CLADOENDESIS OF EPHEMEROPTERA

ABC

z 

[1] Larva

[2] Subimago & imago

[3] Egg

[1.1] Head [1.2] Thorax [1.3] Abdomen [2.1] Head [2.2] Thorax [2.3] Abdomen

INDEX OF CHARACTERS

2.3. SUBIMAGO & IMAGO:
ABDOMEN AND ITS APPENDAGES
Segments I–VIII [2.3.1-5], sternum IX [2.3.6–7], gonostyli [2.3.8–14], penis [2.3.15–17], segment X [2.3.18], caudalii [2.3.19–22]

Abdominal segments I–VIII [2.3.1-5]

[2.3.1] Traces of larval tergalial attachment

— VI transferred anteriorly: Posteritorna (5), Caenotergaliae (1)

— VII transferred anteriorly: Ephemera/fg9 (1)

[2.3.2] Lateral lobes

— on segments V–IX: Siphlonisca (2), part of Oniscigaster/fg2 (VAR)

[2.3.3] Abdominal segment VI

— enlarged: Posteritorna (5)

— with transverse dorsal elevation: Baetisca/f3=g2 (1)

[2.3.4] Abdominal segment VII

— enlarged, with mid-dorsal elevation: Machadorythus (1)

— in female with copulatory pouch: Siphlonurus/fg1 (1)

— in female with copulatory pouch with basal lobes: Parameletus/fg1 (1)

— in female with secondary ovipositor: ?Habrophlebiodes (?) Habrophlebia/fg3 (1), Megaglena (?), Maheathraulus (?), ?Nesophlebia (?), Hagenulus/fg3 (), Borinquena ()

[2.3.5] Abdominal sternum VIII

— in female with pair of impressions: Tortopus/g1 (1)

  

Abdominal sternum IX of subimago & imago [2.3.6–7]

[2.3.6] Postgenital (subanal) plate of female

— normally developed: Siphlaenigma (P)

— reduced: Liberevenata (4)

— bipointed: Baetisca/f3=g2 (2)

[2.3.7] Styliger of male

— normally developed: Siphlaenigma (P)

— with reduced median part and separated gonostyli pedestals: Liberevenata (4), Isonychia/fg2 (1), Polymitarcys/f1=Ephoron/g2 (not all) (7) {Campsurus/fg2 (2), Asthenopus (P)}

— reduced, without gonostyli pedestals: Geminovenata (6)

— medially elongate and contiguous with gonostyli: Brasiliocaenis (1)

— dorsally with membranous area: Ameletus/fg1 (5)

— W-shapely speaks into sternite IX: Caenoptera (5)

— with projection on gonostylus pedestal: Tortopus (1)

— with pair of apical-dorsal processes: Hermanellognatha (3) 

— with unpaired apical-dorsal processes: ??Thraulodes ()

— with median sterno-styligeral muscle: ..... Indobaetis (P), 

— without median sterno-styligeral muscle: part of Liberevenata (4) {?Afroptiloides (), Labiobaetis/f1=Pseudopannota/g1 (1) {Echinobaetis (), Papuanatula (??)} part of Rhodobaetis (), Glossidion (), Pseudocloeon (?)}

 

Gonostyli [2.3.8–14]

[2.3.8] Gonostylus

— absent (among all Ephemeroptera): part of Geminovenata (6) {Homoeoneuria/g2 (6), Fittkauneuria (1)}

— vestigial: Potamanthellus (1), part of Caenis/f5=g3 (1)

— without muscle: Campsurus/fg2 (2)

— grooved: Caenoptera (not all) (5) {Brachycercus/f1=g3 (P), Madecocercus (P), Tillyardocaenis (P), "Afrocercus" (P)}

— non-grooved: part of Caenoptera (5) {Caenis/f5=g3 (1)}

— contiguous with styliger: Brasiliocaenis (1)

[2.3.9] Anlage of subimaginal gonostylus in larva has

— 2nd segment directed laterally (Afroptilum-Cloeon-type): Turbanoculata (not all) (13) {pm. Protopatellata (P), Cloeon/fg1 (P), Centroptilum/g1 (), Baetopus/g1 (P), Cheleocloeon (P), Afrobaetodes (P)}

— 2nd segment directed posteriorly, can be curved or dipped (Nigrobaetis-type): part of Turbanoculata (13) {Baetovectata (not all) (3) {Nigrobaetis/g1 (P), Camelobaetidius/g1 (P), Baetodes/g1 (P), Fallceon (P), Caribaetis (P), Americabaetis (P), Deceptiviosa (P), Mayobaetis (P), Moribaetis (P), Edmundsiops (P), Callibaetis/fg1 (5), Cloeodes/g1 (P)}}

— 2nd segment directed medially-posteriorly (Baetis-type): part of Turbanoculata (13) {Baetofemorata (not all) (2)}

— 2nd segment curved medially-anteriorly (Labiobaetis-type and Acentrella-type): part of Turbanoculata (13) {Baetofemorata (not all) (2) {Acentrella/fg1 (9)}, Labiobaetis/f1=Pseudopannota/g1 (2), Acerpenna (), Indobaetis (), Echinobaetis (), Paracloeodes/g1 ()}

— apices brought together: Rhithrocloeon/fg1 (1.2)

[2.3.10] 1st segment of gonostylus

— movably articulated with 2nd: part of Pantricorythi (2)

— distinct from 2nd: Chromarcys (P), Ichthybotus (P), Ephemera/fg9 (P), Pentagenia (P), Protobehningia (P)

— not distinct from 2nd: Geminovenata (6), Potamanthus/fg1 (9), Euthyplocia/fg1 (5), Behningia/fg2 (4), Palingenia/f3=g2 (5), Polymitarcys/f1=Ephoron/g2 (7), †"Palaeoanthus" (1), Potamanthellus/g1 (1), Caenoptera (5)

— elongate: Pantricorythi (2)

— not elongate: Ephemerella/fg2 (P)

— with apical-ventral protuberance: Rhithrocloeon/g1 (1.1) 

— on inner margin with projection: Habroleptoides (1) 

— on inner margin with angular bend: Habrophlebia/fg3 (2) 

[2.3.11] 2nd segment of gonostylus

— pointed: Rhithrocloeon/g1 (1.1) 

— thickened apically: Cloeon/fg1 (6) 

— with bulbous base: Tricorythodes (1)

— directed laterally: part of Pantricorythi (2) {Tricorythopsis (3)}

[2.3.12] Number of distal segments of gonostylus (among all Ephemeroptera)

— more than 2: Mirawara (1), part of Discoglossata (12), Arthroplea (6), part of Cinygmula (VAR), part of Palingenia/f3=g2 (5), †"Cronicus" (3)

— 2: Siphlonurus/fg1 (P), Dipteromimus (P), Ameletus/fg1 (P), Metretopus/fg1 (P), Acanthametropus/fg2 (P), Ametropus (P), Siphlaenigma (P), Vetulata (P), Nesameletus/f1=Metamonius/g1 (P), Ameletopsis/fg2 (P), Chiloporter (P), Chaquihua (P), Rallidens (P), Coloburiscus/fg1 (P), Isonychia/fg1 (P), Oligoneurisca (P), Oligoneuriella/g1 (not all) (P), Elassoneuria/g1 (not all) (P), Heptagennota (not all) (P), Neoephemera/fg1 (P), Potamanthus/fg1 (9), Ichthybotus (P), Ephemera/fg10 (P), Hexagenia/fg3 (P), Litobrancha (P), Pentagenia (P), part of Palingenia/f3=g2 (5), Polymitarcys/f2=Ephoron/g3 (P), Neoephemera/fg1 (P), Leptophlebia/fg1 (not all) (P), †"Electrogenia" (P), †"Xenophlebia" (P), †"Cretoneta" (P), †"Mogzonurus" (P)

— 1: Posteritorna (12), Analetris (1), Liberevenata (not all) (5), Oligoneuria/f4=g5 (2), Euthyplocia/fg1 (not all) (5) {Euthyplocia/fg2 (P), Proboscidoplocia (P)}, Pseudeatonica (1), Eatonica/fg2 (1), Eatonigenia (3), Ephemerella/fg1 (not all) (8), part of Leptophlebia/fg1 (P) {part of Terpides/fg1 (7) {Fittkaulus (1)}}

— 0: part of Liberevenata (5) {Rhithrocloeon/fg2 (1)}, Campylocia (3), Polyplocia (3), Exeuthyplocia/fg1 (4), Behningia/fg1 (10), Campsurus/fg1 (10), Caenoptera (5), part of Pantricorythi (2) {Afrotricorythi (2), Tricorythopsis (3), Haplohyphes (1)}

[2.3.13] Distal segments of gonostylus

— shortened: Neoephemera/fg1 (P)

— elongate: Miroculis/g1 () 

— narrowed basally: Thraulodes () 

[2.3.14] Single distal segment of gonostylus

— clavate: Cloeon/fg1 (6) 

— small: Pantricorythi (not all) (2)

— elongate: Attenella (1), Indocloeon (1)

Penis [2.3.15–17]

[2.3.15] Penial arms

— articulated with gonostyli pedestals: Campsurus/fg2 (2)

— lost articulation with tergite IX: Campsurus/fg2 (2), part of Caenoptera (5)

[2.3.16] Penis

— with titillators – see [2.3.17]

— with median dorsal condylus: Elassoneuria/g1 (3) {Madeconeuria (P)}, Oligoneuria/f4=g5 (3)

— with extensible lateral and median sclerites: Elassoneuria/g2 (2)

— with telescopic apical process: Oligoneuriella/g1 (1), Homoeoneuria/g2 (7)

— with other characteristic features: Siphlonurus/fg1 (3), Parameletus/fg1 (2), Ameletus/fg1 (6) {Metreletus (3), Ameletus/fg2 (2)}, Isonychia/fg2 (P), Prionoides (1), Borisonychia (), Oligoneuriella/g1 (1) {Oligoneuriella/g2 (1), Oligoneuriopsis (1)}, Pentamerotarsata (5) {Afghanurus/g1 (1), Rhithrogena/fg4 (2), Epeiron (not all) (3), Himalogena (2), Sibirigena (3), Iron/g2 (P), Ironopsis/g1 (2)}, Potamanthus/fg2 (1), Eatonigenia (2), Behningia/fg1 (9), Caenoptera (5), Madecassorythus/fg1 (1), Leptophlebia/fg3 (1), Habrophlebiodes (1), Habrophlebia/fg1 (2), Hagenulus/fg2 () {Careospina (), Traverina ()}, Hermanellonota (2), Kimminsula (?), Petersula (), Castanophlebia (1)

— consists of paired gonovectes [2.3.16] and penial bridge [2.3.16b]: Liberevenata (4)

[2.3.16a] Gonovectes

— fused with penial bridge: Rhithrocloeon/fg1 (6), Cloeon/fg1 (), ?Baetopus/fg1 (?)

— not fused with penial bridge: Baetovectata (2), Centroptilum (), ?Cheleocloeon (P), ?Afrobaetodes (P), 

— strongly arched: Baetovectata (2)

[2.3.16b] Penial bridge

— with projection conic: Centroptilum/g1 (), part of Cloeon/fg1 () {Cloeon/fg2 ()} 

— with projection truncate or rounded: part of Cloeon/fg1 {Similicloeon (), Procloeon/g1 ()}

[2.3.17] Titillators

— median present: Pentamerotarsata (not all) (5) {Arthroplea (8), Heptagenia/f6=g5 (P), Ecdyonurus/fg2 (P), Afghanurus/fg1 (P), Notacanthurus/g1 (P), Cinygmula (P), Paegniodes (P), Proepeorus (P), Belovius/g1 (not all) (VAR), Iron/g2 (P), Caucasiron (P)}

— median contiguous: Ecdyonurus/fg2 (P), Afghanurus/g1 (P), Notacanthurus/g1 (P)

— median separated: Afronurus/g1 (not all) (1)

— median and ventral present: Arthroplea (8), Rhithrogena/fg2 (not all) (4)

— ventral flat: Rhithrogena/fg3 (not all) (5) {Rhithrogena/fg4 (2), Himalogena (2), Sibirigena (not all) (3)}, part of Afromera (VAR)

— ventral rolled: Ephemera/fg11 (1)

— absent: part of Afronurus/g1 (1), part of Rhithrogena/fg3 (5) {Epeiron (3), part of Sibirigena (3)} Bleptus (3), Ironodes (2), Epeorus/fg4 (1), part of Belovius/g1 (VAR), Ironopsis/g2 (1), part of Afromera (VAR)

Abdominal segment X of subimago & imago [2.3.18]

[2.3.18] Abdominal tergite X

— with angles reaching cerci ventro-laterally: Fossoriae (5), Caenotergaliae (12), Geminovenata (8)

Caudalii of subimago & imago [2.3.19–22]

[2.3.19] Caudalii of subimago and imago

— in male imago shorter than abdomen: Acanthametropus/fg2 (5)

— in male subimago (but not imago) shorter than abdomen: Prosopistoma/f1=g2 (9), Geminovenata (7), Caenoptera (6)

— in female do not moult: Caenoptera (6)

— in female diminished: Caenoptera (6), Tricorythodes/fg1 (6)

— without distinct segmentation: Prosopistoma/f1=g2 (9), Behningia/fg1 (11)

[2.3.20] Setae on caudalii

— situated as primary swimming setae of larva: Geminovenata (not all) (7)

— irregular: Chromarcys (P)

— in whorls: Geminovenata (not all) (7), Caenoptera (6)

— absent: Caenoptera (6)

[2.3.21] Cerci of subimago and imago

— contiguous basally: part of Oligoneuria/f4=g5 (6) {Lachlania (3), Spaniophlebia (3)}

[2.3.22] Paracercus of subimago and imago (among all Ephemeroptera)

— absent: part of Oligoneuria/f4=g5 (6) {Spaniophlebia (4)}

— vestigial or markedly diminished: Baetisca/f2=g1 (3), Siphlonurus/fg1 (5), Ameletus/fg1 (7), Metretopus/fg1 (6), Liberevenata (6), Tasmanophlebia/g1 (3), part of Nesameletus/f1=Metamonius/g1 (P), Ameletopsis/fg2 (3), Mirawara (4), Chaquihua (3), Rallidens (9), Coloburiscus/fg1 (9), Isonychia/fg1 (9), Chromarcys (1), part of Oligoneuriella/g1 (VAR), part of Oligoneuria/f4=g5 (6) {Lachlania (3)}, Heptagennota (6), part of Fimbriatotergaliae (P) {part of Rhoenanthus/g1 (VAR), Hexagenia/fg1 (3), Palingenia/f3=g2 (6), Campsurus/fg1 (11), Leucorhoenanthus (1), Potamanthellus/g1 (2), Teloganodes/f1=g3 (2), part of Tricorygnatha (P), Dicercomyzon (5)}, part of Leptophlebia/fg1 (P), †"Cronicus" (2), †"Electrogenia" (2), †"Balticophlebia" (P), †"Baltameletus" (1), †"Cretoneta" (2), †"Proameletus" (1)

— developed: Prosopistoma/f1=g2 (9), Dipteromimus (P), Acanthametropus/fg1 (P), Oniscigaster/fg2 (P), Nesameletus/f1=Metamonius/g1 (not all) (P), Chiloporter (P), Ametropus (P), Siphlaenigma (P), Discoglossata (not all) (P) {Oligoneuriella/g1 (not all) (VAR), Homoeoneuria/g1 (P), Elassoneuria/g1 (P), Oligoneuria/f5=g6 (4), Fittkauneuria (P)}, Fimbriatotergaliae (not all) (P) {Potamanthus/fg2 (P), Rhoenanthus/g1 (not all) (VAR), Campylocia (P), Polyplocia (P), Proboscidoplocia (P), Ephemera/fg10 (P), Neoephemera/fg2 (P), Ochernova (P), Caenoptera (P), †"Palaeoanthus" (P)}, Ephemerella/fg1 (not all) (P), Leptophlebia/fg1 (not all) (P), †Baba (P)

— developed in female, vestigial in male: Rhoenanthus/g1 (not all) (VAR), Euthyplocia/fg2 (VAR), Exeuthyplocia/fg1 (5), Ichthybotus (4), Behningia/fg1 (12), Cryptoprosternata (not all) (5) {Pentagenia (P), Polymitarcys/f2=Ephoron/g3 (P)}, part of Tricorygnatha (P)

— longer than cerci: Caudatella (1)

— with spine-like ventral seta: Allenhyphes (1)