Research Article |
Corresponding author: Leon Gustavo de Miranda Tavares ( leogtav@gmail.com ) Academic editor: André Nel
© 2023 Leon Gustavo de Miranda Tavares, Renato Jose Pires Machado, Adolfo Ricardo Calor.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
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Tavares LGM, Machado RJP, Calor AR (2023) The Neotropical antlion genus Ameromyia Banks, 1913 (Neuroptera: Myrmeleontidae), systematics and redefinition under a phylogenetic approach. Arthropod Systematics & Phylogeny 81: 499-553. https://doi.org/10.3897/asp.81.e89641
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A taxonomic revision and the first phylogenetic hypothesis of the Neotropical genus Ameromyia Banks (Myrmeleontidae: Brachynemurini) is herein presented. The phylogeny is based on 45 morphological characters and recovered the traditional Ameromyia as paraphyletic in respect to the monotypic genus Venezueleon Stange, which is here synonymized under Ameromyia. Three species are synonymized (A. hirsuta Navás and A. stevensi Navás under A. nigriventris (Walker), and A. pentheri Navás under A. strigosa (Banks)) and two new species are described (A. clepsydra sp. nov. and A. explicata sp. nov.). Ameromyia sensu novo is a valid genus with 12 species restricted to South America, and divided into two species groups. Taxonomic keys are also presented to adults and larvae, as well as a discussion on the genus biology.
Brachynemurini, morphology, Myrmeleontinae, phylogeny, taxonomy
Myrmeleontidae Latreille is currently the most diverse family in the order Neuroptera, comprising nearly 2,150 extant species (
The tribe Brachynemurini (Myrmeleontinae) currently comprises 117 valid species in 28 genera, which are restricted to the Americas (
However, the first cladistic analysis of Brachynemurini reinstated its tribal status, but reclassifying its species in three different tribes: Brachynemurini (s.str.), Gnopholeontini and Lemolemini, mainly based on female genitalia and larval characters (
Ameromyia Banks, 1913 is a Neotropical genus of Brachynemurini which today comprises 12 valid species (
Live specimens of Ameromyia. A A. strigosa, Argentina, by Quentin Vandemoortele. B A. muralli, Brazil, by Eli Vieira Araujo-Jnr. C A. modesta, Surinam, by Tom Murray. D A. pubiventris, Brazil, by Carlos José Correia de Santana. E A. clepsydra sp. nov., Brazil. F A. explicata sp. nov., Brazil. Latter photographs by Leon Tavares.
Ameromyia was first described circumscribing the Brachynemurini species that present a long CuP in the forewing and a long CuA in the hind wing, as well as the presence of both banksian lines on both wings (
Ameromyia male genitalia was described as bearing the same structure as its sister genus Venezueleon. However, both genitalia descriptions were conflicting in regard to the homology of both gonarcus and parameres (
A total of 607 Ameromyia specimens were herein analyzed. Type specimens analyzed are discussed under “Remarks” for each species. For species with syntypes, lectotypes were designated from the syntypes series according to the better-preserved specimen. Specimens were provenient from several collecting events throughout 2017–2019, and by loans or photographs from institutions. Collected data for the specimens studied is included in the section “Examined Material” for each species. New distributional records at country level are highlighted in bold in the section “Distribution” for each species. A map showing the distribution of each species, per species group and by collection sites was constructed in QGis 3.4.13 Essen using projection unit GCS, datum WGS 84 (QGIS Development Team, 2015). Specimens whose label locations stated only country or state/district/province were plotted on the capital of the stated location. Specimens collected were preserved in 80% ethyl alcohol, envelopes or pinned. They were identified using the key provided by
Immature specimens selected for rearing were placed in individual plastic containers. Plaster or styrofoam plates were fit into each container, and then coated with sand, soil and sometimes small pebbles from the habitat where the larvae were collected. Larvae were fed two to three times per week with either Drosophila melanogaster Meigen maggots, first to third instar Tenebrio molitor Linnaeus larvae, and/or Nasutitermes Dudley nymphs, depending on prey availability. After pupation, cocoons were transferred to a glass rearing cage with sand and vertical wooden sticks. Rearing boxes were kept in air-conditioned rooms with the temperature between 23–25°C. After adult emergence, the pupal exuvia and the larval sclerites were preserved in 80% ethyl alcohol, allowing for direct association between the different life stages of the same species. Cocoons were stored in vials with naphtalene and posteriorly associated with its respective preserved specimens. Adult specimens were placed in voil fabric enclosures of 60 × 60 cm during mornings, and of 2 × 2 m during afternoons and nights. Voil fabric enclosures were enriched with dry bushes and/or branches, and sand, dirt and rocks from the specimen’s natural habitats. Adult specimens were fed adult Drosophila melanogaster Meigen, and handfed maggots, honey plus water, or gelatin with brown sugar and egg yolk. Hand feeding consists of grabbing adult specimens by their wings, and touching the food to their mouthparts, until they taste, grab the food and start chewing. When senescent, adult specimens were placed into a freezer for posterior pinning, or stored in 80% ethyl alcohol.
Ingroup consisted of all valid Ameromyia species, including the new species. Outgroup was sampled upon specimen availability and taxonomic history according to previous systematic studies (
A total of 45 morphological characters were coded and included in the character matrix (Supplementary Material Table S1). Some characters were adapted from
Head (characters 1–4)
1. Frons: Setose (0) (Fig.
2. Interocular distance, in relation to eye width: shorter (0) (Fig.
3. Interantennal distance, in relation to scape width: shorter (0); longer (1) (Fig.
4. Vertex–clypeus length, in relation to thorax dorsoventral height: longer (0) (Fig.
Thorax (characters 5–12)
5. Mesothorax, mesepisternum, dorsal and ventral areas, coloration: not contrasting (0) (Fig.
6. Prothorax, proportions in dorsal view: broader than long (0) (Fig.
7. Legs, profemoral sense hair: absent (0) (Fig.
8. Legs, length of profemoral sense hair, in relation to profemur length: shorter than profemur (0); longer or as long as profemur (1) (Fig.
9. Legs, Pretarsal claws, length, in relation to distal tarsomere length: shorter than distal tarsomere (0) (Fig.
10. Legs, profemur, decumbent setae, coloration: only black (0) (Fig.
11. Legs, profemur, color pattern: without contrast between external and internal face (0) (Fig.
12. Legs, tarsus, color pattern: without contrast between external and internal face (0) (Fig.
Wings (characters 13–31)
13. Forewing, origin of RP, in relation to CuA fork: before CuA fork (0) (Fig.
14. Forewing, number of crossveins between CuP and posterior branch of CuA: less than 6 crossveins (0) (Fig.
15. Forewing membrane, mediocubital area, coloration: clear (0); infuscated (1) (Fig.
16. Forewing membrane, mediocubital area pattern of infuscation: dotted, infuscated at the base of each crossvein (0) (Fig.
17. Forewing, presectorial and radial sector area, vertical crossveins, degree of infuscations: not suffused with infuscations (0) (Fig.
18. Forewing, Sc vein, color pattern: homogeneous (0) (Fig.
19. Forewing, RA vein, color pattern: homogeneous (0) (Figs
20. Forewing, CuP origin, in relation to basal crossvein: basad to basal crossvein (0); distad to basal crossvein (1) (Fig.
21. Forewing, CuP vein, color pattern: homogeneous (0) (Fig.
22. Forewing, course of vein 2A, in relation to vein 3A: forming a sharp angle towards 3A (0); forming a smooth curve towards 3A (1) (Fig.
23. Forewing, vein 2A in relation to 3A: independent from 3A (0) (Fig.
24. Forewing, independent vein 2A, in relation to 3A: touches vein 3A (0) (Fig.
25. Forewing, Rhegmal area, coloration: hyaline (0) (Figs
26. Forewing, Banksian lines: absent (0); present (1) (Figs
27. Hind wing, apex margin, shape: acute (0) (Fig.
28. Hind wing, crossveins between vein CuA and posterior branch of MP: absent (0); present (1) (Fig.
29. Hind wing, number of crossveins between CuA and posterior branch of MP: 1–2 crossveins (0); 3–4 crossveins (1) (Fig.
30. Hind wing, posterior area cells, length/width ratio: longer than high (0) (Fig.
31. Hind wing, Banksian lines: absent (0); present (1) (Fig.
Abdomen (characters 32–37)
32. Male abdomen, length, in relation to hind wing length: shorter than hind wing (0); barely longer than hind wing (1) (Fig.
33. Male terminalia, ectoproct: without postventral lobe (0); with postventral lobe (1) (Fig.
34. Male terminalia, ectoproct, postventral lobe length, in relation to ectoproct height: shorter (0) (Fig.
Ameromyia male terminalia and genitalia chaetotaxy: A A. protensa lectotype male terminalia, lateral view, photograph by Lara Lopardo (
Female genitalia (characters 36–38)
35. Female terminalia, pregenital plate: insconspicuous (0) (Fig.
36. Female terminalia, gonapophyseal plate, length in relation to lateral gonapophysis: shorter (0) (Figs
Ameromyia female terminalia: A A. protensa, lateral view. B A. protensa, posterior view. C A. nigriventris, ventral view. Photographs by Lara Lopardo (
37. Female terminalia, lateral gonapophyses, distal margins: fused (0) (Fig.
Male genitalia (characters 38–45)
38. Male genitalia, genitalic sac, setae: absent (0); present (1) (Figs
Scanning electron microscope (SEM) micrograph of Ameromyia nigriventris male genitalia: A lateral view. B Posterior view. — Abbreviations: cs, clavate setae; et, ectoproct; go(h), gonarcus, hidden under pleura; hs, hinge setae; pah, paramere hinge; pap, paramere plate; pat, paramere tooth.
Male genitalia of Ameromyia explicata sp. nov., cleared in KOH: A dorsal view, in folded conformation. B Dorsal view, in unfolded conformation. C Lateral view, in folded conformation. D Lateral view, in unfolded conformation. — Abbreviations: cs, clavate setae; go, gonarcus; pah, paramere hinge; pap, paramere plate; pat, paramere tooth.
39. Male genitalia, genitalic sac, setae, shape of distal end: filiform (0); clavate (1) (Figs
40. Male genitalia, genitalic sac, clavate setae, “club” shape: globose (0) (Figs
41. Male genitalia, parameres hinge, presence: absent (0); present (1) (Figs
42. Male genitalia, parameres, lateral tooth ventrally: absent (0); present (1) (Figs
43. Male genitalia, parameres lateral ventral tooth, placement, in relation to paramere plates: distal (0); basal (1) (Figs
44. Male genitalia, gonarcus, arching, in relation to parameres: arching dorsad to parameres (0); arching anterior to parameres, barely dorsad (1) (Figs
45. Male genitalia, mediuncus, presence: absent (0) (Figs
Analyses under equal weighting resulted in four most parsimonious trees, with 82 steps. Two sets of trees were almost identical, differing only in regard to the outgroup (relationship between Dimarella riparia and Myrmeleon sp.), and therefore, only two trees with truly different topologies for Ameromyia internal relationships were produced (Fig.
Phylogenetical hypothesis for Ameromyia: strict consensus cladogram resulting from analyses under equal weighting. Capital letters next to nodes and colored background around nodes and stems indicates the clades and species groups referred in text, respectively. Squares ordered on stems indicates characters used in the analysis: number above square and number under square indicates number of character and state of character as stated on “Character List” section. Filled squares indicates synapomorphies and blank squares indicates homoplasies. Values under branches indicates their support values for Bremer absolute and relative values, and resampling values (GC) respectivelly. A Proposed classificaton for Ameromyia sensu novo. B Alternative tree.
The clade which represents Ameromyia was recovered as paraphyletic in all analyses, with Venezueleon guaricus (~ A. guarica comb. nov.) placed deep into internal branches. This clade is supported by eight synapomorphies (29:1 At least four crossveins between hind wing vein CuA and posterior branch of MP; 34:0 Ectoproct postventral lobe shorter than ectoproct height; 36:1 Female gonapophyseal plate elongate; 37:1 Lateral gonapophysis of female genitalia fused; 39:1 clavate setae on male genitalia; 43:1 Parameres lateral tooth basad to paramere plates; 44:1 Male gonarcus positioned anterior to parameres; and 45:0 Male mediuncus absent).
Two well defined groups were recovered inside the Ameromyia sensu novo clade. The ‘modesta’ group (Fig.
The ‘nigriventris’ group (Fig.
Regarding the remaining species within the nigriventris group clade (clades C and H), our analyses recovered two different topologies. In our proposed classification for Ameromyia (Fig.
In the alternative tree (Fig.
Ameromyia
=Amazoleon
=Foya
=Nemotolus
=Venezueleon
Ameromyia clepsydra sp. nov.; A. dimidiata Navás, 1915a; A. explicata sp. nov.; A. guarica (Stange, 1994) comb. nov.; A. modesta (Banks, 1943); A. muralli Navás, 1932; A. nigriventris (Walker, 1860); A. pleuralis Navás, 1926b; A. protensa (Gerstaecker, 1893); A. pubiventris (Walker, 1860); A. strigosa (Banks, 1909); A. tendinosa (Gerstaecker, 1893).
Fore and hind wings with both banksian lines well developed; hind wings vein CuA with at least four crossveins connecting with posterior branch of MP; male ectoproct postventral lobe shorter than ectoproct height; female gonapophyseal plate elongate; lateral gonapophysis of female genitalia fused; male genitalic sac with clavate setae; parameres lateral tooth basad to paramere plates; male gonarcus positioned anterior to parameres; and male medincus absent; larvae with thread-like setae.
Ameromyia clepsydra sp. nov.; A. guarica comb. nov.; A. modesta; A. nigriventris.
Adult. Head: frons with short setae. Antenna of same length on both sexes, moderately short with about 30 flagellomeres. Antennal fossae separated by about two times pedicel width. Interocular distance less than eye width. Profemur without clavate setae. Vertex with many short decumbent setae. Ocular rim without setae. Clypeus with scattered setae, which are oriented ventrally. Labrum with two rows of short setae on ventral margin, all of which pointing ventrally. — Thorax: Pronotum broader than long when pretarsal claws are shorter than distal tarsomere, and longer than broad when pretarsal claws are otherwise. Pronotum with many long anteriorly oriented bristles on all margins, which are longer on lateral margins. Pronotum often with mirrored “P” shaped markings medially. Prescutum 2 and 3, mesoscutum and metascutum with many short, posteriorly-oriented white setae on posterior margins, and meso and metascutella with longer white setae on posterior margins. Thorax with many decumbent setae laterally, and mesepisternum frequently with a sulcus mesally. Lateral side of thorax sometimes with a longitudinal pale band, which covers episternum and epimeron of meso and metathorax, and contrasts with ventral thorax which is darker. Mesonotum without blade-like setae. — Legs: Legs with many short decumbent setae. Femoral sense hair present or absent. If present, profemoral sense hair as long as femur and much longer than that of mesoleg, which is shorter than mesofemur. Femur with or without black and/or white bristles. Tibiae with many black bristles, which are longer than femoral bristles, when present. Tibial bristles as long as tibial width, when femoral bristles are absent. Tibial spurs present, longer than basitarsus. Five tarsomeres, longer than broad. Distal tarsomere much longer than basitarsus. Pretarsal claws at least 2/3 of distal tarsomere length, more commonly longer than distal tarsomere. — Wings: Pilula axillaris well developed. Forewing as long as hind wing or slightly longer. Both anterior and posterior banksian line well developed and present in both wings. Wing membrane mostly hyaline, sometimes with a reddish or brownish tint. Forewing costal area simple at least until midwing length. Posterior area of forewing about as broad as that of hind wing. Forewing rhegmal area with brown infuscation on rhegmal crossveins. Forewing vein CuP runs along posterior fork of CuA, with at least four crossveins connecting them. Forewing vein 2A running towards posterior wing margin in a smooth curve. Forewing vein 2A connected to 3A by a crossvein, or touching 3A before running towards posterior wing margin. Hind wing CuA runs along posterior branch of MP fork with at least four crossveins connecting them. — Abdomen: Male abdomen longer or much longer than wings, female abdomen shorter than wings, or almost 1.5× longer than wings. Male tergite IX with many long, thick setae ventrally. Male tergite IX subquadrangular, with posteroventral margin slightly swollen and with many posterior oriented setae. Male ectoproct with a developed short postventral lobe, which is subequal to half of ectoproct height, without median or secondary lobes. Parameres posteriorly plate-like, arched, striated on external face of paramere plates, with a basal lateral hollow tooth on each side, and high sclerotized posterior folds. Parameres with a hinge mesally on internal face of each paramere plate, and with a row of short setae on each internal margin. Parameres anterior to hinge smooth, “spoon-like”, positioned between parameres plates (in “folding” position), and fused dorsally with an anterior bifurcated projection. Gonarcus smooth, membranous, with long anteriorly projected arms which are flat and long. Gonarcus positioned anterior to parameres, arching barely dorsad to parameres anterior bifurcation and dovetailing or almost dovetailing behind parameres plates lateral tooth. Genitalic sacs lateral to paramere tooth, with many clavate setae that reach beyond and/or above and posteriorly to paramere plates. Female terminalia with pregenital plate membranous with many setae. Gonapophyseal plate elongate, but shorter than posterior gonapophysis. Posterior gonapophysis digitiform, with many setae. Lateral gonapophysis fused, with well-developed digging setae. Ectoproct with well developed digging setae. — Larva (third instar). Head: Larvae with three teeth. Mandible with distance between mandible base and basal tooth longer than between teeth. Mandible not enlarged at base. Some setae on external margin of mandibles base with length as long or almost as long as mandible base width. Internal margin of mandibles roughly straight until reaching distal teeth, and roughly about the same width. Distal tooth smaller than basal and medial teeth and oblique in relation to other teeth. Distal tooth closer to medial tooth than medial tooth is to basal tooth. Head capsule on dorsal view as broad as long or slightly broader than long. Head capsule on ventral view longer than broad or as long as broad. Head dorsal surface with thread-like setae near posterolateral margins. Head capsule dorsally with many short, thick, blunt dolichasters and very short, almost “sphere-like” dolichasters, ventrally with short, thin dolichasters, and long and thick setae on lateral margins, which are straight on the anterior portion of head, and curved on posterior portion of head. Labrum with a row of thick, cylindrical dolichasters on posterior margin. Palpi with three segments, distal palpomere bigger than basal palpomere, which is bigger than mid palpomere. Distal palpomere enlarged at base with an acute end. Presence of rowed thick, cylindrical, white dolichasters near base of each palpi. — Thorax: Dorsal surface sometimes covered with thread-like setae. Dorsal surface prunescent, covered with short dolichasters. Thoraxic dorsal sclerites with short rowed dolichasters on all margins. Thorax ventral surface with long hair-like setae. Prothorax with many short, curved bristles on anterior margin. Mesothoracic spiracle borne on tubercle, tubercle length half of metathorax width at most. Tubercule covered with thread-like setae. Metathorax with two pedunculated setiferous processess on lateral margins, which bear many long bristles. Metathorax with a pair of eliptical dark spots submedially. — Abdomen: Abdominal tergites with thread-like setae, and sternites sometimes with thread-like setae. Abdominal spiracles not enlarged nor borne on tubercles. Setiferous processes on abdominal lateral margins bearing long white, black, or black and white bristles. Setiferous processes on abdominal lateroventral margins with hair-like setae. First abdominal tergite with two eliptical black spots submedially, almost aligned with metathoraxic dark spots. Odontoid process on sternite VIII longer than basal width. Ninth sternite covered with thread-like setae. Rastra with 4 thick setae, with innermost setae much shorter than remaining setae.
At first look, Ameromyia species can look similar to Argentoleon or to large Austroleon specimens, but can be readily differentiated from both genera by the short male postventral ectoproct lobes (which are much longer in Argentoleon and absent in Austroleon); the presence of both banksian lines in both wings; and the much longer hind wing CuA. Other Brachynemurini species bear a long hind wing CuA vein (such as Brachynemurus fuscus (Banks) and Brachynemurus nebulosus (Olivier)) or both banksian lines in both wings (B. nebulosus), but they are very different regarding distributional ranges and other Ameromyia diagnostic characters, such as the simple forewing costal area cells. Ameromyia male genitalia is also very conspicuous (Figs
Two species groups are determined based on taxonomical characters of wing veins, male genitalia, and cladistic results: the ‘modesta group’ and ‘nigriventris group’ (Fig.
Within groups, however, Ameromyia species are very morphologically similar, differing mostly in coloration. Coloration pattern on main wing veins such as Sc, R, Cu and A veins appears to be consistent across species, but markings pattern on wing membrane can be polymorphic within species. Thorax pronotal markings are also usually very similar within Ameromyia, but some species bear unique markings which are consistent within specimens, such as A. modesta (Figs
The larvae of most Ameromyia species are currently unknown, except for A. clepsydra sp. nov., A. guarica, A. modesta and A. nigriventris.
SEM micrograph of Ameromyia clepsydra sp. nov. third instar larvae: A head, dorsal view. B Thorax, dorsal view. C Mesothoracic tubercle, dorsal view. D Abdomen terminalia, ventral view. — Abbreviations: an, antennae; bt, basal tooth; dgs, digging setae; ds, dolichasters; dt, distal tooth; mdt, medial tooth; met, metathorax; mt, mesothorax; oc, occelli; op, odontoid process; pt, prothorax; ra, rastra; sr, spiracle; st, sternite; ts, thread-like setae; tu, tubercle.
Very few data have been made available regarding Ameromyia biology and/or behaviour.
In this work, no feeding behaviour was observed in the wild, but in captivity, adult specimens of A. explicata sp. nov., A. nigriventris and A. strigosa accepted D. melanogaster as prey. Aphids, maggots, small catterpillars and green lacewings were offered, but none were succesfully preyed upon. Maggots were only eaten when handfed, and small catterpillars were only eaten when handfed, and if some hemolymph or gut content was exposed. All adult specimens accepted artificial diets such as honey + water, boiled egg white, and gelatin with sugar and egg yolk. However, all artificial foods were only eaten when handfed and individuals would starve to death otherwise, even if the food was left inside the enclosures. This suggests these Ameromyia species might rely solely on aerial hunting for feeding. Diets consisting of only sugar and water would only sustain individuals for a week, whether in every other diet (both natural and artificial) the specimens lived for approximately three weeks. Starving adults would die in two or three days.
The larvae of A. modesta and A. nigriventris were described as living in sand dunes, and A. guarica was described as living in shallow sand anchored to bedrock (
Collection site and habitat of Ameromyia clepsydra sp. nov., in Andaraí, Bahia, Brazil: A site surroudings and sandy substrate on bedrock. B Second-instar larvae exposed in its “trench”. D Third-instar larvae exposed on its “trench”. D Empty, open coccoon on bedrock crevice. — Abbreviations: cc, coccoon; sp, sand pools.
Collection site and habitat of Ameromyia nigriventris, in Miguel Calmon, Bahia, Brazil: A site surroundings and sandy substrate on compacted soil. B First-instar larvae exposed in its “trench”. C Third-instar larvae exposed on its “trench”. D Coccoon, exposed on a dug-out “trench”. — Abbreviations: sp, sand pools; tr, trenches.
1 | Posterior area of hind wing narrow, cells delimited by crossveins between CuA and hind wing margin are longer than high (Fig. |
(modesta species group) 2 |
1’ | Posterior area of hind wing broad, cells delimited by crossveins between CuA and hind wing margin are at least as high as long (Fig. |
(nigriventris species group) 5 |
2 | Posterior area of forewing at base higher than prefork area at CuA fork; fore and hind wing RP veins dark coloured (southern Brazil; Argentina; Uruguay) |
Ameromyia protensa (Gerstaecker) (Figs |
2’ | Posterior area of forewing at base narrower than prefork area at CuA fork (Fig. |
3 |
3 | Thorax with dorsal part of pleuron pale brown in contrast to dark brown ventral part (Fig. |
Ameromyia pubiventris (Walker) (Fig. |
3’ | Thorax with pleuron not contrastingly pale above and dark below; legs with anterior face pale brown and posterior face dark brown (Fig. |
4 |
4 | Profemoral sense hair present; pretarsal claws longer than distal tarsomere; pronotum at least as broad as long (Fig. |
Ameromyia modesta (Banks) (Figs |
4’ | Profemoral sense hair absent; pretarsal claws shorter than distal tarsomere (Fig. |
Ameromyia guarica (Stange) (Figs |
5 | Forewing CuP runs parallel to CuA with less than 6 crossveins between them (Fig. |
Ameromyia clepsydra sp. nov. (Figs |
5’ | Forewing CuP runs parallel to CuA with more than 6 crossveins between then (Fig. |
6 |
6 | Forewing with four to five oblique markings along MA vein (Figs |
Ameromyia strigosa (Banks) (Figs |
6’ | Forewing without or with one oblique marking; frons and profemur decumbent setae white and black or all black (Fig. |
7 |
7 | Fore and hind wings falcate (Figs |
Ameromyia muralli Navás (Fig. |
7’ | Fore and hind wings not falcate (Fig. |
8 |
8 | Forewing CuP coloration homogeneous (Fig. |
9 |
8’ | Forewing CuP with dashed pattern (Fig. |
11 |
9 | Forewing CuP pale (Fig. |
Ameromyia dimidiata Navás (Figs |
9’ | Forewing vein CuP dark (Fig. |
10 |
10 | Forewing RA vein pale (Fig. |
Ameromyia pleuralis Navás (Fig. |
10’ | Forewing RA vein dark or dashed (Fig. |
Ameromyia tendinosa (Gerstaecker) (Figs |
11 | Lateral side of thorax with an evident pale brown longitudinal band, with dorsal portion of mesepisternum pale in contrast to darker ventral portion (Fig. |
Ameromyia nigriventris (Walker) (Figs |
11’ | Lateral side of thorax without an evident pale brown longitudinal band, with mesepisternum relatively homogeneous in coloration (Fig. |
Ameromyia explicata sp. nov. (Figs |
1 | Row of dolichasters on clypeolabrum anterior margin all black; presence of black setae on tuft of setae on abdominal setiferous processes; mesothoraxic tubercle longer than broad | 2 |
1’ | Row of dolichasters on clypeolabrum anterior margin black between mandibles and white at mandibles base; tuft of setae on abdominal setiferous processes with white setae only; mesothoraxic tubercle shorter than broad (Figs |
Ameromyia nigriventris (Walker) (Figs |
2 | Thorax and abdominal surface with dark thread-like setae (Figs |
3 |
2’ | Thorax and abdominal surface with only white or pale thread-like setae |
Ameromyia guarica (Stange) (Fig. |
3 | Clypeolabrum conical dolichasters grey/pale (Figs |
Ameromyia clepsydra sp. nov. (Figs |
3’ | Clypeolabrum conical dolichasters black (Fig. |
Ameromyia modesta (Banks) (Fig. |
Black and white coloration. Thorax lateral side black with no apparent longitudinal pale band. Wing veins alternating black and white coloration. Wing membrane with many markings and infuscations, mostly on radial, rhegmal and substigmal areas. Sectorial and presectoral area crossveins sometimes with an infuscated biareolate “8” shaped pattern. Male genitalic clavate setae club subglobose.
Adult. Head: Antennae clubbed (4 mm) with 31 flagellomeres, light brown with yellow rings between the flagellomeres. Vertex white with grey patches, and dark brown spots on lateral margins; with a transversal and a longitudinal dark brown streak at anterior and posterior row, forming a “T” shape. Vertex decumbent setae white. Interantennal area white with a dark streak in the middle, and dark spots under each scape. Frons light yellow near antennae and light grey near clypeus, and median, transversal dark brown patches. Frons setae black. Interocular distance less than eye width. Clypeus white, with two dark brown marks pointing ventrally. Labrum marginally yellow, dark brown centrally. Mandible dark brown. Palpi dark brown, with white joints. — Thorax: Thorax black, with white or light brown markings. Thorax on lateral view dark, with no lateral longitudinal band under wings. Pronotum as broad as long, white with a longitudinal dark brown band on each side. Pronotum marginal setae white. Mesonotum dark, with white markings on mesoscutum and mesoscutellum. Metanotum dark with light brown medially on metascutum and a longitudinal light brown band on metascutellum. — Wings: Wing tip acute, frequently notched on posterior margin or with a strongly falcate and sinuous apex posterior margin. Veins with dark brown and white dashed pattern. Wing membrane hyaline, with dark infuscations along crossveins, with more intensity on forewing than on hind wing, except for infuscations on presectoral and on radial areas, which are evident on both wings. Presectorial and radial areas in both wings sometimes with longitudinal crossveins between vertical crossveins, forming a pattern akin to an “8” shape. Forewing mediocubital area with dark infuscations around vertical crossveins which become sparser along MA vein, culminating on rhegmal area. Forewing CuP and hind wing CuA veins extending well before CuA fork and MP fork respectively, with at least four crossveins connecting them and posterior branch of fork. Pterostigma frosted white, encompassing four crossveins. Cells delimited by transversal veins in hind wing posterior area higher than long. — Legs: Coxae dark. Femur and tibiae with long black bristles. Femur white, suffused with black spots. Femur decumbent setae black. Profemural sense hair longer than profemur, and mesofemural sense hair much shorter than mesofemur. Tibia white with distal joint and near apical joint black. Tibial spurs slightly shorter than pretarsal claws. Five tarsomeres, white, except for distalmost and third to distalmost tarsomeres which are black. Pretarsal claws at least 1.5 times longer than basitarsus. — Abdomen: Abdomen mostly black, turning dark brown at terminalia. Tergites with scarce white setae, and sternites with black setae. Male ectoproct with many long black setae on posterior face, and a very short, slight round postventral lobe, which is less than 1/3 ectoproct height. Parameres hinge with a row of short setae on each side. Gonarcus arch positioned not above paramere plates on lateral view. Genitalic sac with many short, subclavate setae. — Larva (third instar): Head: Head capsule as long as broad in dorsal view, slightly longer than broad in ventral view. Head capsule dorsally grey, with dark brown patches medially, and on anterior margin. Clypeolabrum light brown medially, and dark near mandibles. Head capsule ventrally light brown. Short dolichasters on head dorsal surface cone-like. Head capsule dolichasters mostly black, with a few white dolichasters near anterior and lateral margins. Row of dolichasters on labrum anterior margin black. Dorsal surface with mostly white thread-like setae near posterior and posterolateral margins, and sometimes on mandibles. — Thorax: Thorax dorsally dark, with sparsed light brown patches, and ventrally light brown with small dark spots on setal insertion. Thorax dorsal surface with dark and white thread-like setae, and many short, black dolichasters. Mesothoracic spiracle borne on tubercle, which is about two times longer than broad. — Abdomen: Abdomen with dark thread-like setae on dorsal surface. Abdomen with prunescence, dorsally dark, with sparsed light brown patches, and ventrally light brown with small dark spots on setal insertion and near each basal abdominal margin. Abdominal setiferous processes with black and white setae.
Holotype: BRAZIL • ♂; BAHIA, Andaraí, Rio Piabas; 12°57′02.3”S 41°16′37.4”W; 25.xi.2018; Coleta Noturna, Lençol; Assmar, A;
“Clepsydra” translates to “hourglass” in Latin. The characteristic pattern formed by transverse crossveins in forewing presectoral and radial sector area resembles an hourglass (Fig.
Brazil (Fig.
Larvae were found anchored to bedrock, buried in small pools of shallow water-deposited sand near a river (Fig.
This species is remarkably distinguishable from other Ameromyia species, mostly because of the contrasting black and white coloration (Fig.
Ameromyia clepsydra sp. nov.: A lateral view and B posterior view of male terminalia. C Head, thorax and legs, ventral view. D Illustration of male terminalia, lateral view. — Abbreviations: cs, clavate setae; et, ectoproct; etl, ectoproct postventral lobe; go, gonarcus; hs, hinge setae; pah, paramere hinge; pap, paramere plate; pat, paramere tooth; st, sternite; tg, tergite.
This species wing venation patterns and crossvein density on both wings seems to be very plastic. Specimens collected at Bahia state have a biareolate pattern which resembles an hourglass or an “8” shape on presectorial and radial areas of both wings (Fig.
Ameromyia dimidiata
=Ameromyia baronei
Argentina; Paraguay (Fig.
Wing veins all pale. Cells in mediocubital area at least half brown suffused until at least 1/3 of wing span. Legs pale colored in contrast with dark abdomen and ventral thorax. Male ectoproct posterior margin with a weak dorsal notch.
Adult. Head: Antennae clubbed, 33 flagellomeres, dark brown with flagellum darker than scapus and pedicel. Vertex dark brown anteriorly, and light brown with dark brown patches posteriorly to anterior row. Vertex decumbent setae black, sometimes white on interantennal area. Interantennal area brown, usually with dark brown patches under scapus, in the middle of frons and just above clypeus. Frons pale brown, dark brown, or pale brown with dark brown patches. Frons setae black. Interocular distance less than eye width. Clypeus and labrum light brown. Mandibles dark brown. Palpi light brown, with external face of palpomeres slightly darker. — Thorax: Thorax dark brown, with light brown markings. Thorax on lateral view dorsally light brown and ventrally dark brown, with an evident longitudinal pale brown band under wings. Pronotum as broad as long, dark brown, with lateral margins pale brown, and a longitudinal medial pale brown streak. Pronotum marginal setae white. Mesonotum dark brown, with small light brown patches. Mesonotum with scattered white setae and curved white setae pointing posteriorly on mesoscutellum posterior margin. Mesoscutellum with white curved setae pointing posteriorly on posterior margin. Metanotum medially light brown and laterally dark brown, with white setae on posterior margins of metascutum and metascutellum. — Wings: Wing tip acute. Veins pale brown. Wing membrane hyaline, with dark brown infuscation on forewing mediocubital area, which is heavily suffused at forewing base and fading until apical third. Rhegmal area hyaline. Forewing vein CuP and hind wing vein CuA extending well before CuA fork and MP fork respectively, with at least nine crossveins connecting them and posterior branch of fork. Pterostigma light brown, opaque, encompassing six veins. Cells delimited by transversal veins in hind wing posterior margin higher than long. — Legs: Coxae pale brown. Femur and tibia with black bristles. Femur pale brown, with external face slightly darker than internal face. Meso and metafemur pale brown with internal face slightly darker than external face. Decumbent setae on basal half of profemur white, and black on distal half. Profemural sense hair longer than profemur and mesofemural sense hair much shorter than mesofemur. Tibia pale brown. Tibial spurs slightly shorter than pretarsal claws. Five tarsomeres, pale brown with distal joint slightly darker. Pretarsal claws at least 1.5 times longer than basitarsus. — Abdomen: Abdomen dark brown with many setae, which are white on tergites and sternites I–III and black on remaining sclerites. Male ectoproct with a short round postventral lobes on ventral face. Parameres subtriangular on lateral view, when folded. Gonarcus arch positioned not above paramere plates on lateral view. Male genitalia clavate setae with globose club. — Larva. Larvae unknown.
The type of A. dimidiata has lost both its hind wings (Fig.
A single male specimen labeled from Colombia were identified from the
This species is distinguishable from other Ameromyia species due to its veins on forewing which are all pale, and the absence of a rhegmal infuscation on forewing (Figs
Ameromyia dimidiata: A male habitus, dorsal view. B Head, thorax and legs, lateral view. C Male terminalia, lateral view. D Head and forelegs, frontal view. E illustration of male genitalia, lateral view. — Abbreviations: cs, clavate setae; et, ectoproct; etl, ectoproct postventral lobe; go, gonarcus; hs, hinge setae; pah, paramere hinge; pap, paramere plate; pat, paramere tooth; st, sternite; tg, tergite.
(32♀21♂) ARGENTINA • 1♂; 16.ii.2014; sweep; BMC;
Thorax lateral black, without a lateral pale band; forewing CuP alternating black and white, with nine or more crossveins between CuA posterior fork branch; female terminalia with pregenital plate slightly gibbous.
Adult. Head: Antennae with 32–34 flagellomeres. Flagellomeres black, dorsal face sometimes yellow, light brown between flagellomeres. Vertex yellowish brown, vertex rows with large dark spots and/or patches. Vertex decumbent setae black. Interantennal area mostly dark, sometimes marginally yellowish brown. Frons setae black, or white. Frons dark near antennae, and yellowish brown near clypeus. Frons setae black. Interocular distance less than eye width. Clypeus yellowish brown, sometimes with dark brown patches on lateral margins. Labrum yellowish brown. Mandibles dark brown. Palpi light brown, much darker on external face of palpomeres. — Thorax: Thorax dark with yellowish brown markings. Thorax dark on lateral view, with no evident longitudinal band. Pronotum as broad as long, dark, with yellowish brown on markings and lateral margins. Pronotum marginal setae white. Mesonotum dark, with a medial longitudinal yellowish-brown stripe on each side of mesoscutum and medially on mesoscutellum. Metanotum dark, with yellowish-brown patches on metascutum. — Wings: Wing tip acute. Veins with dashed black and white pattern, except for hind wing CuA, which is dark. Wing membrane mostly hyaline, with many markings. Forewing presectorial area and on RP first branch area with dark infuscation alternating on vertical crossveins. Forewing mediocubital area with dark infuscations around vertical crossveins which become sparser along MA vein, culminating on rhegmal area. Brown faint infuscations around forewing general crossveins and gradates, and on hind wing poststigmal area. Forewing CuP and hind wing CuA veins extending before forewing CuA and hind wing MP fork respectively, with nine or more crossveins connecting them and posterior branch of fork. Pterostigma opaque, dark brown on basal half and white on distal half, encompassing six to seven crossveins. Cells delimited by transversal crossveins in hind wing posterior area higher than long. — Legs: Coxae dark. Legs internal face yellowish brown and external face dark, with many dark brown spots on setal insertion. Femur and tibia with black bristles. Femur decumbent setae white and black. Tibiae with three evident or subtle basal, medial and distal dark brown rings around tibiae circumference. Tibial spurs slightly shorter than pretarsal claws. Tarsomeres light brown, with distal fourth dark brown. Pretarsal claws at least 1.5 times longer than basitarsus. — Abdomen: Abdomen sclerites dark, with yellowish or light brown in posterior margin near terminalia, with brown setae. Male paramere plates subtriangular on lateral view. Gonarcus arch positioned not above paramere plates on lateral view. Male genitalia clavate setae with globose club. — Larva. Larvae unknown.
Holotype: BRAZIL • ♂; MARANHÃO, Caxias, Res. Ecol. Inhamum; 31.x–02.xi.2005; Lençol luz mista; F. Limeira-de-Oliveira; CZMA. Paratypes: BRAZIL – BAHIA • ♂; Miguel Calmon, Mulungu do Chiola; 02.i.2021; 11°24′15.9″S 40°38′58.9″W; Luz; Vieira-Silva, V.;
Brazil (Fig.
(17♀7♂) BRAZIL – AMAZONAS • 3♂2♀; Barcelos, Em Campina; 0°28′38″N 63°28′18″W; vii–viii.2007; Malaise; A.S.Filho and T.Krolow;
The name “explicata”, which means “unfolded”. This refers to the fact that, during this work, the only males whose genitalia were in an “unfolded” state (Fig.
A live female specimen was observed in the wild at Santa Rita de Cássia, BA, Brazil (Fig.
This species is very similar to A. nigriventris. The most evident distinction is the completely dark coloration of the thorax lateral sides (Fig.
Ameromyia explicata sp. nov.: A female paratype, habitus, dorsal view. B Holotype, head and forelegs, frontal view. C Illustration of male genitalia, lateral view. D illustration of female genitalia, ventral view. — Abbreviations: cs, clavate setae; et, ectoproct; etl, ectoproct postventral lobe; go, gonarcus; gp, gonapophyseal plate; hs, hinge setae; lg, lateral gonapophysis; pah, paramere hinge; pap, paramere plate; pat, paramere tooth; pg, posterior gonapophysis; pp, pregenital plate; st, sternite; tg, tergite.
This species has a northern distribution in Brazil, from the Amazon Forest to the Northeastern region. Specimens collected in Miguel Calmon, Bahia, Brazil, cohabit this locality with A. nigriventris. However, all A. explicata sp. nov. specimens were collected at January, and during a field trip to Miguel Calmon in June, only A. nigriventris specimens were able to be found in the wild, which might suggest these species are separated by seasons. The specimens collected in the Amazon were found in “campinas”, which are open areas with sandy soil distributed within the Amazon rainforest. Those specimens have a few differences from the northeastern brazilian specimens, but those are mostly slight variations in thorax and leg coloration and are hereby considered to be the same species despite the biome and geographical disparity. A few female specimens from Maranhão state bear a noticeable posterior projection on the pregenital plate, slender and longer digging setae on ectoproct and lateral gonapophysis, and a different pattern on wing coloration. It is possible those females are in fact a different species. However, until a male specimen is analyzed, we chose to identify these specimens as A. explicata sp. nov., and opted to not include them among the paratypes.
Venezueleon guaricus
Venezuela (Fig.
Pronotum broader than long; femoral sense hairs absent; profemur without leg bristles; tibiae leg bristles equal or subequal to tibial width; pretarsal claws shorter than distal tarsomere; forewing with long rhegmal infuscation connecting both banksian lines.
Adult. Head: Antennae with 30–31 flagellomeres. Antennae dark brown, flagellomeres darker than scape and pedicel. Vertex light brown, with dark brown patches medially, dark brown spots on anterior and posterior rows and light brown markings on middle row. Vertex decumbent setae white, sometimes black on interantennal area. Frons pale brown, sometimes with dark brown patches medially. Frons setae black. Clypeus pale brown, slightly lighter near labrum. Labrum pale brown, slightly darker than clypeus. Mandibles black, completely covered by clypeolabrum. Palpi pale brown, distal palpomeres darker around sensory area. — Thorax: Thorax on lateral view dark brown, with an evident or faint longitudinal pale brown band under wings. Pronotum broader than long, pale brown with longitudinal dark brown markings laterad of a longitudinal medial pale brown stripe. Mesonotum pale brown, with dark brown laterally on mesoscutum. Metanotum pale brown with dark brown laterally on metascutum. Thorax with dark brown spots on setal insertions. — Wings: Wing tip acute. Forewing membrane with a slight brown tint, hind wing membrane hyaline. Dark brown infuscations along forewing crossveins which form the banksian lines; along rhegmal area connecting both banksian lines; along dashed crossveins on radial area; along few crossveins on mediocubital area after CuA fork; along crossveins connecting with CuA posterior branch of fork posterior margin; and on general crossveins and gradates near wing margin. Forewing mediocubital area with dark infuscations around vertical crossveins which become sparser along MA vein, culminating on rhegmal area. Forewing CuP vein and hind wing CuA extending well before forewing CuA fork and hind wing MP fork respectively, with at least four crossveins connecting them and posterior branch of fork. Pterostigma brown on basal half and pale on distal half, opaque, encompassing five to six crossveins. Cells delimited by transversal veins in hind wing posterior area longer than high. — Legs: Coxae brown. Legs pale brown, with posterior face much darker than anterior face. Profemur without black bristles. Meso and metafemur with very few black bristles which are shorter than femoral width. Femoral sense hairs absent. Tibiae with black bristles that are shorter than tibial width. Tibial spurs longer than pretarsal claws. Tarsomeres pale brown, dark brown on ventral surface. Pretarsal claws shorter than distal tarsomere. — Abdomen: Abdomen dark brown, with many brown setae. Male parameres plates on lateral view with a round dorsoposterior margin. Gonarcus arch positioned not above paramere plates on lateral view. Male genitalia clavate setae with sickled clubs. — Larva (third instar). Head: Head capsule slightly broader than long in dorsal view, slightly longer than broad in ventral view. Head capsule dorsally dark brown, with few light brown patches medially. Clypeolabrum dark, darker near mandibles. Head capsule ventrally light brown. Short dolichasters on head dorsal surface cone-like. Head capsule dolichasters mostly black, with a few white dolichasters near anterior and lateral margins. Row of dolichasters on labrum anterior margin black. Head dorsal surface covered with white thread-like setae, mostly on head capsule posterolateral margins, and mandibles base. — Thorax: Thorax dorsally dark brown, with sparsed light brown patches, and ventrally light brown with small dark spots on setal insertion. Thorax dorsal surface with many short, black dolichasters, and white thread-like setae. Mesothoracic spiracle borne on tubercle, which is about two times longer than broad. — Abdomen: Abdomen covered with white thread-like setae dorsally. Abdomen dorsally light brown, with sparsed grey and dark brown patches, and ventrally dark brown with large triangular light brown patches medially at each sternite, with small dark spots on setal insertion and near each basal abdominal margin. Lateral face ventrad of the ventral setiferous processes of spiracles with white thread-like setae. Abdominal setiferous processes with black and white setae.
This species is known only by the specimens presented in the original description. Previously a monotypic genus, Venezueleon is hereby synonymized under Ameromyia. Both genera had very similar diagnosis, differing only in regard of the size of the pretarsal claws (large in Ameromyia and small in Venezueleon) and profemoral sense hairs (long in Ameromyia and absent in Venezueleon) (Fig.
According to
Ameromyia guarica: A paratype fore and hind wings, dorsal view. B Holotype, head and forelegs, frontal view. C Illustration of male genitalia, lateral view. D Holotype male terminalia, lateral view. — Abbreviations: cs, clavate setae; et, ectoproct; etl, ectoproct postventral lobe; go, gonarcus; hs, hinge setae; pah, paramere hinge; pap, paramere plate; pat, paramere tooth; st, sternite; tg, tergite.
Paratype: (1♂, 1 larvae) VENEZUELA – GUARICO • 1♂[paratype], 1 larvae; 15 km S Calabozo, Rio Orituco; 27.ii.1988; R. B. Miller and L. A. Stange;
Amazoleon modesta
Ameromyia modesta
(Banks). –
Brazil; Suriname; Venezuela (Fig.
Prothorax dark brown with a single longitudinal white line in the middle. Dark brown streak in rhegmal area, formed by infuscation around rhegmal crossveins. External and internal face of legs with visible contrast between dark brown and light brown.
Adult. Head: Antennae with 31–35 flagellomeres. Flagellum pale brown, turning dark brown near the club. Vertex dark brown, slightly pale brown laterally. Vertex decumbent setae black, sometimes white on interantennal area. Interantennal area with a large brown spot. Frons pale brown with slight brown medial patches, and grayer near clypeus. Frons setae black. Interocular distance less than eye width. Clypeus and labrum light brown. Mandible light brown on basal half and black on distal half. Palpi light brown, and slightly darker on external face of palpomeres. Head without thread-like setae. — Thorax: Thorax mostly dark brown with a medial longitudinal white band, and light brown around thorax sutures. Thorax on lateral view dark brown, with an evident or faint longitudinal pale brown band under wings. Pronotum as broad as long. Pronotum dark brown, with a longitudinal medial white streak. Pronotum marginal setae white. Mesonotum dark brown, sometimes with pale brown on prescutum, and mesoscutellum with pale brown medially and on posterior margin. Mesonotum with scattered white setae and curved white setae pointing posteriorly on mesoscutellum posterior margin. Metanotum dark brown, with metascutellum with pale brown medially and on posterior margin. Metanotum with scattered white setae and curved white setae pointing posteriorly on metascutellum posterior margin. — Wings: Wing tip acute. Veins with dashed dark and pale brown pattern. Wing membrane hyaline, except on most vein junctures which have brown infuscation, along forewing CuA vein, and on rhegmal area which have dark brown infuscations. Forewing mediocubital area with dotted pattern, with dark infuscations around vertical crossveins. Forewing CuP and hind wing CuA veins extending before CuA and MP fork respectively, with at least four crossveins connecting them and posterior branch of fork. Pterostigma opaque, white, encompassing three to four crossveins. Cells delimited by transversal veins in hind wing posterior area longer than high. — Legs: Coxae dark brown. Femur and tibia with black bristles. Legs with external face of femur, tibiae and tarsomeres dark brown contrasting to internal face which is light brown. Femur decumbent setae black. Profemural sense hair longer than profemur and mesofemural sense hair much shorter than mesofemur. Tibial spurs slightly shorter than pretarsal claws. Five tarsomeres. Pretarsal claws at least 1.5 times longer than basitarsus. — Abdomen: Abdomen sclerites dark brown medially and slightly paler on basal and apical margins, with many dark setae. Male parameres plates on lateral view with a round dorsoposterior margin. Paramere tooth curved upwards on lateral view. Gonarcus arch positioned not above paramere plates on lateral view. Male genitalia clavate setae with sickled club. — Larva (third instar). Head: Head capsule slightly broader than long in dorsal view, slightly longer than broad in ventral view. Head capsule dorsally dark brown, with few light brown patches medially. Clypeolabrum dark, darker near mandibles. Head capsule ventrally light brown. Short dolichasters on head dorsal surface cone-like. Head capsule dolichasters mostly black, with a few white dolichasters near anterior and lateral margins. Row of dolichasters on labrum anterior margin black. Dorsal surface on posterolateral margins with dark thread-like setae. — Thorax: Thorax dorsally dark brown, with sparsed light brown patches, and ventrally light brown with small dark spots on setal insertion. Thorax dorsal surface with dark thread-like setae, and many short, black dolichasters. Mesothoracic spiracle borne on tubercle, which is about two times longer than broad. Metathoraxic surface with whitish prunescence medially. — Abdomen: Abdomen with dark thread-like setae dorsally. Abdomen with a whitish prunescence, dorsally light brown, with sparsed grey and dark brown patches, and ventrally light brown with small dark spots on setal insertion and near each basal abdominal margin. Abdominal setiferous processes with black and white setae.
Ameromyia modesta is recorded from Brazil and Suriname for the first time. The type locality known as “Akuriman” or “Acurima”, refers to a mountain near the border between Venezuela and Brazil (Roraima), on the municipality of Gran Sabana.
This species is very similar in coloration to A. guarica, but it bears profemoral and mesofemural sense hairs, larger pretarsal claws and a longer than broad pronotum. Regarding other species in the same group, A. modesta differs from A. pubiventris in the pronotal markings (in which the pronotum is mostly light brown with a longitudinal stripe) (Figs
Ameromyia modesta: A fore and hing wing, dorsal view. B Holotype head and foreleg, frontal view. C Holotype’s hind wing, dorsal view. D Holotype terminalia, lateral view. E Illustration of male genitalia, lateral view. F Holotype abdomen. Holotype photographs by Charles Farnum, Museum of Comparative Zoology, Harvard University. — Abbreviations: cs, clavate setae; et, ectoproct; etl, ectoproct postventral lobe; go, gonarcus; hs, hinge setae; lg, lateral gonapophysis; pah, paramere hinge, pap, paramere plate; pat, paramere tooth; pg, posterior gonapophysis; st, sternite; tg, tergite.
(24♀8♂, 1 larva): BRAZIL – RORAIMA • 1♀; Beira do rio Uraricoera; 25.iii.2016; 3°20′59″N 61°25′22.3″W; 83m; luz móvel; CC Gonçalves;
Ameromyia muralli
Bolivia; Brazil (Fig.
Forewing mediocubital area with evident continuous or dotted dark suffusion at least until reaching CuA fork. Hind wing shape falcate. Profemur with homogeneous color pattern, with no contrast between external and internal face.
Adult. Head: Antennae with 31–33 flagellomeres. Flagellomeres light brown with slightly darker brown patches. Vertex light brown with a latitudinal dark brown dotted streak. Middle and posterior row of vertex with dark brown spots. Vertex decumbent setae black. Interantennal area light brown. Frons marginally dark brown and light brown centrally. Frons setae black. Clypeus and labrum light brown. Mandibles light brown on basal half, and black on distal half. Palpi light brown, with external face of palpomeres much darker. — Thorax: Thorax dark brown with yellowish-brown markings. Thorax on lateral view dark brown, with an evident longitudinal pale brown band under wings. Pronotum as broad as long. Pronotum dark brown, with a yellowish-brown longitudinal stripe medially and near lateral margins. Mesonotum dark brown, with yellowish-brown on notal junctures, and medially on mesoscutellum. Metanotum dark brown with yellowish-brown on notal junctures. — Wings: Forewing apex weakly falcate, and hind wing falcate. Veins with dashed dark and pale brown pattern. Wing membrane hyaline, with a dark infuscations on crossveins of forewing mediocubital area, along CuA vein, and on rhegmal area. Forewing mediocubital area with continuous or dotted dark infuscations around vertical crossveins at least until CuA fork. Forewing CuP and hind wing CuA veins extending before CuA and MP fork respectively, with nine or more crossveins connecting them. Pterostigma opaque, dark brown on basal half and white on apical half, encompassing six to seven crossveins. Cells delimited by transversal veins on hind wing posterior area higher than long. — Legs: Coxae dark brown. Femur and tibiae with black bristles. Legs light brown, with many dark brown spots. Femur decumbent setae white and black. Profemural sense hair longer than profemur and mesofemural sense hair much shorter than mesofemur. Tarsomeres light brown, with distal fourth dark brown. Pretarsal claws at least 1.5 times longer than basitarsus. — Abdomen: Abdomen dark brown, with white setae on first sclerites, and brown setae on remaining sclerites. Male parameres oblong or subtriangular on lateral view. Gonarcus arch positioned not above paramere plates on lateral view. Male genitalia clavate setae with globose club. — Larva. Larvae unknown.
Ameromyia muralli is recorded from Bolivia for the first time, more precisely in Puerto Suárez, right on the border with Mato Grosso do Sul state in Brazil. The analysis of the holotype was based on old photo slides (Fig.
Ameromyia muralli: A habitus, lateral view. B Dorsal view. C Holotype habitus, dorsal view, photographed by Lionel Stange. D Illustration of male genitalia, lateral view. — Abbreviations: cs, clavate setae; et, ectoproct; etl, ectoproct postventral lobe; go, gonarcus; hs, hinge setae; pah, paramere hinge; pap, paramere plate; pat, paramere tooth; st, sternite; tg, tergite.
This species can be readily recognized by the falcate hind wings (Fig.
(19♀3♂): BOLIVIA – GERMÁN BUSCH • 1♀; Puerto Suarez; J. Steinbach;
Myrmeleon nigriventris
Ameromyia nigriventris
(Walker). –
=Ameromyia hirsuta
=Ameromyia stevensi
=Moza longiventris
=Ameromyia longiventris (Navás). –
=Ameromyia stevensi striolata
Argentina; Brazil; Colombia; Guyana; Suriname; Uruguay; Venezuela (Fig.
Forewing veins Sc, RA and CuP with alternating coloration; lateral side of thorax with an evident longitudinal pale band; male gonarcus positioned obliquus to paramere plates.
Adult. Head: Antennae with 34 flagellomeres. Flagellomeres dark brown, with ventral face of the club light brown. Vertex brown with dark brown spots on all vertex rows, and a dark brown transversal streak on anterior and posterior rows forming a “T” shape. Vertex decumbent setae black, sometimes white on interantennal area. Interantennal area usually brown, with dark brown patches. Frons varies, from completely light brown, to light brown with oblique dark brown streaks and completely dark brown. Frons setae all dark, all white, or both dark and white. Interocular distance less than eye width. Clypeus pale brown. Labrum pale brown. Mandibles light brown on basal half and dark brown on distal half. Palpi pale brown, slightly darker on external face of palpomeres. — Thorax: Thorax dark brown, with pale brown markings. Thorax on lateral view dark, with an evident lateral longitudinal band under wings. Pronotum as broad as long, pale brown with two mirrored longitudinal dark brown “P” shaped bands, and two lateral small dark brown streaks. Pronotum marginal setae white or dark brown. Mesonotum dark brown with a longitudinal light brown streak on mesoscutellum. Metanotum dark brown with two longitudinal light brown patches on metascutum. — Wings: Tip acute. Veins with dashed dark brown and light brown pattern, except for forewing MA vein which has dark brown and white pattern. Wing membrane hyaline, with or without brown infuscations along radial, mediocubital and/or cubital areas. When infuscated, forewing mediocubital area with dark infuscations around vertical crossveins with a dotted uniform pattern, or with a dotted pattern which become sparser along MA vein, culminating on rhegmal area. Wing membrane sometimes with a slight brown tint along costal and radial areas. Forewing CuP vein and hind wing CuA extending well before forewing CuA fork and hind wing MP fork respectively, with nine or more crossveins them and posterior fork branch. Pterostigma frosted white, encompassing three to four crossveins. Cells delimited by transversal veins in hind wing posterior area higher than long. Female hind wing sometimes with an opaque longitudinal streak on apex. — Legs: Coxa dark. Femur and tibiae with black bristles. Profemur pale brown with exterior face dark brown. Meso and metafemur exterior face pale brown and interior face dark brown. Femur decumbent setae white and black. Profemural sense hair longer than profemur, and mesofemural sense hair much shorter than mesofemur. Tibial spurs slightly shorter than pretarsal claws. Tarsomeres pale brown with distal tip dark brown. Pretarsal claws at least 1.5 times longer than basitarsus. — Abdomen: Abdomen dark brown, except for tergite 9 and ectoprocts which are pale brown with dark brown patches. Tergites 1, 2, and 3 with many short white setae, and subsequent tergites with many short dark brown setae. Sternite I with many short white setae, and male sternite III sometimes with many stout white setae. Sternite II, female sternite III and subsequent sternites with short dark brown setae. Male ectoproct with a short, globose postventral lobe. Male paramere plates subtriangular, almost subquadrangular on lateral view. Gonarcus arch positioned slightly above paramere plates on lateral view. Male genitalia clavate setae with globose club. — Larva (third instar). Head: Head capsule as broad as long in dorsal view, slightly longer than broad in ventral view. Head capsule dorsally brown, with few dark brown patches near posterior margin. Clypeolabrum light brown, darker near mandibles. Head capsule ventrally light brown. Short dolichasters on head dorsal surface cylindrical, or almost spherical. Head capsule dolichasters black and white, mostly white near anterior and lateral margins. Dorsal surface on posterior and posterolateral margins covered with short, white thread-like setae. Row of dolichasters on clypeolabrum posterior margin white, black between mandibles. — Thorax: Thorax prunescent, dorsally brown or light brown, with very sparsed brown patches which have no prunescence, and ventrally light brown with small dark spots on setal insertion. Thorax dorsal surface with sparse, short white thread-like setae, and with many short, black dolichasters medially, and many white dolichasters around lateral margins. Dorsal surface covered with very short “star-shaped” dolichasters. Mesothoracic spiracle borne on tubercle, which is much broader than long. Tubercule covered with short, white dolichasters. — Abdomen: Abdomen with white thread-like setae, which are sparse on dorsal surface, and ventrally are present on sternites I–VII, and tangled in circle-like shapes. Dorsal surface covered with very short “star-shaped” dolichasters. Abdomen prunescent, dorsally brown or light brown, with three equally spaced black patches medially on each abdominal segment, and ventrally light brown with small dark spots on setal insertion and near each basal abdominal margin. Abdominal tergites with many short white and black dolichasters, and sternites with white and black dolichasters. Abdominal setiferous processes with only white setae.
Specimens were observed in their natural habitat at the municipality of Miguel Calmon, Bahia, Brazil. Eight adult specimens (three males and five females) and 14 larvae were captured and subsequently reared in captivity. Adults were paired in different sex couples in order to observe possible matings, and females were provided with earth and sand from their natural habitat in order to observe possible oviposition. A single captive female was observed in posture, similar to what as described by
Larvae were found on patches of sandy, loose earth above compacted clay soil. Almost always, the larvae were inside a dug out “trench”, achored to the hard substrate (Fig.
All the type specimens were analyzed, except for A. stevensi striolata holotype. Ameromyia nigriventris previous syntypes consists of one male (paralectotype) (Fig.
Ameromyia nigriventris: A male paralectotype, frontal view. B Lectotype forewing. C Female paralectotype forewing. D A. hirsuta holotype, head, thorax and legs, lateral view. E Illustration of male genitalia, lateral view. — Abbreviations: cs, clavate setae; et, ectoproct; etl, ectoproct postventral lobe; go, gonarcus; hs, hinge setae; pah, paramere hinge; pap, paramere plate; pat, paramere tooth; st, sternite; tg, tergite.
In this work, we partially agree with Walker circumscription for A. nigriventris. The female paralectotype morphotype is most common in central Brazil, and this is the only specimen analyzed with this morphotype outside Brazil. A single male specimen with this morphotype was analyzed, and male genitalia agrees with A. nigriventris. However, the male paralectotype is enigmatic. It has a dark forewing CuP and hind wing CuA, as well as a suffused forewing mediocubital area, which agrees with A. tendinosa, but forewing RP and hind wing MA and MP have dashed patterns, in contrast to A. tendinosa respective wing veins that are completely dark. The completely suffused mediocubital area on forewing was present only in this specimen among all A. nigriventris analyzed during this work. Additionally, A. tendinosa has not been found outside central South America. In fact, no other species that bear a complete suffusion of forewing mediocubital area has been registered for Colombia. Nonetheless, despite the inconsistencies, we are still identifying the male paralectotype as an anomalous specimen of A. nigriventris. However, it is possible that this specimen in fact does represent a different species and further analyses could better elucidate the identity of this specimen. Unfortunately, we were not able to analyse its genitalia.
Ameromyia stevensi holotype (Fig.
We also agree with Stange’s synonymy of A. longiventris under A. hirsuta (
Ameromyia nigriventris larvae was previously described as not having thread-like setae (
Ameromyia nigriventris is the species with the largest degree of variation regarding wing infuscation patterns within the genus. Wings can bear color pattern equal to that of the lectotype, equal to that of the female paralectotype; a combination of both; devoid of infuscation or with an almost imperceptible infuscation in the aforementioned patterns. Ameromyia explicata sp. nov. and A. tendinosa are very morphologically similar to A. nigriventris. However, the first has a fully dark meso and metathorax on latheral view, while A. nigriventris has an evident lateral pale band, and the latter has a complete suffusion on forewing mediocubital area, as well as a dark colored forewing CuP vein, in contrast to A. nigriventris, which suffusion on forewing mediocubital area varies, and has a dashed pattern on forewing CuP.
(115♀62♂4?, 3 larvae) ARGENTINA • 1♀; MISIONES, Loreto; iii.1953; Bought by F.H.Walz;
Ameromyia pleuralis
Brazil (Fig.
Forewing veins C, Sc, and RA all pale, in contrast to MA which has a dashed pale brown and dark brown pattern, and CuP which are dark brown.
Adult. Head: Antennae with 31 flagellomeres. Flagellomeres dark brown, with ventral face of the club light brown. Vertex light brown with dark brown spots on setae insertion, and dark brown spots and patches on anterior and posterior rows. Vertex decumbent setae black. Interantennal area dark brown. Frons light brown, usually with dark brown medially and under scape. Frons setae black. Interocular distance less than eye width. Clypeus light brown, slightly darker medially. Labrum light brown, slightly darker on ventral margin. Mandibles dark brown. Palpi light brown, with exterior face slightly darker. — Thorax: Thorax on lateral view darker ventrally, with an evident longitudinal pale brown band under wings. Pronotum as broad as long, light brown, with two longitudinal dark brown markings medially. Pronotum marginal setae white. Mesonotum dark, with longitudinal light brown streaks on each side of mesoscutum, and one medially on mesoscutellum. Metanotum light brown, with metascutum laterally dark. — Wings: Wing tip acute. Forewing veins Sc and RA vein pale; CuA, anterior branch of CuA fork and MP with a dashed dark and light brown pattern, and remaining veins are dark brown. Hind wing veins C, Sc and RA vein pale, and remaining veins are dark brown. Wing membrane hyaline, except on Costal and Radial areas, which have a brown tint. Forewing mediocubital area with dark infuscations around vertical crossveins, in a continuous pattern or apparently dotted but with connecting suffusions pattern, at least until apical third. Forewing CuP vein and hind wing CuA extending well before forewing CuA fork and hind wing MP fork respectively, with nine or more crossveins connecting them and posterior branch of fork. Pterostigma opaque, light brown, encompassing five to six crossveins. Cells delimited by transversal veins in hind wing posterior area higher than long. — Legs: Coxae light brown on posterior face and slightly darker on anterior face. Femur and tibiae with black bristles. Legs light brown, dark brown on exterior face. Femur decumbent setae white and black. Profemural sense hair longer than profemur, and mesofemural sense hair much shorter than mesofemur. Tibiae with dark brown spots on bristle insertion. Tibial spurs slightly shorter than pretarsal claws. Tarsomeres light brown, with posterior tip dark brown. Pretarsal claws at least 1.5 times longer than basitarsus. — Abdomen: Abdomen dark brown with many dark brown setae. — Larva. Larvae unknown.
Ameromia pleuralis holotype is labelled as “syntypus” (Fig.
This species is very similar to A. tendinosa and A. dimidiata, but the latter has a southern distribution and has a pale forewing CuP vein, while A. tendinosa has a forewing R vein with a dashed coloration distally.
(3♀3♂) BRAZIL – GOIÁS • 1♀1♂; Jataí, Faz. Nova Orlandia; i.1964; Martins, Morgante and Silva;
Myrmeleon protensis
Nemotolus protensis
(Gerstaecker). –
Ameromyia protensa
(Gerstaecker). –
=Foya trapezia
=Ameromyia alterna
=Ameromyia decarloi
Argentina, Brazil, Paraguay; Uruguay (Fig.
Forewing RP veins, basal half of CuP and banksian lines black in contrast to other veins which are mostly pale. Forewing mediocubital area cells at basal third at least 1/3 suffused with dark brown. Hind wing RP and banksian lines veins and black in contrast to other hind wing veins which are mostly pale. Clavate setae on male genitalic sac very long.
Adult. Head: Antennae with 27–29 flagellomeres. Flagellomeres dark brown, except the ones at the club, which are light brown ventrally. Vertex light brown, with dark brown anterior and middle rows, and two brown spots in posterior row. Vertex decumbent setae black. Interantennal area dark brown. Frons light brown, with dark brown patches medially. Frons setae black. Interocular distance less than eye width. Clypeus light brown, with two medial dark brown markings. Labrum light brown, with very faint dark brown patches that connect with clypeus dark brown markings. Mandibles with external face light brown, and internal face dark brown. Palpi light brown, with external face slightly darker. Mandibular and labial distalmost palpomeres with dark brown tip. — Thorax: Thorax light brown, with dark brown markings. Thorax on lateral view dark brown, with an evident longitudinal light brown band under wings. Pronotum as broad as long, light brown with dark brown markings. Pronotum marginal setae white and/or black. Mesonotum light brown, with dark brown patches on prescutum and dark brown laterally on mesoscutum. Metanotum light brown, mesoscutum dark brown laterally. — Wings: Wing tip acute. Forewing RA, MA and A veins pale; RP and Banksian Lines veins dark; CuP dark on basal half and dashed near posterior wing margin; CuA and MP veins with dark and pale sections; remaining veins with dashed pattern. Hind wing veins R and MA pale; RP veinsdark; remaining veins with dashed pattern. Wing membrane hyaline, or sometimes with a faint brown tint. Forewing mediocubital area with dark brown infuscations along CuA vein until at least half of forewing distance, which cover at least 1/3 of the cells. Forewing CuP and hind wing CuA veins extending before forewing CuA and hind wing MP fork respectively, with at least four crossveins connecting them and posterior branch of fork. Pterostigma opaque, dark yellow, encompassing three to four crossveins. Cells delimited by transversal veins on hind wing posterior area longer than high. — Legs: Coxae dark brown anteriorly and light brown posteriorly. Legs light brown and with many dark brown spots on setae insertion. Femur and tibiae with black bristles. Femur light brown, with external face slightly darker than internal face. Femur decumbent setae white and black. Profemural sense hair longer than profemur and mesofemural sense hair much shorter than mesofemur. Tibial spurs slightly shorter than pretarsal claws. Tarsomeres light brown, with distal fourth dark brown. Pretarsal claws at least 1.5 times longer than basitarsus. — Abdomen: Abdomen dark brown ventrally, with brown setae. Tergites I–III light brown, and remaining tergites dark brown. Male paramere plates quadrangular on lateral view. Gonarcus arch positioned not above paramere plates on lateral view. Male genitalia clavate setae very long, with a sickled club. — Larva. Larvae unknown.
The type specimen of Foya trapezia was not analyzed in this work. Foya trapezia has a controversial taxonomical background as it was synonymized under Amazoleon pubiventris along with Myrmeleon tendinosus by
Ameromyia protensa: A male specimen, dorsal view. B Lectotype head and forelegs, frontal view. C Lectotype head, thorax and legs, lateral view. D Male terminalia, lateral view. E Illustration of male genitalia, lateral view. Type photographs by Lara Lopardo (
The type specimens of A. alterna and A. decarloi were not located, and are still missing as previously mentioned by
Ameromyia protensa is the only species from the modesta group that occurs in Argentina, Paraguay and Uruguay (Fig.
(7♀9♂) ARGENTINA • 1♂; S of Corrientes, Parana River; 16.i.2009, M.Snizek;
Myrmeleon pubiventris
Amazoleon pubiventris
(Walker). –
Ameromyia pubiventris
(Walker). –
Brazil; Suriname (Fig.
All wing veins with dashed dark brown and pale brown coloration pattern. Thorax dorsally pale in contrast to ventral face which is dark brown. Legs pale brown, with scattered dark brown spots. Abdomen pale brown with the distal third dark brown. Male genitalic sac extending laterally, and slightly dorsally above the gonarcus.
Adult. Head: Antennae with 26–29 flagellomeres. Flagellomeres pale brown, the ones at the club with many dark brown spots. Vertex light brown, with latitudinal dark brown streaks in anterior and middle row, and two brown spots in posterior row. Vertex decumbent setae black. Interantennal area pale or dark brown. Frons pale brown, frequently with dark brown patches centrally. Frons setae black. Interocular distance less than eye width. Clypeus pale brown. Labrum pale brown with gray patches medially. Mandibles light brown on basal half and dark brown on distal half. Palpi pale brown, with external face slightly darker. — Thorax: Thorax pale brown, with two dark brown longitudinal streaks dorsally. Thorax on lateral view dark brown, with an evident longitudinal pale brown band under the wings. Pronotum as broad as long, pale brown, with small dark brown spots and dark brown markings appearing as three longitudinal pale brown streaks. Mesonotum dark brown, mesoscutellum pale brown. Metascutum pale brown. — Wings: Wing tip acute. Veins with dashed dark and pale brown pattern. Wing membrane hyaline or with a brown tint, and dark brown infuscations on crossvein junctures. Forewing mediocubital area with dark infuscations around vertical crossveins in a uniform dotted pattern, culminating on rhegmal area. Forewing rhegmal area with an evident dark brown insfuscation. Forewing CuP and hind wing CuA veins extending before CuA and MP fork respectively, with at least four crossveins connecting them and posterior branch of fork. Pterostigma opaque, white, dark brown centrally, encompassing three to four crossveins. Cells delimited by transversal veins on hind wing posterior area longer than high. — Legs: Coxae dark brown on anterior face and pale brown on posterior face. Legs light brown with dark brown spots on setae insertion. Femur and tibiae with black bristles. Femur decumbent setae dark brown. Profemural sense hair longer than profemur and mesofemural sense hair much shorter than mesofemur. Tibial spurs slightly shorter than pretarsal claws. Tarsomeres light brown, with distal fourth dark brown. Pretarsal claws at least 1.5 times longer than basitarsus. — Abdomen: Abdomen pale and dark brown, sometimes banded, with many small dark brown spots. Tergites I–IV pale brown with, turning dark brown from tergite V onwards. Sternites II and III pale brown and marginally dark brown, and remaining sternites dark brown. Male paramere plates quadrangular with a round posterodorsal margin, on lateral view. Gonarcus arch positioned not above paramere plates on lateral view. Male genitalia clavate setae with a sickled club. — Larva. Larvae unknown.
Ameromyia pubiventris: A Lectotype, dorsal view. B Lectotype, habitus, dorsal view, with type labels, photographed by Ben Price (
During this work, only specimens from Brazil were analyzed. However,
Despite the records for Surinam and the type locality in the Amazon Region, this species appears to be more abundant in open areas in the Cerrado and Caatinga biomes in Brazil.
Ameromyia pubiventris can be easily differentiated from A. modesta and A. guarica by leg coloration, and from A. protensa by the much smaller posterior area of forewing, and dashed Radial sector veins (Figs
(124♀43♂8?) BRAZIL • 3♀3♂; Bates;
Brachynemurus strigosus
Banks 1909: 2 [Lectotype male, Pedregal, Argentina, 15.i.1906 (MCZ)]. –
Ameromyia strigosa
(Banks). –
=Ameromyia pentheri
=Ameromyia fidelis
Argentina; Brazil; Bolivia; Chile; Paraguay (Fig.
Dark and yellow coloration. Thorax setae all white. Profemur with decumbent setae all white. Wing veins with dashed dark brown and yellowish-brown pattern. Four to five obliquous streaks in forewing, around MA vein. Male ectoproct obliquus in relation to tergite IX.
Adult. Head: Antennae with 34 flagellomeres. Flagellomeres black, slightly lighter at antennae club, with yellow between segments. Vertex yellow, with dark latitudinal bands on anterior and middle rows, and dark spots on posterior row. Vertex decumbent setae all white, or black with white only on interantennal area. Frons yellow, with dark brown patches that can extend to interantennal area. Frons setae white. Interocular distance less than eye width. Clypeus yellow, sometimes with dark brown patches. Labrum yellow, with slightly darker ventral margin. Mandibles dark brown. Palpi yellow, slightly darker on external face of palpomeres. — Thorax: Thorax dark with yellow markings. Thorax dark on lateral view, with a faint longitudinal pale band under wings. Pronotum as broad as long, dark with two lateral and one medial longitudinal yellow stripe. Pronotum marginal setae white. Mesonotum dark, with yellowish-brown on notal junctures. Metanotum dark, with yellowish brown medially on metascutum. — Wings: Wing tip acute. Hind wing rarely falcate. Veins with dashed dark brown and pale yellow or white pattern. Wing membrane hyaline. Forewing with dark brown infuscations in wing membrane alternating around crossveins on presectorial and radial areas, and along crossveins on sectorial, mediocubital and rhegmal areas, forming five evident oblique streaks around MA vein. Forewing mediocubital area with dark infuscations around vertical crossveins which become sparser along MA vein, culminating on rhegmal area. Forewing CuP and hind wing CuA veins extending well before forewing CuA and hind wing MP fork respectively, with nine or more crossveins connecting them and posterior branch of fork. Pterostigma opaque, dark brown on basal half and white on apical half, encompassing six to seven crossveins. Cells delimited by transversal veins on hind wing higher than long. — Legs: Coxae dark. Femur and tibiae with black bristles. Femur dark on external face and light yellow on internal face. Profemur decumbent setae white. Meso and metafemur decumbent setae white and black. Profemural sense hair longer than profemur and mesofemural sense hair much shorter than mesofemur. Tibia light yellow with dark spots on setae insertion. Tibial spurs slightly shorter than pretarsal claws. Tarsomeres light yellow, with distal fourth dark brown. Pretarsal claws at least 1.5 times longer than basitarsus. — Abdomen: Abdomen sclerites black, sometimes with yellow on posterior margins. Abdomen with white setae, brown near terminalia. Male ectoproct obliquous in relation to tergite IX, with postventral lobe slightly elevated. Male paramere plates subtriangular on lateral view. Parameres hinge anterior projections not entirely covered by gonarcus. Gonarcus arch positioned not above parameres plates. Male genitalia clavate setae with globose club. — Larva. Larvae unknown.
A live male specimen was observed in wild at Santa Rita de Cássia, Bahia state, Brazil. The location where the specimen was collected was at the Cerrado biome, with dry vegetation and hard, compacted soil. This specimen hovered in the air almost two meters above the ground.
This species, previously restricted to Chile, Paraguay and Argentina, is now registered for the first time to Bolivia. It is also now recorded to Brazil, as Ameromyia pentheri is hereby proposed as a synonym of A. strigosa. The holotype of A. pentheri (Fig.
Ameromyia pentheri and A. strigosa: A dorsal view and B lateral view of A. pentheri holotype, habitus. C Head and forelegs of A. pentheri holotype, frontal view. D Head and foreleg of A. strigosa lectotype, frontal view, photographed by Charles Farnum, Museum of Comparative Zoology, Harvard University.
The type specimen of A. fidelis was not analyzed in this current work, but the original description greatly fits that of A. strigosa, and therefore we chose to maintain the synonymy proposed by
Ameromyia strigosa is very distinctive as it bears four to five oblique markings along MA vein in the forewing (Figs
Ameromyia strigosa lectotype: A head, thorax and abdomen (broken), dorsal view. B Head, thorax and legs, lateral view. C Fore and hind wing. D Male terminalia, ventral view. E Type labels. F Detached abdomen, ventral view. Photographs by Charles Farnum, Museum of Comparative Zoology, Harvard University. — Abbreviations: et, ectoproct; etl, ectoproct postventral lobe; st, sternite.
Ameromyia strigosa: A fore and hindwing, general shape and color. B Falcate hind wing variation. C Male terminalia. D Illustration of male genitalia, lateral view. — Abbreviations: cs, clavate setae; et, ectoproct; etl, ectoproct postventral lobe; go, gonarcus; hs, hinge setae; pah, paramere hinge; pap, paramere plate; pat, paramere tooth; st, sternite; tg, tergite.
(59♀27♂) ARGENTINA • 1♀; Rio Negro, Cipoletti; 20.i.1960; Bought by F.H.Walz;
Myrmeleon tendinosus
Ameromyia tendinosa
(Gerstaecker):
Brazil (Fig.
Wing veins CuP completely dark brown in contrast to other veins which have a dashed dark brown and pale brown pattern. Forewing cells in mediocubital area almost completely suffused until at least the middle of wing span.
Adult. Head: Antennae with 30–32 flagellomeres. Flagellomeres dark brown, except for the ones at the club, which are light brown dorsally or ventrally. Vertex light brown, with latitudinal dark brown streaks in anterior and middle row, or completely dark brown with faint light brown laterally. Vertex decumbent setae black. Interantennal area dark brown. Frons light brown, frequently with dark brown patches centrally, or entirely dark brown, light brown near clypeus. Frons setae black. Interocular distance less than eye width. Clypeus light brown brown with dark brown patches. Labrum light brown brown. Mandibles light brown on basal half, and dark brown on apical half. Palpi light brown, with external face of palpomeres slightly darker. — Thorax: Thorax dark brown with few light brown markings. Thorax dark brown on lateral view, with an evident longitudinal pale brown band under wings. Pronotum as broad as long, dark brown, with light brown anteriorly and laterally. Pronotum marginal setae white, or dark brown. — Wings: Wing tip acute. Veins with dashed dark and pale brown pattern, except for forewing and hind wing veins CuP which are dark brown. Wing membrane hyaline, except for forewing mediocubital area which is almost completely dark brown infuscated until forewing apical third, and the rhegmal area which is faintly dark brown infuscated. Forewing and hind wing sometimes with a faint dark brown infuscation alongside gradates. Forewing CuP and hind wing CuA veins extending before forewing CuA and hind wing MP fork respectively, with nine or more crossveins connecting them and posterior branch of fork. Pterostigma opaque, dark brown on basal half and white on distal half, encompassing six to seven crossveins. Cells delimited by transversal veins on hind wing posterior area higher than long. — Legs: Coxae dark brown. Legs internal face light brown and external face dark brown, with many dark brown spots on setal insertion, and on internal face in meso and metalegs. Femur and tibiae with black bristles. Femur decumbent setae white and black. Profemural sense hair longer than profemur and mesofemural sense hair much shorter than mesofemur. Tibial spurs slightly shorter than pretarsal claws. Tarsomeres light brown, with distal fourth dark brown. Pretarsal claws at least 1.5 times longer than basitarsus. — Abdomen: Abdomen sclerites dark brown, with light brown in posterior margin near the terminalia, with brown setae. Male paramere plates oblong on lateral view. Gonarcus arch positioned not above paramere plates on lateral view. Male genitalia clavate setae with globose club. — Larva. Larvae unknown.
The taxonomic background for this species is confusing. Decades after its original description by Gerstaecker,
This species is restricted to Brazil. The male paralectotype of A. nigriventris somewhat agrees with this species and can possibly be a specimen of A. tendinosa, which would expand this species distribution to the Amazon region including Colombia. Additionaly, three female specimens labeled as from Colombia were identified from the
Ameromyia tendinosa can be distinguished from other species in the nigriventris group by the dark forewing CuP and by the forewing mediocubital area which is completely suffused with dark brown infuscation from wing base at least to CuA fork (Figs
Ameromyia tendinosa: A fore and hind wings, B head and thorax, dorsal view, C head, thorax and legs, lateral view and D male terminalia of lectotype. E illustration of male genitalia, lateral view. Photographs by Lara Lopardo (
(5♀4♂) BRAZIL: MATO GROSSO • 1♀; Campo Novo do Parecis; xii.1960; M. Alvarenga;
Previously, Ameromyia sensu
The setose frons as well the enlarged pretarsal claws were found to be synapomorphies of the more inclusive group, the sampled Brachynemurini genera (Ameromyia, Argentoleon and Austroleon), and although Ameromyia specimens frequently have much more density of setae on the frons than Argentoleon and Austroleon, this appears to be a variable character and some specimens appear to have fewer setae than others. On the other hand, character state 39:1 “Clavate setae on male genitalia appears as a synapomorphy of Ameromyia by the present analyses.
The long CuA of the hind wing was discussed by
Ameromyia male genitalia was previously described as having no “hinge” (sensu
In Brachynemurini males, parameres have folds that, upon genitalia evertion, produces a movement in which the parameres fold upon themselves and greatly changes the genitalic complex apparent conformation (
The cladistic analyses recovered two species groups as it were previously suggested by Stange (modesta and nigriventris groups) (
The functionality of the thread-like setae is not fully understood, but it might be related to helping the larvae camouflage by holding debris in the same way as conical dolichasters and plumose setae (Acevedo-Ramos et al. 2021;
The modesta group corresponds to Amazoleon, previously delimited by
The second species group, nigriventris group, is composed of species that bear a broad posterior area of hind wing, with cells that are as high as long, or higher than long (Figs
The first species to branch off inside the nigriventris group, A. clepsydra sp. nov., features wing synapomorphies that are common to both groups while having conspicuous autapomorphies. Ameromyia clepsydra has the forewing CuP and hind wing CuA following posterior branch of CuA and MP fork respectivelly for a short distance, which agrees with the modesta group, but hind wing posterior area is much broader, which agrees with the nigriventris group. The course of forewing vein 2A also fits the pattern of the modesta group species (forewing vein 2A connected to 3A by a crossvein). However, a single analyzed specimen had both patterns, one in each wing (Fig.
The remaining species, as in clade C in our proposed classification (Fig.
The placement of A. muralli is different between both trees. In our proposed classification, this species was recovered as sister to clade D, while in the alternative tree it was recovered as sister to clade F. This rogue placement is the result of the great plasticity and homoplasy of the aforementioned forewing characters, which in the case of A. muralli, could allocate this species as sister to either clade under equal weighting analyses. Ameromyia muralli bear a profemur without contrasting colors, a suffused mediocubital area on forewing, and is polymorphic in relation to the state of the presectorial area infuscation pattern. In the proposed classification, the cladistic analysis indicates the profemur with contrasting coloration evolved independently, in both modesta and nigriventris groups (in clades B and D), while the presence of infuscation on forewing presectorial and radial area crossveins, and the multistate character of forewing mediocubital suffusion changed states many times inside Ameromyia. The alternative tree (Fig.
All weighted analyses recovered the same tree as our proposed classification for the genus, suggesting that, although both results under equal weighting show the same number of steps, the character of leg coloration (11) has more congruency than those of wing membrane coloration (16 and 17). Indeed, whilst not verified by many phylogenies, historically, Brachynemurini taxonomy is heavily biased towards leg morphology, and the pattern of wing markings and crossvein infuscation are very variable across the tribe, as it is within Ameromyia as well (
Clade E, although supported by a single synapomorphy (character state 17:1 Forewing presectorial and radial area crossveins with infuscations), is recovered as an unresolved polytomy of A. explicata sp. nov., A. nigriventris and A. strigosa, reflecting the large morphological similarity of the species in this group. Clade F on the other hand is supported by two forewing synapomorphies: a forewing CuP vein with homogeneous coloration and a clear forewing presectorial and radial areas, although the latter is in fact a reversion of the state transformation of this character for the nigriventris group. In clade F, A. tendinosa is sister to a clade comprising A. dimidiata and A. pleuralis (clade G). Although both A. tendinosa and A. pleuralis possess a dark forewing vein CuP, the analysis recovered the latter more related to A. dimidiata based on the pale forewing veins Sc and R.
In this work, we present Ameromyia as a valid Brachynemurini genus, with 12 valid species in two well defined species groups. Venezueleon was recovered deep inside Ameromyia and thus synonymized under the latter, which in retrospect makes taxonomical sense as the two genera had an overlap of their previously diagnostic characters, with some exceptions such as the size of pretarsal claws, which is a known plastic character at species level for Brachynemurini in general. Adult specimens from different species (but from the same species groups) bear almost identical male and female genitalia and are morphologically very similar, differing mostly in leg and wing vein coloration and wing marking patterns, although the latter appears to be very homoplasic and polymorphic within some species. More extensive analysis which utilizes different datasets such as larval morphology and/or molecular data could hopefully better elucidate Ameromyia internal relationships. This study also enlightens the need of better understanding the South American Brachynemurini genera, as results of this work partly contradict some statements from previous works, such as the case of all analyzed genera bearing hinged genitalia.
We would like to thank all the museums and collection curators that made the specimens available to this study by loan or pictures: Ben Price (
Tables S1, S2
Data type: .docx
Explanation note: Table S1. Character matrix (45 characters and 17 terminal taxa). Abbreviations for polymorphies: A = [0&1]; B = [0&2]. — Table S2.