Research Article |
Corresponding author: Carlos Molineri ( carlosmolineri@gmail.com ) Academic editor: Marianna Simões
© 2021 Carlos Molineri, Lucimar G. Dias, María del Carmen Zúñiga.
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Abstract
The family Polymitarcyidae, with a worldwide distribution, includes mayflies with large burrowing nymphs. South America harbors the highest diversity of this family, including the subfamilies Asthenopodinae and Campsurinae. In this work, three new species belonging to the genera Tortopsis and Tortopus (Campsurinae) are described based on adults and nymphs from Colombia: Tortopsis toro sp. nov., Tortopsis andaki sp. nov. and Tortopus coreguaje sp. nov. Additionally, Tortopsis limoncocha is firstly recorded from Colombia. A cladistic analysis of all the species in these genera is presented, using external morphological characters of adults and eggs. Keys to male and female adults of all the species of both genera are presented.
Campsurinae, cladistics, new species, South America, systematic, taxonomic key
The monophyletic group formed by Tortopus Needham and Murphy, 1924 and Tortopsis Molineri, 2010 (Campsurinae) gathers very specialized mayflies with burrowing nymphs and extremely short-lived adults (
The role of some Campsurinae species in bioturbation of lentic habitats, with large nutrient and pollutant movements from the sediment by the nymphs had received some attention (
Tortopus was established by Needham and Murphy in 1924 and seven species are currently recognized (
Tortopsis was established by
The Caquetá River is located in the Amazon region, in the southeastern portion of Colombia and northwestern portion of Brazil. It extends through 2,280 km, with 1,200 km within Colombia and the remaining within Brazil. The river arises in the Colombian Massif known as “Páramo de las Papas”, in the central Andean region of the Cauca department, and flows into the Amazon River in Brazilian territory, where it changes the name to Japurá. It has an extensive water network, and along with the Putumayo River, are the two main tributaries of the Colombian Amazon. The Caquetá River drains a large basin of approximately 267,730 km2, shared between the departments of Cauca, Caquetá, Putumayo, Guaviare, Vaupés and Amazonas within Colombia (
The type locality of the collected specimens here described corresponds to the lowlands of the middle basin of the Caquetá River, in the town of Curillo, located within the limits of the dense Amazon jungle in the department of Caquetá. All the material described here was collected at a single station in the town of Curillo, based on authorization 1166 granted by ANLA (National Authority of Environmental Licenses in Colombia) in the project “Trans–Amazon Aquatic Insects: study of potential areas of endemism in Colombia”.
Nymphs were manually collected on riverbanks, with the help of a blade to separate substrate pieces with the nymphal burrows. Substrate consisted of hard clay, and the pieces picked up were disaggregated on the shore to collect the nymphs. Mature nymphs were settled in rearing cages in the river for a short period and then transported in an ice keeper with river water and a piece of substrate (at ambient temperature). Adults were collected with a light trap at river margin set around sunset. All the collected material was fixed and preserved in ethyl alcohol 96°. Slide mountings were done using Canada Balsam. Photographs were taken with a ZEISS AxioCam ICc 5 mounted on a Stemi 508 stereo microscope and a Leica M205C stereomicroscope with an attached Leica MC–170HD camera. Line drawings were made using a camera lucida mounted on a microscope Olympus BX51. Some photographs are the result of stalking partially focused images with the software CombineZP (
The matrix and characters proposed by
Holotype male imago (
Tortopsis toro sp. nov., known from imagos of both sexes and nymphs, can be distinguished from all other species of the genus by: In adults, 1) forewing length ranges between 12.2–13.2 mm (male) and 15.5–17.0 mm (female); 2) hyaline wings, with brownish veins and costal margin shaded slightly gray; 3) parastyli almost straight in lateral view, slightly curved medially, with a longitudinal ventral furrow (Figs
Male imago. Length (mm): body, 14.0–14.5; forewing, 12.2–13.2; hindwing, 5.9–6.1; foreleg, 5.3–5.9; cerci, 32.0–36.0. General coloration yellowish white shaded with gray dorsally (Fig.
Female imago. Length (mm): body, 18.0–18.5; forewing, 15.5–17.0; hindwing, 6.5–7.5; cerci, 4.3–4.8. General coloration as in male except darker markings. Head. Coloration black between ocelli, with posterior and submedian grayish lines on occiput (similar to Fig.
Egg. Suboval. Length, 410–420 μm; width, 320–340 μm.
Nymph (mature). Length of body (from apex of tusks to apex of abdominal tergum X): female, 22.0–26.0 mm; male, 14.0–19.0 mm. General coloration yellowish white with gray markings dorsally (Figs
Tortopsis toro sp. nov., nymph. 7, labrum, dorsal view; 8, left mandible, inner view, 9, right mandible, inner view; 10, right mandible, detail of tusk, ventral view (arrow indicates stout spine near inner margin); 11, hypopharynx, dorsal view; 12, labium, ventral view; 13, left maxilla, ventral view (arrow indicates membranous gill); 14, right maxilla, ventral view.
This species is dedicated to Dr. Beatriz Toro Restrepo, Universidad de Caldas, in recognition of her work in environmental education, friendship and contributions in fieldwork with aquatic insects of Colombia.
Known only from the type locality.
Tortopsis toro sp. nov., imago: 19, male genitalia, ventral view; 20, detail of apex of penis, ventrolateral view; 21, male genitalia, lateral view; 22, female abdominal sternum VIII. Nymph: 23, head outline and setation, dorsal view (tusks omitted); 24, left mandible, inner view (arrow indicates stout spine near inner margin); 25, detail of right mandibular tusk, inner view (arrow indicates subapical outer indentation); 26, paraprocts, ventral view (right half setation omitted).
Holotype male imago (
Tortopsis andaki sp. nov., known from imagos of both sexes and nymph, can be distinguished from all other species of the genus by: In adults, 1) fore wing length 9.2–9.6 mm (male), 9.5–9.7 mm (female); 2) wings hyaline slightly tinted with purplish gray, veins purplish gray; 3) parastylus curved dorsally, more markedly on apical third, with a longitudinal ventral furrow (Figs
Male imago. Length (mm): body, 10.8–11.0; fore wing, 9.2–9.6; hind wing, 4.0; foreleg, 4.9–5.0; cerci, 25.0–26.0. General coloration whitish shaded dorsally with light purplish gray (Figs
Female imago. Length (mm): body, 10.0–10.1; fore wing, 9.5–9.7; hind wing, 3.9–4.0; cerci, 3.0. General coloration as in male but shading more strongly marked. Head black between ocelli, with small light gray marks on occiput. Wings membrane hyaline slightly tinged with whitish yellow, veins brownish. Abdomen shaded with gray on terga. Parastyli receptors on sternum VIII with sockets (Figs
Nymph (mature female). Length: body, 17.0 mm. General coloration yellowish white shaded with gray dorsally (Fig.
Tortopsis andaki sp. nov., imago. Male: 33, male genitalia, ventral view; 34, genitalia, lateral external view; 35, same, inner view; 36, penis detail, dorsal view; 37, penis, detail, ventral view; 38, details of penis apex, dorsal view above, ventral view below. Female: 39, abdominal sternum VIII, lateral view; 40, same, ventral view.
Egg. Subcircular, white. Length, 330–350 μm; width, 280–300 μm.
This species is dedicated to the Andaki indigenous people, an American ethnic group that inhabited the upper Caquetá River basin.
Known only from the type locality.
Tortopsis andaki sp. nov., female nymph. 41, labrum, dorsal view; 42, left mandible, ventral view; 43, right mandible, ventral view; 44, same, detail (arrows indicate first and last spine of the inner marginal row); 45, hypopharynx, ventral view; 46, labium, ventral view; 47, right maxilla, ventral view (arrow indicates membranous gill); 48, left maxilla, ventral view; 49, fore leg, dorsal view; 50, detail of fore tarsal claw, dorsal view; 51, middle leg, dorsal view; 53, hind leg, dorsal view.
1 male imago (
Male adults (Not keyed: T. bruchianus (Navás) is only known from female; T. parishi (Banks) is poorly known from the holotype male)
1 | Penis flattened and somewhat widened distally ( |
2 |
– | Penis cylindrical, long and slender ( |
4 |
2(1) | Penis slightly widening from base to apex | T. puella (Pictet) |
— | Penis abruptly widened in distal half | 3 |
3(2) | Parastyli entire, without furrow; apical spine of penis pointed ( |
T. obscuripennis (Domínguez) |
– | Longitudinal furrow on parastyli (Figs |
Tortopsis toro sp. nov. |
4(1) | Parastyli with a longitudinal ventral furrow (Figs 63–65, 77 in |
5 |
– | Parastyli entire; apical spine hooked (e.g., Figs |
6 |
5(4) | Apex of penis simple, with a semicircular sclerotized structure ( |
T. spatula Molineri |
– | Apex of penis more complex, with sclerotized two–pointed portion, and a membranous rounded lobe (Figs |
T. andaki sp. nov. |
6(4) | Parastyli relatively straight in lateral view, may be curved on apical 1/4 | 7 |
– | Parastyli curved from its base ( |
8 |
7(6) | Parastyli curved on apical 1/4 (Figs |
T. limoncocha Molineri |
– | Parastyli straight ( |
T. canum Gonçalves et al. |
8(6) | Parastyli strongly curved, the main axis of the apical third forms an angle of 90° with the corresponding axis of the basal third ( |
T. unguiculatus (Ulmer) |
– | Parastyli curved smoothly from its base | 9 |
9(8) | Known distribution restricted to southern Bolivia and Northwestern Argentina | T. sarae (Domínguez) |
– | Known distribution restricted to North America | T. primus (McDunnough) |
Female adults (female of T. unguiculatus is inadequately known and could not be included in the key)
1 | Wings dark, all veins shaded strongly with gray or black; fore wing length 19.5–20.5 mm; color pattern on head and pronotum as in figure 89 in |
T. obscuripennis (Domínguez) |
– | Wings lighter, yellowish to brownish, some veins may be dark, mainly in costal region, fore wing length variable; color pattern on head and pronotum not as above | 2 |
2(1) | Parastyli receptors on abdominal sternum VIII elongated and with slightly sinuous anterior margin ( |
3 |
– | Parastyli receptors on abdominal sternum VIII generally not elongated (except T. spatula), margin not sinuous ( |
4 |
3(2) | Head shaded gray to black between lateral ocelli and with few small markings in occiput ( |
T. puella (Pictet) |
– | Head shaded gray in a V–shaped line between lateral ocelli and another medial thin line running anteriorly toward median ocellus, occiput without traceable marks (but material studied was very faded); abdomen with thin blackish medial line on terga I–VIII; on sternum VIII, anterior and posterior margins of sockets are divergent toward medial line ( |
T. primus (McDunnough) |
4(2) | Parastyli receptors V–shaped (Fig. |
5 |
– | Parastyli receptors C–shaped ( |
6 |
5(4) | Occiput shaded with gray ( |
T. spatula Molineri |
– | Occiput whitish, without shading; parastyli receptors as in Figs |
T. andaki sp. nov. |
6(4) | Head with gray or black markings on occiput (Fig. |
7 |
– | Occiput without marks or with very light gray and small markings near hind margin ( |
8 |
7(6) | Color pattern on head and pronotum as in Fig. 88 (in Molineri, 2010); parastyli receptors small, not reaching more than half of VIII sternum ( |
T. limoncocha Molineri |
– | Color patter on head and pronotum as in Fig. |
T. canum Gonçalves et al. |
8(6) | Eggs orangeish, known distribution: Central Argentina | T. bruchianus (Navás) |
– | Eggs yellowish | 9 |
9(8) | Sockets on sternum VIII relatively small (about 1/3 or less of total sternal length); known distribution: NW Argentina and S Bolivia | T. sarae (Domínguez) |
– | Sockets on sternum VIII relatively large (about 1/2 of total sternal length); known distribution Colombia– Caquetá | T. toro sp. nov. |
Holotype male imago (
Tortopus coreguaje sp. nov., known from male imago and nymph, can be distinguished from all other species of the genus by the following combination of characters: In male imagos, 1) fore wing length 10.9–11.0 mm (male); 2) pale wings, veins translucent gray; 3) pedestal short with relatively large, straight and very long parastyli (Figs
Tortopus coreguaje sp. nov., male imago. 56, genitalia, ventral view (left forceps omitted); 57, genitalia, lateral view. Nymph: 58, head outline and setation, dorsal view (tusks omitted); 59, right mandible, inner view; 60, left mandible, inner view; 61, same, detail of apex (arrow indicates blade-like projection); 62, fore leg, ventral view (rows of filtering setae delineated); 63, fore tarsal claw; 64, hind tarsal claw.
Male nymph (immature). Length: body, 9.5 mm; cerci, 2.5 mm; caudal filament, 2.0 mm. General coloration whitish shaded with gray (Fig.
This species is dedicated to the Coreguaje community inhabiting along the piemont and lower parts of the Caquetá River and its affluents.
Known only from the type locality.
Male adults (not keyed: T. circumfluus Ulmer is only known from female)
1 | Pedestal longer than wide, with small basal extension; parastylus reduced, expressed only as an acute point ( |
T. ipixuna Molineri et al. |
– | Pedestal wider than long (not including the parastylus), with a well-developed basal extension, parastylus at least 1/2 the length of pedestal ( |
2 |
2(1) | Parastylus length shorter than length of its base ( |
3 |
– | Parastylus length subequal to base to more than 2 times length of base ( |
4 |
3(2) | Abdominal terga and sterna shaded brownish gray; parastylus relatively short ( |
T. bellus Lugo-Ortiz & McCafferty |
– | Abdominal terga shaded gray, sterna much less marked; parastylus slightly longer ( |
T. harrisi Traver |
4(2) | Parastylus relatively thin from base ( |
5 |
– | Parastylus thick at the base ( |
6 |
5(4) | Parastylus 1.5–2.5 the length of pedestal base ( |
T. igaranus Needham & Murphy |
– | Parastylus 2.6 the length of pedestal base (A/B in Fig. |
T. coreguaje sp. nov. |
6(4) | Abdomen shaded with gray on dorsum, ventrally paler; penis relatively thin basally ( |
T. zottai (Navas) |
– | Abdomen shaded strongly with brownish gray, dorsally and ventrally; penis with a wider base ( |
T. arenales Molineri |
Searches including all the taxa (under both, equal and implied weights) resulted in many possible resolutions with their strict consensus showing a large politomy that included all the species in Tortopus and Tortopsis. To study the relationship among species in Tortopus and Tortopsis, we deactivated three of them (Tortopus circumfluus, Tortopsis bruchianus and Tortopsis parishi). These species present many missing entries in the matrix, as they are poorly known from their original descriptions and were not collected again. When deactivated, a unique and more resolved tree is found under implied weights (Fig.
Equal and implied weighting strategies found the same synapomorphies for the group Tortopsis + Tortopus, and for each genus, as detailed below. Synapomorphies defining the clade Tortopus + Tortopsis: Female wing veins thickened (character 2:1); female hind wing with anastomosed anal sector (character 3:1); female parastyli receptors on abdominal sternum VIII, formed by paired sockets (character 5:2); female parastyli receptors, U–shaped (character 6:1, later change to other forms); legs of imagos of both sexes (except male forelegs), reduced and distorted (character 9:1); male genitalia with two segments on forceps (character 11:1); with outer projected pedestals, forming a dorsal and pointed parastylus (character 13:3); knob at forceps base present (character 27:1); and egg without polar caps (character 30:0).
The synapomorphies of Tortopus are: Female parastyli receptors with long furrows anterior to sockets (character 8:1); male gonopore associated with a sclerotized margin (character 17:1); penis flattened (character 21:1); and male abdominal sternum IX almost separated in two portions by a median notch (character 28:2). Finally, the synapomorphies of Tortopsis are: Female fore wing veins on R sector without longitudinal or intercalary veins between IR and R1 (some fused crossveins may be present, forming 1 or 2 short attached marginal intercalaries) (character 4:2); female parastyli receptors with sockets opening towards median line (character 7:1); male gonopore associated with a claw–like or spatulated spine (character 16:1); and male genitalia with long parastyli (character 24:3).
Some species groupings were found inside each genus. For example in Tortopus, three species (T. ipixuna, T. harrisi and T. bellus) share male genitalia with very short parastyli (character 24:1). In Tortopsis, T. toro sp. nov. is sister to the pair T. andaki sp. nov. + T. spatula, because the three species present a longitudinal furrow on parastyli (character 25:1). The remaining species in Tortopsis (T. canum, T. primus, T. limoncocha, T. unguiculatus, T. sarae, T. puella and T. obscuripennis) form a monophyletic group defined by female parastyli receptors C–shaped (character 6:0). Finally, two species (T. puella + T. obscuripennis) are joined by their apically wide penis (character 23:1), a feature independently acquired by T. toro sp. nov.
The form of the penis in T. toro sp. nov. is similar to T. obscuripennis and T. puella, because of the abrupt distal widening. On the contrary, the semicircular spine at the apex of penis is similar to T. spatula, as well as the presence of a furrow along the parastyli (this last feature also shared with T. andaki sp. nov. The nymph of T. toro sp.nov. is similar to T. obscuripennis in the shape and setation of tusks, but the subapical outer indentation on the tusks of T. toro sp. nov. is more marked (Figs
Tortopsis andaki sp. nov. is similar to T. spatula, but can be readily distinguished because T. andaki sp. nov. presents a more complex penis apex, with a rounded membranous lobe and two sclerotized projections (Figs
Tortopsis limoncocha was previously known from a single locality in the Ecuadorian portion of the Amazonas basin (
The nymph here described for Tortopus coreguaje sp. nov. is the second nymph known for the genus; the nymph of T. harrisi (
Our results improved the resolution reached in the last published phylogenetic hypothesis (
The biodiversity study in the Caquetá region (Figs
The authors have no conflicts of interest to declare that are relevant to the content of this article. All authors contributed equally to the manuscript.
We thank the Universidad de Caldas for providing funds to LGD to conduct the project “Trans–amazonian aquatic insects: study of potential areas of endemism in Colombia”. We also thank Beatriz Toro Restrepo (Universidad de Caldas) and Juliette Pauline Chaux (Amazon Conservation Team) for assistance with sampling specimens in Caquetá River. To Fabian Barroso–Mena for the hydrological information of the studied zone and to Felipe Ortega (Laboratorio de Imágenes, Programa de Biología, Universidad del Valle) for the photographic record of images No 4, 5, 6. Comments from M. Sartori and I. Gonçalves helped us to improve the manuscript. We thank Jade Black for language revision. CM thanks Argentine Council of Scientific Research.
File 1
Data type: .pdf
Explanation note: Matrix, list of characters and states (modified from