On the fauna of chironomidae (Diptera) of the Hrazdan Basin (Armenia)

A faunistic review of larvae and pupae of chironomid midges (Diptera, Chironomidae) from 15 habitats in the Hrazdan River valley (Armenia) is presented. The revealed fauna includes 40 species from 20 genera and 6 families and tribes. Diploid chromosome numbers are specified for 24 species from 13 genera and 4 subfamilies; 23 species from 14 genera and 4 subfamilies are reported from the region for the first time. Predominant oligotrophic status is confirmed for most of the reservoirs studied.     

Larval morphology enhances phylogenetic reconstruction in Cetoniidae (Coleoptera: Scarabaeoidea) and allows the interpretation of the evolution of larval feeding habits

Abstract.  The Cetoniidae, the showiest of scarabs, comprises some 3900 species in 515 genera, distributed worldwide except for subpolar areas and some offshore New Zealand islands. Parsimony analysis of 76 larval and adult characters and 42 terminal taxa supports the monophyly of Cetoniidae (sensu Krikken, 1984 ), but not of the traditionally considered subfamilies Cetoniinae and Trichiinae (sensu Krikken, 1984 ). In the study taxon, larval characters are shown to be more informative than those of adults for deeper phylogeny. The evolution of some larval characters (head and legs) in relation to feeding habits is discussed on the basis of phylogenetic analysis. The results show an evolutionary shift from wood associations to a higher ecological plasticity that allows the larvae to feed on wide sources of organic matter (including compost, dung, gopher burrows, packrat middens, ant debris piles, etc.).     

Phylogeny of psychodid subfamilies (Diptera: Psychodidae) inferred from nuclear DNA sequences with a review of morphological evidence for relationships

Psychodidae is a diverse family of flies with approximately 3000 described species in six subfamilies, including Phlebotominae vectors of human disease. Psychodidae has been the subject of few phylogenetic investigations and development of a stable classification has been hampered by poor understanding of the morphology of larvae, pupae and adults. Specimens were collected, and we analysed DNA sequence data from two nuclear genes for one or more representatives of all subfamilies. The subfamilies with multiple representatives included were resolved as monophyletic with good support. Placement of Horaiellinae, Sycoracinae and Trichomyiinae remains unclear, whereas Bruchomyiinae is hypothesized as the sister group to (Phlebotominae + Psychodinae). Representatives of some psychodine tribes were resolved in agreement with previous hypotheses. Relationships among and within subfamilies are discussed, and morphological characters supporting these relationships are reviewed. One compelling synapomorphy of the male genitalia supporting a relationship between Phlebotominae and Psychodinae is the presence of articulated surstyli with apical retinacula. Only cerci are present and sometimes developed into clasping structures in males of other subfamilies.     

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.     

Midges of the subfamily Tanypodinae (Diptera,Chironomidae) of Russia: Tribes Coelotanypodini Kieffer, 1913 and Tanypodini Meigen, 1803

It is established that in the territory of Russia the subfamily Tanypodinae (Diptera, Chironomidae) includes 80 species belonging to 10 subgenera of 8 genera and 5 tribes; 4 species are recorded for the first time in the Palaearctic Region, and 16 species, in Russia. In the present study, larvae, pupae, and male adults from different areas of Russia, belonging to 9 species of 2 subgenera, 2 genera, and 2 tribes, are examined. Keys to larvae, pupae, and male adults of these species is provided.     

A phylogenetic analysis of Sciomyzidae (Diptera) and some related genera

Sciomyzidae is a family of acalyptrate flies with 546 species in 61 genera that is among the most extensively studied groups of higher Diptera. Most of the known larvae are obligate enemies of Gastropoda. Hundreds of studies published over the past 50 years have resulted in detailed information concerning morphology of adults and immature stages, biology, development, behaviour, phenology and distribution. However, studies of phylogenetic relationships are based almost exclusively on morphological characters of adults, and no comprehensive molecular analysis across the family has been published. Here we fill this void by generating and analysing molecular data for 54 species of Sciomyzidae (22 genera), including Phaeomyiidae (one genus), and seven representative species of five other families of Sciomyzoidea (Coelopidae, Dryomyzidae, Helcomyzidae, Heteromyzidae and Huttoninidae) as outgroups. The reconstruction is based on morphological characters as well as nucleotide sequences for genes from the mitochondrial (12S, 16S, COI, COII, Cytb) and nuclear genome (28S, EF1α). The results are compared with recent morphological analyses. Our analyses support the monophyly of Sciomyzidae + Phaeomyiidae, and place Phaeomyiinae as a unique lineage within Sciomyzidae. A modified classification comprising three subfamilies is proposed. The major subfamily, Sciomyzinae, consists of two monophyletic and well separated groups, the tribes Sciomyzini and Tetanocerini.     

Monophyly of Euaesthetinae (Coleoptera: Staphylinidae): phylogenetic evidence from adults and larvae,review of austral genera,and new larval descriptions

Abstract We develop a morphological dataset for the rove beetle subfamily Euaesthetinae comprising 167 morphological characters (135 adult and 32 larval) scored from 30 terminal taxa including 25 ingroup terminals (from subfamilies Euaesthetinae and Steninae) and five outgroups. Four maximum parsimony analyses using different sets of terminals and character sets were run to test the monophyly of (1) Euaesthetinae, (2) Steninae, (3) Euaesthetinae + Steninae, (4) euaesthetine tribes Austroesthetini, Alzadaesthetini, Euaesthetini, Fenderiini and Stenaesthetini, and (5) the ten currently known austral endemic genera together. Analyses of adult and larval character sets separately and in combination recovered the monophyly of Euaesthetinae, Steninae, and both subfamilies together, with strong support. Analysis of 13 ingroup terminals for which complete data were available suggests that monophyly of Euaesthetinae is supported by 19 synapomorphies (13 adult, six larval), of Steninae by 23 synapomorphies (14 adult, nine larval), and of both subfamilies together by 24 synapomorphies (21 adult, three larval). Within Euaesthetinae, only the tribe Stenaesthetini was recovered as monophyletic based on adult characters, and in no analyses were the ten austral endemic genera recovered as a monophyletic group. Phylogenetic relationships among euaesthetine genera were weakly supported, although analyses including adult characters supported monophyly of Octavius and Protopristus separately, and of Octavius + Protopristus, Austroesthetus + Chilioesthetus and Edaphus + Euaesthetus. Steninae may include a third genus comprising two undescribed species probably possessing a ‘stick–capture’ method of prey capture, similar to that in Stenus. These two species formed a strongly supported clade recovered as the sister group of Stenus based on adult characters. Diagnoses and a key to adults are provided for the 15 euaesthetine genera currently known from the austral region (Australia, New Zealand, South Africa and southern South America). Euaesthetine larvae previously were known only for Euaesthetus, and we describe the larvae of nine more genera and provide the first larval identification key for genera of Euaesthetinae.     

Larvae of Trichophya and phylogeny of the tachyporine group of subfamilies (Coleoptera: Staphylinidae) with a review, new species and characterization of the Trichophyinae

Abstract. Larvae of the staphylinid subfamily Trichophyinae are described for the first time based on larvae of a new species of Trichophya from the southwestern United States. Adults and larvae of the new species, Trichophya texana Ashe & Newton (type locality Texas, Brewster Co., Big Bend National Park), are described and illustrations of both provided. Also given are a key for separation of the Nearctic species of Trichophya , a checklist of the known World fauna of the Trichophyinae (including first report of the genus from Mexico and Guatemala), and a characterization of the subfamily Trichophyinae based on both larvae and adults. The relationships of major genera and higher taxa in the tachyporine group of staphylinid subfamilies are analysed cladistically using larval characters. No larval characters were found that provide evidence for the monophyly of the tachyporine group; no evidence was found for the monophyly of the Tachyporinae; Charhyphus, Olisthaerus and Phloeocharis (Phloeocharinae + Olisthaerinae) form a monophyletic group; the Trichophyinae and Habrocerinae are sister groups and together probably are the sister group to the Aleocharinae; the Aleocharinae are confirmed to be monophyletic based on larval characters; and Gymnusa + Deinopsis form the sister group to the remainder of the Aleocharinae.     

The family and subfamily classification and New Zealand genera of Pythidae and Scraptiidae (Coleoptera)

ABSTRACT. The family Pythidae is defined on adult and larval characters to include the subfamilies Pythinae and PllipalPlnae. Trachelostenidae stat.n. is excluded. Relationships of Pythidae, Boridae, Trictenotomidae and SalPlngidae (s.l.) are analysed. Strong resemblances between some PllipalPlnae and Pyrochroidae appear to have arisen through convergent evolution. Adults and larvae of the New Zealand genera Techmessa, Techmessodes and Exocalopus are described. In Scraptiidae, adults of the New Zealand genera Nothotelus and Phytilea , and the larva of Nothotelus , are described. Type data are given for all described New Zealand species of these families. The following new synonymies are established: PllipalPlnae (Abdullah, 1964)=Techmessinae Paulus, 1971. Exocalopus pectinatus Broun, 1893= nitidiceps Broun, 1910. Techmessa concolor Bates, 1874= attenuata Broun, 1893= rugicollis Broun, 1910= unicolor Paulus, 1971. Techmessa telephoroides Bates, 1874= varians Broun, 1893. Techmessodes Plcticornis (Broun, 1880)= distans (Sharp, 1882). Techmessodes versicolor Broun, 1893= cephalotes Broun, 1910. The following genera are reassigned to the families indicated: Ischyomius to Trictenotomidae (from Pythidae or Melandryidae). Phytilea to Scraptiidae (from Oedemeridae or Anthicidae). Pseudananca to Aderidae (from Oedemeridae). Scraptogetus (= Metasclera ) to Aderidae (from Scraptiidae, Oedemeridae or Anthicidae).     

Higher level phylogeny for the passion-vine butterflies (Nymphalidae, Heliconiinae) based on early stage and adult morphology

A higher level phylogeny for the passion-vine butterflies (Nymphalidae, Heliconiinae) was generated by cladistic analysis of 146 morphological characters from all life stages. The 24 species studied were selected representatives of the ten currently accepted genera of the sub-tribe Heliconiiti. Analyses of only characters from larvae and pupae did not produce well resolved trees. However, some characters of the immature stages provided critical support for the monophyly of two clades. Analysis of only adult characters yielded a tree that closely resembled that obtained from all data combined. The phylogeny here derived from the combined analysis of early stage and adult characters differed in topology from all previously proposed hypotheses, and supported the monophyly of all currently recognized genera. Characters supporting each clade are described and illustrated, and various hypotheses of phylogenetic relatedness of passion-vine butterfly taxa are discussed.     

Phylogeny and early evolution of the Cynipoidea (Hymenoptera)

Based on several structural and biological characteristics, the Cynipoidea can be divided into two groups, 'macrocynipoids' and 'microcynipoids'. The macrocynipoids (i.e. the family Liopteridae and the genera Austrocynips, Eileenella, Heteribalia and Ibalia ) are generally large insects that parasitize wood- or cone-boring insect larvae. The microcynipoids are smaller insects that are either phytophagous gall inducers and inquilines (Cynipidae) or parasitoids of larvae of Hymenoptera, Neuroptera or Diptera (Figitidae sensu lato , including the families Eucoilidae, Charipidae and Anacharitidae). The phylogenetic relationships among genera of macrocynipoids and between these and a sample of four genera representing the Figitidae and Cynipidae were examined by parsimony analysis of 110 external morphological characters of adults. Within the macrocynipoids, three monophyletic lineages emerged, classified here as different families: the Austrocynipidae, with a single species, Austrocynips mirabilis , the only cynipoid with a true pterostigma; the Ibaliidae, including the genera Eileenella, Ibalia and Heteribalia ; and the Liopteridae, comprising the remaining genera of macrocynipoids. The analysis further supported the monophyly of the microcynipoids and indicated that the macrocynipoids form a paraphyletic group relative to the microcynipoids, with the shortest tree suggesting the relationship (Austrocynipidae, (Ibaliidae, (Liopteridae, microcynipoids))). These results imply that cynipoids were originally parasitoids of wood-boring insect larvae and that the other modes of life evolved secondarily within the group.     

Higher Level Phylogeny of Curculionidae (Coleoptera: Curculionoidea) based mainly on Larval Characters, with Special Reference to Broad-Nosed Weevils

A cladistic analysis of Curculionidae was performed using 49 characters (41 from larvae, three from pupae, and five from adults). Illustrations of characters of immatures are provided. The analysis involved 19 terminal units and a hypothetical ancestor determined by the outgroup comparison method used to root the tree. One most parsimonious cladogram was obtained based on the complete data set and the following phylogenetic hypothesis is proposed: Ithycerinae, Microcerinae, and Brachycrinae sensu stricto are broad-nosed weevils placed sequentially at the base of the cladogram. The remaining weevil subfamilies form two major natural groups: one constituted by the sister taxa Rhynchophorinae—Platypodinae; the other with Erirhininae at the base, as sister taxon of the "Curculionidae sensu stricto " which show an unresolved trichotomy involving Curculioninae, Cossoninae—Scolytinae, and the clade including the Entiminae and allied subfamilies. This latter clade of broad-nosed weevils has Thecesterninae at the base; the next branch is Amycterinae, the sister taxon of the clade comprising two groups: one constituted by Aterpinae, Rhytirrhininae, and Gonipterinae; the other is Entiminae whose units form two main clades: one constituted by the sister tribes Pachyrhynchini—Ectemnorhinini, and the other by Alophini, Sitonini, and Entimini. When the analysis was done using only immature characters, results congruent with those based on the complete data set were obtained, except for the placement of Erirhininae. According to the results the hypothesis of monophyly of broad-nosed weevils is not accepted; the Entiminae are justified as monophyletic and their natural classification into tribes is proposed and the phylogenetic position and relationships of higher taxa of Curculionidae are discussed. This paper shows the importance of immature characters in recognition of natural groups and relationships in Curculionidae.     

A revised subfamily classification of Tenebrionidae (Coleoptera)

Existing classifications of Tenebrionidae are reviewed briefly. The inclusion of the families Alleculidae, Lagriidae, and Nilionidae in Tenebrionidae is confirmed. The splitting off from this complex of a family, Tentyriidae, by Doyen is discussed and rejected. Various taxa which had been included in Tenebrionidae are excluded, amongst which Syrphetodes, Brouniphylax, Exohadrus, Arthopus, Cotulades, Docalis, and Latometus (=Elascus) have not previously been formally excluded. A new family, Archeocrypticidae, is established and defined briefly for Archeocrypticus, Sivacrypticus, and Enneboeus.

Data from matrices based on adult and larval characters comparing Tenebrionidae with most other families of Tenebrionoidea (=Heteromera) are presented for derived characters in common, and for overall similarity. The families most closely related to Tenebrionidae according to these data are Zopheridae, Chalcodryidae, Merycidae, Archeocrypticidae, Synchroidae, Colydiidae, and Monommatidae; none is very close to Tenebrionidae, which has had a long independent history.

Characters of the subfamilies recognised are tabulated, and interpreted in a phylo‐genetic dendrogram. Phylogeny is discussed in relation to adaptive changes in the biology of the various subfamilies, which are Zolodininae new subfamily, Pimeliinae new sense (including Tentyriinae), Toxicinae new sense, Phrenapatinae new sense (including Archeoglenini new tribe), Diaperinae new sense, Gnathidiinae, Tenebrioninae new sense, Alleculinae, Nilioninae, Lagriinae new sense, Cossyphinae, and Cossyphodinae new status.

Biology, economic importance, copulation, orientation of the aedeagus, and distribution are discussed briefly.

Definitions of the family and subfamilies and a key to subfamilies are given, and keys to tribes are included for the smaller subfamilies. The previously unknown larvae of the genera Zolodinus, Menimus, Archeoglenes, Lepispilus, and Nyctoporis are described in detail. Pupae of Zolodinus and Nyctoporis are described. Keys to larvae include many other genera which were hitherto unknown or poorly known.     

Phylogeny of Myrmeleontiformia based on larval morphology (Neuropterida: Neuroptera)

The suborder Myrmeleontiformia is a derived lineage of lacewings (Insecta: Neuroptera) including the families Psychopsidae, Nemopteridae, Nymphidae, Ascalaphidae and Myrmeleontidae. In particular, Myrmeleontidae (antlions) are the most diverse neuropteran family, representing a conspicuous component of the insect fauna of xeric environments. We present the first detailed quantitative phylogenetic analysis of Myrmeleontiformia, based on 107 larval morphological and behavioural characters for 36 genera whose larvae are known (including at least one representative of all the subfamilies of the suborder). Four related families were used as outgroups to polarize character states. Phylogenetic analyses were conducted using both parsimony and Bayesian methods. The reconstructions resulting from our analyses corroborate the monophyly of Myrmeleontiformia. Within this clade, Psychopsidae are recovered as the sister family to all the remaining taxa. Nemopteridae (including both subfamilies Nemopterinae and Crocinae) are recovered as monophyletic and sister to the clade comprising Nymphidae + (Myrmeleontidae + Ascalaphidae). Nymphidae consist of two well‐supported clades corresponding to the subfamilies Nymphinae and Myiodactylinae. Our results suggest that Ascalaphidae may not be monophyletic, as they collapse into an unresolved polytomy under the Bayesian analysis. In addition, the recovered phylogenetic relationships diverge from the traditional classification scheme for ascalaphids. Myrmeleontidae are reconstructed as monophyletic, with the subfamilies Stilbopteryginae, Palparinae and Myrmeleontinae. We retrieved a strongly supported clade comprising taxa with a fossorial habit of the preimaginal instars, which represents a major antlion radiation, also including the monophyletic pit‐trap building species.     

Dactylogyridae 2022: a meta-analysis of phylogenetic studies and generic diagnoses of parasitic flatworms using published genetic and morphological data

Dactylogyridae is one of the most studied families of parasitic flatworms with more than 1000 species and 166 genera described to date including ecto- and endoparasites. Dactylogyrid monogeneans were suggested as model organisms for host-parasite macroevolutionary and biogeographical studies due to the scientific and economic importance of some of their host lineages. Consequently, an array of phylogenetic research into different dactylogyrid lineages has been produced over the past years but the last family-wide study was published 16 years ago. Here, we provide a meta-analysis of the phylogenetic relationships of Dactylogyridae including representatives of all genera with available molecular data (n = 67). First, we investigate the systematic informativeness of morphological characters widely used to diagnose dactylogyrid genera through a parsimony analysis of the characters, character mapping, and phylogenetic comparative methods. Second, we provide an overview of the current state of the systematics of the family and its subfamilies, and summarise potentially poly- and paraphyletic genera. Third, we elaborate on the implications of taxonomic, citation, and confirmation bias in past studies. Fourth, we discuss host range, biogeographical, and freshwater-marine patterns. We found two well-supported macroclades which we assigned to the subfamilies Dactylogyrinae and Ancyrocephalinae. These subfamilies further include 16 well-supported clades with only a few synapomorphies that could be deduced from generic diagnoses in the literature. Furthermore, few morphological characters considered systematically informative at the genus level display a strong phylogenetic signal. However, the parsimony analysis suggests that these characters provide little information on the relationships between genera. We conclude that a strong taxonomic bias and low coverage of DNA sequences and regions limit knowledge on morphological and biogeographical evolutionary patterns that can be inferred from these results. We propose addressing potential citation and confirmation biases through a ‘level playing field’ multiple sequence alignment as provided by this study.     

小粉虫各发育期的临界热极值和失水的研究

本文研究了升高环境温度对小粉虫(Alphitobiusdiaperinus,鞘翅目拟步行虫总科)末龄幼虫、蛹以及成虫的失水和热敏性(临界热极值,Criticalthermalmaximum,CTmax)的影响。小粉虫成虫和蛹的CTmax值显著低于末龄幼虫(末龄幼虫:CTmax=48.5±0.5℃;蛹:CTmax=48.0±0.9℃;成虫:CTmax=47.8±1.1℃)。此外,成虫的适应性对其CTmax没有显著的影响。在20至60℃连续记录失水值中,三个龄期之间存在显著差异。随着超过临界点温度(约为40℃)的快速升高,末龄幼虫与成虫的失水程度相近。蛹具有明显低的失水速率,即使在温度高于40℃的情况下。当温度升高10℃时(25-35℃),会造成小粉虫蛹和成虫失水增加2.0倍和2.6倍,分别为:Q10=2.05±0.70和2.49±1.31,而幼虫则为3.52±1.27。当高于35-45℃时,幼虫和成虫失水增加甚快(Q10=6.85±1.90和8.51±2.32)。而蛹在高于35-45℃时失水也增加(Q10=3.76±1.83),但低于幼虫和成虫。小粉虫的蛹处于静止和不取食状态,代谢速率较低,其具有特殊结构的气孔能较好地保持关闭状态,且蛹的表皮由脂肪和蛋白质组成,以上诸因素可解释其失水减少的原因     

Signal through the noise? Phylogeny of the Tachinidae (Diptera) as inferred from morphological evidence

The oestroid family Tachinidae represents one of the most diverse lineages of insect parasitoids. Despite their broad distribution, diversity and important role as biological control agents, the phylogeny of this family remains poorly known. Here, we review the history of tachinid systematics and present the first quantitative phylogenetic analysis of the family based on morphological data. Cladistic analyses were conducted using 135 morphological characters from 492 species belonging to 180 tachinid genera, including the four currently recognized subfamilies (Dexiinae, Exoristinae, Phasiinae, Tachininae) and all major tribes. We used characters of eggs, first‐instar larvae and adults of both sexes. We examined the effects of implied weighting by reanalysing the data with varying concavity factors. Our analysis generally supports the subfamily groupings Dexiinae + Phasiinae and Tachininae + Exoristinae, with only the Exoristinae and the Phasiinae reconstructed as monophyletic assemblages under a wide range of weighting schemes. Under these conditions, the Dexiinae, which were previously considered a well‐established monophyletic assemblage, are reconstructed as being paraphyletic with respect to the Phasiinae. The Tachininae are reconstructed as a paraphyletic grade from which the monophyletic Exoristinae arose. The Exoristinae are reconstructed as a monophyletic lineage, but phylogenetic relationships within the subfamily are largely unresolved. We further explored the evolution of oviposition strategy and found that the oviparous groups are nested within ovolarviparous assemblages, suggesting that ovipary may have evolved several times independently from ovolarviparous ancestors. This counterintuitive pattern is a novel hypothesis suggested by the results of this analysis. Finally, two major patterns emerge when considering host associations across our phylogeny under equal weights: (i) although more than 60% of tachinids are parasitoids of Lepidoptera larvae, none of the basal clades is unambiguously associated with Lepidoptera as a primitive condition, suggesting that tachinids were slow to colonize these hosts, but then radiated extensively on them; and (ii) there is general agreement between host use and monophyly of the major lineages.     

A reappraisal of the family Comasteridae A. H. Clark, 1908 (Echinodermata: Grinoidea), with the description of a new subfamily and a new genus

The family Comasteridae is divided into four subfamilies on the basis of syzygy positions in the division series and proximal arm. Use of this character results in major differences in perceived relationships between genera compared with that obtained when traditional subfamilial characters are used. The primary character previously used to determine comasterid subfamilies, cirrus ornamentation, is rejected as unimportant in recognizing taxa any higher than species. Seventeen genera, including a new genus from the tropical western Atlantic, and 99 species are now recognized in the Comasteridae. Four of these genera and five species cannot be placed within the newly diagnosed subfamilies. Characters of generic importance are considered to be pinnule comb form, mouth position and, to a lesser extent, the characters traditionally used to distinguish genera.