Taxonomy and phylogeny of the tribePlucheeae (Asteraceae)

The tribePlucheeae (Benth.)A. Anderb., has been analysed cladistically by means of a computerized parsimony program (Hennig 86), using theArctotideae as outgroup. The results of the analysis are presented in a consensus tree and one cladogram. Four major monophyletic subgroups can be recognized: TheColeocoma group (3 genera), thePterocaulon group (3 genera), theLaggera group (6 genera), and thePluchea group (12 genera). All recognized genera are described and most genera are supplied with taxonomical notes including comments on their taxonomic status. Genera such asBlumea, Pluchea, andEpaltes are demonstrated to be unnatural assemblages.Monarrhenus andTessaria are both closely related to thePluchea complex. The old generic nameLitogyne Harv. has been taken up for one species ofEpaltes, the genusRhodogeron is reduced to a synonym ofSachsia, and the following new combinations are made;Litogyne gariepina (DC.)A. Anderb., andSachsia coronopifolia (Griseb.)A. Anderb.     

INDIAN OCEAN NANOPLANKTON. I. RHABDOSPHAERACEAE (PRYMNESIOPHYCEAE) WITH A REVIEW OF EXTANT TAXA1

Family characteristics of the Rhabdosphaeraceae are revised to limit the genera to those having cyrtoliths; genera with placoliths are removed from the family. Rhabdoliths, cyrtoliths bearing a process in the central area, are present in all genera. Coccospheres having monomorphic coccoliths, all being rhabdoliths, form one group within the family, whereas genera with dimorphic coccoliths in the coccosphere comprise a second group. Cyrtoliths without processes in the latter group may be intermixed with rhabdoliths in some genera, whereas other genera have rhabdoliths located only in polar regions of the coccosphere. Two generic nomenclatural changes are proposed, Algirosphaera being the name applied to species previously placed in Anthosphaera, an invalid generic name, and Palusphaera is recognized as a valid monotypic genus, P. vandeli being the name applied to the species that has been named Rhabdosphaera longistylis in recent literature. A new combination is made, Rhabdosphaera xiphos (Deflandre et Fert) comb. nov., recognizing a species formerly known only in sediments as extant. Rhabdosphaera, Acanthoica and Algirosphaera are genera with dimorphic coccoliths in the coccospheres; Discosphaera, Palusphaera and Anacanthoica are genera having monomorphic coccoliths in the coccosphere.     

Molecular phylogeny of fungus gnats (Diptera: Mycetophilidae) revisited: position of Manotinae,Metanepsiini, and other enigmatic taxa as inferred from multigene analysis

The phylogeny of selected genera from four subfamilies of fungus gnats (Diptera: Mycetophilidae) – Manotinae, Leiinae, Sciophilinae and Gnoristinae (including Metanepsiini) – is reconstructed based on the combined analysis of five mitochondrial (12S, 16S, COI, COII, cytB) and two nuclear (28S, ITS2) gene markers. Results of the different analyses all support Manotinae as a monophyletic group, with Leiinae as the sister group. Allactoneura DeMeijere is nested in the monophyletic and strongly supported clade of Leiinae. The tribe Metanepsiini is revealed as paraphyletic and the genera Metanepsia Edwards and Chalastonepsia Søli do not appear to be closely related. The genera Docosia Winnertz, Ectrepesthoneura Enderlein, Novakia Strobl and Syntemna Winnertz were placed with a group of genera included traditionally in the Gnoristinae. The monophyly of Dziedzickia Johannsen and Phthinia Winnertz is not supported. The genera of Sciophilinae (excluding Paratinia Mik but including Eudicrana Loew) form a monophyletic group in the Bayesian model.     

Floral morphology and taxonomic relations among the genera of Amaranthaceae in the New World and the Hawaiian Islands

Twenty-three genera of Amaranthaceae occur in the New World. Two endemic genera occur in the Hawaiian Islands. Among the genera of the subfamily Amaranthoideae, Celosia, Cyathula and Achyranthes have their main distributions in the Old World; the two last-named genera are represented in the Americas only by widespread weeds. All the New World genera of the subfamily Gomphrenoideae are mainly or entirely restricted to this region. Characters of androecium and gynoecium are fundamental in the recognition of genera within the family. Androecia of different genera may be structurally and phylogenetically more similar than would appear from a cursory examination. It is suggested that the type of staminal tubes found in Pseudogomphrena and Froelichia can be derived from that in Alternanthera and Froelichiella by reduction of filament length and a fusion of pseudostaminodia with the filaments. The staminal tube in Gomphrena could result from a further decrease in distance between pseudostaminodia of the Pseudogomphrena type, and a deeper forking of the pseudostaminodia; each so-called apical filament lobe in Gomphrena would then be homologous with half a pseudostaminodium in Pseudogomphrena. Much of the variation in the androecia of these and other genera, as well as within genera such as Pfaffia, can be explained as the combined results of coalescence and splitting-up tendencies. Splitting up of staminal tubes may not necessarily take place along the borders of phylogenetically original filaments and pseudostaminodia. The Amaranthus-type of pollen is found in the majority of genera of the subfamily Amaranthoideae, but also in the Chenopodiaceae. A group of genera within the subfamily Gomphrenoideae also has pollen very similar to, or identical with, this type. Most genera of the subfamily Gomphrenoideae have pollen of the Gomphrena-type. Pseudoplantago has unilocular (at anthesis) anthers, a characteristic of the subfamily Gomphrenoideae, but floral structure as well as pollen morphology connect the genus to a group of genera within the Amaranthoideae, subtribe Achyranthinae. The combination of subcuboidal shape and opercula with radially arranged hooked protuberances, makes the pollen of Pseudoplantago unique among the angiosperms studied so far. Floral morphology and palynological characteristics indicate a close relationship between Pfaffia and Alternanthera. Both genera, as currently accepted, are relatively homogeneous from pollen morphological points of view. There are no correlations between pollen morphology and the variation in the androecium in Pfaffia, nor would pollen structure support recognition of Hebanthe as a distinct genus. Woehleria and Irenella may be derived from, or be of the same origin as, Dicraurus and Iresine. All four genera are placed in the subfamily Gomphrenoideae because of the bisporangiate anthers, but their pollen structure is very close to, or identical with, that of the Amaranthus-type. Pseudogomphrena combines characteristics of Gomphrena and Pfaffia.     

The Phylogenetic Relationships of the Gobioninae (Teleostei: Cyprinidae) Inferred from Mitochondrial Cytochrome b Gene Sequences

The Gobioninae are a group of morphologically and ecologically diverse Eurasian freshwater cyprinid fishes. The intergeneric relationships of this group are unresolved and the possible monophyly of this subfamily remains to be established. We used complete mitochondrial cytochrome b gene sequences from most genera within the gobionine group, in addition to a selection of cyprinid outgroups, to investigate the possible monophyly of this group and resolve the interrelationships within the group. Our results support the monophyly of the Gobioninae and identify four monophyletic groups within the subfamily; the Hemibarbus group, the Sarcocheilichthys group, the Gobio group, and the Pseudogobio group. The morphologically aberrant genera Gobiobotia, Xenophysogobio and Gobiocypris are included in the Gobioninae, with the latter a sister group of Gnathopogon.     

Root anatomy of palms III. Ceroxyloideae,Nypoideae, Phytelephantoideae

Summary The root-anatomy of representatives of nine genera of Ceroxyloideae of the eleven genera accepted by UHL & DRANSFIELD (1987), of Nypa, the onliest genus of Nypoideae, and of all three genera of Phytelephantoideae have been studied. In Coryphoideae and Calamoideae every genus can be distinguished by the anatomy of the roots, whereas within one genus the species do not differ but slightly. This is true for the newly examined genera too and contributes both to define and identify the genera and towards an improved understanding of the relationship between genera and subfamilies. The genera of Ceroxyloideae are obviously related, they form a natural group, and even the somewhat isolated Pseudophoenix is well at home here. Both Nypa and Phytelephantoideae do not exhibit any affinities with other palms studied up to now.     

Systematics of the Neotropical braconid wasps of the Pedinotus genus group (Doryctinae)

The braconid subfamily Doryctinae (Hymenoptera) is a cosmopolitan, species‐rich group of parasitoid wasps whose known species richness and morphological diversity is mainly concentrated in the Neotropical region. Among the doryctine taxa that are endemic to this region, there is a group of six genera (the Pedinotus genus group) that are mainly characterized by having lateral, posteriorly converging or subparallel grooves on the second metasomal tergite (including Gymnobracon Szépligeti, Hybodoryctes Szépligeti, Lamquetia Braet et van Achterberg, Osmophila Szépligeti, Pedinotus Szépligeti and Trigonophasmus Enderlein). Most of these genera were described more than a century ago, and thus their limits and diagnostic morphological features are not well defined. We present an analysis of the phylogenetic relationships among various representative species of five of the six genera belonging to the Pedinotus genus group based on one mitochondrial (cytochrome oxidase I) and two nuclear (28S, Wingless) gene markers. We also estimated the timing of origin and diversification within the group and investigated the external morphology of the genera involved in order to determine reliable identification characters. The group as a whole, whilst being recovered as monophyletic with the inclusion of Acanthorhogas Szépligeti, was not statistically supported. However, the respective monophyly of all of the morphologically supported genera are confirmed, as are the intergeneric relationships (Gymnobracon, Trigonophasmus) ((Acanthorhogas, Osmophila) (Lamquetia, Pedinotus)). We propose that the pattern of sublateral grooves or depressions appears to be a useful systematic character for recognition of an endemic Neotropical radiation that could have originated c. 41–27 Ma. We redescribe six of the seven genera that we recognize for the Pedinotus genus group and provide a key to enable their identification.     

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.     

Phylogeny of xanthopygine rove beetles (Coleoptera) based on six molecular loci

The rove beetle subtribe Xanthopygina (Coleoptera: Staphylinidae: Staphylininae: Staphylinini) is a species‐rich group of 27 neotropical genera that contains some of the largest and most brightly coloured of all staphylinid beetles. The monophyly of the subtribe has never been tested before, using a large dataset of taxa and genes. Bayesian and maximum likelihood analyses are used on individual genes (COI, 28S rDNA, wingless, arginine kinase, CAD and topoisomerase I) and the partitioned concatenated dataset to test for monophyly and examine the relationships among Xanthopygina genera. Xanthopygina (excluding Philothalpus) are shown to be a monophyletic group with strong support values. The genus Philothalpus is removed from Xanthopygina and placed in the tribe Staphylinini as incertae sedis. Four distinct clades of Xanthopygina genera are recognized. The origin of Xanthopygina is hypothesized to be in the Late Cretaceous or later and the origin of myrmecophilous adaptations is discussed.     

Relationships ofLomatium among other genera ofApiaceae

Lomatium is the largest North American genus ofApiaceae (=Umbelliferae). Relations amongLomatium and other genera have not been well understood. TheApiaceae-Araliaceae lineage and genera of North AmericanApiaceae were analyzed phenetically and cladistically in turn, and seven phenetic alliances of genera are identified. Data on frequencies of character-state combinations were assembled and usingSporne's notions of maximal and minimal co-variation of states, two character-states were identified that are derived among Umbellifer genera. Using cladistic analysis of the alliance consisting ofLomatium and six other genera we identified three sister groups toLomatium:Polytaenia alone;Polytaenia withPseudocymopterus, and a group of four genera. Not all genera in the group of seven withLomatium are supported by autapomorphies impeding phylogenetic reconstructions. Both phenetics and cladistics have helped to suggest and evaluate relationships; to call attention to groups requiring further study, and in particular to identify derived character-states.     

A MORPHOLOGICAL STUDY AND TAXONOMIC REASSESSMENT OF THE GENERITYPE SPECIES IN THE GRACILARIACEAE1

We investigated the reproductive morphology of representative material corresponding to the type species of each of the described genera presently placed in synonymy under Gracilaria. From these observations and published studies of recognized genera, 10 species groups are identified in the Gracilariaceae based on spermatangial type and cystocarp development. Actual or potential generic names are given in brackets after each group: 1) abscissa group (Melanthalia), 2) flabellata group (Curdiea), 3) lemaneiformis group (Gracilariopsis), 4) chilensis group, 5) edulis group (Plocaria/Polycavernosa), 6) urvillei group (Hydropuntia), 7) crassissima group, 8) salicornia group (Corallopsis), 9) gracilis group, and 10) bursa‐pastoris group (Gracilaria). Tyleiophora was shown to belong to the bursa‐pastoris group. The type species of the parasitic genera Gracilariophila and Congracilaria are closely related to their host species. Species assemblages recognized here based on morphological evidence received moderate to strong bootstrap support in recently published molecular phylogenies. Further studies may show that some groups correspond to genera, whereas others do not merit generic status.     

Phylogenetic relationships of the Magnoliaceae inferred from cpDNA matK sequences

The coding region of the matK gene was sequenced to infer the phylogeny of the family Magnoliaceae. Phylogenetic analyses of 21 matK sequences representing ten genera of Magnoliaceae and three outgroups suggest relationships among both subfamilies and genera. Monophyly of the subfamily Liriodendroideae (the genus Liriodendron) and the subfamily Magnolioideae is strongly supported, respectively. Within the subfamily Magnolioideae, three clades are formed: (1) the genus Magnlietia, (2) the subgenus Magnolia, and (3) the subgenus Yulania, with the genera Michelia, Paramichelia, Tsoongiodendron, Alcimandra, Kmeria, Parakmeria and Manglietiastrum. However, the genus Magnolia is shown to be a polyphyletic group, and the genus Michelia a paraphyletic group. Relatively low sequence divergences are detected among genera of the the subfamily Magnolioideae, ranging from 0.14% to 1.70%, especially in the tribe Micheliinae (0.14–0.98%). Molecular evidence from matK sequence data suggests that the phylogenetic positions and the delimitation of the eight genera Magnolia, Michelia, Tsoongiodendron, Paramichelia, Alcimandra, Kmeria, Parakmeria and Manglietiastrum need to be reconsidered. Received: 2 January 2000 / Accepted: 12 February 2000     

Genera and species of Baetidae in Japan: <Emphasis Type="Italic">Nigrobaetis,Alainites, Labiobaetis</Emphasis>, and <Emphasis Type="Italic">Tenuibaetis</Emphasis> n. stat. (Ephemeroptera)

We describe morphological characters of the genera Nigrobaetis, Alainites, Labiobaetis, and Tenuibaetis n. stat. and provide generic situations of six Japanese species: Nigrobaetis chocoratus n. comb., N. sacishimensis n. comb., Alainites atagonis, A. florens, A. yoshinensis, and Tenuibaetis pseudofrequentus. To evaluate the polarities of the morphological characters and the monophyly of Nigrobaetis, Alainites, Labiobaetis, and Tenuibaetis, character states of these four genera were compared with the genus Cloeon as an outgroup. Labiobaetis is considered to be a monophyletic group supported by a wide paraglossa. Tenuibaetis is a monophyletic group that is distinguishable from the related genera by robust setae with a medial ridge on the dorsomedian surface of the nymphal femur. We did not find any synapomorphic characters of Nigrobaetis or Alainites. Although we tentatively treat Nigrobaetis and Alainites as distinct genera, they are considered to be paraphyletic taxa.     

Evolutionary relationships in thePodalyrieae andLiparieae (Fabaceae) based on morphological, cytological, and chemical evidence

Taxonomic relationships amongst the genera of the southern African tribesPodalyrieae andLiparieae are discussed. Data gained from morphological, cytological and chemical investigations are analyzed cladistically to determine relationships. The genusCadia (tribeSophoreae) is included in the investigation to establish whether it should be transferred to thePodalyrieae. The results clearly indicate that thePodalyrieae andLiparieae are monophyletic and that they should be united, but thatHypocalyptus andCadia should be excluded. Within the monophyletic group, there are two distinct subclades each supported by three apomorphies. The results also show that there is a strong sister relationship betweenAmphithalea andCoelidium. In the taxonomic treatment theLiparieae are placed into synonymy under thePodalyrieae and two subtribes are recognized. A key to the genera in the tribe is given, followed by a synopsis of the genera.     

Phylogenetic relationships in the Marcetia alliance (Melastomeae,Melastomataceae) and implications for generic circumscription

The Marcetia alliance of Melastomataceae is an exclusively Neotropical group that includes at least 12 genera of mostly herbs and subshrubs, occurring in the cerrado of central Brazil and savannas of the Amazon region and Guayana highlands. This study aimed to test the monophyly of genera in the Marcetia alliance, evaluate their phylogenetic relationships and generic boundaries, and investigate morphological characters as potential synapomorphies for delimiting clades or genera. We used nuclear (ITS, ETS) and plastid (accD‐psaI, atpH‐atpF, trnS‐trnG) DNA sequences of 107 terminals in 12 genera from the alliance. Aciotis, Fritzschia, Marcetia and Siphanthera were shown to be monophyletic and supported by molecular and morphological characters. Other genera with variable morphology and wider distributions, such as Acisanthera, Comolia, Ernestia and Macairea, were recovered as paraphyletic or polyphyletic. Most morphological characters analysed were found to be homoplastic, but when combined they are potentially useful for the diagnosis of genera and infrageneric groups. This study represents a major step in understanding internal relationships and provides the basis for a revision of the generic classification in the Marcetia alliance.     

An introductory ecological biogeography of the Australo-Pacific Meliphagidae

Abstract

The Meliphagidae, that can readily be defined on tongue characteristics, are a monophyletic group centred in the Australo-Pacific region, but with one African genus (Promerops). The classification of Salomonsen (1967) allows 38 genera and 170 species in the former region, and one genus with two species in the latter. Australia and New Guinea jointly have 23 genera and 108 species, and constitute the centre of diversity of the group. Endemic genera are concentrated in Australia and New Guinea, and around the periphery of the Pacific part of the range (Sulawesi, Bonins, Marianas, Hawaii, New Zealand). The meliphagids are diversified in body size and bill form. They are basically nectarivores and insectivores, with most species combining the two roles to varying degrees. There is a good general correlation between bill form and way of life. A few species feed on trunks and aerial flycatching is well developed in many. Morphological modification is only minor in these instances and the meliphagids as a group remain rather generalised in bodily proportions. A long period of coevolution with Australian plant elements is shown by meliphagids being the major pollinators of several tree and shrub genera.

The group combines monotypic genera with restricted ranges and wide-ranging genera with many species. Of the latter; Myzomela, Lichema, and Philemon are centred in the tropics, and Meliphaga and Phylidonyris in Australia. Most of these co-occur over a wide area, this being favoured by differences in body size and bill morphology.

Comparison of three kinds of meliphagid communities, two typical continental ones, two of isolated forest outlyers in Australia, and six insular Pacific ones, shows the first to be rich (10 and 11 genera, 21 and 17 species), and the second impoverished (6 and 7 genera, 9 and 12 species). Individual Pacific island groups, however, have only 2–5 genera, and 3–6 species. Genus to species ratios are 0.55–0.64 in the major continental communities, but are 1.0 in New Zealand and Samoa.

Morphological distance between species, measured as the percent difference in size between successive members along a size gradient is 5.4 and 5.5% for wing length and 4.9 and 9.3% for bill length in the two continental communities. It increases to 7.8–14.9, and 11.3–12.7%, respectively, in the isolated forest outlyers of Tasmania and southwestern Australia. The figures are 23.0 and 35.0% for wing and bill length in New Zealand, and 41.0 and 51.0% in Fijian forms. This accords with current theory that in impoverished insular environments, size separation of co-occurring species must be greater.

The marked success of the Meliphagidae in the Australo-Pacific region can be attributed to their versatility and adaptibility, and dual role of insectivore and nectarivore in an area exceptionally rich in nectar-producing trees and shrubs.     

Systematics ofLavatera andMalva (Malvaceae,Malveae)—a new perspective

Lavatera andMalva are circumscribed by epicalyx characters, but the distinction is untenable on this basis. Morphological characters and phylogenetic trees from rDNA ITS sequence data indicate the presence of two well-supported species groups different from the traditional genera. One group includes a mixture of closely related species in both genera and forms a more broadly circumscribedMalva, including most disjunctLavatera species—notably four of western North American islands, one Australian and one Canarian species. The other group forms a more narrowly definedLavatera. Another CanarianLavatera species may be a common ancestor to the two groups.     

Time Constraints Limit Group Sizes and Distribution in Red and Black-and-White Colobus

Researchers have shown that, in frugivorous primates, a major constraint on group size is intra group feeding competition. The relationship is less obvious in folivorous primates. We investigated whether colobine group sizes are constrained by time limitations as a result of their low energy diet and ruminant-like digestive system. We used climate as an easy to obtain proxy for the productivity of a habitat. Using the relationships between climate, group size, and time budget components for populations of Colobus and Piliocolobus at different research sites, we created 2 taxon-specific models. In both genera, feeding time increased with group size or biomass. The models for Colobus and Piliocolobus correctly predicted the presence or absence of the genera at, respectively, 86% of 148 and 84% of 156 African primate sites. Median predicted group sizes where the respective genera were present are 19 for Colobus and 53 for Piliocolobus. We show that the differences between the 2 genera are due mainly to intrinsic differences in the way each taxon’s digestive physiology interacts with climatic variables to influence resting time requirements. The models may help us explore their responses to climatic change in both the past and the future.     

Ultrastructure of nuclear inclusions and the separation ofVerbenaceae andOleaceae (incl.Nyctanthes)

TEM observations were carried out on 40 taxa of the familyVerbenaceae and 35 taxa of the familyOleaceae, in order to ascertain distribution, ultrastructure and morphology of the intranuclear proteinic inclusions in the mesophyll parenchymatic cells. The investigated genera amount to some 25% and 60% respectively of the genera of the two families. Inside theVerbenaceae, lamellar inclusions (L-type) occur in 6 out of 23 investigated genera: they are mostly present inside the tribesCitharexyleae andVerbeneae (both belonging toVerbenoideae), while they are absent in other subfamilies. All of the investigatedOleaceae genera show intranuclear crystalline inclusions (C 1-type) of three different shapes. Among theAsteridae this is a character peculiar toOleaceae. They appear to be a well defined natural group, including the controversial genusNyctanthes.     

The suprageneric groups ofthe Pimplinae (Hymenoptera: Ichneumonidae): a cladistic re-evaluationand evolutionary biological study

The phylogenetic relationships of the suprageneric groupsof the ichneumonid subfamily Pimplinae (Hymenoptera) are re‐assessedusing 166 morphological and biological characters for 162 species,representing all of the available described genera and subgenera.The cladistic analysis was repeated using abstracted genera, re‐codedfrom the ­initial set of species, as terminal taxa. Thetopology of the resulting cladograms was similar. In the first (primary) analysisseveral genera (including Neotheronia, Itoplectis, Dolichomitus, Dreisbachia, Polysphincta, Oxyrrhexis and Zonopimpla)were not retrieved as monophyletic groups; however, all except thelast were found to be monophyletic in the second analysis. Theseresults suggest that using abstracted taxa may force a ‘falsemonophyly’ on the preselected groups. Thus we reject theuse of such abstractions, preferring instead to use exemplar speciesthat together show much of the variation that occurs within a hypothesizedgenus. Within the Pimplinae three major groupings were recognized,the Delomeristini (including the Perithoini syn. nov.) , thePimplini and the Ephialtini. Within the Pimplini, two generic groupswere recovered, the Xanthopimpla and Pimpla genus‐groups,but a third postulated group, the Theronia genus‐group, wasfound to be paraphyletic. Within the Ephialtini five groups wererecognized, the Pseudopimpla, Alophosternum, Camptotypus, Ephialtes and Sericopimpla genus‐groups.The spider parasitizing complex of genera (the Polysphincta genus‐complex)was found to nest within the Sericopimpla genus‐group confirmingthe placement of Polysphinctini as a synonym of Ephialtini. Problemswith the status of some existing genera are highlighted, but formalnomenclatural changes are not proposed. The ancestral Pimplinaeare hypothesized to have been solitary ectoparasitic idiobiontson weakly concealed immature Hymenoptera. The major radiations withinthe Pimplinae are shown as: (1) a progressive exploitation of cocooned,then weakly cocooned, lepidopterous pupae in the Pimplini leadingto idiobiont endoparasitism; (2) increasing specialization to attackhosts deeply concealed in wood in the Ephialtes genus‐group,and (3) specialization on a variety of cocooned hosts, includingspider egg sacs, leading to koinobiont ectoparasitism of spiders.A brief synopsis of the distribution of the group is given, and somebiogeographical inferences drawn. The group is presumed to haveoriginated and radiated on Laurasia; no evidence for trans‐Antarcticrelationships can be found. © 2002 The LinneanSociety of London, Zoological Journal of the Linnean Society,2002, 136 , 421?485