The three root parasites, Nuytsia, Atkinsonia, and Gaiadendron, are supported as successive sister taxa to the remaining genera, resulting in a monophyletic group of aerial parasites. Three major clades are resolved each corresponding to a subtribe. However, two South American genera (Tristerix and Notanthera) and the New Zealand genus Tupeia, which were previously classified in subtribe Elytranthinae, are weakly supported as part of a clade representing the South American subtribe Psittacanthinae.     

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.     

Floral structure and evolution in the Anacardiaceae

WANNAN, B. S. & QUINN, C. J, 1991. Floral structure and evolution in the Anacardiaceae. Carpel morphology and anatomy is investigated in 17 genera and carpellode morphology in 12 genera. There is an evolutionary sequence in the family from poorly differentiated, nearly apocarpous gynoecia towards syncarpous gynoecia with clearly defined stigmata, styles and ovaries. There has also been marked reduction culminating in pseudomonomery. The carpellodes of the male flowers appear more conservative, and provide evidence of affinities between genera with quite different fertile gynoecia. The characters have been polarized using Burseraceae as a sister group. Data from these sources, as well as from pericarp anatomy, wood anatomy and biflavonoid content indicate that the long standing intrafamilial classification into five tribes is artificial, and that the two small satellite families, Blepharocaryaceae and Julianiaceae should be included in the family. A large monophyletic group is recognized comprised of essentially four of the existing tribes (Anacardiëae, Dobineëae, Semecarpeae, Rhoëae), as well as the two satellite families. This group incorporates two subgroups of more closely allied genera. The remaining genera (mostly Spondiadeae) are very diverse, and for the present are placed in an artificial group characterised by a set of plesiomorphs. Relationships within this group must be resolved before a satisfactory taxonomy of the family can be achieved.     

Phylogeny and classification of the Culicidae (Diptera)

The generic relationships and higher classification of the family Culicidae are examined on the basis of a phylogenetic analysis. New and traditional morphological characters studied and compared throughout the Culicidae resulted in the acquisition of character data relative to the majority of species within each genus. Polymorphisms and morphological observations are discussed and additional information and illustrations are provided for the majority of characters and their character states. The analysis of seventy-three adult, pupal and fourth-instar larval characters coded for the thirty-eight currently recognized genera of mosquitoes resulted in relationships and groupings which differ significantly from traditional hypotheses. The analysis supports the monophyly of the subfamily Anophelinae and the tribes Culicini and Sabethini. The Anophelinae form the most basal clade of the family. The results indicate that Aedini is a paraphyletic assemblage with respect to the Mansoniini, each of which is monophyletic in itself. The Aedini + Mansoniini form a sister group to the Culicini + Sabethini, with the Aedini and the Culicini placed in ancestral relationships to the Mansoniini and the Sabethini, respectively. Based on the topography of generic relationships among more 'generalized' mosquitoes, the boundaries and relationships of the tribes Aedeomyiini, Uranotaeniini, Ficalbiini, Hodgesiini, Orthopodomyiini and Culisetini appear to be problematic. Relationships between genera of the tribe Aedini are generally poorly resolved due to a significant amount of polymorphism, especially within the genus Aedes as currently defined. There is no support for separate subfamily recognition for the genus Toxorhynchites , which is downgraded to tribal status as a result of the analysis. The results are discussed in relation to previous hypotheses based on subjective inference and cladistic analyses.     

A molecular phylogeny of Panicum (Poaceae: Paniceae): tests of monophyly and phylogenetic placement within the Panicoideae

Panicum L. is a cosmopolitan genus with approximately 450 species. Although the genus has been considerably reduced in species number with the segregation of many taxa to independent genera in the last two centuries, Panicum remains a heterogeneous assemblage, as has been demonstrated in recent years. The genus is remarkably uniform in its floral characters but exhibits considerable variation in anatomical, physiological, and cytological features. As a result, several classifications, and criteria of what the genus should really include, have been postulated in modern literature. The purpose of this research, based on molecular data of the chloroplast ndhF gene, is to test the monophyly of Panicum, to evaluate infrageneric classifications, and to propose a robust phylogenetic hypothesis. Based on the present results, previous morphological and molecular phylogenetic studies, and inferred diagnostic morphological characters, we restrict Panicum sensu stricto (s.s.) to the former subgenus Panicum and support recognition of Dichanthelium, Phanopyrum, and Steinchisma as distinct genera. We have transfered other species of Panicum to other genera of the Paniceae. Most of the necessary combinations have been made previously, so few nomenclatural changes have been required. The remaining species of Panicum sensu lato (s.l.) are included within Panicum incertae sedis representing isolated species or species grouped within monophyletic clades. Additionally, we explore the performance of the three codon position characters in producing the supported phylogeny.     

Higher-level phylogeny and morphological evolution of tyrant flycatchers, cotingas, manakins, and their allies (Aves: Tyrannida)

Despite increased understanding of higher-level relationships in passerine birds in the last 15 years, the taxonomic boundaries and phylogenetic interrelationships of the major groups of the Tyrannida (including the cotingas, manakins, tityrines, and tyrant flycatchers) remain unclear. Here, we present an analysis of DNA sequence data obtained from two nuclear exons, three introns, and one mitochondrial gene for 26 genera of Tyrannida and 6 tracheophone outgroups. The analysis resulted in well-supported hypotheses about the earliest evolution within Tyrannida. The Cotingidae, Pipridae, Tityrinae (sensu) [Prum, R.O., Rice, N.H., Mobley, J.A., Dimmick, W.W., 2000. A preliminary phylogenetic hypothesis for the cotingas (Cotingidae) based on mitochondrial DNA. Auk 117, 236-241], Tyrannidae, and the tyrannid subfamiles Tyranninae and Pipromorphinae (sensu) [Sibley, C.G., Monroe, B. L. Jr., 1990. Distribution and Taxonomy of Birds of the World. Yale University Press, New Haven, CT] were all found to be reciprocally monophyletic (given the present taxon sampling). The Cotingidae and Pipridae form a clade that is the sister group to a well-supported clade including Oxyruncus, the Tityrinae, Piprites, and the Tyrannidae. Oxyruncus is the sister group to the Tityrinae, and Piprites is placed as the sister group to the Tyrannidae. The tyrannid subfamilies Tyranninae and Pipromorphinae are monophyletic sister taxa, but the relationships of Platyrinchus mystaceus to these two clades remains ambiguous. The presence of medial (=internal) cartilages in the syrinx is a synapomorphy for the Oxyruncus-Tityrinae-Piprites-Tyrannidae clade. Although morphological synapomorphies currently support the monophyly of both the Pipridae and the Cotingidae, convergences and/or reversals in morphological character states are common in Tyrannida. The relationship between Oxyruncus and the Tityrinae is congruent with additional syringeal synapomorphies and allozyme distance data. Accordingly, we propose the recognition the family Tityridae within the Tyrannida to include the genera Schiffornis, Laniisoma, Laniocera, Iodopleura, Xenopsaris, Pachyramphus, Tityra, and Oxyruncus.