Phylogeny of cardinalfishes (Teleostei: Gobiiformes: Apogonidae) and the evolution of visceral bioluminescence

The cardinalfishes (Apogonidae) are a diverse clade of small, mostly reef-dwelling fishes, for which a variety of morphological data have not yielded a consistent phylogeny. We use DNA sequence to hypothesize phylogenetic relationships within Apogonidae and among apogonids and other acanthomorph families, to examine patterns of evolution including the distribution of a visceral bioluminescence system. In conformance with previous studies, Apogonidae is placed in a clade with Pempheridae, Kurtidae, Leiognathidae, and Gobioidei. The apogonid genus Pseudamia is recovered outside the remainder of the family, not as sister to the superficially similar genus Gymnapogon. Species sampled from the Caribbean and Western Atlantic (Phaeoptyx, Astrapogon, and some Apogon species) form a clade, as do the larger-bodied Glossamia and Cheilodipterus. Incidence of visceral bioluminescence is found scattered throughout the phylogeny, independently for each group in which it is present. Examination of the fine structure of the visceral bioluminescence system through histology shows that light organs exhibit a range of morphologies, with some composed of complex masses of tubules (Siphamia, Pempheris, Parapriacanthus) and others lacking tubules but containing chambers formed by folds of the visceral epithelium (Acropoma, Archamia, Jaydia, and Rhabdamia). Light organs in Siphamia, Acropoma, Pempheris and Parapriacanthus are distinct from but connected to the gut; those in Archamia, Jaydia, and Rhabdamia are simply portions of the intestinal tract, and are little differentiated from the surrounding tissues. The presence or absence of symbiotic luminescent bacteria does not correlate with light organ structure; the tubular light organs of Siphamia and chambered tubes of Acropoma house bacteria, those in Pempheridae and the other Apogonidae do not.     

Subgeneric differentiation within Kurzia (Crustacea: Anomopoda: Chydoridae) and a new species from Central America

The genus Kurzia is divided into two subgenera: Kurzia s. str. and Rostrokurzia n. subg. Subgenus Kurzia includes Kurzia (K.) latissima Kurz, 1874, with a palearctic -, Kurzia (K.) polyspina n. sp. with a neotropic - , and Kurzia (K.) cf. media (Birge, 1879) with a nearctic distribution. Rostrokurzia includes Kurzia (R.) longirostris Daday, 1898 (pantropical distribution), and Kurzia (R.) brevilabris Rajapaksa & Fernando, 1986, from subtropical and tropical Asia. Kurzia latissima Kurz, 1874, from Central Europe is redescribed in detail.     

The natural delimitation ofCentaurea (Asteraceae: Cardueae): ITS sequence analysis of theCentaurea jacea group

Taxonomic complexity has hindered partitioning the genusCentaurea into natural subdivisions, even though it has long been recognized as an unwieldy, artificial assemblage. Most of the remaining difficulties center in theCentaurea jacea group, whose taxa share a common advanced type of pollen. Because it comprises half the species of the genusCentaurea, as well as five other disputed genera previously segregated fromCentaurea (Chartolepis, Cheirolepis, Cnicus, Grossheimia andTomanthea), theCentaurea jacea group is a significant taxonomic challenge. Newer molecular approaches are useful for resolving complex relationships because they provide more precise inferences of evolutionary relationships than traditional morphological characters. Sequences of the Internal Transcribed Spacers (ITS) of nuclear ribosomal DNA were analyzed for a comprehensive sample of this group. Results indicated that theCentaurea jacea group is monophyletic and includes the segregated genera, but not two other genera (Oligochaeta andZoegea), whose inclusion in theCentaurea jacea group was doubtful. In addition to pollen morphology, the ITS phylogeny is also supported by karyological evidence and by good correlation with biogeographic distribution of the species. The monophyly of theCentaurea jacea group suggests that a natural delimitation ofCentaurea that minimizes nomenclatural changes is possible, but only if a new type of the genus is designated.     

Predation potential of three flatworm species (Platyhelminthes: Turbellaria) on mosquitoes (Diptera: Culicidae)

We conducted a field survey for flatworms to select species as potential biological control agents against Aedes aegypti and Culex pipiens (Diptera, Culicidae) breeding in artificial containers. Laboratory experiments were performed to determine the daily predation rate, differential predation on each mosquito larval instar, selective predation on either A. aegypti or C. pipiens, and predator tolerance to water from artificial containers. Girardia anceps (Tricladida, Paludicola, Dugesiidae), Mesostoma ehrenbergii and Bothromesostoma cf. evelinae (Rhabdocoela, Typhloplanoida, Typhloplanidae) were found in temporary puddles and permanent pools. In the laboratory, they killed between 52% and 100% of immature mosquitoes coexisting in the same habitat. No preference of flatworms for mosquito preys was detected. Predation rate was related to predator size and instar of preys. Girardia anceps and B. evelinae survived after a dry period and when re-flooding occurred, they laid eggs. Tolerance to water from artificial containers was highest in G. anceps and this species could be a suitable predator to reduce mosquito populations from artificial containers using an inoculative approach.     

Medfly eradication in California:

The impact of malathion-bait sprays (directed against medfly, Ceratitis capitata [Wiedemann]) on an endemic gall midge (Rhopalomyia californica Felt) and its parasitoids was investigated during 1982–83 in the south San Francisco Bay area of northern California. In a heavily sprayed area (Woodside), a population explosion of the midge was detected following 24 applications of malathion bait. The midge population reached levels ca. 90x greater than those observed in an adjacent unsprayed area (Jasper Ridge). In a moderately sprayed area (Portola Valley), the midge population increased as much as 5x that observed in the adjacent unsprayed area (Jasper Ridge), following 12 applications of malathion bait. In laboratory tests, the malathion bait was toxic to both the midge and its parasitoids. The major parasitoids were Torymus koebelei (Huber), Zatropis capitis Burks, Platygaster californica (Ashmead) and Mesopolobus sp. Population increases of the midge following malathion-bait sprays were attributed to destruction of parasitoids and other natural enemies of the midge. If the environmental impact of malathion-bait sprays is related to the number of applications (as suggested in this study), then it would be worthwhile to determine the appropriate bait-spray strategy for a given situation, so as to minimize adverse effects on nontarget species, yet insure suppression or eradication of medfly.

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Résumé L'impact des pièges tratiés au malathion (destinés à Ceratitis capitata Wiedem) sur Rhopalomyia californica Felt et ses parasitoïdes a été examiné en 1982–1983 dans le sud de la zone de la baie de San Francisco en Californie. Dans une zone fortement traitée (Woodside), une explosion de population a été décelée après 24 traitements. La population de R. californica a atteint des niveaux 90 fois supérieurs à ceux observés dans une zone contiguë non traitée (Jasper Ridge). Dans une zone modérément traitée (Portola Valley), avec 12 traitement, la population de R. californica a atteint jusqu'a 5 fois celle de Jasper Ridge. Au laboratoire, le piège à malathion a été toxique tant pour R. californica que pour ses parasitoïdes, dont les principaux étaient: Torymus koebelei (Huber), Zatropis capitis Burks, Platygaster californica(Ashmead) et Mesopolobus sp. L'accroissement de la population de C. capitata après traitement a été attribué à la destruction de parasitoïdes et d'autres ennemis naturels. Si l'effet su l'environnement du traitement est lié au nombre d'interventions (comme le suggère cette étude), alors cela vaudrait la peine de définir une stratégie de traitement appropriée à une situation donnée, de façon à minimiser les effets négatifs sur des espèces non visées, tout en assurant la suppression ou l'éradication de C. capitata.

     

Development and reproduction of Stratiolaelaps scimitus (Acari: Laelapidae) with fungus gnat larvae (Diptera: Sciaridae), potworms (Oligochaeta: Enchytraeidae) or Sancassania aff. sphaerogaster (Acari: Acaridae) as the sole food source

Stratiolaelaps ( = Hypoaspis) miles (Berlese) (Acari: Mesostigmata: Laelapidae) is a polyphagous soil-dwelling predatory mite that is widely marketed for use in greenhouse production systems to manage populations of dark-winged fungus gnats, Bradysia spp. (Diptera: Sciaridae) and for supplemental control of thrips. The suggestion by Walter and Campbell (2003, Biol. Control 26: 253–269) that North American commercial cultures of S. miles may actually be S. scimitus was confirmed. The development and reproduction at 21–23 °C of S. scimitus provided ad libidum with one of three different prey – the fungus gnat Bradysia aff. coprophila (Lintner), potworms (Enchytraeidae), or Sancassania aff. sphaerogaster (Zachvatkin, 1937) (Acari: Astigmata: Acaridae) – were compared. Developmental duration of the egg and non-feeding larval stages were 2.47 and 1.11 days, respectively; mortalities were 8.3 and 5.5%. Stratiolaelaps scimitus failed to develop beyond the protonymphal stage when provided with S. aff. sphaerogaster alone, although some feeding was observed. Development and reproduction of S. scimitus was successful on both fungus gnat larvae and enchytraeids, with no influence of prey on protonymphal duration (4.70 days) and mortality (8.3%), or on deutonymphal duration (4.61 days) and mortality (6.1%). Adult female S. scimitus feeding on potworms, compared to those feeding on fungus gnat larvae, had a significantly shorter pre-oviposition period (2.69 vs. 4.59 days). However, diet did not influence other adult female developmental or reproductive characteristics (oviposition period, 18.6 days; post-oviposition period, 6.2 days; total adult longevity, 27.3 days; total number of eggs, 26.5). S. scimitus reared on potworms tended (p = 0.06) to have a higher intrinsic rate of increase, a higher finite rate of increase and a shorter doubling time (r _m = 0.142 day⁻¹, λ = 1.153, Dt = 4.85 days) than those reared on fungus gnat larvae (r _m = 0.105 day⁻¹, λ = 1.110, Dt = 6.58 days), but differences in net reproductive rate (R ₀) and generation time (G) were not significant.     

Hectorella: A member of the betalain-suborderChenopodiineae of the orderCentrospermae

Roots ofHectorella caespitosa Hook. f. were induced to produce a red pigment which was shown to be a betalain and not an anthocyanin. These data indicate thatHectorella belongs to theChenopodiineae, the betalain suborder of theCentrospermae, and excludes alignment with the anthocyanin family theCaryophyllaceae.     

Taxonomic status of three nominal species of Indo-Pacific symphurine tonguefishes (Symphurus: Cynoglossidae: Pleuronectiformes)

The taxonomic status of three rarely captured, nominal species of Indo-Pacific symphurine tonguefishes,Symphurus trifasciatus (Alcock, 1894),S. microrhynchus (Weber, 1913), andS. holothuriae Chabanaud, 1948, is resolved.Symphurus microrhynchus, previously known only from three specimens, is redescribed based on 12 specimens.Symphurus holothuriae, previously reported only from the holotype, is a junior subjective synonym ofS. microrhynchus. Symphurus trifasciatus is redescribed based on six specimens, including the lectotype designated in this study.Symphurus microrhynchus is distinguished from congeners by a combination of meristic features, head and opercle shape, dorsalfin position, and features of its pigmentation. The species occurs in relatively shallow tropical waters from Zanzibar through French Polynesia (the only reported occurrence of a shallow-waterSymphurus from a locality on the Pacific Plate), including Indonesia, Papua New Guinea, Vietnam, and northwestern Australia.Symphurus trifasciatus, known from deep waters along both coasts of the Indian subcontinent and adjacent waters, has an overall similarity in body shape and meristic features reminiscent of those inS. microrhynchus, but differs notably in having a longer and wider head, a distinctive opercle shape, number of scale rows on the head, and pigmentation pattern.     

Longevity of multiple species of tephritid (Diptera) fruit fly parasitoids (Hymenoptera: Braconidae: Opiinae) provided exotic and sympatric-fruit based diets

While adult parasitic Hymenoptera in general feed on floral and extrafloral nectars, hemipteran-honeydews and fluids from punctured hosts, Diachasmimorpha longicaudata (Ashmead), an Old World opiine braconid introduced to tropical/subtropical America for the biological control of Anastrepha spp. (Tephritidae), can survive on fruit juices as they seep from injured fruit. An ability to exploit fruit juice would allow such a parasitoid to efficiently forage for hosts and food sources simultaneously. Two New World opiines, Doryctobracon areolatus (Szepligeti) and Utetes anastrephae (Viereck), are also prominent Anastrepha parasitoids and are roughly sympatric. All three species were provided with: (1) pulp and juice diets derived from a highly domesticated Old World fruit (orange, Citrus sinensis L.) that is only recently sympatric with the Mexican flies and parasitoids and so offered little opportunity for the evolution of feeding-adaptations and (2) a less-domesticated New World fruit (guava, Psidium guajava L.), sympatric over evolutionary time with D. areolatus and U. anastrephae. Both sexes of D. longicaudata died when provided guava pulp or juice at a rate similar to a water-only control. D. areolatus and U. anastrephae, presumably adapted to the nutrient/chemical constituents of guava, also died at a similar rate. Survival of all three species on orange pulp and juice was greater than on water, and often equaled that obtained on a honey and water solution. In confirmatory experiments in Mexico, D. areolatus and U. anastrephae, as well as other tephritid parasitoids Doryctobracon crawfordi (Viereck) and Opius hirtus (Fisher), all died at a significantly higher rates when provided guava in comparison to a honey and water diet. Such a result is likely due to guavas being repellent, innutritious or toxic. D. longicaudata clearly consumed guava juice tagged with a colored dye. Dilutions of orange and guava juice resulted in shorter lifespans than dilutions of orange juice and water demonstrating that there while diluted orange juice provided nutrition the addition of guava created toxicity. Given the differences in fruit-food quality, adult opiine food sources would not be obtainable at all oviposition sites and in the case of guava, more additional sites and foraging for food than previously postulated may be required.     

Primitive Defence: The MiAMP1 Antimicrobial Peptide Family

The founder of the MiAMP1 protein family was originally isolated from Macadamia integrifolia and had antimicrobial activity in vitro. MiAMP1 was the first plant protein with a structure containing a βγ-crystallin precursor fold, a structural superfamily associated with antimicrobial proteins in other kingdoms. In recent times, expanding plant genomics information has revealed that genes encoding homologues of MiAMP1 are conserved across the plant kingdom from lycophytes, gymnosperms to early angiosperms (e.g. Amborella, Papaver) and some monocots (e.g. Zantedeschia, Zea, Sorghum). Many studies of plant–pathogen interactions in gymnosperms have demonstrated a potential role for MiAMP1 family members in defence against fungal pathogens. This commentary describes the discovery and diversity of this protein family and considers current evidence supporting, and future opportunities for substantiating, a role in defence in primitive plants, and why this role may have diminished in higher plants.     

Suitability ofScapteriscus moles crickets [Ort.: Gryllotalpidae] as hosts ofEuphasiopteryx depleta [Dipt.: Tachinidae]

Euphasiopteryx depleta (Wied) flies successfully developed in the mole cricketsScapteriscus borellii Giglio-Tos andScapteriscus vicinus Scudder, as well as in the short-tailed cricket,Anurogryllus sp., after artificially placing larvae under their pronota. Using this technique, no relation was found between the number of larvae placed on each cricket and the resulting number of puparia obtained from the cricket. In all cases, the number of flies emerging from the puparia was small, with females failing to mature reproductively. AsE. depleta has been reared from field collectedScapteriscus spp., the exact reasons for lack of maturation remain unknown.Anurogryllus sp. can be considered a probable alternate host forE. depleta.      

A phenetic study of Cassia sensu lato (Leguminosae-Caesalpinioideae: Cassieae: Cassiinae) in Thailand

Cassia L. sensu lato, a large heterogeneous genus of flowering plants, occurs naturally in the tropics around the world. Recent works based on floral morphology have supported a division of this genus into three genera, namely Cassia L. s. str., Chamaecrista Moench and Senna Mill. In order to investigate this new classification, 508 specimens of 18 taxa of the genus Cassia s.l. grown in Thailand were analyzed using cluster analysis and canonical discriminant analysis. The total 32 vegetative and reproductive morphological characters were employed in these analyses. In cluster analysis, Cassia s.l. can be separated into four groups, respectively viz. Chamaecrista, Senna alata, Senna and Cassia s /str. The four-cluster grouping is discussed. From a canonical discriminant analysis using the four-cluster grouping as a priori groups, it can be concluded that Cassia s. str., Senna, and Chamaecrista are indeed distinct taxa. The three most important characters that separate the three genera are filament length, fruit length, and ovary stalk length. These quantitative characters, together with some qualitative characters, were useful in constructing an identification key to these genera. Among the three genera, it was also found that Senna is rather a heterogeneous taxon. The difference between the studied species was discussed.     

Rickettsial Agents from Parasitic Dermanyssoidea (Acari: Mesostigmata)

Mites are often overlooked as vectors of pathogens, but have been shown to harbor and transmit rickettsial agents such as Rickettsia akari and Orientia tsutsugamushi. We screened DNA extracts from 27 mites representing 25 species of dermanyssoids for rickettsial agents such as Anaplasma, Bartonella, Rickettsia, and Wolbachia by PCR amplification and sequencing. DNA from Anaplasma spp., a novel Bartonella sp., Spiroplasma sp., Wolbachia sp., and an unclassified Rickettsiales were detected in mites. These could represent mite-borne bacterial agents, bacterial DNA from blood meals, or novel endosymbionts of mites.     

Ammonium sensing in nitrogen fixing bacteria: Functions of theglnB andglnD gene products

A plentiful supply of fixed nitrogen as ammonium (or other compounds such as nitrate or amino acids) inhibits nitrogen fixation in free-living bacteria by preventing nitrogenase synthesis and/or activity. Ammonium and nitrate have variable effects on the ability ofRhizobiaceae (Rhizobium, Bradyrhizobium andAzorhizobium) species to nodulate legume hosts and on nitrogen fixation capacity in bacteroid cells contained in nodules or in plant-free bacterial cultures. In addition to effects on nitrogen fixation, excess ammonium can inhibit activity or expression of other pathways for utilization of nitrogenous compounds such as nitrate (through nitrate and nitrite reductase), or glutamine synthetase (GS) for assimilation of ammonium. This paper describes the roles of two key genesglnB andglnD, whose gene products sense levels of fixed nitrogen and initiate a cascade of reactions in response to nitrogen status. While work onEscherichia coli and other enteric bacteria provides the model system,glnB and, to a lesser extent,glnD have been studied in several nitrogen fixing bacteria. Such reports will be reviewed here. Recent results on the identity and function of theglnB andglnD gene products inAzotobacter vinelandii (a free-living soil diazotroph) and inRhizobium leguminosarum biovarviciae, hereinafter designatedR.l. viciae will be presented. New data suggests thatAzotobacter vinelandii probably contains aglnB-like gene and this organism may have twoglnD-like genes (one of which was recently identified and namednfrX). In addition, evidence for uridylylation of theglnB gene product (the PII protein) ofR. l. viciae in response to fixed nitrogen deficiency is presented. Also, aglnB mutant ofR. l. viciae has been isolated; its characteristics with respect to expression of nitrogen regulated genes is described.     

Paraveinal mesophyll: Review and survey of the subtribeErythrininae (Phaseoleae,Papilionoideae, Leguminosae)

Paraveinal mesophyll (PVM) is a distinctive anatomical feature of the leaf mesophyll of some plant taxa that may represent a specialized physiological compartment. A comprehensive review of the 42 published references that mention PVM or similar cell layers and a survey of 121 of the 272 species of all nine genera of thePhaseoleae subtribeErythrininae demonstrate that PVM is nearly exclusively found inLeguminosae. InLeguminosae, PVM is either rare or absent in subfamilyCaesalpinioideae, uncommon inMimosoideae, and extensively distributed amongPapilionoideae. In subtribeErythrininae, PVM is ubiquitous inErythrina, and occurs in four other genera. ThreeErythrininae genera (Apios, Mucuna, andCochlianthus) lack PVM. Unique chloroplast-poor, enlarged conical cells (pellucid palisade idioblasts) occur in 80 species ofErythrina but not in any other genus ofErythrininae.     

Recognition of heterospecific parasitism: Competition between Aphidiid (Aphidius ervi) and Aphelinid (Aphelinus asychis) parasitoids of Aphids (Hymenoptera: Aphidiidae; Aphelinidae)

The solitary parasitoids Aphidius erviHaliday (Hymenoptera: Aphidiidae) and Aphelinus asychisWalker (Hymenoptera: Aphelinidae) attacked but generally did not oviposit in pea aphids parasitized by the other species. Wasps selectively oviposited in unparasitized hosts when given a choice. Host discrimination depended on the recognition of internal cues. Females of A. asychiseither could not recognize or ignored A. ervi'sexternal host marking pheromone. Under most conditions, A. ervisurvived in superparasitized hosts, killing competing A. asychislarvae by physical attack and possibly physiological suppression. The outcome of larval competition was not affected by oviposition sequence or age difference between larvae; A. asychissurvived only when it had substantially completed larval development before the host was superparasitized by A. ervi.It is suggested that competition for host resources incurs a cost, for the winner in terms of reduced size or increased development time and for the loser in terms of lost progeny and searching time. Consequently, heterospecific host discrimination can be functional. Internal, and probably general, cues enable wasps to recognize and avoid oviposition in hosts already parasitized by an unrelated species.     

Lipochaeta and Melanthera (Asteraceae: Heliantheae subtribe Ecliptinae): establishing their natural limits and a synopsis

We restrict the genusLipochaeta to the allopolyploid species of the typical section.Lipochaeta s.str. is interpreted to be the result of an intergeneric hybridization betweenMelanthera and a presently unknown taxon, perhaps of the genusWedelia. Lipochaeta is characterized, in addition to its allopolyploidy (n=26), by having both flavonols and flavones, disk corollas with 4 lobes, achenes tuberculate at maturity, the disk achenes flattened to slightly biconvex, and ray achenes obcompressed.Lipochaeta sect.Aphanopappus andWollastonia are here reduced to synonymy underMelanthera. We transfer 14 HawaiianLipochaeta and one New Caledonian species as well as the AsianWedelia prostrata toMelanthera. These transfers, along with the species in Africa and North America, bring the number of species in the genus to 35.Melanthera is delimited by an abruptly narrowed to truncate and flattened top of the achene, (0−)1–15(−20) often unequal, ciliate or barbellate, caducous pappus bristles immediately surrounding the corolla, involucral bracts and receptacular paleae with many veins forming longitudinal striations, and n=15. The florets are 5-merous, the corollas are yellow or white, and rays are absent (in white-flowered species) or present and neutral or fertile. In dealing with species formerly placed inLipochaeta, the GalapagosL. laricifolia is here transferred from the illegitimate generic nameMacraea toTrigonopterum and the BrazilianL. goyazensis is transferred toAngelphytum. We maintain the earlier reduction ofEchinocephalum underMelanthera and reduce all three taxa originally described in it to one,M. latifolia.     

Freshwater gomphonemoid diatom phylogeny: preliminary results

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

Importation,colonization and establishment ofAnagyrus indicus [Hym.: Encyrtidae] onNipaecoccus viridis [Hom.: Pseudococcidae] in Jordan

Anagyrus indicus Shafeeet al. was collected on the island of Guam and released in the Jordan River Valley of Jordan in order to regulateNipaecoccus viridis (Newstead) on citrus.A. indicus was originally released in February 1984 and became colonized by April of the same year. It also dispersed by natural means up to 61 km following the original release and colonized that same year on infestedZizyphus sp. By spring of 1985 it became the dominant parasitoid attackingN. viridis. Another parasitoid,Anagyrus kamali Moursi, was collected in Jordan in June of 1985 for the first time. This latter parasitoid apparently moved into the Jordan River Valley by means of natural dispersal from a neighboring country. Infestations ofN. viridis have been greatly reduced in areas whereA. indicus became established. This paper reports the results of research only. Mention of a proprietary product does not constitute an endorsement for its use by USDA.     

The taxonomy of rhizobia: an overview

The taxonomy of rhizobia, bacteria capable of nodulating leguminous plants, has changed considerably over the last 20 years, with the original genus Rhizobium, a member of the alpha-Proteobacteria, now divided into several genera. The study of new geographically dispersed host plants, has been a source of many new species and is expected to yield many more. Here we provide an overview of the history of the rhizobia, but focus on the Rhizobium–Allorhizobium–Agrobacterium relationship. Finally, we review recent reports of nodulation and nitrogen fixation with legume hosts by bacteria that are outside the traditional rhizobial phylogenetic lineages. They include species of Methylobacterium and Devosia in the alpha- Proteobacteria and of Burkholderia and Ralstonia in the beta-Proteobacteria.