CLADOENDESIS OF EPHEMEROPTERA

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Cloeodes/g1

(Panephemeroptera Euephemeroptera Euplectoptera Anteritorna pm.Tridentiseta  
Tetramerotarsata Liberevenata Turbanoculata
 Anteropatellata Baetovectata - Cloeodes/g1)

Nomen hierarchicum: Cloeodes/g1 [g:1938]

In circumscription fits:

— gen. Cloeodes: Salles & Gattoliatt & Sartori 2016: 100

Nominal taxon included: Notobaetis/g [g:1980]


References. Salles & Gattoliatt & Sartori 2016: ; – Kluge 2017: ;


Autapomorphies of Cloeodes/g1.

(1) On larval tibia, proximal arched row of fine setae is very regular, with setal sockets contiguous; this setal row is short and disconnected with patella-tibial suture (Kluge 2017: Fig.16; Kluge 1991: Fig.3:6; Salles & al. 2015: Fig.7d, 9d, 12a, 14a, 16a, 18a, 20a, 22a, 24a, 26a, 28a; Salles & al. 2016: Fig.6F). The species of doubtful position incus Waltz & McCafferty 1987 [Bernerius] has this character (Waltz & McCafferty 1987:177-184: Fig.11).

In Cloeodes patella-tibial suture retains usual shape, being not shifted distally (Kluge 2017: Fig.16), and is equally developed on all legs on larva (Kluge 2017: Fig.21–23, 60–62, 102) and on all legs of female subimago and imago; it is absent only on fore legs of male subimago and imago. These features are initial for Anteropatellata and different from Centroptella/g1 (incl. Chopralla), whose patella-tibial suture on larval middle and hind legs is greatly shifted distally, and on fore legs is absent in all stages.

Proximal arched row of fine setae is developed on larval tibia in some other taxa, but in taxa other than Cloeodes/g it has spaces between setae sockets (Kluge & Novikova 1992: Fig.1–2; Kluge & al. 2017: Fig.13; ).

(2) On larval abdominal sternum, transverse row of fine setae is very regular, with setal sockets contiguous (Kluge 2017: Fig.20; Waltz & McCafferty 1987:177-184: Fig.5; 1987:191-207: Fig.44; Salles et al. 2015: Fig.10E–F). Setae of these rows are long and simple (not furcate). Pair of these setal rows is present on each of abdominal sterna II–VI. On sternum II these rows can be either well-developed, or vestigial, consisting of few (3–2) setae, which sometimes are situated not close together. In incus [Bernerius] sterna have «long, fine setae randomly scattered over surface, without tufts of setae», that was a reason for placing it to a separate genus Bernerius Waltz & McCafferty 1987.

Transverse rows of long setae on the same places of abdominal sterna are present also in Centroptella/g1 (incl. Chopralla), but their setal sockets are separated, and all setae are bifurcate. Presence of such spaced rows of bifurcate setae is not a reliable apomorphy, because the same rows are present in some non-related taxa, for example in selected Oriental species of Procloeon/g2. Unlike this, rows of non-bifurcate setae with contiguous sockets is not found in taxa other than Cloeodes/g1.

Setae forming transverse rows on sterna, are very delicate and colorless; under light microscope, their structure is well visible only if prepare a dry slide of empty cuticle (molting exuviae can be used for this purpose). 

Characters of Cloeodes/g1 of unclear phylogenetic status. 

() Larval antenna bases are not brought together, frons between them does not form a keel (unlike Paracloeodes and some Baetungulata – see Index of characters [1.1.3]).

() Prostheca of right mandible is bifurcate, with longer branch directed proximally [on left mandible massive stick-shape – see Turbanoculata (6)]. The same in some Centroptella/g1 and some other taxa (see Index of characters [1.1.25]).

() Larval abdominal terga with scales in operculate sockets [see Turbanoculata (12)] (see Index of characters [1.3.4]).

Plesiomorphies of Cloeodes/g1. Larval paracercus is allways developed.

Unlike Centroptella, patella-tibial suture [with conspicuous row of trichobothria – see Cloeodes/g1 (1)] on anterior side of tibia has usual oblique shape, being equally developed on all legs [see Anteropatellata (1)].

In mature larva ready to moult to subimago, buds of subimaginal gonostyli [see Baetovectata (3)] are folded under larval cuticle by the "Nigrobaetis-type" (2nd segment directed posteriorly and curved by its convexity medially and by apex laterally) (see Index of characters [2.3.9]).

Variable characters of Cloeodes/g1. 

() Ability to make rhythmical respiratory movements by tergalii (see Index of characters [1.3.30]) is either present (e.g., in nigrohumeris [Cloeodes]), or absent (Kluge 2017).

?Maxillary palp can be 3-segmented or 2-segmented. 

Claws [initially with two rows of denticles – see Tetramerotarsata (11)] can have two rows of prominent or poorly developed denticles (in selected Centroptella), or lack denticles (in Cloeodes/g1 and selected Centroptella). 

() On fore wing marginal intercalaries are situated mainly in twos in each field [see Baetovectata (1)]; but sometimes, especially in females, one of two veins is lost, and in each field a single eccentric intercalary is present. 

() Hind wings can be present or lost; in the last case vestiges of hind protoptera can be present or lost.

Size. Fore wing length 2–10 mm (see Tetramerotarsata).

Distribution. Neotropical, Oriental and Afrotropical Regions.


Nominal species in Cloeodes/g1:

aiuruoca Massariol & Angeli & Salles (in Salles & Massariol & Angeli & Lima & Gattolliat & Sartori) 2015 [Cloeodes]

amantykyra Massariol & Angeli & Salles (in Salles & Massariol & Angeli & Lima & Gattolliat & Sartori) 2015 [Cloeodes]

anduzei Traver 1943 [Pseudocloeon]  

atlanticus Massariol & Angeli & Salles (in Salles & Massariol & Angeli & Lima & Gattolliat & Sartori) 2015 [Cloeodes]

auwe Salles & Batista (in Salles & Batista & Soares-Cabette) 2004[Cloeodes]  -

aymara Traver 1971 [Baetis]  

aymore Massariol & Salles 2011 [Cloeodes]

barituensis Nieto & Richard 2008 [Cloeodes]  

binocularis Needham & Murphy 1924 [Pseudocloeon]  

boldrinii Massariol & Angeli & Salles (in Salles & Massariol & Angeli & Lima & Gattolliat & Sartori) 2015 [Cloeodes]

caraibensis Hofmann & Thomas (in Hofmann & Sartori & Thomas) 1999 [Cloeodes]  

consignatus Traver 1938 [Cloeodes]  

espinillo Nieto & Richard 2008 [Cloeodes]  

excogitatus Waltz & McCafferty 1987 [Cloeodes]  

guara Massariol & Angeli & Salles (in Salles & Massariol & Angeli & Lima & Gattolliat & Sartori) 2015 [Cloeodes]

hydation McCafferty & Lugo-Ortiz 1995 [Cloeodes]

inferior Kluge 1991 [Cloeodes]  --/

ioachimi Massariol & Angeli & Salles (in Salles & Massariol & Angeli & Lima & Gattolliat & Sartori) 2015 [Cloeodes]

irvingi Waltz & McCafferty 1987 [Cloeodes (Cloeodes)]  

itajara Massariol & Salles 2011 [Cloeodes]

jaragua Salles & Lugo-Ortiz 2003 [Cloeodes]  

lucifer Massariol & Angeli & Salles (in Salles & Massariol & Angeli & Lima & Gattolliat & Sartori) 2015 [Cloeodes]

macrolamellus Waltz & McCafferty 1987 [Cloeodes]  

maculipes Traver 1938 [Cloeodes] — typus nominis Cloeodes  

magnus Massariol & Angeli & Salles (in Salles & Massariol & Angeli & Lima & Gattolliat & Sartori) 2015 [Cloeodes]

maracatu Lima & Pinheiro & Massariol (in Massariol & Lima & Pinheiro & Quieroz & Oliveira & Salles) 2013 [Cloeodes]  

melanotarsus Massariol & Angeli & Salles (in Salles & Massariol & Angeli & Lima & Gattolliat & Sartori) 2015 [Cloeodes]

nigrohumeris Kluge 2017 [Cloeodes] --/

opacus Nieto & Richard 2008 [Cloeodes]

penai Morihara & Edmunds 1980 [Notobaetis] — typus nominis Notobaetis  

peninsulus Waltz & McCafferty 1987 [Cloeodes]  

redactus Waltz & McCafferty 1987 [Cloeodes (Cloeodes)]  --/

spaceki Queiroz & Oliveira & Salles (in Massariol & Lima & Pinheiro & Quieroz & Oliveira & Salles) 2013 [Cloeodes]  

superior Kluge 1991 [Cloeodes]  --/

tracheatus Massariol & Angeli & Salles (in Salles & Massariol & Angeli & Lima & Gattolliat & Sartori) 2015 [Cloeodes]

turbinops Needham & Murphy 1924 [Pseudocloeon]  

vibratorius Kluge 2017 [Cloeodes] --/

waltzi Wiersema & Baumgardner 2000 [Cloeodes]

xyrognathos Massariol & Angeli & Salles (in Salles & Massariol & Angeli & Lima & Gattolliat & Sartori) 2015 [Cloeodes]