Corresponding author: Carlos Molineri (
Academic editor: E. Dominguez
The Neotropical species of
Se revisan las especies neotropicales de
Molineri C, Salles FF, Peters JG (2015) Phylogeny and biogeography of Asthenopodinae with a revision of
The family
The main objective of the present paper is to evaluate in a phylogenetic context the validity of the generic groups of
Additionally we describe and illustrate some unknown stages of previously known species, propose a key to the subfamilies and genera of
Material is deposited in the following Institutions: CUIC (Cornell University Insect Collection, Ithaca, NY), FAMU (Florida A&M University, Tallahassee, FL), IFML (Instituto-Fundación Miguel Lillo, Tucumán), IBN (Instituto de Biodiversidad Neotropical, Tucumán), MACN (Museo Argentino de Ciencias Naturales, Buenos Aires), MECN (Museo Ecuatoriano de Ciencias Naturales, Quito), MUSENUV (Museo de la Universidad del Valle, Cali), RBINS (Royal Belgian Institute of Natural Sciences, Brussels), FCE-Ep (Facultad de Ciencias, Entomología, Montevideo, Uruguay), CZNC (Coleçao Zoológica Norte Capixaba, São Mateus, Espírito Santo), INPA (Instituto Nacional de Pesquisas da Amazônia, Manaus), and ZMH (Zoologisches Museum Hamburg).
Characters are scored from external morphological features of adults (male imago unless otherwise indicated), nymphs and eggs. Dissected parts of the nymphs and adults were mounted on microscope slides using Canada Balsam, except wings that were mounted dried. All the material is preserved in ethyl alcohol 96%. Photographs were taken using a NIKON SMZ-10 stereomicroscope or a microscope, with a Nikon D5000 digital camera; some pictures were modified with Combine ZP (
The measures and ratios used in some characters are explained and illustrated to permit repeatability. To score variation in shape of genitalic structures (chars. 12–17) some measures were defined (see Appendix
A matrix of 17 taxa and 72 characters was constructed (Appendix
Outgroups were scored either from fresh material (
Searches were conducted in TNT (
All the available geographic records (448 records for 17 taxa) of the species or genera included in the phylogeny were compiled. Concerning
The biogeographical analysis was performed through spatial analysis of vicariance (
Parsimony under implied weights (IW) resulted in a single tree (Figs
The type species of
Most parsimonious tree with changes on continuous characters (number above circles are character numbers, see Appendix
Most parsimonious tree (same for Fig.
Group support and vicariant nodes. Numbers above branches indicate GC values (400 replicates, Symmetric Resampling), below branches Relative Bremer supports (from 2000 trees, up to 8 steps longer) are indicated. Support values higher than 50 are bolded. Black squares indicate nodes with allopatric descendants (see text for explanation).
Summarizing, the taxonomic changes needed to adjust formal classification with the phylogeny include: 1) revalidation of
An additional taxonomic change is proposed for
The subfamily
It is interesting to note some characters that simultaneously change in opposite “directions”, showing a certain tendency in
The sister relation between
The clade formed by
In spite of
Variation in the relative length of male foreleg segments has been repeatedly used by different authors in the taxonomy of the family. One of our characters concerning this variation (char. 1, ratio length tarsal segment 2/tibia) presented many changes: most important are those defining the genus
In the nymphs, the robustness of the mandibular tusks is an important feature distinguishing different subfamilies. The optimization of character 27 (ratio length/width of tusk, Fig.
Trends in some charecteristics are markedly manifested, for example the width of male foretibia (char. 0 ratio width tibia/tarsal segment 2, Fig.
The length of male foreleg is another character commonly used to define groups (here represented as the ratio length of FW/foreleg, Fig.
General shape of the forceps in male adults shows opposing tendencies, becoming slender in
The spatial analysis of vicariance found 6 disjoint sister pairs (Fig.
A key to the subfamilies of
Key to the species of
1 | FW with vein Sc ending before the tip of the wing, its apical portion not curved posteriorly; cubital area of FW broadly expanded, usually with 3–5 intercalaries and many cross veins and marginal intercalaries; FW with MA fork 1/3 or more distance from wing base (i.e., a long MA stem is present); HW with convex intercalary between R1 and Rs field (x.i. in Fig. |
|
– | FW (e.g., Figs |
|
2 | Pronotum ring-like (ratio width/length ca. 1/3); FW with CuA relatively straight, ICus subparallel (e.g., Figs |
|
– | Pronotum longer (width similar to length), in male with triangular anterior portion; CuA sigmoid, ICus apically diverging; median remnant of styliger plate absent; female forelegs generally absent in imago (may be present in subimago), if complete much reduced in size and twisted; eggs C-shaped, resembling a sphere with one side pushed in, polar cap single if present, chorion punctuated at least in the concave face | |
3 | Eyes of male enlarged, separated on meson of head by a distance subequal to width of lateral ocellus |
|
– | Eyes of male normal, similar to female, separated on meson of head by a distance greater than 2 times the width of lateral ocellus |
|
4 | Penes blade-like (Fig. |
|
– | Penes variable but commonly cylindrical (Figs |
Neotropical |
5 | Penes with many spines on outer subapical margin (Fig. |
|
– | Penes with smooth outer margin (Figs |
|
6 | Foreleg (FL) much shorter in length than FW (ratio FW/FL length = 1.4–2.0), foretarsal segment 1 distinct (Fig. |
|
– | FL subequal in length to FW (ratio FW/FL length: 1.0–1.2); 3–5 crossveins between R and M, basal to R fork; foretarsal segment completely fused with tibia (Fig. |
|
7 | Penes with a short apical spine-like projection (arrow in Fig. |
|
– | Penes apically rounded or slightly pointed but never with a spine (Fig. |
|
8 | Legs II and III flap-like without tibia and tarsi, in both sexes; forceps 1-segmented; eggs with one polar cap (some species show the cap on the convex face) | |
– | Legs II and III weak and twisted, but complete; forceps 2-segmented (a weak line separates a short basal segment); eggs without polar caps (a long thread coiled around the egg is present in one species, |
|
9 | Male sternum IX not divided medially along its length; penes fused basally; pedestal with relatively short parastylus; female FW with IRs complete; sockets on sternum VIII small and submedian | |
– | Male sternum IX divided medially along its length; penes completely separated; pedestal with long and curved parastylus; female FW with IRs incomplete (2 IR veins are wanting); sockets on sternum VIII larger and sublateral |
1 | Outer edge of mandibular tusks with many large tubercles; tarsus and tibia of foreleg separated |
|
– | Outer edge of tusks without large tubercles (Figs |
|
2 | Dorsum of head mostly glabrous, without large tufts of tightly grouped short setae (Figs |
|
– | Dorsum of head with dense patches of short setae, mainly anteriorly to lateral ocelli; occiput flat, subquadrate in dorsal view; apex of left mandibular tusk with 1 pointed process | |
3 | Mandibular tusks relatively long and slender, without tubercles or large spines on inner margin (Figs |
|
– | Mandibular tusks robust, stout (Figs |
|
4 | Mandibular tusks without large subbasal tubercle on inner margin; foretarsal claw with double row of denticles (Fig. |
|
– | Mandibular tusks with a large subbasal tubercle on inner margin (arrows in Fig. |
|
5 | Apex of left mandibular tusk with 4 pointed processes (Ethiopian and Oriental) |
|
– | Apex of left mandibular tusk with 3 pointed processes (Fig. |
|
6 | Outer margin of mandible with a tooth-like indentation (Fig. |
|
– | Outer margin of mandible smooth; abdominal gill I uniramous |
|
7 | Mandibular tusks with prominent basal or sub-basal tubercle on median margin (rarely tubercle absent), from some to many apical crenulations, numerous setae on outer margin of mandibles; abdominal gill I bifurcated |
|
– | Mandibular tusks with 1 or 2 prominent tubercles on distal third of median margin, few long setae on outer margin of mandibles; abdominal gill I single |
|
8 | Mandibular tusks with 2 tubercles (submedian and subapical) on median margin; distal projection of foretibia-tarsus 2/5 the length of claw |
|
– | Mandibular tusks with a single subapical tubercle on median margin; distal projection of foretibia-tarsus 2/3 the length of claw |
|
Shortly after,
The inclusion of many of these characters in a formal cladistic analysis permitted us to recognize apomorphic from plesiomorphic states, showing that some of the hypothesized synapomorphies for
The revalidation of
Two fossils may falsify these hypotheses.
The second fossil with contradictory information,
Five groups (
But perhaps the most interesting biogeographic pattern found in the present work is the paralogy in tropical South American areas caused by the paraphyletic nature of the Neotropical
Eight autapomorphies define the genus
Male imago. Length (mm): body, 7.0–7.8; FW, 7.5–8.6; HW, 3.1–3.7; foreleg, 6.2–6.9; cerci, 21.6. Antennae: scape slightly longer than pedicel; flagellum bristle-like. Thorax. Pronotum width/length: 1.5–2.5. Legs. Forelegs relatively long, ratio length FW/foreleg = 1.1; tarsal segment 1 fused to tarsal segment 2 (Fig.
Female adult. Length (mm): body, 10.2–10.8; FW, 11.1; HW, 4.3. Thorax. Pronotum width/length = 1.5–2.3. Wings with more crossveins and intercalaries than male. Abdominal sternum VIII with paired anteromedian sockets on an oval and ventrally protruding structure, sockets small, shallow and contiguous. Terminal filament reduced, shorter than tergum VIII, with few thin annuli; cercus 0.5–0.6 times the length of abdomen.
Eggs (Fig.
Nymphs (Fig.
Amazonas subregion (Amazonas river in Colombia and Brazil).
Listed in
Nymphs (Fig.
Amazonas river, from Leticia (Colombia) to Manaus (Brazil).
Male imago. Length (mm): body, 5.0–8.0; forewing, 5.7–8.4; hind wing, 2.6–3.9; foreleg, 3.1–4.5; cerci, 19.0–25.0. Pronotum width/length: 2.1–2.5. Wings (Figs
Female adult. Length (mm): body, 6.5; FW, 8.9; HW, 3.3; cerci, 1.5. Pronotum ring-like. Wing venation (Figs
Eggs (Figs
Arbitrary combination of letters.
Treating this sole species in a new genus, distinct from
Paratype male (IFML TEPH095, slide 041) from URUGUAY, Artigas, San Gregorio, orillas río Uruguay, 29.xi.1959, light trap, C.S. Carbonell col.; 5 male imagos (slide IBN141CM) and 1 male and 1 female (slide IBN471CM) subimagos (IBN) from COLOMBIA, Amazonas, P. N. Amacayacu, río Amacayacu, 93 m,
Male imago. See generic section above and original description in
Female subimago. Length (mm): body, 6.5; FW, 8.9; HW, 3.3; cerci, 1.5. General coloration yellowish white with gray markings. Head cream extensively shaded gray dorsally, the shading is uniform anteriorly but in the form of a fine netted pattern posteriorly to lateral ocelli, occiput with a pair of submedian pale anterior spots and a pair of submedian dark posterior spots; venter of head whitish. Antennae yellowish white shaded with gray on scape and pedicel. Thorax cream. Anterior ring of prothorax very thin, less than 1/4 the dorsal length of posterior ring; ratio width/length: 2.6; pronotum shaded blackish on median area except pale medial line, presternum paler, shaded gray before coxa. Mesonotum shaded very diffusely with gray, darker on longitudinal carinae and between posteroscutal protuberances; mesosternum and pleurae paler, shaded gray on anterior corner of katepisternum. Metanotum shaded gray on posterior half, except on a pale median triangular mark, shaded darker posteriorly to this pale mark; metasternum whitish. Legs whitish except coxae yellowish shaded gray and apex of hind trochanter pointed and yellowish orange. Wings (Figs
Eggs. See generic description.
This species presents a wide geographic range that spans from the Amazon River in the North to the Uruguay River in the South, also extending towards the West in Bolivian Chiquitania.
None.
In the forewings of
Seven autapomorphies define the genus
Male imago. Length (mm): body, 7.3–13.5; FW, 7.0–14.5; HW, 3.7–7.3; foreleg, 5.0–14; cerci, 20.0–38.5. Antennae: scape subequal to pedicel; flagellum bristle-like. Thorax. Pronotum width/length: 1.2–1.9. Legs. Forelegs relatively long, ratio length FW/foreleg = 1.0–1.6; tarsal segment 1 fused to tibiae (Fig.
Female length (mm): body, 7.2–19.0; FW, 12.2–22.5; HW, 5.3–11.5; cerci 1.2–4.0. Thorax. Pronotum width/length = 1.5–2.3. Wings with crossveins and intercalaries more numerous than in male. Abdominal sternum VIII (Fig.
Eggs (Fig.
Nymphs, nearly mature (Fig.
Amazonas and Parana subregions (Argentina, Brazil, Colombia, Guyana, Uruguay).
The characteristics traditionally associated with
The revalidation of the genus
1 | Large species (FW > 14 mm); wings smoky yellowish (Fig. |
|
– | Smaller species (FW < 12 mm); wings hyaline, yellowish areas sometimes present along hind margin; penes much less curved and not twisted (Figs |
|
2 | FW 11.4–11.9 mm; genitalia as in Fig. |
|
– | Smaller species, FW 7.0–8.8 mm; genitalia as in Fig. |
|
1 | FW with single imv in most spaces (Figs |
|
– | FW with double or triple imv (Fig. |
|
2 | Wings (Fig. |
|
– | Wings hyaline (Fig. |
|
Type material was not studied; it consists of the holotype male imago, damaged, with many parts missing including the genitalia. It is a pinned specimen deposited at Naturistorisches Museum Wien,
Male imago. Length (mm): body, 9.3–11.2; FW, 11.4–11.9; HW, 5.8–6.2; foreleg, 10.0–10.3; cerci, 30.1. Described in
Female adult. Length (mm): body, 11.5–12.6; FW, 14.5–19.0; HW, 6.3–9.4. General coloration yellowish light brown. Head dorsally blackish except on median zone, paler; venter of head yellowish white. Antennae light brown, shaded gray on scape. Thorax yellowish brown with blackish membranes, shaded with brownish gray on pronotum and with black on posteromedian marks on meso- and metanotum. Pronotum width/length: 1.5–2.3. Legs whitish yellow shaded brownish on dorsum of leg I and on apex of femur III. Wings (Fig.
Eggs (Fig.
Uruguay, Argentina.
Holotype (IBN) male and paratypes (IBN) 4 males and 22 females (slides IBN468–469CM) from ARGENTINA, Misiones, Parque Provincial Urugua-í, Arroyo Uruzú, 7−11.xii.1999, C. Molineri col.; 1 female imago (“
Non-type material: 1 female (wings on slide) from BRAZIL, Sao Paulo, Jacareí, rio Paraiba do Sul, 21.xi.1987, CG Froehlich et al. cols (deposited at MZSP).
Four autapomorphies characterize this species, all are small changes in continuous characters, except the marked shortening of the median remnant of styliger plate (at the middle, since laterally a tong-like projection is present, Fig.
Male imago. Length (mm): body, 12.2–13.5; FW, 14.0–14.5; HW, 6.8–7.3; foreleg, 12.3–14.0; cerci, 35.5–38.5. General coloration yellowish white. Head shaded black dorsally almost entirely, with a pair of distinct submedian black marks anteriorly to median ocellus; occipital hind margin with pale median zone; head ventrally pale without markings. Antennae: scape and pedicel short, yellowish on venter of pedicel, both slightly shaded with gray; flagellum very thin, hyaline, similar in length to forefemur. Thorax. Pronotum translucent, shaded slightly with gray except on membranes separating anterior and posterior rings, darker laterally; pronotum width/length: 1.3–1.9. Meso- and metanotum yellowish white with gray markings mainly posteriorly but also on sutures. Thoracic pleurae and sterna yellowish white shaded gray only at base of coxae and wings. Legs. Forelegs: coxa dorsally whitish with a gray mark, ventrally yellowish; femur dorsally yellowish shaded gray on apical third, ventrally whitish; tibia whitish translucent shaded gray mainly on dorsum; tarsi translucent shaded slightly with gray; large claws, stalk with brown inner margin, rest whitish; articulations between femur-tibia and tibia-tarsus very sclerotized, brownish. Middle and hind legs yellowish, shaded with gray from half of femur to apex of leg. Wings (Fig.
Female subimago. Length (mm): body, 9.5–19.0; forewing, 15.5–22.5; hind wing, 7.5–11.5; cerci 3.5–4.0. General coloration dark brown shaded widely with black. Head black dorsally, yellowish white ventrally except on remnants of tusks, brownish. Antennae dark brownish except apical half of flagellum whitish. Thorax. Sclerites dark brown, membranes shaded black. Pronotum width/length: 2. Legs brownish except membranous portions, whitish. Wings (Fig.
Eggs (Fig.
The species is dedicated to the great mayfly specialist Jay R Traver, who visited Uruguay and worked with Mr. C. S. Carbonell’s collections at the “Museo de la República” recognizing the females of this species as distinct from
Parana biogeographic subregion in Argentina, Uruguay and Brazil.
Females subimagos were collected (in Misiones Province) while swarming in compact groups at about 3 m above water in pool areas around sunset. The same behavior was reported by
Holotype and 3 paratypes male imagos from Brazil, Amazonas, Barcelos, rio Demene, ´boca´barco,
Male imago. Length (mm): body, 7.3–8.0; forewing, 7.0–8.8; hind wing, 3.7–4.3; foreleg, 5.0–5.8; cerci, 20.0–23.0. General coloration yellowish white. Head shaded gray dorsally almost entirely, frons with black dot at base of antenna, with medial line and irregular black marks; occipital region with pale median zone; head ventrally pale without markings. Antennae: scape and pedicel short, subequal in length, whitish shaded with purplish; flagellum very thin, hyaline. Thorax. Pronotum whitish with anterior and posterior portion subequal in size, shaded gray in a transverse band between both portions, posterior portion black along hind and lateral margins; pronotum width/length: 1.2–1.3. Mesonotum yellowish white shaded with gray on posterior half of medial line, on area between posterolateral protuberances and on anterior margin of these structures. Metanotum yellowish slighlty shaded with gray medially. Thoracic pleurae and sterna yellowish white shaded gray dorsally and anteriorly to mid coxa. Legs. Forelegs: whitish completely shaded with gray; large claws, apically expanded (Fig.
Note: Cornell male (slides, male imago). Length (mm): body missing; FW, 9.0; HW, 4.5. FW with 11 long marginal intercalary veins; 3 cross veins between R and M basad to R stem; IMP fused basally to MP1 or free; MP2 fused to IMP. HW with 6 long intercalary veins on hind margin. Genitalia: penes robust, twisted; median remnant of styliger plate without posterolateral projections; forceps long and slender, ratio length/basal width: 6.3–6.7.
Female imago. Length: body, 7.2 (shrunken, empty)–12.3; FW, 12.2–14.8; HW, 5.3–6.0; cerci, 1.2–1.3. General coloration dark brown. Head dorsally black except on clypeus, whitish with a pair of lateral brownish bands, ventraly much paler brownish white. Thorax brownish with blackish membranes and carinae. Pronotum with a pair of distinct black marks submedially; width/length ratio: 2. Legs pale, brownish white. Wings (Fig.
Eggs. Length 210–240 µ, width 180–200 µ. Subovate, yellowish, with two small whitish polar caps (maximum width, 75–85 µ), polar caps much thinner than the egg and formed by 14–16 threads. Under SEM the larger disk-like chorionic structures are surrounded by many smaller ones, which at their time are surrounded by smooth chorion (Fig.
Nymphs. Length (mm): body, 7.8; cerci, 2.0–2.3; terminal filament, 3.1. General coloration yellowish light gray (Fig.
“Chumuco” is one of the common names applied to river cormorans in some South American countries. The penis lobe of this new species resembles the neck and head of that bird.
Brazil (Amazonas, Espírito Santo), Colombia (Amazonas), Guyana.
The male imago of this species has been known since
Five autapomorphies were recovered for
Male imago. Length: body, 6.5–10.5; FW, 7.0–10.1; HW, 2.9–4.8; foreleg, 5.1–7.5; cerci, 22.0–37.0. Antennae: scape slightly longer than pedicel, flagellum bristle-like. Thorax. Pronotum width/length: 1.7–2.4. Legs. Forelegs subequal to shorter than body, ratio length FW/foreleg 1.4–1.8; longest segment is tibia (ratio length tarsal segment 2/tibia = 0.4–0.7); tarsal segment 1 distinct, not fused, very short (Fig.
Female adult. Length: body, 8.0–19.5; FW, 12.0–18.5; HW, 4.6–7.8; cerci, 3.0–7.0. Thorax. Pronotum width/length = 2–3. Wings with more crossveins and intercalaries than in male. Abdominal sternum VIII with anteromedian keel (Fig.
Eggs (Fig.
Nymphs. Length (mm): body, 9.7–15.0 mm; cerci, 4.0–7.0; terminal filament, 5.0–5.1. Head suboval in dorsal view, smooth (without pilose area); occipital region well developed, strongly convex (Figs
Foreleg of male imago, detail of articulation between tibia and tarsus:
Amazonas and Parana biogeographic subregions (Argentina, Bolivia, Brazil, Colombia, Ecuador, Peru).
The genus
Male
1 | Penile lobe (distad to basal thumb) with a similar width along its length, basal thumb separated by a wide furrow (Figs |
|
– | Penile lobe (distad to basal thumb) wider basally, basal thumb fused to penile lobe (Fig. |
|
2 | Apical spine of penes long and acute (Fig. |
|
– | Apical spine of penes short (Fig. |
|
3 | Penes long, apical spine slightly marked, median remnant of styliger plate projecting laterally (Fig. |
|
– | Penes short, apical spine well marked, median remnant of styliger plate normal (Fig. |
|
4 | FW 9.5–10.1; penile lobe strongly widened basally (ratio length / basal width = 2.9, Fig. |
|
– | FW 7.0–9.5 mm; penile lobe not so wide at the base (ratio length / basal width = 4.0–5.0, Fig. |
|
Female and eggs of
1 | Disk like structures on the equatorial area relatively well separated from each other, separation about 0.6 or more of maximum width of a disk (Fig. |
|
– | Disk like structures on the equatorial area almost touching each other, maximum separation about 0.3 or less of maximum width of a disk (Fig. |
|
2 | With a group of 2–3 very small disks beneath each disk like structure (Fig. |
|
– | Only smooth chorion below the disk like structures (Fig. |
|
Nymphs (only 3 species known, almost undistinguishable, the characters below should be confirmed with the study of more material)
1 | On the inner margin of left mandibular tusk, the space between the subbasal and the submedian tubercles is short and strongly concave (Fig. |
|
– | On the inner margin of left mandibular tusk, the space between the subbasal and the submedian tubercles is longer and straighter (Figs |
|
2 | Ratio total length of mandible/mandibular tusk length: 1.59–1.62 (Fig. |
|
– | Ratio total length of mandible/mandibular tusk length < 1.5 (Fig. |
Photographs of the type at the British Museum were studied.
Male imago. Length: body, 8.0–8.7; FW, 10.0; HW, 4.4; foreleg, 7.5; cerci, 33.0–35.0. General coloration yellowish light brown. Head whitish, heavily shaded black dorsally, paler on posteromedian zone of occiput, black shading extending anteriorly on frons as two parallel lines surrounding median ocellus. Antennae pale, slightly shaded gray on dorsum. Thorax. Pronotum yellowish translucent completely shaded gray, darker on anterior ring; paler on two transverse lines, one separating anterior and posterior rings and another more posterior and obliquely transverse; pleurae shaded with black, sternum with a median gray macula. Pronotum width/length ratio: 2.0–2.3. Mesonotum whitish yellow (or brownish in some males) with a black median triangle between posteroscutal protuberances, metanotum similar in color, also shaded black posteromedially; mesopleurae and sterna paler, shaded with black along anterior margin of katepisternum. Legs yellowish white shaded with gray dorsally on all coxae, femora and tibiae; foretarsal segment 1 blackish (Fig.
Female adult. Length: body, 10.5–13.2; FW, 14.0–18.5; HW, 5.7; cerci, 5.8–7.0. Morphologically very similar to female adults of
Eggs (Fig.
Mature nymph. Length of male: body, 9.5–9.7; cercus, 7.0; terminal filament, 5.0. Length of female: body, 17.0; cercus, 8.0; terminal filament, 7.0. Only characters that differ from
Amazonas River from Leticia (Colombia) to Manaus (Brazil).
Much confusion exists in the literature concerning this species. Many authors mention
As the result of the present study, the female adult, egg, and nymphal stages are described here for the first time. Previous descriptions of female and nymphs in the literature were done from specimens of
Holotype (IBN) male imago from Ecuador, Napo Province, Laguna Limon Cocha, 250 m, 6.iv.1984, E. Domínguez col.(aprox.
Male imago. Length (mm): body, 9.0–10.5; FW, 9.0–10.1; HW, 4.0–4.8; leg I, 6.5–7.4; cerci, 37.0. General coloration yellowish light brown. Head whitish shaded black dorsally on pale median mark on hind margin and along inner margin of eyes; frons pale except paired submedian black lines. Antennae: whitish shaded diffusely with gray on scape and apex of pedicel; length (mm): scape 2.25, pedicel 1.5, flagellum 7.25. Thorax. Pronotum yellowish translucent shaded with black dorsally except at pale median membrane between both pronotal rings, on mediolongitudinal line and along margins; the black shading presents many scattered and small pale spots. Pronotum width/length: 1.7–2.4. Meso- and metanotum yellowish shaded with grayish on carinae, posteromedian triangular mark (on mesonotum), and scutellum (both). Thoracic pleurae and sterna paler, shaded gray on pleural sclerites. Legs yellowish white, shaded gray on all coxae. Leg I shaded gray almost completely, stronger on femur and tibia, paler on tarsal segments (Fig.
Female imago. Length (mm): body, 15.5–19.5; FW, 16.0–17.5; HW, 5.2–7.8; cerci, 3.5–5.0. Pronotum width/length: 2. Morphologically very similar to
Eggs (Fig.
Nymphs. Length of male (mm): body, 10.0–11.0 mm; cerci, 7.0–8.0; terminal filament, 5.0–5.5. Length of female (mm): body, 17.0–20.0 mm; cerci, 4.0–5.0; terminal filament, 5.0. General coloration brownish. Head (Fig.
From Latin “magnus” meaning “large”, noun in apposition. The name alludes to the general size of the individuals, mainly the female adults.
Only known from the type locality in Napo (Ecuador).
The type series described here as
holotype male imago (slide IBN479CM) from Colombia, Amazonas, Puerto Nariño, Loreto Yacu,
Male imago. Length (mm): body, 7.0–7.1; FW, 7.8–9.2; HW, 3.3–4.0; leg I, 5.1–5.2; cerci, 22.0–23.0. General color whitish brown. Head whitish shaded black dorsally except thin line along hind margin, with a pair of blackish short lines anteriorly to median ocellus. Antennae whitish shaded gray at margins of scape. Thorax. Pronotum ratio width/length: 2.1. Pronotum yellowish translucent widely shaded black, except on pale transversal line between anterior and posterior rings, and on mediolongitudinal line of posterior ring, and posterolateral oblique dashes. Meso- and metanotum whitish yellow shaded black on carinae and margins, also shaded on black on posteromedian triangular zone. Legs whitish shaded gray on dorsum of foreleg, all coxae, and on legs II–III on apex of tibiae and dorsum of tarsi; foretarsal segment 1 is shown in Fig.
Female imago. Length (mm): body, 10.5; FW 13.0; HW, 5.0, cerci 3.8. Similar to male, shaded dorsally more uniformly and markedly. Pronotum ratio width/length: 2.7. FW with 4 cross veins between R and M basal to R stem (none of them just below fork). Abdominal sternum VIII with keel as in Fig.
Eggs (Fig.
The species is named for Mike Hubbard who has contributed significantly to the understanding of mayflies throughout the world.
Two near localities in the Amazonas River from Colombia.
This species is very similar to
Holotype male imago (slide IBN473CM) from Argentina, Corrientes, Parque Nacional Mburucuya, Selva Misionera (sector 6), luz, 29.iii.2001, F. Navarro col.
Male imago. Length (mm): body, 6.5–9.0; FW, 8.0–9.0; HW, 3.6–4.0; foreleg, 5.1–5.4; cerci, 25.4–28.0. General coloration yellowish white. Head whitish shaded with black dorsally except on hind margin and posteromedian pale mark; frons pale shaded with a pair of black submedian longitudinal lines; venter of head pale. Antennae whitish shaded with gray on scape; length (mm): scape 1.75, pedicel 1.25, flagellum 7.5. Thorax. Pronotum whitish translucent, shaded black on anterior ring and lateral margins; posterior rings shaded gray, with darker mediolongitudinal line. Meso- and metanotum yellowish white, shaded gray on scutellum; sterna pale shaded gray only on mesokatepisternum. Legs whitish shaded gray on coxae. Foreleg completely shaded gray, paler on base of tarsal segments 2–5 (Fig.
Female subimago. Length (mm): body, 12.5; FW, 16.0; HW, 6.5; cerci broken off and lost. General coloration orangish yellow shaded widely black. Head dorsally blackish except medial line on occiput, and anteriorly to median ocellus. Thorax. Pronotum width 2.5 mm, total length 0.8 mm; cream shaded black on thin anterior ring, with gray on posterior ring, membranes whitish. Mesothorax orangish yellow with gray markings. Wings translucent whitish, veins whitish except basal half of C, Sc and R yellowish. Abdomen uniformely shaded gray dorsally, except on pale medial line. Sternum VIII with long and thin anteromedian keel. Base of caudal filaments whitish (rest broken off and lost).
Eggs (Figs
Nymph (cuticle from reared female described above). Length (mm): body, 16.5 mm; cerci and terminal filament, 7.0 (both broken at apex). Antennae broken off and lost. Mouthparts. Mandibular tusks with relatively large space between the large basal tubercle and smaller subdistal tubercle, not C-shaped (similar to Fig.
The name refers to one of the etnic groups inhabiting the area where the specimens were collected.
Argentina (Corrientes), Brazil (Sao Paulo), Uruguay (Salto).
This species is very distinctive, not only by the long and slender penis lobe, but also because of the presence of relatively long marginal intercalary veins (in male FW and HW). The reared female subimago is tentatively associated with the male, because of similarity in coloration and shared distributional range. Eggs extracted from this female show also some differences from the other known in the genus, mainly the larger extent of smooth chorion around the plates and the presence of small disks below the larger ones (Fig.
(see
Only one autapomorphy was recovered in our analysis for
Argentina, Bolivia, Brazil, Colombia and Peru.
This species was recently described from all the stages (
Optimization of selected continuous characters.
Optimization of selected continuous characters.
World map indicating the distribution of the three subfamilies in
Map of South America (in part) showing the distribution of
Map of South America (in part) showing the distribution of
We acknowledge Dr. Marco Guimarães and Jairo Oliveira from the Laboratório de Ultraestrutura Celular Carlos Alberto Redins (LUCCAR), CCS/UFES, as well as edital MCT/FINEP/CT–INFRA - PROINFRA 01/2006, and to Centro Integral de Microscopía Electrónica (CIME), CONICET, Tucumán, Argentina for the SEM images. Special thanks to Michel Sartori for loan of out-group material, Paula Souto for photographs of
List of continuous characters and their definitions. Characters were scored from male imagos unless otherwise indicated.
List of apomorphies from terminals and nodes. Arrows separate plesiomorphic from apomorphic states.
Matrix of characters and states. Ready to use in TNT.
nstates cont;
xread
‘
72 17
&[continuous]
Ephoron 1.211 0.400 0.932-1.081 1.008-1.123 1.857-1.912 1.846-2.167 16.000 4.188 3.000 0.650 0.250 1.273 5.455 11.667 ? ? ? ? ? 0.778-0.992 1.322-1.551 0.727-1.625 2.000-5.000 ? 1.625 ? ? 4.145
Camp_violaceus 1.765 0.357-0.360 1.758-2.179 0.371-0.436 2.082-2.273 2.236-2.346 2.000-6.000 1.000-1.667 1.000-3.000 1.580-1.830 0.205-0.261 1.333-1.440 ? 19.000-23.000 2.308-2.552 2.727-2.846 1.304-1.500 2.600 1.118-1.410 0.287-0.375 1.091-1.300 2.625-3.333 3.000-4.000 ? 1.712 1.542 1.148-1.443 3.688
Povilla 1.571-2.000 0.645-0.755 1.661-1.736 0.316 1.831-2.214 2.095-2.447 1.000-15.000 0.467-0.700 1.000-2.000 1.324-1.801 0.100-0.235 3.333-3.928 3.375-5.429 4.333-7.428 4.400-6.800 4.667-6.800 4.300-7.000 1.300 2.857-4.167 0.068-0.260 1.457-1.750 ? ? 13.000 1.555 1.787 1.439 2.015
Languidipes ? ? 3.174 0.375-0.464 2.447 3.382 5.000 1.667 1.000 1.653 ? ? ? 4.545 ? 9.333 7.000 ? 2.913 0.100 ? ? ? ? ? ? ? ?
picteti 0.838-0.840 1.205-1.350 1.118-1.150 0.355-0.382 1.828-1.983 1.828-1.983 14.000-17.000 2.000-3.353 4.000-6.000 0.595-0.733 0.337-0.373 1.550-1.875 8.100 6.714-7.400 2.737-2.895 8.000-8.696 3.056-3.059 1.083-1.182 1.500-2.250 0.184-0.206 1.383-1.591 1.222-1.516 3.778-5.167 6.000 ? ? ? ?
traverae 1.133-1.192 1.320-1.478 1.022-1.183 0.364-0.406 1.895-1.986 1.957-2.059 16.000-22.000 0.750-1.950 4.000-5.000 0.838-0.890 0.223-0.366 1.310-1.880 11.800 4.667-4.900 2.600-2.667 5.818-5.909 2.065-2.321 1.000 2.000-2.071 0.180-0.203 1.476-1.605 2.364-3.000 2.143-3.667 5.000-6.000 ? ? ? ?
chumuco 1.097-1.188 0.619-0.667 1.393-1.600 0.318-0.440 1.870-2.000 1.892-2.047 9.000-12.000 1.111-2.833 3.000-4.000 0.560-0.710 0.389 1.243-1.286 6.333-6.667 7.833-9.500 2.380-2.500 2.625-3.143 1.833-2.000 2.333-2.667 2.000-2.133 0.089-0.093 1.313-1.413 2.688-3.067 2.500-3.750 14.000-16.000 1.829-2.055 ? 1.258-1.226 1.978-2.412
crenulatus 3.250-4.000 0.741-0.825 1.087-1.150 0.329-0.347 2.027-2.091 2.419-2.654 10.000-14.000 1.400-1.429 2.000-4.000 1.164-1.795 0.241 1.550-2.462 4.286 8.889-9.000 5.909-6.500 5.000-5.417 3.250 0.600-0.684 2.188 ? 1.462-1.677 1.192-1.375 4.083-6.750 4.000-7.000 1.577 ? ? 2.196
gilliesi 1.411-1.909 0.523-0.652 1.556-1.975 0.300-0.350 1.781-2.079 2.054-2.308 4.000-10.000 0.250-1.167 1.000-2.000 0.780-1.098 0.191-0.308 2.143-2.500 4.500-6.143 6.333-7.692 1.905-2.067 3.647-4.706 1.177-1.905 0.944-1.5 2.647 0.169 1.455-1.833 0.892-1.071 2.800-6.167 14.000-16.000 ? ? ? ?
magnus 1.968-2.341 0.554-0.655 1.365-1.456 0.248-0.253 1.870-2.022 2.104-2.302 10.000-22.000 0.091-0.318 0.000-2.000 1.690-3.370 0.207-0.257 1.719-2.400 3.545-6.167 6.154-6.231 2.000-2.222 2.857-2.933 1.905-2.000 1.556-1.667 1.964-2.200 0.323-0.389 1.594-1.781 1.176-1.261 3.286-4.333 4.000-5.000 1.429-1.489 1.714-1.811 1.319-1.396 2.286-2.500
angelae 1.491-1.833 0.592-0.596 1.359-1.635 0.298-0.309 1.889-2.201 2.122-2.560 5.000-11.000 0.000-0.300 1.000-4.000 1.375-2.890 0.218-0.275 2.083-2.190 3.000-4.400 6.500-7.100 1.958-2.282 4.000-4.909 2.174-3.067 1.429-2.000 2.222-2.889 0.250-0.353 1.533-1.667 1.111-1.320 3.125-4.500 3.000-5.000 1.511-1.624 1.741-1.950 1.389-1.427 2.171-2.303
hubbardi 1.923-2.133 0.530-0.587 1.529-1.769 0.339-0.355 2.000-2.091 2.300-2.600 4.000-14.000 0.143-0.500 2.000-3.000 1.730-1.850 0.262-0.283 2.100 3.571-5.125 4.692-6.000 1.714-1.955 3.583-4.500 1.870-2.118 1.600-1.800 2.714 0.292 1.448-1.741 1.111-1.318 3.143-4.500 3.000-8.000 ? ? ? ?
guarani 1.926-1.974 0.429-0.579 1.381-1.765 0.268-0.354 1.744-2.000 1.829-2.500 4.000-6.000 0.000 1.000-2.000 1.170-1.369 ? 2.150 5.200 4.833-5.789 2.077-2.476 5.778-6.000 2.080-2.600 1.125 3.125 ? 1.250-1.600 1.231 4.333 3.000-8.000 1.408 1.818 1.351 2.630
curtus 1.550-1.751 0.584 1.365-1.457 0.248-0.286 1.870-2.022 2.104-2.341 18.000-25.000 0.222-0.316 0.000-1.000 2.100-2.950 0.171 1.964-2.320 3.222 5.385 1.786 3.846 2.273 2.167 2.706-3.059 0.378-0.500 1.419-1.545 1.128-1.341 4.000-6.286 3.000-5.000 1.573-1.729 1.714-2.371 1.395-1.618 2.182-2.419
Camp_vulturorum 1.864 0.395 1.987 0.370 1.983 2.271 9.000 1.000 1.000 1.825 0.168 1.000 ? 15.000 1.415 5.769 ? 1.071 1.146 0.257-0.319 1.224-1.294 1.833 ? ? 1.825 1.369 1.117-1.237 3.214
Tortopus 1.786 0.536-0.600 2.044-2.268 0.352 ? 2.190-2.514 1.000 0.000 2.000 1.500 0.194 ? ? 16.667 ? 6.364 ? 1.000 ? 0.277 1.226-1.333 ? ? ? 1.639 1.494 1.283 4.000
Tortopsis 1.300 0.533 2.714 0.375-0.417 2.169 2.568 0.000-1.000 0.000 1.000-3.000 0.926 0.222 ? ? 18.700 ? 11.923 ? 1.077 1.407 0.321-0.350 1.147-1.171 ? ? ? 1.562 ? 1.239 4.771