Phylogeny and evolution of hemipteran insects based on expanded genomic and transcriptomic data.

BACKGROUND: Hemiptera is the fifth species-rich order of insects and the most species-rich order of hemimetabolous insects, including numerous insect species that are of agricultural or medical significance. Despite much effort and recent advance in inferring the Hemiptera phylogeny, some high-level relationships among superfamilies remain controversial. RESULTS: We sequenced the genomes of 64 hemipteran species from 15 superfamilies and the transcriptomes of two additional scale insect species, integrating them with existing genomic and transcriptomic data to conduct a comprehensive phylogenetic analysis of Hemiptera. Our datasets comprise an average of 1625 nuclear loci of 315 species across 27 superfamilies of Hemiptera. Our analyses supported Cicadoidea and Cercopoidea as sister groups, with Membracoidea typically positioned as the sister to Cicadoidea + Cercopoidea. In most analyses, Aleyrodoidea was recovered as the sister group of all other Sternorrhyncha. A sister-group relationship was supported between Coccoidea and Aphidoidea + Phylloxeroidea. These relationships were further supported by four-cluster likelihood mapping analyses across diverse datasets. Our ancestral state reconstruction indicates phytophagy as the primary feeding strategy for Hemiptera as a whole. However, predation likely represents an ancestral state for Heteroptera, with several phytophagous lineages having evolved from predatory ancestors. Certain lineages, like Lygaeoidea, have undergone a reversal transition from phytophagy to predation. Our divergence time estimation placed the diversification of hemipterans to be between 60 and 150 million years ago. CONCLUSIONS: By expanding phylogenomic taxon sampling, we clarified the superfamily relationships within the infraorder Cicadomorpha. Our phylogenetic analyses supported the sister-group relationship between the superfamilies Cicadoidea and Cercopoidea, and the superfamily Membracoidea as the sister to Cicadoidea + Cercopoidea. Our divergence time estimation supported the close association of hemipteran diversification with the evolutionary success and adaptive radiation of angiosperms during the Cretaceous period.

Exploring Phylogenetic Relationships of the Mecocephala Group (Hemiptera: Pentatomidae) Using Continuous and Discrete Characters, with Notes on Taxonomy.

The Mecocephala group comprises about 50 species, restricted to the Neotropics and with the highest species richness in Neotropical South America. Several species use rice as host plants and their identification is facilitated by the presence of exaggerated head proportions and a unique male genitalic morphology. The taxonomy of the group has been extensively explored, but inferring its monophyly and especially its internal phylogenetic relationships has been challenging. Here, we inferred the phylogenetic relationships for the group assembling the most complete taxonomic sampling to date, analyzing discrete and continuous morphological characters through equal and implied weighted parsimony analyses. The monophyly of the group was recovered, but internal relationships varied slightly according to the dataset tested. Thus, we propose internal arrangements for the group and provide a formal description of the Mecocephala group, diagnoses for each genus, a dichotomous key to identify its genera, and illustrations of the morphological characters and type species.

The impact of geographic isolation and host shifts on population divergence of the rare cicada Subpsaltria yangi.

The contributions of divergent selection and spatial isolation to population divergence are among the main focuses of evolutionary biology. Here we employed integrated methods to explore genomic divergence, demographic history and calling-song differentiation in the cicada Subpsaltria yangi, and compared the genotype and calling-song phenotype of different populations occurring in distinct habitats. Our results indicate that this species comprises four main lineages with unique sets of haplotypes and calling-song structure, which are distinctly associated with geographic isolation and habitats. The populations occurring on the Loess Plateau underwent substantial expansion at âˆ¼0.130-0.115 Ma during the Last Interglacial. Geographic distance and host shift between pairs of populations predict genomic divergence, with geographic distance and acoustical signal together explaining > 60% of the divergence among populations. Differences in calling songs could reflect adaptation of populations to novel environments with different host plants, habitats and predators, which may have resulted from neutral divergence at the molecular level followed by natural selection. Geomorphic barriers and climate oscillations associated with Pleistocene glaciation may have been primary factors in shaping the population genetic structure of this species. Ultimately this may couple with a host shift in leading toward allopatric speciation in S. yangi, i.e., isolation by distance. Our findings improve understanding of divergence in allopatry of herbivorous insects, and may inform future studies on the molecular mechanisms underlying the association between genetic/phenotypic changes and adaptation of insects to novel niches and host plants.

The First Three Mitochondrial Genomes for the Characterization of the Genus Egeirotrioza (Hemiptera: Triozidae) and Phylogenetic Implications.

(1) Background: Mitochondrial genomes are important markers for the study of phylogenetics and systematics. Triozidae includes some primary pests of Populus euphratica. The phylogenetic relationships of this group remain controversial due to the lack of molecular data. (2) Methods: Mitochondria of Egeirotrioza Boselli were sequenced and assembled. We analyzed the sequence length, nucleotide composition, and evolutionary rate of Triozidae, combined with the 13 published mitochondrial genomes. (3) Results: The evolutionary rate of protein-coding genes was as follows: ATP8 > ND6 > ND5 > ND2 > ND4 > ND4L > ND1 > ND3 > APT6 > CYTB > COX3 > COX2 > COX1. We reconstructed the phylogenetic relationships of Triozidae based on 16 triozid mitochondrial genomes (thirteen ingroups and three outgroups) using the maximum likelihood (ML) and Bayesian inference (BI) approaches. The phylogenetic analysis of the 16 Triozidae mitochondrial genomes showed that Egeirotrioza was closely related to Leptynoptera. (4) Conclusions: We have identified 13 PCGs, 22 tRNAs, 2 rRNAs, and 1 control region (CR) of all newly sequenced mitochondrial genomes, which were the mitochondrial gene type in animals. The results of this study provide valuable genomic information for the study of psyllid species.

Disparities in Genetic Diversity Drive the Population Displacement of Two Invasive Cryptic Species of the Bemisia tabaci Complex in China.

Within the whitefly Bemisia tabaci (Gennadius) (Hemiptera: Aleyrodidae) complex, two cryptic species, namely Middle East-Asia Minor 1 (MEAM1) and Mediterranean (MED), are important invasive pests affecting global agriculture and horticulture. They were introduced into China sequentially in the mid-1990s and around 2003, respectively. Subsequently, the latter invader MED has outcompeted the earlier invader MEAM1, becoming the dominant population in the field. Although extensive studies have explored the underlying mechanisms driving this shift, the contribution of population genetics remains notably underexplored. In this study, we analyzed the genetic diversity and structure of 22 MED and 8 MEAM1 populations from various regions of China using mitochondrial DNA sequencing and microsatellite genotyping. Our results indicate low and moderate levels of genetic differentiation among geographically separate populations of MED and MEAM1, respectively. Median-joining network analysis of mtCOI gene haplotypes revealed no clear geographic structuring for either, with common haplotypes observed across provinces, although MED had more haplotypes. Comparative analyses revealed that MED presented greater genetic diversity than MEAM1 on the basis of two markers. Furthermore, analysis of molecular variance supported these findings, suggesting that while some genetic variation exists between populations, a significant amount is also present within populations. These findings reveal the population genetics of the two invasive cryptic species of the B. tabaci complex in China and suggest that the disparities in genetic diversity drive the displacement of their populations in the field. This work also provides valuable information on the genetic factors influencing the population dynamics and dominance of these invasive whitefly species.

Evolutionary and phylogenetic insights from the mitochondrial genomic analysis of Diceraeus melacanthus and D. furcatus (Hemiptera: Pentatomidae).

The mitochondrial genomes of D. melacanthus and D. furcatus were sequenced and used to investigate the phylogenetic relationships with 54 species of Pentatomidae. Their mitogenomes are 17,197 and 15,444 bp-long, respectively, including 13 protein-coding genes (PCGs), 2 ribosomal RNA genes, and 22/21 transfer RNA genes, with conserved gene arrangement. Leu, Lys, and Ser were the most common amino acids in their PCGs. PCGs evolutionary analysis indicated their mitogenomes are under purifying selection, and the most conserved genes are from the cytochrome complex, reinforcing their suitability as markers for molecular taxonomy. We identified 490 mtSSRs in 56 Pentatomidae species, with large variation and a positive correlation between mtSSR number and genome size. Three mtSSRs were identified in each Diceraeus species. Only the mtSSR in the nad6 (D. melacanthus) and nad4 (D. furcatus) appear to have application as molecular markers for species characterization. Phylogenetic analysis confirmed the monophyly of Pentatomidae. However, our analysis challenged the monophyly of Pentatominae and Podopinae. We also detected unexpected relationships among some tribes and genera, highlighting the complexity of the internal taxonomic structure of Pentatomidae. Both Diceraeus species were grouped in the same clade with the remaining Carpocorini analyzed.

Horizontally transferred glycoside hydrolase 26 may aid hemipteran insects in plant tissue digestion.

Glycoside hydrolases are enzymes that break down complex carbohydrates into simple sugars by catalyzing the hydrolysis of glycosidic bonds. There have been multiple instances of adaptive horizontal gene transfer of genes belonging to various glycoside hydrolase families from microbes to insects, as glycoside hydrolases can metabolize constituents of the carbohydrate-rich plant cell wall. In this study, we characterize the horizontal transfer of a gene from the glycoside hydrolase family 26 (GH26) from bacteria to insects of the order Hemiptera. Our phylogenies trace the horizontal gene transfer to the common ancestor of the superfamilies Pentatomoidea and Lygaeoidea, which include stink bugs and seed bugs. After horizontal transfer, the gene was assimilated into the insect genome as indicated by the gain of an intron, and a eukaryotic signal peptide. Subsequently, the gene has undergone independent losses and expansions in copy number in multiple lineages, suggesting an adaptive role of GH26s in some insects. Finally, we measured tissue-level gene expression of multiple stink bugs and the large milkweed bug using publicly available RNA-seq datasets. We found that the GH26 genes are highly expressed in tissues associated with plant digestion, especially in the principal salivary glands of the stink bugs. Our results are consistent with the hypothesis that this horizontally transferred GH26 was co-opted by the insect to aid in plant tissue digestion and that this HGT event was likely adaptive.

Improving DNA barcoding library of armored scale insects (Hemiptera: Diaspididae) in China.

DNA barcoding is used to identify cryptic species, survey environmental samples, and estimate phyletic and genetic diversity. Armored scale insects are phytophagous insects and are the most species-rich taxa in the Coccoidea superfamily. This study developed a DNA barcode library for armored scale insect species collected from southern China during 2021-2022. We sequenced a total of 239 specimens, recognized as 50 morphological species, representing two subfamilies and 21 genera. Sequencing analysis revealed that the average G + C content of the cytochrome oxidase subunit I (COI) gene sequence was very low (~18.06%) and that the average interspecific divergence was 10.07% while intraspecific divergence was 3.20%. The intraspecific divergence value was inflated by the high intraspecific divergence in ten taxa, which may indicate novel species overlooked by current taxonomic treatments. All the Automated Barcode Gap Discovery, Assemble Species by Automatic Partitioning, Taxon DNA analysis and Bayesian Poisson Tree Process methods yielded largely consistent results, indicating a robust and credible species delimitation. Based on these results, an intergeneric distance threshold of ≤ 5% was deemed appropriate for the differentiation of armored scale insect species in China. This study establishes a comprehensive barcode library for the identification of armored scale insects, future research, and application.

Phylogenomics of endemic Australian Ulopinae (Hemiptera: Cicadomorpha: Cicadellidae).

Ulopinae is a distinctive subfamily of leafhoppers that is widely distributed across the Afrotropical, Palearctic, Indomalayan and Australasian regions. The ulopine fauna of Australia is entirely endemic and includes two tribes of striking appearance, the Ulopini and Cephalelini. Knowledge of these groups is fragmentary and in many instances, no information is available beyond original descriptions. We assess the monophyly, phylogenetic placement and species-level diversity of the Ulopini genus Austrolopa . Phylogenetic analyses based on sequence data from target nuclear loci (18S , 28S , H2A and H3 ) and mitochondrial genomes (15 genes) for 23 membracoid taxa yielded congruent topologies. Our results provide strong evidence for the monophyly of Ulopinae and a clade consisting of Ulopini + Cephalelini. However, a non-monophyletic Cephalelini arises from within a polyphyletic Ulopini. Austrolopa was strongly recovered as monophyletic in all analyses, a result also supported by morphological features. The genus currently includes six species, three of which are described based on morphological and molecular data: Austrolopa botanica , sp. nov. , Austrolopa rotunda , sp. nov. and Austrolopa sublima , sp. nov. A lectotype designation is provided for Austrolopa kingensis Evans, 1937, sp. reval. Our findings illustrate that the Australian Ulopinae is far more diverse than currently circumscribed and several species of Austrolopa are yet to be recognised. ZooBank: urn:lsid:zoobank.org:pub:1480285B-8F61-4659-A929-2B1EF3168868.

Revisionary notes on the genus Adelungia Melichar, 1902 (Hemiptera: Cicadellidae, Megophthalminae), with the description of a new genus.

Leafhopper genus Adelungia Melichar, 1902 is reviewed. Spathicerula Gnezdilov gen. nov. is described with Adelungia calligoni Oshanin, 1908 as the type species, and a new combination-Spathicerula calligoni (Oshanin, 1908), comb. nov., is proposed. Adelungia tridigitata Dlabola, 1984 is placed in synonymy under A. calligoni. Lectotypes are designated for Adelungia elegans Melichar, 1902 and A. breviceps Kusnezov, 1931 to stabilize the nomenclature in the group according to the International Code of Zoological Nomenclature.

Toward a new classification of planthoppers Hemiptera Fulgoromorpha: 2. Higher taxa, their names and their composition.

A synthesis of planthopper suprafamilial and familial classification is provided and discussed, solving some nomenclatural issues for the group, and adopting an enriched presentation with a clear distinction between the taxon name and its concept.

A new genus and species of giant mealybugs (Hemiptera: Coccomorpha: Monophlebidae) from eastern China.

A new genus of giant mealybugs (Hemiptera: Coccomorpha; Monophlebidae), Coronaproctus Li, Xu & Wu gen. nov., is established for the species Coronaproctus castanopsis Li, Xu & Wu sp. nov., collected from Zhejiang province, China on the twigs of Castanopsis eyrei (Fagaceae). The adult female and male are described and illustrated. The new genus is characterized by the adult female possessing quadrilocular disc pores, 13 cicatrices on the venter, and a U-shaped opening of the marsupium, but lacking large bilocular pores.

A new species of planthopper from Costa Rica in the genus Herpis (Hemiptera: Derbidae) from palms in the cloud forest.

Recent survey efforts in Costa Rica have documented many new species of planthoppers, primarily in the families Derbidae and Cixiidae, on palms. Recently, a specimen was collected sweeping palms in the Los Angeles cloud forest in Costa Rica and was identified as belonging to the genus Herpis (Derbidae). It was subsequently determined to represent a previously undescribed species. Herein, the new species, Herpis circumsoros Bahder & Bartlett sp. n. is described with supplemental molecular data for the cytochrome c oxidase subunit I (COI) gene and 18S rRNA gene to support placement of the new species in the genus Herpis.

A new genus, Duffelsa gen. n., with descriptions of three new species and one new combination (Hemiptera: Cicadidae).

One new cicada genus, Duffelsa gen. n., is erected in the tribe Leptopsaltriini Moulton, 1923, with the designation of Pomponia orientalis (Distant, 1912) as the type species. Three new species, D. parvula sp. n., D. grandia sp. n. and D. rubida sp. n., are described from China. A key to species of Duffelsa gen. n. is provided. The relationship of this new genus to related taxa is discussed.

Review of Cyphotes Burmeister, 1835 (Hemiptera: Membracidae) with the description of a related new genus.

The Neotropical treehopper genus Cyphotes Burmeister, 1835 (= Aspona Stal, 1862 syn. nov.) is redefined. Cyphotes contains only two species, the type species Cyphotes nodosa Burmeister, 1835 (= Aspona bullata Stl, 1862 syn. nov.) and Cyphotes quadrinodosa (Fonseca & Diringshofen, 1969) reinstated. comb. Allocyphotes gen. nov. (type species Cyphotes insolita Goding, 1929) is proposed to accommodate two other species previously placed in Cyphotes, Allocyphotes pompanoni (Boulard, 2011) comb. nov. and Allocyphotes colombiensis (Gonzlez-Mozo, 2017) comb. nov., and three new species from Ecuador: A. flavus sp. nov., A. waoraniorum sp. nov and A. robertoi sp. nov. for a total of six species. Illustrations, including genitalia images, new locality records and keys to genera and species are provided.

A new species of planthopper in the genus Anchimothon (Hemiptera: Auchenorrhyncha: Derbidae) on palms from Costa Rica.

Anchimothon is a small genus of derbid planthoppers known from Mesoamerica and northern South America. Recent survey efforts in the Caribbean basin have documented many new taxa of fulgoroids from palms. During this survey, a novel taxon identified as Anchimothon was collected from palm seedlings at La Selva Biological Station in Costa Rica. Here, the novel taxon is described as A. myriei sp. n. with molecular data for the cytochrome c oxidase subunit I (COI) and18S rRNA providing support for placement of the new species in Anchimothon. An updated key is provided for the current species of Anchimothon.

Microbiome analyses of 12 psyllid species of the family Psyllidae identified various bacteria including Fukatsuia and Serratia symbiotica, known as secondary symbionts of aphids.

BACKGROUND: Psyllids (Hemiptera: Psylloidea) comprise a group of plant sap-sucking insects that includes important agricultural pests. They have close associations not only with plant pathogens, but also with various microbes, including obligate mutualists and facultative symbionts. Recent studies are revealing that interactions among such bacterial populations are important for psyllid biology and host plant pathology. In the present study, to obtain further insight into the ecological and evolutionary behaviors of bacteria in Psylloidea, we analyzed the microbiomes of 12 psyllid species belonging to the family Psyllidae (11 from Psyllinae and one from Macrocorsinae), using high-throughput amplicon sequencing of the 16S rRNA gene. RESULTS: The analysis showed that all 12 psyllids have the primary symbiont, Candidatus Carsonella ruddii (Gammaproteobacteria: Oceanospirillales), and at least one secondary symbiont. The majority of the secondary symbionts were gammaproteobacteria, especially those of the family Enterobacteriaceae (order: Enterobacteriales). Among them, symbionts belonging to "endosymbionts3", which is a genus-level monophyletic group assigned by the SILVA rRNA database, were the most prevalent and were found in 9 of 11 Psyllinae species. Ca. Fukatsuia symbiotica and Serratia symbiotica, which were recognized only as secondary symbionts of aphids, were also identified. In addition to other Enterobacteriaceae bacteria, including Arsenophonus, Sodalis, and "endosymbionts2", which is another genus-level clade, Pseudomonas (Pseudomonadales: Pseudomonadaceae) and Diplorickettsia (Diplorickettsiales: Diplorickettsiaceae) were identified. Regarding Alphaproteobacteria, the potential plant pathogen Ca. Liberibacter europaeus (Rhizobiales: Rhizobiaceae) was detected for the first time in Anomoneura mori (Psyllinae), a mulberry pest. Wolbachia (Rickettsiales: Anaplasmataceae) and Rickettsia (Rickettsiales: Rickettsiaceae), plausible host reproduction manipulators that are potential tools to control pest insects, were also detected. CONCLUSIONS: The present study identified various bacterial symbionts including previously unexpected lineages in psyllids, suggesting considerable interspecific transfer of arthropod symbionts. The findings provide deeper insights into the evolution of interactions among insects, bacteria, and plants, which may be exploited to facilitate the control of pest psyllids in the future.

New genera and species of felt scales (Hemiptera: Coccomorpha: Eriococcidae), with descriptions of new species and immature instars of described species.

In this work, emphasis is placed on the Eriococcidae (Hemiptera: Coccomorpha) of the Nearctic Region including the immature instars of many species not previously described, two new genera (Carphococcus Miller and Stocks and Ypofloiococcus Miller and Stocks), and 11 new species (Acanthococcus pennyae Miller and Stocks, Carph. apodus Miller and Stocks, Carpochloroides eugeniae Miller and Stocks, Ovaticoccus betsyae Miller and Stocks, Ov. gordoni Miller and Stocks, Ov. haigi Miller and Stocks, Ov. maryfoleybensonae Miller and Stocks, Ov. telotrichus Miller and Stocks, Ov. tuttlei Miller and Stocks, Ov. villanuevorum Miller and Stocks and Y. libeauae Miller and Stocks). In addition to the new taxa, descriptions and illustrations are provided for adult females, adult males and/or immature instars of Ac. arenosus (Cockerell), Ac. coccineus (Cockerell), Ac. dennoi Miller and Miller, Ac. gallicolus (Cockerell & Rohwer), Ac. hoyi Miller and Miller, Ac. kemptoni (Parrott), Apezococcus idiastes Ferris, Carpo. mexicanus Ferris, Cryptococcus fagisuga Lindinger, Cr. williamsi Kosztarab and Hale, Hypericicoccus hyperici (Ferris), Ov. agavacearum Pellizzari and Kozár, Ov. agavium (Douglas), Ov. cornutus (Ferris), Ov. densus (Miller), Ov. eriogoni (Miller), Ov. exoticus Pellizzari and Kozár, Ov. neglectus (Cockerell), Ov. peruvianus (Granara de Willink & Díaz), Ov. salviae Miller, Ov. sanguineus (Miller), Ov. tippinsi Miller and Miller, Ov. variabilis Miller, Spiroporococcus braggi (Cockerell & Robinson), S. yuccae (Ferris) and Xerococcus foquieriae Ferris. Lectotype designations are made for Ap. idiastes. Carpo, mexicanus and H. hyperici. New collection data are provided for Ov. californicus McKenzie, Ov. johnsoni (Miller), Ov. mackenziei Miller, Ov. parvispina (Chaffin) and Ov. viscosa (Kondo). Dichotomous keys are given for: determination of most eriococcid instars of both sexes; Nearctic genera of the Eriococcidae based on adult females; adult females, second-instar females, second-instar males, and first-instar nymphs of Ovaticoccus Kloet 1944; and the seven species of Acanthococcus Signoret 1875 treated here, based on first-instar nymphs. Oregmopyga Hoy 1963 and Cornoculus Ferris 1955 are treated as junior subjective synonyms of Ovaticoccus. Atriplicia Cockerell and Rohwer 1909 has been placed previously as a junior synonym of Eriococcus but here is regarded as a junior subjective synonym of Acanthococcus. We treat 44 species; new descriptions are provided for 101 treated instars, and detailed line drawings are given for all but three of them. The following species, listed in ScaleNet as occurring in the Nearctic Region, are transferred to Acanthococcus from Eriococcus Targioni Tozzetti 1868: Acanthococcus actius Miller and Miller 1993 rev. comb.; Ac. arenariae Miller and Miller 1993 rev. comb.; Ac. barri Miller 1991 rev. comb.; Ac. beshearae Miller and Miller 1993 rev. comb.; Ac. dennoi Miller and Miller 1993 rev. comb.; Ac. droserae Miller, Liu and Howell 1992 rev. comb.; Ac. epacrotrichus Miller and Miller 1992 rev. comb.; Ac. froebeae Miller 1991 rev. comb.; Ac. gallicolus (Cockerell & Rohwer 1909) comb. n.; Ac. hoyi Miller and Miller 1992 rev. comb.; Ac. mackenziei Miller and Miller 1992 rev. comb.; Ac. macrobactrus Miller and Miller 1992 rev. comb.; Ac. megaporus Miller and Miller 1993 rev. comb.; A. mesotrichus Miller and Miller 1993 rev. comb.; Ac. microtrichus Miller and Miller 1992 rev. comb.; Ac. monotrichus Miller and Miller 1993 rev. comb.; Ac. ophius Miller and Miller 1993 rev. comb.; Ac. stauroporus Miller and Miller 1992 rev. comb.; Ac. washingtonensis Miller and Miller 1992 rev. comb.; and Ac. whiteheadi Miller 1991 rev. comb. In addition, two species are transferred to Acanthococcus from Rhizococcus Signoret: Ac. coccineus (Cockerell 1894a) rev. comb. and Ac. kemptoni (Parrott 1900) rev. comb. Three images of water color paintings by Mary Foley Benson are included.

Five new Brazilian species of Fuminana Freytag (Hemiptera: Cicadellidae: Gyponini) with new records and key to males.

Five new Brazilian species of Fuminana Freytag, 1989 are described and illustrated: F. diaphana sp. nov. from states of Maranhão and Piauí, F. guaricica sp. nov. from Paraná State, F. incurvata sp. nov. and F. retorta sp. nov. from Rondônia State, and F. niluga sp. nov. from states of Acre, Pará, Rondônia and Mato Grosso. The female genitalia is described and illustrated for the first time and a revised genus description is presented. New distribution records are given and seven species are newly recorded from Brazil: F. astra (DeLong & Freytag, 1969), F. conspicua (DeLong & Freytag, 1969), Fuminana curra (DeLong & Freytag, 1969), F. extata Freytag, 1989, F. jaculum (DeLong & Freytag, 1969), F. signa (DeLong & Freytag, 1969) and F. superba (DeLong & Freytag, 1969). A key to males is provided as well as some taxonomic notes about the genus.