Molecular phylogeny of the plant bugs (Heteroptera: Miridae) and the evolution of feeding habits

The first comprehensive cladistic analysis of Miridae, the plant bugs, is presented based on analysis of 3935 base pairs of mitochondrial (16S, COI) and nuclear (18S, 28SD3) DNA for 91 taxa in seven subfamilies. Data were analysed using maximum likelihood (ML), parsimony and Bayesian inference (BI) phylogenetic frameworks. The phylogenetic results are compared with previous hypotheses of higher relationships in the family using alternative hypothesis tests. A Bayesian relaxed molecular clock is used to examine divergence times, and ancestral feeding habits are reconstructed using parsimony and a Bayesian approach. Clades recovered in all analyses are as follows: Cimicomorpha, Miroidea and Miridae; Bryocorinae: Bryocorini; Stenodemini; Mirinae; Deraeocorinae (Clevinemini + Deraeocorini); Cylapinae; Isometopinae; Bryocorinae: Dicyphini; Orthotylini; Phylinae (Phylini + Pilophorini), and Phylinae as sister group to all the remaining mirid taxa. These results are largely congruent with former hypotheses based on morphological data with respect to the monophyly of various subfamilies and tribes; however, our results indicate that the subfamily Bryocorinae is not monophyletic, as the two tribes, Dicypini and Bryocorini, were separated in the phylogenetic results. Divergence time estimates indicate that the radiation of the Miridae began in the Permian; most genus‐level radiations within subfamilies began in the late Cretaceous, probably in response to the angiosperm radiation. Ancestral feeding state reconstructions based on Bayesian and parsimony inference were largely congruent and both reconstructed phytophagy as the ancestral state of the Miridae. Furthermore, the feeding habits of the common ancestors of Mirinae + Deraeocorinae, Bryocorinae + Cylapinae + Isometopinae + Orthotylinae, and the remaining taxa excluding Phylinae, were inferred as phytophagous. Therefore, at least three shifts from phytophagy or polyphagy to predation occurred within the Miridae. Additionally, based on the mirid host‐plant records, we discovered several trends, such as a strong relationship between host‐plant ranges and a facultative feeding habit. © The Willi Hennig Society 2011.     

Larval morphology and biology of oxycorynine weevils and the higher phylogeny of Belidae (Coleoptera, Curculionoidea)

Phylogenetic relationships among members of the family Belidae (Curculionoidea) were reconstructed through cladistic analysis using 58 characters and 17 terminals. The characters were from larval morphology (30), adult morphology (25) and biology regarding larval host-plants and feeding habits (three). They were scored for exemplar taxa in 17 genera, representing different belid subfamilies and tribes, plus two outgroup taxa in Megalopodidae and Nemonychidae. The sampled genera included all those for which larval and adult information is available, and two known only from adults. New information on the larvae and biology of two oxycorynines is provided. These are the Chilean Oxycraspedus cribricollis , whose larvae live in decayed female strobili of the gymnosperm Araucaria araucana , and Hydnorobius hydnorae from Argentina, whose larvae, described and illustrated in the present paper, develop inside the flower and fruit bodies of Prosopanche americana (Hydnoraceae), a root-parasitic angiosperm. The relationships proposed by the single optimal cladogram resulting from simultaneous analysis of all taxa and characters are recovered by one of three optimal cladograms based on the larval data set alone. The cladogram justifies a revised classification of Belidae in two sister subfamilies: Belinae (with tribes Pachyurini, Agnesiotidini and Belini) and Oxycoryninae (with tribes Oxycorynini and Aglycyderini). It summarizes larval and adult synapomorphies defining the family Belidae, subfamilies and tribes. Based on the phylogenetic tree, the evolution of biological traits is traced. Larval development in vegetative organs of conifers is ancestral in Belidae. A shift to reproductive structures characterizes the Oxycorynini, a habit which was conserved while several shifts to distantly related host-plant groups occurred.     

Phylogenetic structure in the grass family (Poaceae) as inferred from chloroplast DNA restriction site variation

A cladistic analysis of chloroplast DNA restriction site variation among representatives of all subfamilies of the grass family (Poaceae), using Joinvillea (Joinvilleaceae) as the outgroup, placed most genera into two major clades. The first of these groups corresponds to a broadly circumscribed subfamily Pooideae that includes all sampled representatives of Ampelodesmeae, Aveneae, Brachypodieae, Bromeae, Diarrheneae, Meliceae, Poeae, Stipeae, and Triticeae. The second major clade includes all sampled representatives of four subfamilies (Panicoideae [tribes Andropogoneae and Paniceae], Arundinoideae [Arundineae], Chloridoideae [Eragrostideae], and Centothecoideae [Centotheceae]). Within this group (the “PACC” clade), the Panicoideae are resolved as monophyletic and as the sister group of the clade that comprises the other three subfamilies. Within the latter group, Danthonia (Arundinoideae) and Eragroslis (Chloridoideae) are resolved as a stable monophyletic group that excludes Phragmites (Arundinoideae); this structure is inconsistent with the Arundinoideae being monophyletic as currently circumscribed. The PACC clade is placed within a more inclusive though unstable clade that includes the woody Bambusoideae (Bambuseae) plus several disparate tribes of herbaceous grasses of uncertain affinity that are often recognized as herbaceous Bambusoideae (Brachyelytreae, Nardeae, Olyreae, Oryzeae, and Phareae). Among eight most-parsimonious trees resolved by the analysis, four include a monophyletic Bambusoideae sensu lato (comprising Bambuseae and all five of these herbaceous tribes) as the sister group of the PACC clade; in the other four trees these bambusoid elements are not resolved as monophyletic, and the PACC clade is nested among these tribes. These results are consistent with those of previous analyses that resolve a basal or near-basal branch within the family between Pooideae and all other grasses. However, resolution by the present analysis of the PACC clade, which includes Centothecoideae, Chloridoideae, and Panicoideae, but excludes Bambusoideae, is inconsistent with the results of previous analyses that place Bambusoideae and Panicoideae in a monophyletic group that excludes Centothecoideae and Chloridoideae.     

A review of the phylogeny and classification of the Asteraceae

The Asteraceae are commonly divided into two large subfamilies, the Cichorioideae (syn. Lactucoideae; Mutisieae, Cardueae, Lactuceae, Vernonieae, Liabeae, Arctoteae) and the Asteroideae (Inuleae, Astereae, Anthemideae, Senecioneae, Calenduleae, Heliantheae, Eupatorieae). Recent phylogenetic analyses based on morphological and chloroplast DNA data conclusively show that the Mutisieae-Barnadesiinae are the sister group to the rest of the family and that the Asteroideae tribes form a monophyletic group. The Vernonieae and Liabeae are sister tribes and the Eupatorieae are nested within a paraphyletic Heliantheae; otherwise tribal interrelationships are still largely uncertain. The Mutisieae-Barnadesiinae are excluded from the Mutisieae and elevated to the new subfamily Barnadesioideae. The two subfamilies Barnadesioideae and Asteroideae are monophyletic, whereas the status of the Cichorioideae remains uncertain. Analyses of chloroplast DNA data support the monophyly of the Cichorioideae; however, morphological data indicate that the subfamily is paraphyletic. Further studies are needed to test the monophyly of the Cichorioideae, as well as to further resolve tribal interrelationships in the two larger subfamilies.     

Evolution of the leucine gene cluster in Buchnera aphidicola: insights from chromosomal versions of the cluster

In Buchnera aphidicola strains associated with the aphid subfamilies Thelaxinae, Lachninae, Pterocommatinae, and Aphidinae, the four leucine genes (leuA, -B, -C, and -D) are located on a plasmid. However, these genes are located on the main chromosome in B. aphidicola strains associated with the subfamilies Pemphiginae and Chaitophorinae. The sequence of the chromosomal fragment containing the leucine cluster and flanking genes has different positions in the chromosome in B. aphidicola strains associated with three tribes of the subfamily Pemphiginae and one tribe of the subfamily Chaitophorinae. Due to the extreme gene order conservation of the B. aphidicola genomes, the variability in the position of the leucine cluster in the chromosome may be interpreted as resulting from independent insertions from an ancestral plasmid-borne leucine gene. These findings do not support a chromosomal origin for the leucine genes in the ancestral B. aphidicola and do support a back transfer evolutionary scenario from a plasmid to the main chromosome.     

Critical Reexamination of Palynological Characters Used to Delimit Asclepiadaceae in Comparison to the Molecular Phylogeny Obtained from PlastidmatK Sequences

The family Asclepiadaceae (Dicotyledones) was created by Brown in 1810 by splitting in two the family Apocynaceae of Jussieu established in 1789. The morphological characters used to make this distinction were mainly palynological, such as presence of tetrads or pollinia and number and orientation of pollinia. Those characters, still used in higher taxonomic delimitation (families, subfamilies, and tribes), are here critically reexamined and compared to a molecular phylogeny obtained with one of the more variable plastid genes (matK) of 46 species in the order Gentianales. In this molecular phylogeny, Asclepiadaceae form a monophyletic group derived from within Apocynaceae. Each of the subfamilies of Asclepiadaceae is monophyletic and based on reliable palynological characters, but palynological characters are not useful to delimit tribes of the subfamily Asclepiadoideae. Based on the molecular data, these tribes have undergone parallelisms in several reproductive traits.     

STUDIES ON THE PHYLOGENY AND HOST PLANT ASSOCIATION OF LACHNIDAE (HOMOPTERA: APHIDINEA)*

Abstract With reference to the material related to the evolution of host-plant associations, phylogenetic analyses of aphids in Lachnidae are carried out by using 20 taxa and 26 characters. The monophyly of Traminae and Cinarinae are recognized. However, Lachninae is proved to be a polyphyletic group. Three traditional subfamilies, Cinarinae, Lachninae and Traminae are still retained. Three tribes within Cinarinae which include Cinarini, Eulachnini and Schizolachnini are suggested. It is concluded that the ancestral host plants of Lachnidae were deciduous plants rather than coniferous trees or other plants.     

The phylogenetic relationships and natural classificition of the Vespoidea (Hymenptera)

Abstract. Phylogenetic relationships of the subfamilies and tribes of the Vespoidea (= Diploptera) are investigated using cladistic methods. A natural classification is proposed, sequencing seven tribes in six subfamilies within the single family Vespidae. Euparagia is the sister-group of the rest of the Vespidae. The Gayellinae and Masarinae are sister-groups, and are reduced in rank to tribes within the subfamily Masarinae. This is the sister-group of the Eumeninae + Stenogastrinae + Polistinae + Vespinae. The Zethinae is a paraphyletic group; both it and the Raphiglossinae are deleted and included in the Eumeninae. The Stenogastrinae is regarded as the sister-group of the Vespinae + Polistinae.     

Phylogeny,classification and biogeography of bombyliine bee flies (Diptera,Bombyliidae)

The Bombyliinae comprises over 1100 described species in 73 known genera distributed worldwide. It is one of the largest subfamilies of bee flies (Diptera: Bombyliidae). We present the first phylogenetic hypothesis for this subfamily, based on 157 adult morphological characters scored for 123 species representing 60 genera, including all the tribes of Bombyliinae, and the related subfamilies Lordotinae and Toxophorinae. Four most parsimonious trees were generated from our analysis under equal weighting schemes. The monophyly of Bombyliinae is supported, and Lordotinae is sister to the Bombyliinae. Within Bombyliinae, Conophorini is sister to the remaining tribes. Five previously recognized tribes are revised and four new tribes are erected. We placed almost all genera in our tribal classification, based on our phylogenetic results and available character evidence. The genus Parabombylius is proposed as a synonym of Bombylius. The Gondwanan origin for the major lineages of Bombyliinae is strongly indicated by our biogeographic analysis which reconstructs ancestral areas. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub: 1EC5C827‐34D5‐4A95‐BA78‐4ACF457F6D40.     

A phylogeny of the chloroplast gene rbcL in the Leguminosae: taxonomic correlations and insights into the evolution of nodulation

Phylogenetic analysis of the chloroplast-encoded rbcL gene in Leguminosae are consistent with previous hypotheses in suggesting that the family as a whole is monophyletic, but that only two of its three subfamilies are natural. The earliest dichotomies in the family appear to have involved tribes Cercideae or Cassieae (subtribe Dialiinae), followed by Detarieae/ Macrolobieae, all of which are members of subfamily Caesalpinioideae. The remainder of the family is divided into two clades: (1) Mimosoideae and the caesalpinioid tribes Caeasalpinieae and Cassieae (subtribes Ceratoniinae and Cassiinae); (2) Papilionoideae. Basal groups within Papilionoideae are, as expected, elements of the grade tribes Sophoreae and Swartzieae. Major clades within Papilionoideae include: (1) a Genistoid Alliance comprising Genisteae, Crotalarieae, Podalyrieae, Thermopsideae, Euchresteae, and also some Sophoreae; (2) a clade marked by the absence of one copy of the chloroplast inverted repeat, with which are associated Robinieae. Loteae, and some Sophoreae; (3) Phaseoleae, Desmodieae. Psoraleeae, and most Millettieae, a group also marked by presence of pseudoracemose inflorescences; and (4) a well-supported clade comprising Aeschynomeneae, Adesmieae, and some Dalbergieae. Nodulation is most parsimoniously optimized on the rbcL strict consensus tree as three parallel gains, occurring in Papilionoideae, the caesalpioioid ancestors of Mimosoideae, and in the genus Chamaecrista (Caesalpinieae: Cassieae).     

A classification of the Monimiaceae

Thirty-two genera of the Monimiaceae are assigned to six subfamilies and eight tribes. Two further genera are insufficiently known to be placed. One subfamily (Glossocalycoideae Thorne, two tribes (Hennecartieae Philipson, Palmerieae Philipson) and one genus ( Laureliopsis Schodde) are described as new.     

Phylogeny and classification of Leptophlebiidae (Ephemeroptera) with an emphasis on Neotropical fauna

Mayflies from the family Leptophlebiidae are cosmopolitan and highly diverse morphologically; they are also the largest family in numbers of genera and the second in number of species in the order Ephemeroptera. In spite of their broad diversity and the efforts employed to understand the evolution of this group, the internal classification of Leptophlebiidae remains controversial at all levels. More recently, important changes have been incorporated into the systematics of the family, increasing the number of subfamilies (from two to six) and recognizing several tribes. We present a phylogeny of the family based on 153 taxa (53 genera) and two molecular markers, representing 1655 bp, and verify the taxonomic status of the subfamilies, tribes and complexes. Based on these results, the number of subfamilies has been increased from six to eight and one new tribes and two new subtribes have been added. In addition, new ranks are proposed and the concept of Atalophlebiinae revised, including genera with distributions in the Australasian and Neotropical regions.     

Untangling the assassin's web: Phylogeny and classification of the spider-associated Emesine complex (Hemiptera: Reduviidae)

Web-building spiders are formidable predators, yet assassin bugs in the Emesine Complex (Hemiptera: Reduviidae: Emesinae, Saicinae, and Visayanocorinae) prey on spiders. The Emesine Complex comprises >1000 species and these web-associated predatory strategies may have driven their diversification. However, lack of natural history data and a robust phylogenetic framework currently preclude tests of this hypothesis. We combine Sanger (207 taxa, 3865 bp) and high-throughput sequencing data (15 taxa, 381 loci) to generate the first taxon- and data-rich phylogeny for this group. We discover rampant paraphyly among subfamilies and tribes, necessitating revisions to the classification. We use ancestral character state reconstructions for 40 morphological characters to identify diagnostic features for a revised classification. Our new classification treats Saicinae Stål and Visayanocorinae Miller as junior synonyms of Emesinae Amyot and Serville, synonymizes the emesine tribes Ploiariolini Van Duzee and Metapterini Stål with Emesini Amyot and Serville, and recognises six tribes within Emesinae (Collartidini Wygodzinsky, Emesini, Leistarchini Stål, Oncerotrachelini trib.n. , Saicini Stål stat.n. , and Visayanocorini Miller stat.n. ). We show that a pretarsal structure putatively involved in web-associated behaviours evolved in the last common ancestor of Emesini, the most species-rich clade within Emesinae, suggesting that web-associations could be widespread in Emesinae.     

Phylogeny and evolution of Mesozoic and extant lineages of Histeridae (Coleoptera), with discovery of a new subfamily Antigracilinae from the Lower Cretaceous

In order to place a newly discovered species Antigracilus costatus gen. sp. n. from the Lower Cretaceous Yixian Formation (China) and to assess previously unplaced fossil taxa, we investigated the relationships of extant and extinct lineages of Histeridae based on three data sets: (i) 69 morphological characters belonging to 48 taxa (representing all 11 subfamilies and 15 of 17 tribes of modern Histeridae); (ii) partitioned alignment of 6030 bp from downloaded nucleotide sequences (28S, CAD, COI, 18S) of 50 taxa (representing 10 subfamilies and 15 of 17 tribes of modern Histeridae); and (iii) a combined morphological and molecular dataset for 75 taxa. Phylogenetic analyses of the morphology and combined matrices recovered the new Lower Cretaceous taxon as a sister group to remaining Histeridae and it is placed in †Antigracilinae subfam. n. †Antigracilinae constitutes the earliest record of Histeridae from the Lower Cretaceous Yixian Formation (∼125 Myr), backdating the minimum age of the family by 25 Myr from the earliest Cenomanian (~99 Myr) to the Barremian of the Cretaceous Period. Our molecular phylogeny supports Histeridae to be divided into seven different clades, with currently recognised subfamilies Abraeinae (sensu lato), Saprininae, Chlamydopsinae, and Histerinae (sensu lato) recovered as monophyletic, while Dendrophilinae, Onthophilinae, and Tribalinae are polyphyletic taxa. The Burmese amber species †Pantostictus burmanicus Poinar & Brown is placed as a sister group to the tribe Plegaderini (Abraeinae) and was assigned as a new tribe Pantostictini trib. n. Both molecular and combined phylogenies recovered the subfamilies Trypanaeinae and Trypeticinae deeply within the subfamily Abraeinae (sensu lato), and they are downgraded into Trypanaeini stat. n. and Trypeticini stat. n.     

Morphology-based phylogeny of the treehopper family Membracidae (Hemiptera: Cicadomorpha: Membracoidea)

A parsimony‐based phylogenetic analysis of eighty‐three morphological characters of adults and immatures of seventy representatives of the tribes and subfamilies of Membracidae and two outgroup taxa was conducted to evaluate the status and relationships of these taxa. Centrotinae apparently gave rise to Nessorhinini and Oxyrhachini (both formerly treated as subfamilies, now syn.n. and syn.reinst., respectively, of Centrotinae). In contrast to previous analyses, a clade comprising Nicomiinae, Centronodinae, Centrodontinae, and the unplaced genera Holdgatiella Evans, Euwalkeria Goding and Antillotolania Ramos was recovered, but relationships within this clade were not well resolved. Nodonica bispinigera, gen.n. and sp.n., is described and placed in Centrodontini based on its sister‐group relationship to a clade comprising previously described genera of this tribe. Membracinae and Heteronotinae were consistently monophyletic. Neither Darninae nor Smiliinae, as previously defined, was monophyletic on the maximally parsimonious cladograms, but constraining both as monophyletic groups required only one additional step. The monophyly of Stegaspidinae, including Deiroderes Ramos (unplaced in Membracidae), was supported on some but not all equally parsimonious cladograms. More detailed analyses of individual subfamilies, as well as morphological data on the undescribed immatures of several membracid tribes and genera, will be needed to elucidate relationships among tribes and genera. A key to the subfamilies and tribes is provided.     

New fossil paussids from Dominican amber with notes on the phylogenetic systematics of the paussine complex (Coleoptera: Carabidae)

The first fossil representative of the Protopaussinae and the second fossil Eohomopterus (Paussinae) are described from amber inclusions of Tertiary age collected in the Dominican Republic on the West Indian island of Hispaniola ( Protopaussus pristinus sp.n., Eohomopterus poinari sp.n.). These finds increase the total number of paussids present in Dominican amber to four species in three genera ( Protopaussus , Eohomopterus , Homopterus ). A key is provided for the identification of the New World genera of Protopaussinae and Paussinae and for the species of Eohomopterus . Based on synapomorphic character states, the paussid beetles are split into two subfamilies, the monobasic Protopaussinae and the more inclusive Paussinae (the twenty-three genera are placed into two supertribes: the Carabidomemnitae comprising two tribes and the Paussitae comprising eight tribes. The extant congeners of Eohomopterus and Homopterus live in the Neotropical Region, while the extant members of Protopaussus are known only from the Oriental Region. This record of New World Tertiary Protopaussus adds another example of the Old World relationships of taxa from Dominican amber. The temporal and geographical distribution of Protopaussus indicates that its present zoogeographical range may represent the relict of a wide Laurasian distribution in the past. It does not support the previously suspected Oriental centre of origin for the ancestral stock of {Protopaussinae + Paussinae}.     

Morphological characters add support for some members of the basal grade of Asteraceae

Recent molecular studies in Asteraceae have divided tribe Mutisieae (sensu Cabrera) into 13 tribes and eight subfamilies. Each of the major clades is well supported but the relationships among them are not always clear. Some of the new taxa are easily characterized by morphological data but others are not, chief among the latter being three subfamilies (Stifftioideae, Wunderlichioideae and Gochnatioideae) and the tribe Hyalideae. To understand evolution in the family it is critical to investigate potential morphological characters that can help to evaluate the basal lineages of the Asteraceae. The data for this study were taken from 52 species in 24 genera representing the basal groups in the family. Many characters were examined but most of the useful ones were from reproductive structures. Several apomorphies supported a few of the clades. For instance, members of subfamily Wunderlichioideae (Hyalideae and Wunderlichieae) share predominantly ten‐ribbed achenes and members of Wunderlichioideae + Stifftioideae share two synapomorphies: 100–150 (200) pappus elements, arranged in (three) four or five series. These apomorphies can be viewed as an indication of a sister‐group relationship between the two subfamilies as the placement of Stifftieae was not well resolved by the molecular data. Members of Wunderlichieae are characterized by having a paleaceous receptacle, style branches that are strongly papillose above and below the bifurcation, and a pappus of scales. Hyalis and Ianthopappus (Hyalideae) share venation type and an apiculate anther appendage but these are also found in Gochnatieae. Other clades have fewer supporting characters. These characters are just a beginning. Cladograms with morphology characters plotted, illustrations and a key to the basal grade of Asteraceae are provided. © 2013 The Linnean Society of London     

Systematic revision of Entomobryidae (Collembola) by integrating molecular and new morphological evidence

Entomobryidae, the largest collembolan family, is traditionally classified at suprageneric level using a limited set of morphological structures, such as scales, antennal segmentation. Most tribal and subfamilial delimitations appear, however, disputable in the light of recent works. Integrating molecular and morphological evidence, we propose here a revision of the systematics of the family. In addition to traditional taxonomic characters, tergal specialized chaetae (S‐chaetae) are newly introduced, and their patterns are shown to be diversified at all levels from species to subfamilies. S‐chaetotaxic pattern on phylogenetic tree shows that evolution of S‐chaetae is not parallel between the different terga and that their patterns coincide well with the known molecular phylogeny, providing a powerful tool for the systematics of Entomobryidae. Orchesellinae sensu Soto‐Adames et al. (Annals of the Entomological Society of America, 101, 2008, 501); is divided into three subfamilies: Orchesellinae s. s., Bessoniellinae and Heteromurinae, the latter two upgraded from the original tribal level. Entomobryinae sensu Szeptycki (Morpho‐Systematic Studies on Collembola. IV. Chaetotaxy of the Entomobryidae and its Phylogenetical Significance, 1979), is no longer divided into scaled and unscaled tribes, and Lepidosira‐group is transferred from Seirinae to Entomobryinae. A key to subfamilies and tribes and a comparison with previous classifications of the Entomobryidae are provided. This study greatly improves the understanding of primary and secondary characters and erects the fundamental framework for the taxonomy of Entomobryidae.     

A molecular phylogeny of fleas (Insecta: Siphonaptera): origins and host associations

Siphonaptera (fleas) is a highly specialized order of holometabolous insects comprising ～2500 species placed in 16 families. Despite a long history of extensive work on flea classification and biology, phylogenetic relationships among fleas are virtually unknown. We present the first formal analysis of flea relationships based on a molecular matrix of four loci (18S ribosomal DNA, 28S ribosomal DNA, Cytochrome Oxidase II, and Elongation Factor 1‐alpha) for 128 flea taxa from around the world representing 16 families, 25 subfamilies, 26 tribes, and 83 flea genera with eight outgroups. Trees were reconstructed using direct optimization and maximum likelihood techniques. Our analysis supports Tungidae as the most basal flea lineage, sister group to the remainder of the extant fleas. Pygiopsyllomorpha is monophyletic, as are the constituent families Lycopsyllidae, Pygiopsyllidae, and Stivaliidae, with a sister group relationship between the latter two families. Macropsyllidae is resolved as sister group to Coptopsyllidae with moderate nodal support. Stephanociricidae is monophyletic, as are the two constituent subfamilies Stephanocircinae and Craneopsyllinae. Vermipsyllidae is placed as sister group to Jordanopsylla. Rhopalopsyllidae is monophyletic as are the two constituent subfamilies Rhopalopsyllinae and Parapsyllinae. Hystrichopsyllidae is paraphyletic with Hystrichopsyllini placed as sister to some species of Anomiopsyllini and Ctenopariini placed as sister to Carterettini. Ctenophthalmidae is grossly paraphyletic with the family broken into seven lineages dispersed on the tree. Most notably, Anomiopsyllini is paraphyletic. Pulicidae and Chimaeropsyllidae are both monophyletic and these families are sister groups. Ceratophyllomorpha is monophyletic and includes Ischnopsyllidae, Ceratophyllidae, and Leptopsyllidae. Leptopsyllidae is paraphyletic as are its constituent subfamilies Amphipsyllinae and Leptopsyllinae and the tribes Amphipsyllini and Leptopsyllini. Ischnopsyllidae is monophyletic. Ceratophyllidae is monophyletic, with a monophyletic Dactypsyllinae nested within Ceratophyllinae, rendering the latter group paraphyletic. Mapping of general host associations on our topology reveals an early association with mammals with four independent shifts to birds. © The Willi Hennig Society 2008.