A revision of the Chilean Brachyglutini – Part 8. Revision of Achilia Reitter, 1890: A. sinuaticornis, A. kindermanni, A. humidula, A. praeclara, A. nigrita, A. rufula species groups, and Achilia incertae sedis species (Coleoptera: Staphylinidae: Pselaphinae)

Abstract: The Achilia sinuaticornis, A. kindermanni, A. humidula, A. praeclara, A. nigrita, and A. rufula species groups sensu Jeannel (1962 and 1963), and Achilia incertae sedis species according to Jeannel (1962) and Franz (1996) of the species-rich genus Achilia Reitter, 1890 are revised. Of the seventeen taxa placed in these seven species groups (including incertae sedis species), seven names are placed in synonymy: A. curta Jeannel, 1962 = A. longiceps (Reitter, 1885) syn. nov., A. elongata Jeannel, 1962 = A. humidula (Reitter, 1885) syn. nov., A. occipitalis Jeannel, 1962 = A. humidula (Reitter, 1885) syn. nov., A. diademata Jeannel, 1962 = A. humidula (Reitter, 1885) syn. nov., A. parvula Jeannel 1962 = A. humidula (Reitter, 1885) syn. nov., A. dicastrii Franz, 1996 = A. bicornis Jeannel, 1962 syn. nov., A. latifrons Raffray, 1904 = A. longiceps (Reitter, 1885) syn. nov. Achilia nigrita Jeannel, 1962 belongs to a different genus and will be treated in forthcoming papers. The remaining ten species are redescribed, and two new species belonging to the A. sinuaticornis group are described: A. fokkata sp. nov. and A. lapsus sp. nov. The lectotypes of A. longiceps (Reitter, 1885), A. convexiceps Raffray, 1904, A. quadraticeps Raffray, 1904, and A. latifrons Raffray, 1904 are designated. For all these species their distribution is detailed and mapped, and habitat/collecting data are summarized.

Paired structures of the body (such as foveae) are treated as plural, while parts of the paired appendages are treated as singular to simplify the characterizations.The body length is measured from the anterior clypeal margin to the posterior margin of the last visible abdominal tergite.The length and width of the body parts were measured between points of maximum extension, e. g. the head length is measured between the anterior clypeal margin and the posterior margin of the neck, the head width includes the eyes, the elytral length along the suture line, and the elytral width is the total width of the two elytra taken together.The abdominal tergites are numbered based on order of visibility.Morphological terminology follows that of Chandler (2001), except our use of 'ventrite' instead of 'sternite' when describing meso-and metathoracic structures, and the sclerotized features of the aedeagus termed "dorsal strips" in Sabella et al. (2017) are here termed "longitudinal struts".

TAXONOMY
Achilia sinuaticornis species group Jeannel (1963: 353, 366) characterized this group as follows: elytra with three basal foveae; first abdominal tergite with long and oblique basal striae separated by one third of tergal width; anterior half of male head modified; male antennomeres modified; aedeagus quadrangular with dorsal longitudinal struts subparallel; parameres relatively wide with one seta on short outer lobe; copulatory pieces bifid, and apically recurved.According to Jeannel (1963: 366-367) this group is closely related to the A. grandiceps group.It included only A. sinuaticornis, to which we add two new species described below -i.e. A. fokkata sp.nov.and A. lapsus sp.nov.-for a total of three species.The species of the A. sinuaticornis group possess the following common features: pubescence decumbent, consisting of long setae sparse over entire body; head wider than long, strongly modified in male, due to expanded vertexal foveae not visible in dorsal view, while for females these vertexal fovea are big; eyes moderately protruding and longer than tempora; maxillary palpi small with last palpomere elongate; antennae with antennomeres strongly modified in male; pronotum wider than long and wider than head with disc slightly convex, disc surface smooth and shiny with only few scattered punctures; median antebasal fovea smaller than lateral foveae; anterior portion of lateral margins of pronotum distinctly convergent and sinuate anteriorly; posterior portion of lateral margins of pronotum slightly convergent; basal margin of pronotum bordered with row of contiguous shallow impressions; elytra together wider than long with protruding humeri; elytral disc smooth, shiny, elytra with three big basal foveae, sutural stria entire; elytral discal stria extending to about elytral midlength; abdomen smooth, with some minute punctures;

MATERIAL AND METHODS
This study is based on the examination of 1616 specimens.The acronyms used in the present study refer to the following collections ( Chandler) Only critical references are cited for the species.Under the sections "type material" or "additional material" the locality data are standardized, with indications of major administrative units (Regions and Provinces) and names of the collectors; for the holotypes of older specimens the labels are also given verbatim.For MHNG material additional information pertaining to sampling sites are enriched from unpublished locality lists when available.For the method of selection of the type material see Sabella et al. (2017).The aedeagi and other body parts illustrated here were mounted in Canada balsam on acetate slides, and drawn using a drawing tube mounted on a Zeiss Axioskop compound microscope.Images were taken using a Leica DFC425 camera in conjunction with a Leica M205-C compound microscope.Zerene Stacker (version 1.04) was used for image stacking.All images were modified and grouped using Adobe Photoshop CC and Illustrator CS6.
first abdominal tergite distinctly longer than second and with diverging basal striae extending to about one-third of paratergal length, separated at base by about one-third of tergal width with short and sparse setal brush between basal striae.In order to keep the text concise, these features are not repeated in their respective descriptions of the species of this group.
Female: Similar to male except head, antennae, metaventrite and legs unmodified.
Collecting data: Collected from August to January; found in different types of forests, sometimes with Araucaria araucana and Chusquea, at elevations ranging from 300 m to about 1300 m.Most of the material come from sifted samples of leaf and log litter; some specimens were collected with windows traps.
Distribution: Achilia sinuaticornis is known from the Araucanía and Valparaíso Regions (Fig. 117 squares edged in blue).

Comments:
The species was described from a single male collected in Cerro El Roble on 18.VIII.1961,not on X.1961 as stated by Jeannel (1963: 353), a contradiction to what he stated on page 351 of the same publication where he reported the correct date of August 1961.We have examined this unique specimen which, however, did not carry any label designating it as a holotype; furthermore, the determination label that was handwritten by Jeannel bears the caption "Achilia torticornis" instead of "Achilia sinuaticornis".antennae (Figs 11,66,71), and aedeagus (Fig. 3).The females of A. sinuaticornis are very similar to those of A. lapsus sp.nov.; for characters to distinguish females of these two species see the "Comments" section of the latter species.(9)(10)(11) antennae, (12)(13)(14) antennomere 11, (15-17) mesofemora, and protibiae in (18, 19, 21, 23) lateral and (20, 22, 24) dorsal views.testaceous with elytra, antennae and legs reddish, palpi yellowish; some specimens with dark, almost blackish color likely resulting from some kind of oxidation during their conservation in alcohol and acetic acid.

Despite
Male: Head as in Figs 77-80 with occipital region very rounded, strongly projecting anteriorly above deep excavation of anterior half of head; occipital region with marked V-shaped medial impression; anterior margin of occipital region with large medial incision with margins densely pubescent; anterior excavation with upwardly oriented medial apophysis originating within incision of anterior border of occipital region, this apophysis apically enlarged and covered with dense pubescence consisting of short yellowish setae; anterior and lateral sides of excavation sharp and densely covered with long setae; frontal lobe flattened and lightly punctate.
Antennae (Figs 10,81) with scape longer than wide; pedicel wider than long; antennomere III-IX wider than long; antennomere X distinctly longer than wide, about as wide as antennomere XI, with posterior margin bearing some thin setae; antennomere XI longer than wide, longer than V-X combined, its surface bearing scattered tubercles.Distal half of metaventrite convex.Legs with mesofemora more or less enlarged with long and sparse setae on entire distal margin (Fig. 17).Aedeagus (Fig.  3).In addition, the distal half of the metaventrite of A. lapsus sp.nov. is convex, while that of A. sinuaticornis possesses a medial ovoidal impression bearing long setae.The females of A. lapsus sp.nov.are very difficult to distinguish from those of A. sinuaticornis; the only difference we noted, which remains difficult to appreciate, pertains to the medial occipital region of the head, which is more elevated and more convex in the new species.
Achilia kindermanni species group Jeannel (1962: 398, 431) characterized this group as follows: elytra with three basal foveae; basal striae of first abdominal tergite separated at most by a quarter of the tergal width; frontal lobe of male head modified; male antennomeres unmodified; funicular antennomeres distinctly longer than wide; aedeagal parameres without long apophysis at insertions of setae.
According to Jeannel (1962) this group of species consists of A. cordicollis Raffray, 1904, A. curta Jeannel, 1962, A. kindermanni (Reitter, 1883), and A. longiceps (Reitter, 1885).However, after examination of the types of these taxa we concluded that A. curta is a junior synonym of A. longiceps (Reitter, 1885) (syn.nov.).In addition, we examined the types of A. latifrons Raffray, 1904, which is treated in this chapter for convenience although it technically belongs to the "Species incertae sedis" group according to Jeannel (1962) and Franz (1996), and concluded that it is also a junior synonym of A. longiceps (Reitter, 1885) (syn.nov.).Consequently the A. kindermanni group contains now only A. kindermani, A. cordicollis, and A. longiceps.The species of the A. kindermanni group possess the following shared features: pubescence decumbent, consisting of long setae sparse over the body, setae very sparse on elytra; head slightly wider than long, vertexal foveae at level of about half length of eyes; eyes longer or slightly longer than temples; antennae with antennomeres III-VIII about same width; pronotum wider than long, slightly wider than head; disc slightly convex, smooth and shiny with some punctures; median antebasal fovea about as wide as lateral ones (smaller than lateral one in A. longiceps); anterior portion of lateral margins of pronotum distinctly convergent and sinuate anteriorly; posterior portion of lateral margins of pronotum slightly convergent; basal margin of pronotum bordered with row of contiguous shallow impressions; elytra together wider than long with very protruding humeri; elytral disc smooth, shiny, with very few punctures; elytra with four basal foveae, outer two closely spaced fovae merged together in some specimens; sutural stria entire; elytral discal stria extended to about elytral mid-length; abdomen smooth, with some minute punctures; first abdominal tergite with subparallel basal striae extending to about one-fourth of paratergal length and separated at base by about one-third of tergal width, with short and very sparse setal brush between basal striae.In order to keep the text more concise, these features are not repeated in the following descriptions.

Achilia cordicollis Raffray, 1904
Achilia cordicollis Raffray, 1904: 136;Jeannel, 1962: 431, 434.Comments: Achilia kindermanni was described by Reitter (1883: 51) based on a single male specimen from Valdivia.We identified the holotype as a male specimen from the Raffray collection of the MNHN that matches the original description, comes from Valdivia, and moreover bears locality and identification labels handwritten by Reitter (see section at the beginning of this species treatment covering "Type material").The males of A. kindermanni are easily distinguished from those of the other species of the A. kindermanni group by the shape of the head (Figs 82-85), antennae (Figs 43,86), and aedeagus (Fig. 25).The females of A. kindermanni are similar to those of A. longiceps; for characters to distinguish females of these two species see the "Comments" section of the latter species.
Achilia longiceps (Reitter, 1885) [39][40][41][42][87][88][89][90][91][92][93][94][95][96]118 Bryaxis longiceps Reitter, 1885: 327 pl. 2 fig.8 (head and antenna of male).Achilia longiceps Jeannel, 1962: 431, 432  Comments: Reitter (1885: 323, 327-328) described A. longiceps based on an unknown number of males collected from Valdivia, specifying that the female was unknown.Jeannel (1962: 433) redescribed this species mentioning that 5 males collected in Valdivia are present in the MNHN collections, and also stated that the female was unknown.In the Raffray collection housed in the MNHN we found indeed 5 males identified as A. longiceps by both Raffray and Jeannel, with the first male of this series bearing a "TYPE" red label.Despite the fact that these 5 specimens are labelled as from "Chili" without further indication, we consider that there is no reason to doubt that these 5 males belong to the series used by Reitter to describe this species.The male bearing the red label "TYPE", which corresponds perfectly to Reitter's original description, is therefore designated here as the lectotype of A. longiceps, and the remaining 4 males as paralectotypes.Jeannel (1962: 433) described A. curta on the basis of one male from Chepu (Chiloé province), and one male and one female from La Esperanza (Llanquihue province), stating that the holotype of A. curta from Chepu was housed in the MNHS.We examined the specimen listed in the catalog of the MNHS holotypes of insects (Camousseight, 1980: 17): it is indeed a male of A. curta, but labeled as from "La Esperanza".On the other hand, we found in the MNHN collection one male of A. curta labeled as from Chepu, identified as such by Jeannel and bearing a "TYPE" red label.So, the latter specimen housed in the MNHN is the holotype of A. curta.
According to Jeannel (1962: 433) the males of A. curta differ from those of A. longiceps by the shorter and narrower head, the shorter antennae, and the aedeagus with copulatory pieces longer and slenderer.We have compared the lectotype and paralectotypes of A. longiceps with the holotype of A. curta, and their aedeagal conformation is nearly identical .Furthemore, after examination of extensive additional material, our opinion is that the morphological differences between these two types fall within the intraspecific variability of what we hold as one species.Consequently, we place A. curta as a junior synonym of A. longiceps (syn.nov.).
For convenience we will deal here with A. latifrons Raffray, 1904, although it technically belongs to the "Species incertae sedis" according to Jeannel (1962) and Franz (1996).Raffray (1904: 138) described A. latifrons based on an unknown number of specimens from Chile, without further geographic indication.In the Raffray Collection housed in the MNHN we found three females identified as such by Raffray and labeled as from "Chili", one of them bearing also a red "TYPE" label.Jeannel (1962: 433) redescribed this species and mentioned only three females from Chile as being present in the MNHN collection.We consider that there is no reason to doubt that these three females belong to the series used by Raffray to describe this species.The female bearing the red "TYPE" label, which corresponds perfectly to Reitter's original description, is here designated as the lectotype of A. latifrons, and the remaing two females as paralectotypes.
We have closely examined the types of A. latifrons, which fit perfectly in our concept of A. longiceps (see treatment of this species above).Consequently, we place A. latifrons as a junior synonym of A. longiceps (syn.

nov.).
The males of A. longiceps are easily distinguished from those of the other species of the A. kindermanni group by the shape of the head (92)(93)(94)(95), antennae 91,96), mesotrochanters , mesotibiae (Fig. 39), and aedeagus (Fig. 26).The females of A. longiceps are easily distinguished from those of A. kindermanni by the subparallel lateral margins of the head and the frontal lobe wide in A. longiceps, while in A. kindermanni the lateral margins of the head are anteriorly convergent and the frontal lobe is narrow.
Achilia humidula species group Jeannel (1962: 398, 435) characterized this group as follows: elytra with three basal foveae; basal striae of first abdominal tergite separated at most by a quarter of tergal width; frontal lobe of male's head modified; male's antennomeres unmodified; funicular antennomeres not longer than wide; aedeagal parameres with denticulate distal margin; aedeagal internal sac with a pair of copulatory pieces.
According to Jeannel (1962) this group contained three species: A. elongata Jeannel, 1962, A. humidula (Reitter, 1995) and A. occipitalis Jeannel, 1962.According to Sabella et al. (2017: 120) we should also add to these species A. parvula Jeannel, 1962.However, after close examination of the holotypes of these taxa as well as that of A. diademata Jeannel, 1962 (the latter species assigned by Jeannel (1962) to the group of A. praeclara) we came to the conlusion that these four names are junior synonyms of A. humidula (syn.nov.).Consequently, the A. humidula group is reduced to only A. humidula.
According to Jeannel (1962: 438-439) A. diademata could belong to the A. praeclara group.However, we have compared the holotype of A. humidula with the types of A. diademata and their aedeagal conformation is nearly identical .In our opinion the few subtile differences between the males of these series fall within the intraspecific variability of what we hold as one species.Consequently, we place A. diademata as a junior synonym of A. humidula (syn.nov.).Jeannel (1962: 435) described A. elongata on the basis of one male collected in Chepu on 13.X.1958,stating that this holotype was housed in the MNHS.However, in the MNHS collections we could not find any specimens of A. elongata and this species is not listed in the catalog of the MNHS holotypes of insects (Camousseight, 1980).Instead, we found in the MNHN one male of this species identified as such by Jeannel and labeled as being from Chepu, and bearing a red "TYPE" label.Our opinion is that there is no reason to doubt that this specimen is the holotype of A. elongata.
According to Jeannel (1962: 435-436) the male of A. elongata differs from that of A. humidula by its larger Female: Body 1.55 mm long, elytra reddish, palpi yellowish.Head shining, with sides anteriorly convergent; frons distinctly raised at middle, frontal lobe with marked transverse sulcus; tempora convex, about as long as eyes.Antennae with scape and pedicel slightly longer than wide; antennomeres III-VIII about as long as wide; antennomeres IX-X wider than long; antennomere X distinctly larger than IX; antennomere XI longer than wide, about as long as VII-X combined.Elytra with three basal foveae; humeral area not very prominent.Distal half of metaventrite slightly impressed at middle.Legs with posterior margin of protrochanters pointed at middle.Comments: Reitter (1885: 325, 329) described A. praeclara based on one male from Valdivia, mentioning that the specimen was not in a good state of preservation, and without further details.Jeannel (1962: 439-440) redescribed this species, mentioning that only one male from Valdivia was present in the MNHN collection, adding that this specimen had the distal half of the elytra damaged.After a careful search through all of the MNHN collections, we found one male labeled as from "Chili", identified by both Raffray and Jeannel as A. praeclara, and bearing a red "TYPE" label.This specimen is indeed in a poor state of preservation (in particular the last antennomeres of both antennae are missing), but its elytra are still intact.The specimen carries only a "Chili" location label without further precision, and its state of conservation doesn't match the details given by Jeannel.However, it is otherwise a male in perfect accordance with Reitter's original description, and we believe that it is indeed likely the holotype of A. praeclara.The males of A. praeclara are easily distinguished by the shape of the head (Figs 102-105), antennae (Figs 54, 106), protrochanters (Fig. 58), mesotibiae (Fig. 61), and aedeagus (Fig. 52).

Achilia quadraticeps Raffray, 1904
Achilia quadraticeps Raffray, 1904: 133;Jeannel, 1962: 438, 440.These two females correspond perfectly with Reitter's original description of A. quadraticeps, and we believe that there is no doubt that they belong to the type series.Consequently, we designate the specimen bearing the red "TYPE" label as the lectotype of A. convexiceps, and the other specimen as a paralectotype.According to Raffray (1904: 137) this species is similar to A. picea Raffray, 1904, which we synonymized with A. elfridae Raffray, 1904(see Kurbatov et al., 2021).We share his opinion that this species resembles more closely species of the A. cosmoptera group than that of the A. praeclara group.Nevertheless, in absence of male specimens, we will follow the current placement until the reliability of the species groups within Achilia are reassessed.
Female: Similar to male except head, antennae, and legs unmodified, and metaventrite slightly impressed.
Collecting data: Collected from September to April; mainly collected in forests of Nothofagus spp.and mixed forest at elevations ranging from 250 m to about 1400 m.Many specimens, especially males, came from FIT and car netting, but also were taken from sifted samples of leaf and log litter.
Distribution: Achilia simulans occurs in the Regions of Araucanía, Bío-Bío, Los Ríos and Maule (Fig. 118, blue triangles).Of the 287 specimens we have examined of this species it appears that the only record for the Los Ríos Región is the holotype, from Valdivia.As the holotype has only "Chili" on its locality label it could be questioned wether it really comes from Valdivia (see paragraph "Comments" below).
Comments: Reitter (1885: 325, 328-329) described A. simulans based on a single male from Valdivia.Jeannel (1962: 433) redescribed the species mentioning only one male from Valdivia that was present in the MNHN collection, stating that the female was unknown.
In the MNHS collections we found one male identified by both Raffray and Jeannel as A. simulans, and bearing a red "TYPE" label.Although this specimen is only labeled as being from "Chili" without any further indication, it corresponds perfectly with the descriptions of Reitter and Jeannel, and we believe that this is the holotype of A. simulans.The males of A. simulans are easily distinguished by the shape of the head , antennae 111,116), mesofemora (Fig. 59), mesotibiae (Fig. 61), and aedeagus (Fig. 53).The females of A. simulans are also easily distinguished by the shape of the head, which has the posterior portion slightly raised, and the anterior portion elongate with slightly convergent margins that are separated from the clypeus by a distinct inverted V-shaped sulcus.

Achilia nigrita Jeannel, 1962
Achilia nigrita Jeannel, 1962: 442.After examination of the holotype and only known specimen of A. nigrita, it appears that it likely belongs to the genus Achillidia Jeannel, 1962.However, we will formally transfer it only in our next paper of this series, which will be dedicated to all the other genera of Chilean Brachyglutini.
Achilia rufula species group Jeannel (1962: 399, 442) the significant variability shown by the males with respect to the morphology of the head (Figs 62-65 and 67-70), antennomeres (Figs 66 and 71), protibiae (Figs 18-24), protarsi (Figs 5-8), and mesofemora (Figs 15-16), the morphology of the aedeagus in all the specimens examined is identical to that shown by Fig. 3.For this reason, we prefer to consider A. sinuaticornis a single species characterized by high variability, rather than describing new taxa possessing undiscriminant genitalia.The males of A. sinuaticornis are easily distinguished from those of the other species of the A. sinuaticornis group by the shape of the head (Figs 62-65 and 67-70),
Jeannel (1962: 434)placed it with uncertainty in the A. kindermanni group, and mentioned that the discovery of the male might result in the separation of this species from the genus Achilia.Achilia kindermanni is widespread in Southern and Central Chile, where it was found from the Los Lagos to the Maule Regions (Fig.117red circles).
Female: Similar to male except head, metaventrite, abdominal sternites, and legs unmodified.Collecting data: Chile that were present in the MNHN collection.The Raffray collection which is housed in the MNHN holds two females from "Chili", identified by both Raffray and Jeannel as A. convexiceps", with one of these two specimens bearing a red "TYPE" label.