CLADOENDESIS OF EPHEMEROPTERA

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Tricorythodes/fg1

(Panephemeroptera Euephemeroptera Euplectoptera Anteritorna Bidentiseta Furcatergaliae  
Ephemerella/fg1 Pantricorythi Tricoryptera Leptohyphes/fg1 - Tricorythodes)

Nomen hierarchicum: Tricorythodes/fg1 [f:2000; g:1920] (incl. Tricorythafer, Tricoryhyphes, Homoleptohyphes, Asioplax)   

In circumscription fits:

— gen. Tricorythodes Ulmer (1919)1920:1–80: 51

— Tricorythodes/fg1: Kluge 2004: 343

Doubtful nominal taxon included:

Tricorythafer/g [g:1942] synn. obj.: Caenopsis/g [g:1920] nom.praeocc.; Needhamocoenis/g [g:1945] (probably erroneously described from Africa as male imago of a single species)

Nominal taxa included: 

Tricoryhyphes/g [g: 1987] 

Homoleptohyphes/g [g:1987] 

Asioplax/g [g:2000]

Macunahyphes/g [g:2005] 

Cabecar/g [g:2006]


References. Ulmer (1919)1920:1–80: ; – Needham & Traver & Hsu 1935: * *; – Kluge 1992c: * * *; – Kluge 2004: * * *


Autapomorphy of Tricorythodes.

(1) 2nd segment of gonostylus is basally strongly swollen, bulbous (Kluge & Naranjo 1990: Fig.26–29, 51–56, 70–71). In other respects gonostylus retains plesiomorphic features: 1st segment [elongate – see Pantricorythi (2)] is well-developed, 3rd (distal) segment is present (unlike Tricorythopsis and some others – see Index of characters [2.3.12]).

Apomorphies of Tricorythodes common with Caenoptera (see Classifications of Furcatergaliae II).

(2) Posterior scutal protuberances [bearing dorsal bases of S.LPNm – see (3)] are strongly enlarged and convergent posteriorly; each of them has a common boundary with medioscutum all over its length, because lateroparapsidal suture lost its curvation and merges with medioparapsidal suture anteriad of transverse interscutal suture [see Leptohyphes/fg1 (4)] (Kluge 2004: Fig.104:A). Similar in Tricorygnatha.

(3) Lateropostnotum [bearing ventral base of S.LPNm – see (2)] is strongly enlarged, stretches up to sternum, reaches coxal conjunctiva and is fused with epimeron (Kluge 2004: Fig.104:B). The same in Tricorythopsis, unlike all other Ephemerella/fg1 (see Index of characters [2.2.16]).

(4) Metathorax is strongly reduced (Kluge 2004: Fig.104:E–G): hind wings [see Leptohyphes/fg1 (2)] are lost in both sexes; in female metathoracic scuto-episternal muscles (S.ESm) are lost (while in male non-functional indirect wing muscles S.ESm and MTm of metathorax are retained). The same in Caenoptera; in other mayflies, even if hind wings are completely lost, metathoracic indirect wing muscles are retained in both sexes.

(5) In imago and subimago 1st tarsal segment of middle and hind legs [initially fused with tibia and shortened – see Furcatergaliae (2)] on its outer side is secondarily distinctly separated from tibia; joining of tibia and tarsus is strongly inclined resembling larval condition, thus outer side of 1st tarsal segment is strongly elongate. Tarsus retains 5 distinctly separated segments (Kluge 2004: Fig.105:A). The same in Caenoptera (Kluge 2004: Fig.86:G); in Tricorythopsis and Tricorygnatha tarsus is separated in the same manner, but has four segments only (see Index of characters [2.2.84]).

(6) In female imago and subimago caudalii [cerci and paracercus – see Plesiomorphies of Ephemerella/fg1] are strongly diminished, shorter than the body. When adult caudalius develops under larval cuticle, it is not crumpled (Kluge 2004: Fig.105:E), unlike Leptohyphes/fg2 (Kluge 2004: Fig.101:G), Allenhyphes, Tricorygnatha and other taxa. This character is not connected with sexual dimorphis of larval caudalii [see Leptohyphes/fg1 (8)].

Apomorphy of Tricorythodes common with Leptohyphes/fg2 and Allenhyphes.

(7) Tergalii completely lost processes of ventral bifurcate lobes, thus each tergalius consists of no more than 3 simple lamellae arising from base [see Leptohyphes/fg1 (1)]. The same on tergalii II of Leptohyphes/fg2 and all tergalii of Allenhyphes.

In selected species (bullus [T.], cubensis [T.], grallator [T.], santarita [T.]) tergalius II (gill operculum) lost anal branch of ventral lobe, thus is 2-lamellate; in santarita [T.] costal branch of its ventral lobe is vestigial; in other species this costal branch is well-developed and has well-developed costal flap [see Leptohyphes/fg1 (1)] (Kluge 2004: Fig.103:A-B) (unlike Leptohyphes/fg2). Tergalii III–V are always 3-lamellate. In various species tergalius VI is 2-lamellate or 1-lamellate (Kluge & Naranjo 1990: Fig.18–65; Molineri 2002: Fig.14–172).

Plesiomorphies of Tricorythodes. Mouth apparatus [see Ephemerella/fg1 (1)–(3), (16)] is non-specialized, maxilla with 3 canines and 2 dentisetae [see Bidentiseta (1)]. Unlike Leptohyphes/fg2: tergalius II (gill operculum) has well-developed costal flap of ventral lobe [see Leptohyphes/fg1 (1)] (Kluge 2004:  Fig.103:A-B); lateropostnotum [see (3)] retains well-developed straight lateropostnotal crest forming one line with infrascutellar-lateropostnotal arm (Kluge 2004: Fig.104:B;).

Variable characters of Tricorythodes. In the group sacculobranchis larval femoral setae [see Pantricorythi (1)] are short and blunt, and gill operculum is oval, with maximum width near apex (Kluge & Naranjo 1990: Fig.1–23) (the same in Leptohyphes/fg2 and Allenhyphes); in other known groups of Tricorythodes femoral setae are pointed, and gill operculum is triangular, with maximum width near base.

Size. Fore wing length 3–9 mm (see Tricoryptera).

Distribution. America (Neotropical and Nearctic Regions).


Nominal species in Tricorythodes/fg1:  

albilineatus Berner 1946 [Tricorythodes]

allecta Needham 1905 [Caenis] --/,/

angulatus Traver 1959 [Tricorythodes]

arequita Traver 1959 [Tricorythodes]

atrata McDunnough 1923 [Tricorythus] — syn.subj. allecta [Caenis]

australis Banks 1913 [Tricorythus] — typus nominis Macunahyphes

barbus Allen 1967 [Tricorythodes]

baumanni Kilgore & Allen 1973 [Leptohyphes] — syn.subj. mirus [Leptohyphes]

bullus Allen 1967 [Tricorythodes]

capuccinorum Emmerich 2007 [Tricorythodes]

caunapi Dias & Bacca & Ferreira 2011 [Tricorythodes]

chalaza Gonçalves & Da-Silva & Nessimian 2010 [Tricorythodes]

cobbi Alba-Tercedor & Flannagan 1995 [Tricorythodes]

comus Traver 1959 [Tricorythodes]

condylus Allen 1987 [Tricorythodes]

corpulentus Kilgore & Allen 1973 [Tricorythodes]

cristatus Allen 1967 [Tricorythodes]

cubensis Kluge & Naranjo 1990 [Tricorythodes] --/

curiosus Lugo-Ortiz & McCafferty 1995 [Leptohyphes]

curvatus Allen & Cohen 1977 [Tricorythodes]

diasae Gonçalves & Da-Silva & Nessimian 2010 [Tricorythodes]

dimorphus Allen 1967 [Tricorythodes]

edmundsi Allen 1967 [Tricorythodes]

explicatus Eaton 1892 [Tricorythus] ,//,///

faeculopsis Belmont & Salles & Hamada 2011 [Tricorythodes]

fallacina McDunnough 1939 [Tricorythodes] — syn.subj. explicatus [Tricorythus]

fallax Traver (in Needham & Traver & Hsu) 1935 [Tricorythodes] — syn.subj. explicatus [Tricorythus]

fictus Traver (in Needham & Traver & Hsu) 1935 [Tricorythodes] ,

fugitans Needham 1920 [Caenopsis]

grallator Kluge & Naranjo 1990 [Tricorythodes] --/

griseus Hofmann & Sartori (in Hofmann & Sartori & Thomas 1999 [Tricorythodes]

hiemalis Molineri 2001 [Tricorythodes]

kirki Baumgardner 2007 [Tricorythodes]

lichyi Traver 1943 [Tricorythodes]

minutus Traver (in Needham & Traver & Hsu) 1935 [Tricorythodes] — syn.subj. explicatus [Tricorythus]

mirca Molineri 2002 [Tricorythodes]

mirus Allen 1967 [Leptohyphes]

molinerii Dias & Salles 2006 [Tricorythodes]  

mosegus Alba-Tercedor & Flannagan 1995 [Tricorythodes]

montanus Kluge & Naranjo 1990 [Tricorythodes] --/

mulaiki Traver 1959 [Tricorythodes]

nicholsae Wang & Sites & McCafferty 1997 [Leptohyphes]

notatus Allen & Brusca 1973 [Tricorythodes]

numinuh Wiersema & McCafferty & Baumgardner 2001 [Asioplax] -,-

ocellus Allen & Roback 1969 [Tricorythodes]

peridius Burks 1953 [Tricorythodes] — syn.subj. allectus [Caenis]

primus Baumgardner 2007 [Tricorythodes]

popayanicus Domínguez 1982 [Tricorythodes]

quizeri Molineri 2002 [Tricorythodes]

sacculobranchis Kluge & Naranjo 1990 [Tricorythodes] --/

sallesi Dias & Cabette & De Sousa 2009 [Tricorythodes]

santarita Traver 1959 [Tricorythodes]

serratus Baumgardner & Ávila 2006 [Cabecar] — typus nominis Cabecar

sierramaestrae Kluge & Naranjo 1990 [Tricorythodes] --/

sordidus Allen 1967 [Tricorythodes]

stygiatus McDunnough 1931 [Tricorythodes]

texanus Traver (in Needham & Traver & Hsu) 1935 [Tricorythodes]

tragoedia Souto & Angeli & Salles 2017 [Tricorythodes]

trifasciatus Molineri & Zuniga 2006 [Tricorythodes]

ulmeri Allen & Brusca 1973 [Tricorythodes]

uniandinus Emmerich 2007 [Tricorythodes]

yapekuna Belmont & Salles & Hamada 2012 [Tricorythodes]

yura Molineri 2002 [Tricorythodes]

zunigae Molineri 2002 [Tricorythodes]


See also: 

Leptohyphes/fg1 INCERTAE SEDIS

Furcatergaliae INCERTAE SEDIS

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