Six new species of Handaoia Seyrig, 1952 (Hymenoptera, Ichneumonidae, Phygadeuontinae): the first to be described from the New World

Handaoia Seyrig, 1952 is a small genus of Phygadeuontinae currently represented by eleven described species from Madagascar, Tanzania and Europe, and can be recognized by the combination of the distally expanded and ventrally flattened antennal flagellum, complete posterior transverse carina of the mesosternum, isolated ‘pit’ (episternal scrobe) in the mesopleuron, and a single bulla in fore wing vein 2m-cu. Most species have a distinctive combined area basalis and area superomedia on the propodeum. The following six new species from Central and South America are described and illustrated: H. cuscoensis Bordera sp. nov. from Peru, H. fritzi sp. nov. from Brazil, H. mercedensis Bordera sp. nov. from Peru, H. plaumanni sp. nov. from Brazil, H. ruizcancinoi Bordera sp. nov. from Mexico, and H. urceus sp. nov. from Brazil. A key to the New World species is provided.


Introduction
is a small genus of Phygadeuontinae Förster, 1869 (sensu Santos 2017), currently represented by eleven described species (Yu et al. 2016): seven from Madagascar (Seyrig 1952), one from Tanzania (Szépligeti 1908) and two from Central and Northern Europe (Horstmann 1979(Horstmann , 1981. Undescribed species have also been reported from the Philippines, Japan, South Africa, Peru (Townes 1970) and Australia (Gauld 1984). Townes (1970) classified Handaoia in the subtribe Chiroticina Townes, 1970 (Cryptinae, Phygadeuontini). Nothing is known about the biology of any species of Handaoia, although by analogy with other Phygadeuontinae, they can be expected to be idiobiont ectoparasitoids of larvae or pupae of holometabolous insects.
While studying specimens from three collections with extensive Neotropical Phygadeuontinae holdings, the Natural History Museum (London, UK), American Entomological Institute (Utah, USA) and Colección Entomológica de la Universidad de Alicante (Spain), we discovered six new species of Handaoia from Central and South America. As the genus had previously been reported only from Peru within the Neotropical region (Townes 1970), and only represented by undescribed species, these specimens significantly extend the known distribution and diversity of Handaoia. The aim of this paper is to describe and illustrate these new species, and to provide a key to the New World.

Material and methods
Material deposited in the following institutions has been studied: AEIC = American Entomological Institute Collection, Utah, USA CEUA = Colección Entomológica de la Universidad de Alicante, Alicante, Spain NHMUK = Natural History Museum, London, UK Morphological terminology follows Broad et al. (2018) with surface microsculpture descriptions based on Eady (1968). Layer (extended focus) photographs of H. fritzi sp. nov., H. plaumanni sp, nov. and H. urceus sp. nov. were taken at NHMUK using a Canon EOS 5DSR digital camera with a Canon MP-E 65 mm Macro Lens attached to a StackShot Macro Rail system controlled by Helicon Remote and Helicon Focus ver. 3.6.6W. Layer photos of H. cuscoensis Bordera sp. nov., H. mercedensis Bordera sp. nov. and H. ruizcancinoi Bordera sp. nov. were taken at the University of Alicante using a Canon EOS 70D digital camera with the same macro lens and system.

Diagnosis
Handaoia can be recognized and separated from other Phygadeuontinae genera by the combination of the distally expanded and ventrally flattened antennal flagellum, complete posterior transverse carina of the mesosternum, isolated ʻpit' (episternal scrobe) in the mesopleuron, and a single bulla in fore wing vein 2m-cu. The following more complete diagnosis is modified from Townes (1970), who included Handaoia in a key to the genera of 'Chiroticina'. Townes' definition applies equally well to the Afrotropical species originally included as well as to the Neotropical and Palaearctic species: flagellum of female antenna fusiform (expanded beyond the middle and flattened ventrally) with a median white ring; genal carina reaching base of mandible; mandibular teeth subequal; maxillary palp long, reaching beyond centre of mesosternum; pronotum with two dorsal pits separated by weak longitudinal carina, usually flanked laterally by 3-4 longitudinal carinae; epomia absent; median lobe of mesoscutum without median longitudinal groove; scutellum without lateral carinae; mesopleural impression (episternal scrobe) ventral to speculum as an isolated pit, some distance from mesopleural furrow; posterior transverse carina of mesosternum complete; fore wing with areolet pentagonal, its outer side open (vein 3rs-m missing), vein 2m-cu with one wide bulla; area superomedia always separated from area petiolaris and usually confluent with area basalis; laterotergites of metasomal tergites 2 and 3 not separated or only weakly separated from tergite. Small species, body length 2-5 mm.

Etymology
The name of the species refers to the Department of Cusco (Peru), the type locality region.

Material examined
Only known from the holotype female.

Etymology
This species is named in honour of Dr Fritz Plaumann, illustrious botanist and entomologist based in Brazil, who collected the type material. A noun in genitive case.

Material examined
Known from five females.

Diagnosis
Handaoia mercedensis Bordera sp. nov. can be distinguished from all other New World species by the combination of the following characters: apophysis of propodeum conspicuously elevated, parallelsided with blunt tip (Fig. 4G, arrow); fore wing with well-defined transverse dark brown band (Fig. 4A); juxtacoxal carina strong, irregular, joining submetapleural carina at mid length (Fig. 4F, arrow); flagellum dark brown (except white band), with first and second flagellomeres light brown ( Fig. 4A-B).

Etymology
The name refers to La Merced, municipality of Chanchamayo in the Department of Junin (Peru), where the holotype was collected.
MesosoMa. Finely granulate and matt (Fig. 4C-D); mesoscutum slightly convex, notauli absent (Fig. 4C); scutellum moderately convex, without lateral carinae; sternaulus deep, exceeding half the length of mesopleuron (Fig. 4D); juxtacoxal carina strong, irregular, joining submetapleural carina at mid length (Fig. 4F, arrow); propodeum (Fig. 1C) with strong and conspicuous carinae; area superomedia confluent with area basalis and separated from area petiolaris; inner side of area externa about 4.7 × as long as inner side of area dentipara; inner side of area dentipara about 0.3 × length of outer side; area spiracularis confluent with area lateralis; posterior transverse carina strong, forming an elevate apophysis joining lateral longitudinal carina, apophysis parallel-sided with blut tip (Fig. 4G, arrow). Length of hind femur about 4.0 × its height. Hind wing with vein cu-a 0.5 as long as abscissa of CU between M and cu-a.

Diagnosis
Handaoia plaumanni sp.nov. can be distinguished from all other New World species by the combination of the following characters (both male and female): apophysis of propodeum as a low transverse crest (Fig. 5I, arrow); sternaulus very short and weak, reaching at most 0.3 × the length of the mesopleuron (Figs 5E, 6D); juxtacoxal carina absent (Figs 5H, 6E); area externa of propodeum confluent with area dentipara (Figs 1D, 6F).

Etymology
This species is named in honour of Dr Fritz Plaumann, illustrious botanist and entomologist based in Brazil, who collected the type series. A noun in the genitive case.

Material examined
Known from three females and two males.
Otherwise similar to female.

Diagnosis
Handaoia ruizcancinoi Bordera sp. nov. can be distinguished from all other New World species by the combination of the following characters: apophysis of propodeum conspicuously high, subtriangular with blunt tip (Fig. 7H, arrow); juxtacoxal carina weak but complete, joining submetapleural carina at its anterior part (Fig. 7G, arrow); inner side of area externa about 7.0 × as long as inner side of area dentipara (Fig. 1E).

Etymology
The species is named in honour of Dr Enrique Ruiz Cancino from Universidad Autónoma de Tamaulipas (Mexico), in recognition of his contribution to the study of the Ichneumonidae of Mexico. A noun in the genitive case.
MesosoMa. Finely granulate and matt (Fig. 7A, C-D); mesoscutum slightly convex, almost flat posteriorly; notauli moderately deep over 0.7 × length of mesoscutum (Fig. 7D); scutellum moderately convex, without lateral carinae; sternaulus as a wide V-shaped depression anteriorly, reaching about 0.3 × the length of mesopleuron (Fig. 7C); juxtacoxal carina weak but complete, joining submetapleural carina at its anterior part (Fig. 7G, arrow); propodeum (Figs 1E, 7C, H) with strong and conspicuous carinae; area superomedia confluent with area basalis and separated from area petiolaris; inner side of area externa about 7.0 × as long as inner side of area dentipara; inner side of area dentipara about 0.3 × length of outer side; area spiracularis confluent with area lateralis; posterior transverse carina strong, forming an elevated apophysis joining lateral longitudinal carina, apophysis subtriangular with blunt tip (Fig. 7C, H, arrow). Length of hind femur 3.9 × its height. Hind wing with vein cu-a 0.77 × as long as abscissa of CU between M and cu-a.

Etymology
From the Latin for 'pitcher', named after the flask-or pitcher-shaped combined area basalis + superomedia. A noun in apposition.

Male
Unknown.

Discussion
This study significantly improves our knowledge of Neotropical Handaoia, previously only known from two undescribed Peruvian species (Townes 1970), to the recognition of six new species from widely scattered locations in Brazil, Mexico and Peru. However, there are undoubtedly more species to be discovered. There are many Neotropical phygadeuontines in collections that have yet to be identified to genus; sorting and identifying specimens is obviously essential to make these accessible, but this is a task that requires available expertise in ichneumonid identification, which is often a severely limiting factor.
The classification of Handaoia requires further discussion. Although Townes (1970) included Handaoia in his 'Chiroticina' subtribe of what is now the subfamily Phygadeuontinae (Santos, 2017), the few phylogenetic analyses of Phygadeuontinae have found that the chiroticine genera do not form a clade (Laurenne et al. 2006;Quicke et al. 2009;Santos 2017). In fact, there is little evidence for the monophyly of most subtribes in Phygadeuontinae or Cryptinae Kirby, 1837. In a combined morphological and molecular analysis (although with all Phygadeuontinae coded identically), Quicke et al. (2009) found that Handaoia and also Austriteles Gauld, 1984 formed a clade with Ateleute Förster, 1869 and Tamaulipeca Kasparyan, 2000, two of the three genera comprising the recently recognised Ateleutinae Townes, 1970(Santos 2017Santos et al. 2018). Could these genera be better classified in Ateleutinae? The morphological evidence is weak: Handaoia shares the complete posterior carina of the mesosternum and a single large bulla in fore wing vein 2m-cu with Ateleutinae; Austriteles shares the single bulla and a first metasomal segment with the spiracle at about mid-length and the sternite ending anterior to the spiracle. There are also numerous differences between the currently included genera within the Ateleutinae and both Handaoia and Austriteles. It would be well worth sequencing additional genes (Quicke et al. 2009 only sequenced one gene) and more thoroughly evaluating the potential synapomorphies of an expanded Ateleutinae. We also take the opportunity to note that the Australian species that Gauld (1984) assigned to Handaoia differ markedly from other Handaoia: the propodeum of the Australian species has the transverse carinae strong, with the area superomedia basically absent, and the dorsal pronotal groove simple. These might be better reclassified in another genus.
We have not examined specimens of other species of Handaoia but we have seen high quality photographs of type material of H. arenacea Seyrig, 1952, H. brevipennis Seyrig, 1952, H. elongata Seyrig, 1952, H. flexibilis Seyrig, 1952and H. gracilior Seyrig, 1952 from the Paris Museum of Natural History (https://science.mnhn.fr/taxon/genus/handaoia) and of Hemiteles bellicornis Thomson, 1888 in the Biological Museum, Lund University (photos on Flickr: https://www.flickr.com/). The species we include in Handaoia are from widely scattered parts of the world, but they share a distinctive suite of characters, unique in combination, so we include them all in Handaoia. Handaoia bellicornis and the unidentified specimen illustrated in Townes (1970) have the area superomedia separated from the area basalis, whereas the Malagasy and New World species have these areas confluent. Without checking specimens it is difficult to evaluate the presence or absence of the carina across the dorsal pronotal groove, which seems to be absent in some species.