A comparative morphological revision of the aphid genus Myzaphis van der Goot, 1913 (Insecta: Hemiptera: Aphididae) revealed a new genus and three new species

The aphid genus Myzaphis van der Goot, 1913 from the tribe Macrosiphini is revised to include eight species. Apterous and alate viviparous females, known fundatrices and known sexual morphs (oviparous females and males) of Myzaphis bucktoni, M. juchnevitschae, M. rosarum, M. tianshanica and M. turanica are re-described and illustrated. Lectotype and paralectotypes of Myzaphis bucktoni and M. turanica are designated. The status of M. komatsubarae nomen dubium is discussed. Myzaphis avariolosa is regarded as a species belonging to the genus Ericaphis. Three new species: M. oezdemirae Kanturski & Barjadze sp. nov., M. tuatayae Kanturski & Barjadze sp. nov. from Turkey and M. rezwanii Kanturski & Barjadze sp. nov. from Iran are described and illustrated. Myzaphis bucktoni is recorded from Portugal for the first time. Diagnosis of the genus Myzaphis van der Goot, 1913 is redefined and a new genus Richardsaphis Kanturski & Barjadze gen. nov. is erected with the type species R. canadensis (Richards) comb. nov. Richardsaphis is for the first time recorded from the USA and hitherto unknown oviparous female and alate male are described and illustrated. Original keys to species of the genus Myzaphis and aphid genera of the tribe Macrosiphini with 2-2-2 first tarsal chaetotaxy are also provided.


Introduction
The aphid genus Myzaphis is represented by eight species worldwide [1,2]. Members of this genus are characterized by a small, elongate-oval body and characteristic rugose or wrinkled dorsal cuticle [3]. They live on undersides of leaves of Dasiphora and Rosa spp. (Rosaceae), are monoecious holocyclic, rarely anholocyclic and not visited by ants [4]. Myzaphis rosarum (Kaltenbach, 1843) and M. turanica Nevsky, 1929 are pests of the cultivated rose [5]. Five species-M. avariolosa David, Rajasingh & Narayanan, 1970, M. canadensis Richards, 1963, M. juchnevitschae , M. komatsubarae Shinji, 1922   are characterized by a well-developed socket, the basal part round and the apex flattened, often wider than the base (Fig 2C). Rostrum is pointed, with few setae: two on segment III, and 12 on segment IV+V, four of which are accessory setae (Fig 2D). Tarsi are characterised by short HT I and HT II with short setae (Fig 2E). On the ventral side of HT I two kinds of setae are visible: four long, hair-like and pointed and one short and rigid central "sense peg" (Fig 2F). The end of HT II is characterized by normal shaped and pointed claws and long parempodia which are rounded from the basal part to about half of their length and then flattened with slightly spatulate apices ( Fig 2G). In M. rosarum the dorsal sculpture seems to be strongly wrinkled (Fig 1) but higher magnification shows it is more or less regular, small, rounded or oval cavities or depressions (Fig 2H and 2I). The dorsum is also characterized by few and inconspicuous setae which are similar to those on the head. Abdominal setae are also very short, rigid, robust or thick with wide, rounded and high sockets, basal part tubular and apices rounded or clubshaped ( Fig 2J). The SIPH are more or less straight or slightly curved with the surface imbricated or wrinkled ( Fig 2K). Apex of the SIPH with a well-developed, strong flange and welldeveloped operculum on the siphuncular pore. The SIPH without apical reticulation ( Fig 2L). Cauda can be differently developed but always is more or less tongue-shaped, triangular in dorsal and ventral view (Fig 2M and 2N) and oval from lateral side (Fig 2O), with six or seven long, fine and pointed setae. Anal plate and genital plate are covered by fine and pointed setae, but those on genital plate are less numerous and much shorter (Fig 2N and 2O). Sensilla of the genus Myzaphis. The antennal sensilla are general divided into campaniform, trichoid, placoid and coeloconic sensilla. On the pedicel, there are two kind of sensilla: single rhinariolum on the ventral side, which is characterised by small (diameter about 2μm) and rounded opening and one peg-like sunken coeloconic sensillum with 6-8 very short projections (Fig 3A and 3B). Dorsal side of the ANT II bears flat and rounded campaniform sensillum with rounded central part ( Fig 3C). Surface of antennal segments is covered by clearly visible rings of folds, wrinkles or imbrications that are poorly separated from each other ( Fig  3D, 3E and 3G). Antennal segment III and IV are covered only by very few type-I trichoid sensilla which are very similar as those on other body parts. The mentioned sensilla are very short, tubular on the basal part with rounded or club-shaped apices. The sockets are strongly developed, rounded basally and slightly cone-shaped ( Fig 3F). On ANT V, near the apex there is a rounded sclerotic ring with very well developed, thick and rigid projections of different length ( Fig 3G). The long projections are slightly spatulate and the shorter ones are characterized by more pointed apices. Deep inside the sclerotic ring a big placoid multiporous sensillum is present (Fig 3H and 3I). On ANT VI, on the border between the basal part and the PT there is also a cavity, surrounded by very well-developed sclerotic ring with different shaped projections around and inside. Inside the cavity three kinds of sensilla are present and lie tightly next to each other (Fig 3K). The rounded, big placoid multiporous sensillum is visible under the separate and rounded ring of projections and its structure is the same as in the big placoid sensillum on ANT V (Fig 3L). Near the big placoid sensillum two slightly oval small placoid sensilla can be noted (Fig 3K and 3M). Between and next to the small placoid sensilla there are also 2-3 sunken coeloconic sensilla with 6-7 projections visible (Fig 3K, 3M and 3N).
The mouthparts are covered mainly by trichoid sensilla. URS is covered by three types of sensilla: one pair of fine and pointed type II basiconic sensilla on the proximal part of segment IV, three pair of long, fine and pointed type II trichoid sensilla ( Fig 3O) and eight pairs of very short, rigid and pointed type III basiconic sensilla on the poorly separated segment V (Fig 3P). SEM of antennal and mouthparts sensilla of the genus Myzaphis: A. very short and rigid type I trichoid sensilla and small rounded rhinariolum (white arrow) on the pedicel. B. structure of the rhinariolum showing sunken peg-like sensillum with 6-8 projections (dotted arrow). C. rounded campaniform sensillum on the pedicel (star) and structure of imbrications on ANT III (asterisks). D. very short, rigid type I trichoid sensilla on ANT III with very scarcely developed border between ANT III and IV (arrow). E. very short, rigid type I trichoid sensilla on ANT III with very scarcely developed border between ANT III and IV (arrow). F. structure of the type I trichoid sensillum on ANT. G. very short and rigid type I trichoid sensilla and big placoid sensillum on ANT V. H. structure of big placoid sensillum on ANT V under the very well developed sclerotic ring. I. structure of the sclerotic ring projections on big placoid sensillum on ANT V. J. type I trichoid sensilla and placoid sensilla on base (BASE) and very short type II trichoid sensilla on the end of processus terminalis (PT) on ANT VI. K. one big placoid sensillum (arrow head), two small placoid sensilla (arrows), and sunken coeloconic sensilla (dotted arrow) on the BASE of ANT VI. L. structure of big placoid sensillum on ANT VI under the very well developed sclerotic ring. M. structure of small placoid sensillum on ANT VI under the very well developed sclerotic ring. N. structure of sunken coeloconic sensillum on ANT VI under the very well developed sclerotic ring. O. URS with type II basiconic sensilla (arrow), type I trichoid sensilla (white asterisk) and type II basiconic sensilla (black asterisk). P. anterior view of the fifth rostral segment showing eight pairs of type III basiconic sensilla around the stylet opening (star). https://doi.org/10.1371/journal.pone.0193775.g003

Myzaphis van der Goot, 1913
Myzaphis van der Goot, 1913 [12]: 96. Francoa Del Guercio, 1917 Type species: Aphis rosarum Kaltenbach, 1843: 101 [11], by original designation Diagnosis. Small, spindle-shaped, or oval aphids with short appendages. Head with weakly developed antennal tubercles, but a characteristic feature of the genus is a strongly projecting quadrate or rounded median tubercle. ANT only about half of body length, without secondary rhinaria in apterae. Alatae have secondary rhinaria on ANT III only, or on ANT III-IV. Dorsal body setae are blunt and somewhat capitate. First tarsal segments all have 5 setae. The dorsum of the aptera is sclerotic and wrinkled or ornamented with numerous small rounded depressions. Alatae have dusky or dark sclerotic markings, often forming a central dorsal abdominal patch. Spiracular apertures are partly covered by opercula. SIPH are elongated, cylindrical for much of their length with the distal part often curved outwards and slightly swollen, and with a small, but distinct flange. The cauda is tongue-shaped or elongated triangular [18].
List of species: 1. M. bucktoni Jacob, 1946 2. M. juchnevitschae  3. M. komatsubarae Shinji, 1922   Apterous viviparous females. BL 1.04-2.45 mm long. Body shape oval or spindle-shaped. Head with well-developed compound eyes and triommatidia. Antennal tubercles weaklydeveloped or undeveloped. Frons with median tubercle or flat. Median tubercle or the median area of frons with 2-4 setae, longer or shorter than BD III, blunt or pointed. ANT 6-segmented, without secondary rhinaria. ANT III longest, ANT IV shorter or longer than ANT V. ANT V with one small, rounded primary rhinarium with ciliated edge. ANT VI with one small, rounded primary rhinarium with ciliated edge and 5-6 very small accessory rhinaria, tightly adhering to the primary rhinarium. ANT setae very short, never longer than the width of segments, with blunt apices. ANT I with 5-9, ANT II with 3-6 setae. PT with 3-4 apical setae. Rostrum reaching to middle coxae. Dorsal side of head, thorax and abdomen slightly sclerotized, more or less but evidently wrinkled. Setae on the dorsal side of body extremely short, inconspicuous, with blunt apices. Setae on legs short, never longer than the width of tibiae. First segment of tarsi with 5-5-5 setae. SIPH always longer than cauda, tubular, almost straight, inner side sometimes slightly curved. Cauda tongue-shaped with 6-7 pointed setae.
Alate viviparous females. Head and thorax strongly sclerotized. Head with big compound eyes, triommatidia and ocelli. ANT with secondary rhinaria on ANT III and very rarely also on ANT IV. Secondary rhinaria rounded or slightly oval, with smooth edge, arranged irregularly or in 2-3 rows on the whole length of the segment. ANT III longest, ANT IV shorter or longer than ANT V. ANT V with one small, rounded primary rhinarium with ciliated edge. ANT VI with one small, rounded primary rhinarium with ciliated edge and 5-6 very small accessory rhinaria, tightly adhering to the major rhinarium. ANT I with 3-9, ANT II with 3-6 setae. PT with 3-4 apical setae. ANT setae very short, never longer than the width of segments, with blunt apices. Body setae very short and inconspicuous, setae on legs never longer than the width of tibiae. Fore wings hyaline or slightly pigmented (yellow or pale yellow) with brown veins. Media twice branched. Hind wings with two oblique veins. Rostrum reaching to mesosternum. First segment of tarsi with 5-5-5 ventral setae. SIPH always longer than cauda, tubular, slightly swollen from the middle of their length. Cauda tongue-shaped with 6-7 pointed setae.
Oviparous females. General characters like in apterous viviparous females. Dorsal side of body much more membranous than in apterous viviparous females, only slightly wrinkled. ANT I with 3-7, ANT II with 4 setae. TIBIAE III much darker than other parts of body, swollen or normal with rounded pseudosensoria (scent plaques) arranged on almost whole length.
Alate males. Small apterous (then main morphological features like in apterous viviparous females) and very small alate (then main morphological features like in alate viviparous females). Dorsal side of body more or less sclerotized as irregular sclerites or cross-bars. ANT III-VI with not numerous, small rounded secondary rhinaria like in alate viviparous female. ANT I with 3-7, ANT II with 4-5 setae.
Apterous viviparous female-re-description (n = 94) (Figs 5A-8A; Table 2) Colour. Colour in life: pale yellow to pale green with light to dark brown dorsal markings in the form of a brown head, two large brown patches of pronotum and a pair of broad pleural stripes extending from the mesonotu almost to the base of cauda [19]. Colour in mounted specimens: ANT, legs and SIPH pale to yellow with brown to dark brown head, pronotum and two broad longitudinal stripes from mesonotum to the end of abdomen, ANT V, ANT VI and cauda ( Fig 5A).  long; on abdomen 0.005-0.031 mm long. SIPH 1.55-1.94 × cauda, almost straight, slightly curved inner side in 1/3 length. Cauda broadly tongue shaped ( Fig 8A).
Alate viviparous female-re-description (n = 13) (Figs 9A and 10A; Table 3) Colour. Colour in life: head, ANT and thorax dark brown to black. Legs brown to black. Abdomen pale green with light brown dorsal patch and darker green longitudinal pleural stripes [19]. Colour in mounted specimens: head, ANT and thorax brown to dark brown with    [19]. Colour in mounted specimens: head brown, pronotum light brown, the rest of thorax and abdomen yellow. ANT brown except ANT III. Femora yellow with light brown dorsal parts, fore and middle tibiae yellow with distal parts and tarsi brown. Hind tibiae and tarsi brown to dark brown, SIPH brown with paler basal part, cauda brown ( Fig 11A). Apterous male-re-description (n = 7) (Figs 13A and 14A; Table 5) Colour. Colour in life: Abdomen dusky olive green, legs dark brown. All other body parts black [19]. Colour in mounted specimens: Body mostly sclerotized. Head and thorax and sclerotized parts of abdomen brown. ANT dark brown almost black. Legs brown with only slightly paler inner sides. Genitalia, SIPH and cauda dark brown (Fig 13A).
Morphometric characters. Front of head broadly convex in dorsal view, with no distinct median frontal tubercle and two slightly pointed setae (Fig 6B) (Fig 8B).
Alate viviparous female-re-description (n = 1) (Figs 9B and 10B; Table 3) Colour. Colour in life: head, thorax and dorsal abdominal patch brown. Rest of abdomen and SIPH greenish yellow, cauda black [10]. Colour in mounted specimens: head, ANT and thorax brown. Fore and hind wings pale with brown veins. Pterostigma pale brown with slightly darker edges. Legs with light brown femora and yellowish tibiae with darker distal parts and tarsi. Abdomen pale with light brown dorsal sclerotic patch, marginal sclerites, SIPH and cauda (Fig 9B).
Morphometric characters. Head setae 0.012-0.015 mm long, HLS about 1.00 × BD III. ANT (Fig 14C) Table 2) Colour. Colour in life: unknown. Colour in mounted specimens: body in general pale brown or yellowish with slightly darker narrow longitudinal stripes in pleural region of abdomen ( Fig 5C).
Oviparous female-re-description (n = 4) (Figs 11B and 12B; Table 4) Colour. Colour in life: unknown. Colour in mounted specimens: head, ANT IV-VI, fore and middle legs, hind femora yellow to pale brown. Hind tibiae brown. The rest of body pale or yellowish (Fig 11B). Alate male-re-description (n = 3) (Figs 13E and 14E; Table 5) Colour. Colour in life: unknown. Colour in mounted specimens: Head and thorax sclerotized, brown. ANT III and genitalia light brown. The rest of ANT segments, legs, abdominal patch and SIPH pale to yellow. Fore and hind wings pale yellow with light brown veins. Pterostigma pale brown with very slightly darker edges (Fig 13E).
Morphometric characters. Head setae 0.007-0.017 mm long, HLS 1.00-1.41 × BD III. ANT (Fig 14E)  Remarks. We re-describe here the sexuales, because the first brief descriptions of these morphs were given by Tuatay and Remaudière [20] as sexual morphs of M. turanica.
Etymology. The authors have the pleasure in naming the new species to honour our colleague Dr. Isil Ö zdemir, aphid taxonomist in Turkey. Fundatrix-description (n = 21) (Fig 4B; Table 1) Colour. Colour in life: unknown. Colour in mounted specimens: all body parts pale to yellowish. (Fig 4B).  Table 2) Colour. Colour in life: unknown. Colour in mounted specimens: all body parts pale yellow, with slightly darker cauda (Fig 5D). In some specimens, also ANT VI and tarsi darker.
Alate viviparous female-description (n = 2) (Figs 9C and 10C; Table 3) Colour. Colour in life: unknown. Colour on mounted specimens: head, ANT and thorax brown. Fore and hind wings yellowish with brown veins. Pterostigma pale brown with darker edges. Legs pale brown with distal parts of femora, tibiae and tarsi. Abdomen pale with dorsal sclerotic patch, SIPH and cauda pale brown (Fig 9C).
Alate viviparous female-re-description (n = 21) (Figs 9D and 10D; Table 3) Colour. Colour in life: unknown. Colour in mounted specimens: head and thorax dark brown to with lighter pronotum. ANT uniformly brown. Fore and hind wings pale yellowish with brown veins. Pterostigma pale brown with dark edges. Legs light brown with lighter proximal parts of femora and darker distal parts of tibiae and tarsi. Abdomen pale with pale brown dorsal sclerotic patch and marginal sclerites. SIPH and cauda pale brown (Fig 9D).

Colour.
Colour in life: yellow-green [10]. Colour in mounted specimens: body in general pale, only distal parts of ANT, tibiae and cauda yellow (Fig 5F).
Alate viviparous female-re-description (n = 2) (Figs 9E and 10E; Table 3) Colour. Colour in life: head, ANT, thorax, abdominal patch, SIPH and cauda brown. Unsclerotized part of abdomen and legs yellow [10]. Colour in mounted specimens: head, ANT, thorax and legs brown to light brown. Fore and hind wings pale with light brown veins. Pterostigma pale brown with slightly darker edges. Abdomen pale with light brown dorsal sclerotic patch, SIPH and cauda (Fig 9E).
Morphometric characters. Head setae 0.025-0.027 mm long, HLS 2.85-3.35 × BD III. ANT0.57-0.59 × BL and 0.28-0.29 × HW. ANT III with 3-18 secondary rhinaria (Fig 10E). Diagnosis. Among all known species of Myzaphis, known morphs of M. tianshanica differ from the others by having long, conspicuous and pointed setae on head and the dorsal side of thorax and abdomen, as opposed to short, inconspicuous and blunt or slightly capitate setae in other Myzaphis species.
Host plants.  Table 2) Colour. Colour in life: unknown. Colour in mounted specimens: body pale to yellowish with slightly darker cauda ( Fig 5H).
Alate viviparous female-re-description (n = 6) (Figs 9F and 10F; Table 3) Colour. Colour in life: unknown. Colour in mounted specimens: head and thorax brown, ANT and femora yellow to light brown, tibiae pale with distal yellow to light brown distal parts and tarsi. Wings pale with brown veins. Pterostigma pale brown. Abdomen pale with abdominal patch, SIPH and cauda yellowish (Fig 9F). Oviparous female-re-description (n = 8) (Figs 11E and 12E; Table 4) Colour. Colour in life: unknown. Colour in mounted specimens: head, femora, tibiae and SIPH yellow to light brown. ANT V, tarsi and cauda light brown to brown (Fig 11E). Apterous male-description (n = 2) (Figs 13D and 14D; Table 5) Colour. Colour in life: unknown. Colour in mounted specimens: Body mostly sclerotized. Head, ANT, legs, SIPH and cauda light brown to brown. Sclerotization of thorax and abdomen light brown (Fig 13D).
Morphometric characters. Head setae 0.005-0.017 mm long, HLS 1.00-1.06 × BD III. ANT (Fig 14D)  Distribution. In Europe, mostly in Mediterranean region (France, Spain, Italy-Sicilia, and Portugal-FIRST RECORD); the record in Sweden needs conformation. Also in the Middle East (Israel), Central Asia (Kazakhstan, Kirgistan, Tajikistan, Uzbekistan) and other parts of Asia (India and Mongolia). The detailed records for countries in Palaearctic are given in Holman [20].
Remarks on the taxonomic status of M. komatsubarae Shinji. From the very short and general description, "Characteristics: Body green to pale. Antennae infuscated throughout, III longer than IV and V taken together with about 19 subcircular sensoria, flagellum of VI about as long as base. Rostrum black throughout. Siphunculi black, cauda infuscated or black" [6], only little can be deduced. From the information about the secondary rhinaria on ANT III it is likely that the species was described from an alate viviparous female. Even if the antennae of alate some species of Myzaphis could be more or less brown to dark, neither the SIPH or cauda are dark or black. Also, the length of ANT III in contrast to ANT IV and V is a variable character not only among aphid genera but also among the species of Myzaphis, but in all alate viviparous females it is more or less slightly longer that the length of the two next ones.
It is impossible to establish the identity of the species as well as its generic affinity. Its hostplant being Sorbus commixta hints that it could be a species of Dysaphis Börner, 1931 [4].
Due to the absence of any available material, drawings or good description to evaluate this species, we consider Myzaphis komatsubarae Shinji, 1922 [8]. Colour in mounted specimens: head, ANT, legs, SIPH and cauda yellow. Thorax and abdomen pale (Fig 5I).
Morphometric characters. body spindle shaped with long ANT and legs. Head without median frontal tubercle with low but well developed and evident ANT tubercles. Dorsal and ventral side of head, especially near the ANT tubercles with numerous well-developed spicules ( Fig 6I). Dorsum not sclerotized, membranous without any sclerotized markings, smooth, never rugose or corrugated (Fig 7I). Dorsal setae on thorax 0.004-0.005 mm long; on abdomen 0.004-0.037 mm long, inconspicuous with blunt apices. SIPH 2.21-2.47 × cauda, almost straight, very slightly swollen in about of their length, imbricated. Cauda tongue-shaped with 4 setae (Fig 8I).
Diagnosis. Ericaphis avariolosa comb. nov. differs from all known species of the genus Myzaphis in development and shape of frons and ANT tubercles. This species is characterized by completely undeveloped median frontal tubercle and prominent, steep-sided ANT tubercles (completely undeveloped in remaining Myzaphis species, also these with low or undeveloped frontal tubercle, but then the frons often rounded). There is also one important difference between E. avariolosa comb. nov. and all known Myzaphis species-head surface with prominent, numerous spicules also on the ANT tubercles (head completely smooth in Myzaphis). Ericaphis species are known to have membranous or more or less sclerotic dorsal side of body, but like E. avariolosa comb. nov. their cuticle never forms any pattern, is not rugose nor wrinkled, which characterizes the members of Myzaphis. Other differences between this species and Myzaphis include: clearly longer femora and tibiae relative to body length, ANT longer than half of body length-0. We think that this species has affinity to Ericaphis Börner, 1939 based on combinations of the following characters: (1) well developed antennal tubercles; (2) presence of numerous Description. Apterous viviparous female. Small, spindle-shaped aphids with rather short appendages. ANT 6-segmented, only 1/3 of body length, without secondary rhinaria. PT shorter than BASE. The head has low antennal tubercles and rounded median tubercle. Rostrum short, not reaching to middle coxae. URS oblong triangular with blunt apices. First tarsal segments all have 2 ventral setae. Dorsal body setae are blunt and somewhat capitate. The dorsum is sclerotic and wrinkled. Subgenital and anal plates and cauda with spinulose imbrications. SIPH are rather long, with the distal part often slightly swollen, and with a small, but distinct flange. The cauda is tongue-shaped.
Alate viviparous female. It has secondary rhinaria on antennal segments III-IV and often V, and dusky sclerotic markings on abdomen or a central dorsal abdominal patch. Presence of central dorsal abdominal patch in alate viviparous females was not mentioned in the original description [7], but investigation of type material (2 alate viviparous females) shows us that they have a pale abdominal patch.
Etymology. The generic name Richardsaphis is of feminine gender and the authors have the pleasure in naming the new genus to honor Dr. W.R. Richards, who was an outstanding aphid taxonomist at the Canadian National Collection of Insects, Ottawa, Canada and "aphis" (plant louse).
Diagnosis. New genus resembles the genus Myzaphis by shape of antennal tubercles, siphunculi and cauda, absence of secondary rhinaria in apterae and presence of central dorsal abdominal patch on the abdomen in alatae. Both genera live on Rosa spp. They differ from each other by: (1) first tarsal chaetotaxy. Richardsaphis gen. nov. has 2:2:2, while it is 5:5:5 in Myzaphis; (2) dorsum of abdomen in apterae: sclerotic and wrinkled without numerous small rounded depressions in Richardsaphis gen. nov., while numerous, small, rounded depressions always present in Myzaphis; (3) presence/absence of secondary rhinaria on ANT V in alate females: secondary rhinaria often present on ANT V in Richardsaphis gen. nov., while they are always absent on ANT V in Myzaphis.
Morphometric characters. Head with low, rounded median frontal tubercles (Fig 15D). abdomen, SIPH and cauda yellowish to pale brown. Wings pale with brown veins. Pterostigma pale (Fig 16A). Colour. Colour in life: yellow-pink. Colour in mounted specimens: head, thorax and ANT yellow. Legs yellow with pale tibiae with darker distal parts. Wings pale to yellowish with yellowish veins. Pterostigma pale. SIPH and cauda pale. (Fig 18A).
Morphometric characters. Head setae 0.007-0.017 mm long, HLS 1.30-1.41 × BD III. ANT long, 0.89-0.92 × BL and 0.23-0.24 × HW. ANT III with 21 ( Fig 18B), ANT IV with 7-9 ( Fig 18C), ANT V 10-11( Fig 18D) and ANT VI with 7 secondary rhinaria ( Fig 18E). Distribution. This species was previously known only from the type locality in Canada but was recently also collected in the Jemez Mountains in New Mexico, USA. Environmental Protection, University of Silesia in Katowice, Poland) for the full access to the laboratory equipment. Mariusz Kanturski gratefully acknowledges the Scholarship for Outstanding Young Scientists from the Ministry of Science and Higher Education of Poland (1165/E-340/STYP/12/ 2017). The funder had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.