Twenty-four new species of Aleiodes Wesmael from the eastern Andes of Ecuador with associated biological information (Hymenoptera, Braconidae, Rogadinae)

Abstract Aleiodes Wesmael is the most diverse rogadine genus worldwide, with specialized koinobiont endoparasitic development in Lepidoptera caterpillars resulting in mummification of the host remains. This paper focuses on describing new Aleiodes species from the Yanayacu Biological Station, with special interest in those with biological information. We describe 24 new species (Aleiodes albidactyl sp. n., Aleiodes albigena sp. n., Aleiodes albiviria sp. n., Aleiodes bimaculatus sp. n., Aleiodes cacuangoi sp. n., Aleiodes colberti sp. n., Aleiodes delicatus sp. n., Aleiodes dyeri sp. n., Aleiodes elleni sp. n., Aleiodes falloni sp. n., Aleiodes frosti sp. n., Aleiodes kingmani sp. n., Aleiodes longikeros sp. n., Aleiodes luteosicarius sp. n., Aleiodes marilynae sp. n., Aleiodes mirandae sp. n., Aleiodes napo sp. n., Aleiodes nubicola sp. n., Aleiodes onyx sp. n., Aleiodes shakirae sp. n., Aleiodes stewarti sp. n., Aleiodes townsendi sp. n., Aleiodes tzantza sp. n., and Aleiodes yanayacu sp. n.) from Napo Province in Ecuador, 16 of which were reared from host caterpillars. With these results 89 species of Neotropical Aleiodes are now known, with 41 of them having host records. The most commonly reared species were in the circumscriptus/gastritor species-group, and mostly associated with Geometridae hosts (six of ten species). Three species of seriatus species-group, in contrast, were each reared from a different family. One of these species (i.e. A. frosti sp. n.), reared from Notodontidae, cuts a posterior radial opening in the mummy for emergence, a unique behavior in Aleiodes, recorded here for the first time. A. luteosicarius sp. n. is the first described species from Ecuador in the pallidator species-group. Differing from previously described pallidator species, which attack only Lymantriinae larvae, A. luteosicarius sp. n. attacks several species of Arctiinae larvae, being both subfamilies within Erebidae with densely setose caterpillars. We also describe new species of the gressitti and pulchripes species-groups.


Introduction
Aleiodes is the most common and speciose rogadine braconid genus worldwide. Species richness estimations have changed dramatically in last decade. In the Taxapad catalog (Yu et al. 2012) 431 described species are recorded, but Butcher et al. (2012), in a single work published after that catalog, described 179 new species from Thailand. Those authors estimated the Aleiodes fauna of Thailand alone to include more than 400 species, pushing the diversity of Aleiodes to a much higher level than previously estimated, especially for the tropical fauna, previously considered not as rich as the Holarctic (S. Shaw 2006). Quicke (2012) discussed evidence suggesting a greatly unknown tropical Ichneumonoidea fauna, where the lack of studies on small body sized groups and highly speciose genera are the main reasons for the underestimation. For Neotropical Aleiodes, Delfin-Gonzalez and Wharton (2002) estimated about 200 undescribed species, but in light of these recent works, this number is also likely to be much higher (in addition, for example, there are at least 160 undescribed species from Brazil. EMS, previously unpublished data). Prior to this study, there are 65 described species of Aleiodes in Neotropical region, 25 of them with host records (Fortier 2009, Townsend and Shaw 2009, Shimbori and Penteado-Dias 2011.
The first division of Aleiodes into species-groups (S. ) accounts for fifteen different groups. After phylogenetic analyses three additional groups were proposed (Fortier and Shaw 1999). Townsend and Shaw (2009) found the species of the closely-related gastritor and circumscriptus groups in Ecuador to be difficult to separate, and recommended treating these as one single group for Neotropical fauna. Although the existing species-group system presents some limitations when applied to Neotropical fauna (Townsend and Shaw 2009), it provides a working framework to investigate this diverse group and to improve our taxonomic knowledge. Most species-groups are represented in Neotropical Region, except for the compressor, praetor, procerus, rugulosus, ufei and unipunctator groups (S. , S. Shaw et al. 1998a, Marsh and S. Shaw 1998, S. Shaw et al. 1998b, Marsh and S. Shaw 1999, Marsh and S. Shaw 2001, Marsh and S. Shaw 2003, S. Shaw et al. 2006, S. Shaw et al. 2013, being the pallidator species-group represented only by undescribed species in Neotropical Region. Based on undescribed material from Neotropical collections, the seriatus species-group is the most diverse group in this region, followed by circumscriptus/gastritor species-group. Rogadinae is a cosmopolitan subfamily of koinobiont endoparasitoids of Lepidoptera (M. Shaw 1983, M. Shaw and Huddleston 1991, M. Shaw 1994, S. Shaw 1997, currently divided in five tribes: Stiriopini, Clinocentrini, Yeliconinae, Rogadini and Aleiodini (=Aleiodes+Heterogamus) (van Achterberg 1995, Zaldívar-Riverón et al. 2008, the later containing Aleiodes, the most speciose genus with about 612 described species of the current 1,141 named Rogadinae species (Yu et al. 2012, Butcher et al. 2012, S. Shaw et al. 2013. All Rogadinae induce the hardening of the host larva before pupation, producing the so-called "mummy," with the mummification of the host larva been considered the only biological synapomorphy of this subfamily (van Achterberg 1995). Aleiodes is known to attack almost exclusively exposed-feeding macrolepidepterans, especially the superfamilies Noctuoidea and Geometroidea, and to a lesser extent, Sphingoidea and Papilionoidea (S. , S. Shaw 2006. A few exceptions include some exposed-feeding microlepidopterans (e.g. Zygaenidae) and, in rare cases, such as Aleiodes compressor (Herrich-Schäffer, 1838), macrolepidopterans with semi-concealed habit (M. Shaw 2002). Aleiodes commonly attacks second and third instar caterpillars, but first instar records are known, though less successful (M. Shaw 1983). The host is mummified in later instars, and within the hardened skin of the host the parasitoid pupates and eventually emerges through a hole on posterior region of the mummy.
The present work presents descriptions of 24 Aleiodes new species, 16 of which with information on biology and photographs of the mummified host larvae. These comprise mostly exposed-feeding host records, with the parasitoids attacking hosts living in small trees and bushes, rather than herbaceous plants near ground level, or higher in the canopy.

Methods
Specimens for this study were collected during the Caterpillars and Parasitoids of the Eastern Andes of Ecuador project (Miller and Dyer 2009), and reared at the Yanayacu Biological Station and Center for Creative Studies (YBS). The YBS is situated on the northeastern slope of the Andes in Napo Province, Ecuador, the watershed streams at the YBS flow to the Amazon basin through the Río Napo, a major tributary of the Amazon River (S. Shaw 2012). The reserve comprises both primary-and secondary-growth montane forests (Miller and Dyer 2009). The plant community at Yanayacu is very diverse and complex, comprising species from at least 76 families (Rab Green et al. 2011).
Specimens were sampled using varied methods including yellow pan trap, Malaise traps situated on the ground and suspended in the canopy, Maxi net, hand collected with aspirators at a light sheet, ultraviolet light trap (= U.V. light trap or black light trap), and during daylight off vegetation with net or vials. Most of the specimens were reared from caterpillars (Greeney 2007), from 2007 to 2013. Caterpillars were sampled by walking through various habitats using two different methods: by hand collecting after inspecting herbs, shrubs and trees up to a height of approximately 2.5 m; or beating plants over a white cloth square of 1×1 m size. Caterpillars were collected in clear plastic bags with their food plant, assigned identification codes, and transported to the rearing shed at YBS. Reared parasitoid specimens are associated with the voucher number of the respective host larva (e.g. YY-00000). Caterpillars and host plants were identified and recorded. Rearing took place in plastic bags in an open-air shelter with ambient temperatures and natural day length. Frass and decaying plant material were removed every other day and new plant material was provided as necessary. While cleaning out the bags, the caterpillars were inspected to note the date of caterpillar pupation or date of parasitoid pupation. Parasitoid pupae were inspected daily for emergence. All emerging adult parasitoids were kept with the original code given to the caterpillar to preserve host data. The parasitoids were preserved in alcohol and transferred to the University of Wyoming where they were dried and point mounted for identification.
Type specimens are deposited at University of Wyoming Insect Museum, University of Wyoming, Laramie, Wyoming, U.S.A. (UWIM). Additional specimens from Canadian National Collection, Ottawa, Canada (CNC) were added whenever appropriate to the scope of this work.
All the species proposed here are satisfactorily distinguished by morphological features. Nevertheless, for some of species, a comparison of ribosomal DNA sequences in gene COI by Donald Quicke was helpful in the definition of species limits.
Because of the supporting grant funding, this work focused on species sampled at the YBS. Some paratypes outside the boundaries of this refuge were included, but only if at least one type specimen is from YBS. All specimens included in this project are from the eastern face of the Andes Mountains, in Napo Province. The altitude of sampling sites ranged from 2,000 to 2,800 meters. In previous work on Aleiodes from Napo Province by Townsend and Shaw (2009) the altitudinal range was from 1,383 to 2,837m. Most of additional material is also from Napo province, with altitude ranging from 1,700 to 2,000 meters; only two type specimens are from lowland Manabí Province, at 400 meters elevation.
For recognition of subfamily Rogadinae see van Achterberg (1993) and Sharkey (1997). The definition of Aleiodes follows that of van Achterberg (1991) and S. Shaw (1993). Species-group definitions follow S. , Fortier and S. Shaw (1999) and Townsend and Shaw (2009). Terminology follows Sharkey and Wharton (1997) and S. . Microsculpture terminology follows that of Harris (1979). Wing vein terminology follows the system adopted by Sharkey and Wharton (1997).
We present descriptions for 24 new species from Northeastern Andes in Ecuador. Along with the description of the new species, we provide summarized taxonomic information on the nine previously described species from the same region (Townsend and Shaw 2009), and also a set of additional characters, not discussed in the original description. New biological information for these species is presented whenever available. Images were captured with a 3MP Leica video camera and a Leica M205C stereomicroscope running Leica Application Suite (LAS) software, and focus-stacked using the same software. Some minor adjustments in images and plate preparation were performed in Adobe Photoshop version CS6. Pictures of caterpillars and host mummies before parasitoid emergence were taken, at the rearing site, by Wilmer Rosendo Simbaña, José Arturo Simbaña and Luis Alberto Salagaje. Ocell-ocular distance about equal to diameter of lateral ocellus ( Figure 19); metasomal tergum 1 at most 1.4× longer than its apical width; species with known biology with normal emergence behavior, cutting a postero-dorsal exit hole in the mummified host caterpillar (Figures 40, 57)  Ocelli larger, ocell-ocular distance 1/2 diameter of lateral ocellus; metasomal tergum 1 unusually long and narrow, more than 2× longer than its apical width ( Figure 47); species with a unique emergence behavior, cutting the whole posterior tip away from the mummified host caterpillar (Figure 51 Figure 7); fore wing vein 1M almost straight or weakly and evenly curved (as in Figure 119)

Aleiodes aclydis Townsend, 2009
http://species-id.net/wiki/Aleiodes_aclydis Figure 1 Diagnosis. Body length 6.1 mm; antenna with 44 segments; head with vertex black, occiput light orangish brown; ocelli large, ocell-ocular distance less than width of lateral ocellus; occipital carina interrupted at vertex; mesosoma mostly light orangish brown, except propodeum black; wings slightly darkened; mesopleuron granulate; apex of hind tibia without comb of modified setae; propodeum without median propodeal carina; metasomal terga entirely black; metasomal tergum 3 costate on anterior 2/3, with median carina along with this sculpturing; ovipositor short, about 0.25× length of hind basitarsus. Additional characters. Last flagellomere with "bottle nipple"-like tip; mesoscutum with carina only in front of scutellar sulcus; scutellar sulcus with complete median carina plus two pairs of weak and incomplete lateral carina; fore wing vein 1M only slightly curved at base; hind wing vein 2-1A absent, vein m-cu present and well pigmented, antefurcal to r-m in left wing and interstitial in the right wing; ovipositor sheaths about as long as hind tarsomere II, 0.6× hind basitarsus.
Distribution. Known only from the type locality, Isla de Las Palmas, Napo province, ECUADOR, at 1,883 meters elevation.
Discussion. From the newly described species, Aleiodes aclydis is very similar to A. albigena sp. n. in color features, including a mostly blackish head with a lighter gena. However, in aclydis the gena is yellow instead of the white gena of albigena sp. n. Both species belong to circumscriptus/gastritor species-group but aclydis is the unique species with a malar space as short as 0.7 times the basal width of mandi-Figures 1-9. 1-3 head, frontal. 1 Aleiodes aclydis Townsend 2 Aleiodes capillosus Townsend; Aleiodes marilynae sp. n. 4-6 Aleiodes albidactyl sp. n. 4 habitus 5 fore wing, detail of 1 st subdiscal and subbasal cells 6 metasoma, dorsal 7-9 Aleiodes albigena sp. n. 7 habitus 8 host larva, nr. Desmotricha (Erebidae) 9 host mummy after parasitoid emergence. bles. No additional A. aclydis specimens were found since Townsend and Shaw's (2009) work. The dorso-medially elevated area on mesopleuron is well-demarcated posteriolly. In the original description the sulcus demarcating this region is called the "sternaulus" (Townsend and Shaw 2009). We now consider that this sulcus is not a true sternaulus, as defined by Sharkey and Wharton 1997, and neither is it the precoxal sulcus, as defined by other authors (van Achterberg 1991, Wharton 2006. Additionaly, the absence of sternaulus is a common feature in all species treated in this work. Therefore, this term is avoided in the descriptions of the new species. The same matter is found in the descriptions of Aleiodes atripileatus Townsend and Aleiodes nebulosus Townsend.
Color. Distinctive black and white color pattern. Head mostly pale yellow with small dark brown spot just above clypeus; occiput laterally and stemmaticum dark brown. Antenna dark brown, scape and pedicel brown. Mesosoma mostly black; mesoscutum mostly pale yellow with postero-lateral borders dark brown; metapleuron, propodeum posteriorly based triangular area, thumb shaped area posteriorly on mesopleuron, ventral half of pronotum, and propleuron white. Metasoma mostly black dorsally and completely white ventrally; T1 apical 1/3 and small mid-basal spot white; T2 mostly white with two round antero-lateral spots; T3 with white semicircular basal spot; lateral borders of remainder terga white; ovipositor sheaths basal 1/5 white, remainder dark brown. Fore and mid legs pale light brown. Hind legs brown; basal half of femur and coxa, trochanter and trochantellus white. Wings very weakly infuscate, veins dark brown.
Head. Antenna 50 antennomeres, apical flagellomere with "bottle-nipple"-shaped apex; malar space slightly longer than basal width of mandible, 0.4× eye height; in dorsal view eyes 2.75× longer than temples; occipital carina incomplete, directed toward vertex, getting close to lateral ocelli, well defined laterally and meeting hypostomal carina; oral space small and circular, maximum width slightly smaller than basal width of mandible; clypeus not swollen; ocell-ocular distance as long as diameter of lateral ocellus; maxillary palp not swollen; head surface sculpture shining granulate, occiput smooth and shining; vertex with few wrinkles; higher face with few transverse rugosity; frons smooth with pairs of wrinkles concentric to each toruli, frons excavated with excavation bordered by a "W-shaped" carina.
Discussion. This species belongs to seriatus species-group. It differs from all other species of this group by the white posterior thumb like marking on the black mesopleuron, and the long vein 1CUa on fore wing, about 1.5× longer than 1CUb and 3.5× vein 1cu-a. This species is most closely related to dyeri sp. n., but the color patterns of both species are quite distinct (see comments for dyeri sp. n.).
Etymology. From the Latin roots meaning "white finger", a reference to the white thumb like mark on mesopleuron.
Color. Mostly black. Head and antenna dark brown; gena, mandibles and palp white, mandible tips dark brown. Pronotum, propleuron, mesoscutum, most of mesopleuron and scutellum honey yellow; metanotum, propodeum, metapleuron and dor-sal 1/4 of mesopleuron, including the border with metapleuron, black; mesopleuron with whitish longitudinal stripe. Legs: all coxa, trochanter and trochantellus, mid femur and tibia, and most hind femur white; fore femur yellowish; fore tibia and tarsi and mid tarsi light brown; hind tibia and tarsi dark brow to black, but tibia with white basal band and fourth and fifth tarsi lighter; hind coxa, trochanter, trochantellus and femur black dorsally, except basal 1/5 of femur; tip of fore and mid femur and mid tibia with infuscate stains. Metasoma dark brown to black dorsally, laterally and ventrally white, but some lateral spots and the last sternites dark brown.
Mesosoma. Sculpturing mostly granulate; pronotum with few laterally running wrinkles laterally; mesopleuron rugose on dorso-anterior corner; propodeum rugosegranulate, with complete mid-longitudinal carina; notauli very shallowly indicated anteriorly, posteriorly disappearing in a depressed rugose area; posterior margin of mesoscutum with very short carina, just anterior to scutellar sulcus; scutellar sulcus with five short carina, not reaching anterior margin of sulcus.
Legs. Hind tibia without comb of modified setae; tarsal claw simple, with a comb of thin bristles medially; hind basitarsus 2.5× length of inner apical spur of hind tibia.
Female. Unknown. Mummy. Length 9.0 mm, body reddish brown, head yellowish, setae pale brown, covered with setae, thorax compact and wrinkled, glue hole located ventrally on the thorax, exit hole irregular, located postero-dorsally, posterior to hind abdominal prolegs.
Biology. Reared from a species near Desmotricha Hampson (Erebidae) larvae (voucher number YY-47082), feeding on Chusquea scandens (Poaceae). Parasitoid took three weeks to emerge after host mummification. This is the only described circumscriptus/gastritor-group species in Neotropical region known to attack Arctiinae hosts, producing a densely setose mummy.
Discussion. Aleiodes albigena sp. n. belongs to circumscriptus/gastritor speciesgroup. Some diagnostic characters are the very shallow notauli present only anteriorly; scutellar sulcus without bisecting carina; occipital carina complete dorsally; face bulging; and frons surface granular, not excavated, without a lateral carina. This species is similar to townsendi sp. n. and shakirae sp. n., but it can be distinguished from both by the mostly black head with contrasting whitish gena, yellowish in townsendi sp. n. and shakirae sp. n., and the almost straight vein 1M on fore wing, strongly curved in townsendi sp. n. and shakirae sp. n. A. albigena sp. n. resembles A. arbitrium in color pattern but differs in the shorter ocell-ocular distance relative to lateral ocelli diameter.
Etymology. From the Latin roots meaning "white cheeks," named in reference to the contrasting white gena, as compared with the black head of this species.
Additional characters. Last flagellomere lanceolate; mesoscutum with complete carina on posterior margin though not well defined; scutellar sulcus with a strong complete median carina and some irregular carina laterally; fore wing vein 1M slightly and evenly curved; hind wing vein 2-1A present as a very short stub, vein m-cu present and distinctly postfurcal to vein r-m; ovipositor sheaths 1.3× longer than hind basitarsus.
Distribution. Known only from the type locality, Rio Chalpi Grande, Napo province, ECUADOR, at 2,837 meters elevation.
Discussion. For standardization reasons we provide the measurement of the ovipositor sheaths instead of the ovipositor itself. The ovipositor and the sheaths length in A. albiterminus are unusually long for Aleiodes species. This is the only species from Ecuadorian Northeastern Andes with the ovipositor sheaths longer than its hind basitarsus. A. albiterminus can be distinguished also by the absence of a median carina on the first to third metasomal terga, and the distinctive off-white markings at the apex of metasomal tergum 3. Aleiodes albiviria sp. n. http://zoobank.org/099A4E94-7013-4B08-930B-1DB134C7EB3E http://species-id.net/wiki/Aleiodes_albiviria Figures 18-21 Description of holotype. Female (holotype). Body length 7.1 mm; antenna length 9.0 mm; fore wing length 5.8 mm.
Head. Antenna with 57 antennomeres, mid flagellomere roughly 2.0× longer than wide, apical flagellomere with small pointed apex; malar space moderate, about 1.25× longer than basal width of mandible, 0.33× eye height; in dorsal view eyes 3× longer than temples; occipital carina incomplete, directed toward vertex, well defined laterally and meeting hypostomal carina; oral space small and circular, maximum width equal to basal width of mandible; clypeus not swollen; ocelli moderate, ocell-ocular distance slightly shorter than diameter of lateral ocellus; maxillary palp not swollen; head surface sculpture finely shining granulate, occiput smooth and shining; higher face with a small longitudinal ridge and transverse rugosity directed to it; frons excavated, excavation bordered by a weak "W-shaped" carina.
Mesosoma. Sculpturing shining granulate; pronotum foveate laterally; mesopleuron with antero-dorsal corner rugose; propodeum granular with some longitudinal diverging wrinkles laterally, mid-longitudinal carina present only on anterior 1/3; notauli with few crenulae and shallow anteriorly, meeting on depressed rugose area posteriorly; posterior margin of mesoscutum with complete carina; scutellar sulcus with median carina plus two pairs of complete lateral carina and one irregular. Wings Legs. Hind tibia with comb of modified setae; tarsal claw simple, with a comb of relatively long thin setae basally; hind basitarsus 3.5× longer than inner apical spur of hind tibia.
Metasoma. T1, T2 and basal 2/5 of T3 costate, longitudinal carina present along this sculpturing; remainder visible terga smooth; ovipositor sheaths short and lanceolate, about as long as hind tarsomere IV (half length of tarsomere II); T1 1.4× longer than its apical width. Discussion. This species is assigned to the seriatus species-group, where it most resembles Aleiodes greeneyi Townsend because of the dorsally incomplete occipital carina. Aleiodes albiviria sp. n. differs from other New World species of this species group by the mostly honey brown body with dark brown notauli and mid-posterior mesoscutum, and the white middle band on blackish antenna. A. albiviria sp. n. also resembles the Brazilian Aleiodes scriptus (Enderlein, 1920), by the costate sculpturing on metasomal tergite 1, which is rugose-costate in all other Neotropical species in seriatus species-group, but differs from scriptus by the shape of hind wing vein RS (parallel to wing margin basally and bent downward apically, as opposed to sinuate at middle in scriptus).
Etymology. From the Latin albus, meaning "white," and viria meaning "bracelet," a reference to the white band on the antenna. Townsend, 2009 http://species-id.net/wiki/Aleiodes_arbitrium Figures 11, 12 Diagnosis. Body length 5.7 mm; antenna with 43 segments; head mostly black; malar space slightly wider than basal width of mandible; ocelli small, ocell-ocular distance slightly wider than width of lateral ocellus; occipital carina at least shortly interrupted at vertex; mesosoma mostly light yellowish brown, except propodeum black; wings clear; mesopleuron granulate; apex of hind tibia without comb of flattened setae; propodeum rugulose basally, granulate apically, with median propodeal carina present; metasomal terga mostly black, except terga 2 and 3 often each with an oval-shaped off white marking; metasomal terga 1-3 costate, median carina complete to end of tergum 2; ovipositor sheaths slightly shorter than half of hind basitarsus.

Aleiodes arbitrium
Additional characters. Last flagellomere with short pointed tip; mesoscutum with carina on posterior margin only in front of scutellar sulcus; scutellar sulcus with a median carina barely complete plus two or three more or less weak and incomplete carina laterally; fore wing vein 1M slightly and evenly curved; hind wing vein 2-1A present as a very short stub, vein m-cu position varying from distinctly antefurcal to interstitial to vein r-m; ovipositor sheaths 0.45 to 0.49 times longer than hind basitarsus.
Morphological variation. All specimens examined agree well with original description of Aleiodes arbitrium, however the occipital carina in all specimens is at least shortly interrupted at vertex, and not complete but weak at vertex as described originally. The two spots on terga vary in size. In some females it is darker, not so contrasting, while in others it is larger, with the T2 spot reaching apical margin of tergite. In some males the spots tends to meet and form one single irregular elongate spot. In all specimens the gena is black-dark brown but the borders with mandibles are light brown.
The holotype and eight non-type specimens were reared from Psaliodes sp. larva feeding on D. costale or D. cornuta. The two paratypes and four other non-type specimens were reared from Psaliodes castanea (Warren) feeding on D. costale. A single non-type specimen was reared from unknown Pyralidae larva on D. cornuta. One paratype collected from Urticaceae did not feed, so may have simply wandered there prior to mummification.
Distribution. Known only from the Yanayacu Biological Station, Napo province, ECUADOR, at 2,163 meters elevation.
Discussion. After publication of Townsend and Shaw (2009) paper, 11 new specimens were reared and ten collected by traps, all from Yanayacu Biological Station, Napo Province, S00°35.9', W77°53.4', 2163. This additional reared material provided good biological information at least on feeding preferences of the host caterpillars. It is likely that A. arbitrium attacks mostly Geometridae species feeding on polypod ferns. In the circumscriptus/gastritor group, this species resembles A. albigena sp. n. in most color patterns; however, the smaller ocelli of A. arbitrium and the honey brown marks on vertex, absent in A. albigena, distinguish these two species. Other species with small ocelli and interrupted occipital carina, as A. onyx sp. n. and A. atripileatus, have more extensively dark bodies than arbitrium.
Additional characters. Last flagellomere with short "bottle-nipple"-like tip; mesoscutum posterior margin with carina only in front of scutellar sulcus; scutellar sulcus with a median plus two or three pairs lateral carina well defined and almost complete; fore wing vein 1M only slightly and evenly curved; hind wing vein 2-1A absent, vein m-cu absent; ovipositor sheaths about 2/3 length of hind basitarsus, one specimen in type series have 2.5/3, and another 1.8/3 proportions for this character.
Biology. Aleiodes atripileatus has been reared from a species of Hypena Schrank (Noctuidae) caterpillars feeding on Urticaceae, including Phenax rugosus, Boemeria bullata, Miriocarpa sp., and three other unidentified urticaceous plants. Other noctuids on the same host plants might be utilized as hosts.
Distribution. Known only from the YBS, Napo province, ECUADOR, at 2,163 meters elevation.
Discussion. Aleiodes atripileatus is one of the most commonly reared Aleiodes species in YBS. The species belongs to the circumscriptus/gastritor species-group. It is very similar to A. nubicola sp. n. and A. cacuangoi sp. n., differing from these two species in having the occipital carina distinctly interrupted at vertex. This character is shared with A. onyx sp. n., from which it differs by having a black vertex, a thinner metasoma, and the vein m-cu of hind wing absent. Different than originally described, the true sternaulus is absent.
Color. Mostly honey brown. Antenna brown, scape and pedicel honey brown as head; cheeks and palp light yellow; ocellar triangle brown; fore and mid coxa, and all trochanter and trochantellus whitish; metanotum and propodeum dark brown; metasoma dark brown dorsally with mid-apical pale yellow spots on T1 and T2; ovipositor sheaths brown. Wings hyaline with brown veins and stigma, parastigma contrasting darker-black.
Head. Antenna 44 segmented, apical flagellomere lanceolate, without pointed tip; malar space about 1.3× longer than mandible basal width, and 0.6× eye height; occipital carina complete; oral space moderate and circular, maximum width about equal to basal width of mandible; clypeus swollen; ocelli small, ocell-ocular distance 1.4× diameter of lateral ocellus; in dorsal view temples almost as long as eye height; head sculpturing mostly granular, face coarsely granular to rugose, occiput smooth.
Mesosoma. Sculpturing mostly granulate; pronotum covered with wrinkles; mesopleuron mostly rugose otherwise granulate, wrinkles stronger on sternaulus area; metapleuron with rugosity posteriorly; propodeum rugose, with complete mid-longitudinal carina; notauli deep and mostly smooth anteriorly, with two or three crenulae, meeting on rugose area posteriorly; posterior margin of mesoscutum bordered by short carina just anterior to scutellar sulcus; scutellar sulcus with median carina plus one pair of lateral carina.
Paratype variation. Body length 5.0-6.0 mm; antennomeres 39-46; occipital carina is only very shortly interrupted in some paratypes, but never curved toward vertex; vein 2-1A of hind wing varying from short to absent. The patratypes from outside the YBS are distinctly smaller (body length 5.0-5.3 mm) than the type specimens from the YBS (5.4-6.0 mm), with fewer segments on antenna (39-40 vs. 43-46). The metasoma in this specimens is lighter than the holotype and paratypes from YBS: the apical terga beyond T3 are mostly honey yellow, and the spots on T1 and T2 are frequently larger, forming one large spot covering apical T1 and all T2 medially. We consider the specimens from Baeza as a geographical variant within bimaculatus sp. n. Since all but one females were collected at once and shows virtually none variation, the variation could be just an artifact. Further samplings could both confirm this hypothesis with some intermediate forms or support an alternative hypothesis (e.g. speciation process). The metasoma in one of the females from Manabí is almost entirely honey brown, the light spots are not contrasting but still visible.
Male. Body length 4.7-5.0 mm. Antennomeres 48-42. Considerable color pattern variation in males: antenna dark brown, pedicel brown, scape honey brown, face brown, hind coxa whitish, all tibia and tarsi darker, ocellar triangle black, metanotum dark brown as propodeum, T2 pale yellow spot varying from smaller than in female to covering most of the tergite, T3 also with pale yellow spot. The metasoma is narrower, T1 up to 1.4× longer than its apical width; eyes in dorsal view 1.55× longer than temple; occipital carina weak dorsally, barely interrupted at vertex. The males from Baeza follow the same pattern of the females, with the apical metasomal terga honey brown instead of dark brown, however the body length in these males is not distinctly shorter than the specimens from YBS.  Napo, Baeza, 2000m, February 19793♂, Napo, 5km South Baeza, 1700m, February 9, 1983, Masner &Sharkey;3♂, Napo, Baeza, 1900m, February 9, 19832♀ Manabi, Montecristi, 400m, February 6, 1983, Masner & Sharkey. Discussion. Aleiodes bimaculatus sp. n. belongs to circumscriptus/gastritor species group. It resembles nubicola sp. n. and cacuangoi sp. n. because of the complete occipital carina and the small ocelli. A. bimaculatus sp. n. differs from these species by the honey brown body color, with propodeum and metasoma dark brown and two pale yellow spots on T1 and T2, while nubicola sp. n. and cacuangoi sp. n. are mostly black. The stigma is yellowish in bimaculatus sp. n., but brownish in cacuangoi sp. n. and light brown to whitish in nubicola sp. n., and the body larger, 5.0-6.0 mm length while cacuangoi sp. n. and nubicola sp. n. are shorter, with maximum body length of 4.6 mm.
Etymology. From the Latin roots bi=two and macula=stain, refers to the two distinctive yellow spots on the dark brown metasomal terga of this species.
Head. Antenna comprising 43 antennomeres, flagellomeres roughly 2.0× as long as wide, apical flagellomere with short pointed apex; malar space moderate, length 1.6× basal width of mandible, and approximately 0.55× eye height; occipital carina complete and well defined, bordered by a more or less deep sulcus on temples and vertex, reaching hypostomal carina; oral space small and circular, maximum width slightly smaller than basal width of mandible; clypeus not swollen; ocelli small, ocell-ocular distance about 1.5× diameter of lateral ocellus; temples about 1.8× eye height in dorsal view; maxillary palp not swollen; head surface sculpturing mostly granulate, vertex more coarsely granulate, occiput very weakly shining coriaceous, apparently smooth.
Mesosoma. Sculpturing mostly granulate; pronotum rugose laterally, foveate dorsally; mesopleuron rugose anteriorly, with elevated area centrally smooth and shining, and some wrinkles centrally, otherwise coriaceous; propodeum rugose posteriorly, with long mid-longitudinal carina on anterior 2/3; notauli deep and renulated anteriorly, meeting a coarsely punctate area posteriorly; scutellar sulcus with well defined median carina and two pairs of lateral irregular carina.  Legs. Hind tibia without comb of modified setae; tarsal claw simple, not pectinate, with a comb of relatively long thin setae basally; hind basitarsus about 3.4 times the length of inner apical spur on hind tibia.
Male. Similar to females but body slightly smaller, antenna shorter, with 40-41 segments, metasoma slightly thinner, and ocelli relatively larger: ocell-ocular distance 1.25× diameter of lateral ocellus. In one specimen the occipital carina is shortly interrupted but the occiput and vertex limits are easily distinguishable. One male paratype have considerably darker head and metasoma: head brown to dark brown, gena at borders with mandibles pale yellow, middle face darker, clypeus and most of temples honey brown, white spots on metasoma reduced, on tergite 2 yellowish and covering less than half its length.
Mummy. Length 6.7-8.0 mm in males and 7.3-10.0 mm in females, mottled gray and yellowish, with lateral bands more or less defined mottled dark brown and brown, thorax compact and wrinkled, exit hole irregular, located postero-dorsally between anal and abdominal prolegs.
Discussion. This species belongs to gastritor/circumscriptus species-group. It is similar to A. atripileatus, differing from it by having the occipital carina complete (interrupted at vertex in atripileatus), smooth area on mesopleuron (granulate-coriaceous in atripileatus), 43 antenomeres (maximum 39 in atripileatus), longitudinal carina complete on T1-T3 (incomplete in atripileatus), and one basal light spot on T1 and one on T2 (absent in atripileatus). This species is also very similar to A. nubicola sp. n. The most evident character to distinguish these species being the color patterns on meso-and metasoma. A. cacuangoi sp. n. have a mostly black metasoma dorsally, with one small to tiny basal white spot on T1 and a finger like mid-basal white spot on T2, the size of the spot on T2 varies from 1/3 to 2/3 of the tergite length. In nubicola sp. n. the metasoma varies from entirely black to black with apical whitish spots, these spots are larger on apical terga and extends to T4 in females, but one male paratype has the spots extending from the apical terga throughout apex of T1. There are not apical spots in none of the terga in cacuangoi sp. n., while in nubicola sp. n. there is not basal white spot on T1. The mesosoma in cacuangoi sp. n. is entirely black except for some reddish brown-brown stripe on mesopleuron, which is distinctly lighter close to mid coxal insertion. A. nubicola sp. n. have similar color pattern but with a postero-median square on mesoscutum and scutellum medially orangish. In cacuangoi sp. n. the head is mostly yellowish with occiput and vertex medially always black, the face varies from pale yellow to dark brown, and temples and gena are pale yellow to brownish orange. In nubicola sp. n. the head is mostly dark brown with a crescent moon-shaped honey yellow area on temples, bordering eyes, and the color of gena is variable. Mummies of atripileatus, cacuangoi sp. n., and nubicola sp. n. are very distinctive: cacuangoi sp. n. mummies are mottled gray and yellowish with more or less defined dark brown lateral stripes, atripileatus mummies are black with extended anal prolegs and head yellow, and nubicola sp. n. mummies are entirely brown and gradually narrowing anteriorly.
Etymology. The species is named in honor to Dolores Cacuango, for her pioneering, outstanding brave efforts for the indigenous rights in Ecuador.
Additional characters. Last flagellomere lanceolate; mesoscutum with carina on posterior margin varying from present only in front of scutellar sulcus to completely absent; scutellar sulcus with one to five incomplete carina; fore wing vein 1M slightly to moderately curved at base; hind wing vein 2-1A present and relatively long, vein m-cu present and pigmented but never tubular, interstitial to vein r-m; ovipositor sheaths about 2/3 as long as hind basitarsus.
Morphological variation. The non-type material fits well in original description, but most of the mummies are considerably lighter.
Discussion. Aleiodes capillosus is very similar to A. marilynae sp. n. (for a discussion on distinguishing characters see discussion section under A. marilynae sp. n.). These are the only two species in the gressitti species-group from Neotropical Region. As well as the morphological similarities in the adult parasitoids, the host mummies of A. capilosus and A. marilynae are very similar. Both produce a relatively swollen mummy with a strongly shrunken thorax, where it is attached to a leaf or branch in a distinct angle.
Female unknown.  (UWIM) Discussion. Aleiodes colberti sp. n. belongs to the pulchripes species-group. This species can be distinguished from other New World species of this group by the infuscate band on the forewing just below the pterostigma, and the white middle band on antenna. Despite these characters, it is similar to Aleiodes earinos Shaw, 1997, differing from which in its mostly black head and some dark stains on the mesoscutum and scutellum, as compared with the unicolored body in earinos. The pattern of wing veins resembles Aleidoes arizonensis Marsh & Shaw, 1997, mainly by the very long second submarginal cell. The infuscation below the stigma is also present in the Cuban and Costa Rican species Aleiodes pedalis Cresson, 1869, but pedalis also has distinct infumation apically, not present in colberti sp. n., as well as the apical hind tibia black, as opposed to reddish in colberti sp. n. In the key to New World pulchripes species (S. , colberti does not run easily to any of the described species. Considering the body color it will be forcibly run to Aleiodes notozophus Marsh & Shaw, 1997, but differs from that species by the large gap between apical claw and basal pectination, which is absent in notozophus. The male of colberti sp. n. has small setose pits on terga 4-6 (4-7 in notozophus). Disregarding the presence of the pits it will run to Aleiodes vaughani Muesebeck, 1960, but the ocelli are larger in the new species.
Etymology. This species is named after Stephen Tyrone Colbert, an American comedian, political satirist, writer, actor, and host of The Colbert Report.
Color. Mostly dark brown; legs pale brown but fore and mid coxae and trochanter whitish, hind coxa mostly dark brown, whitish on basal 1/4; all palp whitish; scape and pedicel, notauli and ventral border of mesoscutum, and mandibles yellowish; scutellum brown mid-anteriorly; propleuron pale brown, anteriorly darker; mesopleuron pale brown laterally, ventrally whitish with a roughly defined inverted "heart-shaped" honey brown infuscation; metasomal terga 3 and 4 slightly lighter than remainder metasoma and with whitish lateral borders; metasoma ventrally white except for last two sternites mostly light brown. Wing veins brown, but basally whitish, stigma pale brown, tegula white.
Head. Antenna with 45 segments, flagellomeres roughly 2.0× as long as wide, apical flagellomere with small pointed apex; malar space wide, about 1.6× basal width of mandible, 0.6× eye height; in dorsal view eyes 2.4× longer than temples; occipital carina complete dorsally, well defined laterally and meeting hypostomal carina; oral space small and circular, maximum width about as long as basal width of mandible; clypeus not swollen; ocelli small, ocell-ocular distance 2× diameter of lateral ocellus; maxillary palp not swollen; head surface sculpture coarse granulate, occiput smooth and shining, frons not excavated, higher face with well defined mid-longitudinal ridge.
Wings.  Marsh & Shaw, 1998, from Florida -USA, by the very small ocelli and the wing venation, remarkably the short second submarginal cell of the forewing, and also the longitudinal carina incomplete on propodeum. A. delicatus sp. n. is mostly dark brown with some lighter regions on mesoscutum and mesopleuron (legs also lighter and metasoma ventrally white), being the color pattern very similar to pectoralis, while akidnus is entirely honey yellow with black stemmaticum; however, the gena of delicatus sp. n. is dark brown, compared to light yellowish in pectoralis. Though short, the second submarginal cell is not almost square as in akidnus and pectoralis. The vein r is 0.75× vein 3RSa in delicatus sp. n., but in akidnus and pectoralis r is slightly longer than 3RSa. In delicatus sp. n. the longitudinal carina is absent on metasomal tergite 3, but it is complete in akidnus and present on basal half in pectoralis, and the first metasomal tergite is 1.6× longer than apical width but roughly as long as wide in akidnus and about 1.3× in pectoralis.
Etymology. From the Latin, meaning delicate.
Color. Head honey yellow, ocellar triangle black; antenna dark brown, scape slightly lighter. Mesosoma honey yellow, some lighter parts on metapleuron and dorsal mesopleuron. Fore leg honey yellow, tarsi slightly darker but 5 th tarsomere brown. Mid leg with same pattern of fore leg, but all tarsi brown; coxa, trochanter and trochantellus whitish, coxa with dark lateral stains. Hind leg: coxa mostly dark brown, basal third pale yellow; trochanter and trochantellus white with infuscate stains dorsally; femur black except for narrow basal whitish band; tibia and tarsi dark brown, tibial spurs honey brown. Metasoma black dorsally, ventrally white. Wings hyaline basally, becoming weakly infuscate apically; veins dark brown.
Head. Antenna with 53 antennomeres, flagellomeres roughly 2.0× as long as wide; malar space short, just slightly longer than basal width of mandible, and 0.3× eye height; eyes large, in lateral view temple very narrow, in dorsal view eyes 4.7× longer than temples; occipital carina incomplete, not meeting dorsally and curving toward lateral ocelli, well defined laterally and meeting hypostomal carina; oral space small and circular, maximum width slightly smaller than basal width of mandible; clypeus slightly swollen; ocell-ocular distance about 0.7× diameter of lateral ocellus; maxillary palp not swollen; head surface sculpturing shining granulate, occiput smooth and shining, frons also smooth with small weak concentric wrinkles; higher face with a small longitudinal ridge and transverse rugosity directed to it; frons excavated with excavation bordered by a weak "W-shaped" carina.
Legs. Hind tibia with comb of modified setae; tarsal claw pectinate, bristles relatively long and tightly arranged, with a short gap between pectination and claw base; hind basitarsus 3.4× longer than inner apical spur of hind tibia.
Male unknown.
Discussion. Aleiodes dyeri sp. n. belongs to the seriatus species-group, where it resembles A. greeneyi, because of the dorsally incomplete occipital carina. It can be distinguished from greeneyi by the honey yellow mesosoma (dorsally black in greeneyi), the fore wing vein r 1.5× longer than RS+Mb (1.0× in greeneyi), and the hind wing vein r-m longer than 1M (shorter in greeneyi). A. dyeri sp. n. is similar to longikeros sp. n. in color patterns. These two species differ in the sculpturing of mesopleuron, being entirely granular in dyeri sp. n. but with a smooth elevated area in longikeros sp. n. The hind wing vein 1M is shorter than r-m in dyeri sp. n., as opposed to being 2.4× longer in longikeros sp. n., and the shape of fore wing vein 1M is weakly sinuate in dyeri sp. n., as compared with strongly curved in longikeros sp. n. Within the Nearctic species, dyeri sp. n. is more similar to Aleiodes preclarus Marsh & Shaw, 1998, from which it differs in the entire yellowish head but ocellar triangle black (several black spots in preclarus), wing veins mostly dark brown except fore wing veins M+CU and 1A, and hind wing veins 1M and M+CU proximally yellowish (pterostigma and fore wing vein C+SC+R with yellow spots in preclarus), and frons smooth (porcate in preclarus).
Comments. The antenna tips of the type specimen have a withered aspect, which makes impossible to measure the exact length of the antenna or describe the shape of the apical flagellomere.
Etymology. This species is named after Dr. Lee Dyer, of the University of Nevada (Reno), the lead investigator of the Caterpillars and Parasitoids of the Eastern Andes of Ecuador (CAPEA) project.
Head. Antenna with 45 segments, flagellomeres roughly 2.0× as long as wide, apical flagellomere with short "bottle-nipple"-shaped tip; malar space moderate, about 1.3× basal width of mandible, 0.4× eye height; in dorsal view eyes 2.5× longer than temples; occipital carina complete dorsally, well defined laterally and meeting hypostomal carina; oral space small and circular, maximum width equal to basal width of mandible; clypeus slightly swollen; ocell-ocular distance about as long as diameter of lateral ocellus; maxillary palp not swollen; head surface sculpturing shining granulate, occiput smooth and shining, frons also smooth with weak concentric wrinkles; frons excavated, excavation bordered by a weak "W-shaped" carina, but stronger laterally.
Mesosoma. Sculpturing shining granulate; pronotum foveate; propodeum rugulose with granular background and complete mid-longitudinal carina. Meso-and metapleuron surface rugose on dorsal dark brown areas; notauli shallow and crenulate anteriorly, posteriorly meeting on depressed rugose area; posterior margin of mesoscutum bordered with complete carina; scutellar sulcus with complete median carina plus two pairs of poorly defined lateral carina.
Wings. Legs. Hind tibia with comb of modified setae; tarsal claw simple, not pectinate; hind basitarsus 3× longer than inner apical spur on hind tibia.
Male. Antenna with 42-43 segments; metasoma narrower, T1 about 1.5-1.6× longer than its apical width; ocell-ocular distance shorter, 0.7× diameter of lateral ocellus; stigma narrower, 4.2× longer than high; one male has the white median stripe on metasoma interrupted on posterior half of tergite 2. Discussion. This species belongs to the seriatus species-group. Aleiodes elleni sp. n. is the only newly described species in this group with a strong, complete occipital carina on vertex. This character is present in two other Neotropical species: A. scriptus (from Brazil) and A. nebulosus (from Ecuador) from which elleni sp. n. differs by having the hind wing vein RS straight. A. elleni sp. n. also differs from scriptus by having the sculpturing of metasoma rugose-costate, as compared with widely costate in scriptus. It differs from nebulosus mostly in color patterns: pronotum yellow (white in nebulosus), apical 2/3 of hind coxa, propodeum, metanotum, metapleuron dorsally and mesopleuron on antero-dorsal corner black (white in nebulosus), and white medial marking on metasoma extending throughout tegite 1 and 2 (only anteriorly on tergite 1 in nebulosus). A. elleni sp. n. also differs from nebulosus in having the 2 nd submarginal cell in fore wing long and rectangular (short and trapezoidal in nebulosus), 2RS and 3RSa forming a right angle (obtuse in nebulosus), and vein r less than half length of vein 3RSa (0.85× in nebulosus), wings hyaline (wings moderately infuscate in nebulosus), and frons with lateral ridges (absent in nebulosus). The straight vein RS of the hind wing is shared with two other species of this species group: A. frosti sp. n. and the Nearctic species Aleiodes femoratus Cresson 1869; however, elleni sp. n. differs from frosti sp. n. by its smaller ocelli and relatively shorter petiole, and from both by the mostly black and medially whitish metasoma (yellowish in frosti sp. n. and femoratus). It also differs from femoratus by the smooth frons, as compared with porcate frontal sculpture in femoratus.
Etymology. This species is named after the American actress, comedian, and television host Ellen Lee DeGeneres.
Color. Body gold-honey brown to orangish brown. Flagellum dark brown, scape and pedicel laterally brown; ocellar triangle black; tarsal claws brown; metasoma ven-trally, mandibles, fore and mid coxa light yellowish. Wings weakly infuscate; veins dark brown but C+SC+R honey brown; stigma and vein R1 pale yellow-pale honey brown.
Head. Antenna with 50 antennomeres, flagellomeres roughly 2.0× as long as wide, apical flagellomere with short pointed apex; malar space as long as basal width of mandible, and 0.35× eye height; in dorsal view eye height 2.2× temple; occipital carina incomplete dorsally (but not curved toward vertex), otherwise complete but not touching hypostomal carina; oral space small and circular, maximum width equal to basal width of mandible; clypeus bulging; ocell-ocular distance 0.86× diameter of lateral ocellus; maxillary palp not swollen. Head surface sculpturing finely granulate, higher face with small longitudinal ridge and transverse rugosity directed to it, occiput smooth and shining.
Mesosoma. Sculpturing mostly granulate; pronotum foveate laterally; mesopleuron rugose on anterior corner; propodeum rugose-granulate, posteriorly with diverging and some transverse wrinkles and weakly rugose laterally, with mid-longitudinal carina present on anterior 2/3; notauli present anteriorly, wide and shallow, posteriorly disappears in a depressed area with striations running antero-laterally from mid-posterior region; posterior margin of mesoscutum with short carina, just anterior to scutellar sulcus; scutellar sulcus with median carina plus two pairs of well defined lateral carina.
Wings. Fore wing: stigma 3× longer than high; vein r 0.55× vein 2RS, as long as vein RS+Mb, and 0.5× as long as vein m-cu; vein 3RSa about 0.5 times vein 3RSb, and as long as vein 2M; vein 1CUa 2× vein 1cu-a; vein 1CUb 1.8× vein 1CUa; vein 1M evenly slightly curved. Hind wing: m-cu indicated as short pigmented vein antefurcal to vein r-m (in this species and others, the vein m-cu is very short but is also indicated by a slight bent on vein M, were these veins meet); vein M+CU about 1.4× 1M; vein 1M 1.4× vein r-m; vein RS smoothly curved at middle; vein M straight; vein 2-1A absent.
Legs. Hind tibia without comb of modified setae; tarsal claw pectinate at base, with a distinct gap between apical claw and basal pectination, pectin with 5-6 bristles; hind basitarsus 3× longer than inner apical spur on hind tibia; few rugositie dorsolaterally on outer side of hind coxa.
Male unknown. Discussion. This species belongs to circumscriptus/gastritor species group. It differs from the described species in this group by its mostly honey brown body color. A. falloni sp. n. resembles Aleiodes speciosus Townsend, from which it can be distinguished by the entire honey brown body color (mostly black-dark brown dorsally with first tergite white in speciosus) and the hind wing vein 2-1A absent (present in speciosus). It is very similar to A. luteosicarius sp. n., which belongs to the pallidator species-group, especially in color pattern (for distinguishing features see discussion at luteosicarius sp. n. section). Morphological distinction between A. falloni sp. n. and A. luteosicarius sp. n. is difficult due to their general resemblance. Separation of specimens in two different entities was supported by comparison of ribosomal COI sequences, resulting in two groups with considerably different genetic information.
Etymology. This species is named after James Thomas Fallon, known as Jimmy Fallon, an American television host, comedian, actor, singer, musician and producer.
Body color. Yellowish to honey yellow, except for the ocellar triangle and antenna dark brown. Wings tinged yellowish; veins honey brown, parastigma blackish with central yellowish spot.
Head. Antenna 65 antennomeres, antenna 1.3× longer than body, flagellomeres roughly 2× longer than wide, apical flagellomere with long and narrow "bottlenipple"-shaped apex; malar space short, about as long as basal width of mandible, and about 0.33× eye height; in dorsal view eyes 5× longer than temples; occipital carina incomplete dorsally, well defined laterally and meeting hypostomal carina; oral space small and circular, maximum width equal to basal width of mandible; clypeus large, not swollen; ocelli large, ocell-ocular distance about 0.4× diameter of lateral ocellus; maxillary palp not swollen; head surface sculpture finely shining granulate, occiput smooth and shining; higher face with a small longitudinal ridge and transverse rugosity directed to it; frons polished and excavated, without lateral ridges.
Mesosoma. Sculpturing shining granulate; pronotum with few wrinkles posteriorly and dorso-laterally; mesopleuron with small area at antero-dorsal corner rugose, and some wrinkles dorsally; propodeum longitudinally rugose posteriorly, mid-longitudinal carina almost complete; notauli with few crenulae and shallow anteriorly, meeting on depressed rugose area posteriorly; posterior margin of mesoscutum with complete carina; scutellar sulcus shallow and smooth except for the median carina.
Legs. Hind tibia with comb of modified setae; tarsal claw simple, not pectinate; hind basitarsus 3× longer than inner apical spur of hind tibia.
Metasoma. T1, T2 and basal 2/3 of T3 rugose-striate, longitudinal carina complete on T1 and almost complete on T2, but not reaching posterior margin; ovipositor sheaths parallel sided and truncate, about as long as hind tarsomere III; metasoma unusually long and narrow, T1 2.2× longer than its apical width.
Male. Body length about 8.5 mm; antenna with 63 segments. Virtually identical to female, but ocell-ocular distance 0.3× diameter of lateral ocellus.
Mummy. Length 14.0 mm, black, thorax brown, head honey brown mottled brown, tubular in shape, thorax compact and wrinkled, exit mode unique within Aleiodes: the parasitoid cuts a radial opening at posterior side of the mummy, just behind the hind abdominal prolegs, releasing a "lid" with the anal prolegs. Biology. Host plant Chusquea scandens (Poaceae); host Lepidoptera: Scoturopsis Hering sp. (Notodontidae); time span from pupation to emergence: about 5 weeks for females, unknown for the male. The parasitoid cuts a radial opening at posterior side of the mummy releasing a "lid", comprinsing the anal apex of mummified caterpillar, before emergence. The mummy exit mode of this species is unique for the genus, since all previously known mummies produced by Aleiodes species had a posterior hole cut for emergence (Zaldívar-Riverón et al. 2008).
Discussion. Aleiodes frosti sp. n. belongs to the seriatus species-group. This species resembles Aleiodes nigricosta (Enderlein 1920) because of its entirely yellowish to honey yellow body, black stemmaticum and brown antenna, but differs in the honey brown fore wing vein C+SC+R, black in nigricosta. A. frosti sp. n. also differs in the extension of median longitudinal carina on metasoma, which is incomplete on tergite 2, but extends to half of tergite 3 in nigricosta, and the exceptionally elongate metasoma. A. frosti sp. n. is also similar to elleni sp. n. by the nearly straight hind wing vein RS, enclosing a marginal cell gradually widening toward wing apex, but it can be readily distinguish by the interrupted occipital carina on vertex, compared to the complete occipital carina of elleni sp. n. The diameter of lateral ocelli, 3-4× longer than ocell-ocullar distance, is also a diagnostic character shared only with one Neotropical species, Aleiodes nigribasis (Enderlein 1920); however, most of the already mentioned diagnostic features for frosti sp. n. (e. g. shape of metasoma and hind wing vein RS, and color patern) are also useful to distinguish it from nigribasis. Within the Yanayacu species in the seriatus group it is similar to greeneyi because of the incomplete occipital carina at vertex. It differs from greeneyi by the entire yellowish body and its unusual long and narrow metasoma.
Etymology. The species is named after the American poet Robert Frost (1874 -1963), author of the poem "The Road Not Taken." This species name is also a reference to that poem, and to the unusual emergence mode of this species, recorded here for the first time. The following quotation extracted from this poem summarizes its idea: "Two roads diverged in a wood, and I -I took the one less traveled by," Robert Frost, 1920. This Aleiodes species takes a "road not taken" by other species, to its adulthood, by emerging in a different and unique way.
Additional characters. Last flagellomere with "bottle-nipple"-like tip; mesoscutum with carina at posterior margin complete although not well defined; scutellar sulcus with complete median carina plus one pair of slightly weaker lateral carina; fore wing vein 1M moderately curved at base; hind wing vein 2-1A indicated as a very short stub, vein m-cu present, interstitial to vein r-m; ovipositor sheaths about as long as hind tarsomere II, 0.6× hind basitarsus.
Distribution. Known only from the type locality, YBS, Napo province, ECUADOR. Discussion. Aleiodes greeneyi is known only by the holotype. It belongs to the seriatus species-group. The incomplete occipital carina at vertex and the moderate ocelli size are similar to those presented in A. longikeros sp. n. and A. dyeri sp. n.; however, the wing vein pattern is somewhat intermediate between these two species. A. greeneyi differs from longikeros sp. n. and dyeri sp. n. in having the pronotum and mesoscutum mostly black (yellow in longikeros sp. n. and dyeri sp. n.), and the hind coxa light brown (bicolored black and white in longikeros sp. n. and dyeri sp. n.). A. greeneyi was reared from geometrid larva. Within the seriatus-group species from Ecuador, A nebulosus and longikeros sp. n. were also reared from Geometridae hosts.
Color. Mostly black. Head honey yellow, ocellar triangle black; antenna dark brown except apical border of pedicel honey yellow. Mesosoma almost entirely black; propleuron and ventral quarter of pronotum, mesoscutum mid-posteriorly and scutellum medially yellowish to honey yellow; posterior border of propodeum white. Fore and mid legs with whitish coxa, darkening apically to honey yellow apical femur and tibia, and brown tarsi; hind leg coxa, trochanter and trochantellus black, but apical border of trochanter and trochantellus, and a small ventral spot on trochantellus white; femur black on basal 2/5 and dorso-apically, otherwise yellowish; tibia and tarsi brown, subbasal whitish small band on tibia. Metasoma black dorsally except for the white T1; apical borders of T4-T7 whitish; basal T5-T7 brownish. Wings moderately infuscate; veins dark brown.
Head. 40 antennomeres, flagellomeres roughly 2.0× as long as wide, apical flagellomere with short pointed apex; malar space moderate, length 1.3× basal width of mandible, and approximately 1/3 eye height; in dorsal view eye 2.6× temples; occipital carina incomplete dorsally, curving toward lateral ocelli, well defined laterally but not meeting hypostomal carina; oral space small and circular, maximum width equal to basal width of mandible; clypeus slightly swollen; ocellus moderate, ocell-ocular distance short, about half diameter of lateral ocellus; maxillary palp not swollen; head surface sculpturing shining granulate, occiput smooth and shining; frons excavated with short lateral ridges.
Mesosoma. Sculpturing finely granulate; pronotum with some wrinkles laterally; mesopleuron coriaceous on central elevated area and some irregular latero-ventral parts; propodeum with mid-longitudinal carina incomplete, granulate postero-laterally with irregular wrinkles, and triangular rugose area mid-anteriorly with diverging wrinkles; notauli well defined anteriorly, narrow and crenulate, meeting a depressed rugose area posteriorly; posterior margin of mesoscutum bordered by complete carina; scutellar sulcus shallow, with median carina plus two pairs of poorly defined lateral carina. Wings Legs. Hind tibia without comb of modified setae; tarsal claw simple, with a comb of thin bristles medially; hind tibial spurs relatively long, hind basitarsus 2.4× longer than inner apical spur.
Discussion. This species belongs to circumscriptus/gastritor species-group. It produces an unusual curled mummy, "J-shaped", and emerge through a large and almost round exit hole at the posterior end. The white first tergite of A. kingmani sp. n., contrasting to the mostly black metasoma, is a useful diagnostic character shared only with speciosus and townsendi sp. n. A. kingmani sp. n. differs from townsendi sp. n. by the mostly black mesosoma (mostly yellowish in townsendi sp. n.), and the entirely black hind coxa (bicolored black and white in townsendi sp. n.). A. kingmani sp. n. differs from speciosus in having mesopleuron black (mostly yellow in speciosus) and entirely granulate (mostly smooth in speciosus); black region on head restricted to ocellar triangle (covering most of vertex and occiput dorsally in speciosus); hind coxa, trochanter and trochantellus black (yellow in speciosus); and occipital carina not meeting the hypostomal carina.
Etymology. This species is named after Eduardo Kingman (Loja, February 23, 1913-Quito, November 27, 1997, one of the greatest Ecuadorian artists, who dedicated his art to portray the indigenous people of Ecuador.
Head. Antenna with 60 antennomeres, about 1.6× as long as body, flagellomeres narrower than in other species, most flagellomeres 2.5× longer than wide, apical flagellomere with long "bottle-nipple"-shaped apex; malar space 1.1× longer than basal width of mandible, 0.4× longer than eye height; in dorsal view eyes 3× longer than temples; occipital carina barely incomplete, almost meeting dorsally and directed toward vertex, well defined laterally and meeting hypostomal carina; oral space small and circular, maximum width 0.8× basal width of mandible; clypeus not swollen; ocellocular distance about as long as diameter of lateral ocellus; maxillary palp not swollen; head surface sculpturing shining granulate, occiput smooth and shining; higher face with a small longitudinal ridge and transverse rugosity directed to it; frons only shallowly excavated.
Mesosoma. Sculpturing shining granulate; pronotum smooth laterally, dorsally granulate and foveate; mesopleuron with central elevated area smooth, antero-dorsal corner rugose; propodeum coarsely shining granular with complete mid-longitudinal carina; notauli very shallow anteriorly, virtually absent. Mesoscutum with some transverse wrinkles on anterior region of notauli, mid-posterior depressed area with few longitudinal wrinkles; posterior margin of mesoscutum with complete carina; scutellar sulcus with median carina plus two pairs of complete lateral carina.
Wings. Legs. Hind tibia with comb of modified setae at apex; tarsal claw simple, not pectinate, with a comb of relatively long thin setae basally. Hind tibial spurs relatively short, hind basitarsus 3× longer than inner spur.
Male unknown Mummy. Length 12.5 mm, light reddish brown, head and prolegs light yellow, head with two longitudinal brownish stripes, thorax compact and wrinkled, posterior apex withered, glue hole located ventrally on the thorax, exit hole irregular, located postero-dorsally, posterior to hind abdominal prolegs.
Discussion. Aleiodes longikeros sp. n. belongs to the seriatus species-group. This species has the longest antenna of any of these 24 newly described species, even though this is a medium sized species. Its antenna is about 1.6× longer than body, as compared with at most 1.3× in other species. The fore wing vein 1M strongly curved basally is also seen in townsendi sp. n. and shakirae sp. n., both belonging to circumscriptus/gastritor speciesgroup, therefore both distinct from longikeros sp. n. by the absence of apical comb of flattened setae on hind tibia. A. longikeros sp. n. is similar to dyeri sp. n. (distinguishing features are discussed on diagnosis section on dyeri sp. n.). It also resembles greeneyi because of the dorsally incomplete occipital carina, from which it can be distinguished by the honey yellow mesosoma (dorsally black in greeneyi), and white mid-apical spots on metasomal terga 1-4 (mostly black to dark brown in greeneyi); mesopleuron smooth on dorsal elevated area (granulate in greeneyi). A. longikeros sp. n. can be distinguished from all New World species by the following combination of characters: fore wing second submarginal cell long and narrow, vein 2RS 0.35× longer than vein 2M; the long antenna; and strongly curved vein 1M on fore wing. In the key to Nearctic species of the seriatus group (Marsh and Shaw 1998), longikeros sp. n. runs to preclarus. The new species differs from preclarus by the above mentioned character combination, and also by the entirely yellowish head and mesosoma, with several dark spots in preclarus, and the smooth frons, as compared with porcate frontal sculpturing in preclarus.
Etymology. From the Latin, meaning "long horned," being a reference to the unusually long antenna of this species.
Color. Entire body honey brown to bronze, notum slightly darker; ocellar triangle black; antenna dark brown; wings hyaline; veins brown except C+SC+R, parastigma centrally, stigma, and R1 yellow; ovipositor mostly with same color of body, only weakly darkening apically.
Head. Antenna with 48 antennomeres, flagellomeres roughly 2.0× as long as wide, apical flagellomere with "bottle-nipple"-shaped apex; malar space as long as basal width of mandible, and 0.3× eye height; in dorsal view eye height 2.7× temple; occipital carina incomplete dorsally but not curved toward vertex, otherwise complete but not touching hypostomal carina; oral space small and circular, maximum width equal to basal width of mandible; clypeus not swollen; ocell-ocular distance 0.9× diameter of lateral ocellus; maxillary palp not swollen. Head surface sculpturing finely granulate, higher face with small longitudinal ridge and transverse rugosity directed to it, vertex coarsely granulate with some transverse wrinkles, occiput smooth and shining.
Mesosoma. Sculpturing mostly granulate; pronotum foveate laterally; mesopleuron rugose on anterior corner; propodeum mostly coarsely granulate with few longitudinal wrinkles posteriorly, laterally weakly rugose, with mid-longitudinal carina present on anterior 2/3; notauli present anteriorly, wide and shallow, posteriorly disappears in a depressed longitudinally rugose-striate area; posterior margin of mesoscutum with short carina just in front of scutellar sulcus; scutellar sulcus with strong median carina plus two pairs of strong but incomplete lateral carina. Legs. Hind tibia without comb of modified setae; tarsal claw pectinate with distinct gap between apical claw and bristles; hind basitarsus 3× longer than inner apical spur on hind tibia; few rugositie dorso-laterally on outer side of hind coxa.
Metasoma. T1-T2 and basal half of T3 striated; remainder terga coriaceous; mid longitudinal carina complete from T1 throughout T2; ovipositor sheaths slightly shorter than hind tarsomere II, relatively thick and somewhat rounded at tip, uniformly and densely covered with relative short regular sized pubescence, but dorso-basally bare, setae length no longer than width of sheaths; T1 slightly longer than apical width.
Paratype variation. Body length 6.7-7.4 mm; antenna 48-50 segments; 3 to 5 carina on scutellar sulcus; about half of paratypes have a lighter body color, otherwise very similar to holotype.
Mummy. All mummies densely setose, setae mostly yellowish to light brown contrasting with dark brown body, head varying from dark brown to yellowish brown; morphology of mummies variable according to different host species, exit hole irregular, postero-dorsal.
Discussion. Aleiodes luteosicarius sp. n. is the only species herein described which belongs to pallidator species-group. It is very similar to falloni sp. n., despite falloni sp. n. belongs to circumscriptus/gastritor species-group. A. luteosicarius sp. n. differs from falloni sp. n. in the following characters: ocell-ocular distance 0.3× diameter of lateral ocellus (about 0.9× in falloni sp. n.); at least half of tergite 3 striated, as in tergite 2 (tergite 3 mostly smooth-coriaceous); m-cu of hind wing slightly postfurcal to almost interstitial (antefurcal in females of falloni sp. n.); clypeus not swollen (swollen in falloni sp. n.); ovipositor sheaths relatively thick and somewhat rounded at tip, uniformly and densely covered with relative short regular sized pubescence, but dorso-basally bare, setae length no longer than width of sheaths (ovipositor sheaths sharpening at apex, setae concentrated apically, and not regular sized, longest setae about 1.5× longer than width of sheaths in falloni sp. n.); fore wing as long as body length (distinctly shorter in falloni sp. n.); stigma narrower (4× longer than high in luteosicarius sp. n. vs. 3× in falloni sp. n.); hind wing vein r-m as long as vein 1M (distinctly shorter in falloni sp. n.); depressed area on mesoscutum longitudinally rugose-striate (striations running antero-laterally from mid-posterior region); propodeum mostly coarsely granulate with few longitudinal wrinkles posteriorly, laterally weakly rugose (distinct pattern of rugosity on propodeum in falloni sp. n.); occipital carina strong and abruptly interrupted dorsally (weaker and gradually disappearing in falloni sp. n.); vertex coarsely granulate with some transverse wrinkles (finely granulate in falloni sp. n.). In the key to species of pallidator species-group from North America (S. Shaw et al. 2013) it runs to Aleiodes pallidator (Thunberg, 1822). The new species closely resembles pallidator, differing from it by the following: propodeum surface is entire rugose (laterally coriaceous in pallidator) and the longitudinal carina is present on anterior 3/4 (complete in pallidator); vein 1CUa about 2× length of 1cu-a (just slightly longer in pallidator); basal cell of hind wing very broad, vein r-m as long as vein 1M (narrower in pallidator); metasomal tergite 1 granular-rugose and tergite 2 striate-rugose (costate in pallidator); parasitoid on Arctiinae (Erebidae) (pallidator attacks Lymantriidae).
Comments. All previous known species of the pallidator species-group are parasitoids on Lymantriinae caterpillars (S. Shaw et al. 2013, as Lymantriidae). This is the first record of a species in this group attacking Arctiinae, and also the first species of the group with known host from Neotropical region. The status of subfamily for these groups is relatively recent, and both belong to Erebidae. Species within the pallidator group has been consistently reared from the setose mummified caterpillars of Lymantriinae in Japan, Europe and North America. The host associations reported here broaden the known host range for this species-group, but also denotes its ecological preferences for attacking densely setose caterpillars.
Etymology. From the Latin meaning "yellow killer," referring to the main color of this parasitoid.
Color. Head whitish, except for the ocellar triangle black, and the light yellow face and palp. Mesosoma black but pronotum and propleuron light yellow, and small posterior spot on metapleuron and posterior 1/5 of propodeum whitish; fore legs coxa, trochanter, trochantellus and mostly femur yellow; fore femur with dorso-apical brown stain, reminder fore legs brown; mid legs dark brown, but trochanter and trochantellus white; hind legs black. Metasomal tergite 1 with a black oval spot medially spanning from just behind basal carina to posterior edges of the tergite, remainder of tergite 1 white; tergite 2 black medially, dark region almost quadrate in shape with lateral borders convex at 2/3 posterior, white laterally; tergite 3 mostly white, with basal almost semi-circular black mark; reminder terga white apically, basally black, more or less concealed by the preceding tergite; metasoma ventrally white with a pair of laterobasal spots on each sternite, the spots larger on the second sternite; ovipositor sheaths black, ovipositor yellow.
Head. 50 antennomeres; most flagellomeres roughly 2× longer than wide, apical flagellomere with small pointed apex; malar space wide, about 2.5× times basal width of mandible and almost as long as eye height; temple wide, in dorsal view about as long as eye; occipital carina absent; oral space small and circular, diameter about equal to basal width of mandible; clypeus weakly swollen; ocelli very small, ocell-ocular distance 2.2× diameter of lateral ocellus; head polished (smooth and shining).
Male unknown. Mummy. Length 14.9 mm, mostly pale brown with some dark spots, dorsally at middle of abdomen with faint "X-shape" mark and four withered expansions on each tip of the mark, thorax very strongly compact and abdomen angled upward, very similar to A. capillosus mummies, glue hole ventrally on thorax, exit hole postero-dorsal, between abdominal and anal prolegs.
Discussion. This is the fourth species of the gressitti species-group described. Of the former three, two are from New World, Aleiodes lissos Marsh & Shaw, 2003 and Aleiodes capillosus Townsend, and one from Campbell Island in the South Pacific: A. gressitti Muesebeck, 1964. This new species most resembles the Neotropical capillosus because of the absence of occipital and epicnemial carina, and also in the mostly black mesosoma and infuscate wings. It differs from capillosus in the whitish head, except for the ocellar triangle being black and the face light yellow, pronotum and propleuron, mid trochanter and trochantellus white, and fore leg coxa, trochanter, trochantellus and mostly femur yellow (while all these parts are black in capillosus). The malar space is very large, being about as long as the eye height (at most 1/2 in other gressitti-group species). The mummies produced by marilynae sp. n. and capillosus are very similar in the extremely contracted thorax and relatively swollen abdomen, and also in the characteristic acute angle at which the mummy is attached to the substrate.
Etymology. This species is named in honor of Marilyn Rieden Shaw, wife of the co-author, Scott R. Shaw, in gratitude for her support for his entomological studies over many years.
Color. Mostly black. Head orangish yellow, including mandibles and palp, but mandibles tip brown; ocellar triangle black; antenna dark brown except apical border of pedicel honey yellow. Mesosoma almost entirely black; propleuron, pronotum, and scutellum medially honey yellow; posterior border of propodeum white. Fore leg yellowish; outer apical 1/3 of femur infuscate; basal half of tibia and tarsi brown. Mid leg with same pattern of fore leg but ground color whitish. Hind leg black; light yellow markings on apical border of trochanter and trochantellus, small ventral spot on trochantellus, apical half of femur latero-ventrally, and small sub-basal band on tibia. Metasoma black dorsally, bordered white; T1, triangular mid-basal area on T2 and apical borders of T3-T7 white; ventrally white but infuscate medially; ovipositor sheaths basal 1/3 whitish, apical 2/3 black. Wings weakly infuscate; veins dark brown.
Head. Antenna with 46 antennomeres, flagellomeres roughly 2.0× as long as wide, apical flagellomere with very short pointed tip; malar space wide, about 1.8× basal width of mandible, and 0.6× eye height; in dorsal view eye 1.4× temple; occipital carina incomplete, close but not meeting dorsally and curving toward lateral ocelli, well defined laterally and meeting hypostomal carina; oral space small and circular, maximum width equal to basal width of mandible; clypeus slightly swollen; ocelli small, ocell-ocular distance about 1.8× diameter of lateral ocellus; maxillary palp not swollen; head surface sculpturing granulate, occiput smooth and shining; frons excavated with short lateral ridges.
Mesosoma. Sculpturing finely granulate; pronotum foveate; mesopleuron central disc mostly smooth and bare, posteriorly and ventrally coriaceous, antero-dorsal corner rugose; propodeum coarsely granular with complete mid-longitudinal carina; notauli well defined and crenulate anteriorly, barely defined but traceable posteriorly, meeting a depressed rugose area; posterior margin of mesoscutum bordered with complete carina; scutellar sulcus with median carina plus two pairs of incomplete lateral carina.
Legs. Hind tibia without apical comb of modified setae; tarsal claw simple, not pectinate, with a comb of relatively long thin setae basally; hind tibial spurs relatively short, about 1/4 basitarsus length.
Discussion. Aleiodes mirandae sp. n. belongs to circumscriptus/gastritor speciesgroup. It is similar to A. napo sp. n. in the very small ocelli (ocell-ocular distance about 2× diameter of lateral ocelli), and also the mostly smooth mesopleuron. It differs from napo sp. n. in the rugose depressed mid-posterior area on mesoscutum, flat and granular in napo sp. n., the head, except black ocellar triangle, pronotum, propleuron and scutellum orangish yellow, all black in napo sp. n. except for small reddish marking on temples. The color pattern is very similar to kingmani sp. n., but additionally to already mentioned diagnostic characters mirandae sp. n. have hind wing vein M+CU shorter than 1M, while in kingmani sp. n. 1M is more than 2× longer than r-m. The host species "palito café chusquea" (Geometridae) is the same species attacked by Aleiodes nubicola sp. n. and Aleiodes shakirae sp. n.
Color. Mostly black. Head with a small brown stain on temples, just behind eyes; palp, fore and mid coxa, trochanter and trochantellus pale yellowish. Metasomal tergite 1 white, with a black oval spot medially spanning from just behind basal carina to posterior edges of the tergite; remainder terga mostly black, white laterally; metasoma ventrally white with a pair of latero-basal spots on each sternite, the spots larger on the second sternite; ovipositor sheaths black, ovipositor yellow.
Head. 47 antennomeres; most flagellomeres about 1.5× longer than wide, apical flagellomere with small pointed tip; malar space moderate, about 1.5× times basal width of mandible, and 0.5× eye height; temple wide, in dorsal view slightly shorter than eye; occipital carina absent dorsally, running toward vertex, ventrally almost meeting hypostomal carina; oral space small and circular, diameter about equal to basal width of mandible; clypeus with distinct line separating dorsal and ventral surfaces, dorsally coriaceous, ventrally convex and smooth; ocelli very small, ocell-ocular distance about 2.5× diameter of lateral ocellus; head, including frons, granulate.
Mesosoma. Mesoscutum and scutellum granular coriaceous; pronotum dorsally and laterally on anterior half granular with scrobiculate sulcus, reminder lateral pronotum smooth except for the posterior corner coarsely rugose; propleuron granularcoriaceous; mesopleuron mostly smooth centrally to finely coriaceous, antero-dorsal corner weakly rugose-punctate; metapleuron and propodeum coarsely granular; longitudinal carina on propodeum almost complete; notauli present only anteriorly, narrow and crenulate, mid-posterior area of mesoscutum not depressed and granulate; entire posterior margin of mesoscutum bordered by carina; scutellar sulcus with median carina plus two pairs of lateral carina.
Male unknown.
Mummy. Length 12.0 mm, head honey yellow, thorax pale brown, abdomen dark reddish brown, almost tubular in shape, exit hole located postero dorsally.
Type material. Type-locality: ECUADOR, Napo Province, Yanayacu Biological Station, YY-48553, S00°35.9', W77°53.4', 2163m, cloud forest, July 14, 2010 Type-specimen: Holotype female and mummy, point mounted separately. Top label: "ECUADOR: Napo Province / Yanayacu Biological Station / S00°35.9', W77°53.4' 2163m / CAPEA -NSF-BSI-07-17458 / REARED / 2010 (hand written) May 48553"; back (hand written): "14-Jul-2010". (UWIM) Biology. Reared from a Noctuidae caterpillar (common name "raya roja a los lados chusquea") collected on Chusquea scandens (Poaceae). The parasitoid took six weeks from host mummification until emergence. Discussion. This species belongs to circumscriptus/gastritor species-group. The color pattern of Aleiodes napo sp. n. is similar to those of capillosus; however, it does not belong to the gressitti species group because of the sculpturing on metasomal tergite 3, which is granular coriaceous on basal half. A. napo sp. n. differs from capillosus also in the presence of both occipital and epicnemial carina, though the former is incomplete dorsally, the presence of a complete longitudinal carina on metasomal terga 1 and 2, body sculpturing mostly granular-coriaceous, and longitudinal carina on propodeum almost complete (in capillosus all the above mentioned carina are absent, and the body sculpturing mostly smooth). Within the circumscriptus/gastritor group, A. napo sp. n. is similar to mirandae sp. n. in having a smooth central disc of mesopleuron and hind wing vein M+CU shorter than 1M, but differs from it in the almost entirely black head and thorax, mostly orangish yellow in mirandae sp. n., and the posterior central region of mesoscutum flat and granular, depressed and rugose in mirandae sp. n.
Etymology. This species is named after the indigenous inhabitants of the eastern Ecuador, for whom the Province (locality of the type) is also named: the Napo Runas.
Additional characters. Last flagellomere with short pointed tip; mesoscutum with complete and well defined carina bordering posterior margin; scutellar sulcus with seven carina; fore wing vein 1M slightly curved at base; hind wing vein 2-1A absent, vein m-cu present, distinctly antefurcal to vein r-m; ovipositor sheaths about as long as hind tarsomere II, 0.5× hind basitarsus. Different than originally described, the sternaulus is absent.
Distribution. Known only from the type locality, Isla de Las Palmas, Napo province, ECUADOR, at 1,885 meters elevation.
Discussion. Aleiodes nebulosus is known only by the holotype. In the seriatus speciesgroup, A. nebulosus is similar to A. elleni sp. n. in having the occipital carina complete at vertex. These two species are also similar in color pattern because of the most yellowish head and mesoscutum, and the mostly dark brown to black metasoma with a whitish mark antero-medially. Distinguishing features between nebulosus and elleni sp. n. are presented in the discussion section for elleni sp. n.
Paratype variation. Body length 3.8-4.6 mm; antenna with 38-40 antennomeres; head in most specimens mostly dark brown, except most of gena and temples just behind eyes honey yellow, but the gena is entirely dark brown in two type specimens; the clypeus color is also variable, in most specimens it is contrasting honey yellow, but in two specimens the clypeus has the same color of face; scutellum color varies from entire black to yellow on apical half, scutellar sulcus varies from yellow to black; light lateral stripes on mesopleuron varies from brown, reddish brown to yellow, in one specimen the stripes are connected by ventral yellowish stripe; most paratypes with metasomal terga 4 and 5 mostly blackish; position of hind wing vein m-cu varies from just postfurcal to just antefurcal; hind basitarsus 4-5× longer than inner apical spur on hind tibia.
Male. Essentially as in female, but eyes slightly smaller; antenna with 35 and 38 segments; metasomal terga entire dark brown in on specimen, the other with whitish central markings throughout all metasomal terga from apical tergite 1. Mummy. Length 8.5-10.8 mm, entire brown, mummy with elongate aspect and thin skin, widening gradually from neck to posterior exit hole, thorax wrinkled, mummy attached to the substrate by silk posteriorly at prolegs region, exit hole irregular, located postero-dorsally anterior to abdominal prolegs, but in two specimens the hole is located postero-ventrally.
Discussion. Aleiodes nubicola sp. n. is similar to A. cacuangoi sp. n. and A. atripileatus (see diagnosis of A. cacuangoi for differences). This species also resembles A. arbitrium in the size of ovipositor and the mostly dark brown head. It differs from arbritrium by the longer malar space, about 1.6× longer than mandible width at base (about 1.0× in arbitrium), the mostly black pronotum and mesonotum (mostly brownish orange in arbitrium), and the position of light marks on head bordering eyes on temples (same marks on vertex in arbitrium). The host species "palito café chusquea" (Geometridae) is the same species attacked by Aleiodes mirandae sp. n. and Aleiodes shakirae sp. n.
Etymology. From Latin, means "cloud inhabiting", a reference for the cloud forest habitat.
Color. Mostly black. Head orangish except black ocellar triangle; cheeks, palp and mandibles whitish; mesocutum with postero-median yellowish square mark; scutellum lighter; fore coxa and femur honey yellow; mid coxa, and all trochanter and trochantellus whitish; most of mid and hind femur whitish; all femur with small brown spot apical-dorsally; all tibia and tarsi brown; hind coxa black; hind tibial spurs and base of basitarsi honey brown; tip of metasoma behind T5 brown; ovipositor sheaths dark brown on apical half, basal half whitish.
Head. Antenna with 33 antennomeres, flagellomeres roughly 2.0× as long as wide, apical flagellomere with small pointed tip; malar space moderate, length about 1.5× basal width of mandible, and half eye height; in dorsal view eye height 2× temples; occipital carina interrupted at vertex, ventrally reaching hypostomal carina; oral space small and circular, maximum width equal to basal width of mandible; clypeus not swollen; ocelli moderate, ocell-ocular distance as long as diameter of lateral ocellus; maxillary palp not swollen; head surface sculpturing finely granulate, but occiput smooth and shining.
Mesosoma. Sculpturing mostly granulate; pronotum with some wrinkles; mesopleuron with anterior corner rugose; propodeum more coarsely granulate, with long mid-longitudinal carina on basal 2/3; notauli deep and crenulate anteriorly, meeting on depressed area posteriorly; posterior margin of mesoscutum with short carina, just anterior to scutellar sulcus; scutellar sulcus with median carina plus two pairs of poorly defined lateral carina. Wings Legs. Hind tibia without comb of modified setae; tarsal claw simple, not pectinate, with a comb of relatively long thin setae basally; hind basitarsus 3× longer than inner apical spur on hind tibia.
Paratype variation. None observed, virtually identical to holotype. Male unknown.
Mummy. Length about 6.5 mm, body entire graphite metallic black color, head orangish yellow, mummy aspect robust, body with one row of setal sockets on each segment and sparse setae except dorsally, wrinkles on thorax; exit hole irregular, located postero-dorsally.
Discussion. This species belongs to circumscriptus/gastritor species group. A. onyx sp. n. is similar to A. atripileatus; however, it can be distinguished by color patterns: head entirely orangish except ocellar triangle black (occiput mostly black in atripileatus), propleuron and ventral 1/4 pronotum honey brown (dark brown-black in atripileatus), mesopleuron whole black (ventral 1/2 honey brown in atripileatus), mesoscutum postero-central region honey yellow (whole dark-brown-black in atripileatus); as well as sculpturing features: mesopleuron central elevated area smooth (granulate in atripileatus), propodeum extensively rugose (granulate in atripileatus); and antenna with fewer flagellomeres: 31 in onyx sp. n. vs. 34 or more in A. atripileatus. Other diagnostic characters fo onyx sp. n. are the occipital carina interrupted on vertex, the very short ovipositor, about as long as hind 3 rd tarsomere, metasoma stout, T1 about 0.9× as long as its apical width and slightly wider than mesosoma. The host mummy is similar to that of atripileatus, but the anal prolegs are not posteriorly extended in mummies made by onyx sp. n. The base color of the mummy is a metallic graphite-like tone, as opposed to opaque black in A. atripileatus mummies. A. onyx sp. n. is the first Aleiodes species known from Ecuador to be reared from Zygaenidae caterpillars.
Etymology. From the Greek, the word onyx means "nail". It is the name of a rock, used in adornments since ancient times, with several colors, being the black ones the most appreciated. The name is a reference for the main black color of this parasitoid mummy, which resembles the color of the black onyx rocks.
Head. Antenna with 47 antennomeres, flagellomeres roughly 2.0× as long as wide, apical flagellomere with long "bottle-nipple"-shaped apex; malar space as long as basal width of mandible, and approximately 1/4 eye height; in dorsal view eyes 3.4× temples; occipital carina incomplete dorsally, curving toward lateral ocelli, laterally complete and meeting hypostomal carina; oral space small and circular, maximum width slightly equal to basal width of mandible; clypeus not swollen; ocellus moderate, ocell-ocular distance short, about 0.45× diameter of lateral ocellus; maxillary palp not swollen; head surface sculpturing finely granulate, occiput smooth and shining; higher face with some wrinkles just bellow toruli.
Mesosoma Legs. Hind tibia without comb of modified setae; tarsal claws pectinate basally, with several very short and tight bristles, longer and sparser apically, wide gap between apical bristles and claw; hind basitarsus 3× longer than inner apical spur on hind tibia.
Paratypes variation. Body length 4.5-6.2 mm; antenna with 47-50 segments; legs color somewhat darker in few paratypes, and/or mid trochantellus laterally infuscate; scutellar sulcus with 3 to 5 carina; other color and proportions with only minimal variation.

Aleiodes speciosus
Additional characters. Last flagellomere with short pointed tip; mesoscutum with carina at posterior margin almost complete but weakly defined; scutellar sulcus shallow, with incomplete median carina; wings moderately infuscate; fore wing vein 1M moderately curved at base; hind wing vein 2-1A present, vein m-cu absent.
Distribution. Known only from the type locality, Camino a Loreto, Napo province, ECUADOR, at 1,383 meters elevation.
Discussion. Aleiodes speciosus is known only by the male holotype. This species belongs to the circumscriptus/gastritor species-group. A. speciosus is similar to kingmani sp. n. mainly by color pattern, but also in having relatively large ocelli. These two species have a white metasomal tergite 1, contrasting with the reminder dark brown terga; however the mesosoma laterally and ventrally, and hind coxa of speciosus is mostly yellowish, compared with being almost entirely black in kingmani sp. n. The mummy produced by speciosus, although decapitated, is noticeably distinct from the ventrally bent "J-shaped" mummy of kingmani sp. n.
Color. Entire body yellowish brown to honey brown, darker dorsally; antenna basally dark brown, lightening gradually toward apex, apical 1/3 pale brown, scape light brown dorsally; face pale yellow, ocellar triangle black; lateral borders of mesoscutum, notauli and posterior depressed area brown; ovipositor sheaths mostly dark brown, basally whitish; wings slightly brown infuscate; veins dark brown except C+SC+R, stigma and R1 honey yellow.
Head. Antenna with 61 segments; flagellomeres about as long as wide, except apical 1/3 and basal 1/6 slightly longer than wide, apical flagellomere with small pointed apex; malar space as long as basal width of mandible, and 0.3× eye height; temple narrow, in dorsal view about eyes 5× longer than temples; occipital carina complete, reaching hypostomal carina; oral space small and circular, diameter about equal to basal width of mandible; clypeus weakly swollen; ocelli moderate, ocell-ocular distance about 1/2 diameter of lateral ocellus; face and gena rugose-costate, with midlongitudinal ridge just bellow toruli, frons smooth and excavated, bordered by weak "W-shaped" carina; temples and vertex granulate.
Mesosoma. Sculpturing mostly granular; pronotum with median scrobiculate line; mesopleuron mostly shining granular-coriaceous, antero-dorsal corner rugose, central elevated area sharply defined and smooth, epicnemial carina complete; propodeum on are the only two species in pulchripes species-group found so far from Yanayacu. A. stewarti sp. n. differs from colberti sp. n. in the antenna dark brown basally, gradually lightening toward pale brown apex (black with mid white band in colberti sp. n.), wings uniformly weakly infuscate (with dark band bellow stigma in colberti sp. n.), ocelli about 2× ocell-ocular distance (about 8× in colberti sp. n.); body mostly honey brown (reddish brown in colberti sp. n.), tarsal claw pectination with several short bristles extending to base of claw (in colberti sp. n. the pectination have less and larger bristles, and a distinct gap with claw base).
Comments. Since stewarti sp. n. is described based on several females and colberti sp. n. is described based on one male, and considering the geographical distribution and sexual dimorphism in the group, there is a possibility of these two species are one single species with very extreme sexual dimorphism. However, we do not think that it is likely because none of the known species in pulchripes species-group, having both males and females described, exhibit anything close to such extreme variation, which compels us to maintain these two entities as distinct species.
Etymology. This species is named after Jon Stewart (John Stuart Leibowitz), an American comedian, political satirist, writer, director, actor, and television host of The Daily Show.
Additional characters. Last flagellomere with "bottle-nipple"-shaped tip; mesoscutum with carina at posterior margin present only in front of scutellar sulcus; scutellar sulcus with three strong and short carina; fore wing vein 1M slightly curved at base; hind wing vein 2-1A present and relatively long, vein m-cu absent.
Distribution. Known only from the type locality, YBS, Napo province, ECUADOR. Discussion. Aleiodes stilpnos is the only species treated in this work in the albitibia species-group. The albitibia-group is relatively rarely collected in the Neotropical region, with only two described -A. stilpnos from Ecuador and Aleiodes fuscipennis (Szépligeti, 1904) from Peru, Venezuela (Torres and Briceño 2005) and Chile (deposited at UWIM), and at least two undescribed species from Costa Rica. A. stilpnos differs from other species in Ecuador by the following characters: head, mesosoma and legs mostly black contrasting with light orange metasoma, and wings infuscate; mesopleuron central disc smooth and bare; tarsal claws strongly pectinate; and costate sculpturing on metasomal terga 1-3.
Color. Head yellow, ocellar triangle dark brown; antenna dark brown-black, scapus lighter dorsally, extreme base of scapus and first flagellomere yellow; mesosoma yellow, anterior corner of mesopleuron, lunules, metanotum, propodeum and dorsal 1/4 of metapleuron dark brown, remainder metapleuron and mesopleuron, at border with metapleuron, white; latero-ventral mesopleuron with slightly lighter stripe; fore leg yellow, telotarsus brown; mid coxa, trochanter, trochantellus and femur basally white, remainder mid leg yellow, darkening toward apex, telotarsi and tibial spurs brown; hind leg: coxa black on 1/2 basal and white on 1/2 apical, trochanter and trochantellus black, but apical border of trochanter, and trochantellus ventral-apical 1/3 plus a small ventral spot white; femur black on basal 2/5 and at extreme apex, otherwise white; tibia and tarsi black, small white subbasal band on tibia. Metasoma white with black dorsal triangle beginning on central apex of T1 and covering most dorsal portion of subsequent terga; ovipositor sheaths dark brown on apical half, basal half whitish. Wings weakly infuscate; most veins and stigma dark brown; vein C+SC+R black (extreme base whitish) connecting to a black parastigma with whitish central spot.
Head. Antenna with 46 antennomeres, flagellomeres roughly 2.0× as long as wide, apical flagellomere with "bottle-nipple"-shaped apex; malar space as long as basal width of mandible, and 1/3 eye height; in dorsal view eye height 2.8× temple; occipital carina incomplete dorsally, curving toward lateral ocelli, laterally complete and meeting hypostomal carina; oral space small and circular, maximum width slightly smaller than basal width of mandible; clypeus not swollen; ocellus moderate, ocell-ocular distance 0.8× diameter of lateral ocellus; maxillary palp not swollen; head surface sculpturing finely granulate, occiput smooth and shining; higher face with some transverse wrinkles just bellow toruli.
Mesosoma. Sculpturing finely granulate; propodeum more coarsely granulate, with mid-longitudinal carina complete and some diverging wrinkles posteriorly; notauli weak, shallow and smooth; posterior margin of mesoscutum with complete carina; scutellar sulcus with median carina plus one pair of weak lateral carina. Wings Legs. Hind tibia without comb of modified setae; tarsal claw pectinate basally, with several very short and tight bristles, longer and more sparse apically, wide gap between apical claw and basal pecitnation; hind basitarsus about 3× longer than inner apical spur on hind tibia.
Paratype variation. Antenna broken, otherwise essentially as holotype. Male unknown. Mummy. Length 17.0 mm, entire mummy mottled with gray and brownish, thorax narrow and wrinkled, mummy withered posteriorly behind exit hole, mummy as- pect long and narrow, curved upward "V-shaped", exit hole irregular, located posterodorsally, anterior to prolegs.
Discussion. Aleiodes townsendi sp. n. belongs to the circumscriptus/gastritor species group. It is similar to A. shakirae sp. n. because of the presence of a strongly curved vein 1M in the fore wing, some similar color patterns, and the elongate and curved "V-shaped" mummy. It differs from shakirae sp. n. by having the metasomal tergite 1 mostly white with a small mid-apical black spot and about as long as apical width, entirely black or dark brown and about 2× longer than apical width in shakirae sp. n. The metapleuron in townsendi sp. n. is bicolored, black and white, while entirely black in shakirae sp. n. A. townsendi sp. n. hind coxa is black basally and apically white, compared with the inverse color pattern in shakirae sp. n., and the wings are moderately infuscate, while hyaline in shakirae sp. n. (additional features are cited in discussion section for A. shakirae sp. n.). The ocelli in A. townsendi sp. n. are relatively small, but its large eyes, almost 3× longer than temple in dorsal view, make the ocell-ocular distance shorter, the width of lateral ocellus being roughly equal to ocell-ocular distance.
Etymology. This species is named after Andrew Townsend, collector of one of the type specimens, for his contributions to the knowledge of the Ecuadorian Braconidae fauna.
Color. Dorsally dark brown to black, except head; laterally and ventrally mostly yellowish to whitish. Head honey yellow; mandibles, cheeks, and palp whitish, but teeth brown; black stain dorsally, from ocellar triangle through occiput mid-dorsally. Mesosoma black dorsally; pronotum and propleuron whitish, but mid-dorsally pronotum dark brown; mesopleuron honey yellow; metapleuron black. Legs mostly Biology. The holotype was reared from a mummified larva collected on Palicourea ulloana (Rubiaceae). The sampling date is listed in the database as May 12, 2010, and the adult emergence date is listed as May 5, 2010, so clearly one of these dates must be incorrect. Since this caterpillar record is nested within a large group of other caterpillar records also collected on May 12, 2010, it seems most likely that the emergence date was recorded incorrectly. Since the pupation date is assigned as May 18, 2010, this also corroborates that the emergence date could not possibly have been May 5, 2010. It seems most likely that the emergence month was recorded incorrectly and perhaps the actual emergence date was June 5, 2010. The holotype host is probably a Noctuidae due to similarity with the paratype mummy, which is possibly conspecific, identified as a Noctuidae. The host plant for the paratype is unknown, and it is not possible to determine if the holotype host caterpillar fed or not on P. ulloana.
Discussion. This species belongs to circumscriptus/gastritor species group. Aleiodes tzantza sp. n. is similar to A. atripileatus, from which it can be distinguished by the larger ocelli: ocell-ocular distance about half diameter of lateral ocelli (1.5 to 1.7× in A. atripileatus), and the color of mesopleuron entirely yellow (dark brown at least dorsally in A. atripileatus). A. tzantza sp. n. mummies are very similar to those of atripileatus, but the projecting anal prolegs are much longer in this species, and the body is distinctly longer. Its short ocell-ocullar distance is similar to A. speciosus, from which it differs by having the mesopleuron surface granular (mostly smooth with anterior quarter rugose in speciosus) and entirely honey yellow (anterior 1/4 black in speciosus), pronotum whitish laterally (mostly dark brown in speciosus), and metasomal terga all dark brown (first tergite white in speciosus).
Etymology. "Tzantza" is the Shuar word for the ritual of reducing heads by a mummification process used by the Shuar, a people native from the current Ecuadorian Amazon territory, resulting in a shrunken mummy as the ones produced by the Aleiodes species.
Color. Mostly black. Propodeum with mid-apical white mark; metapleuron mostly whitish, but brown dorsally. Metasoma with T1 and T2 pale light yellow, reminder terga dark brown. Fore legs mostly dark brown, trochanter whitish with brown mark on inner side; mid legs mostly dark brown, coxa, trochanter and trochantellus yellowish, femur mostly light brown; hind legs yellowish basally, apex of femur with brown stain dorsally, tibia and tarsi dark brown except for whitish stain covering basal half of inner side of tibia. Wings infuscate with dark brown veins and stigma.
Head. Antenna with 42 antennomeres, flagellomeres roughly 2.0× as long as wide, apical flagellomere with pointed tip; malar space about as long as basal width of mandible, and 0.4× eye height; in dorsal view eye 1.4× temple; occipital carina incomplete, interrupted at vertex, well defined laterally and meeting hypostomal carina; oral space small and circular, maximum width roughly equal to basal width of mandible; clypeus slightly swollen; ocell-ocular distance about 1.3× diameter of lateral ocellus; maxillary palp not swollen; head surface sculpturing granulate, occiput smooth and shining; frons slightly excavated, shining coriaceous, sculpturing concentrically arranged; face with some transverse rugae medially, and a short but well defined mid-longitudinal carina higher on face, carina extending dorsally between toruli.
Mesosoma. Sculpturing finely granulate; pronotum foveate; mesopleuron central disc smooth and bare, antero-dorsal corner rugose; metapleuron smooth; propodeum coarsely granular with complete mid-longitudinal carina and some longitudinal rugosity posteriorly;  notauli well defined and crenulate anteriorly, barely defined but traceable posteriorly, meeting a rugose area; posterior margin of mesoscutum bordered with complete carina; scutellar sulcus with median carina plus two pairs of incomplete lateral carina.
Biology. Reared from a Geometridae species, feeding on Phenax rugosus (Urticaceae). Three weeks elapsed from mummification until adult emergence.
Discussion. Aleiodes yanayacu sp. n. belongs to circumscriptus/gastritor speciesgroup. This species is similar to A. mirandae sp. n. and A. napo sp. n. in the mostly smooth mesopleuron, and the occipital carina, interrupted at vertex. A. yanayacu sp. n. differs from both species in having a slightly larger ocellus, the ocell-ocular distance is about 1.3× the diameter of lateral ocellus, while in mirandae sp. n. and napo sp. n. it is about 2.0×. The hind wing vein M+CU is about as long as vein 1M in yanayacu sp. n., but distinctly shorter in mirandae sp. n. and napo sp. n. The sculpturing of metasomal terga 1 and 2 is also distinctive in yanayacu sp. n., with widely spaced costa, as compared with the finely rugose-costate sculpturing in mirandae sp. n. and napo sp. n. The metasomal tegite 2 is entirely withish in yanayacu sp. n. (black and white in mirandae sp. n. and napo sp. n.) and the hind coxa yellowish (black mirandae sp. n. and napo sp. n.).
Etymology. The species is named after the sampling and rearing location, theYanayacu Biological Station.