Ameromyia, Banks, 1913

Tavares, Leon Gustavo de Miranda, Machado, Renato Jose Pires & Calor, Adolfo Ricardo, 2023, The Neotropical antlion genus Ameromyia Banks, 1913 (Neuroptera: Myrmeleontidae), systematics and redefinition under a phylogenetic approach, Arthropod Systematics & amp; Phylogeny 81, pp. 499-553 : 499

4.1. Ameromyia

Ameromyia Banks 1913: 227. [Type species: Brachynemurus strigosus Banks, by original designation]. - Banks 1943: 165 [key]. - Penny 1977: 38 [species list, distribution]. - Stange 1994: 90 [taxonomy, species list, phylogeny of tribe]. - Stange 2004: 227, 383, 392, 393 [species catalog, diagnosis, habitat; genera identification key (adults), genera identification key (larvae)]. - Machado et al. 2019: 4, 7, 14 [family phylogeny]. - Oswald 2021 [in genera catalog]. - Machado and Martins 2022 [faunal catalog].

= Amazoleon Banks 1913: 229 [Type species: Myrmeleon pubiventris Walker, by original designation]. - Navás 1914a: 205 [ Amazoleon = Moza ]. - Banks 1943: 164 [key]. - Stange 1967: 45 [synonymy] - Oswald 2021 [in genera catalog].

= Foya Navás 1914b: 53 [Type species: Foya trapezia Navás, by original designation and monotypy]. - Esben-Petersen 1920: 190 [synonymy = Amazoleon ].

= Nemotolus Banks 1943: 163 [Type species: Myrmeleon protensis Gerstaecker, by original designation]. - Stange 1967: 45 [synonymy].

= Venezueleon Stange 1994: 88 [Type species: Venezueleon guaricus Stange, by original designation and monotypy]. - Stange 2004: 249 [species catalog, diagnosis, habitat; genera identification key (adults), genera identification key (larvae)]. - Machado et al. 2019: 24 [genera list]. - Oswald 2021 [in genera catalog] [NEW SYNONYMY]

Distribution.

South America (Figs 12 , 13 ).

Species included.

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).

Diagnosis.

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.

Larvae known (Figs 14 , 15B, C , 16B, C , 19 , 26 , 27 , 30 , 36 ).

Ameromyia clepsydra sp. nov.; A. guarica comb. nov.; A. modesta ; A. nigriventris.

Description.

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.

Remarks.

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 9 , 10 ), as the paramere plates are enlarged and the paramere hooks are short and stout, and basad to paramere plates. The gonarcus is membranous and arches anterior to the parameres, with no mediuncus. The most similar-like genitalia to be found among Brachynemurini in previous studies is in Peruveleon Miller and Stange ( Stange 1970, 1994), but this genus is easily distinguished from Ameromyia by many morphological characters such as leg chaetotaxy, the length of the hind wing CuA vein, and also by molecular data ( Machado et al. 2019).

Two species groups are determined based on taxonomical characters of wing veins, male genitalia, and cladistic results: the ' Ameromyia modesta group’ and ' Ameromyia nigriventris group’ (Fig. 11 ). These two species groups were previously named and identified by Stange (1994), but were classified only by larval characters. The Ameromyia modesta group can now be characterized by the abdomen length, which is longer than wings in females and almost double the length of wings in males, as well as the sickled club of the clavate setae on male genitalia (Fig. 7C ), while the Ameromyia nigriventris group is characterized mostly by the hind wing posterior area cells, which are as high as long or higher than long.

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 1C , 2C , 28A ) and A. protensa (Fig. 39A ). Leg characters are usually informative, and although badly conserved specimens can show misleading color patterns as they can faint or darken over time, leg chaetotaxy is very consistent within species. Genitalic structures are little informative. Ameromyia male specimens have hinged parameres ( Ameromyia sensu Adams 1956) and a highly membranous gonarcus, and thus genitalic structure can be complex to visualize as the sclerites can bend and flex. At dorsal view, all known species bear almost identical male genitalia (Fig. 10A ), and female genitalia presents close to no variation between species, even when across different species groups (Fig. 8 ). The digging setae on female terminalia can present some slight variation on thickness and/or length across specimens, but this is probably a regional adaptation for the granulometry of available sand pools across different habitats. Due to this great genitalic similarity between species, female terminalia and genitalia are only described in each description section when that species bear some morphological variation, and male genitalia is illustrated only in lateral view, in folding position, for each species section, as the lateral view provides more information as to the shape of the parameres ( Adams 1956).

The larvae of most Ameromyia species are currently unknown, except for A. clepsydra sp. nov., A. guarica , A. modesta and A. nigriventris . Stange (2004) described the presence of thread-like setae on dorsal segments (Fig. 14A-C ) as a character for differentiating the two groups, as it is present in A. modesta but described as absent in A. nigriventris . However, it is present in all known Ameromyia larvae.

Biology.

Very few data have been made available regarding Ameromyia biology and/or behaviour. Stange (1994) stated the known larvae lives on sand dunes, and later (2004) stated the adults rests on grass stems during the day. In this study, we have observed and/or reared in captivity adults from four species ( A. clepsydra sp. nov.; A. explicata sp. nov.; A Ameromyia nigriventris and A. strigosa ), and larvae from two species ( A. clepsydra sp. nov. and A Ameromyia nigriventris ). Adults from all species observed except for A. clepsydra sp. nov. were capable of flying high, at at least 2 m above ground, and when in captivity, they almost always perched at the top of the rearing enclosure. In their natural habitat, A. nigriventris females were seen flying close to the ground, presumably scouting for good ovipositioning spots, while all observed males which were near the ground were found slowly hovering in place, just above patches of loose earth which were apparently suitable for larvae. It is unknown if this is somehow a courting behaviour. Regardless of sex, when disturbed, specimens would quickly try to outmanouver their catcher, then rapidly ascend in the air and gain speed with the wind. Wild specimens of A. clepsydra sp. nov. were always observed flying very low, close to the bedrock, and no such ascending behaviour could be seen.

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 ( Stange 1994). Of the larvae reared in the present work, that of A. clepsydra sp. nov. (Fig. 15B, C ), lives in the same microhabitat as described for A. guarica : shallow, water-deposited sand on top of a river bedrock (Fig. 15A ). The A. nigriventris larvae collected during this work were found on patches of loose, sandy wind-deposited earth on top of compacted clay soil (Fig. 16A ). Despite the habitat differences, there is much similarity to the analyzed larvae microhabitat. Ameromia nigriventris larvae stays submerged in the loose, sandy soil, while using the compacted clay as a hard substrate which they anchor themselves to (Fig. 16B, C ), and A. clespydra sp. nov. anchor themselves to the bedrock itself or the shallow compacted sand on top of the same bedrock (Fig. 15B, C ). Almost always, larvae were found on small “trenches”, presumably dug out by the larvae themselves (Figs 15B, C , 16B-D ). All known Ameromyia larvae bear many long bristles on thoraxic and abdominal setiferous processes, which would help in escavating suitable holes in hardened substrate. When exposed, the larvae would slightly contract themselves into their holes, and remain still for a while. Then, they would try to dig backwards, or leave their spot and walk forwards, towards a new hiding place. When moving forwards, regardless of instar, A. nigriventris larvae are very quick and highly agile, while A. clepsydra sp. nov. are very slow and lethargic, sometimes refusing to move even if poked or disturbed. Coccoons (empty or with pupae still inside) in the wild were found in the same said “trenches” (Fig. 16D ), or protected by bedrock (Fig. 15D ). When in captivity, Ameromyia larvae were highly sensitive about the presence or absence of both the hard substrate to achor themselves to, and to said “trenches”. Larvae from both reared species would easily startle and hardly feed in the absence of a similar hard substrate, and A. clepsydra sp. nov. larvae in the late third instar would not spin a coccoon nor pupate, and instead walk incessantly until it found an appropriate substrate or until death. To simulate the hard achorage of their natural habitat, plaster or styrofoam was "dug out" and then fit into the larvae plastic enclosures, which highly decreased larvae mortality. This highly suggests Ameromyia species are much more sensible to microhabitat selection and alterations, as other larvae collected from the same sites and in similar microhabitats (such as Austroleon immitis and Dimarella riparia ) were able to feed, develop and pupate normally even in the absence of these specific microhabitat conditions. Although larvae from other species were not found during this work, as the morphology and chaetotaxy of female terminalia is consistently similar throughout the genus, larvae habitat might also be similar, as the characteristics of female terminalia affects oviposition and the selection of larvae habitat ( Stange and Miller 1990).

Figures

Figure 1. 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 Jose Correia de Santana. E A. clepsydra sp. nov., Brazil. F A. explicata sp. nov., Brazil. Latter photographs by Leon Tavares.

Figure 2. Ameromyia specimen views: A A. strigosa, head and forelegs, frontal view. B A. guarica and C A. modesta, head and thorax, dorsal view.

Figure 7. Ameromyia male terminalia and genitalia chaetotaxy: A A. protensa lectotype male terminalia, lateral view, photograph by Lara Lopardo (EMAU). B A. explicata sp. nov., globose club of genitalic sac clavate setae. C A. guarica, same but club sickled. D A. clepsydra sp. nov., same, but club subglobose. - Abbreviations: et, ectoproct; etl, ectoproct postventral lobe.

Figure 8. Ameromyia female terminalia: A A. protensa, lateral view. B A. protensa, posterior view. C A. nigriventris, ventral view. Photographs by Lara Lopardo (EMAU). - Abbreviations: et, ectoproct; ds, digging setae; gp, gonapophyseal plate; lg, lateral gonapophysis; pg, posterior gonapophysis; pp, pregenital plate; st, sternite, tg, tergite.

Figure 9. 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.

Figure 10. 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.

Figure 11. 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 Ameromyia sensu Ameromyia novo. B Alternative tree.

Figure 12. South America map showing the current distributional records for Ameromyia species in the Ameromyia nigriventris species group.

Figure 13. South America map showing the current distributional records for Ameromyia species in the Ameromyia modesta species group.

Figure 14. 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.

Figure 15. Collection site and habitat of Ameromyia clepsydra sp. nov., in Andarai, 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.

Figure 16. 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.

Figure 19. Ameromyia clepsydra sp. nov., third instar larvae: A dorsal view. B Ventral view.

Figure 26. Ameromyia guarica, preserved in alcohol, third instar larval specimen: A Dorsal view. B Ventral view. C Lateral view. D Dorsal view, dried up. E Lateral view of head and thorax, dried up. Photographs by Jonathan Bremer (FSCA). - Abbreviations: ts, thread-like setae; tu, tubercle.

Figure 27. Larval specimen of an unknown species of Venezueleon (Ameromyia sensu Stange): A dorsal view. B Ventral view. C Lateral view. Photographs by Jonathan Bremer (FSCA).

Figure 28. Ameromyia modesta holotype: A dorsal view. B Lateral view. C Type labels. Photographs by Charles Farnum, Museum of Comparative Zoology, Harvard University.

Figure 30. Ameromyia modesta, third instar larva: A dorsal view. B Ventral view. C Lateral view.

Figure 36. Ameromyia nigriventris, third instar larvae, preserved in alcohol: A dorsal view, B ventral view and C lateral view, photographed by Jonathan Bremer (FSCA). D head capsule dorsal view, dried specimen. - Abbreviation: ts, thread-like setae.

Figure 39. 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 (EMAU). - 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.