Seven new species of spider-attacking Hymenoepimecis Viereck (Hymenoptera, Ichneumonidae, Pimplinae) from Ecuador, French Guiana, and Peru, with an identification key to the world species

Abstract Seven new species of Hymenoepimecis Viereck are described from Peruvian Andes and Amazonia, French Guiana and Ecuador: H. andina Pádua & Sääksjärvi, sp. nov., H. castilloi Pádua & Sääksjärvi, sp. nov., H. dolichocarinata Pádua & Sääksjärvi, sp. nov., H. ecuatoriana Pádua & Sääksjärvi, sp. nov., H. longilobus Pádua & Sääksjärvi, sp. nov., H. pucallpina Pádua & Sääksjärvi, sp. nov., and H. rafaelmartinezi Pádua & Sääksjärvi, sp. nov. In addition, the male of the Hymenoepimecis kleini Pádua & Sobczak, 2015 is described, new faunistic records for the genus provided, as well as an illustrated identification key to all known species of the genus.


Introduction
Hymenoepimecis Viereck, 1912 is a moderately large genus of spider-attacking Darwin wasps. It is confined to the New World where it occurs from Mexico and Central America to Southern Brazil (Gauld 1991;Kumagai and Graf 2002;Yu et al. 2012;. The species of Hymenoepimecis are known to be koinobiont ectoparasitoids attacking sub-adult and adult orb-web-spinning spiders belonging to the families Araneidae and Tetragnathidae (summarised in Pádua et al. 2016). Hymenoepimecis is closely related to Acrotaphus Townes and Ticapimpla Gauld and it can be easily separated from these two genera by possessing a pocket-like structure on the mediodorsal part of the pronotum (Gauld 1991).
Currently, there are 20 described species in the genus (Yu et al. 2012; and the majority of them have been discovered from Brazil (Brullé 1846;Kriechbaumer 1890;Loffredo and Penteado-Dias 2009;Sobczak et al. 2009; and Central America (see Cresson 1865;Brues and Richardson 1913;Townes and Townes 1966;Gauld 1991Gauld , 2000. Very little is known about the occurrence of the genus in other Latin American countries. The main aim of this work is to describe seven new species of Hymenoepimecis from Peru, French Guiana and Ecuador. In addition, we provide new faunistic records for the genus and an illustrated identification key to all species.

Materials and methods
The specimens studied in this work are deposited at the Florida State Collection of Arthropods (FSCA), Gainesville, USA, The Natural History Museum, University of San Marcos (MUSM), Lima, Peru, The Natural History Museum, London, UK (NHM), and the Biodiversity Unit, Zoological Museum, University of Turku (ZMUT), Turku, Finland. The specimens of MUSM are currently on loan in ZMUT.
Specimens were examined with an Olympus SZ61 and SZX10 stereomicroscopes and measurements were made through a millimetre ocular. Digital layer images were taken using a Canon DS126461 digital camera attached to an Olympus SZX16 stereomicroscope and combined by using the software Zerene Stacker (Version 1.04 Build T201706041920). Drawings were vectorized digitally using Adobe Illustrator.
The morphological terminology and style of descriptions follow those of . In this study, the measures and proportions between the structures are given as the value of the holotype [in brackets], followed by the minimum and maximum number of variations. The new country records of the genus are marked with a double asterisk and new country records of the species were marked with a single asterisk. The maps were produced through the website Simplemappr (https://www. simplemappr.net).

9
Mesosoma entirely orange (Fig. 34); metasoma orange, with posterior margins of tergites II-IV narrowly black, tergites V+ black (Fig. 64) Gauld, 1991 10 Sternite with a high, nasute ventral protuberance (Fig. 47)  Fore wing yellowish hyaline, with black apex (Fig. 62)  Metasoma blackish or darkish brown, some tergites with whitish anterior margin (e.g., Fig. 61) or metasoma orange, some tergites with posterior margin blackish (Fig. 69)  Occipital carina reduced in dorsal part (Fig. 43); metasoma orange, with tergites VI+ black (Fig. 73) -6, 8-11, 13, 14); head with occipital carina projected and curved upwards, without a concavity in the apex dorsal (23)(24)(25)(26)28,29); pronotum with the pocket-like structure not reduced longitudinally (38)(39)(40)(41)43,44); sternite I with a low, rounded swelling posteriorly (38)(39)(40)(41)43,44) or with a high, laterally compressed, nasute ventral protuberance (Fig. 47)  Comments. According to Gauld (1991Gauld ( , 2000 Hymenoepimecis is very similar with the sister genus Acrotaphus Townes, 1960. They both have the occipital carina strongly raised (flange-like) and projecting posteriorly to surround the anterior reflexed end of the pronotum; head rounded with the genae strongly narrowed from the eyes to the occipital flange; and the pronotum unusually elongated, with a long horizontal part mediodorsally. It differs from Acrotaphus by having a unique forwardly directed pocket-like structure on the pronotum mediodorsally. Diagnosis. This species can be distinguished from all other Hymenoepimecis by the combination of the following characters: 1) head, in dorsal view, with gena short and slightly convex behind eyes; 2) posterior ocelli separated from eyes by 0.7-0.8 times its own maximum diameter; 3) occipital carina only slightly projected and not curved upwards dorsally; 4) ovipositor 1.1-1.2 times as long as hind tibia; 5) metasoma darkish brown, with whitish lateral marks on anterior margins of tergites II-V.  8] times as broad as high, smooth, slightly convex with few spaced bristles laterally; head in dorsal view, with gena short and slightly convex behind eyes; posterior ocelli separated from eyes by approx. [0.7] 0.7-0.8 times its own maximum diameter; occipital carina lit-tle projected and not curved upwards dorsally. Pronotum more or less long, smooth and polished, with distance from tegula to head greater than [0.5] 0.5-0.6 times distance from tegula to hind margin of propodeum, and in anterior part with opening pocket-like structure not reduced longitudinally; mesoscutum smooth and polished; scutellum, in profile, convex; mesopleuron smooth and polished, with anterodorsal and posterodorsal parts bearing sparse, fine setiferous punctures; metapleuron smooth and polished, with few sparse, fine setiferous punctures; propodeum smooth, polished, with sparse, fine setiferous punctures and with lateral longitudinal carina present only posteriorly. Fore wing approx. [6.0] 6.0-7.0 mm; cu-a more or less interstitial to the base of Rs&M; 2rs-m approx. [0.3] 0.3-0.5 times as long as abscissa of M between 2rs-m and 2m-cu; hind wing with abscissa of Cu1 meet cu-a equidistant between 1A and M. Hind leg with tibia + tarsus [0.6] times fore wing length; tarsal claw with more or less square basal lobe with apex of claw slightly overtaking the lobe. Metasoma slender; tergite I [1.7] 1.6-1.7 times as long as posteriorly width, centrally quite strongly convex with lateral carinae present only at extreme anterior end flanking the anterior concavity; sternite I with a low, rounded swelling posteriorly; tergite II approx. [1.4] 1.3-1.4 times as long as posteriorly width; tergites III and IV approx. [1.2] 1.2-1.3 times as long as posteriorly width; ovipositor [1.1] 1.1-1.2 times as long as hind tibia.

Hymenoepimecis andina
Colour. Head black with apical margin of clypeus and mouthparts (except apex mandible black) yellowish; antenna blackish. Mesosoma orange. Fore and mid leg orange, the hind leg black. Wings hyaline slightly darkish; pterostigma brownish. Metasoma darkish brown, with whitish lateral marks on anterior margins of tergites II-V; ovipositor brownish with base and apex whitish, sheath blackish.
Male. (Fig. 97). Similar to female in structure and colouration, but with body with 6.0-8.0 mm; face approx.1.1 times as width as high; posterior ocelli separated from eyes by 0.7-0.9 times its own maximum diameter. Fore wing 4.5-6.5 mm. Tarsal claw simple. Metasoma slender; tergite I 1.6 times as long as posteriorly width; tergite II approx.1.4 times as long as posteriorly width; tergites III and IV 1.2-1.3 times as long as posteriorly width.
Distribution. Peru (Andes) (Fig. 107). Biological notes. Host unknown. Etymology. The specific name refers the locality of the specie type, in the Peruvian Andes, Cusco Peru.

Hymenoepimecis bicolor
Description. Female. Body approx. [6.0] mm; face [0.9] times as broad as high (from supraclypeal suture to base of antenna), smooth, flat with few spaced bristles laterally; head in dorsal view, with gena short and slightly convex behind eyes; posterior ocelli separated from eyes by [0.7] times its own maximum diameter; occipital carina little projected and not curved upwards dorsally. Pronotum as long as high, smooth and polished, with distance from tegula to head greater than approx. [0.7] times distance from tegula to hind margin of propodeum, and in anterior part with opening pocket-like structure not reduced longitudinally; mesoscutum smooth and polished; scutellum, in profile, convex; mesopleuron smooth and polished, with posterodorsal part bearing sparse, fine setiferous punctures; metapleuron smooth and polished, with a few sparse, fine setiferous punctures; propodeum smooth, polished, with sparse, fine setiferous punctures and with lateral longitudinal carina present only posteriorly. Fore wing [5.0] mm; cu-a more or less interstitial to the base of Rs&M; 2rs-m approx. [0.3] times as long as abscissa of M between 2rs-m and 2m-cu; hind wing with abscissa of Cu1 meet cu-a equidistant between 1A and M. Hind leg with tibia + tarsus approx. [0.6] times fore wing length; tarsal claw with more or less square basal lobe with apex of claw slightly overtaking the lobe. Metasoma slender; tergite I [2.0] times as long as posteriorly width, centrally quite strongly convex with lateral carinae present only at extreme anterior end flanking the anterior concavity; sternite I with a low, rounded swelling posteriorly; tergite II approx.
Etymology. This species is named in honour of Carol Castillo, collector of the type specimen. Carol is a Peruvian entomologist and has studied Darwin wasps in the tropical Andes. She has discovered and/or described a large number of new species.
Description. Female. Body approx. [13.0] 11.5-13.0 mm; face [1.0] 0.9-1.0 times as broad as high, smooth, slightly convex with few spaced bristles; head in dorsal view, with margin of gena flat behind eyes; posterior ocelli separated from eyes by approx. [0.6] 0.5-0.6 times its own maximum diameter; occipital carina little projected, slightly curved upwards in the mediodorsal part. Pronotum more or less long, smooth and polished, with distance from tegula to head greater than [0.6] 0.5-0.6 times distance from tegula to hind margin of propodeum, and in anterior part with opening pocket-like structure not reduced longitudinally; mesoscutum smooth and polished; scutellum, in profile, convex; mesopleuron smooth and polished, with anterodorsal and posterodorsal parts bearing sparse, fine setiferous punctures; epicnemial carina present ventrally, extending until reaching the level of lower corner of pronotum laterally; metapleuron smooth and polished, rather uniformly covered with sparse, fine setiferous punctures. Fore wing approx. [10.0] 9.0-11.0 mm; cu-a interstitial to the base Rs&M; 2rs-m approx. [0.6] times as long as abscissa of M between 2rs-m and 2m-cu; hind wing with abscissa of Cu1 meeting cu-a equidistant between M and 1A. Hind leg with tibia + tarsus [0.6] 0.55-0.6 times the fore wing length; tarsal claw with more or less square lobe, with apex slightly overtaking the lobe. Metasoma slender; tergite I [1.6] 1.5-1.6 times as long as posteriorly width, centrally quite strongly convex with lateral carinae present only at extreme anterior end flanking the anterior concavity; sternite I with a low, rounded swelling posteriorly; Male. (Fig. 99). Similar to female in structure and colouration, but with body with 5.0-7.0 mm; face 0.9-1.1 times as broad as high; head in dorsal view, with margin of gena short, slightly convex behind eyes; posterior ocelli separated from eyes by approx. 0.8 times its own maximum diameter; occipital carina not projected. Fore wing 4.0-6.0 mm; cu-a more or less interstitial to the base Rs&M; 2rs-m 0.3-0.4 times as long as abscissa of M between 2rs-m and 2m-cu. Tarsal claw simple. Metasoma slender; tergite I 1.6-1.7 times as long as posteriorly width; tergite II approx. 1.3 times as long as posteriorly width; tergites III and IV 1.2 times as long as posteriorly width.
Variation. Tergite V black with anterior margin orange or apical half orange and basal half-black. Three females from French Guiana (Saül city) presented margin of gena slightly convex. We think they are conspecific, but we are not treating them as paratypes.
Distribution. Ecuador**, French Guiana and Peru (Fig. 107) Diagnosis. This species can be distinguished from all other Hymenoepimecis by the combination of the following characters: 1) fore wing hyaline yellowish, with two blackish bands; 2) pronotum orange; 3) female with tarsal claw with more or less square lobe, and with apex overtaking the lobe; 4) female with ovipositor 1.2 times as long as hind tibia.
Description. Female. Body [9.0] mm; face [1.0] times as broad as high, smooth, slightly convex with few spaced bristles; head in dorsal view, with gena strongly narrowed behind eyes; posterior ocelli separated from eyes by approx. [1.0] times its own maximum diameter; occipital carina projected and weakly reduced and curved upwards dorsally. Pronotum long, smooth and polished, with distance from tegula to head approx. [0.8] times distance from tegula to hind margin of propodeum, and in anterior part with opening pocket-like structure not reduced longitudinally; mesoscutum smooth and polished; scutellum, in profile, convex; mesopleuron smooth and polished, with anterodorsal and posterodorsal parts bearing sparse, fine setiferous punctures; metapleuron smooth and polished, rather uniformly covered with sparse, fine setiferous punctures; propodeum smooth, polished, with sparse, fine setiferous punctures and with lateral longitudinal carina present only posteriorly. Fore wing [7.0] mm; cu-a more or less interstitial to the base of Rs&M; 2rs-m [0.5] times as long as abscissa of M between 2rs-m and 2m-cu; hind wing with abscissa of Cu1 meeting cu-a equidistant between M and 1A. Hind leg with tibia + tarsus [0.6] times the fore wing length; tarsal claw with more or less square lobe, with apex of claw overtaking the lobe. Metasoma slender; tergite I [1.4] times as long as posteriorly width, centrally quite strongly convex with lateral carinae present only at extreme anterior end flanking the anterior concavity; sternite I with a low, rounded swelling posteriorly; tergite II approx. Colour. Head black; clypeus black with apex yellowish; mouthparts yellowish, with apex mandible black; antenna brownish. Mesosoma orange. Fore and mid leg orange, the hind entirely blackish brown. Fore wing hyaline yellowish, with apex blackish and with a blackish preapical band; pterostigma with basal half black and apical half yellow; hind wing with slightly blackish band in median part. Metasoma orange, with tergites VI+ black; ovipositor and sheath brownish.

Hymenoepimecis kleini
Male. (Fig. 101). Similar to female in structure and colouration, but with hind leg black, except coxa, with basal part orange; body approx. 8.0 mm; face a approx. 1.1 times as broad as high; posterior ocelli separated from eyes by approx. 0.8 times its own maximum diameter. Fore wing approx. 6.0 mm; 2rs-m approx. 0.2 times as long as abscissa of M between 2rs-m and 2m-cu. Tarsal claw simple. Metasoma slender; tergite I approx. 1.6 times as long as posteriorly width; tergite II approx. 1.4 times as long as posteriorly width; tergites III and IV approx. 1.3 times as long as posteriorly width.
Distribution. Brazil, Ecuador** and Peru* (Fig. 107).  Figures 8,23,38,53,68,83,102 Diagnosis. This species can be distinguished from all other Hymenoepimecis by the combination of the following characters: 1) fore wing hyaline yellowish, with two blackish bands; 2) pronotum orange (Peruvian specimens with anterior margin of the pronotum blackish); 3) female with tarsal claw with a longitudinally elongated lobe, and with apex overtaking the lobe; 4) female with ovipositor 1.5-1.6 times as long as hind tibia.
Description. Female. Body approx. [15.0] 12.0-15.0 mm; face [1.1] 1.0-1.1 times as broad as high, smooth, slightly convex with few spaced bristles; head in dorsal view, with gena strongly narrowed behind eyes; posterior ocelli separated from eyes by approx. [0.9] 0.8-1.0 times its own maximum diameter; occipital carina projected and curved upwards dorsally. Pronotum long, smooth and polished, with distance from tegula to head greater than approx. [0.7] 0.6-0.7 times distance from tegula to hind margin of propodeum, and in anterior part with opening pocket-like structure not reduced longitudinally; mesoscutum smooth and polished; scutellum, in profile, convex; mesopleuron smooth and polished, with anterodorsal and posterodorsal parts bearing sparse, fine setiferous punctures; metapleuron smooth and polished, rather uniformly covered with sparse, fine setiferous punctures; propodeum smooth, polished, with sparse, fine setiferous punctures and with lateral longitudinal carina present only posteriorly. Fore wing approx. [13.0] 9.0-13.0 mm; cu-a interstitial to the base of  Colour. Head black; clypeus black with apex yellowish; mouthparts yellowish, with apex mandible black; antenna brownish. Mesosoma orange. Fore and mid leg orange, the hind entirely blackish brown except base of coxa orange. Fore wing hyaline yellowish, with apex blackish and with a blackish preapical band; pterostigma with basal half black and apical half yellow; hind wing with slightly blackish band in median part. Metasoma orange, with posterior margin black in tergite V and tergites VI+ black; ovipositor and sheath brownish.
Male. (Fig. 102). Similar to female in structure and colouration, but with hind coxa orange body with 10.5-12.0 mm; face 1.0 times as broad as high; posterior ocelli separated from eyes by 0.8-1.0 times its own maximum diameter. Fore wing 8.0-10.0 mm; 2rs-m 0.3-0.5 times as long as abscissa of M between 2rs-m and 2m-cu. Tarsal claw simple. Metasoma slender; tergite I 1.6-1.7 times as long as posteriorly width; tergite II 1.2-1.5 times as long as posteriorly width; tergites III and IV 1.0-1.3 times as long as posteriorly width.
Variation. The Peruvian specimens have anterior margin of pronotum blackish. Some specimens have tergite V entirely black or black with anterior margin orange; hind coxa orange with basal part blackish.
Description. Female. Body approx. [16.0] mm; face [1.0] times as broad as high, smooth, slightly convex with few spaced bristles; head in dorsal view, with gena slightly narrowed behind eyes; posterior ocelli separated from eyes by approx. [1.2] times its own maximum diameter; occipital carina projected and curved upwards dorsally. Pronotum long, smooth and polished, with distance from tegula to head greater than [0.6] times distance from tegula to hind margin of propodeum, and in anterior part with opening pocket-like structure not reduced longitudinally; mesoscutum smooth and polished; scutellum, in profile, convex; mesopleuron smooth and polished, with anterodorsal and posterodorsal parts bearing sparse, fine setiferous punctures; metapleuron smooth and polished, with a few sparse, fine setiferous punctures; propodeum smooth, polished, with sparse, fine setiferous punctures and with lateral longitudinal carina present only posteriorly. Fore wing approx. [13.0] mm; cu-a interstitial to the base of Rs&M; 2rs-m approx. [0.75] times as long as abscissa of M between 2rs-m and 2m-cu; hind wing with abscissa of Cu1 meet cu-a equidistant between 1A and M. Hind leg with tibia + tarsus [0.6] times the fore wing length; tarsal claw with more or less square basal lobe with apex of claw overtaking the lobe. Metasoma slender; tergite I [1.6] times as long as posteriorly width, centrally quite strongly convex with lateral carinae present only at extreme anterior end flanking the anterior concavity; sternite I with a low, rounded swelling posteriorly; tergite II approx. [1.3] times as long as posteriorly width; tergites III-IV approx. [1.2] times as long as posteriorly width; ovipositor [1.6] times as long as hind tibia.
Colour. Head black with apical margin of clypeus and mouthparts (except apex mandible black) yellowish; antenna blackish. Mesosoma entirely orange. Fore and mid leg orange, the hind leg black. Fore wing black, with basal region yellowish and with a yellowish band between junction of vein R1 up to pterostigma until half vein M; pterostigma black, except apical margin yellowish; hind wing black with basal region and apex yellowish. Metasoma entirely black; ovipositor brownish with apex reddish brown, sheath blackish.
Male. Unknown. Distribution. Peru (Fig. 109) Diagnosis. This species can be distinguished from all other Hymenoepimecis by the combination of the following characters: 1) fore wing hyaline yellowish, with two blackish bands; 2) face sculptured below the insertion of antennae, with a longitudinal carina in the middle part; 3) occipital carina projected, not curved upwards, with a concavity in the apex dorsally; 4) pronotum with the pocket-like structure reduced longitudinally; 5) sternite I with a ventral spine-like projection posteriorly; 6) metasoma orange, with tergites VI+ black; 7) hind leg orange, with femur, tibia and tarsus back; 8) female with tarsal claw with basal lobe more or less square and apex of claw overtaking the lobe; 9) female with ovipositor 1.0-1.2 times as long as hind tibia.
Description. Female. Body approx. [10.0] 9.0-10.0 mm; face [0.9] 0.8-1.1 times as broad as high, sculptured below the insertion of antennas, with longitudinal carina in the middle part and with few bristles spaced on the lower face; head in dorsal view with gena strongly narrowed behind eyes; posterior ocelli separated from eyes by [1.0] 0.8-1.0 times its own maximum diameter; occipital carina not curved upwards, with a concavity in the apex dorsally. Pronotum long, smooth and polished, with distance from tegula to head greater than [0.5] 0.5-0.6 times distance from tegula to hind margin of propodeum, and in anterior part with opening pocket-like structure reduced longitudinally; mesoscutum smooth and polished; scutellum, in profile, convex; mesopleuron smooth and polished, with anterodorsal and posterodorsal parts bearing sparse, fine setiferous punctures; metapleuron smooth and polished, rather uniformly covered with sparse, fine setiferous punctures; propodeum smooth, polished, with sparse, fine setiferous punctures and with lateral longitudinal carina present only posteriorly. Fore wing approx. [7.0] 7.0-8.0 mm; cu-a interstitial to the base of Rs&M; 2rs-m [0.3] 0.3-0.5 times as long as abscissa of M between 2rs-m and 2m-cu; abscissa of Cu1 meeting 1m-cu equidistant between Cu1a and Cu1b; hind wing with abscissa of Cu1 meeting cu-a closer to 1A that M. Hind leg with tibia + tarsus [0.55] 0.5-0.6 times the fore wing length; tarsal claw with basal lobe slightly more or less square, with apex of claw overtaking the lobe. Metasoma slender; tergite I approx. [1.2] 1.2-1.4 times as long as posteriorly width, centrally quite strongly convex with lateral carinae present only at extreme anterior end flanking the anterior concavity; sternite I with Colour. Head black; clypeus with apical margin yellowish, labrum and mouthparts yellowish, except apex mandible black; antenna brown. Mesosoma orange. Fore and mid leg orange, the hind leg orange, with femur, tibia and tarsus black. Fore wing bristles spaced on the lower face, and by sternite I with a ventral spine-like projection posteriorly. It differs from H. uberensis mainly by having the fore wing hyaline yellowish with two black bands and hind leg orange, with femur, tibia and tarsus black (fore wing blackish, with yellowish hyaline band between junction of vein R1 up to pterostigma until half vein M and hind leg entirely black in H. uberensis). Pádua & Sobczak, 2015 Figures 13, 28, 43, 58, 73, 88, 105 Hymenoepimecis ribeiroi Pádua & Sobczak, 2015: 188. Diagnosis. See .

Discussion
The present study increases the number of recognized Hymenoepimecis species to 27 species. In addition, is presented a large amount of faunistic records from different parts of Central and South America. It is also shown that H. rafaelmartinezi sp. nov. belongs to the H. jordanensis species group  which is characterised by the following set of characters: 1) face sculptured below the insertion of antennae, with a longitudinal carina in the middle part of the face; 2) head with occipital carina projected, not curved upwards, with a dorsal concavity in the apex; 3) pronotum with the pocket-like structure reduced longitudinally; 4) sternite I with a ventral spine-like projection posteriorly. After the present study this species group is composed of five species (H. amazonensis, H. jordanensis, H. kleini, H. rafaelmartinezi sp. nov., and H. uberensis), A noteworthy new character state was recorded for Hymenoepimecis: in H. dolichocarinata sp. nov., the epicnemial carina is ventrally present and extending to the level of the lower lateral corner of the pronotum. This character state is shared with most species of Acrotaphus (Pádua et al. 2020), and likely represents a homoplastic trait in the two genera.
Finally, the genus is reported for the first time from the Andes (Figs 107, 108). Two Peruvian species, both collected by Carol Castillo, were found from the Andean-Amazonian interface (Cusco, San Pedro, Cosñipata valley, 1302-1520 m). Hymenoepimecis andina sp. nov. and H. castilloi sp. nov. are both remarkable because they resemble Polysphincta Gravenhorst (and Ticapimpla amazonica Palacio, Broad, Sääksjärvi & Veijalainen, 2010) by the shape of the occiput. All other species of Hymenoepimecis have the occipital carina strongly raised and produced into a neck-like structure.