scholarly journals Phylogeny of Anisepyris Kieffer (Hymenoptera: Bethylidae: E pyrinae), with investigation of diagnostic features

2021 ◽  
Vol 79 ◽  
pp. 189-204
Author(s):  
Diego N. Barbosa
Keyword(s):  
Cladistic Analysis ◽  
Morphological Characters ◽  
Neotropical Region ◽  
Diagnostic Features ◽  
Dense Forest ◽  
Original Distribution ◽  
Mandible Shape ◽  
Recent Origin ◽  
Species Groups ◽  
Transformation Series

Abstract A cladistic analysis based on 120 morphological characters and 72 OTUs was conducted for the flat wasp genus Anisepyris Kieffer. The genus is mainly Neotropical region with few Nearctic species. The analysis retrieved well-supported relationships among the Anisepyris species and exposed the distribution of synapomorphies among the lineages, particularly concerning the mandible shape, dorsal pronotal area sulcus, and mesopleural foveae. The main diagnostic characters, described by Barbosa and Azevedo (2018) for the 13 species-groups of Anisepyris, were scrutinized. Transformation series and evolutionary hypotheses were also discussed. This discussion elucidates the importance of each character in the evolution and diversification along the different lineages. A discussion on the species-groups distribution is proposed, in relation to their diagnostic features. A hypothesis about the original distribution and the subsequent diversifications was also discussed. In conclusion, due to a possible recent origin for Anisepyris species, the dispersion could be related primarily to dense forest areas; and the retrieved synapomorphies could be related to posterior distributions of the species in disparate areas from their original environment.

Morphotaxonomic revision of Simulium (Gomphostilbia) (Diptera: Simuliidae) in the Oriental Region

Zootaxa ◽  
2012 ◽  
Vol 3577 (1) ◽  
pp. 1 ◽  
Author(s):  
HIROYUKI TAKAOKA
Keyword(s):  
Geographical Distribution ◽  
Classification Scheme ◽  
Cladistic Analysis ◽  
Morphological Characters ◽  
Species Group ◽  
Morphological Identification ◽  
Oriental Region ◽  
Preliminary Attempt ◽  
The Third ◽  
Species Groups

Simulium (Gomphostilbia) Enderlein, the third largest (206 named species included) in the genus Simulium Latreille s. l., is one of the two most abundant and diverse subgenera in the Oriental Region. To provide a classification scheme to facilitate morphological identification of the species within this subgenus, its diagnostic characters are redefined, and nine known species-groups within it are reviewed. Based on putative lineages explored by using certain adult morphological characters, seven more species-groups are proposed: asakoae, darjeelingense, epistum, gombakense, heldsbachense, hemicyclium and palauense species-groups, while the trirugosum species-group is merged in the varicorne species-group. Subgroups are also introduced to represent apparently different lineages within certain species-groups based on certain pupal morphological characters: two in the banauense species-group, seven in the batoense species-group redefined, four in the ceylonicum species-group redefined, four in the epistum species-group, two in the hemicyclium species-group, two in the sherwoodi species-group and four in the varicorne species-group redefined. A new checklist of species of the subgenus Gomphostilbia, and a key to all 15 species-groups within it are provided. The eastward expansion of the geographical distribution of the subgenus Gomphostilbia is inferred on the basis of the more frequent occurrence of apomorphic characters of certain adult and pupal morphological features in insular species-groups than in continental species-groups. A preliminary attempt using a cladistic analysis of morphological characters shows that among 10 subgenera examined, Gomphostilbia has a sister-taxon relationship with the Australasian subgenus Morops Enderlein, and this clade, together with the Central-Western Pacific subgenus Inseliellum Rubtsov, is positioned closest to the most derived clade formed by Daviesellum Takaoka & Adler and Simulium Latreille s. str.

Pedrowygomyia, a new Neotropical genus of Prosimuliini (Diptera: Simuliidae): Gigantodax s. lat. split into two genera

1998 ◽  
Vol 29 (2) ◽  
pp. 161-167 ◽  
Author(s):  
Daniel Rafael Miranda-Esquivel ◽  
Sixto Coscarón
Keyword(s):  
New Genus ◽  
Cladistic Analysis ◽  
Morphological Characters ◽  
Species Group ◽  
New Combinations ◽  
Species Groups

AbstractA cladistic analysis of the species groups recognized in Gigantodax Enderlein, using morphological characters, shows that Gigantodax s. lat. is paraphyletic. From a phylogenetic standing Gigantodax must be split into two monophyletic entities: Gigantodax s. str. and the G. cortesi species group. The new genus Pedrowygomyia is proposed for the G. cortesi species group. It presently contains four species, P. cortesi, P. jatunchuspi, P. punapi and P. chacabamba (all new combinations).

scholarly journals Taxonomy and phylogeny of New World Polyplectropus Ulmer, 1905 (Trichoptera: Psychomyioidea: Polycentropodidae) with the description of 39 new species

Zootaxa ◽  
2010 ◽  
Vol 2582 (1) ◽  
pp. 1 ◽  
Author(s):  
MARIA LOURDES CHAMORRO ◽  
RALPH W. HOLZENTHAL
Keyword(s):  
New Species ◽  
Bayesian Methods ◽  
New World ◽  
Morphological Characters ◽  
Neotropical Region ◽  
Species Group ◽  
Biogeographic Region ◽  
Parsimony Tree ◽  
Group A ◽  
Species Groups

The taxonomy of the New World species of Polyplectropus (Ulmer, 1905a) is revised to include detailed male and female diagnoses, descriptions, illustrations, distribution records, and keys to males of all species and species groups. A phylogenetic analysis based on 59 morphological characters, 89 of 92 (97%) New World Polyplectropus species, and 2 outgroup taxa was inferred using parsimony and Bayesian methods, which resulted in minor topological differences. Conflicting estimates of relationship among and within most species groups led to a less resolved Bayesian tree (vs. parsimony tree) due to high variation in rates of change among characters and an overall low number of characters. A new classification for New World Polyplectropus is proposed with revised characterization of 10 recognized species groups, 6 newly established. Four species remain unassigned to species group. A key to genera of New World Polycentropodidae, including a redescription of Polyplectropus is provided. The homology of the male genitalia of Polyplectropus is discussed. Ninety-two species are treated. The following 39 new species are described: Polyplectropus adamsae (Peru), P. alatespinus (Brazil), P. amazonicus (Brazil), P. andinensis (Argentina, Bolivia), P. blahniki (Venezuela), P. bolivianus (Bolivia), P. brasilensis (Brazil), P. brborichorum (Ecuador), P. cressae (Venezuela), P. colombianus (Colombia), P. corniculatus (Peru), P. cuzcoensis (Peru), P. ecuadoriensis (Ecuador), P. flintorum (Venezuela), P. gaesum (Brazil), P. guyanae (Guyana, Venezuela), P. hollyae (Brazil), P. hystricosus (Brazil), P. insularis (Panama), P. juliae (Brazil), P. kanukarum (Guyana), P. maculatus (Venezuela), P. manuensis (Peru), P. matatlanticus (Brazil), P. minensium (Brazil), P. novafriburgensis (Brazil), P. peruvianus (Peru), P. petrae (Brazil), P. pratherae (Brazil), P. puyoensis (Ecuador), P. robertsonae (Bolivia), P. rodmani (Brazil), P. rondoniensis (Brazil), P. tragularius (Brazil), P. tripunctatum (Peru), P. venezolanus (Venezuela), P. woldai (Panama), P. zamoranoensis (Honduras), and P. zuliae (Venezuela). Polyplectropus buchwaldi (Ulmer, 1911) is designated as a nomen dubium.Key words: phylogeny, taxonomy, Neotropics, Nearctric, Bayesian methods, parsimony, classification, cladistics, morphology, Trichoptera, Polycentropodidae, Polyplectropus, new speciesThe polycentropodid caddisfly genus Polyplectropus Ulmer contains 219 species found primarily in the tropical and subtropical regions of the world, with 125 species recorded from the Oriental biogeographic region, 53 from the Neotropical region (6 species extending their ranges into the Nearctic region), 37 from the

Simulium (Hearlea) Vargas, Mart nez Palacios, & D az N jera (Diptera: Simuliidae): Taxonomic revision and cladistic analysis

Zootaxa ◽  
2004 ◽  
Vol 396 (1) ◽  
pp. 1 ◽  
Author(s):  
S. COSCARÓN ◽  
D. R. MIRANDA ESQUIVEL ◽  
J. K. MOULTON ◽  
C. L. COSCARÓNARIAS ◽  
S. IBAÑEZ BERNAL
Keyword(s):  
Phylogenetic Analysis ◽  
New Species ◽  
Cladistic Analysis ◽  
Taxonomic Revision ◽  
Morphological Characters ◽  
Character Weighting ◽  
Species Groups ◽  
Geographic Distributions ◽  
Weighting Methods

Simulium (Hearlea) Vargas, Mart nez Palacios, & D az N jera 1957 is comprised of 20 known species that are largely confined to the area between M xico and Guatemala. Herein, all currently recognized species within this subgenus are described, illustrated, keyed (except males), and subjected to phylogenetic analysis. Geographic distributions for each species are also updated. Simulium deleoni Vargas, 1945, is synonymized with S. capricorne De Le n, 1944. Simulium paracarolinae new species, from Guatemala, is described from the larva. A cladistic analysis of all Hearlea species and representatives from three outgroups was conducted using 39 morphological characters. Although different character weighting methods yielded different topologies, all trees agreed that Hearlea is monophyletic is comprised of two species groups, based mainly upon characters of the larva and pupa: the Simulium juarezi group S. ayrozai Vargas, S. burchi Dalmat, S. canadense Hearle, S. capricorne De Le n, S. chiriquiense Field, S. contrerense D az N jera & Vulcano, S. dalmati Vargas & D az N jera, S. delatorrei Dalmat, S. estevezi Vargas, S. ethelae Dalmat, S. juarezi Vargas & D az N jera, S. microbranchium Dalmat, and S. nigricorne Dalmat and the S. carolinae group Simulium (Hearlea) carolinae De Le n, S. gorirossiae Vargas & D az N jera, S. johnsoni Vargas & D az N jera, S. larvispinosum De Le n, S. menchacai Vargas & D az N jera, S. paracarolinae n. sp., and S. temascalense D az N jera & Vulcano.

A phylogeny of the family Fanniidae Schnabl (Insecta:Diptera:Calyptratae) based on adult morphological characters, with special reference to the Austral species of the genus Fannia

2008 ◽  
Vol 22 (5) ◽  
pp. 563 ◽  
Author(s):  
M. C. Domínguez ◽  
S. A. Roig-Juñent
Keyword(s):  
Cladistic Analysis ◽  
Morphological Characters ◽  
External Morphology ◽  
Phylogenetic Position ◽  
Phylogenetic Hypothesis ◽  
Parsimony Analysis ◽  
Male And Female ◽  
Southern South America ◽  
The Family ◽  
Species Groups

The present study proposed a phylogenetic hypothesis of the family Fanniidae based on a cladistic analysis using characters from adult external morphology and female and male terminalia. The main purpose of this study was to clarify the phylogenetic position of newly described or poorly known species, mostly from southern South America, the Neotropics, Africa, Australia and New Zealand. In total, 151 characters from adult male and female external morphology and terminalia were scored for 78 species of Fanniidae. Ten continuous characters were included and analysed as such. Three genera of Fanniidae and all the species-groups and subgroups proposed for the genus Fannia, except for the admirabilis-group and the setifer-subgroup were included as terminal taxa. An heuristic parsimony analysis under implied weights was performed. The analysis recovered the monophyly of the Fanniidae and the genus Fannia, as well as the monophyly of several species-groups within Fannia. Male and female external morphological characters were, in general, highly homoplasious, whereas characters from male terminalia showed low level of homoplasy and provided resolution at suprageneric nodes and species-groups.

Cryptic species diversity and character congruence: review of the tribe Anthracini (Diptera : Bombyliidae) in Australia

1998 ◽  
Vol 12 (6) ◽  
pp. 977 ◽  
Author(s):  
David K. Yeates ◽  
Christine L. Lambkin
Keyword(s):  
New Species ◽  
Cryptic Species ◽  
Cladistic Analysis ◽  
Morphological Characters ◽  
Species Group ◽  
Morphological Data ◽  
Distribution Maps ◽  
Eastern Australia ◽  
Wing Patterns ◽  
Species Groups

The Australian Anthracini are revised. In all, 28 new species are described, bringing the total fauna to 34 species. The previously described species of Anthrax Scopoli – A. maculatus Macquart, A. incomptus Walker, A. confluensisRoberts, A. lepidiotus Roberts and A. proconcisus Hardy – are diagnosed and the following eight new species of Anthrax are described: A. argentia, A. asciculus, A. clinatus, A. crenatus, A. dolabratus, A. funestus, A. opacus and A. torulus. This taxonomic study reveals a group of at least 20 cryptic species previously included in collections under the name Anthrax angularis Thomson. A new genus, Thraxan, is erected to contain this cryptic group of species and the following 20 new species are described: T. acutus, T. abditus, T. caligneus, T. cinctus, T. cornuatus, T. depressus, T. echinatus, T. ebenus, T. emicatus, T. hamulus, T. luteus, T. misatulus, T. nodus, T. norrisi, T. obstipus, T. patielus, T. planus, T. prolatus, T. simulatusand T. spiculus. Many of these cryptic species have been collected sympatrically, hill topping together in eastern Australia. A key is provided to the species of Anthrax and Thraxan, genitalia drawings are presented for most species and distribution maps of all species are presented. A cladistic analysis of the species of Anthrax and Thraxan is also presented. A total of 26 of the species is compared for 125 synapomorphies in 39 adult morphological characters. Three species-groups were found: Thraxan, and two species-groups within Anthrax, the A. proconcisus species-group and the A. maculatusspecies-group. Previous authors divided Anthrax into species-groups on the basis of wing patterns, but found that these species-groups were not confirmed when other characters were taken into consideration. We studied the congruence of seven different character sets within the clade comprising Anthrax and Thraxan – antennae, venation, wing patterns, vestiture, genitalia, male genitalia and female genitalia – using several incongruence indices. Significance of incongruence was measured using a randomisation procedure. Results of these studies indicate that the wing-pattern character set is significantly incongruent with the other morphological data. These quantitative cladistic results explain the difficulty previous authors experienced in finding suites of characters to support species-groups in Anthrax on the basis of wing patterns. A relationship is found between the level of incongruence and the distance over which mate-recognition signals operate.

CLASSIFICATION AND EVOLUTION OF THE EUMENINE WASP GENUS SYMMORPHUS WESMAEL (HYMENOPTERA: VESPIDAE)

1989 ◽  
Vol 121 (S148) ◽  
pp. 5-168 ◽  
Author(s):  
Jeffrey M. Cumming
Keyword(s):  
Cladistic Analysis ◽  
Rio Grande Valley ◽  
Rio Grande ◽  
Neotropical Region ◽  
Type Locality ◽  
Nesting Behavior ◽  
Phylogenetic Position ◽  
Species Groups ◽  
Three New Species ◽  
New Synonymy

AbstractThe genus Symmorphus Wesmael is distributed throughout the Oriental, Palearctic, and Nearctic regions, as well as the northernmost Neotropical region, and appears to occupy a relatively isolated phylogenetic position within the Eumeninae. Females of Symmorphus, for those species in which nesting behavior is known, mass-provision cells in pre-existing cavities with chrysomelid, curculionid, or microlepidopterous larvae.Thirty-six species are recognized, including three new species and two new subspecies. Keys are provided for the identification of adult specimens, all species are described, and their geographical distributions mapped.Two subgenera, S. (Parasymmorphus) Cumming and Vecht and S. (Symmorphus) Wesmael, are recognized. Symmorphus (Parasymmorphus) includes S. parvilineatus (Cameron), S. momunganensis (Schulthess), and S. negrosensis Cumming and Vecht. Within S. (Symmorphus) three informal species groups are recognized, with the following included species and subspecies: (1) S. hoozanensis group — S. hoozanensis (Schulthess), S. alkimus alkimus Cumming and Vecht, S. alkimus dialeukus ssp.nov. (type-locality Sibolangit, N. Sumatra), S. ambotretus sp.nov. (type-locality Mt. Omei, Sichuan Province, China), and S. canlaonicus sp.nov. (type-locality Mt. Canlaon, Negros Oriental, Philippines); (2) S. declivis group — S. declivis Harttig; (3) S. murarius group — S. murarius (Linnaeus), S. foveolatus Gussakovskij, S. glasunowi Morawitz, S. sichuanensis Lee, S. crassicornis (Panzer), S. captivus (Smith), S. angustatus (Zetterstedt), S. albomarginatus albomarginatus (Saussure), S. albomarginatus midas ssp.nov. (type-locality Bentsen Rio Grande Valley State Park, Texas, USA), S. decens (Kostylev), S. violaceipennis Soika, S. mizuhonis Tsuneki, S. bifasciatus (Linnaeus), S. debilitatus (Saussure), S. cliens Soika, S. connexus (Curtis), S. lucens (Kostylev), S. sublaevis (Kostylev), S. canadensis (Saussure), S. projectus Bohart, S. tukvarensis (Meade-Waldo), S. nipteroides sp.nov. (type-locality N. Yourdou, India), S. fuscipes (Herrich-Schaeffer), S. apiciornatus (Cameron), S. aurantiopictus Soika, S. allobrogus (Saussure), S. cristatus (Saussure), S. gracilis (Brullé), and S. paralleliventris Soika.The following new synonymy is proposed: S. calciatii (Zavattari) = S. glasunowi Morawitz; S. nursei (Bingham), S. hospes (Nurse), and S. hospitans (Dalla Torre) = S. crassicornis (Panzer); S. hakutozanus Tsuneki and S. nansetsurei Tsuneki = S. angustatus (Zetterstedt); S. walshianus (Saussure) and S. meridionalis Viereck = S. albomarginatus albomarginatus (Saussure); S. kurentzovi Kurzenko, S. iiyamai Tsuneki, S. shiroyamai Tsuneki, S. piceanus Tsuneki, and S. sassai Tsuneki = S. mizuhonis Tsuneki; S. sinuatus (Fabricius), S. sinuatus mutinensis (Baldini), S. sparsus Morawitz, S. sinuatissimus Richards, S. mutinensis auster Soika, and S. mutinensis yezoanus Tsuneki = S. bifasciatus (Linnaeus); S. ishikawai Soika = S. lucens (Kostylev); S. seoulensis Tsuneki = S. apiciornatus (Cameron); S. pumilus (Saussure) and S. trisulcatus Cameron = S. cristatus (Saussure); S. elegans libanicus Soika = S. gracilis (Brullé); and S. allobrogus (Saussure) is removed from synonymy with S. bifasciatus (Linnaeus). Odynerus latipennis Smith and Odynerus cilicius Cameron, previously considered to belong within Symmorphus, are excluded from the genus, and S. sounkionis Tsuneki is transferred as Ancistrocerus sounkionis (Tsuneki) comb.nov. Neotypes are designated for S. crassicornis (Panzer) and S. fuscipes (Herrich-Schaeffer), and lectotypes are designated for S. foveolatus Gussakovskij, S. glasunowi Morawitz, S. nursei (Bingham), S. laeviventris (Thomson), S. albomarginatus (Saussure), S. decens (Kostylev), S. sinuatus (Fabricius), S. sinuatus mutinensis (Baldini), S. debilitatus (Saussure), S. lucens (Kostylev), S. sublaevis (Kostylev), S. debilis (Saussure), S. apiciornatus (Cameron), S. philadelphiae (Saussure), and S. elegans (Wesmael).A cladistic analysis of Symmorphus, based on 26 characters, is presented, including a discussion of the relationships of the genus to the remainder of the Eumeninae. Hypotheses outlining the historical zoogeography of Symmorphus are presented in light of current paleogeographic and paleoclimatic reconstructions, by comparing these cladistic relationships with the chorological patterns observed within the genus and other Vespidae.

A revision of the Australian taxa previously attributed to Assiminea buccinoides (Quoy & Gaimard) and Assiminea tasmanica Tenison-Woods (Mollusca:Gastropoda:Caenogastropoda:Assimineidae)

2005 ◽  
Vol 19 (4) ◽  
pp. 325 ◽  
Author(s):  
Hiroshi Fukuda ◽  
Winston F. Ponder
Keyword(s):  
New Genus ◽  
Cladistic Analysis ◽  
River Estuary ◽  
South Australia ◽  
Single Species ◽  
Morphological Characters ◽  
South Eastern ◽  
Eastern Australia ◽  
Species Groups ◽  
Western Victoria

A new genus, Cryptassiminea, is introduced for the taxon previously known as Assiminea buccinoides (Quoy & Gaimard). These small gastropods are abundant in mangrove and salt marsh habitats in south-eastern and subtropical eastern Australia. Seven species (five new) are recognised using morphological characters in the complex previously treated as a single species. Five taxa have rather narrow ranges while the other two are widespread and often sympatric. Two groups of species are recognised. One contains Cryptassiminea buccinoides, widespread in south-east and east Australia, and two closely related allopatric taxa from South Australia and south-eastern Tasmania (C. adelaidensis, sp. nov. and C. kershawi, sp. nov.). A second group of species is typified by Cryptassiminea tasmanica (Tenison-Woods), also widespread in east and south-east Australia and often sympatric with C. buccinoides. Allied to C. tasmanica, are two closely related taxa from western Victoria: C. glenelgensis, sp. nov. from the Glenelg River estuary and C. surryensis, sp. nov. from the Surry River estuary and Western Port, in the vicinity of Geelong. A distinctive species, Cryptassiminea insolata, sp. nov. from the east coast of Queensland, also has similarities with C. tasmanica. A cladistic analysis using morphological characters of the Cryptassiminea taxa and three other genera of Assimininae, with an omphalotropidine as the outgroup, resulted in a single tree. The new genus has rather poor support, possibly because many of its characters appear to be plesiomorphic within Assimineinae. Cryptassiminea is defined by a unique combination of characters but lacks any obvious synapomorphy. Two clades within Cryptassiminea are well supported, each containing the species-groups referred to above.

Molecular phylogeny of the limacoid snail family Dyakiidae in Southeast Asia, with the description of a new genus and species

2020 ◽  
Author(s):  
Parin Jirapatrasilp ◽  
Piyoros Tongkerd ◽  
Ekgachai Jeratthitikul ◽  
Thor-Seng Liew ◽  
Arthit Pholyotha ◽  
...  
Keyword(s):  
Molecular Phylogeny ◽  
Southeast Asia ◽  
New Genus ◽  
Cladistic Analysis ◽  
Morphological Characters ◽  
The Other ◽  
New Genus And Species ◽  
Snail Family ◽  
Geographical Distribution Pattern ◽  
Transformation Series

Abstract Members of the terrestrial snail family Dyakiidae from Southeast Asia show a distinct geographical distribution pattern and possess different degrees of complexity in their amatorial organ gland. This study is the first molecular phylogeny of ten of the 12 genera in this family, performed to provide insights into the origin of Dyakiidae and the evolution of their shells and amatorial organ gland structure. A new genus and new species, Pseudoquantula lenticularis Jirapatrasilp & Panha gen. & sp. nov., was uncovered based on its distinct morphological characters and molecular divergence. All other genera were retrieved as monophyletic except for Dyakia. Mainland Southeast Asia was inferred to be the ancestral range of the Dyakiidae, and the lineages then dispersed to and diversified in Borneo. Cladistic analysis showed that all 14 morphological characters used in this study were homoplastic. These results disagree with the previous amatorial organ transformation series, in which neither Pseudoplecta nor Quantula was ancestral to the other genera. The enigmatic genus Pseudoplecta, which lacks an amatorial organ gland, exhibited secondary loss.

Taxonomy and evolution of asymmetric male genitalia in the subgenus Ashima Chen (Diptera: Drosophilidae: Phortica Schiner), with descriptions of seven new species

Zootaxa ◽  
2020 ◽  
Vol 4789 (1) ◽  
pp. 1-54
Author(s):  
MASANORI J. TODA ◽  
HANS BÄNZIGER ◽  
PRADEEP C. SATI ◽  
RAJENDRA S. FARTYAL ◽  
AWIT SUWITO ◽  
...  
Keyword(s):  
New Species ◽  
Male Genitalia ◽  
Species Complex ◽  
Cladistic Analysis ◽  
Morphological Characters ◽  
Mirror Image ◽  
Directional Asymmetry ◽  
Ancestral State ◽  
Antisymmetric State ◽  
Species Groups

The taxonomy of the subgenus Ashima of the genus Phortica is revised. A cladistic analysis of 66 morphological characters is conducted, covering 35 species (28 known and 7 new species: Phortica efragmentata sp. nov., P. andreagigoni sp. nov., P. watabei sp. nov., P. halimunensis sp. nov., P. akutsui sp. nov., P. kerinciensis sp. nov., and P. takehiroi sp. nov.) which correspond to 71.4% of 49 total spp. of Ashima. The resulting cladogram shows that the studied species are separated into several clades/subclades/cluster each highly supported with specific synapomorphies. Those clades/subclades/cluster are newly defined as the following species groups, subgroups or complex: the afoliolata, foliiseta, nigrifoliiseta and angulata species groups; the foliiseta, tanabei, nigrifoliiseta and glabra species subgroups; and the foliiseta species complex. The subgenus Ashima is peculiar in having the asymmetric male genitalia as the ground plan and showing the antisymmetry (i.e. intraspecific mirror-image variation) in some species but the directional asymmetry (i.e. side-fixed asymmetry) in others. The evolution of genital asymmetry in this subgenus is estimated by mapping the states (symmetry, directional asymmetry and antisymmetry) of bilateral structures of male genitalia on the cladogram. This ancestral state reconstruction estimates that the directional asymmetry of male genitalia has evolved at the ancestor of this subgenus and then changed to the antisymmetric state independently in two lineages, the angulata + nigrifoliiseta species groups and the foliiseta species complex. In this study, a standardized terminology recently proposed for the male terminalia of Drosophila melanogaster is extendedly adopted to describe the morphology of male terminalia in the subgenus Ashima, one group of the subfamily Steganinae. 

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